Articulated Coralline Algae
of the

Gulf of California, Mexico,

I: Amphiroa Lamouroux

Taran ek ~ JAMES: N. NORRIS -
ie WILLIAM [ JOHANSEN" =

seni OOO gabe Beenie ® i - eR

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SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES ¢e NUMBER 9

Articulated Coralline Algae
of the
Gulf of California, Mexico,
I: Amphiroa Lamouroux

James N. Norris
and H. William Johansen

A an c i all a
j 4
i he i eo}

PUBLICATIONS

SMITHSONIAN INSTITUTION PRESS
City of Washington
1981

JX 183 fS) WIRE AC IE

Norris, James N., and H. William Johansen. Articulated Coralline Algae of
the Gulf of California, Mexico, I: Amphiroa Lamouroux. Smithsonian Contributions
to the Marine Scrences, number 9, 29 pages, 18 figures, 1981.—Amphiroa (Coral-
linaceae, Rhodophyta) is a tropical and subtropical genus of articulated
coralline algae and is prominent in shallow waters of the Gulf of California,
Mexico. Taxonomic and distributional investigations of Amphiroa from the
Gulf have revealed the presence of seven species: A. beauvoisii Lamouroux, A.
brevianceps Dawson, A. magdalensis Dawson, A. misakiensis Yendo, A. rigida
Lamouroux, A. valonioides Yendo, and A. van-bosseae Lemoine. Only two of
these species names are among the 16 taxa of Amphiroa previously reported
from this body of water; all other names are now considered synonyms. Of the
seven species in the Gulf of California, A. beauvoisi, A. misakiensis, A. valonioides
and A. van-bosseae are common, while A. brevianceps, A. magdalensis, and A. rigida
are rare and poorly known. None of these species is endemic to the Gulf, and
four of them, A. beauvorsi, A. misakiensis, A. valonioides, and A. rigida, also occur
in Japan.

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 of eastern North America.

Library of Congress Cataloging in Publication Data

Norris, James N.

Articulated coralline algae of the Gulf of California, Mexico.

(Smithsonian contributions to the marine sciences ; no. 9)

Bibliography: p.

Includes index

Contents: 1. Amphiroa Lamouroux.

1. Corallinaceae. 2. Algae—Mexico—California, Gulf of. I. Johansen, H. William. I.
Title. III. Series.

QOK569.C8N67 589.4’1 81-607063 AACR2

Contents

PNG INOW eG onmn Cine wear teres eae eh ae ee ec. a a8
UE LUO MIG ATINOUIROUX aS lL Wary Ayres He ee ee cis aes ME ot
Key to the Species of Amphiroa
Amphiroa beauvoisi: Lamouroux
Amphiroa brevianceps Dawson
Amphiroa magdalensis Dawson
Amphiroa misakiensis Yendo
Amphiroa rigida Lamouroux
ALA DTODB, CONDOS SUSUGO) ask on 5s ebb ag a wee eee ene ee ee
Amphiroa van-bosseae Lemoine
Literature Cited
Index

ill

NM Ne He
SRO UO OOSINS SCs Ci Comoe INS

OD w

NO

Articulated Coralline Algae
of the
Gulf of California, Mexico,

I: Amphiroa Lamouroux

James N. Norris
and H. William Johansen

Introduction

Species of Amphiroa Lamouroux (Corallinaceae;
Cryptonemiales) are common in intertidal and
shallow subtidal zones in tropical and subtropical
areas of the world. In the Gulf of California,
Mexico, they are the most prominent articulated
coralline algae; two other coralline genera are
found there, Janza Lamouroux and Corallina Lin-
naeus. The fronds of Amphiroa are purplish, bushy
clumps | to 12 cm long, or turfs 1 to 2 cm thick.
They grow in a variety of habitats, but usually
require at least some degree of wave action.

A terminology has arisen with respect to artic-
ulated coralline algae (Johansen, 1974). The
fronds are made up of numerous jointed branches
consisting of calcified segments called “intergen-
icula.” The intergenicula are separated from one
another by uncalcified nodes, or “‘genicula” (e.g.,
Figures 2 and 5). Extending through a branch is
a core of medullary filaments with the cells in
tiers (e.g., Figure 4), which is surrounded by a
cortex and epithallus.

Of the three genera in the Gulf of California,
only Corallina Linnaeus is represented by a single

James N. Norns, Department of Botany, National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 20560.
H. William Johansen, Department of Biology, Clark University,
Worcester, Massachusetts 01610.

species, C. pinnatifolia var. digitata Dawson (1953),
a small plant distinguished from Amphiroa by its
percurrent axes, which are densely clothed with
lateral branches. /anza Lamouroux, the other ge-
nus, is represented in the Gulf by four species
having delicate fronds composed of branches less
than 0.5 mm in diameter. These species have
regular dichotomous branching and conceptacles
borne in single swollen chambers at the apices of
the intergenicula. Whereas, in Amphiroa, the
fronds are coarser, branches generally over 1.0
mm in diameter (except in A. valonioides Yendo
where they are less than 0.5 mm in diameter),
and usually the dichotomous branching 1s irreg-
ular. Also the conceptacles of Amphiroa are in-
variably borne on intergenicular surfaces, al-
though sometimes they may be nearly invisible,
especially where branches are relatively thick.
Amphiroa Lamouroux (1812:187) is a member
of the subfamily Amphiroideae Johansen (1969a:
47), tribe Amphiroeae Cabioch (1972:266) (see
also, Johansen and Silva, 1978, and Johansen,
1981, for review of supergeneric classification).
The species are distinguished from one another
frond
height, intergenicular width, intergenicular shape

by the following taxonomic features:

in cross-section, the number of medullary cell
tiers in genicula and plant habit (e.g., turfy,

2 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

recumbent, or erect and bushy). Although the
branching is basically dichotomous, it is often
irregular and congested, with the branches tend-
ing not to lie in one plane. As in other members
of the Corallinaceae (e.g., Johansen, 1976a), the
reproductive cells are produced within concepta-
cles which, in Amphiroa, are always borne on the
surfaces of the intergenicula (Figure 14). Each
conceptacle opens by a single pore, through which
the reproductive cells exit.

The Gulf of California is a discrete body of
water with unique characteristics (van Andel and
Shor, 1964, for review), representing the Cortez
Province of Briggs (1974) or the Gulf of California
biogeographical region of W. H. Adey and R. S.
Steneck (pers. comm.); yet none of the species of
Amphiroa are now considered endemic to the Gulf.
Furthermore, four of the species are present in
areas other than the eastern Pacific. Amphiroa
beauvorsit (as A. zonata), A. misakiensis, A. rigida, and
A. valonwoides are all present in Japan (Okamura,
1936:515-521; Chihara, 1970:71—72). It is inter-
esting to note that Dawson (1960:97) found sev-
eral non-corallinaceous species, e.g., Pachydictyon
coriaceum (Holmes) Okamura (Dictyotales), also
common to both the Gulf of California and Ja-
pan. Hommersand (1972) has also suggested a
close relationship between the algae of the Gulf
of California and Japan.

The first report of articulated coralline algae
occurring in the Gulf of California was by Hariot
(1895) who reported Amphiroa linearis Kutzing
(1858). It was more than one-half century before
another account of Amphiroa was reported in the
Gulf, with Dawson (1944:276-277) reporting
three species: Amphiroa pusilla Yendo (1902), A.
zonata Yendo (1902), and A. rigeda Lamouroux
(1816). Of these, the first two were originally
described from Japan, and the third from the
Mediterranean Sea. The bulk of information on
coralline algae from the Gulf of California, and
the Pacific coast of Mexico, is due to the efforts
of E. Yale Dawson (1944, 1953). The first detailed
treatment of these algae from the Gulf and Pacific
Mexico was Dawson’s (1953) report of 11 taxa of
Amphiroa from the Gulf of California: A. annulata

Lemoine (1929), A. zonata Yendo (1902), and
seven others, which were described as new, A.
annulata var. pinnata, A. taylorit, A. droueti, A. brev-
anceps, A. magdalensis, A. subcylindrica and A. fran-
ciscana var. robusta.

Shortly thereafter, Dawson (1959) listed the
algae collected on a cruise into the southern Gulf
of California, reporting several species of Amphiroa
previously recorded for this region and described
a new variety, A. diumorpha var. digitiformis. Finally,
Dawson (1966b) extended the known range of
Amphiroa franciscana Taylor (1945) var. franciscana
into the Gulf making a total of 16 taxa of Amphiroa
reported from this body of water. Dawson (1953:
136-137) however, recognized only 13 species
from the Gulf, noting that Hariot’s (1895:169)
report of A. linearis was probably A. zonata, in-
cluded A. rigida in A. subcylindrica, and assigned
specimens previously reported as A. pusilla (Daw-
son, 1944) to A. drouetti, A. dimorpha, and A. zonata.
Later papers reporting taxa of this genus from
the Gulf of California are by Dawson (1966a,
1966b) and Norris (1972).

Our study of the articulated coralline algae
revealed that in the Gulf of California there are
seven species of Amphiroa that can be separated
on morphological and anatomical bases. After
surveying the described taxa, we conclude that
the entities of Gulf Amphiroa are: A. beauvoisi
Lamouroux, A. brevianceps Dawson, A. magdalensis
Dawson, A. misakiensis Yendo, A. rgida Lamour-
oux, A. valonioides Yendo and A. van-bosseae Le-
moine. Four of the seven species in the Gulf, A.
beauvoisit, A. misakiensis, A. valonioides and A. van-
bosseae, are common and represented by many
collections. The remaining three, A. brevianceps, A.
magdalensis, and A. rigida, are known only from a
few collections, and less confidence can be ex-
pressed in defining them.

MaterRIALs AND MetHuops.—Most of the collec-
tions studied were dried or preserved in 5 percent
Formalin/seawater; individual pieces of the
branches were fixed and decalcified in Susa fixa-
tive (Suneson, 1937:5). For anatomical studies,
material was embedded in paraffin, sectioned (8-

NUMBER 9

10 zm) and stained in Delafield’s haematoxylin
(Humason, 1967:142).

Most of the specimens studied were collected
by James N. Norris (JN), E. Y. Dawson (EYD),
H. William Johansen (HWJ), Katina E. Bucher
(KB), or W. R. Taylor (WRT). These specimens,
as well as those collected by others, are housed in
the following herbaria (abbreviations from Holm-
gren and Keuken, 1974):

AHFH_ Allan Hancock Foundation Herbarium
University of Southern California, Los Angeles
ARIZ _ Algal Collection (with Dr. Robert W. Hoshaw)

University of Arizona, Tucson
BM British Museum (Natural History)
London
CN Laboratoire de Botanique
Faculté des Sciences, Caen

CUW — Clark University
Worcester, Massachusetts

K Herbarium, Royal Botanical Garden
Kew:

L Rijksherbarium,
Leiden

LAM Algal Collection (now at AHFH)

Los Angeles County Museum

Los Angeles, California
MEXU Instituto de Biologia
Universidad Nacional
Mexico D.F.
University of Michigan
Ann Arbor, Michigan
PC Museum National d’Histoire Naturelle

Laboratoire de Cryptogamie

Autonoma de México,

MICH

Paris
UC University of California, Berkeley
US United States National Herbarium

Smithsonian Institution

Washington, D.C.

In our attempts to examine type specimens of
all the taxa, we were frustrated by an inability to
locate those of Yendo’s species (1902). It appears
that Yendo’s type specimens are not extant, a
problem compounded by the fact that he often
did not give a single type locality or ever designate
type specimens for his taxa. Therefore, we have
chosen herein the pertinent illustrations of
Yendo’s as lectotypes for his taxa (Stafleu et al.,
1978, Art. 9). Only selected specimens or those of

distributional importance are cited under “Spec-
imens Studied.”

ACKNOWLEDGMENTS.—We wish to thank Drs.
Robert W. Hoshaw and Donald A. Thomson and
the University of Arizona for supporting the se-
nior author during his tenure at Laboratorio de
Biologia Marina, Puerto Penasco, Sonora. Sup-
port for field work in Baja California was pro-
vided by a National Science Foundation grant
(NSF-BMS- 73-07000) to J. Norris and M. Neu-
shul. We thank Dr. Roger Meslin of the Labora-
toire de Botanique, Caen, and Mr. Robert Setzer
of the Allan Hancock Foundation, Los Angeles,
for the loan of herbarium material. The senior
author is especially grateful to Alexander “Ike”
Russell of Tucson, Arizona, for flying him to Isla
Angel de la Guarda, Desemboque de los Seris,
and Bahia de los Angeles, and assistance in ob-
taining critical material for this study. Assistance
in collecting by Katina Bucher and Eric Johan-
sen, and in the preparation of specimens by Paul
J. Matty is appreciated. Thanks go to Mr. Ed-
ward Davidson of the Smithsonian Office of Fel-
lowships and Grants for financial assistance for
travel of the junior author to the Smithsonian to
complete this study, and to Dr. Walter H. Adey
and Robert S. Steneck, National Museum of
Natural History, Smithsonian Institution, for dis-
cussions on marine biogeography. We are grateful
for the reviews of this paper by Dr. Dan H.
Nicolson and Katina E. Bucher of the National
Museum of Natural History, Smithsonian Insti-
tution, Dr. Robert W. Hoshaw of the University
of Arizona, Tucson, and Roberta A. Townsend
of the University of Sydney.

Amphiroa Lamouroux, 1812

Description.—Holdfasts: crusts, sometimes ob-
scure or missing in plants in turfs. Fronds: in
clumps several centimeters high, or in turfs with
sand and other algae mixed in. Branching: basi-
cally dichotomous, but often irregular and some-
times with branches arising from intergenicula in
a seemingly random manner; branches sometimes
more or less in one plane or they may form a

4 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

bushy, erect clump; the branches may be erect or
recumbent to varying degrees. Jntergenicula: flat to
terete, usually 3 or more times as long as broad.
Genicula: one or more tiers of uncalcified, deeply
staining medullary cells; demarcation between
intergenicula and genicula distinguished by the
presence or absence of calcite in cell walls, cell
tiers may be partly intergenicular and genicular;
genicular cortices usually present, but often dis-
organized and ruptured. Medulla: formed by syn-
chronous divisions of apical cells, hence resulting
in arching tiers within each of which the cells are
the same length, divisions such that tiers vary in
height from 10-20 wm to more than 120 pm;
secondary pit-connections forming between the
cells of a tier. Cortex: thick or thin, sharply de-
marcated from medulla, continuing to grow
slowly in thickness as intergenicula age.

Data on reproductive structures are summa-
rized from these studies: Suneson, 1937; Segawa,
1940a, 1940b; Ganesan, 1968; Johansen, 1968;
and Murata and Misaki, 1978. Conceptacles: orig-
inating near branch apices and protrude from or
are sunken into cortical tissue on surfaces of in-
tergenicula; usually several conceptacles per in-
tergeniculum; older conceptacles becoming bur-
ied by cortical growth in some species. 7etraspor-
originating as dome-shaped
swellings on intergenicular surfaces by excessive

angial conceptacles:

growth of small cortical cells (the future roof of
the conceptacle) over a lens of elongated cells
(cavity cells, Johansen, 1968), subsequent devel-
opment involving the initiation and growth of
tetrasporangia, or bisporangia in some, in the
periphery (and sometimes also the center) of the
lens, meanwhile the cavity cells degenerating to
form a chamber; also degenerating are a group of
small cortical cells in the center of the roof, this
space is the pore of the conceptacle; tetrasporan-
gia 40 to 110 um long. Sexual plants: dioecious,
conceptacles generally more crowded than in
tetrasporangial plants. Male conceptacles: low pro-
file, chambers broad and with low ceiling, fertile
surface restricted to flat floor of chamber. Female
conceptacles: roofs of conceptacles consisting of fil-

aments that have grown centripetally from tissue
surrounding the fertile layer; 1 to 3, 2-celled
carpogonial filaments arising from each support-
ing cell and constituting, along with undeveloped
carpogonial systems, the fertile layer; trichogynes
from the more central carpogonia projecting
through pore at maturity. Carposporangial concep-
tacles: fusion cell 6-10 wm thick and as much as
100 wm or more in diameter, carposporangial
filaments arising from margins of fusion cell and
sometimes also from its upper surface near the
margin, carposporangia 25 to 65 wm in diameter.
Type-Species.—Amphiroa tribulus (Ellis and So-
lander) Lamouroux, 1812:186. Basionym: Coral-
lina tribulus Ellis and Solander, 1786:124.
Remarks.— To distinguish among the morpho-
logically variable species of Amphiroa we place

| es
:

Ficure 1.—Branch transections and genicular longitudinal
sections of the Gulf of California species of Amphiroa (all to
same scale): a, A. beauvorsiz; b, A. brevianceps; c, A. magdalensis;
d, A. misakiensis; e, A. rigida; f, A. valonioides; g, A. van-bosseae.

NUMBER 9

Tasie 1.—Summary of the distinguishing features of the species of Amphiroa in the Gulf of
California (rnd = round, com = compressed, ere = erect, usually as small clumps in which the

branches are free from one another, rec = usually spreading or recumbent, tuf = tufts or turfs,

usually compact, sand-filled)

| Frond Intergenicula Genicula
Sues | f Diameter or
DEcles | Maximum width in No. of tiers
length upper frond Shape in of medullary
| (cm) Habit (mm) cross-section cells
A. beauvoisi 12 ere 0.5-1.0 rnd/com 35
A. brevianceps | 4 ere 1.5-3.0 com >5
A. magdalensis | 5 ere 1.0=2.5 com ~10
A. misakiensis 4 rec 2.0-4.0 com 5 or
more
A. rigida 2 ere 0.4—1.0 rd 2
A. valonioides 2 tuf <0.5 rnd/com 1
A. van-bosseae 10 ere 1.0—2.0 rnd 5 or
more

most reliance on the internal structure of genicula
and on the external form of intergenicula (Table
1, Figure 1). In the Gulf of California species
there are four types of genicula: (1) a single
medullary tier: A. valonioides; (2) two tiers of med-
ullary cells, with oblique transverse walls between
the tiers: A. rigida; (3) 3 to 5 tiers: A. beauvorsiz; (4)
more than 5 tiers: A. brevianceps, A. magdalensis, A.
misakiensis, A. van-bosseae. In groups 1 to 3 the
distinctions are clear-cut, but in the more massive
genicula of the species in group 4, it has not been
possible to find genicular distinctions, and reli-
ance must be placed on intergenicular characters.

When analyzing intergenicular form, the fol-

lowing characteristics must be considered: (1)
terete- or flat-shaped (circular or compressed in
transection), (2) the dimensions, and (3) in flat
branches, the intergenicular configuration adja-
cent to the genicula (Figure 1). These features are
often not as taxonomically decisive as are those
of genicula. Although mostly terete, intergenicula
in A. beauvoisu and A. valonioides are sometimes
slightly compressed. In the other species, either
the terete or the flat character seems to hold
(Table 1). Only intergenicula in the upper parts
of the fronds can be used; basal intergenicula are
usually terete, even in species where upper inter-
genicula are flat.

Key to the Species of Amphiroa

1. Intergenicula distinctly flat; rarely cylindrical 2

Intergenicula cylindrical or nearly so

2. Fronds more or less erect and regularly branched; intergenicula all approx-

imately similar in appearance ...

Fronds spreading, recumbent and irregularly branched; intergenicula vari-

able in appearance

Sere en Cen ane dl acwhee Oe A. misakiensis

3. Intergenicula more than | mm wide; genicula consisting of 6 or more tiers

of medullary cells

Intergenicula less than 1 mm wide; genicula consisting of 3 to 5 tiers of

medullary cells

A. beauvoisii

4. Intergenicula less than 4 mm long
Intergenicula more than 5 mm long
a:
tiers of medullary cells
more tiers of medullary cells
6
tiers of medullary cells
1 tier of genicular cells
ie

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

oli TRE AER aie Rice Loree A. brevianceps
5 OI Sea ES a A. magdalensis
Intergenicula less than | mm in diameter; genicula consisting of 4 or fewer

Intergenicula more than | mm in diameter; genicula consisting of 5 or
2658 aS een A. van-bosseae

. Intergenicula 0.5 to | mm in diameter; genicula consisting of 2 or more

Intergenicula less than 0.5 mm in diameter; genicula usually consisting of
hast Eg edt SRS A. valonioides
Fronds less than 2 cm tall, rigidly branched in any plane; genicula

consisting: off 2 itierstof medulllanyacclls eee Een rnnee A. rigida
Fronds mostly 2 to 12 cm tall, more or less branched in one plane; genicula

consisting of 3 to 5 tiers of medullary cells ............... A. beauvoisii

Amphiroa beauvoisii Lamouroux
Figures la, 2, 3, 4a—c, 5, 6b, 7, 8, 146, 156

Amphiroa beauvoistt Lamouroux, 1816:299 [no illustration].

Amphiroa linearis Kutzing, 1858:22, pl. 46: figs. 2a—c [type
locality: “Ad litora Africae occidentalis” (holotype, L
938,334... 357)).

Amphiroa zonata Yendo, 1902:10, pl. 1: figs. 11-14, pl. 4: fig.
9 [type locality: not specified, “Misaki, Shimoda and
Sunosaki” are listed (lectotype, Yendo’s illustrations are
herein chosen)].—Dawson, 1944:276, 1953:146; 1959:22;
1966a: 18.

Amphiroa pusilla sensu Dawson, 1944:276 [in part] [not Am-
phiroa pusilla Yendo, 1902:13].

Amphiroa peninsularis Taylor, 1945:188, pl. 48: fig. 1 [type
locality: South Bay, Isla Cedros, Baja California (WRT-
34-646A, holotype, AHFH 90)].

Amphiroa crosslandi sensu Dawson, 1953:149; 1954:136 [not
Amphiroa crosslandit Lemoine, 1929:50].

Amphiroa drouetaa Dawson, 1953:140, pl. 27: figs. 5-6 [type
locality: “Intertidal on reef at north end of Isla Turner,
off Isla Tiburon, Sonora” (EYD-717-40, holotype, AHFH
1555; isotype, UC 940256)].—Dawson, 1959:21; 1966a:
18.

Amphiroa franciscana var. robusta Dawson, 1953:150 [no illus-
tration] [type locality: Acapulco, Guerrero, Mexico (EY D-
3881, holotype, AHFH 55177) 1966b:59.

Description.—Fronds: up to 12 cm high, more
or less erect and open, but sometimes in compact,
pulvinate clumps (Figure 2). Branching: dichoto-
mous, more or less in one plane, sometimes irreg-

ularly so, dichotomies sometimes separated by
one or more unbranched intergenicula. Jntergeni-
cula: near base terete or subterete, up to 1.3 (—1.7)
mm diameter and 3(—4) mm long, intergenicula
in upper parts of fronds terete; subterete, or flat,
especially near branch apices, becoming more
terete with age because of cortical thickening,
mostly 0.5-1 mm broad and 3-5 or sometimes
more than 10 mm long, branching intergenicula
sometimes Y-shaped (Figure 4). Genzcula: devel-
oped by cracking and sloughing of calcified cor-
tices overlying uncalcified tissues; fully formed
genicula usually barely visible between intergen-
icula, consisting of 3-5 tiers of medullary cells
and irregularly disposed patches of cortical cells
(Figure 5). Conceptacles: scattered over intergeni-
cular surfaces, protruding slightly; tetrasporan-
gial and bisporangial conceptacles 200-250
(—300) wm inside diameter; sexual plants not
encountered in the Gulf of California.

Type Loca.ity.—‘‘Cotes du Portugal” (La-
mouroux, 1816:299).

Ho.otyre.—In Lamouroux’s herbarium (CN).

GuLF OF CALIFORNIA DistRiBUTION.—Puerto
Penasco to La Paz (Figure 6).

Paciric Coast DistrisuTion.—Santa Catalina
Island, off southern California (Johansen, 1976b:
400, as A. zonata) to Ecuador (Dawson, 1953:146,
as A. zonata); Galapagos Islands (Silva, 1966:152,
as A. zonata).

NUMBER 9

SPECIMENS STUDIED.—Gulf of California. SONORA:
Punta Pelicano, vicinity of Puerto Penasco, 2 Jul

6 mm

Ficure 2.—Amphiroa beauvoisi: a, fronds from a single clump
JIN & HWJ-73-7-3) (note variation from cylindrical to
slightly compressed); 6, genicula (JN-28 III 1973) (note they
are more evident here than in “a,” probably due to greater
flexing).

~

1973, JN & HWJ-73-7-3 (US, CUW); Playa Ar-

enosa, vicinity of Puerto Penasco, 15 Feb 1965,

100 pm
|

Ficure 3.—Longitudinal sections through apex of a branch
of Amphiroa beauvoisu (JN-4384).

A. E. Dennis D-81 and D82 (both US), 8 Apr

1966, EYD-27360 (US); Cumpleanos Tide Pool,
Playa Estacion, Puerto Penasco, 4 Jul 1973, JN
& HW J-73-7-32 (CUW); Playa Estacion, in front
of Laboratorio de Biologia Marina, Puerto Pen-
asco, 27 Jul 1965, EYD-27470 (US), 7 Apr 1966,
EY D-27299 (US), 8 Apr 1966, EYD-27360 (US),
and 20 Oct 1972, JN-3583 (US); Isla San Jorge,
20 Feb 1946, EY D-846 (US); Puerto Lobos (Cabo
Tepoca), 26 Mar 1937, Remple Sta. 724-37
(AHFH); Puerto Libertad, 19 Feb 1946, EYD-
720 (US); Ensenada Bocochibampo, near Guay-
mas, 16 May 1946, EYD-1755 (US), 12 Feb 1946,
EYD-490 (US); shallow lagoon, Guaymas Bay,
23 Jan 1940, EYD-5827 (US, UC, AHFH); Punta
Colorado, near Guaymas, 15 Feb 1946, EY D-556
(AHFH); Punta San Pedro, Guaymas, 22 Dec
1939, Drouet & Richards 3386 (UC, AHFH);
Ensenada de San Francisco, near Guaymas, 18
May 1946, EYD-1973 (US), 12 Jun 1958, EYD-
11039 (US); Punta Prieta, Bahia Topolobampo,
10 Jun 1952, EYD-10966 (US). Baja CALIFOoR-
NIA: Punta La Gringa, Bahia de Los Angeles, 22
May 1972, JN-3049 (US); Islas de los Gemelos,
Bahia de Los Angeles, 21 May 1972, JN-3008b
(US); Bahia Agua Verde, 12 Feb 1940, EY D-528-
40 (AHFRH), 11 Jul 1965, EYD-25852a (US); Isla

8 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

ae Sa) Hie yale A { ita

on aa 0

AGT Hipetlt wht

| Hi hi |

a isl an es veonateanale

ii

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ate haan it

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me ip! * et

a Pp eet be i :
+% ‘ i?
a, an via ve oUt Atlee

Ficure 4.—Longitudinal sections of intergenicula: a, Amphiroa beauvoisi: (JN & HWJ-73-7-31)
(note arching lines of primary pit connections (solid arrow) and secondary pit connections
(open arrow), which are about one-third below tops of cells); b, A. beauvoist: (JN-4384) (note
thin cortex); c, A. beauvoisa (JN & HWJ-73-7-14) (note different heights of medullary cell tiers);
d, A. van-bosseae (JN & HWJ-73-7-31) (note unusual arrangement of alternating long and short
tiers of medullary cells).

Ficure 5.—Longitudinal sections of successively older genicula of Amphiroa beauvoisiz: a, apex of
branch with a young geniculum (JN-4384) (note that specimen was decalcified in preparing
the slide and that the uncalcified genicular cells stain more intensely than the calcified
intergenicular cells); 6, close-up of geniculum in “a” (note that young genicular cortex is still
intact and unruptured); c, older 5-tiered geniculum with ruptured genicular cortex (JN-4384):
d, portion of branch showing geniculum and conceptacle (JN-4023).

OP aan.
aera D

oe

%

10 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Cabo Tepoca
oO OD
<J Bahia de

,» Los Angeles

]. Raza ©) aaa
b (F sla Tiburon
I. San eye ON sia ane 80°
I. Tortuga
\ (3) ODD2aD. Guaymas

Bahia

Sh
Concepcion

} Via Carmen 250
Puerto
Escondido #
—I. Monserrate
g
Bahia Agua ()
Verde a
I. San Diego
Topolobampo
vA La Paz~ 26°
Eureka
“4 \ a)
Punta Palmilla CS
Cabeza Ballena Zs abo Pulmo
Cabo San Lucas. \ Los Frailes
e
7
Mazatlan
) % ©
» © aS) ge

C1» Puerto Libertad i

Ficure 6.—Distribution of species of Amphiroa in the Gulf of
California (1 = A. beawoisi, 2 = A. brevianceps, 3 = A.
magdalensis, 4 = A. misakiensis, 5 = A. rigida, 6 = A. valonioides,
7 = A. van-bosseae).

Ildefonso, 19 Jul 1965, EYD-25877 and EYD-
25891 (both US); Isla Cholla, off Isla Carmen, 23
Apr 1958, EYD-18677 (US); Bahia Concepcion,
26 Mar 1949, EYD-7112 (AHFH); Isla Monser-
rate, 21 Apr 1958, EYD-18795 (US); Bahia La
Paz, 10 Nov 1946, EYD-3464 (AHFH). Las 1s-
LAS DE LA CINTURA: Puerto Refugio, Isla Angel de
la Guarda, 26 Jan 1940, EYD-222 (AHFH); Isla

Patos, off Isla Tiburon, 17 Feb 1946, EYD-808
(US); Bahia Agua Dulce, Isla Tiburon, 21 Feb
1946, EYD-951 (US); Isla Turner, off Isla Tibu-
ron, EYD-108-40 (AHFH), 18 Jul 1940, EYD-
717-40 (holotype AHFH). California. CHANNEL
ISLANDS: Catalina Harbor, Santa Catalina Island,
14 Feb 1949, P.C. Silva-4636 (AHFH); Little
Harbor, Santa Catalina Island, 8 Mar 1971, R.
Setzer-5102 (AHFH); West Cove, San Clemente
Island, 12 Oct 1971, R. Setzer-5803 (AHFH).
ORANGE COUNTY: Corona del Mar, 21 Jan 1951,
EYD-9598 (US, UC, AHFH); Laguna Beach, 8
Nov 1942, N. Cooper-151 (AHFH). san preco
county: Cardiff, 29 Jan 1949, EYD-6039 (UC,
AHFH); La Jolla, 15 Jan 1946, EYD-318
(AHFH), 16 Feb 1958, EYD-18414 (AHFH), and
11 Nov 1958, EYD-19746 (AHFH). Pacific Coast
of Mexico. BAJA CALIFORNIA: Isla Coronado del
Sur, 20 Jun 1947, EYD-4247 (UC, AHFH) and
11 Aug 1948, C. Hubbs-48-217 (AHFH); Isla
Guadalupe, 18 Dec 1949, EYD-8367 (AHFH);
Punta Santa Rosallita, 10 Oct 1946, EYD-2852
(UC, AHFH); Miller’s Landing, 11 Oct 1946,
EY D-2946 (UC, AHFH, K); Punta Malarrimo,
Bahia Sebastian Viscaino, 17 Apr 1951, EYD-
9944 (UC, AHFH); ‘Campito’, east of Punta San
Eugenio, 31 Oct 1951, EYD-10487 (AHFH);
Punta Norte, Isla Cedros, 30 Oct 1951, EYD-
10582 (UC; AHFH); South Bay, Isla Cedros, 10
Mar 1934, WRT-646A (UC) and 19 Apr 1951,
EYD-9864 (UC, AHFH); Punta San Eugenio, 29
Aug 1957, EYD-20841 (US); Bahia San Barto-
lome (= Bahia Tortuga), 7 Nov 1949, EYD-6584
(AHFH); Punta Thurloe, 3 Apr 1955, EYD-
13313 (US); Bahia Ascuncion, 28 Apr 1950,
EYD-9178 (US, UC, AHFH); Isla Ascuncion, 25
Aug 1957, EYD-20379, EYD-20432a, and EYD-
20439 (all US); Punta Abreojos, 30 Apr 1950,
EYD-9456 (US, UC, AHFH, K) and 9 Feb 1964,
EYD-26302 (US); Bahia San Hipolito, 18 Aug
1957, EYD-20336, EY D-20338, and EYD-20361
(all US); Punta Entrada, Bahia Magdalena, 2
May 1950, EYD-9296 (UC, AHFH). REvILLA
GIGEDO ARCHIPELAGO: Binder’s Cove, Isla Socorro,
16 Apr 1955, EYD-13602 (US). IsLAs TRES MAR-
1as: Isla Maria Magdalena, 9 May 1939, WRT-

NUMBER 9

reg
Waldo L. Schmitt, No. A7O— (7H 1933, oe
«

Seven tanenneereer reenter te

|

5mm 8)

Ficure 7.—Type specimens: a, Amphiroa mexicana
(AHFH 91); b, A. beauvoisiz (CN).

642A (paratypes US, UC). jatisco: Bahia Cha-
mela, 14 Apr 1959, EYD-21258 (US). coxima:
Bahia Santiago, 12 Apr 1959, EYD-21313 (US).
GUERRERO: Morro de Petatlan, 17 Mar 1933, W.
L. Schmitt-120C-33 (UC); Bahia Petatlan, 2-3
Mar 1934, WRT-568 (paratypes US, UC,
Ewtiiil)- Puerto Guatulco, 8 Apr 1959, EYD-
21440a (US); Salina Cruz, 10 Jan 1947, EYD-
3826 (UC), 12 May 1952, EYD-10760 (US); Isla
Grande, 10 Apr 1959, EYD-20954 (US); Aca-

13.
C

Ficure 8.—Lectotype selected herein of Amphiroa zonata
Yendo [= A. beauvorszz| (see also Figure 156): a, habit of frond;
b, longitudinal section through part of intergeniculum; c,
oblique section through intergeniculum showing a young
conceptacle; d, cross-section of branch. (All from Yendo,
1902, pl. 1.)

pulco, 3 Feb 1947, EYD-3881 (holotype AHFH).
El Salvador. El Tunco, west of Puerto Libertad, 10
Sep 1960, EYD-21939 (US); El Cuco, 8 Sep 1960,
EYD-21817 (US); Acajutla, 4 Sep 1960, EYD-
21882 (US). Pacific Coast of Costa Rica. Bahia
Ballena, 30 Mar 1959, EYD-21191 (US); Isla del
Cano, 28 Mar 1959, EYD-21059 and EYD-21060
(both US). Pacific Coast of Panama. Isla Jicaron, 25
Mar 1959, EYD-21139 (US); Isla del Rey, Bahia
San Telmo, 23 Mar 1959, EYD-21163 (US).
REMARKS.—Many specimens of Amphiroa beau-
voisit have been collected in the Gulf of California
and reported under other names. Dawson (1953:
134-135, 140-141) considered A. zonata and A.
droueti as distinct from each other on the basis of
differences in the relative visibility of genicula in
whole branches, in intergenicular breadth, in de-
gree of flatness or roundness of intergenicula in

12 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

cross-sectional view, in relative lengths of inter-
genicula, and in the relative amounts of “forking”
of intergenicula. This complex represents one
widely variable species. On morphological bases,
it was impossible to categorize them into the taxa
that Dawson recognized in the Gulf.

We were unsuccessful in our efforts to locate
type material of A. zonata Yendo (1902). In ac-
cordance with Article 9.3 of the /nternational Code
of Botanical Nomenclature (Stafleu et al., 1978), we
selected the illustrations of Yendo (i.e., pl. 1: figs.
11-14; pl. 2: fig. 9) as the lectotype of A. zonata
(Figure 8). Furthermore, the examination of this
and other type specimens of Amphiroa shows that
the earliest validly published name applicable for
this morphologically variable complex is Amphiroa
beauvoista Lamouroux, 1816 (Figure 7d).

In all likelihood, A. beauvoisiz is the most wide-
spread species of Amphiroa, perhaps occurring in
most tropical and subtropical areas where the
genus may be found. Although we are as yet not
prepared to consider entities other than those of
the Gulf of California, it is probable that several
other species described subsequent to Lamouroux
(1816) may also belong to A. beauvoisi. For ex-
ample, in the Pacific Ocean, Amphiroa galapagensis
Taylor (1945), A. mexicana Taylor (1945) (Figure
6a), and A. echigoensis Yendo (1902) may be syn-
onymous with A. beawvoisit.

Amphiroa crosslandi Lemoine (1929) was men-
tioned as occurring in the Gulf at La Paz, Baja
California del Sur, by Dawson (1953:149). This
is the only Gulf of California collection labeled
A. crosslandii that we are aware of. Unfortunately,
Dawson did not cite specific collections, but it is
probably based on EY D-3464 (AHFH 44178, UC
974659), which we conclude represents a collec-
tion of several small aberrant plants of A. beau-
voisit. While we have examined an isotype of A.
crosslandii (BM) and found it to superficially re-
semble A. beauvoisiz, this taxon must await further
study before any conclusion can be made on its
taxonomic status.

Amphiroa linearis was reported by Hariot (1895)
from the Gulf of California. While we have not

seen the specimens on which Hariot based his
record, Dawson (1953:137) noted that it was
probably Amphiroa zonata. Further, the holotype
of A. linearrs Kutzing (from Gabon, West Africa),
represented by fragments in L 938, 334. . .357 is
in agreement with our concept of A. beauvoisiz. We
now consider A. linearis as a synonym of A. beau-
vowsie.

Our study of the type specimen of Amphiroa
franciscana var. robusta Dawson (AHFH) from Aca-
pulco reveals that it too is within the morpholog-
ical range we recognize for Amphiroa beauvoisit.
When Dawson (1953:150) erected the variety, he
also listed a single Gulf of California specimen
(paratype, EYD-4455), which we did not study;
it may also represent A. beauvozsze.

Unlike other species in the Gulf of California,
A. beauvoisit is widespread in distribution in trop-
ical and subtropical areas of the Atlantic and
Indian Oceans. In the western Atlantic, it occurs
from North Carolina (Schneider, 1976:138) south
to Uruguay (Taylor, 1960:405-—406, in part as A.
brasiliana). It also is reported from Portugal (type
locality), the Mediterranean Sea (Hamel and Le-
moine, 1953:42), and is abundant in the south-
western Indian Ocean (Johansen, unpublished
data).

Amphiroa brevianceps Dawson
Ficures 16, 6, 9
Amphiroa brevianceps Dawson, 1953:142, pl. 31: fig. 2.

DescripTion.— Fronds: in small clumps up to 4
cm high. Branching: somewhat regularly and
densely dichotomous, in one plane. Jntergenicula:
near base subterete or flat and up to 2 mm broad,
intergenicula in upper parts flat, up to 3 mm
broad and short, usually less than 4 mm long, not
differentiated into midrib and wings. Genicula:
externally prominent, dark colored, bracketed by
calcified extensions of the adjacent intergenicula,
6 or more tiers of cells per geniculum. Conceptacles:
Dawson (1953:143) reported tetrasporangial and
carposporangial conceptacles on both intergeni-
cular surfaces.

NUMBER 9

Ficure 9.—Type specimen of Amphiroa brevianceps
(EY D-3825, AHFH).

Type Locatity.—“On rocky shore just east of
Salina Cruz, Oaxaca, Mexico” (Dawson, 1953:
143).

Ho.otype.—EYD-3825, 10 Jan 1947 (AHFH
55180; isotype UC 925637).

GuLr OF CALIFORNIA DistRIBUTION.—Guay-
mas; Los Frailes.

Paciric Coast DistrrpuTion.—Punta San Hi-
polito, Baja California (Dawson, Neushul, and
Wildman, 1960b:16):; Salina Cruz, Oaxaca.

SPECIMENS STUDIED.—Gulf of California. SONORA:
Punta de Las Cuevas, Ensenada de San Fran-
cisco, near Guaymas, 14 Feb 1946, EYD-593
(paratype AHFH). saya CALIFORNIA: Punta
[Los] Frailes, 13 Mar 1949, EYD-6895 (paratype

13

AHFH). Pacific Coast of Mexico. Oaxaca: Salina
Cruz, Oaxaca, 10 Jan 1947, EYD-3825 (holotype
AHFRH) and 12 May 1952, EYD-10765 (US).

Remarks.—In the Gulf of California, there are
three species of Amphiroa in which the intergeni-
cula are flat, A. brevianceps, A. magdalensis, and A.
misakiensis. Of these, A. brevianceps and A. magda-
lensis are represented by few specimens, with A.
brevianceps the most poorly known. Amphiroa brev-
anceps is Closely related to Amphiroa anceps (La-
marck) Decaisne, a widespread species in subtrop-
ical and tropical areas of the world (Weber-van
Bosse, 1904:93, pl. 16: figs. 6-8).

Weber-van Bosse (1904) and Johansen (1969b)
described the genicula of A. anceps; those in A.
brevianceps are identical. The flexing of the fronds
and the thickness of the intergenicular cortex
results in a patch of calcified cortical tissue break-
ing away so that in surface view a genciulum is
bracketed by calcified tissue. Dawson (1953:143)
described a geniculum in A. brevianceps as a un-
calcified “window” appearing “embraced” [by
intergenicular tissue]. Amphiroa anceps is noted for
this feature, but here the intergenicula are longer
than in A. brevianceps. This genicular character is
most evident in younger parts of the fronds before
erosion and secondary growth take place. More
specimens are needed so the specific relationships
of these taxa can be resolved.

Amphiroa magdalensis Dawson
Ficures lc, 6, 106
Amphiroa magdalensis Dawson, 1953:143, pl. 30: fig. 2.

Description.—Fronds: up to 5 cm high in loose
tufts. Branching: sparsely dichotomous or irregu-
lar. Intergenicula: basal intergenicula terete, others
markedly flattened and 1-2.5 mm broad, 3 to 6
times as long. Genicula: conspicuous because of
broken out pieces of adjacent intergenicula in
center of branch; up to or more than 10 cellular
tiers per geniculum. Conceptacles: tetrasporangial
conceptacles scattered on intergenicular surfaces,
usually more on one side than the other; sexual
plants not encountered in the Gulf of California.

14 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Type Locarity.—“Rocky shore at Punta En-
trada, Isla Magdalena, Baja California, Mexico”
(Dawson 1953:144).

Ho. otype.—EYD-6688, 8 Mar 1959 (AHFH
55165).

GuLF OF CairorniA Distripution.—Isla Ide-
fonso (Dawson, 1959:21) to Punta Palmilla; Ma-
zatlan.

Paciric Coast Distrrpution.—Punta Abreojos
to Bahia Magdalena, Baja California.

SPECIMENS STUDIED.—Gulf of California. BAJA
CALIFORNIA: Isla Tortuga, 25 Apr 1958, EYD-
18646 (US); Cabeza Ballena, 11 Mar 1949, EYD-
6831 (AHFH); Punta Palmilla, 7 Nov 1946,
EYD-3224 (AHFH). sinatoa: Mazatlan, 7
Jun 1952, EYD-10828 (US). Pacific Coast of Mex-
ico: BAJA CALIFORNIA: Punta Abreojos, 9 Feb
1964, EYD-26301 (US); Punta Entrada, Bahia
Magdalena, 8 Mar 1949, EYD-6688 (holotype
AHFH). Punta Redondo, Isla Santa Margarita,
3 Feb 1964, EYD-26294 (US); Cabo Tosco, Isla
Santa Margarita, 4 Feb 1964, EYD-26316 (US).

RemMarks.—Among the more robust species of
Amphiroa, A. magdalensis may be distinguished by
having flat, thin, long intergenicula (Figure 108).
The genicula have more cellular tiers than other
Gulf of California species, with 10 tiers present in
some specimens (Figure lc).

Few specimens of A. magdalensis have been col-
lected and the relationship of this species to other
species is poorly known. Dawson labeled a Gulf
specimen from near Guaymas, Sonora (EYD 650,
US) as Amphiroa foliacea Lamouroux, probably
because of the wings flanking the slightly raised
midrib in some of the intergenicula, a feature
ascribed to A. foliacea (Ganesan, 1968:16). As
mentioned earlier, A. magdalensis and A. brevianceps
are similar; in the latter the intergenicula are
shorter. Amphiroa anceps, an older species common
in other subtropical areas (e.g., southeast Africa,
Johansen, 1969b:120), is also related structurally
to these two Gulf species. For the time being,
these taxa should be considered distinct until
population studies, particularly on those from
type localities, and more collections have been
made. It may be that they represent a single

a
TNF
mae TN eae
N Ph. TEN
re bet NG

Ficure 10.—Type specimens: a, Amphiroa subcylindrica (= A
van-bosseae, EY D-555, AHFH); b, A. magdalensis (EY D-6688,
AHFH).

polymorphic species. Dawson (1953:145) stated,
with reference to specimens of A. magdalensis that
“these are representative of the wide range of
variation in size which may be encountered in
species of Amphiroa and of which one must be well
aware in attempting determinations of these per-
plexing plants.”

In addition to the Gulf, A. magdalensis is known
from the Pacific coast of Baja California and Isla
Guadalupe (Dawson, 1953:144). Hommersand
(1972) suggested that this species and Amphiroa
ephedraea (Lamarck) Decaisne (as interpreted from
Japanese specimens) are closely related or possi-
bly conspecific. In this connection, it should be
noted that A. ephedraea as recognized in Japan
(Okamura, 1936:518) is not the same species ex-
emplified by the type of the species (PC).

NUMBER 9

Amphiroa misakiensis Yendo
Ficures ld, 6, 11, 12a, 13a, 15¢

Amphiroa misakiensis Yendo, 1902:14, pl. 1: figs. 24, 25; pl. 6:
fig. 1.

Amphiroa pusilla sensu Dawson, 1944:276 [in part] [not Am-
phiroa pusilla Yendo, 1902:13].

Amphiroa dimorpha sensu Taylor, 1945:192, pl. 55.—Dawson,
1953: 141; 1966a:18 [not Amphiroa dimorpha Lemoine,
1929:76].

Amphiroa dimorpha var. digitiformis Dawson, 1959:21 [as ‘digi-
tiforme’|, fig. 4 [type locality: “...at a depth of about 5
feet, Isla Cholla, off Isla Carmen, April 23, 1958,” EYD-
18684 (holotype LAM now AHFH 81912;
AHFH)].

isotype

DescripTion.—Fronds: up to 4 cm long; usually
growing more or less recumbent and spreading
from the substrate so that one branch surface is
uppermost. Branching: basically dichotomous, but
this pattern often obscure when one to several
branches arise irregularly from intergenicula. Jn-
tergenicula: near base small and subterete, in upper
parts of fronds flat, irregularly shaped, size vary-
ing greatly but usually up to 7 mm long and 3-4
mm broad. Genicula: developing by cracking and
sloughing of calcified cortical tissue overlying
uncalcified genicular tissue, genicula quite evi-
dent externally, comprised of 5 or more tiers of
medullary cells plus patches of uncalcified corti-
cal tissue. Conceptacles: several per fertile intergen-
iculum scattered over uppermost (or lowermost
according to Dawson, 1953:142) surfaces, pro-
truding slightly, tetrasporangial conceptacles
about 200 um inside diameter; sexual plants not
encountered in the Gulf of California.

Type Loca.ity.—‘‘Misaki” Japan (Yendo,
1902:14).

Lecrotypre.—In the absence of a known holo-
type, Yendo’s illustrations (1902, pl. 1: figs. 24,
25, & pl. 6: fig. 1) are selected as the lectotype of
A. misakiensis Yendo (Figure 15c).

GuLr oF Ca.irorniA Distripution.—Puerto
Penasco to Cabo San Lucas; Mazatlan (Figure
6).

Paciric Coast Distrigution.—Baja California
to Costa Rica; Peru.

SPECIMENS STUDIED.—Gulf of California. SONORA:

Punta Pelicano, vicinity of Puerto Penasco,
6 Apr 1966, EYD-27269 (US) and 2 Jul 1973,
JN & HWJ-73-7-2 (US, CUW); Playa Aren-
osa, vicinity of Puerto Penasco, 15 Feb 1965,
ee Dennis -/2 (US) 130) june 19655 EYaD-
27430 (US), 5 Sep 1972, JN-3439 (CUW), and
27 Oct 1972, JN-3606 (US); Cumpleanos Tide
Pool, Playa Estacion, Puerto Penasco, 8 Sep 1972,
JN-3531 (US), and 4 Jul 1973, JN & HWJ-73-7-
27 (ARIZ); Playa Estacion, in front of Labora-
torio de Biologia Marina, Puerto Penasco, 27 Jul
1965, EYD-27471 (US), 7 Apr 1966, EYD-27339
(US), 11 Jul 1972, JN-3146b (ARIZ), and JN-
3173 (GUW), 22 Jan 1973, JN-3684 (MEXU),
and 25 Nov 1972, JN-3806 (US); Guaymas, 22
Dec 1939, Drouet & Richards-3387 (AHFH);
Ensenada de San Francisco, near Guaymas, 17
Apr 1946, EYD-1865 (AHFH) and 20 Nov 1946,
EY D-35552 (AHFH). Baja Ca.irornia: Bahia
Agua Verde, 20 Apr 1958, EYD-18902 (US); Isla
IIdefonso, 19 Jul 1965, EYD-25906 (US); Isla
Cholla, off Isla Carmen, 23 Apr 1958, EYD-18684
(holotype, LAM); Punta[Los] Frailes, 13 Mar
1949, EYD-7088 (AHFH); Cabeza Ballena, 9
Nov 1946, EYD-3391 (AHFH); Cabo San Lucas,
20-23 Aug 1957, EYD-20173 (US). Las Isras
DE LA CinTuRA: Isla Patos, off Isla Tiburon, EY D-
807 (US). sitnaLtoa: Mazatlan, 7 Dec 1946,
EYD-3665 (AHFH). Pacific Coast of Mexico.
BAJA CALIFORNIA: Chester Islets, Bahia Sebastian
Viscaino, 16 Aug 1957, EYD-20493 (US); Bahia
San Hipolito, 18 Aug 1957, EYD-20352 (US);
Punta Abreojos, 9 Feb 1964, EYD-26303 (US);
Punta Entrada, Bahia Magdalena, 19 Aug 1957,
EY D-20213 (US). REVILLA GIGEDO ARCHIPELAGO:
Binder’s Cove, Isla Socorro, 19 Nov 1953, EYD-
12139 (US); Isla San Benedicto, 17 Nov 1953,
EYD-12046 (AHFH). tstas tres martias: Isla
Mania Gleotas Om heb 9545 YD=I2 3115
(US). NAyarRitT: Mira Mar, 20 Dec 1946, EYD-
3703 (AHFH). coutma: Bahia Carrizal, 4 Dec
1959, EYD-21008 (US). oaxaca: Puerto Gua-
tulco, 4 Jul 1959, EYD-21398 (US). Ei Salva-
dor. Punta Acajutla, 4 Sep 1960, EYD-21777
(US). Pacific Coast of Costa Rica. Isla del Cano,
28 Mar 1959, EYD-21063 and EYD-21102 (both

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Ficure 11.—Amphiroa misakiensis: a, branches and conceptacles (JN-3684) (note irregularity in
branching); 6, JN & HWJ-73-7-2; c, flat crust-like piece from which arises several branches (JN
& HWJ-73-7-2a); d, specimen resembling Dawson’s A. dimorpha var. digitiformis (JN-5102) (note
numerous small branches arising from some of the large flat intergenicula).

NUMBER 9

Ficure 12.—Longitudinal sections of genicula: a, Amphiroa misakiensis (JN & HWJ-73-7-27)
(note two branches that developed from dichotomously divided intergeniculum before the
genicula formed; b, A. van-bosseae (JN-4859) (note multi-tiered geniculum in which adjacent
intergenicula are well separated by genicular tissue); c, magnified view of the geniculum shown
in “6” (note remnants of genicular cortex); d, A. valonioides (JN & HW J-73-7-4) (note geniculum
of single-tiered cells).

18

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

oad

ugh ete es
100 pm » < b
L

Ficure 13.—Sections of conceptacles: a, two tetrasporangial conceptacles of Amphiroa misakiensis
(JN & HWJ-73-7-2a) (note cortex is thicker on the dorsal surface); 6, A. valonioides (JN-4044)
(note protruding conceptacle, cf. “a”); c, old tetrasporangial conceptacle of A. van-bosseae (JN-
4859) (note copious cortical tissue over the conceptacle chamber).

NUMBER 9

US); Bahia Uvita, 30 Mar 1959, EYD-21092
(US).

Remarks.—As with other taxa described by
Yendo (1902), a type specimen seems not to be
extant. However, Segawa (1940b) published an
anatomical study of Amphiroa misakiensis based on
specimens from Japan and, on the basis of the
data he provided and Yendo’s original descrip-
tion, illustrations and habit photograph, we refer
the Gulf of California material, identified as Am-
phiroa dimorpha by Dawson (1953, 1966a), to A.
misakiensts.

Of all the species of Amphiroa in the Gulf, this
seems to be the most polymorphic. Generally
many intergenicula are broad and flat, but oc-
casionally small cylindrical intergenicula arise
from the large flat ones, as desribed and figured
for Amphiroa dimorpha var. digitiformis Dawson
(1959), which we now consider to be a synonym
of A. misakiensis. This species grows in tufts in
which the branches are often recumbent or only
partially erect (Table 1). Flat intergenicula may
often have the upper surface slightly convex and
the lower concave. Sometimes the edges of broad
intergenicula are slightly bent toward the dorsal
sumface(Vendo, 1902, pl. 1: figs. 24,25). The
uppermost cortices are thicker than the lower
cortices (Figure 13a), a feature also seen in Bos-
stella californica ssp. schmitt: (Manza) Johansen
(1973), another species of articulated coralline in
which the intergenicula are horizontally disposed.

Widely distributed in the eastern Pacific, A.
misakiensis (as A. dimorpha) has been reported from
Isla Cedros, off the Pacific coast of Baja California
(Dawson, Neushul, and Wildman, 1960b:16) Pa-
cific Costa Rica (Dawson, 1957:19) and from
Peru (Dawson, Acleto, and Foldvik, 1964:47).
Dawson, Neushul, and Wildman (1960a:44, as A.
dimorpha) stated that in central Baja California
this species is “an exclusively deepwater plant of
frequent occurrence at depths of 25-65 feet [7.6-
19.8m] in the southernmost kelp areas.” Else-
where, it may occur in “‘severely surfy habitats,”
such as, El Salvador (Dawson, 1961b:411, as A.
dimorpha).

Amphiroa rigida Lamouroux
Ficures le, 6, 14a,c

Amphiroa ngida Lamouroux, 1816:297, pl. 11: fig. 1.

Amphiroa taylor: Dawson, 1953:138, pl. 26: fig. 1 [type local-
ity: Bahia Braithwaite, Isla Socorro, Revilla Gigedo Ar-
chipelago, Mexico (WRT-34-27, AHFH
55156)]; 1959:22; 1966a:18.

holotype,

Description.—Fronds: up to 2 cm high, isolated
or sometimes in more or less erect clumps. Branch-
ing: irregular and usually at wide angles, branch
junctions usually not coinciding with genicula.
Intergenicula: terete, to 0.5 (—1.0) mm diameter
and variable in length. Genicula: often nearly in-
visible in young parts of fronds, becoming ex-
posed by separation of calcified cortices overlying
uncalcified genicula, made up of two tiers of cells
each 75-100 um high, end walls of cells oblique
where they meet and cells appearing imbricated.
Conceptacles: not studied in Gulf of California
specimens (see Suneson, 1937, for species).

5mm ©) 10 pm (4

FicureE 14.—a, Fronds of Amphiroa ngida from a single collec-
tion (EYD-27400) (note that genicula are seldom at the
branch dichotomy); 6, genicula of A. beawvorsi (JN & HWJ-
73-7-3) (note squared-off way that cells of adjacent tiers are
joined); c, genicula of holotype of A. taylor (= A. mgida,
WRT-34-27) (note unique imbrication of cell ends, a feature
not found in other Gulf of California species of Amphiroa).

20 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

5)

Type Locarity.—‘‘Mediterranee” (Lamou-
roux, 1816:297).

Ho.otype.—Fragments in Lamouroux’s her-
barium in CN.

GuLr OF CALIFORNIA DisTRIBUTION.—Puerto
Penasco; Puerto Escondido (Dawson, 1959:27);
Cabeza Ballena.

Paciric Coast DistrrutTion.—Isla Socorro,
Revilla Gigedo Archipelago; Isla Maria Magda-
lena; Costa Rica; Nicaragua (Dawson, 1962).

SPECIMENS STUDIED.—Gulf of California. SONORA:
Playa Estacion, vicinity of Puerto Penasco, 9 Apr
1966, EYD-27400 (US). BAJA CALIFORNIA: Ca-
beza Ballena, 11 Mar 1949, EY D-6833 (paratype,
A. tayloru, AHFH). Pacific Coast of Mexico.
REVILLA GIGEDO ARCHIPELAGO: Caleta Binmer, Isla
Socorro, 19 Nov 1953, EYD-12141 (US); Bahia
Braithwaite, Isla Socorro, 2 Jan 1934, WRT-34-
27 (holotype, A. taylor, AHFH). Pacific Coast of
Costa Rica. Isla del Cano, 29 Mar 1959, EYD-
21096a (US).

Remarks.—The distribution of Amphiroa rigida
is widespread, occurring in such diverse areas as
the Mediterranean Sea (type locality), in Japan
(Segawa, 1940a), and now the Gulf of California,
Pacific Mexico, and Pacific Costa Rica. A more
robust variety, A. rigida var. antillana Bgrgesen
(1917:182), was described from the Caribbean
Sea. The type specimen of Amphiroa taylorii
(AHFH) from Isla Socorro, Revilla Gigedo Ar-
chipelago, as well as the Gulf of California spec-
imens previously identified as A. taylori, all belong
to A. ngida. The only notable difference being
that the Mexican plants are smaller than the type
of A. regeda Lamouroux (CN).

Amphiroa rigida is a unique species of articulated
coralline algae. Particularly charactersitic are the
two-tiered genicula, in which the end walls of the
cells are slanted and appear to overlap (Figure
14c; Suneson, 1937:48, fig. 28A). This type of end
wall has not been observed in other species of
Amphiroa, but is clearly evident in the type speci-
men of A. rigida and in the type of A. taylori. In
all other species of Amphiroa, the end walls are
transverse rather than oblique (see e.g., Figure
146). In fact, Yendo (1904:17) in his study of

coralline genicula incorrectly considered the gen-
icula of A. rigida to consist of single tiers in which
each cell was “twisted” at its midpoint so as to
form what appeared to be an oblique crosswall.
The developmental sequence of the genicula in
A. rigida has not been studied.

Plants of A. rigida in the Gulf are relatively
small, inconspicuous and easily confused with A.
valonioides. However, a character that is usually
sufficient to set A. regida apart is the irregularly
disposed, rigid branches (Figure 14a). Rarely do
the junctions of branches coincide with genicula
in A. rigida, as they often do in other species of
Amphiroa. This species is apparently not common
in the Gulf of California. In Puerto Penasco and
possibly elsewhere, it is found growing sympatri-
cally with A. valonioides.

Cabioch (1969) found that A. rgeda from the
Mediterranean Sea has a unique mode of early
growth and development that results in the bases
of the fronds becoming embedded in the crustose
coralline Neogoniolithon notarist: (Dufour) Hamel
and Lemoine. In this relationship the crustose
plants serve as holdfasts for the articulated plants.
This phenomenon has not been reported else-
where, but Dawson (1953:138) did state that A.
taylor’ (now considered a synonym of A. rgzda)
occurs on crustose coralline algae.

It is impossible at present to give an idea of the
distribution of A. rgeda in the eastern Pacific.
Some specimens that may represent this taxon
are those reported as A. rigida var. antillana from
Atlantic Costa Rica (Dawson, 1962:383), and A.
taylor from Pacific Costa Rica and Nicaragua
(Dawson, 1957:19, 1962:395).

Amphiroa valonioides Yendo
Ficures If, 6, 12d, 136,15a, 16, 17

Amphiroa valonioides Yendo, 1902:5, pl. 1: figs. 1-3, pl. 4:
fig. 1.

Amphiroa annulata Lemoine, 1929:78, fig. 34; pl. 4: fig. 1 [type
locality: “Galapagos. Ile James, James Bay, Station 2,
drage a 15 brasses (27 metres)” (holotype, BM) ].—Daw-
son, 1953:136, pl. 29: fig. 3; 1966a:18.

Amphiroa franciscana Taylor, 1945:187, pl. 48: fig. 2, pl. 49
[type locality: “Esmeraldas, dredged off Bahia San Fran-

NUMBER 9

cisco,” Ecuador, WRT-34-484 (holotype AHFH 93, iso-
type MICH)].

Amphiroa annulata var. pinnata Dawson, 1953:137 [no illustra-
tion] [type locality: Cabeza Ballena, Baja California, Mex-
ico, EYD-3374 (holotype, AHFH 55116)].

DescripTion.—Fronds: up to 2 cm high, more
or less erect, often in sand-filled turfs. Branching:
dichotomous, lateral adventitious
branches arising from sides of intergenicula,
branches several un-
branched intergenicula, especially those in turfs.
Intergenicula: terete or rarely flat, up to 0.5 mm
diameter (rarely more) and 1-3 mm long. Genvr-
cula: developing by cracking and sloughing of
calcified cortices overlying uncalcified genicular
tissues, fully formed genicula barely visible in
terete branches,

often with

sometimes containing

visible in flat

more easily

Ficure 15.—Habit of lectotypes selected herein: a, Amphiroa
valomordes (from Yendo, 1902, pl. 4: fig. 1; see also Figure
17a—c); b, A. zonata (= A. beauvoisiz) (from. Yendo, 1902, pl. 4:
fig. 9; see also Figure 8a,b); c, A. misakiensis (from Yendo,
1902, pl. 6: fig. 1).

21

branches, consisting of one tier of cells (possible
2 tiers in some instances). Conceptacles: often in a
single row on intergenicula, protruding markedly,
tetrasporangial and _ bisporangial conceptacles
about 200 um inside diameter; carposporangial
plants of this species (as Amphiroa franciscana) re-
ported but not studied by Dawson (1953:149).

Type Locarity.—Not specifically given; Prov-
ince of Hiuga and Misaki, Japan, listed in Yendo
(i902:5):

Lectotype.—In the absence of a known holo-
type specimen, we select Yendo’s illustrations
(1902, pl. 1: figs. 1, 3; pl. 4: fig. 1) as the lectotype
(Figure 17).

GuLr oF CALirorNiIA DistRiBuTION.— Puerto
Penasco to Cabeza Ballena; Mazatlan.

Paciric Coast Distripution.—Baja California
to Panama; Ecuador; Galapagos Islands.

SPECIMENS STUDIED.—Gulf of California. SONORA:
Punta Pelicano, vicinity of Puerto Penasco,
6 Apr 1966, EYD-27272 (US) and 2 Jul
1973, JN & HWJ-73-7-4 (US); Playa Arenosa,
vicinity of Puerto Penasco, 8 Apr 1966, EYD-
27362 (US), 5 Sep 1973, JN-3441 (US) and JN-
3442 (CUW); Cumpleanos Tide Pool, Playa Es-
tacion, Puerto Penasco, 4 Feb 1973, JN-3772
(ARIZ), 4 Jul 1973, JN & HWJ-73-7-33 (US);
Playa Estacion, in front of Laboratorio de Biolo-
gia Marina, Puerto Penasco, 29 Jun 1965, EYD-
27488 (US), 11 Jul 1972, JN-3146 (MEXU), and
25 Nov 1972, JN-3675 (US); Ensenada de San
Franciso, near Guaymas, 17 May 1946, EYD-
1866 (US); Ensenada Bocochibampo, near Guay-
mas, 16 May 1946, EYD-1780 (US). BAJA CAL-
IFORNIA: Bahia Agua Verde, 11 Jul 1965, EYD-
25852 (US); Isla San Diego, 19 Apr 1958, EYD-
18919 (US); Cabeza Ballena, 11 Mar 1949, EY D-
6834 (AHFH); Punta Palmilla, 7 Nov 1946,
EYD-3258 (US); Cabeza Ballena, 9 Nov 1946,
EYD-3374 (holotype, AHFH). sinaLtoa: Maza-
tlan, 7 Jun 1952, EYD-10824 (AHFH). Las 1s-
LAS DE LA CINTURA: Isla Patos, north end of Isla
Tiburon, 17 Feb 1946, EYD-812 (US); Isla Raza,
21 Nov 1964, EYD-26129 (US). Pacific Coast of
Mexico. BAJA CALIFORNIA: Isla Piedra, Laguna
Ojo de Liebre (Scammon’s Lagoon), 30 Apr 1946,

22 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

(JN-3146) (note
characteristic tuft of fronds); 6, separated branches from a
tuft (JN & HWJ-73-7-33) (note protruding conceptacles).

Figure 16.—Amphiroa valonioides: a, habit

EYD-2499 (AHFH); “Campito,” east of Punta
San Eugenio, 31 Oct 1951, EYD-10438 (AHFH);
Punta Malarrimo, 17 Apr 1951, EYD-9951
(AHFH); Laguna San Ignacio, 11 Feb 1950,
EYD-9005 (AHFH). Nayarit: Mira Mar, 20
Dec 1946, EYD-3702 (AHFH). Guerrero: Aca-
pulco, 3 Feb 1947, EYD-3908 (AHFH)._ REvILLA
GIGEDO ARCHIPELAGO: Binder’s Cove, Isla Socorro,
19 Nov 1953, EYD-12144 (US) and 16 Apr 1955,
EYD-13566 (US); Braithwaite Bay, Isla Socorro,
2=4 Jan 11934, WR®-27 (UG, US): LAs IsrAs
TRES MARIAS: Isla Maria Magdalena, 9 May 1939,
WRT-39-643A (UC). El Salvador. Acajutla, 4
Sep 1960, EYD-21919 (US); Isla Meanguera,

HTT YT

Tn

c| Kt A? ih spe: 4
iy sl Yad 4 4% oe a; GQ t

| rth x 4
| + oe ‘
LW TTL 1) 3 jAZzz
+ re . Nak
| a aa x i oA
yO VV) i} uf ur =: “=
ry AA WX) , A vo
eT 4 ye? ek
Ly iY > oe.
A Sima Oo
9 a Se
672893
ee a8 4 Ty Ni”
i ret Benne
- PAL, >
3. CM

Ficure |7.—Lectotype selected herein of Amphiroa valonioides
(from Yendo, 1902, pl. 1; see also Figure 15a): a, part of
fronds with secund branching; 6, part of frond with protrud-
ing conceptacles; c, longitudinal section showing genicula of
single tier of cells.

Gulfo de Fonseca, 7 Sep 1960, EYD-21899
(US). Pacific Coast of Costa Rica. Bahia Ballena,
30 Mar 1959, EYD-21201 (US); Gulfo Dulce, 26
Mar 1939, WRT-39-112 (US) and WRT-39-
116A (UC, US). Pacific Coast of Panama. Isla
Jicaron, 25 Mar 1959, EYD-21122 (US); Isla
Brincanco, Isla Contreras, 25 Mar 1959, EYD-
21052 (US). Ecuador. Esmeraldas, Bahia San
Francisco, 11 Feb 1934, WRT-34-484 (holotype

NUMBER 9

AHFH, isotype MICH). GALAPAGOsS_ ISLANDS:
Isla Santiago (=I. James), Crossland s.n. (holo-
type, A. annulata, BM). Isla Wenman, 11 Jan
1934, WRT-88a (US). Isla Fernandina, 25 Jul
1938, W. L. Schmitt-16-38 (US).

ReMARKS.—Among the most diminutive plants
in Amphiroa are those of this species, with most
specimens having branches less than 400 fm in
diameter. In the field the best way to distinguish
Amphiroa valonioides is by plant size and the fre-
quent occurrence of markedly protruding concep-
tacles. The genicula are comprised of single tiers
of medullary cells. The primary branching may
be sparse, particularly in plants growing in com-
pact tufts. More often fronds have secondary
branches with laterals arising from the intergen-
icula surfaces. This latter feature led to the estab-
lishment of Amphiroa annulata var. pinnata by Daw-
son (1953:137), a taxon which we do not consider
distinctive.

According to Dawson (1953, as A. franciscana,
p. 149, and A. annulata p. 136), A. valonioides has a
wide distribution in the eastern Pacific. From
central Pacific Baja California, it occurs at least
as far south as Ecuador (Taylor, 1945:187, as A.
franciscana), and in Japan (Okamura, 1936:516).
Furthermore, this species may occur in tropical
Pacific islands, but verification of its presence
there remains for future study.

Amphiroa van-bosseae Lemoine
Ficures lg, 4d, 6, 10a, 126,c, 13c, 18

Amphiroa van-bosseae Lemoine, 1929:73 [as ‘van Bosseae’| fig.
30, pl. 3: fig. 7.

Amphiroa ngida sensu Dawson, 1944:276 [in part] [not Am-
phiroa rigida Lamouroux, 1816:297].

Amphiroa subcylindnca Dawson, 1953:139, pl. 29: fig. 1 [type
locality: Punta Colorado, near Guaymas, Sonora, Mexico,
EYD-555 (holotype AHFH 4277, isotype, UC 940252)];
1959:22; 1966a:18.

DescripTion.—Fronds: up to 10 cm high, more
or less erect, often in clumps. Branching: basically
dichotomous, often obscure and irregular. Jnter-
genicula: terete to subterete, 1-2 mm diameter and
variable in length but up to or more than | cm

23

long, length difficult to discern because of geni-
cula that are barely visible, thickening with age.
Genicula: developing by cracking and sloughing of
calcified cortical tissue overlying uncalcified gen-
icula, fully formed genicula usually barely visible
between intergenicula near branch apices, con-
sisting of 5 (rarely 4) or more tiers of medullary
cells and patches of cortical cells. Conceptacles:
scattered over intergenicular surfaces, protruding

E mm

Ficure 18.—Branches of Amphiroa van-bosseae: a, cylindrical
intergenicula (JN-3050) (note they are very long and coarser
than other Gulf of California Ampzroa species); 6, intergeni-
cula with conceptacles (JN & HWJ-73-7-1).

24 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

only slightly, becoming buried by continuing cor-
tical growth, tetrasporangial and _ bisporangial
conceptacles 200-300 um inside diameter, sexual
plants not encountered in the Gulf of California.

Type Locauity.—‘‘Galapagos. Ile Charles,
Post Office Bay, aout 1924 (recueillie juste au-
dessous de la limite de la mer)” (Lemoine, 1929:
US).

Ho.otypre.—St. George South Pacific Expedi-
tion, D. C. Crossland s.n., August 1924 (BM).

GuLF OF CALIFORNIA DiIsTRIBUTION.—Puerto
Penasco to Punta Palmilla.

Pactric Coast Distrrpution.—Galapagos Is-
lands.

SPECIMENS STUDIED.—Gulf of California. SONORA:
Punta Pelicano, vicinity of Puerto Penasco,
2 Jul 1973, JN & HW)J-73-7-1 (US); Playa
Arenosa, vicinity of Puerto Penasco, 20 Oct 1973,
JN-3602 (ARIZ) and 15 Feb 1965, A. E. Dennis-
D62 (US); Cumpleanos Tide Pool, Playa Esta-
cion, Puerto Penasco, 4 Jul 1973, JN. & HWJ-73-
7-28 (CUW); Playa Estacion, in front of Labor-
atorio de Biologia Marina, Puerto Penasco, 27
Jul 1965, EYD-27472 (US), 7 Apr 1966, EYD-
27300 (US), 29 Apr 1972, JN-2957 (US), 20 Oct
1972, JN-3584 (MEXU), 25 Nov 1972, JN-3682
(US), and JN-3808 (CUW); Bahia Tepoca, 4 Feb
1940, EYD-395 (AHFH); Bahia Carrizal, near
Cabo Arco, vicinity of Guaymas, 15 May 1946,
EYD-1693 (AHFH); Guaymas Harbor, 9 Feb
1940, EYD-Sta. 37 (UC); Punta Colorado, near
Guaymas, EYD-555 (holotype AHFH; isotype
UC). BAJA CALIFORNIA: Puertecitos, 17 May
1972, JN-3296 (US); Punta La Gringa, Bahia de
Los Angeles, 22 May 1972, JN-3050 (US); Punta
Concepcion, 15 Jul 1965, EYD-25918 (US); Isla
Ildefonso, 19 Jul 1965, EYD-25890 (US); Isla
Monserrate, 21 Apr 1958, EYD-18794 (US);
Bahia Agua Verde, 20 Apr 1958, EYD 18885
(US); between Eureka and La Ribera, 5 Nov
1946, EYD-3179 (AHFH); Cabo Pulmo, 4 Nov
1946, EY D-3114 (AHFH); Punta Palmilla, 7 Nov
1946, EYD-3219 (AHFH). Las IsLAS DE LA CIN-
TURA: Isla Patos, Isla Tiburon, 17 Feb 1946, EY D-
806 (US); Bahia Agua Dulce, Isla Tiburon, 21
Feb 1946, EYD-979 (AHFH); Isla San Esteban,

5 Feb 1940, EYD-460a (AHFH); Isla Turner, off
Isla Tiburon, 26 Jan 1940, EYD-112a (AHFH).

ReMARKS.—In accordance with Article 73.9
under examples (Stafleu et al, 1978), “a hypen is
correctly used in an epithet after a word which
could stand independently. ...” Therefore, we
spell the species epithet as “van-bosseae,’ and do
not use the original spelling of Lemoine (1929:
73), “van Bosseae,” or the spelling of Dawson (e.g.,
1961a:421) as “vanbosseae.”

Specimens belonging to Amphiroa van-bosseae
may usually be recognized by the robust fronds
and terete branches; it is the largest cylindrical
species in the Gulf of California. Small specimens
of A. van-bosseae can be difficult to separate from
Amphiroa beauvoisii based solely on gross morphol-
ogy, but examination of their genicula readily
allows them to be separated. In A. van-bosseae the
genicula are of 5, or more often, 6 to 10 cellular
tiers (Figure 126) whereas in A. beawvorsii there are
3 to 5 cellular tiers (Figure 5). In the Gulf only A.
van-bosseae combines many-tiered genicula with
terete intergenicula (rarely are they compressed).

Among the species of Amphiroa containing large
plants with mostly terete intergenicula, A. ephed-
raea (Johansen, 1968, fig. 1) 1s most striking in
appearance and superficially resembles A. van-
bosseae. Amphiroa ephedraea occurs in the western
part of the Indian Ocean (Johansen, 1968:319).
The reports of this species from Japan (Okamura,
1936:518; Chihara,_1970:72) may be plants of
Amphiroa magdalensis or possibly Amphiroa beauvoisit.
The unique genicula in the Indian Ocean A.
ephedraea (Johansen, 1969b:122) do not corre-
spond to the Gulf of California species, in which
development is as in Amphiroa anceps and as de-
scribed by Johansen (1969b:120) for the Type I
category of geniculum development. To the na-
ked eye, branches of A. ephedraea consist of inter-
genicula of more or less uniform length separated
by conspicuous genicula, whereas those of A. van-
bosseae have intergenicula of greatly varying
lengths (Figure 18), and the genicula become
visible only after overlying calcified tissue breaks
away (Type I genicular development, Johansen,
1969b:120). In the Gulf species, genicula are most

3”

NUMBER 9

conspicuous in older parts of the branches and
may, in fact, be externally invisible where they
are buried in calcified tissue near branch apices.
The tendency for intergenicular cortices to con-
tinue growing is more evident in A. van-bosseae
than in the other Gulf species. Hence, older in-
tergenicula are greater in girth than are young
intergenicula (Figure 6), and conceptacles may

ibe)
(Sn

become buried by cortical growth (Figure 14c), a
phenomenon not occurring in the other species.
Amphiroa van-bosseae is plentiful in the Gulf, but
the lack of published reports or specimens known
to us from elsewhere suggest a limited distribu-
tion. The type locality is Post Office Bay, Isla
Floreana (=Ile Charles), Galapagos Islands, and

more material from this area would be welcome.

Literature Cited

Bgrgesen, F.

1917. The Marine Algae of the Danish West Indies, Part
III: Rhodophyceae (3). Dansk Botanisk Arkiv, 3(1c):
145-240.
Briggs, J. C.
1974. Marine Zoogeography. 457 pages. New York: Mc-

Graw-Hill Book Co.
Cabioch, J.
1969. Sur le mode de développement de quelques Am-
phiroa (Rhodophycees, Corallinacées). Compte
Rendu de Academie de Science (Paris), 269D:2338-
2340.

1972. Etude sur les Corallinacées, II.—La morpho-
genese: consequences systematiques et phylogeéne-
tiques. Cahiers de Biologie Marine, 13:137-288, 7
plates.

Chihara, M.
1970. Common Seaweeds of Japan in Color. xviii + 173 pages,
64 plates. Osaka: Hoikusha Publishing Co., Ltd.
Dawson, E. Y.

1944. The Marine Algae of the Gulf of California. Allan
Hancock Pacific Expeditions, 3(10):189-454.

Marine Red Algae of Pacific Mexico, Part I:
Bangiales to Corallinaceae Subf. Corallinoideae.
Allan Hancock Pacific Expeditions, 17:1—240, plates
1-33.

Resumen de las Investigaciones recientes sobre

1953.

1954.
algas marinas de la costa Pacifica de Mexico, con
una sinopsis de la literatura, sinonimia y distri-
bucion de las especies descritas. Revista de la Soctedad
Mexicana de Historia Natural, 13:97-197 + i—x.
[Reprint of 1953, with corrections, index, pagina-
tion and addenda. |

Marine Algae from the Pacific Costa Rican Gulfs.
Los Angeles County Museum Contributions in Science,
15:1-28.

Marine Algae from the 1958 Cruise of the Sted/a
Polans in the Gulf of California. Los Angeles County

Museum Contributions in Science, 27:1—39.
A Review of the Ecology, Distribution and Affin-

ities of the Benthic Flora. Jn The Biogeography of
Baja California and Adjacent Seas. Part II: Ma-
rine Biotas. Systematic Zoology, 9 (3/4):93-100.

A Guide to the Literature and Distribution of
Pacific Benthic Algae from Alaska to the Gala-
15(3):370-461.

Plantas marinas de la zona de las mareas de El

1957.

1959}

1960.

196 1a.

pagos Islands. Pacific Science,

196 1b.
Salvador (Intertidal Marine Plants of El Salva-
dor). Pacific Naturalist, 2(8):389-461.

1962. Additions to the Marine Flora of Costa Rica and
Nicaragua. Pacific Naturalist, 3(13):375-395.
1966a. Marine Algae in the Vicinity of Puerto Penasco,
Sonora, Mexico. Gulf of California Field Guide Series,
1: ii + 57 pages, map. Tucson: The University of
Arizona.
1966b. New Records of Marine Algae from the Gulf of
California. Journal of the Arizona Academy of Sciences,
4:55-66.
Dawson, E. Y., C. Acleto, and N. Foldvik
1964. The Seaweeds of Peru. Bezhefte zur Nova Hedwigia,
13:[5] + 111 pages, 81 plates.
Dawson, E. Y., M. Neushul, and R. D. Wildman
1960a. Seaweeds Associated with Kelp Beds along South-
ern California and Northwestern Mexico. Pacific
Naturalist, 1(14):1-81.
New Records of Sublittoral Marine Plants from
Pacific Baja California. Pacific Naturalist, 1(19):1-
30.
Ellis, J., and D. Solander
1786. The Natural History of Many Curious and Uncommon
Zoophytes, Collected from Various Parts of the Globe by
the Late John Elhs . . . Systematically Arranged and De-
scribed by the Late Daniel Solander...., xii + 208
pages, 63 plates. London: Benjamin White and
Son.
Ganesan, E. K.
1968. Studies on the Morphology and Reproduction of
the Articulated Corallines, III: Amphiroa Lamour-
oux emend. Weber van Bosse. Phykos, 6:7—28.

1960b.

Hamel, G., and P. Lemoine

1953. Corallinacées de France et d’Afrique du Nord.
Archives du Museum National Histoire Natural (Paris),
series 7, 1:17-136.
Hariot, P.
1895. Algues du Golfe de Californie recueillies par M.

Diguet. Journal de Botanique, 8:167-170.
Holmgren, P. K., and W. Keuken
1974. Index Herbariorum, Part I: The Herbaria of the World.
Sixth edition, vii + 397 pages. Utrecht, Nether-
lands: Oosthoek, Scheltema & Holkema.

Hommersand, M. H.
1972. Taxonomic and Phytogeographic Relationships of
Warm Temperate Marine Algae Occurring in
Pacific North America and Japan. In K. Nisizawa,
editor-in-chief, Proceedings of the Seventh International
Seaweed Symposium, Sapporo, Japan, August 8-12,
1971, pages 66-77. New York: John Wiley & Sons.

26

NUMBER 9

Humason, G. L.

1967. Animal Tissue Techniques. Second edition, ix + 569

pages. San Francisco: W. H. Freeman and Co.
Johansen, H. W.

1968. Reproduction of the Articulated Coralline Amphi-
roa ephedraea. Journal of Phycology, 4:319-328, 26
figures.

1969a. Morphology and Systematics of Coralline Algae
with Special Reference to Calliarthron. University of
California Publications in Botany, 49:1-78, 19 plates.

1969b. Patterns of Genicular Development in Amphiroa
(Corallinaceae). Journal of Phycology, 5:118-123.

1973. Ontogeny of Sexual Conceptacles in a Species of
Bossvella (Corallinaceae). Journal of Phycology, 9:
141-148.

1974. Articulated Coralline Algae. Oceanography and Ma-

rine Biology Annual Reviews, 12:77—127.
1976a. Phycological Reviews, 4: Current Status of Ge-
neric Concepts in Coralline Algae (Rhodophyta).
Phycologia, 15:221—244.
1976b. Family Corallinaceae. Jn I. A. Abbott and G. J.
Hollenberg, Marine Algae of California, pages 379-
419. Stanford, California: Stanford University
Press.
1981. Coralline Algae, a First Synthesis. 256 pages. Baco
Raton, Florida: CRC Press.
Johansen, H. W., and P. C. Silva
1978. Janieae and Lithotricheae: Two New Tribes of
Articulated Corallinaceae (Rhodophyta). Phycolo-
gia, 17:413-417.
Kutzing, F. T.

1858. Tabulae phycologicae oder Abbildungen der Tange. Vol-
ume 8, 11 + 48 pages, 100 plates. Nordhausen,
Germany.

Lamouroux, J. V. F.
1812. Extrait d’un memoire sur la classification des Po-

lypiers coralligenes non entierement pierreux. Nou-
veau Bulletin des Sciences, publié par la Societe Philom-
atique de Paris, 3(63):181-188.

1816. Histoire des Polypiers Coralligenes flexibles, le vulgairie-
ment nommes zoophytes. \xxxiv + 560 pages, 19
plates. Caen: F. Poisson.

Lemoine, Madame Paul

1929. Les Corallinacees de l’Archipel des Galapagos et
du Golfe de Panama. Archives du Muséum d’Histoire
Naturelle (Paris), series 6, 4:37-86 + [2], 35 figures,

4 plates.
Murata, K., and T. Masaki
1978. Studies of Reproductive Organs in Articulated
Coralline Algae of Japan. Phycologia, 17(4):403-
412.
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.

27

1976. Resena Historica de las Exploraciones Marinas
Botanicas en le Golfo de California. /n B. Braniff
C. and R. S. Felger, editors, Sonora: Anthropologia
del Desierto, pages 79-84. Mexico, D. F.: Instituto
Nacional de Antropologia y Historia [Coleccion
Cientifica Diversa, 27].

Okamura, K.

1936. Nippon Kaiso-shi [Marine Algae of Japan]. Frontis-
piece, 9 + 6 + 964 + 11 pages. Tokyo.

Schneider, C. W.

1976. Spatial and Temporal Distribution of Benthic
Marine Algae on the Continental Shelf of the
Carolinas. Bulletin of Marine Science, 26(2):133-151.

Segawa, S.

1940a. Systematic Anatomy of the Articulated Coral-
lines (1): Amphiroa rigida Lamouroux. Journal of
Japanese Botany, 16:219-225.

1940b. Systematic Anatomy of the Articulated Corallines
(II): Amphiroa misakiensis Yendo. Journal of Japanese
Botany, 16:488-595.

Silva, P. C.

1966. Status of Our Knowledge of the Galapagos
Benthic Marine Algal Flora Prior to the Galapa-
gos International Scientific Project. Jn R. I. Bow-
man, editor, The Galapagos: Proceedings of the Sym-
posta of the Galapagos International Scientific Project,
pages 149-156. Berkeley: University of California
Press.

Stafleu, F. A., et al.
1978. International Code of Botanical Nomenclature. xiv + 457
pages. Utrecht: Bohn, Scheltema & Holkema.

Suneson, S.
1937. Studien uber die Entwicklungsgeschichte der Cor-
allinaceen. Lunds Universitets Arsskrift, Ny Foeljd

Avdeling 2, 33(2):1-101.
Taylor, W. R.

1945. Pacific Marine Algae of the Allan Hancock Ex-
peditions to the Galapagos Islands. Allan Hancock
Pacific Expeditions, 12:[vi] + iv+ 528, 100 plates.
Manne Algae of the Eastern Tropical and Subtropical
Coasts of the Americas. xi + 870 pages. Ann Arbor:
University of Michigan Press.
van Andel, T. H., and G. S. Shor

1964. A Symposium: Marine Geology of the Gulf of California.

408 pages. Menasha, Wisconsin: George Banta
Company [American Association of Petroleum

1960.

Geologists Memoir, 3].
Weber-van Bosse, A.

1904. II: Corallineae Verae of the Malay Archipelago.
In A. Weber-van Bosse and M. Foslie, The Cor-
allinaceae of the Siboga-Expedition. Szboga-Expe-
ditie: Urtkomsten op Zoologisch, Botanisch, Oceanogra-

28 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

phisch en Geologisch Gebied. . . uatgegeven door M. We- Science, Imperial University, Tokyo, Japan, 16 (part 2,
ber... Monographie, 61:78-110, plates 14-16. Lei- art. 3):1-36 + [2], 7 plates.
den: E. J. Brill. 1904. Study of the Genicula of Corallinae. Journal of the

Yendo, K. College of Science, Imperial University, Tokyo, Japan,
1902. Corallinae Verae Japonicae. Journal of the College of 19(14):1-44 + [2], 1 plate.

Index

(Principal taxa in roman letters; synonyms or taxonomic misidentifications in zlalics)

Amphiroa, 1, 3
anceps, 13, 14, 24
annulata, 2, 20, 23
annulata var. pinnata, 2, 21, 23
beauvoisii, 2, 5, 6, 11, 12, 24
brasiliana, 12
brevianceps, 2, 5, 6, 12, 13, 14
crosslandu, 6, 12
dimorpha, 2, 15, 19
dimorpha var. digitiformis, 2, 15, 19
drouetti, 2, 6, 11
echigoensis, 12
ephedraea, 14, 24
foliacea, 14
franciscana, 2, 20, 21, 23
franciscana var. robusta, 2, 6, 12
galapagensis, 12
linearis, 2, 6, 12
magdalensis, 2, 5, 6, 13, 14, 24
mexicana, 12

28)

misakiensis, 2, 5, 13, 15, 19
peninsularis, 6
pusilla, 2, 6, 15
rigida, 2, 23
rigida, 2, 5, 6, 19, 20
rigida var. antillana, 20
subcylindrica, 2, 23
tayloriz, 2, 19, 20
tribulus, 5
valonioides, 1, 2, 5, 6, 20, 23
van-bosseae, 2, 5,
zonata, 2, 6, 11, 12
Bossiella, 9
california ssp. schmitt, 19
Corallina, \
pinnatifolia var. digitata, |
tribulus, 5
Jama, |
Neogoniolithon notarisii, 20

ig:

bc,

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