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of the Museum of

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CIROLANIDAE

(CRUSTACEA: ISOPODA: FLABELLIFERA)

OF THE TROPICAL EASTERN PACMC

by

Richard C. Brusca

Regina Wetzer

Scott C. France

r

No. 30

15 March 1995

Proceedings of the

San Diego Society of Natural History

ISSN 1059-8707

MCZ

LIBRA, .;y

APR 1 0 \rJb

PROCEEDINGS

of the ,,.... -^

San Diego Society of Natural History

HARVARD

Founded 1874

Number 30

15 March 1995

Cirolanidae (Crustacea: Isopoda: Flabellifera) of the Tropical Eastern Pacific

Richard C. Brusca'

Regina Wetzer'

Scott C. France

San Diego Natural Hisian Museum. P. O. Box 1390. San Die,i>o. California 92112

Dedicated to the memory of J. Laurens Barnard

ABSTRACT.-The cirolanid isopods of the tropical eastern Pacific (central western Baja California, Mexico, to the Peru-Ecuador border)

encompass 18 species in 8 genera, including 7 species newly described here: Anopsilana alec. Cirolana namelessensis. Cirolana nielhnuei

Comlera bulks,. Metacrolana calypso. Nauuolana carlenae. and Onc,lorphe:,s jernbarnardi. Complete species lists are provided for all genera

treated. The tropical eastern Pacific genera and species are descnbed and keyed, and their ecology and distnbution are discussed. The second known

species ot 0„c,lorphe„s and third known species of Conilera are descnbed, both representing the first records of the genera from the Pacific Ocean

hitteen ot these 18 species (83%) occur primanly in the tropical eastern Pacific zoogeographic region. Four species (22%) are amphi-American in

distnbution. One species is endemic to the Gulf of California (O>oto,« nielbrucei n. sp.), and one is endemic to the Galapagos Islands ^Metacaolana

calypso n. sp.). Compared to that of the tropical Canbbean and tropical Indc^west Pacific, the cirolanid fauna of the tropical eastern Pacific is

relatively depauperate.

INTRODUCTION

This paper is one in a series of regional monographs and shorter

papers (jescribing the shallow-water marine isopods of the tropical

eastern Pacific [the "Panamic region" of Ekman (1953), the "east-

ern Pacific zoogeographic region" of Brusca and Wallerstein

( 1979a)]— central western Baja California and the Gulf of Califor-

nia (Mexico) to the Gulf of Guayaquil, Ecuador The family

Idoteidae was treated by Brusca (1983, 1984). Brusca and

Wallerstein ( 1 977, 1979b), and Wallerstein and Brusca ( 1982). The

family Cymothoidae was treated by Brusca (1977, 1978a, 1978b,

1981), Brusca and Gilligan (1983), and Thun and Brusca (1980),

The fainilies Corallanidae and Excorallanidae were covered by

Delaney ( 1 982, 1984, 1 986, 1 989), Bruce et al. ( 1 982). and Delaney

and Brusca (1985). The family Aegidae was treated by Brusca

(1983 land Brusca and France ( 1992). Several analytical and sum-

mary papers have also appeared (Brusca and Wallerstein 1979b,

Brusca and Iverson 1985, Brusca 1987, Brusca and Wilson 1991).

The earliest references to eastern Pacific tnarine isopods date

from the mid- 1800s and the work of James Dana, William

Lockington, Jaines Benedict, and Pearl Lee Boone. However, it was

the pioneering 20-year research program of Harriet Richardson that

laid the foundation of our modern knowledge regarding this fauna.

Milton Miller, Robert Menzies, and George .Schullz built on

Richardson's foundation through the 1970s. However, almost all of

this work was concerned with the temperate eastern Pacific, and

there have been only a few studies of the Cirolanidae of the tropical

'Cun-ent address: Gnce Manne Biological Laboratory. University of Charleston. Charleston, SC

eastern Pacific and until now no attempt at synthesizing the fauna.

The history of cirolanid taxonomy in general was reviewed in

Bruce's (1986a) excellent treatment of the Australian fauna.

METHODS AND TERMINOLOGY

Specimens upon which this study is based were obtained by

field collecting and by loans from various museums. Field collec-

tions were made by Brusca and/or Wetzer in California, Mexico,

Guatemala. Nicaragua, Costa Rica. Panama, Peru, the Galapagos

Islands. Brazil. Uruguay, and various localities in the Caribbean.

Field samples were taken by SCUBA and shore collecting, and

included washes and 0.24-0.5-mm screening.s of material from

various habitats (rocks, algae, coral, coral rubble, sediment, etc.).

All material was fixed and preserved in 70% ethyl alcohol and

examined with dissecting and compound microscopes. Appendages

were drawn with the aid of a caiTiera lucida. Institutional abbrevia-

tions used in this paper are as follows:

AHF — Allan Hancock Foundation, University of Southern Cali-

fornia. Los Angeles. California (all AHF specimens are now housed

atLACM)

BMNH — The Natural History Museum. London [formerly the

British Museum (Natural History 1|

LACM — Los Angeles County Museum of Natural History, Los

Angeles. California

ISSN 1059-8707

R. C. Brusca, R. Wetzer. and S. C. France

SDNHM — San Diego Natural History Museum, San Diego.

California

SIO — Scripps Institution of Oceanography, University of Cali-

fornia. La Jolla, California

SOSC — Smithsonian Oceanographic Sorting Center (USNM),

Washington. D. C,

SMF — Senckenberg Museum, Frankfurt, Germany

UA — University of Arizona, Tucson, Arizona

UCR — Natural History Museum, Universidad de Costa Rica.

San Jose. Costa Rica

USL — University of Southwestern Louisiana. Lafayette. Loui-

siana

USNM — National Museum of Natural History. Smithsonian

Institution, Washington. D.C.

ZMUC — Zoological Museum. University of Copenhagen, Den-

mark

Other abbreviations used include PMS, plumose marginal setae:

SEM, scanning electron microscope; coll.. collector; det.. deter-

mined by; m. meters; mm. millimeters; mi., miles; N.. north; S..

south; E., east; W.. west.

The literature regarding terms used to identify the surfaces of

pereopods is often confusing. Surfaces referred to by some workers

as "anterior" have been referred to by others as "posterior." Simi-

larly, workers have vacillated between the terms "inner" and

"outer," and "medial" and "lateral." In part, the reason for this

confusion is that in most isopods the anterior legs are usually

oriented differently from the posterior legs (Brusca and Wilson

1991 ). In flabelliferans pereopods l-IIl are generally directed for-

ward at an anterolateral angle, whereas pereopods IV-VII are gen-

erally directed backward at a posterolateral angle. To avoid confu-

sion, we use the terms inferior (= ventral), superior (= dorsal),

anterior, and posterior as indicated in Fig. 1 when describing pereo-

pod anatomy. Thus, "inferior" refers to the pereopodal surface

forming the concave curve of the ischium-dactylus. We use the

term "medial" to describe the position of structures near or toward

the animal's midline, and we reserve the term "lateral" for struc-

tures located away from the midline of the body, not in reference to

the axis of particular appendages.

Similarly, the terms "setae" and "spines" have been used vari-

ously by different authors. Arthropod setae are generally defined as

arising from well-developed sockets and being movable, except

those which are secondarily immobilized because of heavy

cuticularization. The sockets are formed by invaginations of the

cuticle, not of the whole integument. Arthropod spines, on the other

hand, regardless of their shape or size, are generally defined as not

being articulated and are characterized by an evagination by the

whole integument. The degree of development of setal sockets

varies from elaborately sunken for heavy setae to delicately in-

serted for the fine setae on the tergal surfaces of many crustaceans.

Although these definitions of "seta" and "spine" are widely ac-

cepted, they are rarely adhered to in the literature on isopods. and it

is not always easy to discern articulations or the presence of sockets

by light microscopy. Furthermore, at least in the isopod literature,

there has been a tradition of referring to large, smooth, robust setae

as "spines," especially if the socket is more or less immobilized. In

keeping with terminology currently used in isopod systematics, we

use the term "seta" for a long, thin, flexible, aniculating cuticular

process, and we reserve the term "spine" for a large, robust, rigid,

cuticular process (whether or not a socket is discernable). We

standardize the terminology used for spines by describing them as

simple, apically forked, basally senate, serrate trident, molariform,

squamate, or circumplumose (Fig. 2). We standardize the terminol-

ogy used for setae by describing them as simple, plumose,

circumplumose. comb, serrate, serrate trident, or palmate (Fig. 2).

The term PMS is restricted to the plumose marginal setae occurring

SUPERIOR

ANTERIOR

SURFACE

DISTAL

Figure 1. A cirolanid isopod (Buihyimmus gigaiueus). illustrating terms used in the text to describe pereopods. In tlabelliferan isopods pereopods I-III

are generally directed forward at an anterolateral angle, whereas pereopods IV-VII are generally directed backward at a posterolateral angle.

long fleshy spine

stout circumplumose spine

serrate spine

Figure 2. Some types of setae and spines found on cirolanid isopods. A, acute and long fleshy simple spines (mandible oi MetiiciroUma costaricensis).

B, stout circumplumose spines (maxillule of AnopsiUiiui oaxaca). C, plumose setae and coupling spines(maxillipedof/l. oaxaca). D, aesthetascs (antennule

of A. ouxaca). E. palmate setae (antennule of Naunnlaiui curlenae n. sp.). F. circumplumose setae (maxilla of A. oiixtiia). G, aesthetascs (antennule of N.

iuriemie n. sp). H, comb setae (maxilliped of CiroUma hai-fonti). 1, comb setae (maxillipedof C Iniiinnli). J, serrate trident setae (maxillipedofA. oaxuca).

K. molariform spines (pereopod 1 of C. Iiurfurdi). L. plumose setae and coupling spines (peduncle of pleopod 3 of C. harfonli). M, simple spines and PMS

(uropod of A. (Htxacii). N, serrate trideni spines (pereopod VII of A. oaxiicu). O, simple spines, serrate spines, and palmate setae (pereopod 1 of C. huifonli).

R. C. Brusca, R. Wetzer, and S. C. France

on the rami of the pleopods and uropods and on the posterior margin

of the pleotelson. The term coupHng spine is reserved for the

specialized spines of maxillipedal endiles and pleopodal peduncles.

In most Peracarida the dactylus terminates in a spinelike "claw,"

usually called the unguis, which may he marked off by a suture or

immovable articulation. In many other malacostracans this terminal

spine is distinct and freely articulating. In his early work, Hansen

(190^) considered this free terminal spine an additional leg article,

but he later to came to agree with Caiman ( 1 909) that it is simply an

enlarged spine. In his revision of the isopod subfamily Lipomerinae

(Asellota: Munnopsidae), Wilson (1989) described this fixed spine,

or unguis, as "a modified seta on the tip of the dactylus." In most

isopod suborders there are some genera in which this fixed terminal

spine, or unguis, may be accompanied by one or more additional

stout, fixed, clawlike spines arising near the base of the unguis. The

size of these "secondary spines" ranges from very short to as large

as the primary spine, or unguis, itself.

For example, in many Gnathiidea a "secondary spine" is present

but small, while in the Calabozoidea and many Phreatoicidea,

Oniscidea, Anthuridea, Sphaeromatidae, Cirolanidae, Limnoriidae.

and others a "secondary spine" one-quarter to one-half the size of

the unguis often occurs. In some Asellota the "secondary spine"

may also be half the size of the unguis (e. g., Munnogonium). or

even fully as large as the unguis (e. g., Stenetrium). In the asellotan

genus Joeropsis there are often two secondary spines, both as large

as the unguis.

Many authors refer to these "secondary spines" as "secondary

ungui" and to such legs as biungulate (or triungulate when two

"secondary spines" are present). However, the point at which small

spines become large enough to warrant being called "secondary

ungui" is subjective, and the literature is therefore unclear and

inconsistent regarding the terms "secondary ungui," "bigungulate,"

and "triungulate." Because of this confusion, and because we pre-

sume all these stout fixed spines to be homologous in a general

sense, we avoid the terms "secondary ungui" and "biungulate" and

instead simply note the occurrence of additional spines at the base

of the ungui when they are present.

Lengths of isopods were measured from the frontal margin ot

the cephalon to the posterior margin of the pleotelson by holding

the animal (not unnaturally) flat and using a calibrated ocular

micrometer. Numerical comparisons of structures given in the text

(such as "endopod width 0.5 times exopod width") were obtained

by measuring the greatest dimensions of the structures concerned,

unless otherwise noted.

We give complete synonymies except where recent monographs

have already done so. In the latter case, we provide synonymies

subsequent to (and often expanding upon) the cited reference.

Species descriptions are based on primary type material, with

supplemental data (i. e., polymorphism) from other material exam-

ined. Primary types of most species are illustrated, and, for some,

additional (Pacific) specimens are also figured. Some older type

material is damaged, with obviously missing spines, setae, etc.

Body measurements and ranges are based on all material examined.

Taxa are treated in alphabetical order within each higher taxon.

Localities listed under Material Examined are ordered from north to

south and numbered sequentially. The species lists for each genus

are updated from Bruce ( 1986a).

Several authors (e. g., Racovitza 1912, Menzies 1962a, Bruce

1986a, Kensley and Schotte 1989) have suggested various generic

groupings within the Cirolanidae. However, as the number of gen-

era has grown, discerning possible monophyletic groupings intu-

itively has become increasingly difficult [note Kensley's (1987)

comments in this regard]. Furthermore, several of the existing

genera could be nonmonophyletic (e. g., Anopsilana. Cirolana.

Melacimlami). Therefore, until an analytical phylogenetic analysis

is accomplished, we feel it is best to treat the genera separately

rather than to contribute to the creation of numerous new names and

combinations that will only continue to be altered and shuffled

about until phylogenetic relationships have been properly assessed.

TAXONOMY

Order Isopoda Latreille, 1817

Suborder Flabellifera Sars, 1882

Family Cirolanidae Dana, 1853

SyntiiiyiiiY. — Emended and subsequent to Bruce ( 1986a;7).

Barnard 1914; 35()A (as Eurydicidae), Brusca and Iverson 1985: 30.

Kensley and Schotte 1989: 122.

Descriinion. — Body symmetrical, usually sleek but occasion-

ally ornamented, rough or nodulose, 2-6.5 times longer than wide.

Eyes lateral, small to moderate in size, vestigial or absent in many

subterranean and deep-water species. Frontal margin of cephalon

evenly convex or produced into short rostrum (= rostral process);

cephalon usually with posterolateral grooves ("incisions") and an

interocular carina. Antennular peduncle 3-articulate, occasionally

reduced to 2 articles; "scale" present (on third article) only in

Balhynonuis: in most taxa the third article is followed by a short

fourth article of uncertain homology; flagellum multiarticulate.

Antennal peduncle 5-articulate, occasionally reduced to 4 articles;

flagellum multiarticulate. Frtintal lamina, clypeus, and labrum dis-

tinct; free posterior margin of labrum concave. Mandible with

fundamentally tridentate incisor, frequently with an accessory tooth

on right or left mandible; spine row well developed and lobelike;

lacinfa apparently absent (see below); molar process flat, elongate

and bladelike, with row of stout submarginal spines along upper

(anterior) margin; palp ,3-articulate (reduced to 2 articles in some

.species). Maxillule lateral (= outer) lobe with 10-14 apical spines;

medial (= inner) lobe with 3-4 circumplumose spines; neither lobe

ever forms a stylet (as in many flabelliferan families, e. g., Aegidae,

Corallanidae, Cymothoidae, Tridentellidae). Maxilla setose, bi-

lobed, outer lobe divided (maxilla of a single lobe in Oncilorpheus).

Maxillipedal palp typically 5-articulate, articles never with hooked

or strongly recurved spines; endite distinct, minute to large. Maxil-

lipeds with epipod present only in gravid females.

Coxal plates on pereonites II-VII well developed, fused to

tergites but with distinct suture lines; usually with oblique-trans-

verse grooves running from anterior to posterior, but absent in

many species. Pereopods ambulatory; I-III tend toward a grasping

subprehensile form with dactyli moderately developed but almost

always shorter than propodi.

Pleon usually with 5 free pleonites plus pleotelson; fusion and

thus reduction in number of free pleonites occur in a number of

genera. Lateral margins of pleonite 5 often overlapped by pleonite

4. Pleopods membranous, without ridges and only rarely with folds,

often with lateral incisions and accessory lobes. Pleopod 5 endopod

usually naked. Uropodal rami well developed, almost always lamel-

lar and mobile, fornnng "tail fan" with pleotelson (exopod absent or

reduced in some genera). Pleotelson and uropods usually with

marginal setae and/or spines.

Remarks.— Ihe name Cirolaninae was first used by Dana (pub-

lished 1853, but dated 1852) and thus has precedence over

Cirolanidae of Harger (1880) and Eurydicidae of Stebbing ( 1905).

In their phylogenetic analysis of the Isopoda, Brusca and Wilson

( 1991 ) were unable to identify any unique synapomorphies of this

family, suggesting the possibility that it is a paraphyletie group.

Bruce ( 1986a) provided an excellent discussion of the morphologi-

cal characters used in cirolanid taxonomy, and there is no need to

repeat that information. However, we will comment on the antennal

peduncles and mandibles.

Tropical Eastern Pacific Cirolanidae

The nature of the antennal peduncle in the Cirtilanidae (indeed,

the Isopoda in general) has been somewhat confused in the litera-

ture. Milne Edwards and Bouvier (1902) described the antennal

peduncle of Hciihviwnm.s as 6-articulate. However, they appear to

ha\e niistaken the large articulating (arthroidal ) menibrane between

articles 1 and 2 for an extra article, as noted by Bruce (1986a).

Hansen ( 1 903) also described the antennal peduncle o{ Bcithynomus

as 6-articulate. focusing on a minute strip of sclerotized cuticle at

the base of the peduncle, at the edge of the ailiculating membrane,

that he considered to be the vestige of a proximal antennal article, or

precoxa. Hansen's conclusion that this piece of cuticle is homolo-

gous to a preeoxal article was based on his observation that it

moved ("articulated") within the antennal socket when the antenna

was moved. Hansen (1903) noted similar conditions in several

species of 0'/o/(»u;, and later (1 905. 1916) he added Coiulerci to the

list of cirolanid genera with 6-articulate antennal peduncles. In his

1925 review. Hansen concluded that the 6-articulate condition was

primitive for the Isopoda as a whole. Thus. Hansen's view was that

the isopods" antennal peduncle primitively constituted a 3-articu-

late sympod (Hansen's "basipodite") combined with 3 additional

basal articles of the flagellurn. This homology was primarily based

on the presence of a "scale" (or "squama") on the third antennal

article of the many Asellota that have 6-articulate antennae. Caiman

( 1909) and Wagele( 1983) agreed with Hansen's conclusion that the

6-articulate condition is primitive for isopods.

We are not entirely convinced that the 6-articulate antennal

peduncle is primitive in the Isopoda. or that it occurs among the

Cirolanidae. Brusca and Wilson ( 1991 ) examined the basal cuticu-

lar pieces noted by Hansen (and others) on the articular membrane

of the antenna of Biitliynoniiis gigaiUeus and B. doederUni and also

found it to move when the antenna is moved. However, they noted

that this piece does not articulate with any other article, or with the

head, but simply floats free on the membrane. Similar free-tloating

cuticular pieces occur in many other genera of Cirolanidae. Bruce

( 1986a) commented on these .structures, noting them in at least 12

Australian genera. Splitting and fragmentation of the proximal

antennal article is common in many isopod groups (e. g., Ligia-

morpha, Anthuridea. Phreatoicidea). and in cirolanids these

arthrodial sclerites may be free-floating pieces fractured off the first

peduncular article (or off the head). We believe that the

homologization of these arthrodial sclerites with a preeoxal article

is still speculative. Among the Malacostraca. an antennal precoxa

(and hence a 3-articulate sympod) occurs unquestionably only in

the mysidaceans. tanaidaceans. and asellotan and microcerberid

isopods. In all other malacostracans the sympod comprises 2 ar-

ticles, and the rami (or scale) arise from the second article. Brusca

and Wil.son ( 1991 ) discussed this unresolved matter in more detail.

The nature of the spinose lobe on the cirolanid mandible, be-

tween the molar and incisor processes, is also still unsettled. Some

researchers (e. g., Kensley 1984, 1987, 1989; Muller 1991 ) refer to

it as a "spine row," while others (e, g., Bruce and Javed 1987,

Bowman and Iliffe 1983. 1987. 1991) have called it a lacinia

mobilis. Most recently. Bruce (1991. in tin.) has concluded that it is

a true spine row. the lacinia being absent in most cirolanids. In

Bruce's view, a lacinia may occur only in Bathynoinus. some spe-

cies of Metacirolana. Polilolana dasyprion, Cimhinii hinyciiui. and

perhaps a few others, and it is represented by the distal biting edge

on the lobe bearing the spine row. which takes the form of a large

blunt tooth in these taxa. However, a similar distal sclerotized biting

edge occurs in other flabelliferans as well. e. g.. Plioiaiopiis reiue.x

(see Bruce 1981c). Among cirolanids, specialization of the distal

region of the spine row is not uncommon, but it presents no clear

pattern among the species or genera. The distal region may be

sclerotized, or bear a sclerotized tooth or biting edge, as in Balliy-

nomus, Neocimlana mcuidata (see Bruce 1986a). Mehicinilana

basteni (see Bruce 1980a). and Metacirolana riigosa (see Bruce

19S0b). Furthermore, the most distal spine group of the spine row

often differs from the more proximal spines, as seen in Natalolanu

pastorei (see Wagele and Bruce 1989) and Alarholana exoconui

(see Bruce and Javed 1987). The precise homology of these struc-

tures is not yet understood. However, because the distal sclerotized

biting edge on Badiynomus and others does not articulate, and

because specialization of the most distal spines is common, we tend

to regard these simply as a specialized region or specialized spines

of the spine row. Thus it appears that a lacinia mobilis is absent in

the Cirolanidae.

The majority of cirolanid species are free-living predators or

carnivorous scavengers (Johnson 1976a, Holdich 1981. Stepien and

Brusca 1985. Bruce 1986a). Species of Cimlana. Conilera. and

Polilolana ire often captured in offshore benthic traps baited with

dead animals (e. g.. Wetzer et al, 1987, Biernbaum and Wenner

1993). Species of Cimlana and Natalolana are often reported at-

tacking both living and dead animals, as reviewed by Stepien and

Brusca (1985). Certain predaceous species are occasionally re-

ferred to as parasites, because they are often found temporarily

feeding on the surface of fishes or large invertebrates. However,

most if not all of these species cling to the prey only long enough to

take a meal, and they are therefore best considered micropredators.

The mandibles of cirolanids are well suited for biting and chewing,

and many can inflict a reasonably painful bite on swimmers (e, g..

see discussion of Excirolana). Many species burrow in sand or live

under rocks, while others spend much of their lives in algal turfs,

mussel beds, kelp holdfasts, the chambers of sponges, or burrows of

other animals. In almost all cases these habitats are utilized oppor-

tunistically. Most cirolanids are good swimmers, entering the water

column at various times. Trapping suggests that many species are

nocturnal. Experimental evidence suggests that at least one species.

Cirolana dimimita. emerges nocturnally into the water column to

prey on certain nearshore fish species (Stepien and Brusca 1985).

Many species are known to have endogenous activity rhythms [see

Hastings ( 1981 ) for an introduction to that literature, and DeRuyck

et al. ( 1991 ) for a recent summary], Cirolanids are most common in

shallow-water habitats (intertidal and subtidal). where they are

often abundant.

Forty-six genera are currently recognized in the family

Cirolanidae. There has probably been some measure of over-split-

ting, however, and several genera may not be monophyletic. A

careful phylogenetic analysis of the group is needed. A number of

species are troglobitic (19-20 species in North America), others

occur in fresh water, and still others are found in the deep sea. Of the

46 genera. 8 are known from the tropical eastern Pacific at this time.

Biogeography. — The historical biogeography of the eastern Pa-

cific Cirolanidae cannot be adequately addressed until phylogenetic

analyses of this large family and its Pacific genera have been

completed. However, some generalities are apparent from the data

derived from this study. Eighteen species in 8 genera have been

identified from the tropical eastern Pacific zoogeographic region.

Seven of these are newly described in this paper. Of these 18

species, 15 (83%) may be regarded as tropical species; 3 of these

(Emydice caiidala, Natalolana carlenae n. sp,. and Excirolana

braziliensis) extend their ranges .somewhat into the warm temperate

region adjacent to the tropics. The 3 remaining species are Cirolana

liarfordi. a temperate species (eastern Pacific distribution: British

Columbia to central western Baja California (with a single record

from the southern Gulf of California). Cirolana diminitta. a Califor-

nia and Gulf of California species (warm temperate-subtropical),

and Natalolana californiensis. a southern California species for

which we have a single record from the Gulf of California.

Excirolana braziliensis is an essentially tropical species in the

western Atlantic. Caribbean, and eastern Pacific, but in the last

region it extends its range south to central Chile. Four species

(22%) are amphi-American in distribution (Anopsilana browni.

R. C. Brusca. R. Wetzer. and S. C. France

Cirolana parva, Excirolana bntzjliensis. and E. inaycma). One spe-

cies is endemic to the Galapagos Islands (Metaciwlana calypso n.

sp.), and one is endemic to the Gulf of California {Cimlaiui

nielbrucei n. sp.).

Compared to the Caribbean cirolanid fauna, which contains

approximately 36 species in 16 genera (Kensley and Schotte 1989).

that of the tropical eastern Pacific is relatively depauperate. All

tropical eastern Pacific genera except Coiiileni also occur m the

Caribbean. The biogeographic history of Conilera is enigmatic, as

its only two undisputed species are currently known from the

northern Atlantic (C. cylindracea) and Ecuador (C hiillisi n. sp.).

That the eastern Pacific isopod fauna has strong ties to the Carib-

bean fauna is evinced by the number of aniphi-American species (4

species of Cirolanidae; at least 14 isopod species altogether) and the

large number of probable sister species (i. e.. geminate species),

such as OncilDiplieiis slehhiiifii (Caribbean) and O. jenyhcinuiidi

n. sp. (Pacific). Two species marginally occuning in the tropical

eastern Pacific. CiroUiiui harfonli and Ncilaroltiiui caUforniensis.

are clearly northern Pacific temperate intruders in the tropics.

Although some Cirolanidae occur in deep water, and others are

fresh water or cave dwellers, the majority of species in the family

inhabit shallow marine waters, living from the littoral region to

depths shallower than 500 m. Of the eastern Pacific species, all but

four {Nalatolana califoniiensis, N. carlenae n. sp., Oncdorpheits

jerryhcinicinU n. sp., and Melacirokinu calypso n. sp.) are littoral or

shallow-water forms, and six may be regarded essentially littoral

(rarely occumng in the sublitloral region): Cirolana haifoixli. C.

namelesseiisis n. sp., Metacirolana coslaricensis, Excirolana

hraziliciisis, E. chamensis. and E. inayana.

World list of genera. —

1. A/»i/)!« Budde-Lund, 1908.

2. Anopsilana Paulian and Deboutteville, 1956.

3. Antrohina Bowman. 1964.

4. /ln(/7«/(/»;(( Botosaneanu and Stock, 1979.

5. Atarbohma "Qmce and ]iX\td, 1987.

6. 6a/w/fl»!fl Carpenter. 1981.

7. fi(j%/a/i(; Kensley. 1989.

8. B((//;\7i«/»i»i Milne Edwards, 1879.

9. Booralana Bruce. 1986.

10. Calypiolana Bruce. 1985.

11. Cartetohma Btuce. 1981.

12. Ceratolana Bowman. 1977.

13. C(m/(j/i(/ Leach, 1818.

14. Ow/rt/i/Jo- Benedict. 1896.

15. Cotop/jr/;»i Richardson, 1902.

16. Co«(7<?ra Leach, 1818.

17. Con/7or/j/)ei« Stebbing, 1905.

18. CreaseriellaRm]&, 1953.

19. Dolicholana Bruce. 1986.

20. £Hnc/;c<' Leach, 1815.

21. &/n7«(i(( Jansen, 1981.

22. £vc/(«/<»i(/ Richardson, 1912.

23. Faucheria Dollfus and Vire, 1905.

24. G/;<:;;/!(>/((/ia Barnard, 1920.

25. Hansenolana Siebh'\r\%, 1900.

26. Haptolana Bowman, 1966.

27. L/»Hco/rt«fl Bruce, 1986.

28. Metacirolana YMi'iokin. 1979.

29. Mexiluna Bowman, 1975.

30. Nalatolana Bruce, 1981.

31. Neocirolana Hd]e. 1925.

32. Oncdorpheits Paul and Menzies, 1971.

33. Orphelana Bruce, 1981.

34. Parabalhynomus Barnard, 1924.

35. Po/i;o/am/ Bruce. 1981.

36. Pontogelos Slebh'mg. 1910.

37. Pseudaega'X'\\orr\sor\, 1884,

38. Pseiidolana Bruce, 1979.

39. Saharolana Monod. 1930.

40. Skolobaena Fenara and Monod, 1 972.

41. 5|)e()(;7>()/<»i(/ Bolivar and Pieltain, 1950.

42. 5/)/((/<'/'()/((/k( Cole and Minckley, 1970.

43. 5/)/w('(Y)/»/(/('.v Dollfus. 1897.

44. 7i/n(»/(;/!(; Argano and Pesce, 1980.

45. Txphlocirolana Racovilza, 1905.

46. Xyto/«/(n Kensley, 1987.

Key to the Genera of Cirolanidae Known from the Eastern Pacific

1 . Body elongate (length more than 4 times width); exopod of

pleopod 1 indurate and operculiform to pleopods 2-5 2

— Body not elongate (length 2.0-3.5 times width); exopod of

pleopod 1 not indurate and operculiform to pleopods 2-5 3

2. Endopods of pleopods 2-5 without PMS; maxillary lobes re-

duced and fused into single piece; first article of anlennule not

articulated at right angle to rest of antennule; outer (lateral)

angle of uropodal peduncle not produced, apex rounded (not

acute) Oncdorpheus

— Endopod of pleopod 5 only without PMS; maxillary lobes not

reduced and fused, medial and lateral lobes distinct (outer lobe

bifurcate); antennule article 2 arising at right angle to article 1;

outer (lateral) angle of uropodal peduncle produced, with acute

apex, almost as long as inner (medial) angle Conilera

3. Antennule article 1 longer than articles 2 or 3; antennule article

2 arising at right angle to article 1; maxillipedal endite barely

reaching (or extending barely beyond) first palp article;

maxillipedal endite without coupling spines; frontal lamina nar-

row, anterior (apex) freely projecting; ciypeus forming a ven-

trally projecting triangular blade; antenna with 4 peduncular

articles; antenna long, extending beyond pereonite VII; lateral

margins of pleonite 5 not encompassed by pleonite 4 Eurydice

— Antennule article 2 or 3 longest; antennule article 2 not arising at

right angle to article 1; maxillipedal endite extending well be-

yond first palp article, usually to distal margin of 2nd palp

article; maxillipedal endite with coupling spines; frontal lamina

variable; elypeus variable; antenna with 4 or 5 peduncular ar-

ticles; antenna length variable; lateral margins of pleonite 5 may

or may not be encompassed by pleonite 4 4

4. Antennule peduncle article 2 or 3 longest; ciypeus projecting

ventrally 5

— Antennule peduncle article 3 always longer than 1 or 2; ciypeus

short, broad, fiat, and sessile, not projecting ventrally 6

5. Mandibular incisor usually with accessory tooth on left man-

dible; rostrum large and usually apically spatulate, separating

antennules and confluent with (or fused to) frontal lamina;

frontal lamina long and narrow; pleotelson with a pair of shal-

low submedian pits; pleopod endopods 2-5 or 3-5 without

marginal PMS; medial angle of uropod only weakly produced;

uropodal endopod outer (lateral) margin with a small pit or

notch Excirolana

— Mandibular incisor usually with accessory tooth on right man-

dible; rostrum small to moderate, never spatulate and never

separating antennules; frontal lamina broad, apex dilated and

projecting; pleotelson without a pair of shallow submedian pits;

pleopod endopod 5 only lacking PMS; medial angle of uropod

strongly produced into an acute process; uropodal endopod

without a pit or notch on the outer (lateral) margin

Metacirolana

6. Pereopods V-VII with basis markedly flattened and provided

with long setae, pereopod VII with medial row of long setae on

Tropical Ea.slern Pacific Cirolanidae

posterior siirt'uce; Irontal lamina long and narrow, length 3— (

times width: male appendix maseulina arising basally

or submedially on endopod: pleopod endopod 3 only without

PMS Nciliitolciiui

— Pereopods V-VII not markedly flattened and bearing long setae;

frontal lamina rectangular, pentangular, or round, length no

more than 2 times width; male appendix maseulina always

arising basally on endopod; pleopods 3-5, or only 5. without

PMS 7

7, Pleopods 3-5 without PMS on endopods; posterolateral region

of pleonite 3 extends posteriorly to or beyond posterior margin

of pleonite 5; endopods of pleopods 3— f reduced in si/.e (in

comparison to those of Cirolana); frontal lamina pentagonal or

subquadrate, barely longer than broad; short rostrum always

present; brackish or fresh water only Anopsitami

— Only pleopod 5 without PMS on endopods; po.sterolateral re-

gion of pleonite 3 variable; endopods of pleopods 3—4 not much

smaller than exopods; frontal lamina subquadrate, round, or

pentagonal, usually distinctly longer than broad; with or without

a short rostrum; marine, brackish, or fresh water Cirolana

Aiwpsilana Paulian and Deboutteville, 1956

Type species. — Anopsilana poissoni Paulian and Deboutteville.

1956. by monotypy. Type material at the Institut Scientifique de

Madagascar. Tsimbazaza. Antananarivo. Madagascar.

Synonymy. — Emended and subsequent to Bruce ( 1992: 1 ).

Aiwpsilana. Monod 1976: 135. Bowman and Fran/ 19S2: 522.

Bolosaneanu el af 1986: 412. Bowman and Iliffe 1987: .^47.

Description. — Body 2.5-3.0 times longer than wide. Eyes

moderate in size; troglobitic species usually blind. Frontal mar-

gin of cephalon produced into a minute or short acute rostrum;

cephalon weakly to strongly immersed in pereonite I. Frontal

lamina subquadrate. pentagonal, or subpentagonal/ovate. as long

as broad or longer than broad, apex projecting or sessile, rounded

or truncate; clypeus tlal (sessile), short, and broad; labruni equal

to or narrower than clypeus. Antennular peduncle 3- or 2-articu-

late (i. e., articles 1 and 2 often fused); article 3 longest, about 5

times as long as wide; article 2 not arising at right angle to article

I; flagellum short but not compressed. Antennal peduncle 5-

articulale; articles 1-3 short and subequal in length, articles 4-5

elongate (length greater than 2.0 times width). 5 longest; flagel-

lum very setose. Mandible with broad tridentate incisor, but

middle cusp on left mandible usually reduced; palp extends about

to apex of incisor, its middle article longest, middle and distal

articles with setae; spine row forms a well-developed rounded

lobe with short simple spines. Maxillule medial lobe with 3 stout

circumplumose spines and I or 2 small simple setae; lateral lobe

gnathal surface with 10-14 stout spines. Maxilla medial lobe

short and truncate, usually with 2 long circumplumose setae and

numerous shorter plumose setae and/or simple setae.

Maxillipedal palp 5-articulate. article 3 wider than 4; endite with

2 or 3 coupling spines and long plumose setae.

Pereonite 1 longest, usually twice as long as pereonite II; coxal

plates 11-Vll becoming larger posteriorly, posterolateral angles in-

creasingly acute posteriorly. Pereopods increasing in length posteri-

orly; pereopods V-VII spinose. with bases not markedly flattened.

Penial papillae (penes) absent or minute.

Pleon of 5 free pleonites; pleonite 1 entirely or largely covered

by pereonite Vll; posterolateral region of pleonite 3 extends poste-

riorly to or beyond posterior margin of pleonite 5; pleonite 4 lateral

margins rounded, completely encompassing lateral margins of

pleonite 5. Peduncles of pleopods 1-5 wider than long, lateral

margins without accessory lobes; endopods always slightly smaller

than exopods. especially on pleopods 3-5. Pleopod 1 not opercu-

late; appendix maseulina stout, arising basally or subbasally on

endopod of pleopod 2. Pleopodal endopods 1-2 with PMS. 3-5

without PMS; endopods of pleopods 3-5 smaller than exopods.

Proximomedial angle of pleopodal endopod 5 produced and

lobelike; peduncle of pleopod 5 small, without coupling spines or

plumose setae on medial margin. Pleotelson apex acute or subacute,

never indented. Uropodal peduncle medial angle strongly produced

and acute; medial margin of exopod with numerous simple spines

and PMS.

Remarks. — Bruce (1986a. 1992) synonymized Trof^locirolana

and Haitilana with .\n(>psilana. treated the Australian species, and

reviewed the genus. Anopsilana is distinguished from the closely

related genus Cirolana by its different pleopod morphology (.see

key). The 15 species in the genus inhabit brackish, anchialine. and

fresh water, including 7 species known from caves, wells, and other

subterranean habitats. A number of authors have stated that

Anopsilana species lack penes; however, penes are present, though

small, in mature males of at least A. oa.xaca and A. hrowni. Females

tend to be larger and less ornamented than males. Three species

occur in the tropical eastern Pacific region. Anopsilana oa.xaca. A.

hrowni. and A. aleci n. sp. In all the species of Anopsilana we have

examined, a stout simple seta occurs on the proximolateral margin

of the exopod on pleopod 1. The genus appears to lack any unique

synapomorphies, suggesting it may not be a monophyletic group.

World list of species. —

1. A. acanthiiro {Nolenhoom. 1981). Haiti.

2. A. aleci n. sp. Miraflores Locks. Panama Canal.

3. A. harnardi Bruce. 1992. Queensland. Australia.

4. A. hrowni (Van Name. 1936). Cuba, Belize, and Pacific Costa

Rica.

5. A. conditoria Bruce and Iliffe. 1993 |but dated 1992]. Philip-

pines (from an anchialine cave pool).

6. A. crenala Bowman and Franz, 1982. Grand Cayman Island.

7. A. cuhensis (Hay. 1903). Cuba.

8. A.jonesi Kensley. 1987. Belize.

9. A. lingita Bowman and Iliffe. 1987. Palau (from a cave pool).

10. A. htciae (Barnard. 1940). South Africa.

11. A. oa.xaca Carvacho and Haasmann. 1984. Oaxaca. Mexico,

and Clipperton Island.

12. A. poissoni Paulian and Deboutteville, 1956. Madagascar.

13. A. pustidosa (Hale. 1925). Australia. East Africa. Mozam-

bique. Madagascar. Aldabra. India, and Papua New Guinea.

14. A. radicicola (Notenboom. 1981 ). Haiti.

15. A. u77/tv/(Stebbing. 1904). East Africa to Australia, including

Sri Lanka and India.

Key to Tropical Eastern Pacific Anopsilana Species

1. Left maxilliped with ,^ coupling spines: pleotelson with 10-14 apical

spines; pleopod I peduncle with 3 coupling spines: uropodal exopod's

lateral (outer) margin with 9-11 spines; uropodal endopod with 6-8

spines on lateral (outer) margin A. oa.xacu

— Left maxilliped with 2 coupling spines: pleotelson with 8 apical spines:

pleopod 1 peduncle with 4-5 coupling spines: uropodal exopod with 0

or 8 spines on lateral (outer) margin; uropodal endopod with 2 spines on

lateral (outer) margin : 2

2. Dorsum of pereon ornamented with submedial tubercles: pleotelson

with median longitudinal ridge: uropodal rami extend to or slightly

beyond pleotelson apex: pleonite 1 entirely hidden by pereoniie VII;

antennae reach pereonite IV; pereopod VII ischium's inferior

margin rugose; uropodal exopod with 5 spines on medial (inner)

margin A. bnnvni

— Dorsum of pereon smooth, not ornamented; pleotelson without medial

ridge; uropodal rami much longer than pleotelson: pleonite I's lateral

margins visible in dorsal aspect: antennae reach pereonite II or III:

pereopod VII ischium not rugose: uropodal exopod with 3 spines on

medial (inner) margin A. aleci n. sp.

R. C. Brusca, R. Wetzer, and S. C. France

Anopsiluna aleci n. sp.

Figs. 3B, 4, 5

Type material examined.— MaXe holotype (USNM 252744) and

3 paratypes (1 male. 2 females) (SDNHM Cat. No. 2222): Panama.

Panama Canal, Miratlores Locks, upper east chamber, formalin

wash of sticks, rocks, etc., on bottom of chamber; 17 Jan. 1971;

coll. P W. Glynn.

Di'scriphiiii (ifiinile. — Cephalon devoid of tubercles (Fig. 3B).

Antennules short, extended just past posterior margin of cephalon;

bases separated by rostrum, flagellum of 10 to 1 2 articles (Fig. 4A).

Antennae extend to posterior margin of pereonite 11; flagellum of

about 25 articles (Fig. 4B). Frontal lamina subrectangular, anterior

margin rounded; labium slightly narrower than clypeus (Fig. 4C).

Mar^dibular spine row with 6 stout simple spines; molar process

with about 19 small acute spines; proximal palp article with 1

Figure 3. Amipsiluiui of the tropical eastern Pacific, dorsal views: A, A. browni

(SDNHM A.(X)07), male. B,A. aleci n. sp. (USNM 252744), holotype, male. C, A. oaxaca

(USNM 102239), male. D, A. oaxaca (SIO C3826), female.

Tropical Eastern PacifiL' Cirolanidae

simple seta; middle palp aiticle with 4 plumose and about 1 1 sitnple

setae; distal artiele with 2 comb and about II simple setae (Fig.

4D). Maxillule medial lobe with I short simple spine and inner

protuberance; lateral lobe with about 12 stout apical spines, many

with short barbs, and about 6 minute subapical spines (Fig. 4E).

Maxilla medial lobe with 2 long circumplumose setae, 4 shorter

plumose setae, and about 8 simple setae; lateral lobes with about 9

and 4 simple setae, respectively; I simple seta lies proximal to base

of lateral lobes, and simple setae lie on their lateral margin (Fig.

4F). Maxillipedal palp with simple setae and comb .setae as figured;

endite short, with 2 coupling spines and about 4 apical plumose

setae on both left and right endilcs (Fig. 4G).

Pereon devoid of dorsal tubercles or setae. Posterior angles of

coxae VII very acute and produced past anterior margin of pleonite

.■?. Coxae IIl-VII visible in dorsal view (Fig. 3B). Pereopod I short;

inferior distal angle of i.schium with 1 simple seta; inferior margin

of carpus with about 7 short blunt simple spines and simple setae;

inferior distal angle of carpus with I spine and I seta; inferior

margin of propodus with 3 spines and a cluster of distal setae;

dactylus with small setae at base of unguis (Fig. 5A). Pereopod IV

Figure 4. Anopsikma aleci n. sp. (USNM 252744), hololype, male. A, amennule (left). B, antenna (left). C, frontal lamina,

clypeus, and labrum. D, mandible (right). E, maxillule (right). F, maxilla (right). G, maxilliped (right).

10

R. C. Brusca, R. Wetzer, and S. C. France

Figure 5. Anopsilana aleci n. sp. (USNM 252744), holotype, male. A, pereopod 1 (right). B, pereopod IV (right). C, pereopod VII (right). D, ventral view

of uropod (right). E, pleopod I (right). F, pleopod 2 (right). G, pleopod 3 (right). H, pleopod 4 (right). I, pleopod 5 (nght).

Tropical Easlem Pacific Cirolanidae

long. ambLiUuory. v\ ith setae and simple and seixate Indent spines as

figured (Fig. 5B). Pereopod VII long, ambulatory, with many simple

and trident spines as figured (Fig. 3C). Penes absent or at least not

observed.

Pleon devoid of dorsal tubercles or setae. Pleopodal rami v\ ith

PMS as figured (Figs. 5E-I). Pleopod 1: peduncle's medial margin

with 1 plumose seta, 4 coupling spines, and numerous simple setae;

lateral margin with 1 large seta and numerous simple setae; exopod

l.S times width of endopod; exopod with I simple seta on

proximolateral margin (Fig. 5E). Pleopod 2: peduncle's medial

margin with 3 coupling spines. 4 plumose setae, and many simple

setae; lateral margin with 1 simple spine and many simple setae;

exopod 1.4 times width of endopod; appendix niasculina just

reaches or extends barely beyond tip of exopod. apex nanow with

small subapical lobe (Fig. 5F). Pleopod 3: peduncle's medial mar-

gin with 3 coupling spines. 3 plumose setae, and many simple setae;

lateral margin with I plumose seta and many simple setae; exopod

1.4 times width of endopod. with short incisions on lateral and

medial margins (Fig. ."^G). Pleopod 4: peduncle's medial margin

with 3 coupling spines. 2 plumose setae, and many simple setae;

lateral margin with 1 plumose seta and many simple setae; exopod

1.5 times endopod width, with short incisions on medial and lateral

margins (Fig. 5H). Pleopod 5; peduncle with 1 plumose seta on

lateral margin; exopod 1.4 times width of endopod. with incisions

on lateral and medial margins (Fig. 51).

Pleotelson subtriangular, slightly sinuate near base, with PMS

and 8 apical spines, without dorsal tubercles (Fig. 3B). Uropods

much longer than pleotelson and taper distally; each ramus with

small apical notch containing long simple setae. Uropodal exopod

0.4 times width of endopod and slightly shorter; lateral margin with

8 simple spines and PMS. medial margin with 3 large simple spines

and PMS. Uropodal endopod: medial margin with 8 large simple

spines and PMS; lateral margin with 2 simple spines and PMS.

Uropodal peduncle with PMS on medial margin and 1 simple spine

on lateral margin. 2 large ventral spines and 1 simple seta near base

of exopod (Fig. 5D).

Female. — Similar to male.

Size. — Maximum length to 1 1.5 mm.

Distribution. — Known only from the Miraflores Locks of the

Panama Canal. It is unlikely that this species occurs only in the

Miraflores Locks, a small and continually disturbed habitat. Future

collecting efforts may find it upstream, in the Gatun Lake region.

Remarks. — The long distally tapering uropods of this species

are similar to, though even longer than, those of Anopsilana hrowni.

Anopsilami aleci differs from A. browni in lacking the dorsal and

pereopodal ornamentation of the latter. A. aleci differs from A.

oaxaca in many characters, including the length and spination of

the uropods and pleotelson and the shape of the frontal lamina.

Et\molog\. — This is species is named for the senior author's

son. Alec, a consummate fisherman who, in his youth, provided

companionship on countless collecting trips to the shores of the Sea

of Cortez.

Anopsilana browni (Van Name, 1936)

Figs. 3A, 6-8

Cirolana browni Van Name, 1936: 423. Bruce 1985: 714;

1986a: 219. Kensley and Schotte 1989: 125.

Type material e.xamined. — ( I ) Male holotype and female allo-

type (AMNH 6519): Cuba, Santa Clara Province. Danny's River. 3

mi. below Rodas; June 1918; coll. B. Brown. (2) 6 paratypes

(AMNH 6536): Same data as holotype.

Other material examined. — Eastern Pacific specimens: (3l

Costa Rica. Puntarenas Province, Gulf of Nicoya. Punta Morales.

from burrows of Sphaeroma peruvianum (Isopoda: Sphaero-

malidae) in roots of Rlii-opliora iiumi;le (red mangrove); SDNHM

A.()()()5. 8 Sept. 1987. 1 specimen, and SDNHM A.OOOT. 16 Dec.

1987; I specimen; both coll. W. S/elistowski; the latter apparently

with 2 Sphaeroma in buiTow.

Description of male (based on Facijic specimens). — Cephalon

only moderately wide; anterior margin steeply inclined; a large,

low. medial bifurcate tubercle and a medial transverse incision

between eyes; rostrum minute. Eyes subquadrate in outline, sepa-

rated by nearly 3 times their diameter. Antennules short, extended

to middle of pereonite I. with simple and palmate setae and

aesthetascs as figured; peduncular article 3 twice as long as article

2; tlagellum of 10-16 articles (Fig. 7A). Antennae long, reaching

posterior margin of pereonite 111 or IV; nagellum of 30-36 articles

with proximal articles shorter than distal articles; proximal articles

with very long setae as figured (Fig. 7B). Frontal lamina pentago-

nal; labrum as wide as clypeus; anterior margin of clypeus concave,

posterior margin of labrum broadly concave (Fig. 6A). Mandibular

spine row with about 9 simple spines; molar process with about 26

small acute spines; middle palp article with 4 long serrate setae and

numerous shorter serrate and simple setae; distal palp article with

about 8 simple and 2 plumose setae as figured; right incisor strongly

tridentate. middle cusp of left incisor reduced to a wavy edge (Fig.

6B). Maxillule medial lobe with protuberance on lateral margin and

2 small simple setae; lateral lobe with about 10 long simple spines

(Fig. 6C). Maxilla medial lobe with about 4 long simple setae and

15 circumplumose setae, 2 most posterior circumplumose setae

very long; lateral lobes with about 5 and 14 long simple setae,

respectively (Fig. 6D). Left and right maxillipedal endites with 2

and 3 coupling spines, respectively, and 4 long plumose setae (only

3 figured, one broken off); palp articles with long simple setae,

apical article also with comb setae (Fig. 6E).

Body stout, strongly convex, broadest at pereonites III-IV.

Pereonites II-VII subequal in length. Pereon with low tubercles; a

row of 4 small medial transverse tubercles ornaments anterior third

of pereonite L becoming smaller laterally and tapering off into a

faint transverse ridge; a row of 4— 6 medial, posterior, submarginal

tubercles occurs on pereonites III-VI, being largest medially and

increasing in size on posterior pereonites. Coxae IV-VII visible in

dorsal view. Pereopod I short; inferior margin of merus with 5 large

molariform spines and a few simple setae, distal superior margin

with simple setae; inferior margin of carpus with I long simple seta;

propodus with 2 small distal spines and simple setae; dactylus with

several simple setae at base of unguis (Fig. 7C). Pereopod IV long

and ambulatory, with many simple spines as figured; in addition,

carpus has I large distal serrate tridentate seta, and propodus has I

palmate seta (Fig. 7D). Pereopod VII long and ambulatory, with

many simple spines as figured; inferior margin of ischium with

rugose surface sculpturing (Fig. 7E). Penes short and small, set

somewhat apart near posterior margin of sternite VII (Fig. 7F).

Pleon with minute dorsal tubercles, forming a longitudinal ca-

rina on the pleotelson; pleonite 3 widest, laterally encompassing

pleonite 4. Pleonite I almost entirely covered by pereonite VII (Fig.

3A). Pleopodal rami with PMS as figured (Figs. 8A-E). Pleopod I:

peduncle's medial margin with 5 coupling spines; exopod 1.3 times

width of endopod; exopod with short simple seta on proximolateral

margin (Fig. 8A). Pleopod 2: peduncle's medial margin with 4

coupling spines. 4 plumose setae, and 3 simple setae; exopod 1.4

times width of endopod; appendix niasculina narrows to acute apex.

I.I times exopod length (Fig. 8B). Pleopod 3: peduncle's medial

margin with 3 coupling spines and 4 plumose setae; exopod 1 .4

times width of endopod. with short incisions on lateral and medial

margins (Fig. 8C). Pleopod 4: peduncle with 3 coupling spines. 2

plumose setae, and 4 simple setae; exopod 1.3 times width of

endopod. with short incisions on medial and lateral margins (Fig.

8D). Pleopod 5: peduncle with I long simple seta on lateral margin;

12

R. C. Brusca, R. Wetzer, and S. C. France

exopod 1 .2 times width of endopod. with short incisions on medial

and lateral margins (Fig. 8E).

Pleotelson subtriangular with 8 apical spines and PMS. lacking

tubercles but ornamented with narrow v-shaped ridge, center of "v"

depressed slightly lower than rest of pleotelson (Fig. 3A). Uropodal

endopod and exopod with spines and PMS; lateral margin of exopod

without spines, medial margin with 5 spines, apex without notch

but with long, simple setae; lateral margin of endopod with 2

spines, medial maigin with 8 spines, apex with small apical notch

containing long simple setae (Fig. 8F).

Female. — Similar to male, but with reduced surface sculpturing

and with dorsal tubercles confined primarily to posterior pereonites.

Size. — Maximum length to 11.1 mm.

Distribution. — Anopsilana browiii is an amphi-American spe-

cies. It has been collected in fresh water (a river in Santa Clara

Province, Cuba), brackish water (Sittee River and Salt Creek, Stann

Creek District, Belize), and low-salinity mangrove habitats (Gulf of

Nicoya. Pacific Costa Rica).

Remarks. — The dorsal sculpturing of specimens of A. hrowni

from Costa Rica is less prominent than thai of the type series, but

otherwise there appear to be no differences. Aside from brief men-

tions by Bruce (1985. 1986a) and Kensley and Schotte (1989). there

are no reports of this species since Van Name's original description.

The two specimens we acquired for the present study both came

from bunows of Sphaeroma peniviaiuim. a common sphaeromatid

borer in tropical eastern Pacific red mangrove trees (Peiry 1988,

Peny and Brusca 1989). This is the first record of cohabitation by

these two species.

Anopsikma oa.xaca Carvacho and Haasman, 1 984

Figs. 3C, D. 9. 10

Anopsikma oaxaca Carvacho and Haasman. 1984: 16. Bruce 1986a:

219.

Material examined. — ( 1 ) Mexico, Guerrero, near Acapulco, La-

guna Pie de la Cuesta (LagunaCoyuca). on beach of more westerly

of two granitic islands; Sta. H46-249. LACM; 16 Sept. 1946; coll.

C. Hubbs; 7 specimens. Clipperton Island (eastern Pacific), ""fresh-

water lagoon" samples: (2)offCalix; USNMAcc. No. 213947. Cat.

No. 102239; 1936; coll. C. Limbaugh; 14 specimens. (3) I0°18' N,

1 09 ' 1 3' W. 0- 1 m in lagoon; Sta. 3. Ace. No. B180-3. SIO Cat. No.

C4799; 17 Mar. 1980; coll. J. Cousteau and S. Luke; approx. 30

specimens. (4) 10°18' N, 109°I3' W. reef fiat at N. end. intertidal;

Sta. 2, Ace. No. BI80-2. SIO Cat. No. C4793: 15 Mar. 1980; coll. J.

Cousteau and S. Luke; I specimen. (5) SIO Cat. No. C3826, USNM

Ace. No. 213947; Oct. 1956: coll. C. Limbaugh; 9 specimens. (6)

Sta. 277, USNM Ace. No. 230730; Sept. 1958; coll. C. F Harbison;

approx. 145 specimens. (7) Sta. 225, USNM Ace. No. 230730;

Sept. 1958; coll. C. F Harbison; approx. 113 specimens. (8) Sta.

113. USNM Ace. No. 230730; Sept. 1958; coll. C. F Harbison;

approx. 56 specimens.

Description of male. — Anterior margin of cephalon medially

produced into a short rostrum that folds ventraily to meet frontal

lamina, appearing truncate in dorsal view; cephalon devoid of

tubercles (Figs. 3C, D). Antennules short, extended to middle of

pereonite 1, bases separated by rostrum, flagellum of 7 to 10 articles

(Fig. 9A). Antennae extended to middle of pereonite II, flagellum

Figure 6. Anopsikma browni (SDNHM A.()007), male. A, frontal lamina, clypeus, and lahrum. B, mandible (left). C, maxillule (left), D, maxilla (left). E,

maxilliped (left).

Tropical Easlcrn Pacific Cirolanidae

13

of 13 to 19 articles (Fig. 9B). Frontal lamina Mihiiuadratc; kibruni

and cIvpcLis Mibec|iial in length and width (Fig. 9C). Mainlibular

spine row with 13 short simple spines; molar process with about 15

small acute spines and many short simple setae; middle palp article

with 4 seiTate trident setae and about 10 simple setae; distal article

with about 11 simple setae and 1 comb seta (Fig. 9D). Maxilkile

medial lobe with I short simple seta; lateral lobe with about 1 1 stout

simple spines, many barbed (Fig. 9E). Maxilla medial lobe with 2

large circumplumose setae, about 10 simple setae, and numerous

short subapical hairs; lateral lobes with about 1 1 and 4 simple setae.

respecti\'ely; simple subapical seta proximal to base of lateral lobes

(Fig. 9F). Maxillipedal palp margins with simple setae as figured;

lateral margin of basis with 3 plumose setae, lateral margin of

middle article w ith I comb seta; endite short, with 3 coupling spines

and 2 apical plumose setae (Fig. 9G).

Pereon devoid oi tubercles or carinae (Figs. 3C. D). Posterior

angles of coxa VII very acute, produced past anterior margin of

pleonite 2; coxae III-VIl to V-VII visible in dorsal view. Pereopod

I short; superior margin of ischium with 2 simple setae; inferior

margin of merus with 5 stout, somewhat blunt spines and I simple

seta; inferior margin of carpus with 2 simple setae and I simple

spine; inferior margin of propodus with 3 simple spines and 1 or 2

simple setae; dactylus with I small spine and several simple setae at

base of unguis (Fig. IDA). Pereopod IV long and ambulatory, with

many simple spines as figured (Fig. lOB). Pereopod VII long and

ambulatory, with many simple and senate trident spines as figured

(Fig. IOC). Penes short and squat, set close together on posterior

margin of sternite VII (Fig. lOD).

Pleon devoid of dorsal tubercles or carinae. Pleonite 3 encom-

passes pleonite 4 laterally (Figs. 3C, D). Pleopodal rami with

Figure l.AnopsiUmu hrowni (SDNHM A. 0007). male. A. antennulc (left). B, antenna (left). C. pereopod 1 (left). D. pereopod IV (left). E, pereopod VII

(left). F. penes.

14

R. C. Brusca, R. Wetzer, and S. C. France

PMS as figured (Figs. lOF-J). Pleopod I; peduncle's medial

margin with 3 coupling spines and 2 plumose setae, lateral mar-

gin with I small simple seta; exopod 1.8 times width of endopod;

exopod with a stout 1 simple seta on proximolateral margin (Fig.

lOF). Pleopod 2: peduncle's medial margin with 3 coupling

spines and 4 plumose setae, lateral margin with 1 small simple

spine; exopod 1.6 times as wide as endopod; appendix masculina

narrows to acute apex with small barbs. 1.3 times exopod length

Figure S.Anopsilana browni (SDNHM A.0007). male. A, pleopod I (left). B. pleopod 2 (left). C. pleopod 3 (left). D, pleopod 4 (left). E. pleopod 5 (left).

F, ventral view of uropod (left).

Tropical Eastern Pacific Cirolanidae

15

(Fig. lOG). Pleopod 3: peduncle's medial margin with 3 coupling

spines and 4 plumose setae, lateral margin with I simple seta;

exopod 1.5 times as wide as endopod. with short incisions on

lateral and medial margins (Fig. lOH). Pleopod 4: peduncle's

medial margin with .^ coupling spines and 4 plumose setae, lateral

margin with 1 simple seta; exopod 1.6 tiiues as wide as endopod.

with short incisions on medial and lateral margins (Fig. 101).

Pleopod 5: peduncle with 1 long distally plumose seta on lateral

margin; exopod 1.4 times width of endopod, with short incisions

on medial and lateral margins (Fig. lOJ).

Pleotelson subtriangular, lateral margins convex, narrowing to

apex; apex with 10-14 marginal simple spines, interspersed with

simple setae; dorsum devoid of tubercles or carinae (Figs. 3C, D).

Uropods extend beyond pleotelson apex, distally acute; both rami

with small apical notch containing long simple setae. Uropodal

endopod and exopod with spines and PMS; apex of each ramus with

bundle of simple setae set in notch. Uropodal exopod slightly

shorter and 0.3 width of endopod ( at widest point of each); exopod's

medial margin with 3 or 4 stout spines and PMS, lateral margin with

9-11 stout spines, decreasing in size toward base. Uropodal

endopod's medial inargin with 6 or 7 stout spines interspersed with

PMS, lateral margin with 6-8 stout spines and PMS. Uropodal

peduncle's lateral margin with 1 small simple spine, medial margin

with PMS (Fig. lOE).

Female. — Similar to male, body generally shorter and slightly

wider

Size. — Maximum length 1 1. .3 mm.

Distiihulion. — A shallow-water species found in brackish inter-

tidal and freshwater habitats; so far known from the states of

Oaxaca and Guerrero, Mexico, and from Clipperton Island.

Remarks. — Anopsiiaua oa.xava has been found in brackish man-

grove and lagoon habitats in Mexico and in the "freshwater" lagoon

inside Clipperton Island, an atoll. In larger individuals, particularly

males, the rostral process of the cephalon is slightly larger or more

produced (note the difference in Figs. 3C, D). Carvacho and

Haasmann (1984) stated that males lack penes; however, short

squat penes are present in the adult male specimens we have exam-

ined.

The type locality oi Anopsilana oa.xaca is Manialtepec Lagoon,

1 .3 km west of Puerto Escondido, Oaxaca, Mexico. The holotype

and allotype are apparently deposited in the collections of the

Institute of Biology, Universidad Nacional Autonoma de Mexico

(UN AM). Paratypes are located in the reference collection of

Centro de Investigacion Cienti'fica y Educacion Superior de

Ensenada (CICESE), Baja California, Mexico, and in the Museum

National d'Histoire Naturelle, Paris. Several attempts to borrow

type material from UNAM and CICESE met with no success.

Figure 9. Anupsikma ocuacci (SIO C.^826), non-oostegial female. A, antennule. B,

antenna. C, frontal lamina, clypeus, and labrum D, mandible (left). E, maxillule. F, maxilla.

G, maxilliped.

16

R. C. Brusca, R. Wetzer, and S. C. France

Figure 10. Anopsilaiia ocixaca {USNM 102239). male. A. pereopod 1. B, pereopod IV. C, pereopod VII. D, penes. E, dorsal view of uropod; note detail

of endopod's medial margin. F. pleopod 1. G, pleopod 2. H, pleopod 3. 1, pleopod 4. J. pleopod 5.

Tropical Eastern Pacific Cirolanidae

17

Cirolaiui Leach, 1818

Type species. — Cirolana craiuiiii Leach, 1818. hy riioiiolypy.

Type material at BMNH. .See Bruce and Elhs (I98,V) for a rede-

scription of Ciioliiiin cnincliii.

Syiuiiiymy. — Emended and subsequent to Bruce ( l986a:L^9).

Cimlaiw. Sars 1897: 69. Moore 1902: 166. Brusca and Ninos 197S;

379. Brusca 1980: 228. Holdich et al. 1981: 5?7. Bruce and Bowman 1982:

i25. Brusca and Iverson 1985: .15. Delaney 1986: 731. Kensley and .Scholle

1987:227.

Description. — Body 2.5-3,0 times longer than broad, smooth or

variously ornamented with tubercles, ridges, carinae, or grooves.

Eyes small to moderate sized. Cephalon with rostrum absent or

present as a small median point, which may e.\tend ventrally to

overlap frontal lamina; posterior region of cephalon often with

partial incision lines on lateral margins. Frontal lamina flat,

subquadrate to about twice as long as broad, but approaching round

or pentagonal In many species; sessile or with anterior margin

freely projecting; clypeus flat (sessile), short, and wider than long

[Bruce ( 198 la) noted that the clypeus of Cirolana furcata "has two

projecting lobes"]; labrum flat, as narrow or narrower than clypeus.

Antennular peduncle usually 3-articulate. often 2-articulate by fu-

sion of first 2 articles; articles I and 2 do not form right angles to

one another (as in Eurydice): article 3 longest; relative fiagellum

length varies with age and body length in many species. Antennal

peduncle ."i-articulate; articles 1-3 short. 4 and 5 long; articles 4 and

5 combined usually longer than 1-3 combined. Mandible with

broad, tridentate incisor, middle cusp often reduced, especially on

left mandible; spine row a well-developed rounded lobe with many

small simple spines and simple setae; palp 3-articulate, middle

article longest, middle and distal articles with setae. Maxillule

medial lobe slender, with 3 stout circumplumose spines and some-

times a single minute simple seta; lateral lobe with 10-14 stout

simple spines, spines often barbed. Ma.xilla medial lobe short and

truncate, typically with a few large circumplumose setae and many

simple and plumose setae; lateral lobe bifurcate, with many apical

simple setae. Ma.\illiped broad, palp 5-articulate, article wider than

others; endite often with distal plumose and/or simple setae, usually

with 2 coupling spines (rarely 1 or 3).

Pereonite 1 usually 1.4-1.5 times longer than pereonite II. Coxae

II-VII increasing in size posteriorly, becoming larger and often

more acute posteriorly. Pereopods generally spinose on inferior and

distal superior margins; inferior margin of merus of pereopod 1 with

several blunt spines; pereopods I-III grasping, shorter than IV-VII.

with inferior margins of merus and ischium not produced or only

modestly produced; pereopods IV-VII long and ambulatory, ar-

ticles neither markedly flattened nor expanded; all pereopodal

dactyli with small, blunt, stout spine at base of unguis. Penes small

and knoblike or moderate in size.

Pleon of 5 pleonites. Pleonite I often concealed or partly con-

cealed by pereonite VII; pleonite 5 narrower than 1-4, and with

lateral margins covered by pleonite 4. Peduncles of pleopods 1-5

broader than long, with coupling spines and plumose setae on

medial margin, often without lateral lobes; rami similar, not mark-

edly elongate, endopods usually shorter and narrower than exopods,

and with fewer PMS. Pleopods I and 2 similar to one another;

exopod of pleopod I of many species with a prominent simple seta

on proximolateral margin; appendix masculina arising basally from

endopod of male pleopod 2. Pleopods 3-5 often with partial or

complete incisions across exopods. Endopod of pleopod 5 with

proximomedial angle produced, lobelike, without PMS; peduncle's

medial margin without plumose setae or coupling spines. Pleotelson

with or without dorsal tubercles or carinae; apex subacute, rounded

or truncate, rarely incised or excavated; usually with marginal

spines and PMS. Uropods with inner angle of peduncle strongly

produced and acute, apex with or without marginal spines but

iilways with PMS; uropodal rami with or without apical notches;

lateral and medial inargins with or without .spines and PMS.

Remarlis. — The genus Cirolana was the second genus described

in what is now the family Cirolanidae. Although with 84 species it

is still the largest genus in the family, many species were transferred

to six other genera by Bruce (1981a): .Anopsilana. Curtetolana.

Melacirotana. NalaUilana. NeociroUma. and Piilitolana.

Bruce ( 1986a) stated that Auopsilana and NeociroUma arc the

genera most closely related to Cirolana. Indeed, these 3 genera are

extremely similar. So far as we are aware. .-Xnop.silana clearly

differs from Cirolana only in lacking PMS on the endopods of

pleopods 3 and 4. Neocirolana differs from Cirolana in its reduced

mandibular inci.sor and the occasional reduction of the maxilla (the

degree of reduction in both mouthparts varies from species to

species).

Cirolana is worldwide in distribution, though most species have

been described from tropical and subtropical regions, at depths

ranging from the intertidal zone to about 200 m. There are several

deep-water species, including one (Cfornicala) from about 2000 m.

Five species occur in the tropical eastern Pacific; haifordi,

iliniinuta. parva. nielhnicei n. sp.. and namelessensis n. sp.

Eiirylana arcuata (Hale, 1925), a Southern Hemisphere species

introduced into San Francisco Bay, California (Bowman et al.

1981). was until recently placed in Cirolana but was removed to

Eiirylana by Jansen ( 1 98 1 ).

In this study, careful uropod and pleotelson spine counts were

made, using both transmitted and reflected light. An effort was

made to determine whether setae and spines were broken off or

missing by examining the margin of the pleotelson or uropod

cuticle for sockets or cuticular fragments. We believe that the

ranges of spine counts we specify for the various Cirolana species

in this study indicate consistent species-specific characters; hence

they figure prominently in species discrimination and the key.

World lisl of .specie.^. —

C. alhida Richardson. 1901. Florida.

C albidoida Kensley and Schotte. 1987. The Bahamas.

C arafiirae Bruce. 1 986. Kai Islands. Indonesia.

C. argentina Giambiagi. 1931 [but dated 1930] (should per-

haps be transferred to E.xcirolana). Argentina.

C. avida Nunomura. 1988. Japan.

C aiistraliensis Hale. 1925. South Australia.

C hisiilcaia Hobbins and Jones. 1993. Red Sea.

C. hoiigaardti Kensley. 1984. South Africa.

C hovina Barnard. 1940. South and East Africa to India.

C. hroclia Bruce, 1986. Queensland, Australia.

C canaliculata Tattersall, 1 92 1 . Off New Zealand.

C capricornica Bruce, 1986. Queensland. Australia.

C. carina Jonea. 1976. Kenya.

C chaloti Bouvier. 1901. Zaire.

C. cingulata Barnard. 1920 South Africa.

C. coonia Bruce. 1986. Queensland. Australia.

C. coronala Bruce and Jones. 1981. Japan.

C corriigi.s Jones. 1976. Kenya and Red Sea.

C cranchii Leach, 1818. Atlantic Ocean and Mediterranean

Sea.

C. cremditelson Kensley and Schotte. 1987. Belize.

C. curtensis Bruce. 1986. Queensland. Australia.

C. diminuta Menzies. 1962. Southern California to Galapagos

Islands.

C epinierias Richardson. 1910. Philippines.

r. erodiae Bruce. 1986. Queensland. Australia.

C.Jhiviatili.s Stebbing, 1902. South Africa.

C. fornicata (Mezhov, 1981 ). North-central Pacific Ocean.

C furcata Bruce. 1981. New South Wales, Australia.

1.

~t

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13.

14.

15.

16.

17.

18.

19.

20.

21.

22.

23.

24.

25.

26,

27.

R. C. Brusca. R. Wetzer. and S. C. France

28.

29.

30.

31.

32.

33.

35.

37.

38.

39.

40.

41.

42.

43.

44.

45.

46.

47.

48.

49.

50.

51.

52.

53.

54.

55.

56.

57.

58.

59.

60.

61.

62.

63.

64.

65.

66.

67.

68.

69.

70.

71.

72.

73.

74.

75.

76.

77.

78.

C. garuwa Bruce. 1986. Western Australia.

C. luilei Bruce, 1981. New South Wales, Australia.

C. harfordi (Lockington. 1877). British Columbia to noilh-

western Me,\ico; also Australia, Hong Kong, and Japan (prob-

ably introduced).

C. hesperia Bruce, 1986. Western Australia.

C. iinposha Barnard, 1955. South Africa.

C. impmceros Bruce. 1986. Queensland, Australia.

C iiwisicaiuhi Barnard, 1940. South Africa.

C. indica Nierstrasz, 1931. Indonesia.

C. kendi Bruce. 1986. Queensland, Australia.

C koinbona Bruce, 1986. Queensland, Australia.

C. lata Haswell, 1881 [Bruce (1986a) synonymized C. lata

var. Integra Miers, 1884, with Cartetolana liiieala Potts, 1915,

as Cartetolana intregra (Miers, 1884)|. New South Wales,

Australia.

C. latistxlis Dana, 1 853 (species inqiilrenda. see Bruce 1986a).

Indo-Pacific.

C. Ilgnicola Nunomura, 1984. East China Sea.

C. Iltloralls Barnard, 1920. South Africa.

C. magdalalna Bruce, 1980. Australian Coral Sea.

C. manorae Bruce and Javed, 1987. Northern Indian Ocean.

C. niascarenensls MuWer, 1991. Reunion Island. Indian Ocean.

C. nielnerti Barnard, 1920. South Africa.

C. meklsla Bruce, 1986. Queensland, Australia.

C. meseda Hobbins and Jones, 1993. Red Sea.

C. nilniila Hansen, 1890. West Indies.

r. morilla Bruce, 1986. Queensland, Australia.

C. namelessensis n. sp. Galapagos Islands. Ecuador.

C. nielhnicel n. sp. Gulf of California. Mexico.

C oblnmcata Richardson, 1901. Caribbean.

C oreonota Bruce, 1986. Torres Strait, Australia.

C. palifrons Barnard, 1920. South Africa.

C. paraerodlae Miiller and Salvat, 1993. French Polynesia.

C. parva Hansen, 1890. Tropical amphi-Anierican.

C. perlata Barnard, 1936. Burma.

C. plconastica Stebbing, 1900. Solomon Islands, West Pacific.

C. porcellana Barnard, ]9?ib{lncertae sedls. see Bruce 1986a).

Burma.

C. portula Bruce, 1986. Victoria, Australia.

C quadrlpustiilata Hurley, 1957. (incertae sedls. see Bruce

1986a). New Zealand.

C. rugicaiida Heller. 1861. South Africa and southern Indian

Ocean.

C. sadoensis Nunomura. 1981. Japan.

C. suldanhae Barnard, 195 1 . South Africa.

C. schioedtel Miers, 1884 (Incertae sedls. see Bruce 1986a).

Australia and Arafura Sea.

C. sinillls Bruce, 1981. New South Wales, Australia.

C. solltaria Bruce, 1 986. New South Wales, Australia.

C. sp. Bruce, 1986. {Incertae sedls, see Bruce 1986a). Austra-

lia.

C. slebbingi Nierstrasz, 1931 (incertae sedls. see Bruce

1986a). Indonesia.

C. stenoiira Bruce, 1986. Lizard Island, Australia.

C. sulcata Hansen, 1890. South Africa.

C. sulcaticaiida Stebbing, 1904. East Africa to India and Sri

Lanka.

C. theleceps Barnard, 1940. South Africa and Red Sea.

C. iranscoslata Barnard, 1959. South Africa.

C. triloba Bruce, 1981. Australia.

C. tuberciilata (Richardson. 1910). Philippines.

C. tuberculosa Bruce, 1986. Queensland, Australia.

C. tiimidosa Holdich, Harrison, and Bruce, 1981. Queensland.

Australia.

79. C. undiilata Barnard, 1914. South Africa.

80. C. iirostxlls Menzies, 1962 {incertae sedls. see Bruce 1986a).

Chile.

81. C. vanhoffeni Nierslrasz, 1931. Indonesia.

82. C. i'('/i((.vr/(Y(;((/(( Stebbing, 1902. South Africa.

83. r. viclna Barnard. 1914. South Africa.

84. C. victorlae Bruce. 1981 . Victoria, Australia.

Key to Tropical Eastern Pacific Cirolana Species

1 . Uropodal rami without apical notch; pleotelson margin with 8-

36 spines (in large males, pleotelson with 2 large dorsal tu-

bercles and pleonite 5 with 2 mediodorsal tubercles); merus of

pereopod I with 6 stout blunt spines on inferior margin; rostrum

meets but does not overlap frontal lamina; mandibular palp

article 3 with distinctly excavated or scoop-shaped comb setae;

antennular peduncle articles 1 and 2 free, not fused C. harfordi

— Both uropodal rami with apical notch; pleotelson margin with

6-10 spines (with or without dorsal tubercles); merus of pereo-

pod 1 usually with fewer than 6 stout blunt spines on inferior

margin; rostrum overlaps frontal lamina; mandibular palp ar-

ticle 3 with simple comb setae; antennular peduncle articles 1

and 2 entirely or largely fused 2

2. Pleonites 3-5 each with a pair of submedian dorsal tubercles;

pleotelson with 2 submedian rows of tubercles; labrum approxi-

mately 3 times as long as clypeus C. namelessensis n. sp.

— Pleonites 3-5 without submedian dorsal tubercles; pleotelson

smooth, devoid of tubercles; labrum and clypeus subequal in

length 3

3. Pleotelson margin with 6-7 spines; inferior margin of propodus

of pereopod 1 tuberculate; pleopod 1 with lateral margin of

endopod strongly concave; pleopod 2 endopod markedly shorter

than exopod C. nlclbnicel n. sp.

— Pleotelson margin with 8-9 spines; inferior margin of propodus

of pereopod 1 not tuberculate; pleopod 1 with lateral margin of

endopod not concave or only weakly concave; pleopod 2

endopod and exopod subequal in length 4

4. Maxillipedal endite reaches only to first palp article; appendix

masculina acute, often with filamentous apex; penes moderate

in size, set close together in line with medial margins of pleopod

I peduncle C. pana

— Maxillipedal endite reaches second palp article; appendix

masculina bluntly round to subacute; penes small, set apart in

line with middle of pleopod 1 peduncle C. dliiiiniita

Cirolana dimimtta Menzies,

Figs. IIA.B, 13-17

962

Cirolana diminiita Menzies, 1962b: 343. Schultz 1969: 184.

Bowman 1977: 653 (as C. par\'a). Brusca 1980: 228. Bruce and

Bowman 1982: 327. Brusca and Iverson 1985: 35. Stepien and

Brusca 1985: 91. Bruce 1986a: 220.

Type material examined. — (1) Holotype (USNM 108647):

Mexico, Baja California, San Quintin Bay, on gray sandy silt with

shell fragments and clay, depth less than 7 m; Sta. SQ 13, USNM

Ace. No. 245146; 22 Apr. 1960; coll. R. J. Menzies; I postnianca.

Paratypes (all from San Quintin Bay; coll. R. J. Menzies; 1960-

1961 ): (2) Sta. SQ 181, intertidal; USNM No. 109271, USNM Ace.

No. 245146; 10 specimens. (3) Sta. SQ 175, intertidal; USNM

109271, USNM Ace. No. 245146; 40 specimens. (4) Sta. SQ 177,

intertidal; USNM 109265, USNM Ace. No. 245146; 1 specimen.

(5) Sta. SQ 28, from dark gray silt and worm tubes, at 6-8 m; 23

Apr, 1960; USNM 109269; 2 specimens. (6) SQ Hubbs trawl;

USNM 109264, USNM Ace. No. 245146; 2 specimens. (7) Sta. SQ

179, intertidal: USNM 109270, USNM Ace. No. 245146; 3 speci-

mens. (8) Sta. SQ 178, intertidal; USNM 109268, USNM Ace. No.

Tropical Easlem Pacific Cirolanidae

Figure 1 1. Cirolana of the tropical eastern Pacific, dorsal views: A, C. dimimila (USNM 108647), holotype, poslmanca. B, C iliimniita (AHF, Sta. SQ

181), paratype, male. C, C. harfordi (BMNH 1878:9), probable syntype, male. D, C. harfordi (SIO CI856), male.

20

R. C. Brusca, R. Wetzer, and S. C. France

Figure 12. Ciwlana of the tropical eastern Pacific, dorsal views, continued; A, C namelessensis n. sp. (USNM 232734), holutype, male. B, C.

nielbnicei n. sp. (LACM). paratype. male. C, C panxi (ZMUC), syntype, male.

245146: 10 specimens. (9) (LACM/AHF) Sta. SQ 181, intertidal;

Menzies Cat. No. 67; 30+ specimens, (10) USNM 109266: inter-

tidal; 2 specimens.

Other imiterial examined. — California specimens; (11) Santa Bar-

bara Co., Point Conception Light. 34° N, 120° W; RA' Veleru /V'Sta,

No. 4822-57, AHF Cat. No. 548-1; 17 Jan. 1957; 3 specimens; (12)

Santa Barbara Co., Santa Cruz Island, Smuggler's Cove, night light:

RA' Vc/ew /V Sta. No. 166()-48, AHFCat. No. 517-2; 29 Dec. 1948;

1 male; (13) Los Angeles Co., Santa Catalina Island, Famsworth

Bank; AHF Ace. No. ^1971-3; 1-2 June 1970; coll. C. Gage and K.

Hooker; 10 specimens: (14) San Diego Co., LaJolla, from kelp bed at

16m;SIOC3821;2Dec.l958andl3Mar. 1959: 25-1- ,specimens;(15)

San Diego Co., La Jolla, from kelp bed: AHF 957- 1 ; 8 specimens; (16)

San Diego Co., off Point Loma, from kelp holdfast, ca. 80 feet:

SDNHM; 9 Sept. 1990: coll. C. Gramlich; 20-i- specimens.

Pacific Baja California specimens: (17) Cedros Island, south

bay, reef and tidepool:R/V VWt'/o /V Sta. 2035-5 1 : 19 Apr. 1951; 1

specimen; (18) Scammon's Lagoon, from rock at 5-8 m; Sta. KG-3:

13 Sept. 1953; coll. "K. G."; 1 male; ( 19) Thurloe Bay. off Thurloe

Point. 18 m; Sta. 283. USNM Ace. No. 128938: 9 Mar. 1934: 2

specimens; (20) Abrejos Point: AHF Cat. No. 777-1; 2 damaged

specimens; (21) Abrejos Point Estero, mid-intertidal of mangrove

lagoon, in filamentous green algae; Sta. "ECCE" MEPA-16C, SIO

C1782, SIO Ace, No. BI 175-29; 26 July 1975: coll. Hemingway

and Luke: 1 specimen; (22) San Ignacio Lagoon. North Whale

Island, 3.25-3.75 m; Sta. H50-71. AHF 956-1; 11 Feb. 1950; coll.

Hubbs. Johnson, and Allanson: 2 males; (23) San Ignacio Lagoon;

AHF 956-1; 2 males: (24) San Ignacio Lagoon, rocky intertidal.

26°45' N. 11 3° 12' W: SIO Ace. No. Bl 76-5. SIO C2477; Mar.

1976; coll. S. Luke: 1 female and 1 juvenile.

Gulf of California specimens (all from Baja California Sur):

(25) Espiritu Santo Island, San Lorenzo Channel, from corallines at

43 m; USNM Ace. No. 1 39772. R/V Velen, 111 Sta. 607-36; 2 1 Mar.

1936; 1 specimen; (26) Espiritu Santo Island, S. San Lorenzo

Channel, from coralline algae at 9-27 m; R/V Vetera III Sta. 498-36,

USNM Ace. No, 139772; 19 Feb. 1936; 2 females: (27) Espiiitu

Santo Island, off Ballenas Bay, 14 m: #643, USNM Ace. No.

144492; 8 Mar. 1937; 1 female; (28) El Bajo. near Loreto. "from

formalin wash of Halimeda sp.": EWI-2: 20 Aug. 1980, LACM:

coll. E. W. Iverson: 1 specimen.

Description of male. — Cephalon without tubercles, with or

without interocular furrow (Figs. IIA, B). Small rostral process

folding ventrally to separate antennules and partly overlap frontal

lamina(Figs. 13A,B,C. I4C). Antennules short, reaching just to. or

beyond, posterior margin of cephalon: peduncle with simple and

palmate setae and groups of minute spinelets. peduncular articles I

and 2 fused: flagellum of 9-12 articles, distal articles with

aesthetascs (Fig. 14A). Antennae reaching posterior margin of

pereonite III or IV. with simple and palmate setae as figured:

tlagellum of 16-29 articles (Fig. I4B). Frontal lamina short and

pentagonal (Figs. 13A, B. 14C). Mandibular spine row forms large

lobe with comblike row of 12 long fleshy spines and 2 small simple

spines above and 6 long simple spines below fleshy spines: molar

process with about 24 small acute marginal spines and many small

simple setae on posterior margin; middle article of mandibular palp

with serrate and simple setae, distal article with about 22 setae of

which 18 are simple and the 4 most distal are comb (Fig. 14D).

Maxillule medial lobe as figured: lateral lobe with about 12 stout

apical spines, the 5 largest spines armed with small barbs (Fig.

14E). Maxilla medial lobe with about 8 plumose setae and 9 simple

setae: lateral lobes with about 14 and 5 simple setae, respectively

(Fig. I4F). Maxillipedal endite very short, with 1 or 2 coupling

spines, small proximal lobe, and about 5 distal circumplumose

setae; palp with simple and comb setae as figured (Fig, 14G).

Tropical Eastern Pacific Cirolanidae

21

Figure 13. Cirotana ciiniinuki (AHF, Sta. SQ 181). paratype. male, scanning electron micrograpiis: A, frontal lamina, clypeus, and labrum. 120x. B,

rostrum and frontal lamina (note overlap of frontal lamina by rostrum), 240x. C, distal margin of rostrum, I800x. D, seta in pit on distal margin of rostrum,

9000X.

22

R. C. Brusca. R. Wetzer. and S. C. France

Figure 14. Cirotana Jimifiiiki ( AHF, Sla. SQ 1811. paratype. male: A, antennule. B, antenna. C, frontal lamina, clypeus, and

labrum. D. mandible. E. niaxillule. F, maxilla. G. maxilliped (right).

Pereon widest at pereonites IV and V (Fig. 1 1 A, B). Coxae V-

VII vi.sible dorsally. Pereon without dorsal tubercles, carinae. or

setae. Pereopod I stout and short, with simple and stout setae;

inferior margin of merus with 5 short, blunt spines and several

simple setae (Figs. 15A, I6B); dactylus with 1 small spine and 1

simple seta at ba.se of unguis (Figs. 15A, I6A). Pereopod IV with

many large spines as figured; dactylus with I small spine and 1

simple seta at base of unguis (Fig. I.'^B). Pereopod VII long, with

many large spines as figured (Fig. 15C, I6C); ischium-propodus

with long serrate spines and stout, basally denticulate spines, as

well as simple spines; dactylus with 1 small spine and 1 long simple

seta at base of unguis (Fig. ISC). Penes short, very small, set

soinewhat apart and in line with middle of peduncles of first pleo-

pods (near posterior margin of sternite VII) (Fig. I5D).

Lateral margins of pleonites 2^ expanded ventrolaterally;

pleon widest at pleonite 2, devoid of dorsal tubercles, carinae, or

setae (Figs. IIA, B). Pleopodal rami with PMS as figured (Figs.

15F-J, I6D). Pleopod I: peduncle's inedial margin with 5 long

Tropical Eastern Pacific Ciroianiiiae

23

Figure 1 5. CiroUma iliminuui {AHF. Sta. SQ 1 8 1 ), paratype, male: A, pereopod I. B, pereopod IV. C, pereopod VII. D, penes. E. uropod. F, pleopod

1. G, pleopod 2. H. pleopod 3. I. pleopod 4. J, pleopod 5.

24

R. C. Brusca. R. Wetzer, and S. C. France

Figure 16. Cirohma diminiita (AHF, Sta. SQ 181), paratype. male, scanning electron micrographs; A, distal portion of pereopod I, 240x. B. marginal

spines on merus of pereopod I, 1 20x. C, ventral view of spines on posterior pereopods, 60x. D, ventral view of PMS on pleopod 1 , 240x.

Tropical haslcni I'acilic Cirolanidae

25

Figure 17. Ciwlana diminuui {AHP. Sla. SQ 181), paratype, male, scanning electron micrographs: A, pleotelson (note straight lateral margins and dorsal

setae), 60x. B, dorsal setae of pleotelson (note scalelike appearance of cuticle in background), 3000x. C, terminal notch on uropodal endopod, 420x. D.

ventral view of pleotelson and uropods. 60x.

26

R. C. Brusca, R. Welzer, and S. C. France

coupling spines, lateral margin with 1 small simple seta; endopod

0.6 times as wide as exopod, lateral margin slightly concave:

exopod with I stout spine on proximolateral margin (Fig. 15F).

Pleopod 2: peduncle's medial margin with 4-5 coupling spines and

6 plumose setae, lateral margin with 1 simple spine; appendix

masculina 1 .3 times asl long as exopod. apex broadly rounded or

occasionally subacute, but never with threadlike extension; appen-

dix masculina with short stout distal setae on margins and occasion-

ally on medial surface as well (Fig. I5G). Pleopod .3: peduncle's

medial margin with 5 coupling spines and 6 plumose setae, lateral

margin with 1 simple spine; exopod with complete or partial mar-

ginal incisions (Fig. I5H). Pleopod 4: peduncle's medial margin

with 4 coupling spines and 6 plumose setae, lateral margin with 1

simple spine; exopod with incomplete marginal incisions (Fig. 151).

Pleopod 5: peduncle with 1 simple spine on lateral margin; exopod

with incomplete marginal incisions (Fig. I5J).

Pleotelson triangular, narrowing strongly near rounded apex,

lateral margins straight or slightly concave (Fig. 17A); apex usually

with 8 stout spines, occasionally 9, interspersed with numerous

PMS (Fig. 1 1 A); dorsum with scattered simple setae (Figs. 1 7 A, B).

Uropodal exopod extends beyond pleotelson (Fig. 17D), not

rounded distally as in C. harfoidi. and more narrow than in C.

harfordi; both rami with large apical notch, long slender setae

arising from each notch (Fig. 17C). Both uropodal rami with spines

and PMS on medial and lateral margins. Uropodal exopod 0.5 as

wide as endopod and slightly shorter; medial margin with 3 stout

spines, lateral margin with 7 or 8 stout spines interspersed with

numerous PMS. Uropodal endopod's medial margin with 5-7 stout

spines; lateral margin with 2^ stout spines (Fig. 15E). Uropodal

peduncle's medial margin with PMS; lateral margin with I medial

spine and a group of spines and setae near the articulation of the

exopod (Fig. 15E).

Female. — Similar to male.

Ultrastnictiiral features. — When viewed with SEM the dorsal

surface of this species, like that of many other isopods, has a

somewhat imbricated, "shingle roof appearance due to scalelike

cuticular structures, which are also present on the mouthparts.

frontal lamina, clypeus. and labrum. In addition, the dorsal surface

of the cephalon, pereon, pleon, and pleotelson have many setae

arising from small cuticular pits (Figs. 17A, B). The distal part of

the cephalic rostrum has peglike cuticular structures or setae arising

from pits along the margin (Figs. 13C, D).

Size. — To maximum length of 10.5 mm.

Di.snibiition. — Cirolana diminuta ranges from southern

California (Point Conception, Santa Barbara Co.) south along the

west coast of Baja California and into the central and southern Gulf

of California. It is a littoral and shallow subtidal species, occuixing

in deeper water (to 43 m) in the warm southern extent of its range.

RciiuuLs. — Menzies' ( 1962b) described C. diminuta as the most

abundant and widespread isopod in San Quinti'n Bay (the type

locality). His designated holotype specimen is a postmanca (noted in

his text as "young") and is undissected, so the original illustrations

and descriptions of mouthparts must have been from a paratype

specimen; there are no descriptions of the pereopods and pleopods in

the original text. Cirolana diminuta and C. pan'a Hansen, 1890, are

very similar and difficult to distinguish. Menzies and Glynn ( 1968)

placed Cirolana diniinula as a junior synonym of C. parva. stating

that the species was circumtropical in distribution. Bruce and Bow-

man (1982) removed C dimi)iuia from synonymy with C. pan'a

without describing precisely how the two could be differentiated.

They stated only that the species "can be separated by the differences

in the characters figured" and by the "vasa deferentia open[ing] flush

with the surface of sternite 7" in C. diminuta. None of Bruce and

Bowman's figures provides reliable means of distinguishing these

two species, and the vasa deferentia do not open flush with sternite in

either species but exit through distinct penes.

In both C. diminuta and C. pan-a length of the antennae, relative

to the body, decreases with increasing body length. In most adults

the antennae extend to pereonite III or IV, although in small speci-

mens of C. parva they may extend to pereonite V. In both species

the margins of the pleotelson may be straight or slightly convex,

although C diminuta is straight most of the time and C pan-a is

convex most of the time. In both species the exopod of pleopod 3

may be completely or incompletely divided by lateral sutures, and

in both species the endopodal lateral margin of pleopod 1 is slightly

concave. A consistent feature distinguishing these two species is the

position of the penes. In C. panxi the penes are moderate in size and

set very close together in a line even with the medial margins of the

peduncles of the first pleopods; in C diminuta the penes are very

small and set farther apart, in line with the middle of the peduncles

of the first pleopods. Also, the apex of the appendix masculina of C.

parra is always acute and often tapers to a filamentous or threadlike

extension; in C. diminuta the apex of the appendix masculina varies

from bluntly rounded to subacute but never has a threadlike or

filamentous projection. In both species short stout distal spines may

occur on either margin (and more rarely across the medial surface)

of the appendix masculina.

Cirolana diminuta belongs to the C. pana complex of sibling

species .'iensu Bruce and Bowman (1982) and Bruce (1986a).

Stafford ( 1912) described an isopod from sponges at Laguna Beach,

California, as Cirolana haifordi var. spongicola. distinguishing it

primarily by the straight lateral margins of the pleotelson and

serrate uropodal rami. Stafford did not designate type specimens or

their deposition. Her figures and description, including a pleotelson

with straight lateral margins and a count of 8 marginal spines on the

pleotelson apex, suggest that her specimens were C. dimitntta.

However, without any of Stafford's material to examine, we feel it

best not to synonymize C. diminuta with C. harfordi var.

spongicola, thus leaving the latter as a nomen dubium.

Cirolana diminuta is primarily a shallow subtidal species, living

in rocks, sandy silts, and algae to a depth of about 50 m. In some

localities it is common in the littoral zone. This species is a member

of the California demersal zooplankton community. Stepien and

Brusca ( 1985) recently presented experimental evidence of its noc-

turnal emergence to prey on epibenthic fishes.

Cirolana harfordi {Lockington, 1877)

Figs. lie, D, 18-20

Synonymy. — Emended and subsequent to Bruce ( 1986a: 146).

Cirolana harfordi japonica. ^unomma 1985a: 132.

Cirolana haiifordi japonica Nunoniura. 1981 : 46 [lapsus calami).

Cirolana haifordi. Miller 1975: 296. 307. Brusca and Ninos 1978: 383.

Momsetal. 1980: 538. Ricketlsetal. 1985: 49, 229, 533. Bruce 1986b: 549.

Non-Cirolana harfordi: Cirolana harfordi var spongicola Stafford,

1912; probably synonym of C diminuta.

Type material e.xamined. — ( 1 ) Probable syntypes (BMNH Reg.

No. 1878:9): U.S.A., California, Santa Rosa Island, under stones,

"MT," in muddy places; coll. W. G. N. Harford: 2 males (1 dam-

aged).

Other material examined. — California specimens: (2) San

Miguel Island, Harris Point, collected under intertidal rocks;

SDNHM; 15 Apr. 1985: coll. T. D. Stebbins: 1 specimen.

Pacific Baja California specimens: (3) Banda Point (ca. 20 mi.

S. of Ensenada), under rocks, low tide: SDNHM A.0009: 5 Dec.

1987; coll. R. Wetzer and R. C. Brusca; 9 specimens. (4) 2 mi. S. of

Rosarito, shore on rocky reef; RA' Velero /VSta. 1597-47, LACM;

6 Mar. 1947; I male. (5) Descanso Point, 7 mi. N. of Halfway

House, rocky shore; R/V Velero /VSta. 1594-47, LACM; 3 Mar.

1947; 2 specimens. (6) second headland S. of Descanso Point,

Tropical Haslem Pacitic Cirolanidae

27

Figure 18. Cirotana harfordi (SIO C1856). male: A, antennule (left). B. antenna (left). C, frontal

lamina, clypeus, and labrum. D, mandible (left). E, maxillule (left). F, maxilla (left). G, maxilliped

(left).

among algae: LACM; 14 Jan. 1946; coll. E. Y. Dawson; 1 female.

(7) N. of Descanso Point, within sight of S. Coronado Island,

collected on rocky reef, tide -1.2 ft.; RA^ Velero /VSta. 1597-47,

LACM; 6 Mar. 1947; 1 1 specimens. (8) Descanso Point, collected

on rocky reef, tide -1.2 ft.; LACM; 3 Mar. 1947; 2 specimens. (9)

Salsipuedes Point, littoral; SDNHM; 1 Sept. 1976; coll. R. C.

Brusca; 4 specimens. ( 10) Salsipuedes Point, rocky intertidal algae

zone; SDNHM; 1 Sept. 1976; coll. R. C. Brusca; 30+ specimens.

(11 ) mouth of Santo Tomas River between San Jose Point and Santo

Tomas Point, rocky reef, tide -1.4 ft.; RA' Velero /V'Sta. 1596-47,

LACM; 5 Mar. 1947; I adult male and 1 juvenile. (12) San Jose

Point; R/V Velero /V Sla. 1596-47, LACM; I gravid female. (13)

Cape Colonet, rocky shore intertidal, 30°57'00" N, 1I6°17'30" W;

RA' Velero /VSta. 1 504-46, LACM; 3 specimens. (14) San Ramon

Bay, Camalti Point; SDNHM; 19 June 1939; coll. Harbison and

Bilderback; 20-1- specimens. (15) S. shore of San Ramon Bay, on

coast over from Colonia Guerrero, 12 mi. N. of San Quinti'n Bay.

littoral; SDNHM; 4 Aug. 1976; coll. R. C. Brusca and B.

Wallerstein; 5 specimens. ( 16) San Quintin Bay, rocky outcropping

in lagoon; R/V Velero /V Sta. 2013-51. LACM; 14 Apr. 1951; 1

specimen. (17) San Quintin Bay, intertidal algae and rock washes;

SDNHM A.OIOl; II Nov. 1989; coll. R. Wetzer and H. Wetzer; 30-t-

specimens. (18) Guadalupe Island, Melpomene Cove. 28°52'05" N,

1 18°I9'05" W, algae wash; R/V Velero /V'Sta. 1912-49, LACM; 17

Dec. 1949; I specimen. ( 19) Guadalupe Island, Melpomene Cove at

landing, rock-algae wash; R/V Velero /V'Sta. 1915-49, LACM; 18

Dec. 1949; coll. R. J. Menzies and D. Reish; 5 specimens. (20)

Santa Rosalillita Point, 28 4()' N. 1 14' 14' W, rocky intertidal;

"ECCE" SR-34, Ace. No. BI 75-39, SIO CI 856; 6 Aug. 1975; coll.

Luke and Hemingway; 20-h specimens. (21 ) 2 mi. S. E. of Cedros

Island Light, 28°20' N, 1I5°10' W, 99 m; R/V VWe/o /// Sta. 1265-

41, LACM; 28 Feb. 1941; 1 specimen. (22) Cedros Island, S. bay,

shore collecting; Sta. 28834, USNM Ace. No. 128938; 10 Mar.

1934; 6 specimens. (23) W. side of Middle San Benito Island,

28' 18'55" N, 1 15'=34'06" W; R/V Velero IV LACM; 8 May 1950; 1

specitnen. (24) W. side of San Benitos Island, 28° 18' N, 1 15°34' W;

28

R. C. Brusca. R. Wet/.er, and S. C. France

RA' VelemlVSla. 1976-50, LACM; 8 May 1950; 7 specimens. (25)

Tortugas Bay, 27°4r N, 114°53' W. collected on rocky point via

SCUBA, 3-5 m; Sta. "ECCE" BT-23. Ace. No. BI75-21, SIO

C1755; coll. G. Hemingway and S. Luke; 1 specimen. (26) San

Bartolome Point, sulphur rock, 27°37'46" N, 114°52'38" W; R/V

Velem IV Sta. 2605-54, LACM; 12 Feb. 1954; 10 specimens,

including a gravid female. (27) Asuncion Point, 27°08' N. 1 14°17'

W. taken" by surface trawl; RA' Velem /VSta. 1950-50. LACM; 28

Apr. 1950; 5 specimens. (28) Magdalena Bay. Entrada Point. 24°32'

N. I12°04' W. shore; R/V Velero /V Sta. 1961-50. LACM; 2 May

1950; lO-t- specimens. No definite locality specified: (29) USNM

86669, Ace. No. 109185; Feb. 1930; coll. E. F Ricketts; 4 speci-

mens; (30); AHF Ace. No. 1146; Nov. 1947; coll. Dr. Gentry; 1

specimen.

Gulf of California specimens: (31 ) Mexico. Baja California. La

Paz Bay; Dawson Sta. 54, LACM; 10-11 Nov. 1946; coll. E. Y.

Dawson; 1 specimen.

Description of male. — Cephalon devoid of tubercles, often with

interocular fun'ow, length about 2.5 times width (Figs. IIC, D).

Antennule short, reaching just beyond cephalon; flagellum with 8-

12 articles, aesthetascs on most or all articles; middle articles of

tlagellum broader than long (Fig. 18A). Antenna long, reaching to

pereonite IVor V; tlagellum with 30-34 articles (Fig. 18B). Frontal

lamina pentagonal, broader anteriorly, length approximately 1.5

times width (Fig. 18C). Mandibular spine row with 14 stout simple

spines; molar process with about 22 small acute spines and up to 6

long simple setae that arise from proximal region of molar; middle

palp article with 22 senate setae, distal article with 19 comb setae,

each hollowed out or excavated (Fig. 18D). Maxillule lateral lobe

with 12 stout spines, many barbed, and 6 small subapical spines

(Fig. 18E). Maxilla tnedial lobe with 13 plumose setae and 8 simple

setae; lateral lobes with simple and comb setae as figured (Fig.

18F). Maxillipedal endite short, with I coupling spine and 5 plu-

mose setae; palp articles with simple and comb setae (Fig. 18G).

Pereon broadest at pereonites IV-V. Posterior margin of

pereonites with row of minute tubercles in larger specimens. Coxae

Figure 19. Ciioluiui haifordi (SIO C1856), male: A, pereopod 1 (left). B, pereopod IV (left). C.

pereopod VII (left). D, penes. E, apices of uropod and pleotelson.

Tropical Eastern Pacific Cirolanidae

29

Q.

1

•a

a

o

U

O

53

30

R C. Brusca, R. Wetzer, and S. C. France

II-VII visible dorsatly (Figs. 1 IC, D). PereopodI stout, with simple,

stout, and molariforni (squat, truncate) spines; ischium with I

molariform spine and merus with 6 niolariform spines on inferior

margin; merus with 1 very large spine on superior margin; propodus

with 4 blunt spines on inferior margin (Fig. 19A). Pereopods IV-

VII long, with many simple and serrate spines and slout. basally

denticulate spines as figured (Fig. 19B, C). Penes very small,

wartlike processes, set somewhat apart near posterior margin of

sternite VII (Fig. I9D).

Pleonites 1-2 often covered by pereonite VII. Pleonites 3-3

with row of small tubercles on posterior margin, and occasionally a

pair of minute dorsal submedian tubercles on pleonite 5 in larger

specimens (Figs. 1 IC, D). Pleopodal endopods 1-4 with PMS only

apically; exopods 1-5 with PMS entirely around lateral margin and

extending to distal third of ramus on medial margin (Figs. 2()A-E).

Pleopods 1-5: peduncles with 1 large circumpluniose seta on lateral

margin; peduncles of pleopods 4-5 with small lateral lobe. Pleo-

pods 3-5: exopods with ctenate scales on medial side. Pleopod 1:

exopod with 1 large proximolateral spine; lateral (outer) margin of

endopod straight; peduncle with 5 coupling spines. Pleopod 2:

appendix masculina. with distal spinelets, extends well beyond

apex of endopod; peduncle with 4 coupling spines and 5 plumose

setae (Fig. 20B). Exopod of pleopod 3 with marginal incisions

(Figure 20C). Pleopod 4: peduncle with 4 coupling spines and 6

plumose setae; exopod with marginal incisions (Fig. 2()D). Pleopod

5: incisions on exopod nearly complete (Fig. 20E).

Pleotelson subtriangular. apex narrowly rounded, with 9-36

marginal spines, interspersed with PMS; adult males with 2 large

dorsal submedian tubercles or carinae, usually absent in females

(Figs, lie, D). Uropods longer than pleotelson, distally rounded,

without apical notches. Uropodal endopod twice as broad as

exopod, with 12-15 marginal spines, interspersed with PMS; apical

spines large and blunt. Uropodal exopod with 13-17 marginal

spines, interspersed with PMS; apical spines large and blunt (Fig.

19E). PMS of pleotelson and uropods with very short setules.

Fciinilc. — Similar to male, but usually lacking the 2 large dorsal

submedian tubercles or carinae on the pleotelson and the rows of

small tubercles on posterior margins of some pereonites and

pleonites.

Size. — To maximum length of 20.0 mm.

Distrihiition. — C. harfonli is a widely distributed species in the

eastern Pacific. It occurs intertidally at least from Vancouver Island.

British Columbia, south about to Magdalena Bay on the west coast

of Baja California. There is also a single record from the .southern

Gulf of California (La Paz Bay). The species has a discontinuous

distribution in the North Pacific, according to Bruce and Jones

(1981 ) and Bruce ( 1986a). being found on the coasts of the Russian

Far East, Japan, and Australia.

Remarks. — Bruce and Jones (1981) concluded that CiroUma

harfonli jiiptmicaTh\e\emann. 1910, was not sufficiently distinct to

warrant subspecific status. Bruce (1986a) synonymized C.

toyawaensis Nunomura. 1982. and C. ihiehnanni Kussakin. 1979.

with C. harfonli. noting that C. harfonli may have been introduced

to Australia. C. harfonli var. spongicola Stafford. 1912, is probably

conspecific with C. diiniiuaa (see remarks for C. diniiiuita).

Occasional males lack the dorsal tubercles on the pleotelson,

and the dorsal tubercles of pleonite 5 are most commonly seen on

specimens from the California Channel Islands. All females we

have examined lack these tubercles.

Cirolana harfordi is one of the most common littoral isopods in

warm and cool temperate waters in the northeastern Pacific, becom-

ing rare in subtropical waters south of west-central Baja California.

It is common and often abundant beneath rocks (especially on

sandy substrates) and in mussel beds, in the littoral zone and

shallow sublittoral region. It infrequently occurs subtidally to

depths of 135 m. Hewatt (1937) recorded densities as high as

1 1.521 per square meter in Monterey mussel beds. Johnson ( 1974.

I976a.b) reported C. harfordi preying most commonly on small

polychaetes (0. 1-0.5 mm in diameter), said to be consumed whole.

Johnson also found them preying on a variety of small crustaceans,

particularly amphipods and copepods. and also scavenging on dead

animal matter. Johnson's work suggested prey/food location by C.

harfordi is primarily by olfactory means. He also found that

ovigerous females seldom, if ever, leave the safety of their under-

rock shelter or other hiding places to feed, fasting for the entire 3^

month brooding period. Johnson claimed that most females "prob-

ably" died (or were preyed upon) after the first brood, few surviving

to produce a second; laboratory populations, however, survived

through 3 or more broods under "optimal conditions" (Johnson

1976a). Fecundity ranged from 18 to 68 eggs per brood. Johnson

proposed a mean 2-year lifespan for C. haifordi in nature, with

year-round breeding and an average population generation time of

13.9 months.

Lockington ( 1877) did not designate type specimens or indicate

a place of specimen deposition in his original description of

Cirolana harfordi, published in the Proceedings of the California

Academy of Sciences. No type specimens are at the California

Academy of Sciences or at other museums where Lockington's

material was deposited, such as the Academy of Natural Sciences of

Philadelphia and the Museum National d'Hisloire Naturelle, Paris.

The BMNH does house four probable synlype specimens. These

specimens are from the type locality (Santa Rosa Island in the Santa

Barbara Channel. California) and were given to the BMNH by

Lockington shortly after publication of the species. These speci-

mens conform with others we have examined.

Cirolana namelessensis n. sp.

Figs. 1 2A. 21.22

Type material examined. — ( I ) Male holotype (USNM 252734)

and 30 male, female, and manca paratypes (USNM 252735): Ecua-

dor, Galapagos Islands. Nameless Island, intertidal; Sta. 26A,

USNM Ace. No. 330765; 22 Feb. 1978; coll. W. B. Hope.

Additional paratypes. Colombian specimens: (2) Pacific coast,

Ensenada de Utria. in algae, on dead Pocillopora robusta, 5 m;

SDNHM A.0I47; 12 Aug. 1984; coll. G. E. Ramos-Tafur; 1 speci-

men. (3) Malpelo Island, coll. for P W. Glynn; SDNHM A.0I49; 29

Feb. -3 Mar. 1972; 12 specimens. (4) Pacific Colombia. Malpelo

Island, dive #5. N.E. wall. ca. 10 ft.; LACM (00000); 2 Mar 1972;

23 specimens. (5) Malpelo Island, intertidal; SDNHM A.0148; 9

Mar. 1972; 8 specimens. Galapagos specimens: (6) Tower Island,

Darwin Bay, coral gathered at Seal Beach No. 1 by "Carl"; RA'

Velero III Sta. 94-33. USNM; 22 Feb. 1933; 8 postmanca speci-

mens. (7) Albemarle Island. Tagus Cove, coral from north shore by

"Cari"; USNM 152-19.34; 14 Jan. 1934; 1 male and 3 females. (8)

Albemarle Island. Albemarle Point, shore collecting about Pontes

heads, low tide; R/V Velero III Sta. 69-33. USNM Ace. No. 1 22445;

II Feb. 1933; I specimen. (9) Indefatigable Island. Gordon Bay;

Allan Hancock Expedition; R/V Velero III Sta. 1 35-34. USNM Ace.

No. 131571; 8 Dec. 1934; coll. W. L. Schmitt; 3 females. (10)

Charles Island (= Isia Santa Maria), near Post Office Bay. from

coral, inside crater: RA' Velero III Sta. 804-38, USNM Ace. No.

148041; 23 Jan. 1938; 2 males and 3 females. (II) Hood Island,

from rocky spit. Gardiner Bay; R/V Velero III Sta. 27-33. USNM

Ace. No. 122445; 25 Jan. 1933; 2 specimens.

Description of male. — Cephalon devoid of tubercles or carinae;

width 2.0 times length (Fig. I2A). Small rostrum separates anten-

nules. produced ventrally to partly overlap frontal lamina (Fig.

21C). Antennules short, reaching middle of pereonite 1, with

aesthetascs and simple and palmate setae as figured: first 2 articles

Tropical Eastern Pacific Cirolanidae

31

Figure 21. Cirolana namelessensis n. sp. {USNM 2?2734), holotype. male: A, antennule. B, antenna. C,

frontal lamina, clypeus. and labrum. D. mandible. E, maxiilule. F. maxilla. G, maxiliiped.

of peduncle largely fused; flagellum of 10-12 articles (Fig. 21 A).

Antenna long, reaching anterior margin of pereonite V. with simple

and palmate setae as figured; tlagellum of 25-31 articles (Fig.

21B). Frontal lamina pentagonal; labrum much longer, but slightly

narrower than clypeus (Fig. 2IC). Mandibular spine row with II

large simple spines and 1 long seta; molar process with stout spines,

many short setae on posterior margin, and 3 very long setae arising

from proximoanterior region; middle palp article with about 9

comb and 1 1 simple setae, distal article with many simple setae and

3 terminal comb setae (Fig. 2 ID). Maxiilule medial lobe with 3

stout circumplumose spines and 1 small seta; lateral lobe with 1 1

stout and 5 small spines; 3 of the stout spines armed with barbs (Fig.

21E). Maxilla medial lobe (gnathal surface) with 2 fully plumose, 6

distally plumose, and 5 simple setae; lateral lobes with 8 and 5

simple setae, respectively (Fig. 21F). Maxillipedal endite short,

with 2 coupling spines. 2 apical and 1 subapical plumose setae, and

2 simple apical setae; palp margins with simple, plumose, and comb

setae as figured (Fig. 2IG).

Pereon widest at pereonite V. Pereonite VII with row of small

tubercles along posterior margin. Coxae IV-VII visible in dorsal

view; VI-VII with oblique posterior carina; VII produced posteri-

orly to midlength of pleonite II (Fig. 12A). Pereopod I stout, with

simple setae as figured; inferior margin of merus with 5 short, blunt,

molariform spines; propodus' distal inferior margin with 3 very

short blunt molariform spines; dactylus with small spine and setae

set below base of primary unguis and cluster of setae on superior

margin at base of unguis (Fig. 22A). Pereopod IV: margins of

ischium-propodus with large stout spines bearing basal denticles;

dactylus with 1 small spine (Fig. 22B). Pereopod VII long; inferior

and distal margins of ischium-propodus with serrate spines and

stout spines bearing basal denticles; dactylus with I small spine

(Fig. 22C). Penes short, small, set somewhat apart near posterior

margin of sternite VII (Fig. 22D).

Pleon broadest at pleonites 2 and 3. Pleonite 1 almost com-

pletely overlapped by pereonite VII (visible medially). Postero-

lateral angles of pleonite 3 produced, partly encompassing pleonite

32

R. C. Brusca, R. Wetzer, and S. C. France

Figure 22. Cirolana luimelessensis n. sp. (USNM 252734), hololype. male: A, pereopod I. B, pereopod IV. C, pereopod Vll. D, penes. E, dorsal

■iew of uropod. F, pleopod I . G. pleopod 2. H, pleopod 3. 1, pleopod 4. J. pleopod 5.

Tropical Eastern Pacific Cirolanidae

33

4. Pleonites 3-5 with 2 submediun tubercles, becoming larger pos-

teriorly (Fig. I2A). Pleopodal rami with PMS as figured (Figs.

22F-J). Pleopod I: peduncle's medial margin with 4 coupling

spines, lateral margin with I simple spine; endopod 0.5 width of

exopod. lateral (outer) margin concave; e.xopod with single simple

spine on proximolateral margin (Fig. 22F). Pleopod 2: peduncle's

medial margin with 4 coupling spines and 3 plumose setae, lateral

margin with I small spine; appendix niasculina length 1.2 times

length of exopod. narrow ing to acute apex with short barbed setae

(Fig. 22Gl. Pleopod 3: peduncle's medial margin with 3 coupling

spines and 1 plumose seta; lateral margin with I short simple spine;

exopod w ith short marginal incisions and ctenate .scales on medial

margin (Fig. 22H). Pleopod 4: peduncle's medial margin with 3

coupling spines and 4 plumose setae, lateral margin with 1 simple

spine; exopod with short marginal incisions and ctenate scales on

medial margin (Fig. 221). Pleopod 5; exopod with short marginal

incisions and ctenate scales on medial margin (Fig. 22J).

Pleotelson triangular, lateral margins straight, apex rounded,

with 6-9 stout marginal spines interspersed with many simple

setae; posterior pleonites and pleotelson with 2 longitudinal

submedian tuberculate carinae, most prominent in larger individu-

als, occasionally with .several smaller tubercles located between the

2 longitudinal carinae (Fig. I2A). Uropods with scattered dorsal

chromatophores; inner angle of peduncle with distal PMS; rami

with small apical notches and a group of long simple setae arising

from each notch (Fig. 22E); margins of uropodal rami weakly

seiTale and somewhat rounded apically (especially endopod), not

lanceolate as in C. nielbrucei: both rami with spines and PMS on

medial and lateral margins. Uropodal exopod 0.67 times width of

endopod, extending almost to pleotelson apex; medial margin with

3—4 spines, lateral margin with 5-8 spines, interspersed with PMS.

Uropodal endopod extends barely beyond pleotelson apex; medial

margin with 5-8 spines, lateral margin with 2-3 spines, inter-

spersed with PMS.

Figure 23. Cirolana nielbrucei n. sp. (LACM 77-294.2), paratype, male: A, antennule (right). B,

antenna (right). C. frontal lamina, clypeus, and labrum. D, mandible (left). E, maxillule (left). F, maxilla

(left). G, maxilliped (left).

34

R. C. Brusca, R. Wetzer, and S. C. France

Figure 24. Cinilana nielbnicei n. sp. (LACM 77-294.2), paratype, male: A, pereopod I (left). B, pereopod IV (left). C, pereopod VII (left). D.

penes, E, ventral view of uropod (left). F, pleopod I (left). G. pleopod 2 (left). H, pleopod 3 (left). 1, pleopod 4 (left). J, pleopod 5 (left).

Tropical Eastern Pacific Cirolanidae

35

Female. — Similar to male. Dorsal luhcrclcs on pIcon and

pleolelson somewhat variable in size; occasionally larger in the

female than in the male.

Size. — Small, to maximum length of 8.4 mm.

Disirihiilion. — An intertidal-shallow subtidal species prefer-

ring coral rubble, coral, and rocky substrates, and so far known

from only three areas: the Galapagos Islands (Nameless.

Albermarle, Tower. Indefatigable. Hood, and Charles islands);

Malpelo Island. Colombia; and Ensenada de Ulria. Colombia (on

the Pacific coast).

Remarks. — Cimlana luimelessensis has affinities with both the

C. paiTa group of sibling species, sensii Bruce and Bowman ( 1982)

and Bruce ( 1986a). and the "tuberculate" group of C//o/fl/ia species

(Bruce. 1986a). The characters most readily distinguishing

Cimlana namelessensis from similar appearing species in the tropi-

cal eastern Pacific are as follows: the tuberculation of the posterior

pleonites and pleotelson (the only other tropical eastern Pacific

Ciidlaini species with dorsal tuberculation is C. harfonii, which can

immediately be distinguished from C. namelessensis by its lacking

apical notches on the uropodal rami and the shape and number of

spines of the pleotelson and uropods. The apical notches of the

uropodal rami are smaller (not as deep) than in C. parva and C.

diminuta.

Etymology. — Named for the type locality: Nameless Island.

Galapagos Islands. Ecuador.

Cirolana nielhnicei n. sp.

Figs. I2B, 23. 24

Type material e.xamined. — (1) Holotype (LACM 77-294.1).

male, and 45 paratypes. 41 at LACM (77-294.2) and 4 at SDNHM

(A.Ol.'iO): Mexico. Gulf of California. Punta Chivato; transect #1.

quadrat #7. I of 4; 16 July 1977: coll. R. C. Brusca. R. Zimmerman,

and R. Winn.

Additional paratypes: (2) Mexico. Sonora. Tiburon Island; Ace.

No. 159124. USNM 8631.^; 3 Apr. 1940; coll. E. F. Ricketts; 9

specimens. (3) Mexico. Baja California (Gulf). El Bajo, nearLoreto.

formalin washing of shallow subtidal rocks; EWI80-I8. LACM; 20

Aug. 1980; coll. E. W. Iverson; 50-h specimens. (4) Mexico. Baja

California (Gulf); 25°3I' N. 1 1 1°04' W, taken with "Chemfish" and

scuba at 3-5 m; RR65-37. Ace. No. BI-6549. SIO C2444; 1 1 July

1965; coll. R. Rosenblatt; I female. (5) Mexico. Baja California

(Gulf), Espiritu Santo Island, shore collecting; R/V Velero III Sta.

634-37. USNM Ace. No. 144492; 6 Mar. 1937; I male.

Deseriplion of male. — Cephalon devoid of tubercles. 2.2 times

wider than long (Fig. 12B). Rostrum small, barely overlapping

frontal lamina (Fig. 23C). Antennule short, reaching just beyond

anterior margin of pereonite I. with simple and plumose setae and

aesthetascs; flagellum of 8 or 9 articles (Fig. 23 A). Antenna long,

reaching middle of pereonite IV; flagellum of 26-28 articles; pe-

duncular article 4 with characteristic subapical denticle (difficult to

see on small specimens) (Fig. 23B). Frontal lamina pentagonal

(Fig. 23C). Mandibular spine row forms a large round lobe with 13

robust spines. 2 of them very long; molar process with 26 small

acute marginal spines and many short simple setae on posterior

margin; middle article of mandibular palp with 10 plumose setae

(not all figured); distal article with approximately 20 comb setae

(Fig. 23D). Maxillule lateral lobe with 10 stout barbed spines. 1

simple spine, and 4 small slender subapical setae (Fig. 23E). Max-

illa medial lobe with 4 simple setae, 6 plumose setae, and 2 long

circumplumose setae; lateral lobes with 1 1 and 4 simple setae,

respectively; lateral margin of basal article with 3 small setae (Fig.

23F). Maxillipedal endite very short, with 2 coupling spines. 3

plumose, and I simple seta apically; palp margins with simple and

comb setae as figured (Fig. 23G).

Pereon broadest at pereonites V and VI. Pereon dorsum devoid

of tubercles or earinae. Posterior angles of coxae VII very long and

acute, produced past anterior margin of pleonite 2; coxae IV-VII

visible in dorsal view (Fig. I2B). Pereopod I short and stout, with

several simple setae; inferior margin of merus with 5 blunt

molariform spines and 3 simple spines; inferior margin of propodus

tuberculate. with large blunt distal spines and 3 small spines;

dactylus with small blunt apical spine (Fig. 24A). Pereopod IV

ambulatory with numerous large spines on all articles as figured;

dactylus with I seta and I apical spine at base of unguis (Fig. 24B).

Pereopod VII with robust serrate spines and stout, basally denticu-

late spines on all articles as figured; dactylus with I seta and I

apical spine at base of unguis (Fig. 24C). Penes short, small, set

.somewhat apart near posterior margin of sternite VII (Fig. 24D).

Pleonites I^: po.sterolateral angles produced, each partly en-

compassing following pleonite. Pleon without dorsal tubercles or

earinae (Fig. I2B). Pleopodal rami with PMS as figured (Figs. 24F-

J). Pleopod 1 : peduncle's medial margin with 5 coupling spines and

2 plumose setae, lateral margin with I small simple spine; endopod

0.67 width of exopod. lateral (outer) margin concave; exopod with

1 stout seta on proximolateral margin (Fig. 24F). Pleopod 2:

peduncle's medial margin with 5 coupling spines and 4 plumose

setae, lateral margin with I spine: appendix masculina 1 .25 times as

long as exopod, narrowing to acute apex with short barbed setae

(Fig. 24G). Pleopod 3: peduncle's medial margin with 4 coupling

spines and 6 plumose setae; lateral margin with I simple spine;

exopod with short marginal incisions (Fig. 24H). Pleopod 4:

peduncle's medial margin with 4 coupling spines and 5 plumose

setae; lateral margin with a simple spine; exopod with short mar-

ginal incisions (Fig. 241). Pleopod 5: peduncle with 1 simple spine

on lateral margin; exopod with short marginal incisions (Fig. 24J).

Pleotelson triangular, lateral margins straight, not convex as in

C. par\'a\ apex nartow. rounded, with 6-7 stout marginal spines,

interspersed with many simple setae; dorsum without tubercles or

earinae (Fig. 12B). Uropods extend slightly beyond pleotelson

apex, distally acute; margins strongly serrate (much more so than in

C, parva or C. diminuta): large apical notch on both rami, a group of

long simple setae arising from each notch: both rami with large

spines and PMS on medial and lateral margins. Uropodal exopod

0.50 width of endopod and slightly shorter than endopod; medial

margin with 2-3 stout spines, lateral margin with 7-8 stout spines

(medial spines larger than lateral spines), interspersed with many

PMS. Uropodal endopod's medial margin with 4-5 stout spines,

lateral margin with 3 stout spines (lateral spines much longer than

medial spines), interspersed with many PMS. Uropodal peduncle's

medial margin with distal PMS. lateral margin with 3 stout spines,

one subapical and 2 apical (Fig. 24E).

Female. — Similar to male.

Size. — Small, to maximum length of 7.85 mm.

Distribution. — An intertidal and shallow subtidal species, so far

known only from the central and southern Gulf of California.

Remarks — Cirolana nielhnicei belongs to the C. pana group of

sibling species, sensii Bruce and Bowman (1982). and Bruce

(1986a). The characters most readily distinguishing C. nielhnicei

from similar appearing species in the tropical eastern Pacific are as

follows: spination of the pleotelson (always 6 or 7 marginal spines

in C. nielhnicei. never more); shape of pleotelson (lateral margins

straight): shape of uropodal rami (apices very acute, margins very

serrate); presence of a subapical denticle on article 4 of antennal

peduncle; and spination of the uropodal rami (varies only slightly

from C. pana. although the spines on the medial margin of the

endopod are much longer and stouter in C. nielhnicei). Sessile

loricated peritrich ciliates have been observed among the spines on

the medial margin of pereopod VII in some specimens.

Ftymology. — We take pleasure in naming this species after the

36

R. C. Brusca, R. Wetzer. and S. C. France

indefatigable Southern Hemisphere isopodologist, Niel L. Bruce.

Cirolaiui pana Hansen. 1890

Figs. 12C. 25, 26

Synoimnx. — Emended and subsequent to Bruce and Bowman

1982:325.

Menzies and Glynn 1968: 38. Schultz 1969: 185. Brusca 1980: 228.

Brusca and Iverson 1985: 35. Bruce 1985: 714; 1986a: 220; 1986b: 549.

Kensley and Schotte 1989: 135. Scholte et al. 1991: 225.

Non-0>o/a/to pan-a: see Bruce and Bowman 1982: 325,

Type material examined. — (1) "Lectotype" (ZMUC), male.

West Indies. St. Thomas. "Syntypes" (ZMUC): (2) female. West

Indies; (3) West Indies, 5 specimens; (4) West Indies. St. Croix, 2

specimens (in very poor condition); (5) 25°N. 34°E. 2 specimens:

(6) [no locality]. 1 specimen; (7) Pacific Ocean. Sainoa. 1 speci-

men.

Other material examined. — Atlantic specimens. Mexico: (8)

Campeche. 5 mi. N. of Sebaplaya; USL-TFE-I. SDNHM; 6 Jan.

1977; 3 females; (9) Campeche. 5 mi. N. of Sebaplaya; USL-TFE-I.

SDNHM; 6 Jan. 1977; coll. J. Martin; I specimen; (10) Campeche,

10 mi. N.E. of Chompoton, "errant collections off grass beds";

USL-TFE-IIB, SDNHM; 7 Jan. 1978; 1 male and 1 gravid female;

( 1 1) Campeche. 10 mi. N. of Chompoton; USL-TFE-I. SDNHM; 6

Jan. 1977; 1 manca; (12) Quintana Roo, Cozumel Island; USNM

Ace. No. 229190; 8 Apr. I960; 50-K specimens; (13) Quintana Roo,

between Lawrence Point and Fupar Point; USNM Ace. 229190; 5

Apr 1960; 24 specimens. (14) Puerto Rico; Ace. No. 259321,

USNM ll2841;coll. R W. Glynn; 11 specimens. (15) Costa Rica,

Linion Province. Cahuita. infauna of intertidal coral rock, from in

front of "Jenny's house," with 2 pink sphaeromatids; PMD-81-2,

SDNHM; 22 Aug. 1981; coll. P M. Delaney. C. Stepien. P. Pepe; 1

male.

Pacific specimens: (16) Mexico. Oaxaca. Sacrificios Island,

15 = 40' N. 96° 14' W. taken with scuba at 13 m; Ace. No. B 178- 15.

TEPE 78-15. RA' Alpha Helix. SIO C4123; 7 Apr. 1978; coll. W.

Newman and S. Luke; 1 female. Costa Rica: (17) Guanacaste

Province, Salinas Bay. shore collecting on beach beyond reef, on

sand and stone; R/V Velero III Sta. 474-35. USNM Ace. No.

131571; 10 Feb. 1935; 3 females; (18) Puntarenas Province.

Quepos. Cocal Beach, sandy beach, with Exciroluna; USNM

150034; 1 female; ( 19) Cocos Island, around point to northwest of

Chatham Bay. from "roach trap" in morning; Ace. No. 122445,

USNM 68433; 1 Mar. 1933; coll. W. L. Schmitt; 80-1- specimens;

(20) Cocos Island, from "roach trap"; USNM Ace. No. 122445; I

Mar. 1933; 50-1- specimens; (21 ) Cocos Island. Bajo Alcyone. 35 m;

LACM; 27 Mar. 1989; coll. K. Kaiser; 3 specimens; (22) Parker

Bay. off small island at N. shore, shore collecting; R/V Velero HI

Figure 25. Cirolana pur\-a (VSNM Ace. No. 93322), male: A, antennule. B, antenna. C, frontal lamina,

clypeus, and labrum. D, mandible. E, maxillule. F, maxilla. G, maxilliped.

Tropical Eastern Pacific Cirolanidae

37

Sta. 466-35. USNM Ace. No. 1.^1571; 9 Feb. \9?,5: I specimen.

(23) Panama. Ladrones Island. 7"52' N, 82°27' W, taken interlidally

with scuba: TEPE 70-13. R/V Alplui Helix. Ace. No. 70-12, SIO

C3814; 19 Sept. 1970; coll. W. Newman and S. Luke; I male.

Ecuador. Galapagos Islands: (24) Tower Island, Darwin Bay. from

"roach trap." 2 mornings: Ace. No. 122445. USNM 68418: 23-24

Feb. 1933; coll. W. L. Schmitt; 150^- specimens; (25) Tower Island.

Darwin Bay. from coral collected at Seal Beach. No. I. by "Carl":

RA' Velt'io III Sta. 94-33, USNM Ace. No. 122445: 22 Feb. 1933: 1

specimen; (26) Tower Island, Darwin Bay, from "roach trap": Ace.

No. 122445, USNM 68437; 21 Feb. 1933; eoll. W. L. Schmitt: KXH

specimens; (27) Tower Island, Darwin Bay, coral gathered at Seal

Beach, No. 1: RA' Velem III Sta. 94-33, USNM Ace. No. 122445;

22 Feb. 1933; 1 specimen; (28) Seymour Island, from "roach trap"

attached to lobster trap; Ace. No. 122445. USNM 68424; 18 Feb.

1933; 60-1- specimens: (29) Albemarle Island, Albemarle Point,

shore collecting; R/V Velem III Sta. 69-33. LACM; 1 1 Feb. 1933; 4

specimens: (30) Indefatigable Island, Gordon Bay; R/V Velem III

Sta, 315-35, USNM Ace. No. 131571; 8 Dee. 1934; 1 specimen;

(31) Indefatigable Island, Academy Bay. dredged; R/V Velem III

Sta. 169-34. USNM Ace. No. 128938; 20 Jan. 1934; 1 specimen;

(32) Indefatigable Island, opposite Gordon Rocks, from coral in

shallow water; R/V Velem III Sta. 315-35, LACM; 8 Dec. 1935; 1

female; (33) Chase Island, Post Office Bay, from "roach trap" on

fish trap; Ace. No. 122445, USNM 68434; 6 Feb. 1933; coll. W. L.

Sehmitt; 1 female: (34) Hood Island, Gardiner Bay, rock spit;

USNM Ace. No. 122445; 25 Jan. 1933: 1 male: (35) Hood Island,

Gardiner Bay, Osborne Island: Ace. No. 122445, USNM 68435: 26

Jan. 1933: coll. W. L. Schmitt: 1 specimen. Ecuador (mainland):

(36) Manta. shore; #400. USNM Aec. No. 131571; 19 Jan. 1935: 1

male; (37) 1 mi. S. of Manta. from shore, reef west of breakwater:

#403, USNM Ace. No. 124571; 20 Jan. 1935; 1 male and I female:

(38) Salinas, with worm galleries; #1^, USNM Aee. No. 93322:

12-15 Sept. 1926: eoll. W. L. Sehmitt; 3 specimens: (39) Salinas:

#2, USNM Ace. No. 93322: 13 Sept. 1926: coll. W. L. Schmitt: 1

male;(40)Salinas;#l.USNMAee. No. 93322; 12 Sept. 1926: coll.

W. L. Schmitt; 2 males: (41) S. side of Santa Elena Point: #7.

USNM Ace. No. 93322; 17 Sept. 1926; I specimen: (42) La

Libertad. Bravo Point. S. side of Santa Elena Point; #7. USNM Ace.

No. 93322; 1 7 Sept. 1926; coll. W. L. Schmitt: 5 specimens: (43) La

Libertad. Bravo Point, S. side of St. Elena Point, shore collecting;

WW Velem III Sla. 19-33, USNM Ace. No. 122445; 21 Jan. 1933: 1

female.

Description of male. — Cephalon devoid of tubercles, about 2.5

times wider than long, with small rostral process, often with

interoeular furrow. Posterior region of cephalon often with short

lateral incisions on margins. Posterior portion of eyes covered by

anterolateral angles of pereonite 1 (Fig. 12C). Antennule very short,

reaching beyond anterior margin of pereonite I. with simple and

palmate setae and aesthetases as figured; flagellum of 7-12 articles

(Fig. 25A). Antenna long, reaching pereonite IV. in small speci-

mens reaching pereonite V. with simple setae; flagellum of 22-33

articles (distal portion not figured) (Fig. 25B). Frontal lamina pen-

tagonal, overlapped anteriorly by distal portion of rostral process

(Fig. 25C). Mandibular spine row forms large lobe with comblike

row of long spines, with 2 long setae at the base; molar process with

about 30 small acute marginal spines and numerous small simple

posterior marginal setae: middle article of palp with serrate setae:

distal article with comb and plumose setae (Fig. 25D). Maxillule

lateral lobe with 10 stout simple spines. 5 largest spines armed with

small barbs (Fig. 25E). Maxilla basal article with 2 short simple

spines on lateral margin: medial lobe with 9 plumose and 6 simple

setae. 1 simple seta near insertion of lateral lobes: lateral lobes with

11 and 4 long simple setae, respectively (Fig. 25F). Maxillipedal

endite very short, with 2 coupling spines, minute proximal lobe.

and 3 distal cireumplumose setae; palp with simple and comb setae

as figured (Fig. 25G).

Pereonite I 1.5 times length of others. Pereon broadest at

pereonites IV and V. Coxae IV-VII projecting beyond posterior

margins of pereonites and visible dorsally. Pereon without dorsal

tubercles, carinae, or setae (Fig. 12C). Pereopod I short, stout;

merus armed with 5 blunt robust niolariform spines and several

simple spines on inferior margin; propodus with 2 sitnple spines on

inferior margin and 1 blunt spine and 5 simple setae on distal angle

of inferior margin: daetylus with small simple seta at base and at

junction with unguis on inferior margin and a group of 5 simple

setae on superior margin near unguis (Fig. 26A). Pereopod IV long

and ambulatory, with large stout simple, serrate, and ba.sally den-

ticulate spines as figured (Fig. 26B). Pereopod VII long, with many

simple, serrate, and basally denticulate spines as figured (Fig. 26C).

Penes small (but larger than in C. diininuta) and set close together

on posterior region of sternite VII, in line with the inner (medial)

margins of peduncles of first pleopods (Fig. 26D).

Pleon widest at pleonite 2; pleonites 2-4 with posterolateral

angles somewhat acute, partly encompassing pleonites 3-5 respec-

tively (Fig. 12C). Pleopodal rami with PMS as figured (Figs. 26F-

J). Pleopod I: peduncle's medial margin with 4 coupling spines,

lateral margin with 1 large simple spine: endopod's lateral (outer)

margin very weakly concave; exopod with 1 stout spine on

proximolateral margin: endopod 0.75 width of exopod (Fig. 26F).

Pleopod 2: peduncle's medial margin with 4 coupling spines and 5

plumose setae, lateral margin with 1 simple spine: appendix

masculina 1.2 times as long as exopod. distal part narrowing to

slender apex, usually with a threadlike extension and small mar-

ginal (and occasionally medial) spines (Fig. 26G). Pleopod 3;

peduncle's medial margin with 4 coupling spines and 5 plumose

setae: lateral margin with 1 simple spine; exopod with short mar-

ginal incisions (Fig. 26H). Pleopod 4: peduncle's medial margin

with 4 coupling spines and 5 plumose setae; lateral margin with I

simple spine; exopod with short marginal incisions (Fig. 261).

Pleopod 5: peduncle with 1 simple spine on lateral margin: exopod

with short marginal incisions (Fig. 26J).

Pleotelson subtriangular. lateral margins straight or slightly con-

vex, apex rounded, with 8 stout marginal spines interspersed with

many PMS (occasional specimens can have 5 spines on one side of

the pleotelson. for a total of 9 marginal spines): pleotelson lacks

carinae or tubercles (Fig. I2C). Uropods extend to or slightly

beyond apex of pleotelson. margins serrate and notched apieally,

not distally rounded as in C. harfordi: rami more nartow than in C

harfonti: group of long simple setae arises from each apical notch;

both rami with spines and PMS on medial and lateral margins.

Uropodal exopod half as wide and slightly shorter than endopod;

medial margin with 3 stout simple spines, lateral margin with 7 or 8

stout simple spines. Uropodal endopod's medial margin with 4 or 5

stout simple spines, lateral margin with 2 or 3 stout simple spines.

Uropodal peduncle's medial margin with di.stal plumose .setae; I

mediolateral marginal spine; a group of spines and setae lies near

the articulation of the exopod (Fig. 26E).

Female. — Similar to male.

Size. — Small, to maximum length of 8 mm.

Distiihittion. — L'ntil recently. C. pcirva was thought to be a

eurythermal. circumtropical. and warm-temperate species (Miller

1968: Menzies and Glynn 1968). Bruce and Bowman (1982) re-

deseribed the species, noting that reliable records existed only for

the Caribbean and Gulf of Mexico. However. C. pana is also a

common member of the tropical eastern Pacific fauna, ranging from

southern Mexico (Oaxaea) south to the Galapagos Islands and

Ecuador, in shallow subtidal and intertidal habitats. Atlantic locales

for specimens examined in this study include the "West Indies," St.

Thomas. Puerto Rico. Yucatan Peninsula, and Costa Rica. As noted

38

R. C. Brusca, R. Wetzer, and S. C. France

Figure 26. Cirolaiia pana (USNM Ace. No. 93322). male: A, pereopod I. B. pereopod IV. C, pereopod VII. D. penes. E, ventral view of uropod.

F, pleopod 1 . G, pleopod 2. H, pleopod 3. 1, pleopod 4. J, pleopod 5.

Tropical Eastern Pacific Cirolanidae

39

by Hansen ( 1890) and Bruce and Bowman ( 1982). one oi' the type

specimens is labeled as being from "Samoa."

Renuiiks. — Bruce and Bowman (1982) raised the probabMily

that what has passed for Cimlana par\-a in the past Is In fact a

complex of closely related sibling species. They confirmed records

of C. pan'a from the Caribbean and Gulf of Mexico, and removed

C. diminiita from synonymy with it. After examination of the type

specimens. Bruce and Bowman's material, and eastern Pacific

specimens, we conclude that C. pana does occur in the tropical

eastern Pacific and is thus a tropical amphl-Ameiican species. The

characters distinguishing C. parva from the very similar C.

dimiiiiita are discussed above (see under "Remarks" for C.

iliniiiuiui). Hansen (1890) did not designate a type locality for

CiroUina pcirva. Bruce and Bowman (1982) noted this and desig-

nated St. Thomas. West Indies, as the type locality, basing this on

label data of dissected syntypes. Hansen (1890) did not figure the

pleopods. but Bruce and Bowman's (1982) figures of "syntypes"

show pleopods 4 and 5 with a complete transverse suture across the

endopod. However, the ZMUC specimen labeled "lectotype" and

other material we have examined lack complete endopodal sutures

(see Fig. 261 and J).

Conileni Leach. 1818

Type species. — Onisciis cylindraceus Montagu, 1 804. The

deposition of the type is unknown.

Ccmilera Leach. 1818: 348. Desniarest 1825: 304, Milne Edwards 1840:

24:. Bate and Weslwood 1867: 302. Hansen 1890: 338. Richardson 190.5:

1 16. Kussakin 1979: 216. Bruce 1985: 714; 1986a: 220. Wetzer. el al. 1987:

1, Kensley and Schotte 1989: 139.

Description. — Body 4.8-5.2 times longer than broad; dorsum

smooth, without ornamentation. Eyes small, separated by greater

than 2 eye-widths. Rostrum minute or absent. Frontal lamina,

clypeus. and labrum sessile, not projecting; narrow, anteriorly ex-

panded frontal lamina separates bases of antennules and antennae;

clypeus wider than long, and wider than labrum. Antennular pe-

duncle 3-articulate; basal article articulated at right angle to remain-

ing articles. Antennal peduncle 5-articulate; articles 3 and 4

subequal; 4th peduncular article with an elongate seta on distal

posterior angle. Mandible tridentate; spine row a well-developed

rounded lobe with stout spines; palp 3-articulate. middle article

longest. Maxillule medial lobe with 3 stout circumplumose spines;

lateral lobe with 10-12 stout, apical spines. Maxilla medial lobe

short, truncate, with many plumose setae; lateral lobe bifurcate with

apical simple setae. Maxilllpedal palp 5-articulate. with middle

article broader than others; endite with 1 coupling hook.

Pereonite 1 longest. Pereopodal dactyli without secondary

ungui. although there is often a small accessory spine at ba.se of

unguis. Pereopods l-lll: distal superior margins of ischium and

merus produced as a spoon-shaped process into which adjacent

distal articles fit. Pereopods V-VII with sparse setation; bases

without median longitudinal row of setae along outer surface. Penes

small.

Pleon of 5 free pleonites; pleonite 1 not concealed by pereonite

Vll; pleonite 4 encompassing lateral margins of pleonite 5. Pleopod

1 operculate to pleopods 2-5 and Indurate; peduncle elongated,

longer than wide; endopod elongate and narrow, medial margin

straight, thick, and almost bare but with moderately long PMS on

distolaleral margin; exopod shorter than the endopod. ovale, distal

margin with moderately long PMS. Pleopod 2: peduncle .scarcely

wider than long, rami with long PMS; appendix mascullnum arises

basally or submedially on endopod. Pleopod 5: endopod without

PMS. Pleotelson triangular with serrate posterior margin. Uropodal

peduncle with laterally produced angle; medial posterior angle

somewhat less produced; endopod with notch on lateral margin.

Remarks. — Until now. only 2 species had been assigned to

Ciinileia. C. cylindniccii (Montagu. 1804) and C. slxyici Packard,

1900. Conileia cyliitdnicea is a European species reported by

Richardson ( 1905) as also occurring in the western North Atlantic.

However, the true Identity of the specimens reported by Richardson

as C. cylindnicea from North American waters (South Carolina,

Mississippi, and Florida) Is uncertain. The type material of C.

cylindnicea is from Naples, Italy, and Richardson (1905) reported

additional records froin England and France. Richardson's (1905)

description and figures, taken from Sars ( 1890). are from Naples

specimens, not from the two specimens she claimed to have exam-

ined from North America. Hence, the American species has never

actually been figured or described. Tattersall (1906) did not ac-

knowledge this species as occurring in North America.

Conilera stygia is a blind form reported only once, from a

freshwater well in Monterrey, Nuevo Leon, Mexico. Cole and

Minckley (1966) suggested that this species may belong to

Speocirolana. The original description lacks figures and is inad-

equate for assessment of its proper generic assignment.

Little has been written on this genus, and the distribution of the

known species is rather enigmatic. Specimens have been collected

with dredges and trawls to about 300 meters, indicating a benthic

habitat. Bate and Westwood (1867) reported collecting Conilera

cylindracea "feeding together within the orbit of the eye of a

whiting, the eyeball of the fish being nearly detached from the

surrounding parts." Day (1884) reported that the entrails of a dog-

fish {.Acanthias vulgaris) "had been entirely eaten out by [C

cylindracea]." He further reported that this species "lives generally

on soft and sandy bottoms, hunts in shoals and. when abundant, will

drive away the congers and other fish." Biernbaum and Wenner

( 1993) reported collecting It in baited traps off the coast of South

Carolina, in depths of 194—212 m.

World list of species. —

1 . C. cylindracea (Montagu, 1 804). Europe and northwest Atlan-

tic (and perhaps southeastern U.S.A.)

2. C. stygia Packard. 1 900 (species incpurenda). Freshwater wells

from Monterrey. Nuevo Leon. Mexico.

3. C. bullisi n. sp. Ecuador.

Conilera bullisi n. sp.

Figs. 27-29

Type material examined. — Female holotype (USNM 252745)

and 3 paratypes (male, postmanca. manca) (USNM 252746); Ecua-

dor. Gulf of Guayaquil, 3° 1 9' S, 80°43' W, Menzies trawl; Anton

Bruun Cruise 18B. Sta. 769-D; 70 m; 10 Sept. 1966.

Description of female. — Cephalon smooth and unornamented,

1 .4 times wider than long, somewhat immersed in pereonite 1; eyes

small, separated by more than 2 eye-widths (Fig. 27). Antennule

short, not reaching posterior margin of cephalon; first article "L"-

shaped. second article arising at right angle from article 1 ; flagel-

lum of about 8 articles; proximal articles wider than long (Fig.

28A). Antennae longer than antennules but barely extending to

pereonite 1; flagellum of about 8 articles (Fig. 28B). Mandibular

incisor tricuspidate. outer cusp widest, inner cusp longe.st, middle

cusp indented; molar process with short, stout spines along upper

margin; spine row well developed with acute spines; palp 3-articu-

late, middle article longest and with 3 simple setae (Fig. 28D).

Maxillule medial lobe with 3 robust, circumplumose apical spines;

lateral lobe with 11 stout apical spines (Fig. 28E). Maxilla with

plumo.se setae on medial lobe, siiiiple apical setae on bifurcate

lateral lobe (Fig. 28F). Maxilllpedal palp 5-articulate; endite reach-

ing middle of palp article 2, with 1 coupling spine (Fig. 28G).

Pereon smooth, not sculptured; pereonites subequal in width;

coxal plates not visible in dorsal aspect; in lateral aspect coxal

plates II-IV subrectangular, V-VIl with acute posterior margins.

Pereopods l-III; propodi with 2 spines on inferior margin and

40

R. C. Brusca, R. Wetzer, and S. C. France

Figure 27. Conilem hullisi n. sp. (USNM 252745),

holotype, female.

clump of simple setae on superior distal margin: caipus with 1 or 2

spines on inferior margin; merus of pereopod I with 3 hlunt and 3

acute spines on inferior margin; merus of pereopod III with only 2

blunt spines; pereopods l-III with 1 blunt spine among setae on

produced distal-superior margin of merus; dactyli shorter than

propodi. slightly curved (Figs. 29A. B). Pereopod VII: merus with

somewhat produced superior distal margin, bearing setae; inferior

margin of merus, carpus, and propodus with thin, acute spines,

sparsely setose; dactylus about 0.5 times length of propodus (Fig.

29C).

Pleonite I longer and wider than subsequent pleonites. Pleonite

5 completely enclosed by pleonite 4; pleonite 5 with a dense row of

simple spines on posterior margin (Figs. 27, 29K). Pleopodal rami

with PMS as figured (Figs. 29E-I). Pleopod 1: peduncle about 1.4

times longer than wide, with 5 coupling spines inset on medial

margin (Fig. 29E). Peduncles of pleopods 2— t with 3 coupling

spines on medial margin. Pleopod 5 without coupling spines.

Pleopodal endopods 1—4 with PMS on apical margin, e,\opods with

long PMS on apical and lateral margins. Pleopods 3-5: exopods

with notch in lateral margin. Uropodal endopod and exopod mar-

gins strongly serrate, with a single PMS arising between each tooth

(Fig. 29D); endopod 1.3 times longer than exopod; exopod nar-

rower than endopod; endopod does not extend beyond posterior

margin of pleotelson.

Description of male. — Penes small, bean shaped, almost touch-

ing (Fig. 29L). Appendices masculinae arise on endopods of pleo-

pods 2 (Fig. 29J). Otherwise similar to female.

Size. — To maximum length of 7 mm.

Dislrihiilion. — Known only from the type locality. Gulf of

Guayaquil. Ecuador.

Remarks. — Conilera hullisi is only the second (the third if C.

stygia is eventually retained in the genus) species described in this

genus, and it is the first Conilera species recorded from the Pacific.

Etymology. — This species is named for William Bullis, practi-

tioner of law, adventurer, friend, and surfer par excellence, in deep

gratitude for the assistance provided to us for our expeditionary

work.

Eiirydice Leach, 1815

Txpe .species. — Eurxdice puUhra Leach. 1815. Type material at

BMNH.

Synonymy. — Emended and subsequent to Bruce ( 1986a: 1 1 ).

Eundice. Brusca 1980: 228. Menzies and Kruczynski 1983: 84. Brusca

and Iverson 1985: 34. Kensley and Schotte 1989: 147.

Description. — Body 2.5-3.5 times longer than broad; dorsum

smooth, without ornamentation. Eyes small to large; onimatidia

extend to ventral lateral position on cephalon. Rostrum minute or

absent. Frontal lamina narrow, anterior part projecting ventrally, not

joined with clypeus; clypeus short, broad, wider than long, with a

freely projecting triangular blade; labrum broadly joined to clypeus

and slightly narrower than clypeus. Antennule short; peduncle 3-

articulate; article 1 longest; article 2 arises at right angle to article I ;

proximal flagellar articles typically fused. Antennal peduncle 4-

articulate (articles 4 and 5 fused?); article 4 longest. Mandible with

broad, tridentate incisor; palp extends well beyond cutting edge;

spine row a large, rounded lobe with stout simple spines; middle

article of palp longest. Maxillule medial lobe with 3 stout

circumplumose spines; lateral lobe with 11-13 stout spines. Max-

illa small, simple medial lobe absent or weakly developed.

Ma)iillipedal palp articles 3 and 4 subequal in length and width;

endite reduced, failing to reach or barely reaching beyond first palp

article, and without coupling spines.

Pereonites I and II subequal in length. Coxae II-VII with

posteroventral angles becoming increasingly acute posteriorly.

Pereopods I-III short, grasping, spinose, with distal superior mar-

gins of ischium and merus more or less produced. Pereopods IV-

Tropical Easlem Pacific Cirolanidae

41

Figure 28. Coiiilem htillisi n. sp. (USNM 252745). holotype, female: A. antennule (left). B. antenna (left). C. frontal lamina,

clypeus. and labrum. D. mandible (left). E. maxillule (left). F, maxilla (left). G, maxilliped (left).

VII long, spinose, setose; ischiuin-propodus flattened. Penes large,

well developed, 1 .5-3.5 times longer than broad, lobes somewhat

flattened.

Pleon of 5 free pleonites; pleonite 5's lateral margins not over-

lapped by pleonite 4. Peduncles of pleopods 1-5 wider than long,

pleopod I 's peduncle subquadrate. Pleopods rounded, only endopod

of pleopod 5 lacking setae; appendix masculina inserted medially

or submedially on endopod on male's pleopod 2. Pleopod 5: pe-

duncle without coupling spines or plumose setae on medial margin;

endopod with proximomedial margin produced, lobelike.

Pleotelson with apex rounded, truncate, or subacute, usually emar-

ginate; dorsum with anteromedial depression. Uropodal peduncle

with inner angle not greatly produced, with row of PMS on lateral

margin; exopod lateral margin without PMS.

Remarks. — Species in this genus are superficially similar in

appearance to those of Cirolaiui and Anopsilana but may be

42

R. C. Brusca, R. Wetzer, and S. C. France

Figure 29. Coiiileni huUisi n. sp. (USNM 2?2745|. holotype, female (except J. L, paratype, male): A, pereopod I. B. pereopod III. C. pereopod VII. D,

dorsal view of uropod. E, pleopod I . F, pleopod 2. G, pleopod 3. H, pleopod 4. 1, pleopod 5. J. pleopod 2 of paratype male ( USNM 252746). K, dorsal view

of pleonite 5 and pleotelson. L, penes of paratype male (USNM 252746).

Tropical Eastern Pacific Cirolanidae

43

quickly distinguished by the unique morphology of the antennu-

lar peduncle (the right-angle relationship between articles 1 and

2), the fused basal articles of the antennular tlagellum, the 4-

articulate antennal peduncle, the reduced maxillipedal endite and

absence of coupling spines, the large penes, and characters of the

pleotelson and uropods. Bruce (1986a) discussed sexual dimor-

phism in this genus.

Eiiixdice also shares some features with MeuuiroUiiui. another

cirolanid genus present in the tropical eastern Pacific. Characters

common to both genera include the short first pereonite (subequal

in length to pereonite II), all pleonites being free, prominent penes,

and a projecting clypeus.

Eiiiydice is a cosmopolitan genus containing approximately 46

described species. Soika ( 1955) proposed the subgenus Pehigonice

for the pelagic species lacking pleotelsonic spines. Like Bruce

(1986a) and Jones and Naylor ( 1967). we doubl that this subgenus

is monophyletic or has any systematic utility.

Only a single species oi Eiirydice (£. caitdata) is known from

the tropical eastern Pacific. Its ecology has not been studied.

Eleftheriou and Jones (1976) have reviewed the ecology and sys-

tematics of the Indian Ocean species; Jones ( 1974) discussed the

ecology of Saudi Arabian species; Soika (1955). Singarajah ( 1966).

Knight-Jones and Qusim (1967). Jones and Naylor (1970). Jones

(1968. 1970a. 1970b). Salvat (1966). Fish and Fish (1972), Albeit

and Naylor (1976), Hastings and Naylor ( 1980), Hastings (1981a.

1981b). and Tully and O'Ceidigh ( 1986) have all reported on the

ecology of European species; Bruce ( 1986a) discussed the Austra-

lian species; and DeRuyck et al. ( 1991 ) reported on South African

species. It appears that most species are predators and/or carnivo-

rous scavengers, and most prefer intertidal and shallow subtidal

habitats, where they maintain endogenous circadian activity

rhythms. Bruce (1986a) noted that there are no intertidal species of

Euiydice in Australia, all eight Australian species being exclusively

subtidal. The eastern Pacific species. E. caudata, is most commonly

collected by night lighting in shallow water.

World list of species. —

1. E. aculicaiida Bruce, 1981. Victoria. Australia.

2. E. affhiis Hansen, 1905. Northeast Atlantic and Mediterra-

nean.

3. E. (/g///.v Jones, 1971. Kenya.

4. E. akiyimiai Nunomura. 1981. Central Japan.

5. E. arahica Jones. 1974. Red Sea.

6. E. haihypelagica Schultz. 1977. Subantarctic. Incertae sedis

according to Bruce ( 1986a).

7. E. hiiuhi Bruce. 1986. New South Wales. Australia.

8. E. caeca Hansen. 1916. North Atlantic. Incertae sedis accord-

ing to Bruce (1986a).

9. E. caudata Richardson. 1899. Southern California to Ecuador.

10. E. cavicaitdata Jones. 1971. Kenya.

11. E. chelifer Jones. 1 97 1 . Kenya.

12. E. clymeneia Monod. 1926. Morocco.

13. E. convexa Richardson. 1900. Florida.

14. E. t'ccra/(n'.?AT Bacescu. 1948. Mediteiranean.

15. E. dollftisi Monod, 1930. Adriatic. Mediterranean, and Black

seas.

16. E. eloiifiata Moreira, 1972. Brazil.

17. £. emarginata Moreira, 1972. Brazil.

18. £. grimaldii Dollfus. 1888. Northeast Atlantic.

19. E. luimilis S(ebb\no. 1910. Maldives.

20. E. indicis Eleftheriou and Jones. 1976. Southwestern India.

21. E. iiiermis Hansen. 1 890. Atlantic and Mediterranean coasts of

Europe.

22. E. inornata iones. 1971. Kenya.

23. E. littoralis ( Moore. 1 902 ). Atlantic coast, from North America

to Brazil.

24. E. longianleimala Nunomura and Ikehara. 1985. Sea of Japan.

25. E. longicornis (Studer, 1883). Namibia.

26. E. longipes ioncs. 1971. Kenya.

27. E. longispina ionefi. 1969. Mediterranean.

28. E. muuritanica DeGrave and Jones. 1991. Mauritania.

29. £. »(//iv« Bruce. 1986. Australia.

30. E. nipponica Bruce and Jones. 1981 . Japan.

31. E. (irientalis Hansen. 1890. Australia. Philippines. Indonesia,

Papua New Guinea. Sri Lanka, and Indo-China.

32. E. peraticis Jones. 1974. Red Sea to western India.

33. E. perstDuild Kcnsley. 1987. South Carolina to Florida. Greater

Antilles. Turks and Caicos, and Venezuela.

34. E.piperata Menzies and Frankenberg. 1966. Georgia to Gulf

of Mexico.

35. E. pontica (Czerniavsky. 1 868). Black Sea and Mediterranean.

36. E. pulchra Leach. 1815. Atlantic coast of Europe and North

Africa.

37. E. racnvitzae Bacescu. 1949. Mediterranean.

38. E. rotundicauda Norman. 1906. Eastern North Atlantic and

Mediterranean.

39. E. spenceri Bruce. 1981. South Australia.

40. E. spinigera Hansen. 1 890. Atlantic and Mediterranean coasts

of Europe.

41. E. stdytruncataJaXXeviaW, 1921. New Zealand.

42. E. tarti Bruce. 1986. Victoria, Australia.

43. E. trimcata (Norman. 1 868). Eastern North Atlantic and Medi-

tenancan.

44. E. valkanovi Bacescu. 1949. Black Sea.

45. E. u'oAa Bruce. 1986. Queensland. Australia.

46. E. wyuiui Bruce. 1986. Queensland. Australia.

Euiydice caudata Richardson,

Figs. 30-34

1899

Euiydice caudata Ricliardson, 1899a: 824; 1899h: 164; 1900: 217;

1901:.'ii6; 1905: 124. Steinbeck and Ricketts 1941: 424. Schultz 1969: 173.

Bowman 1977: 654. Bnisca 1980: 228. Brusca and Ivenson 1985: 34. Bruce

1986a: 221.

Eurvdice branchuropus Menzies and Barnard. 1 959: 32. Schultz

1966: 14; 1969: 173.

Type material examined. — Syntypes (USNM 22565): U.S.A.,

California. Santa Catalina Island. Isthmus Cove; R/V Albatross; 6

Apr. 1897; 20 specimens.

Other material examined. — California specimens: (2) Monterey

Co.. Pacific Grove, in compound tunicate mass; LACM/AHF [no

catalog number]; 1 Apr. 1929; 5 specimens; (3) Los Angeles Co.,

Santa Catalina Island. Emerald Cove. 33°28' N. 1 18°3r W. 27.5 m,

among school of squid copulating under night light; R/V Velero IV

Sta. 2047-51, LACM; 27 July 1951; I specimen; (4) Los Angeles

Co., Santa Catalina Island, off Howland's Landing, 40 m; R/V

Vt'/cwSta. 2 141-52, AHF Cat. No. 770-01. LACM; 6 Aug. 1952; 1

specimen; (5) San Diego Co.. La Jolla. Scripps Pier; No. H46-63.

SDNHM; 2 specimens. Pacific Baja California specimens: (6)

Guadalupe Island (south end). Melpomene Cove, taken at night

with light; AHF [no catalog number]; 27 Jan. -3 Feb. 1950; coll. C.

Hubbs and party; 3 1 specimens; (7) San Benito Islands; RA' Alba-

tro.ts Sta. 5767. USNM 69719; 9 Mar. 191 1; 20 .specimens.

Gulf of California specimens: (8) Sonora. San Jorge Island,

anchorage off west side, dip net under night light; UA 72-60.

SDNHM A.0146; 10 Dec. 1972; 25 specimens; (9) Baja California

(Norte). Puertecitos; 13 Apr. 1973; SDNHM A.0068; coll. D.

Dexter; 30+ specimens; ( 10) Sonora. Tiburon Island. Risco Colo-

rado Point, on Sargassuin at night with light. 10 m; HC-IO-III-

1985. SDNHM; 10 Mar. 1985; coll. H. Chancy; 50+ specimens;

(II) Baja California. Las Animas Island, on sand. 17 m; AHF Cat.

No. 1282-02. LACM; 9 Sept. 1978; coll. A. Kerstitch; 20-(- .speci-

mens; (12) Baja California. San Francisquito Bay. taken with elec-

tric light; R/V Albatross Sta. A5767. USNM 69720; 1911; 15-t-

specimens; (13) Baja California. San Pedro Nolasco Island, sur-

face, by dip net just after dark; AHF Cat. No. 1997-1, LACM; 28

44

R. C. Brusca. R. Wetzer, and S. C. France

Figure 30. Eunilice caudala (USNM 22563). syntype. male.

June 1976; coll. M. Gilligan; 11 specimens; (14) Sonora, San

Carlos Bay; Ace. 160366, USNM 86346; 30 Mar. 1940; coll. E. F.

Ricketts; 15 specimens; (15) Sonora, San Carlos Bay. "pelagic

haul." light over side at anchorage; Ace. 160366, USNM 86342; 30

Mar. 1940; coll. E. F. Ricketts; 30+ specimens; (16) Baja Califor-

nia, Tortuga Island; 12 Mar 1936; night light; USNM Ace. No.

139772; 300+ specimens (in 3 vials); (17) Baja California, S. of

Tortuga Island, 27"24'50" N, 111°53'35" W, 137 m, sand; RA'

Velem HI Sta. 695-37; 17 Mar. 1937; 50+ .specimens; (18) Baja

California, Concepcion Bay, light over side at anchorage; USNM

86343; 28 Mar 1940; coll. E. F. Ricketts; 25 specimens; (19) Baja

California, 1 mi. off Pulpito Point, taken at night with dip net and

night-light, 2115-2215 hrs.; LACM; 22 June 1976; coll. R. H.

Behrstock; 1 specimen; (20) Baja California, Puerto Escondido

(across from Carmen Island). 25'=48' N, 1 1°18' W, 31 m, taken with

night light at 0900 hrs.; RA' Velero IV Sta. 1751-49, LACM; 19

Mar. 1949; 17 specimens; (21) Baja California, Carmen Island,

taken with electric light; R/V Albatross Sta. A5767, USNM 69718;

191 1; 20+ specimens; (22) Baja California, Carmen Island, Salinas

Bay; SDNHM; 14 May 1939; coll. R. W. Mindte, M/S St. Mary\

500+ specimens; (23) Baja California, Agua Verde Bay, with elec-

tric light; R/V Albatross Sta. A5767, USNM 69722; 15+ speci-

mens; (24) Baja California. Agua Verde Bay near Marcial Point,

25°3r N, 1 1 1°()4' W. night light off ship, 2200 hrs.; RA' Velero IV

Sta. 1741-49. LACM; 16 Mar 1949; 50+ specimens; (25) Baja

California, off Marcial Point, "pelagic." 24 m. at anchorage, 2200

hrs.; Ace. 159124, USNM 86337; 24 Mar. 1940; coll. E. F Ricketts;

100+ specimens; (26) Sinaloa, Topolobampo, 25°36' N. 109°04' W;

SDNHM A.0068; 8 July 1972; coll. D. Dexter; 4 specimens; (27)

Baja California, San Jose Island, Ostiones Point, night, at 10 m on

algae; HC-25-II-1985, SDNHM; 25 Feb. 1985; coll. H. Chaney;

150+ specimens; (28) Baja California. San Jose Island, taken with

electric light; R/V Albatross Sta. A5767, USNM 69724; 1911; 10

specimens; (29) Baja California. San Francisco Island, east coast,

taken at night, over 1 8 m bottom with red and green algae, sand, and

ExciroUma sp.; HC-24-11-1985. SDNHM; 24 Feb. 1985; coll. H.

Chaney; 100+ specimens; (30) Baja California, San Francisco Is-

land, S.W. cove, surface; R/V Velero III. LACM; 5 May 1932; 1

specimen; (31) Baja California, Inner Gorda Bank, off San Jose del

Cabo, dredge, 128-143 m; R/V Velero IV 1 135-40, LACM; 20 Jan.

1940; 1 specimen; (32) Baja California, Cape San Lucas; R/V

Albatross Sta. A5767, USNM 69723; 1911; 30+ specimens; (33)

Baja California, Cape San Lucas, "pelagic," with night light;

USNM 86344; 29 Mar 1940; coll. E. F Ricketts; 4 specimens.

Central eastern Pacific specimens: (34) Mexico, Jalisco,

Tenacita Bay, 25°58' N, 1 13°08' W, shore and land collecting; R/V

Velero III Sta. 2-33. LACM; 3 Jan. 1933; 1 specimen. (35) Costa

Rica, Cocos Island, on fish; USNM Ace. 122445; coll. W. L.

Schmitt; 1 specimen. (36) Costa Rica, Cocos Island, Chatham Bay;

Ace. 148787. USNM 86345; 3 Aug. 1938; coll. W. L. Schmitt; 2

juveniles. (37) Costa Rica, Cocos Island. Chatham Bay, 5°33.3' N,

87°02.63' W. Porites reef, 12-15 m; CRC-88-1, LACM 88-7; coll.

R. W. Peck and H. G. Kuck; 10 specimens. (38) Ecuador, La

Libertad; R/V Velero III Sta. 13-33. LACM; 19 Jan. 1933; 3 speci-

mens. (39) Isabel Island, "2 1 ''52' N, 102 "5 1 ' W," taken at anchorage

with electric light; AHF 748-37. LACM; 10 specimens (coordinates

evidently in error].

Description of male. — Cephalon width 3.2 times length; ante-

rior margin of cephalon evenly rounded, without rostral point. Eyes

large and well developed (Fig. 30). Antennule short, barely reach-

ing posterior inargin of cephalon, flagellum with a large basal piece

comprising numerous fused articles and aethestasc bundles and 3

free articles (Figs. 3 1 A, B. 32A). Antenna very long, usually reach-

ing to pereonite VII, occasionally to pleotelson (Fig. 33A); flagel-

lum of 20-25 articles, distal articles 3—4 times longer than broad

(Fig. 32B). Frontal lamina greatly elongated, posteriorly acute

Tropical Eastern Pacific Cirolanidae

45

Figure 31. Eiindice caiidaui iLACM specimens from Mexico, Sonora.Tiburon Island, coll. H.Chaney, 10 Mar. 1985). scanning electron micrographs: A.

anterior view of cephalon and antennules, 50x. B. detail of aniennule peduncle. lOOx. C. frontal lamina, clypeus, and labnim, lOOx. D. frontal lamina. 200x.

46

R. C. Brusca. R. Wetzer, and S. C. France

Figure 32. Eurydice caiidaia. A, B, USNM 22565 syntype, male, C-G, AHF 770-01. male: A,

antennule. B. antenna. C. frontal lamina, clypeus. and labrum. D. mandible. E. maxillule. F, maxilla. G,

maxilliped.

(Figs. 3 IC, D, 32C); posterior margin of labrum rounded. Mandibu-

lar spine row forms a rounded lobe with 5-1 1 simple spines: molar

process with 16-18 acute marginal spines: middle palp article with

7 simple setae, distal article with 11 simple setae as figured (Fig.

32D). Maxillule lateral lobe with 11 large simple spines. 2 with

small distal barbs {Fig. 32E). Maxilla medial lobe with 2 plumose

setae and 4 simple setae: lateral lobes with 4 and 2 simple setae,

respectively (Fig. 32F). Ma.xillipedal endite with 1 plumose seta

and 2 simple setae; palp articles with marginal simple setae and

hairs as figured (Fig. 32G).

Pereon widest at pereonites IV-VI. Coxal plates V-VII visible

in dorsal view; posterior angle of coxa VI elongate and narrowed,

extending nearly to posterior margin of coxa Vll; posterior angle of

coxa VII not elongate and naiTowed but rather blunt (Fig. 30).

Pereopod I short, distal superior margin of merus and ischium

weakly produced (not strongly produced as in NatalolaiwY. distal

superior and inferior inargins of basis with a few long simple setae;

inferior margin of ischium with row of simple setae, distal superior

margin with 1 large spine and several simple setae; superior distal

angle of merus with 4-8 long simple setae, inferior margin with 3

short blunt spines and several simple setae; inferior distal angle of

carpus with 1 blunt spine and several setae; propodus long, more

than twice length of dactylus, superior margin with numerous long

simple setae, inferior margin with 3 kinds of setae, simple, comb,

and serrate, and 1 large stout spine at di.stal angle; dactylus with 1

comb seta and several short simple setae spine at base of unguis

lacking (Fig. 34A). Pereopod IV short; ischium-propodus with

large stout spines on superior and inferior margins as figured;

without spine at base of unguis (Fig. 34B). Pereopod VII long;

merus, carpus, and propodus flattened, longer than broad, and

wider than propodus; with many simple setae and spines as figured

(Fig. 34C). Penes 1.5-2.0 times longer than broad, rounded apically,

with a "joint" or fold about quarter distance from base, set close

together, and separated by 1 minute simple seta (Figs. 33B, 34D).

Pleon elongate, nearly as wide as pereon. Pleonite 5's posterolat-

eral margins slightly produced (Fig. 30). Pleopodal rami with PMS

as figured (Figs. 34F-J). Pleopod 1 smaller than pleopods 2-5;

medial margin with 4 coupling spines and 2 plumose setae; endopod

Tropical EaMem Pacific Cirolanidae

47

Figure 33. Eiirydice cauduui (LACM specimens from Mexico, Sonora,

Tiburon Island, coll. H. Chaney. 10 Mar 1985), scanning electron micro-

graphs: A, lateral view, 15x. B, ventral view of penes on stemite VII. 89x.

0.78 times width of exopod, lateral (outer) margin concave (Fig.

34F). Pleopod 2: peduncle's medial margin with 3 coupling spines

and 2 plumose setae, lateral margin with 1 simple seta; endopod

width 0.92 times exopod width; appendix masculina widening

subapically. scythe-shaped, length 1.19 times length of exopod (Fig.

34G). Pleopod 3; peduncle's medial margin with 3 coupling spines

and 2 plumose setae, lateral margin with 1 simple seta; endopod

width 0.89 times width of exopod; exopod with complete medial

incision (Fig. 34H). Pleopod 4: peduncle's medial margin with 3

coupling spines and I plumose seta, lateral margin with I simple

seta; endopod width 0.73 times e.xopod width; exopod with complete

medial incision (Fig. 341). Pleopod 5: endopod width 0.69 limes

exopod width; exopod with complete medial incision (Fig. 34J).

Pleotelson broadly rounded, wider than long, lateral margins

convex; apex truncate, with shallow transverse indentation set off

by acutely produced corners; indentation with PMS and 4 short

simple spines. Pleotelson dorsum with shallow depression near

base, without carinae or tubercles (Fig. 30). Uropodal peduncle not

visible dorsally; lateral margin fringed with long PMS. Uropodal

rami do not extend beyond pleotelson apex: rami without apical

notches; fringed with long PMS distally; outer distal angle of each

ramus with 2-3 small spines. Uropodal endopod 1.4 times exopod

width, distal lateral margins slightly rounded (Fig. 34E).

Female. — Similar to male, although antennal Oagella may be

somewhat shorter than in males and pleon may be slightly narrower

than pereon.

Ullmslnictiiriil fealiires. — When viewed with SEM. the surface

of the body and appendages has a scalelike cuticular structure. The

body dorsum has many cuticular sensillae set inside small pits (Fig,

3 1 A).

Size. — Small, to maximum length of 9 mm.

Dislrihitlion. — Eiiryilice cciiicldtd ranges from southern Califor-

nia to the Gulf of Guayaquil, Ecuador, including the islands of

Guadalupe, Reviilagigedos, Tres Marias, and Cocos. It is apparently

common throughout its range, both in the warm temperate Califor-

nian Province and in the tropics. The northernmost record (Pacific

Grove, California), the only one north of the California Province,

seems to be an anomaly. The southernmost record is for La Libertad,

Ecuador (on the north shore of the Gulf of Guayaquil). This species

is very common throughout the Gulf of California. Its depth range,

based on the material we have examined, is from the low intertidal

region to 160 m; most records are from 0 to ."iO m depth.

Reniark.s. — Eurydice hranchiiropiis was described from southern

California by Menzies and Barnard ( 1939) but reduced to a synonym

of the Atlantic species E. litlorulis (Moore) by Menzies and Glynn

( 1 968 ). Bowman ( 1 977 ). however, removed E. hninchiiropus from E.

liuoialis and placed it in synonymy with E. caiidata.

Ewydice caiidata is most often taken by night-lighting off ships

anchored over shallow soft bottoms. In California and western

Me.xieo we have collected it regularly from coarse sands and silts,

at depths of 5 to 1 60 m.

Exciwlana Richardson, 1912

Type species. — Cirolana oiientalis Dana, 1853, from the Sulu

Sea, by original designation (Richardson 1912). Location of type

material unknown; specimens possibly lost when the sloop "Pea-

cock" sank off the mouth of the Columbia River (Bruce 1986a).

Synonymy. — Emended and subsequent to Bruce ( 1986a;39).

E.\ciwlwui. Brusca 1980: 227. Jones 1983: 309. Brusca and Iverson

1985: 31. Kensley and Schotte 1989: 149.

PonJogeloides Barnard, 1914: 355.

Description. — Body 2.5-3.0 times longer than broad; dorsum

evenly convex and unsculptured; pereonite I almost always longer

than pereonite II. Eyes moderate in size. Posterior region of

cephalon often with lateral incisions; cephalon with prominent

rostrum; in all but three described species the rostrum is strongly

dilated distally as a spatulate process separating antennules (only

moderately dilated in E. monodi and E. hirsuticauda): rostrum

contlucnt with (or fused to) frontal lamina. Frontal lamina varies

from narrow and linear to somewhat wide (but always longer than

wide); clypeus short, wider than long, sublriangular, weakly pro-

jecting, narrower than labrum; labrum broad. Antennular peduncle

3-articulate, but articles 1 and 2 are fused in many species; article 3

never as long as 1 and 2 combined; tlagellum may vary in length

with age. but always longer than peduncle; basal flagellar articles

longer than wide. Antennal peduncle 4- or 5-articulate; if 5-articu-

late. article 5 never as long as 1—4 combined. Mandible with broad

tridentate incisor, quadridentate on left mandible in many species;

2- or 3-articulate palp extends beyond incisor's cutting edge; spine

row and molar process well developed; spine row lobelike, with

numerous long, acute, flexible spines. Maxillule medial lobe with 3

stout circumplumose setae and occasionally 1 or 2 smaller

circumplumose setae; lateral lobe with several to many stout simple

spines (often barbed). Maxilla medial lobe moderately developed

and setose, with bifurcate lateral lobe well developed, with many

long simple setae. Maxilliped slender, palp of 5 articles, article 3

wider than 4; endite with I coupling spine.

48

R. C. Brusca, R. Welzer, and S. C. France

Figure 34. Eurydice caudata. A-C. USNM 22565 synlype. male. D-I. AHF 770-01, male: A, pereopod I. B, pereopod IV. C. pereopod VII.

D, penes. E, dorsal view ofuropod (inset: details of apices). F, pleopod 1. G, pleopod 2. H, pleopod 3. I, pleopod 4. J. pleopod 5.

Pereopods all ambulatory; l-III grasping: bases of V-VII nar-

row, not markedly flattened; all legs usually bear spines on the

propodus, carpus, merus. and ischium; pereopods often with large

accessory spine at base of ungui. Penes well developed on sternite

of pereonite VII.

Pleon of 5 free pleonites; pleonite 1 may be largely or partly

hidden by pereonite VIl; pleonite 5 as wide as 4. with lateral margins

not overlapped (or only partly overlapped) by 4. Pleopods with

elongate rami; pleopod 1 not indurate oroperculate; pleopods 1 and

2 generally similar, with PMS on both rami; exopods of pleopods 3-

Tropical Eastern Pacific Ciroianidae

49

3 with PMS. endopods of 3-5 (or 2-3) without PMS: appendix

mascuhna of male arises basally or submedially on margin of

endopod of pleopod 2. Pleopodal peduncles I— I with coupling

spines and plumose setae on medial margins and usually on acces-

sory lobes on lateral margins: pleopodal peduncle 3 without cou-

pling spines. Pleotelson apex subacute or rounded, never indented or

excavated; with or without spines, but always with PMS. Dorsum of

pleotelson with 2 shallow submedian depressions, one either side of

midline (in some species these pits are prominent, in others they are

shallow and difficult to discern). Uropodal peduncle's inner angle

not produced or only modestly produced, never long and acute; rami

usually extend to or beyond posterior border of pleotelson; exopod

equal to or longer than endopod; medial distal margins of uropodal

exopod and often endopod usually with stout spines and PMS, outer

margins without spines in most species (present in some); lateral

margin of endopod usually with a small marginal pit.

Remarks. — Bruce (1986a) felt that the genus Ewirolana was

difficult to describe and lacked constant morphological features. He

also noted the unsettled synonymy of Exciiolana with Ponto-

gehides (see Monod 19.^0. 193.3; Jones 1971; Carvacho 1977).

Some authors have regarded Pontogeloides as a subgenus of

Excimhiiui (Monod 1930. 1931; Nierstrasz 1931). There seems to

be no compelling reason to maintain ExciwUma and Pontogeloides

as separate genera, and we agree with Bruce that the latter is a

junior synonym of the former. Only the presence of naked endopods

on pleopod 2 distinguishes Pontogeloides.

There are two synapomorphies unambiguously defining

Exciiolana (including Pontogeloides). First is the prominent rostral

process, which is apically dilated and usually spatulate — it is

confluent with (or indistinguishably fused to) the frontal lamina.

The rostrum separates the antennules. and the frontal lamina sepa-

rates the antennae. Second, the genus is ovoviviparous. with

enlarged oviducts serving as uteri and with reduced oostegites.

(However, internal brooding also occurs in Annina; see below). In

addition to these two attributes, there are two other, nearly consis-

tent features of the genus. First, the lateral margin of the uropodal

endopods usually bears a small pit or notch. This structure is often

difficult to see through the light microscope, and it is impossible to

judge from the literature if it occurs in all species, but we suspect

that it does. In some species (e. g.. E. hirsuticauda) the pit manifests

itself as a minute marginal pocket, flush with the margin of the

endopod and very difficult to see; in others (e. g., E. brazdienisis) it

forms a deep obvious notch in the endopod margin. The function of

this curious pit is unknown. Second, almost all species have distinct

paired submedian depressions on the dorsum of the pleotelson —

these appear to be poorly developed or wanting in at least E.

hirsuticauda and E. Uitipes.

These four features, in combination with those listed in the

general description, unambiguously distinguish this genus. Menzies

(1962a) claimed that the rostrum of £. hirsuticauda was acute, not

expanded. However, his own illustrations of this species, and our

examination of specimens from Chilean sand beaches, indicate that

the rostrum is indeed expanded, albeit weakly. We also note that

numerous authors have mistakenly referred to the rostrum in

Excirolana as the "frontal lamina."

The genus Annina is very similar to Excirolana. Both Annina

lacustris Budde-Lund, 1908 (= Excirolana Innvinani Jones and

Iceley, 1981) and A. kiiinari ( Bowman, 1 97 1 ) ( = E.xcirolana kiimari

Bowman, 1971 ) have previously been assigned to Excirolana. and

Messana (1984) suggested that Excirolana should be considered a

junior synonym oi Annina. However, as Jones (1983) and Bowman

and lliffe (1991) pointed out, several features of Annina clearly

distinguish it from E.xcirolana. \n Annina. the eyes are divided into

dorsal and ventral parts by a nonfaceted gap, article 3 of the

antennule is always as long as or longer than articles 1 and 2

combined, article 5 of the antenna is equal to or longer than articles

I -4 combined, pleonites I and 2 are narrower than 3-3 and covered

laterally by the coxal plates of pereonites VI and VII. the uropods

have a strongly produced mediodistal angle, the cephalon or

pereonites of males bear prominent dorsal horns, and the character-

istic spatulate rostrum and uropodal endopod notch of E.xcirolana

are apparently wanting. However, the two genera are probably

closely related. Their being the only two cirolanid genera known to

brood internally suggests that they might be sister groups, although

the precise nature of the brood pouch in Annina has yet to be

described.

So tar as is known all species o( Excirolana are ovoviviparous,

w ith the eggs retained and the embryos brooded in enlarged paired

oviducts functioning as uteri. Females of at least some species

develop small, reduced oostegites, although gravid and oostegite-

bearing females are rare in collections. The oostegites in some may

be rather large (in E. inayana they meet in the midline). Oostegites,

when present, are easily overlooked because they lie tightly pressed

against the sternum.

Brusca and Iverson (1985) briefly summarized the natural his-

tory of Excirolana. A characteristic that has been occasionally

noted in the literature is the ability of some species to inflict a

painful bite, especially on swimmers" thin skin. We have been

bitten on many occasions by E. inayana. a voracious species that

often emerges in high numbers near shore. Most species of

E.xcirolana are tropical. Bally ( 1983) studied the respiratory activ-

ity of E. natalensis. a southern African species inhabiting both

temperate and tropical shores. DeRuyck et al. ( 1991 ) documented

endogenous circadian rhythms in two South African species.

Three species of Excirolana occur in the tropical eastern Pacific:

£. hraziliensis. E. inayana. and E. chaineiisis. E.xcirolana salvado-

rensis Schuster, 1954, and E. koepckei Bott, 1954, were synony-

mized with E. hraziliensis by Glynn et al. ( 1975), a synonymy that

we first doubted but after a lengthy analysis of several hundred

specimens finally came to concur with. In addition, 2 species are

known from the temperate northeast Pacific: E. linguifroiis and E.

chilloni [= E. kincaidi (Hatch. 1947); = E. vancoiiverensis (Hatch.

1947); = E. japonica Richardson. 1912]. Three species have been

reported from the temperate southeast Pacific: E. chilensis, E.

hirsuticauda. and E. inonodi. Bruce ( 1986a) listed 12 species world-

wide, overlooking E. carangis and E. hirsuticauda. inadvertently

attributing authorship of E. chilensis to Menzies. and omitting E.

inayana and E. brazdiensis from the eastern Pacific fauna. Menzies

( 1962a) incorrectly noted the type species as E.xcirolana chilensis.

World list of species. —

E. affinis (Jones. 1971 ). Kenya.

E. arinata (Dana, 1833). Argentina and Brazil.

E. bicornis Kensley. 1978. South Africa.

E. hraziliensis Richardson. 1912. Caribbean to Brazil; tropical

eastern Pacific.

E. chamensis Brusca and Weinberg. 1987. Panama.

E, chilensis Richardson. 1912. Chile.

E. chiltoni (Richardson. 1903). Japan. Taiwan, Hong Kong; in

eastern Pacific. British Columbia to California.

£. geniculata Jones. 1 97 1 . Kenya.

E. hirsuticauda Menzies. 1962. Central southern Chile.

E. latipes (Barnard. 1914. as Pontogeloides latipes; includes

Excirolana carangis (Van Name. 1920)]. West and South Af-

rica.

E. ///i,t;"(7'"".s (Richardson, 1903). California.

E. inayana (Ives, 1891). Caribbean and tropical eastern Pa-

cific.

E. inonodi Carvacho. 1977. Chile.

E. natalensis (Vanhoffen, 1914). Southern Africa and Mada-

gascar.

E. orientalis (Dana, 1853). Indo-West Pacific, from Madagas-

I.

2

3.

4.

3.

6.

7.

8.

9.

10.

II.

12.

13.

14.

50

R. C. Brusca, R. Wetzer, and S. C. France

f^J

1-iiff

j-y-jtjfjjj

'"),

Figure 35. E\cirol(iiui of the tropical eastern Pacific, dorsal views: A. E. mayana (Mexico, Sonora, Puerto

Penasco, 17 June 1972), adult female. B, E. mayana (Pacific Panama, Farfan Beach, 11 Feb. 1985), juvenile

female. C, E. braziUensis (USNM 43655), holotype, male (Brazil). D, E. braziliensis (AHF Cat. No. 2000-01 ),

holotype morph (Mexico, Sinaloa, Mazallan). E, E. chamensis (LACM Type No. 3013), holotype. male (Pacific

Panama).

car to tropical Australia, Philippines, Japan, and India (in-

cludes Cirolana bomhayensis Joshi and Bal. 1959).

Key to Tropical Eastern Pacific Excimlana Species

Antennular peduncle articles 1 and 2 not fused (3 free peduncu-

lar articles); antennae of adults brushlike (anterior margin of

proximal flagellar articles highly setose); pleotelson"s posterior

border round or subacute, with two small, terminal (subinedial)

marginal spines; coxal plates with oblique groove .. E. mayana

Antennular peduncle articles 1 and 2 fused; antennae not

brushlike; pleotelson's posterior border round, without marginal

spines; coxal plates smooth 2

Mandibular palp of 2 articles; endopod of pleopod 5 divided;

with large stellate chromatophores; interocular distance much

greater than one eye width; length to 5 mm E. chamensis

Mandibular palp of 3 articles; endopod of pleopod 5 not divided

(but with shallow or deep lateral incision); stellate chromato-

phores lacking or minute; interocular distance usually equal to

width of one eye; length to 9 mm E. braziliensis

Tropical Eastern Pacilii: Cirolanidae

51

Figure 36. EMiniliiiin hraziUensis. Two syntypes of Ciwhina salvadorensis Schuster. 1954 (from El Salvador): A, holotype morph. B. fossorial morph.

Excirolana braziliensis Richardson, 1912

Figs. 35C. D. 36-48

Excirolana braziliensis Richardson, 1912b: 203. Nierstrasz 1931: 149.

Wade 1967: 512. Lemos de Castro and Silva Brum 1969: 7. Glynn. Dexter,

and Bowman 1975: 509. Brusca 1980: 227. Carvacho and Haasmann 1984:

16. Brusca and Iverson 1985: 31. Zufiiga et al. 1985: 9. Weinberg and

Starczak 1988: 296; 1989: 143. Kensley and Schotte 1989: 150. Schotte et

al. 1991:255.

Excirolana braziliensis (in part). Dexter 1976: 479; 1979: 543.

Cirolana koepckei Bott. 1954: 107. Carvacho 1977: 30.

Cirolanu mayana (in part). Dexter 1972: 449.

Cirolana salvadorensis Sc\\u^lef, 1954: 105. Dexter 1974: 51.

Cirolana n. sp. Gonzalez-Liboy, 1971 (unpublished master's thesis).

Type material examined. — (1) Male holotype (USNM 43655)

of Excirolana hraziUensis: Brazil, off Cape St. Roque, 12 ni; RA^

Albatross Sta. 2758; 16 Dec. 1887. (2) Paratypes (SMF 3606) of

Cirolana koepckei: Peru, Strand de Barranca; 1952; "holotype

morphs" (includes P-VII acute and blunt spine/dactyli forms; see

below); 9 specimens. (3) Syntypes (SMF 3601) of Cirolana

salvadorensis: El Salvador, Dept. La Paz, Los Blancos; 17 Oct.

1952; coll. O. Schuster; this portion of the syntype series contains

"holotype moiphs" (P-VII acute and blunt spine/dactyli forms),

"fossorial morphs," and "transitionals" (8 adults, 4 juveniles); 12

specimens.

Other material examined. — (unless otherwise indicated lots are

"holotype morphs" only — see below). Atlantic specimens: (4) Bra-

zil, Rio de Janeiro, Copacobana Beach; USNM 86335; Nov. 1943;

coll. Dr. Carlos Moreira; 1 specimen. (5) Panama. San Bias Islands;

SDNHM; 19 Feb. 1980; coll. James G. Morin; 50-(- specimens. (6)

Panama. Shimmey Beach; USNM 227013; 1985; coll. James

Weinberg; 8 specimens.

Gulf of California specimens: (7) Baja California, San Felipe;

SDNHM; 26-30 Mar. 1951; coll. W. Evans and party (W-51-84);

(includes both P-VII acute and blunt spine/dactyli forms); 10 speci-

mens. (8) Baja California, San Felipe area (Campo Ensenada). ca.

31° N, 1I4°48' W. collected with hand net in 0-1 m, over sand

bottom; (DFH 67-5 and 6), Ace. No. BI-67-2, CAS Cat. No. C-

3924; 1967; coll. D. F. Hoese; (includes both P-VII acute and blunt

spine/dactyli forms); 3 specimens. (9) Sonora, El Golfo de Santa

Clara, taken from grunion during grunion run/spawn; AHF Cat. No.

2056-01; 15 Jan. 1975; coll. N. Moffat and D. A. Thomson; I

specimen. (10) Sonora, Puerto Pefiasco. night lighting; SDNHM;

winter, late 1970s; coll. J. R. Hendrickson; 13 specimens. (11) Baja

California, Angel de la Guarda Island, swarming at water's edge;

AHF Cat. No. 2287-01; 1 Apr. 1972; coll. L. T. Findley; 18 speci-

mens. (12) Baja California, San Francisco Island, night lighting,

bottom sand with red and green algae, 16 m; SDNHM; 24 Feb.

1985; coll. Hank Chaney; 2 specimens removed from this lot for

52

R. C. Brusca, R. Wetzer, and S. C. France

Figure 37. Excirolana hrazUieiisis ( from Mexico. Baja California Sur. Isia

San Francisco: 24 Feb. 1985. coll. H. Chaney), scanning electron micro-

graphs. A, B, frontal views showing fused rostrum and frontal lamina and

cuticularsensillaeinpits (A. 74x. B. 148x1. C. pleotelson and uropods.37x.

SEM by R. C. Brusca; 20+ specimens. ( 13) Baja California, Pulpito

Point, 1 mi. offshore, night lighting. 2 11 5-22 15 hrs.; SDNHM; 22

June 1976; coll. R. H. Behrstock; 1 specimen. ( 14) Baja California,

Palmas Bay, N. of Los BaiTiles. Hermosa Beach, in front of trailer

park; EWI 81-7, SDNHM; 30 June 1981; coll. E. W. Iverson; 20-i-

specimens. (15) Sonora. 9 mi. S.E. El Tornillal (ca. 25 mi. S.E. El

Golfo de Santa Clara), water temp. 21.2' C. air temp. 20.4°C; UA

fish collection Cat. No. 69-80. SDNHM; 21 Mar. 1969; coll. K.

Muench; 3 specimens. ( 16) Baja California. Chivato Point, in water

over sand beach; SDNHM; 12 Aug. 1976; coll. R. Brusca, D. Perry;

3 specimens. (17) Sonora, Guaymas area, near San Carlos Bay, "in

tidepools, with Conns sp."; SDNHM; 9 Apr. 1974; coll. M.

Wicksten; 1 specimen. (18) Sonora, ca. 14 mi. E.S.E. of Guaymas,

Estero Yasicuri, 27°51.5' N, 115°38.5' W; SDNHM A.0115; 13

Aug. 1988; coll. L. T. Findley; 1 female with oostegites. ( 19) Baja

California. Evarislo Point, night light over soft bottom; LACM; 16

Mar. 1985; coll. H. Chaney; 5()-t-. (M) Baja California. Agua Verde

Bay, night light over sand and algae 9 m; LACM; 27 Feb. 1985;

coll. H. Chaney; 25 specimens. (21) Baja California, Agua Verde

Bay, night light over 15 m sand bottom; LACM; 15 Mar. 1985; coll.

H. Chaney; 6 specimens. (22) Baja California, Los Frailes, sandy

beach; USNM 150033; 27 Jan. 1972: coll. D. Dexter; 50-(- speci-

mens. (23) Baja California, Los Frailes. night light over 18 m;

LACM; 17 Feb. 1985; coll. H. Chaney; 2 specimens. (24) Baja

California. Cabo San Lucas; R/V Albatmss. USNM 69565; 1911;

50+ specimens. (25) Baja California, Pichilinque Bay (near La

Paz): night light; R/V Albatross Sta. No. A-5767. USNM 69710;

1911; (includes both P-VII acute and blunt spines/dactyli forins); 2

specimens. (26) Baja California, S.E. side Carmen Island, night

lighting; R/V Albatross Sta. No. A— 5767, USNM 69699; 1911;

200+ specimens. (27) Baja California. S.E. side Carmen Island; R/

V Albatross Sta. No. A-5761. USNM 69700; 191 1 ; 30+ specimens.

(28) Sinaloa. Estero El Verde (near Mazatlan); AHF Cat. No. 2000-

01; (P-Vll blunt spine/dactyli form); 9 specimens.

Central Eastern Pacific specimens; (29) Costa Rica, Quepos.

Cocal Beach, sand beach; USNM Ace. No. 309774, USNM 1 50034;

27 Feb. 1971; coll. D. Dexter; label in jar reads "Cirolana

salvadorcnsis Schuster, 1954; found in C. braziliensis vial"; in-

cludes "holotype morphs" (both P-VII acute and blunt spine/dactyli

forms), "fossorial morphs," and "transitionals"; 50+ specimens.

(30) Panama, Naos Island. Boy Scout Beach and Lab Beach, sand

beaches; USNM 227012; 1985; coll. Jaines Weinberg; ca. 20 speci-

mens. (31) Panama, Naos Island. Boy Scout Beach; SDNHM; Feb.

1986; coll. James Weinberg; 6 specimens. (32) Panama, Naos Is-

land. Lab Beach, sand beach; SDNHM; Feb. 1986: coll. James

Weinberg; 6 specimens. (33) Colombia, Depto. Valle de Cauca,

Juanchaco Beach, sandy intertidal; SDNHM: 7 Mar. 1987: coll. G.

A. Ramos: 2 specimens. (34) Ecuador, Puerto Mono, beach transect;

SDNHM; 1 June 1975; coll. B. A. F Hammond; 27 specimens. (35)

Ecuador. Salinas, sand beach; USNM 155475; coll. B. A. F.

Hammond; 20+ .specimens. (36) Ecuador; USNM Ace. No. 327410;

P-PT-Il: accessioned 31 May 1975; includes "holotype morphs."

"fossorial morphs," and "transitionals": 10 specimens. (37) Chile,

Concepcion. Playa Celulosa. sand beach, intertidal; 27 Mar. 1979;

coll. O. Aracena et al. ("Univ. Concepcion Pollution Study

Project"); 50+ specimens.

Description of male. — Cephalon somewhat immersed in

pereonite 1. Eyes large but variable in size, mean interocular width

usually about same as width of one eye, but variable from less than

to slightly greater than width of one eye (Figs. 35C, D). Antennule

and antenna lengths vary with age (at least in Pacific specimens);

Glynn et al. (1975) noted, "antennule shorter (with 6-12 flagellar

articles) than antenna ( 10-12 articles) up to a body length of 3^

mm: between 3 and 4 mm body length the antennules are com-

monly subequal in size; in individuals exceeding 5 mm in length.

Tropical Eastern Pacific Cirolanidae

53

Figure 38. Excirokma bnizitiensis. A-C, antennules: A. hoiotype (USNM

43655), male. B, syntype of Cimlaiui salvadorensis (from same specimen as

Fig. 42A; hoiotype morph). C, from Cimluiui siilvudorensis syntype (same

specimen as Fig. 42B; fossorial morph). D-F, antennae: D, hoiotype (USNM

43655), male. E, syntype of Cirolana salvadorensis (same specimen as Fig.

42A; hoiotype morph). F, from Cirolana salvadorensis syntype (from same

specimen as Fig. 42B; fossorial morph).

the antennule is longer (15-20 articles) than the antenna (11-16

articles): in some small individuals the antennules may vary from

longer, shorter or equal to the antennae." Antennular peduncle

articles I and 2 fused together; peduncle article 3 much wider than

but subequal in length to first flagellar article; first flagellar article

and all others except terminal article with distal aesthetascs (hoio-

type is in poor condition and missing many setae) (Figs. 38A-C).

Antennal peduncle 4-articulate. article 4 about twice length of 3; all

antennal articles with simple setae, those of peduncular articles may

be quite long (Figs. 38D-F). Mandible with innermost incisor cusp

variable in size but always larger than others; spine row with 11-15

spines; molar process with 1 8-23 acute marginal spines and simple

distal setae; palp 3-articulate, middle article longest, with setae on

middle and distal articles (setae missing from hoiotype) (Fig. 39A).

Maxillule's medial lobe with 3 large circumplumose spines and a

small subapical stout circumplumose seta; lateral lobe with 11-13

stout barbed spines (Fig. 39B). Maxilla's medial lobe short, with

plumose setae, simple setae, and short hairs; lateral lobe with

simple and comb setae as figured (Fig. 39C). Maxillipedal endite

with 5 or 6 plumose setae and one large coupling spine; palp 5-

articulate, all articles with simple setae as figured (Fig. 39D).

Pereon and pleon margins evenly convex. Coxal plates usually

visible dorsally on IV-VII, but visibility varies among specimens

(Figs. 35C, D) Coxae smooth, without carinae or grooves. Pereo-

pod I with stout acute spines and simple setae as figured; basis with

up to 6 long thin simple setae on distal inferior margin; ischium

with up to 4 simple setae on superior distal angle; merus expanded

as distal superior lobe, with up to 5 long setae and with 4-9 spines

on inferior margin; carpus short, subtriangular, with 2 or 3 spines on

inferior margin; propodus with 3-6 spines on inferior margin and 2

long comb setae on distal margin (often broken off); dactylus

elongate, nearly as long as propodus, and with comb setae at base of

unguis (often broken off) (Fig. 40). Pereopod III with simple spines

as figured; basis with up to 8 long simple setae on distal inferior

margin; ischium with short and long spines on inferior margin and

distal row of long submarginal setae on superior margin; merus

expanded into a large scoop-shaped distal lobe on superior margin

with stout spines and setae as figured; carpus with short and long

spines on inferior margin as figured; dactylus subequal in length to

propodus (Fig. 41). Pereopod VII with many spines and long simple

setae as figured (Figs. 42—45); number and shape of spines and

shape of articles highly variable (see below); basis with 1 or 2

spines and many long thin simple setae on distal inferior margin;

ischium long, longer than merus or carpus, usually longer than

propodus (except in certain cases, see below), and with 1 or 2 spine

clusters on inferior margin and numerous spines on distal margin;

merus and carpus with cluster of 1-3 spines on inferior margin and

numerous spines on distal margin; propodus usually with one clus-

ter of spines on inferior margin and long spines on distal margin (or

greatly elongated and modified as described below); dactylus short;

54

R. C. Brusca. R. Welzer, and S. C. France

Figure 39. Excirokma hiaziliensis (AHF Cat. No. 2000-01 ). female, hololype niorph. A. mandible

(left). B, maxillule (left). C, maxilla (left). D. maxilliped (left).

unguis typically acute (but see below). Penes small, ovate, with

senate outer margins; set close together (Fig. 471).

Pleon with 5 free pleonites, but first often partly covered by

pereonite VII (Figs. 35C, D). All pleopods with lobe on lateral

margin of peduncle, bearing a simple spine (often broken off); 3

coupling spines and several plumose setae on medial margin of

pleopods 1^ (plumose setae often broken off); exopod of pleopods

3-5 with short lateral incisions; endopod of 5 with ruffled lateral

margin (Figs. 46, 47). Appendix masculina of male short and stout,

articulated, arising about 0.25 distance up from base but not reach-

ing tip of endopod (Figs. 46F, G. 47F, G). Uropodal exopod reaches

or extends beyond posterior margin of pleotelson (Figs. 35C, D,

37C); endopod reaches or falls slightly short of posterior margin of

pleotelson; endopod with 2 apical spines and a deep notch on lateral

margin; e.xopod with 3 clublike apical spines; both rami (and pe-

duncle) with PMS as figured (Fig. 48). Posterior border of

pleotelson broadly rounded; dorsal depressions rounded and con-

nected by transverse ridge (Figs. 35C. D).

Variulions in leg morphology. — The exact numbers of spines

and setae on the pereopods. and the shape of pereopod VII, are

highly variable. In some cases this variability is clearly due to

damage (setae and spines being broken off) or probably to simple

genetic polymorphism (e. g., the number of spines in any given

spine row or spine cluster). In the case of pereopod VII, however,

the variations are dramatic, as noted below. The key (stable) fea-

tures of pereopod I are the lobed merus, subtriangular carpus, long

dactylus, apical comb setae on the propodus. and strong spination

of the inferior margin in general; variations in spine numbers can be

seen in the Fig. 40. The key (stable) features of pereopod III are the

row of submarginal simple .spines on the superior or distal rnargin

of the ischium, the large apical lobe on the superior margin of the

merus, and the long spines of the inferior margin of all articles

(Fig. 41).

Pereopod VII is enormously variable. This variation was briefly

noted by Glynn et al. (1975), who figured the two most extreme

forms of these legs but did not discuss them. In the holotype (Fig.

45 A ) and the majority of material we have examined (e.g.. Figs. 40,

41, 42A, B, 43A, B, 45A), the inferior margins of the ischium,

merus, carpus, and propodus bear long acute spines, the dactylus is

large and acute, and the propodus is about 1.5 times the length of

the dactylus, with one medial spine cluster on the inferior margin.

Most of the spines on pereopod VII bear a single sensory sensillum

arising from an inner neural core. We refer to individuals with these

types of seventh legs as "holotype morphs." In some specimens the

spines and dactylar unguis appear to be eroded, rough, and blunt-

tipped, and the spines may lose the sensory sensillae (e. g.. Figs.

42C, D, 43G, H, 458). In one premolt individual with blunt spines

and blunt ungui the new cuticle can be seen beneath the old outer

cuticle, and the blunt-tipped ungui were apparently to be replaced

with typical acute-tipped ungui (Fig. 45C). This suggests that the

bluntness of spines and dactyli on pereopod Vll of some individuals

may be due to simple mechanical erosion, perhaps as a result of

digging in sand. The common occurrence of individuals with both

the acute-spine/acute-dactyli and blunt-spine/blunt-dactyli mor-

phologies in single samples (see Material Examined) further sug-

gests an environmental-behavioral rather than a genetic explana-

tion for this variation. This hypothesis could be tested in laboratory

cultures.

Another different and dramatic variation seen in pereopod VII is

a marked alteration in the shape of the articles themselves. In the

holotype, all "holotype morphs," and most blunt spine/blunt dactyli

specimens the leg articles are relatively narrow and symmetrical

Tropical Eastem Pacific Cirolanidae

55

Figure 40. Excirotana braziliensis, pereopod 1. posterior surfaces. A {Panama. Naos Island, "Lab Beach"),

holotype morph (left). B (USNM Cat. No. 150034), fossorial morph (left). C (USNM Cat. No. 150034),

holotype morph (left). D (AHF Cat. No. 2001-01 ) details of spines, female. E (USNM 43655). holotype (left).

(e.g.. Fig. 36A). However, in some samples are individuals in which

the seventh legs are greatly enlarged, the ischium, merus, and

carpus become markedly broadened, expanded, and dislally scoop-

shaped, the propodus becomes enormously enlarged and elongated,

reaching 10 limes the length of the dactylus, and the dactylus is

reduced and essentially vestigial. Figures 36B, 42C, D, and 45C

illustrate these variations. For lack of a better term, we refer to these

individuals as "fossorial morphs" (see Material Examined). Figures

43-45 show some intermediate stages in a "transition" from the

"holotype" morphology of pereopod VII to a fully "fossorial" mor-

phology. We refer to such intermediate individuals as "transitionals"

(see Material Examined). In transitional individuals the dactylus

varies from large and acute to somewhat reduced, the propodus is

elongated (2-3 times the length of the dactylus) and bears 2 medial

spine clusters along the inferior margin (rather than a single medial

spine cluster, as in the holotype morph. or 3-5 spine clusters, as in

the fossorial morph). and the merus and carpus vary from weakly

expanded to slightly scoop-shaped. There are mixed morphs in

many samples (see Material Examined), and some individuals even

have one leg morphology on the left side and another on the right.

We are unable to discern any relationship to age. size. sex. or

.season in these variations of the seventh pereopod. nor do any other

aspects of the animals" morphology appear to be correlated to these

leg variations. Thus we suggest that the different leg morphologies

may arise during the life history of a single individual, and that the

variations reflect intraspecific polymorphism, rather than repre-

senting a group of closely related or sibling species. The occurrence

of mixed morphs in single samples (e. g.. the syntype series of

Cirolana scilvadorensis). the lack of relationship to size or sex, the

presence of what appear to be transitional individuals, and some

individuals with one leg type on the right and another on the left, all

suggest to us that the modifications of the seventh pereopod may

develop in response to ecological factors, rather than being geneti-

cally determined or fixed throughout the life of an individual. The

seventh leg of the fossorial moiph is strikingly similar to the dig-

ging legs of certain burrowing amphipods of the Haustorioidea,

such as Urohaiislorius Sheard and Pranlinus Barnard and Drum-

tuond (Barnard and Karaman 1991). Because ExciroUiiui brazili-

ensis is also a burrower. it seems reasonable to hypothesize that the

modificaitons of the seventh leg reflect a stage in the life history of

this species specialized for intensive burrowing, hence our coining

the term "fossorial morph."

56

R. C. Brusca, R. Wetzer, and S. C. France

Figure 41. Exciwlanu braziliensis. pereopod III (right): A (Mexico. Baja California. San Felipe), holotype morph.

B (USNM 43655), holotype, male. C (Pacific Costa Rica, Quepos. Playa Cocall. holotype morph. D (Mexico, Sinaloa,

Mazatlan), holotype morph. E, syntype of Cimluiui salvculorensis (female), fossonal morph (from same specimen as

Fig. 42B).

Female. — Similar to male. Females of this species are capable

of developing on pereopods Il-IV small oostegites that are very

thin and tightly pressed against the sternum.

Size. — To maximum length of 1 1 mm.

Disiribiition. — Atlantic coast: Gulf of Mexico and Caribbean to

Uruguay. Pacific coast: Gulf of California to central Chile. Littoral

to at least 16 m, probably somewhat deeper

Remarks. — E.xcimlana hrazilien.'iis is a widespread amphi-

American species. On the Pacific coast it is the most common

species of the genus living in the sand just below the water line. Our

attempts to resample this species at the type locality, at the now

heavily trafficked beaches near Rio de Janeiro (December 1988),

Tropical Eastern Pacific Cirolanidae

57

Figure 42. Exciwlnna hriiziliensis. extremes of morphological variation in

pereopod VII. A, B (Pacific Panama, Naos Island, "Lab Beach"), anterior (A) and

posterior (B) surfaces of left pereopod VII from holotype morph. C, D (Pacific

Costa Rica, Quepos, Playa Cocal ), antenor (C, nght) and posterior (D, left) surfaces

of pereopod VII from fossorial morph.

from Copacabana Beach south to Mangaraiiba, met no success.

Collections in Uruguay, between Montevideo and Punta del Este,

also failed to recover E. hraziliensis. although a different,

undescribed ExciroUmu was found on a sand beach at Maldonado.

just south of Punta del Este.

Weinberg and Starczak (1988, 1989) measured morphological

variation in populations of £. hra:iliensis on both the Atlantic and

Pacific coasts of Panama and South America. They found consider-

able morphological divergence along coastlines as well as between

oceans, although this divergence was greatest between populations

from opposite oceans. It is important to note that the ecological

literature has consistently confused (or overlooked) the three east-

ern Pacific species of E.xcirohina. whose ranges overlap.

Exciwiana chainensis Brusca and Weinberg,

Figs. 35E, 49-5 1

1987

ExiiriiltiiHi ihiimfii.si.s Brusca and Weinberg, 1987: 1 1.

Type mulerial examined. — ( I ) Male holotype (LACM Type No.

3013): Panama, near Panama City, polluted beach near old part of

town by National Theater; 15 Dec. 1984; coll. J. Weinberg; length

4.3 mm. (2) Male paratype (LACM Type No. 3014): same locality

and collection as holotype; head broken from body. (3) Paratypes

(LACM Type No. 3015): Panama, near Panama City, Chame Point

Bay, fine sand beach; 25 Sept. 1984; coll. J. Weinberg; 5 adults

(lengths 3.1-3.6 mm) and 2 mancas (lengths 1.6. 2.3 mm).

Other material examined. — (4) Panama, near Panama City,

Chame Point; USNM 365595; coll. J. Weinberg; I specimen.

Description oj male. — Dorsal surface ornately pigmented with

stellate chromatophores distributed in band between eyes on

cephalon, in median row on pereon, and in lateral rows on pleon

(Fig. 35E). Eyes small, interocular distance greater than width of one

eye. Antennules longer than antennae; antennular peduncle 2-articu-

late (due to fusion of articles I and 2); article 3 small, not much larger

than first flagellar article (Fig. 50A). Antennal peduncle 4-arliculate,

articles 1-3 subequal. article 4 longest (Fig. 50B). Mandible with 2-

articulale palp (Fig. 49D). Maxillule's lateral lobe blunt, with about

10 stout spines; medial lobe with 3 long slender circumplumose

spines and 1 apical seta (Fig. 49A). Maxilla's lateral lobes with 3 and

6 comb setae, respectively; medial lobe with 8 simple and 3 plumose

setae (Fig 49B). Maxilliped with single coupling spine (Fig. 49C),

Pereopods with acute spines along inner margin and acute

dactyli (never with blunt or truncated spines or dactyli). Pereopod I:

ischium with several stout spines (Fig. 5()C). Pereopod VII: ischium

without lateral spines (Fig. 50E). Coxal plates smooth, without

carinae or grooves.

58

R. C. Brusca, R. Wetzer, and S. C. France

Figure 43. Excirokma braziliensis. morphological variation in pereopod VII. A. B (Mexico, Baja California. Punta Pulpito),

posterior (A. left) and anterior (B, right) surfaces of pereopod VII. C, D (Pacific Costa Rica, Quepos, Playa Cocal). posterior

(C, right) and antenor (D, left) surfaces of pereopod VII. E, F (Pacific Costa Rica), anterior surfaces of pereopod VII (E. right;

F. left). G, H (Mexico. Sinaloa, Mazatlan), posterior (G) and anterior (H) surfaces of left pereopod VII.

Pleonite 1 not hidden by pereonite VII (Fig. .35E). Pleopod 5"s

endopod fully divided (Fig. 5 IE). Pleotelson's posterior border

round, without marginal spines. Uropodal exopod nearly twice as

long as endopod, with 6 long, thin, simple, apical spines, 2 large

medial spines, and numerous very short medial spines; with plu-

mose setae on medial margin and apex (Fig. 5 IF). Uropodal

endopod extends barely to posterior margin of pleotelson; with 3

distomedial spines and PMS along distal inner and outer margins.

Flat surfaces of both uropodal rami covered with very fine setae.

Appendix masculina of males stout and short, arising off proximal

lamellar lobe of endopod, one third distance up base and not reach-

ing apex of endopod (Fig. 5 IB). (For expanded description see

Brusca and Weinberg 1987.)

Female. — Similar to male.

Tropical Easlern Pacific Cirolanidae

59

Figure 44. Excirolana braziliensis. morphological variation in pereopod VII. A-C (3 individuals from

Pacific Costa Rica. Quepos, Playa Cecal, USNM 1300.34): A. anterior surface (left). B, posterior surface (left).

C, anterior surface (left). D (Pacific Panama. Naos Island. "Boy Scout Beach"), anterior surface (right). E (AHF

2000-01), anterior surface (left). F (USNM 86335), anterior surface (right).

Size. — To maximum length of 4.3 mm.

Distribution. — We have seen specimens only from the littoral

region of Chame Point Bay (Panama City) and the Pearl Islands.

Panama. H. A. Garces (In Litt. ) reports the species at "National Theater

Beach" and "under the Bridge of the Americas," in Panama City.

Remarks. — E. cinimeiisis superficially resembles E. hrcizilieiisis

but can be quickly differentiated by its smaller eyes, distinct chro-

matophore pattern, 2-articulute mandibular palp, fully divided

endopod on pleopod 5, and acute spines on all pereopods,

E.xcimlana maxana (Ives, 1891 )

Figs. 35A, B, 52-58

Cirolami maxana Ives 1891: 186. Richardson 1901: ?I2; 1903: 87,

Moore 1902: 166.

E.xcirolana maxana Richardson 1912: 201b. Nierstras/ 1931: 149.

Lemos de Castro and Silva Brum 1969; 3. Schultz 1969: 174. Menzies and

Kruczynski 1983: 48. Bruce 1986a: 221. Kensley and Schotte 1989: 153.

^on-Excirolana maxana: see Menzies and Glynn 1968: 39, Fig. 15.

Dexter 1972: 449 (uncertain identification).

Type material. — None available (see below).

Material examined. — Gulf of California (Sonora) specimens:

( 1 ) Puerto Pefiasco, Cholla Bay. shelly sand, high tide zone;

SDNHM; II June 1971; coll. J. Kudenov; 1 specimen. (2) Puerto

Pefiasco, sand beach at high tide, in about 1 m; SDNHM; 16 June

1972; captured while biting swimmer (C. A. Brusca); coll. R. C.

Brusca; 1 specimen. (3) Puerto Penasco, sand beach in front of

"Garcia House," 0-1 ni (high tide); SDNHM; 17 June 1972; col-

lected biting bodies of R. C. and C. A. Brusca; 1 1 specimens. (4)

Puerto Penasco, sand beach in front of "Garcia House," on dead

fish; SDNHM; 17 June 1972; coll. R. C. Brusca; 30-i- specimens. (5)

Puerto Penasco, night lighting, collected with E.xcirolana

hraziliensis; SDNHM; winter, late 1970s; coll. J. R. Hendrickson; 1

60

R. C. Brusca, R. Welzer. and S. C. France

Figure 45. Excirotami braziliensis. morphological vanation in pereopod VII. A, holotype (right). B, synlype of

Cirolana salvadoiensis (hololype morph). C. syntype of Cimlami sahaclorensis (fossorial morph, female).

specimen. (6) Estero de los Seris (= Estero Vibora); SDNHM; 26

June 1975; coll. R. C. Brusca; 5 specimens. (7) New Kino,

Connolank Bay, rocky shore; SDNHM; 22 June 1966; coll. D. A.

Thompson; 1 specimen.

Gulf of California (Baja California |Norte|) specimens: (8)

Sargento Point, intertidal; SIO Ace. No. BI60-18. Cat. No. C3531;

4 Apr. 1960; coll. A. Flechsig; I specimen. (9) 10-15 mi. N. of San

Felipe, otter trawl; DGL 72053 1 -2, SDNHM; 3 1 May 1972; coll. D.

Lindquist; 1 specimen. (10) Canipo Enscnada (N. of San Felipe),

ca. 3 r N, 1 14° 48' W. 0-1 m, hand net and sieve; CAS Ace. No. BI-

67-2, Cat. No. C-3924; 22-23 May 1967; coll. D. F Hoese; 4

specimens. (11) Campo Coloradito (ca. 38 mi. S. of San Felipe),

attacked fish being cleaned in surf-line at high tide, also biting

swimmers; SDNHM; 15 June 1982; coll. P. M. Delaney; 20+ speci-

mens. (12) Campo Coloradito, rocks on loose sand and gravel, mid-

interlidal; SDNHM A.0139.2; 17 June 1991; coll. R. C. Brusca; 3

specimens. (13) Los Angeles Bay, attracted to German sausages at

water line. 0900 hrs.; SDNHM A.0034; 20 May 1988; coll. R. C.

Brusca and R. Wetzer; 100+ specimens. (14) Estero La Ramada.

31°17'N, 114 53'W;SDNHM;31 May 1972; coll. R G. Lindquist;

1000+ specimens. ( 15) 20 mi. N. of San Luis Gonzaga Bay, pebble-

cobble beach in front of ruins of El Almacen (across the channel

from El Muerto Island), taken by hand while swimming in 0.5-1 m

water; SDNHM; 23 Mar. 1972; coll. L. T. Findley; 6 specimens.

(16) San Luis Gonzaga Bay, Okie's Landing, inside jellyfish

stranded on beach; SDNHM; Mar 1967; coll. E. Robinson; 4

specimens. (17) Los Angeles Bay, SE side of bay, coarse sand

beach; EWI-9, LACM; ?6 Aug. 1980; coll. E. W. Iverson; 10+

.specimens. (18) Los Angeles Bay; SDNHM A.0034; 24 May 1988;

coll. R. C. Brusca and R. Wetzer; 100+ specimens.

Gulf of California (Baja California Sur) specimens: (19) S. of

Chivato Point, from cove just north of Santa Ines Point, beach

screening in surf zone; SDNHM; 26 June 1981; coll. E. W. Iverson;

1 specimen. (20) Concepcion Bay. rinsed from algae washed ashore

on beach; SDNHM; 6 Dec. 1971; coll. R. Brusca and J. L. Barnard;

22 specimens. (21) Concepcion Bay; EWI-12, LACM; 19 Aug.

1980; coll. E. W. Iverson; 20+ specimens. (22) Concepcion Bay, El

Coyote Beach, in sand at wave-wash line; SDNHM; 9 Sept. 1985;

coll. J. P. Donahue; 30+ specimens. (23) Espiritu Santo Island,

"parasitic on fish." 1 6 m; CAS Ace. No. 0690; Sept. 1 97 1 ; coll. A. J.

Ferreira; I specimen. (24) Cerralvo Island, 24" 08.75' N. 109° 51.5'

W. "Pro-no\fish" sample, 5 m; RRG 1-37. SIO 61-259, Cat. No. C-

3905; 22 June 1961 ; 4 specimens.

Central eastern Pacific specimens: (25) Mexico, Baja California

Sur, Sebastian Vizcaino Bay. Miller's Landing. 28° 38' N. 1 14° 04'

12" "W. cobble reef; SDNHM; 12 Dec. 1945; 5 specimens. (26)

Costa Rica, Gulf of Nicoya, Morales Point, from gut of 178-mm

catfish ^Arms dowi). 0430 hrs.; SDNHM; 30 Oct. 1987; coll. W.

Swelistowski; 1 specimen. (27) Panama, small beach near Farfan

Beach, near entrance to Panama Canal; SDNHM; 11 Feb. 1985;

coll. J. Weinberg; 2 specimens. (28) Panama. Uva Island, pebble

beach, caught with rotten bonito left on beach for 3 minutes;

SDNHM; 9 Nov. 1973; coll. P Glynn; 200+ specimens. (29) Co-

lombia, Ensenada de Utria, Isia Playa Blanca, intertidal, in coralline

sand; SDNHM; 28 July 1986; coll. G. E. Ramos; 4 specimens.

Caribbean specimens: (30) Belize, Carrie Bow Cay, lagoon

trap; AC-CBC-314L SDNHM; June 1978; coll. A. Cohen; 20+

specimens. (31) Tobago. W. of Pigeon Point, beach debris; USNM

Cat. No. 15 1601; 4 Apr 1959; 3 specimens.

Descriplioii of male. — Cephalon not immersed in pereonite I;

eyes moderate in size, interocular distance greater than width of one

eye (Figs. 35A. B). Antennules extend to pereonite I or II; peduncle

3-articulate. all articles with simple setae at outer distal corner;

articles I and 2 with short acute spines on inner margin; 9-14

flagellar articles, most distal articles with aesthetascs (Fig. 55A).

Antennae extend to pereonite III or IV; peduncle 4-articulate. first

article with acute spines on inner distal margin; flagellum with 16-

21 articles, as many as 10 of which, in adults, may bear long.

Tropical Eastern Pacific CIrulanidae

61

Figure 46. Excirulana braziliensis, A-E (USNM 86335), pleopods (right side) from a female collected at the type locality: A. pleopod 1 . B, pleopod

2. C. pleopod 3. D, pleopod 4. E, pleopod 5. F, G (USNM 15(X)34), second pleopods of two males: F. hololype morph (right side). G. fossorial morph

(left side).

conspicuous, anteriorly directed setae (see Remarks) arising froin

two separate setal tracks on each article (Fig. 55B). Mandible with

inner incisor cusp largest, outer cusp rounded; spines of spine row

fleshy and blunt; palp 3-articulate, middle article much longer than

proxmial or distal articles; middle and distal articles highly setose

as figured (Fig. 55C). Maxillule's outer lobe with about II stout,

barbed spines, largest with visible row of denticles; inner lobe with

3 large circumplumose spines and I small seta (Fig. 55E). Maxilla's

inner lobe with proximal plumose setae and stout simple distal

setae; outer lobe with simple and comb setae as figured (Fig. 55D).

Maxillipedal endite short, with 1 or 2 coupling spines and numer-

ous distal plumose setae; palp 5-articulate, with long simple setae

on ailicles 2-5 (Figs. 54D. 35F).

Pereon and pleon margin evenly convex. Coxal plates visible

dorsally on III-VII or IV-VII (Figs. 35A, B); coxae with oblique

grooves running from middorsal point to top of posterior angle.

Pereopod I with molariform spines on inferior margin of ischium

and merus as figured (Figs. 56A, 57D). Pereopod III with

molariform spines on merus only (Fig. 36B). Pereopod VII without

molariform spines but with numerous acute spines and simple setae

(Fig. 56C). Penes large, fleshy, lanceolate, set close together.

Pleon with 5 free pleonites, the first largely overlapped by

pereonite VII (Figs. 35A. B). All pleopods with lobe on lateral

margin of peduncle, bearing a simple spine at the base (Fig. 58);

62

R. C. Brusca, R. Wetzer, and S. C. France

Figure 47. Excirokma brcizilieiisis I. AHF 2000-01 ), pleopods from a female ( A-E) and a male (F. G); endopod and exopod separated, endopod on right.

A, pleopod 1 (right). B, pleopod 2 (right). C, pleopod 3 (right). D, pleopod 4 (right). E, pleopod 5 (right). F, pleopod 2 (right). G, appendix mascuMna of

pleopod 2 (right). H, couphng spine from pleopod 3, female, right. I, penes from male.

Tropical Eastern Pacific Cirolanidae

63

Figure 48. Excirolana braziliensis (AHF 2000-01 ) uropods from a male: A, left; B, nght.

pleopods 1-4 with 3 or 4 coupling spines and plumose setae on

medial margin of peduncle; pleopods 1 and 2 with PMS on both

rami as figured; exopods of pleopods 4 and 5 with lateral incisions.

Appendix masculina large and scythe-shaped, articulating

subbasally. about 0.2 distance from base, not quite reaching lip of

endopod (Fig. 38F). Pleotelson lateral margins somewhat convex,

posterior margin rounded, with PMS set in notches and with 2 very

small submedian apical spines (Figs. 33A, B. 57A). Both uropodal

rami extend beyond margin of pleotelson; peduncle with 1 large

spine on outer margin and 2 large and 2 small spines at outer distal

angle; inner margin with PMS as figured. Uropodal exopod with 3

medial, I apical, and I lateral (subapical) spines and PMS as

figured. Uropodal endopod with 5 medial, 1 apical, and one lateral

(subapical) spines and PMS as figured (Fig. 56D).

Female. — Similar to male. Brooding females may have arising

from coxae III, IV, and V well-developed oostegites that overlap in

midline; oostegites are very thin and lie tightly pressed against the

sternum, making them difficult to see.

Size. — To maximum length of 15 mm.

Disthhiiliim. — Florida to Venezuela (Atlantic) and northern

Gulf of California to Colombia (Pacific). £. nuiyana is a littoral and

shallow-water (to 16 m) subtidal species occurring in the same

general area as E. braziliensis.

Remarks. — Despite its distinctive appearance, Excirolana

nuiyana has been an enigmatic and often overlooked species. In his

original description, Ives (1891) noted that the most distinctive

feature of this animal is its brushlike antennae. Moore ( 1902) and

Richardson ( 1901, 1905) also relied on the dense antennal setae to

distinguish this species in their keys to Cirolana. We have found

this "antennal brush" to be absent in juveniles and only weakly

developed in young adults, a feature also noted in the original

description. In the material we examined, the smallest female with a

fully developed "antennal brush" was 7.8 mm long, the smallest

male 5.7 mm. E. mayana also varies considerably in the expansion

and contraction of dorsal chromatophores.

No adequate figures of E. mayana have been previously pub-

lished. Richardson (1905), in copying the drawings of Ives (1891)

and Moore ( 1902), appears to have accidentally copied figures of

the penes of Cymodocea caiuiata. labeling the drawing as Cirolana

mayana. Ives' ( 1891 ) drawings of this species are crude, especially

his depiction of spines and setae. His drawing of the pleotelson

shows a crenulate margin with 20-22 minute spines set in the

notches and long setae arising between these spines. His description

states that the pleotelson is "minutely crenulate on its posterior

border, with very short spines inserted in the notches." Ives' draw-

ings were from a small specimen (9 mm), and the lack of detail

64

R. C. Brusca. R. Wetzer. and S. C. France

|4^ fMf

Figure 49. Excirolami cluimensis (LACM Type No. 3014).

paratype. male: A, maxillule (right) B. maxilla (nght). C. maxil-

liped (right). D. mandible (left).

Figure 50. Excirolami chamensis (LACM Type No. .3014).

paratype, male: A, antennule. B, antenna. C, pereopod 1. D. pereopod III.

E, pereopod VII.

raises suspicion ubotil the accuracy of his illustrations and descrip-

tion. In the material we have examined, from both the eastern

Pacific and Caribbean, the pleotelsonic notches house long setae

and the only spines are a single minute pair at the apex. The

specimens identified as E. ma\ami by Kensley and Schotte ( 1989)

agree with our observations. The two apical spines on the pleotelson

are so small that it is very likely Ives overlooked them (neither are

they figured or mentioned by Kensley and Schotte 1989). Unless

Ives' type material can be located, we cannot be positive his ani-

mals and ours are the same. However, we are reasonably confident

they are, and we suspect that Ives simply misinterpreted the

pleotelsonic margin. Ives also stated, "Spines and bristles upon the

legs are not nun)erous." yet he illustrated the legs as being highly

spinose — which they are.

Menzies and Glynn (1968) listed this species from Puerto Rico

but overlooked Richardson's (1912a) genus Exciinlaiui. citing it as

Ciwhina mayana. However, judged by their description and fig-

ures, Menzies and Glynn clearly misidenlified their specimens

(their material was probably a species of Cirolami). Menzies and

Glynn also claimed that E. mayana is "probably a pantropical

cosmopolite" but provided no data in support of this statement.

Bruce (1986a) listed it as a possible Indo-West Pacific species but

gave no records. Menzies and Kruczynski (1983) included this

species in their key to Caribbean Flabellifera (incorrectly stating

that the pleotelson lacks the submedian depressions characteristic

of the genus) but did not describe or figure it. Dexter (1972) listed

Cirolami maxami as the most abundant organism on both Pacific

and Atlantic sand beaches in Panama. However, she later claimed

(Glynn et al. 1975) that her material had actually been E.

braziliensis, not E. mayami.

Remarkably, Richardson (1905) appears to have been the only

person to have previously reported this common species from the

Pacific. She noted in passing that she had been unable to distinguish

some "dried specimens from San Francisco Bay, Lower California

[presumably Bahi'a San Francisquito. Baja California], sent by Dr.

Ritter" from Caribbean specimens of E. mayana. Excirolana

mayana is one of the most abundant animals in the quiet bays of the

Gulf of California, where it often occurs in vast numbers on sandy

beaches. We have collected hundreds of specimens in a few min-

utes, simply by placing a dead fish or piece of meat at the water line,

whereupon the isopods soon swarm out of the water or sand to

begin consuming the bait. This is the largest Excirolami known

from the tropical eastern Pacific.

Ives (1891) based his description on three specimens taken in

1890 at "Port of Silam." Yucatan Peninsula, by the "Expedition in

charge of Professor Angelo Heilprin. sent by the Academy of

Natural Sciences of Philadelphia to investigate the natural history

of Yucatan and Mexico." Ives was a member of the expedition, but

he gave no indication where the type material was deposited and we

have been unable to locate it (it has not been found at the Academy

of Natural Sciences of Philadelphia, the USNM. or the BMNH).

Metacirolana Kussakin, 1979

Type species.— Cirolami japonica Hansen, 1 890, by subsequent

designation (Kussakin 1979: 212). Holotype at ZMUC.

Sxnonxinv. — Emended and subsequent to Bruce (1986a:31).

Mehiamlami. Kensley 1984a; 33. Brusca and Iverson 1985: 36.

Kensley and Schotte 1989: 153.

Description.— Uo&y 2.0-3.0 times longer than broad; posterior

pereonites and pleon sometimes with dorsal tubercles or carinae.

Eyes usually well developed, moderate in size; absent in some

species. Cephalon short with small to moderate rostral process.

Frontal lamina anteriorly broad and dilated, freely projecting, often

overlapping bases of antennules, often visible in dorsal aspect.

Tropical Eastern Pacific Cirolanidae

65

%fiMt

Figure 5 1 . Excirnluna chumensis (LACM Type No. 3014), paralype. male: A. pleopod I (left). B. pleopod 2 (left). C, pleopod .3 (left). D, pleopod 4 (left).

E, pleopod 5 (left). F, uropod (left).

66

R. C. Brusca, R. Wetzer, and S. C. France

Figure 52. Excirolana mayaiui. scanning electron micrographs: A, antennule aethetascs, 740x. B. antennule aethetascs, 1480x. C. aethestasc surface,

I4,800x. D, three setal types on antenna! peduncle, 1332x.

posteriorly narrowed; clypeus wider than long, with ventrally pro-

jecting median triangular blade; labrum about as wide (or slightly

narrower) but longer than clypeus. Antennule short, never extending

beyond pereonite I; peduncle 3-articulate; article 2 not articulating at

right angle to article 1 ; peduncular article 2 or 3 longest; flagellum

reduced, basal articles longer than broad. Antennal peduncle 5-

articulate; article 5 longest, proximal two articles may be partially

fused. Mandible with broad tridentate incisor and an additional small

accessory tooth on medial (inner) margin of right mandible; middle

lobe of left mandible usually low and bladelike; palp 3-articulate.

extending beyond incisor; spine row a well-developed rounded lobe

with long stout spines. Maxillule's medial lobe with 3 or 4

circumplumose spines, sometimes with reduced setulation, also

occasionally with 1 or 2 short simple spines; lateral lobe with large

stout spines, often barbed. Maxilla's medial lobe somewhat reduced

(but more developed than in Cirolana and Anopsilana) and truncate,

with bifurcate lateral lobe often reduced. Maxilliped slender; palp 5-

articulate; palp article 3 much wider and longer than article 4; endite

with 1 or 2 coupling spines and plumose setae.

Pereonite I usually short ( subequal in length to pereonite II ). All

pereopods ambulatory, less spinose and setose than those of most

other genera of the Cirolanidae. Pereopods I-III short; distal supe-

rior angle of ischium and merus not produced; carpus short, often

triangular. Pereopods IV-VII slender, longer than pereopods I-III,

with articles not markedly flattened. Penes flattened lobes, small to

moderate in size.

Pleon with 5 free pleonites; lateral margins of pleonite 5 not

overlapped or barely overlapped by pleonite 4. Pleopods 1 and 2

similar; peduncles subrectangular and broader than long, without

lateral accessory lobes; appendix masculina inserted (about one-

third distance from base) on endopod of male's pleopod 2. Exopods

of posterior pleopods completely or nearly divided by medial trans-

verse incision. Pleopod 5's endopod without PMS, with or without

lobe on proximomedial angle, and without plumose setae or cou-

pling spines on peduncle. Pleotelson and uropods with or without

marginal spines. Pleotelson apex rounded, truncate or subtriangular,

never indented. Uropodal peduncle inner angle acutely produced.

Remarks. — Metacirolana contains 28 species. The genus was

resurrected by Bruce (1 98 1 a) to house a group of reasonably distinc-

tive cirolanid species, including Metacirohina joaimae (Schultz,

1966) of California waters andM sphaeromifonnisi,\\din%tn. 1890),

a species reported from the Caribbean (Menzies and Glynn 1968)

and the Indian Ocean and western Pacific (Nordenstam 1946).

Menzies and Glynn (1968) and Bruce (1981a) suggested that the

latter species is circumtropical, but we have not found it in the

eastern Pacific.

Tropical Eastern Pacific Cirolanidae

67

Figure 53. ExciroUma mayana, scanning electron micrographs; A, cephalon, 74x. B, rostrum, 370x. C, anterior view of rostrum, 700x. D. rostral

setae. 192x.

Bruce ( 1986a) noted that Metacirohma can be identified by the

projecting clypeus. anteriorly dilated frontal lamina (often visible

in dorsal view), pleonal and mouthpart morphology, and the long

second article of the antennular peduncle. While in many species

the second peduncular article is long, in M. costaricensis. M. ca-

lypso n. sp., and M. joanneae (and possibly others) the third pedun-

cular article is the longest. However, none of these characters is

unique to the genus. A possibly unique feature of Metaciiolaim is

the small accessory tooth on the medial (inner) side of the right

mandibular incisor, giving the incisor a four-pronged appearance.

This condition occurs in at least M. calypso n. sp., M. costaricensis

Brusca and Iverson, 1985, M. basreiii Bruce, 1980, M. bicornis

(Kensley, 1978), and M. japonica (Hansen. 1890) (see Bruce

1986a). Many published drawings of Meracirolana mandibles show

this accessory tooth, but most authors have not indicated whether

the figured mandible is the left or right. Bruce (1980. 1986a)

apparantly confused the left and right mandibles in his figures and

descriptions.

Nierstrasz ( 1931 ) created the name Melacirokma for Ciiolana

japonica Hansen. 1 890. and CiroUma hanseni Bonnier. 1 896. How-

ever, he did not designate a type species, thus rendering his name

invalid (ICZN Article 1 3b). Kussakin (1979) was the first to desig-

nate a type species and thus stands as the valid author of the genus.

World list of species. —

M. agaricicola Kensley, 1984. Belize.

M. agiijae Miiller, 1991. Atlantic Colombia.

M. iinatolci Bruce, 1986. Queensland, Australia.

M. anocula (Kensley. 1984). South Africa.

M. aniaudi Kensley. 1989. St. Paul and Amsterdam islands,

southern Indian Ocean.

M. hasteni {Bvncc. 1980). Australia.

M. bicornis (Kensley, 1978). South Africa.

M. calypso n. sp. Galapagos Islands. Ecuador.

M. convexissinni (Kensley. 1984). South Africa.

M. costaricensis Brusca and Iverson. 1 983. Pacific Costa Rica.

M. fishelsoni (Bruce and Jones. 1978). Red Sea.

M. /;((//(/ Kensley. 1984. Cozumel (Mexico). Belize, and

throughout Caribbean.

M. hanseni (UonmtT. 1896). Kuropc.

M. japonica (Hansen, 1890). Japan, Australia. Tasmania, and

New Guinea.

M. joanneae (Schultz, 1966). California.

M. mbudva Bruce, 1981. Tanzania.

68

R. C. Brusca, R. Wetzer, and S. C. France

Figure 54. Exciwiana mayima. scanning eleclron micrographs: A, rostrum and frontal lamina,

96x. D, maxillipeds. 192x.

I78x, B, clypeus and labrum, 285x. C. buccal field.

17. M. menziesi Kensley. 1984. Belize.

18. M. miyamoltii Nunomwix. 1991. Japan.

19. M. nwiioilHiones. 1976). Aldabra.

20. M. nwoiii^ati Mtiller and Salvat. 1993. French Polynesia.

21. M. nana (Bruce, 1980). Australia.

22. M. pigmenlaia Muller and Salvat, 1993. French Polynesia.

23. M. po/i.s7 Jaume and Garcia, 1992. Balearic Islands, Mediterra-

nean Sea.

24. M. riobaldoi (Lemos de Castro and Lima, 1976). Brazil.

25. M. rotunda (Bruce and Jones, 1978). Red Sea to Tanzania.

26. M. rugosa (Bruce, 1980). Great Barrier Reef, Australia.

27. M. serrata (Bruce, 1980). Lizard Island, Australia.

28. M. sphaeromifonnis (Hansen, 1890). Florida, Caribbean Sea,

Indian Ocean, and possibly central/west Pacific Ocean.

29. M. spinosa (Bruce, 1980). Lizard Island. Australia.

Key to Tropical Eastern Pacific Metacirolana Species

1 . Pleotelson of male without dorsal tubercles or carinae; posterior

margin of pleotelson broad, convex, and strongly serrate: pereo-

pod I's merus with long acute spines on inferior margin;

uropodal endopod shorter than pleotelson M. calypso n. sp.

— Pleotelson of male with dorsal tubercles or carinae; posterior

margin of pleotelson narrow, subtruncate, and not serrate; pereo-

pod I's merus with squamate molariform spines on inferior

margin; uropodal endopod extends to pleotelson ape.x

M. costaricensis

Metacirolana calvpso n. sp.

Figs. 60C. 61,62

Type material examined. — Male holotype (LACM 84-287.1):

Ecuador, Galapagos Islands, near Wolf Island (approx. 1° 18'N,9r'

45' W), neuston tow, 0545-0610 hrs.; 12 May 1984: coll. R. J.

Lavenberg et al.; bottom depth not recorded by collectors but

estimated (R. Lavenberg. in litt.) as approximately 2000 m.

Description of male. — Entire dorsum with striking ornate chro-

matophore pattern (Fig. 60C). Cephalon length, from rostrum to

posterior margin. 1.5 times length of pereonite I: lateral margins

evenly convex, not forming subacute angles. Cephalon devoid of

dorsal tubercles or carinae: rostrum very short, rounded anteriorly

(Fig. 60C). Antennule not quite reaching posterior margin of

pereonite I; flagellum of 6 articles (Fig. 61 A). Antenna extends to

posterior margin of pereonite VII: flagellum of 14 articles (Fig.

6 1 B ). Frontal lamina narrow, almost quadrate posteriorly, expanded

anteriorly to partly overlap proximal antennular article (Fig. 6IC).

Tropical Eastern Pacific Cirolanidae

69

Figure 55. Excirokma mayaiui. A, B (Mexico, Sonora, Puerto Pefiasco), adult: A, antennule (left). B, antenna (left). C-F (Mexico, Baja California Sur.

Concepcion Bay), juvenile: C, mandible (right). D, maxilla (right). E, maxillule (right). F, maxilliped (right). All drawings from female specimens.

70

R. C. Brusca, R. Wetzer, and S. C. France

Figure 56. Exciwhma mayami (Mexico, Sonora, Puerto Penasco, 17 June 1972), adult: A, pereopod I (left). B. pereopod III (left). C, pereopod VII

(left). D, uropod(left).

Mandibular .spine row with 19 long thin spines; niolar process with

about 20 small marginal spines; terminal article of palp with simple

setae and 1 coinb seta; middle article longest, with simple and comb

setae (Fig. 6 ID). Ma.\illule"s lateral lobe with 12 stout spines,

largest spines armed with barbs; medial lobe with 1 small apical

seta in addition to the 3 circumplumose spines (Fig. 61E). Maxilla's

lateral lobes with 3 and 5 plumose setae, respectively; medial lobe

with 1 simple and 7 plumose setae (Fig. 61F). Maxillipedal palp

articles 4 and 5 are margined with plumose setae, other articles with

simple setae; left and right endites small, each with 1 coupling spine

and 4 plumose setae (Fig. 61 G).

Pereonites unequal in length, pereonite I longest; pereon widest

at pereonites IV and V. devoid of dorsal tubercles or carinae. Coxae

usually visible dorsally, extending beyond posterior margins of their

respective segments; coxa VII extends almost to posterior margin of

pleonite 2 (Fig. 60C). Pereopod I short, stout; superior distal angles

of ischium and merus each with 1 long seta; inferior margins of

merus. carpus, and propodus with acute spines as figured; inferior

margin of propodus also with 1 serrate spine; carpus short; dactylus

without small spine at base of unguis (Fig. 62A). Pereopod IV longer

than pereopod I. ambulatory, with simple and serrate spines and

setae as figured; dactylus without small spine(s) at base of unguis

(Fig. 62B). Pereopod VII quite long, ambulatory, with simple and

serrate spines and setae as figured (Fig. 62C). Penes large, about 6

times longer than wide, extended roughly 0.66 length of stemite.

Pleon broadest at pleonite 2. devoid of dorsal tubercles or

carinae. Pereonite VIFs coxae overlap lateral margins of pleonite 1 ;

pleonites 2-A expanded laterally (Fig. 60C). Pleopod rami with

PMS as figured (Figs. 62F-J). Pleopod 1 : peduncle's medial margin

with 4 coupling spines, lateral margin with 2 simple spines;

Triipical Eastern Pacific Cirolanidae

71

Figure 57. Excimkmu nuiyami. scanning electron micrographs: A, pleotelson. 74x. B. seta next to right depression on pleotelson.462x. C, seta in right

depression on pleotelson, 3330x. D. pereopod, y6x.

endopod width 0.68 times width of exopod (Fig. 62F). Pleopod 2:

peduncle's medial margin with 2 coupling spines, 2 plumose setae,

and many short simple setae; lateral margin with 1 simple spine;

endopod width 0.85 times width of exopod; appendix masculina

widest basally. tapering to serrate medial margin near pointed apex,

medial margin with many small setae, length 0.95 times endopod

length (Fig. 62G). Pleopod 3: peduncle's medial margin with 3

coupling spines and 1 plumose seta, lateral margin with 1 simple

spine; endopod width 0.72 times exopod width, exopod with short

marginal incisions (Fig. 62H). Pleopod 4: peduncle's medial mar-

gin with 3 coupling spines and 1 plumose seta, lateral margin with

1 simple spine and tuany short setae; endopod 0.73 times width of

exopod, exopod with complete medial incision (Fig. 621). Pleopod

5: peduncle smaller than peduncles of pleopods 1^, with 1 simple

spine and many short setae on lateral margin; endopod width 0.80

times exopod width; exopod with complete medial incision

(Fig. 62J).

Pleotelson with straight lateral tnargins. apex widely rounded

with sharply crenulate (saw-toothed) margin; without apical spines

but with PMS; dorsum without longitudinal carinae (Fig. 60C).

Uropods shorter than pleotelson, with small terminal notch on each

ramus, 5-7 simple setae arising from each notch. Uropodal exopod

0.70 times as wide as endopod, shorter than endopod. medial mar-

gin with 1 short spine and many PMS, lateral margin with 3 short

spines and setae. Uropodal endopod's medial margin with 2 short

spines and many PMS. lateral margin with 1 short spine and many

PMS. Uropodal peduncle with short apical spine, medial margin

with PMS (Fig. 60C).

Female. — Not known.

Size. — Small, holotype 5.2 mm long.

Disirihiiiion. — So far known from only the type locality, near

Wolf Island, the Galapagos Islands, Ecuador

Remarks. — This species is immediately distinguished from the

only other known eastern Pacific MetaciroUma (M. co.<ilaricensis)

by features noted in the key. It is known from only a single speci-

men collected in a neuston (surface) plankton tow near Wolf Island

in the Galapagos Islands. However, the distinct morphology of the

specimen warrants formal species recognition.

Etymiilogy. — This species is named after Calypso, daughter of

Occanus in Greek mythology. Just as this beautiful island isopod

charmed its describers, the charms of the island nymph Calypso

were so powerful they detained Odysseus seven years on his jour-

ney home from Troy.

72

R. C. Brusca, R. Wetzer, and S. C. France

Figure 58. Excimlana mayana. A-E (Mexico, Sonora, Puerto Peiiasco), pleopods of adult female: A, pleopod 1 (left). B, pleopod 2 (left). C, pieopod 3

(left). D, pleopod 4 (left). E, pleopod 5 (left). F (Mexico, Gulf of California, Cerralvo Island), adult male, pleopod 2 (left).

Metacirnlana costaricensis Brusca and Iverson, 1985

Figs. 60A, B, 63, 64

Metacirolami costaricensis Brusca and Iverson 1985: 36. Fig

Bruce 1986a: 222.

IID.

Type material examined. — Holotype (LACM 80-60.1. AHF

Type No. 801 M ard 15 paratypes (LACM 80-60.2): Co.sta Rica,

Guanacaste Province, Parque Nacional Santa Rosa, rocky littoral

approximately 1 km from mouth of mangrove estuary, ca. 1 0° 48'

N, 86° 57' W. formalin washes of rocks and turf algae, water

temperature 26° C, large surf; 26 Apr. 1980; coll. R. C. Brusca, A.

M. Mackey, M. Murillo, A. Dittle.

Tropical Eastern Pacific Cirolanidae

73

Figure 59. Metacirokma joannae (AHF 2250-1), scanning electron

micrographs: A. ventral view of frontal lamina, clypeus. and labrum, 1 18x

B, ventrally projecting clypeus, 370x.

Other material examined. — Central American specimens: (1)

Costa Rica, Puntarenas Province, just outside mouth of Gulf of

Nicoya. Playa Tarcoles, 9° 45' N. 84° 50' W, dark sand with scattered

rocky points, formalin washes of rocks and associated sediment,

onuphid polychaete tubes abundant, water temperature 29° C;

SDNHM: 22 Feb. 1980; coll. R. C. Bru.sca; 29 specimens. (2) Costa

Rica, Puntarenas Province, Panama Bay, "infauna, 2 meters marea

alto, transect #2 (Vallejo), station #2'"; SDNHM; 3 Feb. 1977; 2

specimens (gift of M. M. Murillo). (3) Panama, Culebra Island;

PWG-67, SDNHM; 23 Jan. 1967; coll. P W. Glynn; 1 female.

Galapagos specimens: (4) Just west of Isabela Island, 0° 15' 43"

S, 91° 26' 38" W, taken by hand; RA^ Anton Briiim Cruise 16; 25

May 1966; SEPBOP program; 9 specimens (Sta. ST66142); 3

males and 1 female (Sta. ST66141 ) (precise coordinates question-

able). (5) Wolf Island, on shore with green algae; Sta. 144. USNM

Ace. No. 128938; 11 Jan. 1934; 1 male. (6) South side of Pinta

Island, near Ibbetson Point, 0° 32' 25 " N, 90° 43.5' W, near national

park marker on shore, noon, low tide, large shallow tidepool, fixed

lava rock with loose lava stones and some sand, 0-1 ft. standing

water, from under loose rocks; LACM; 20 May 1984; coll. A.

Cohen; 6 specimens. (7) Tower Island, tidepool on north side of

Darwin Bay. 0° 1 8' 58" N. 90" 57' 23" 'W. lava rock in large pool 3 m

deep with deeper crevices, low tide, under loose rubble (rock and

dead coral) in exposed (low tide) channel at entrance to pool; AC-

GAL-27,LACM;21 May 1984;coll. A. Cohen; 2 specimens. (9) no

precise locality; AC-GAL-8-A. LACM; 1984; coll. A. Cohen; I

specimen.

Description of male. — Cephalon devoid of tubercles and cari-

nae; length from posterior margin to rostrum subequal to length of

pereonite I. rostrum moderate in si/e; lateral margins forming

subacute angles (Fig. 60A). Antennule short, reaching posterior

margin of cephalon; fiagellum of 3-6 articles, distal flagellar ar-

ticles compressed and short (Fig. 63A). Antenna reaching pereonite

II; peduncular articles 1 and 2 partially fused; fiagellum of 8-12

articles (Fig. 63B). Frontal lainina narrow posteriorly, expanded

and rounded anteriorly; anterior expansion overlaps basal articles

ol'antennules (Fig. 63C). Both mandibular incisors tridentate; right

incisor with accessory tooth; left incisor indistinctly tiidentate,

somewhat bladelike and without accessory tooth; spine row with 14

long thin spines; palp 3-articulate. terminal article with comb setae,

middle article longest and with simple and comb setae (Fig. 63D).

Maxillule's medial lobe with 1 small apical spine, in addition to the

3 circumplumose spines; lateral lobe with 10 stout spines, many

strongly barbed (Fig. 63E). Maxilla's medial lobe with 6 plumose, 1

large circumplumose, and 2 small simple setae; lateral lobes with 3

comb and 4 plumose setae, respectively (Fig. 63F). Maxillipedal

palp articles subrectangular, margins with many long simple and

comb setae; endite small, with 2 plumose setae and 1 simple seta,

with 1 or 2 coupling spines (usually 2) on both left and right endites;

small epipod present in both sexes (Fig. 63G).

Pereon widest at pereonites V and 'VI, devoid of dorsal tubercles

or carinae. Pereonite VII with short rounded process on posterolat-

eral margins. Coxae III-VI carinate, visible in dorsal view and

extending beyond posterior margins of their respective pereonites;

coxa VII large, extending to posterior margin of pleonite 5 (Fig.

60A). Pereopod I short and stout; distal margins of articles not

produced; inferior margin of merus with 3 very short blunt squa-

mate spines. 2 simple setae, and 1 small serrate spine; carpus very

short, inferior margin with 1 squamate spine. 1 serrate spine, and 3

simple setae; inferior margin of propodus with 3 large basally

serrate spines and simple setae; dactylus with I small simple spine

at base of unguis (Fig. 64A). Pereopod IV short, stout, ambulatory,

with simple spines and setae as figured (Fig. 64B). Pereopod VII

ambulatory with simple and senate spines and setae as figured (Fig.

64C). Penes about 3 times longer than wide.

Pleon broadest at pleonites 3 and 4. Pleonite 1 overlapped

laterally by pereonite VII, visible medially. Pleonite 5 with large

median tubercle on posterior margin (Fig. 6()A). Pleopodal rami

with PMS as figured (Figs. 64F-J). Pleopod 1: peduncle's medial

margin with 4 coupling spines, lateral margin with 1 seta; endopod

width 0.6 times width of exopod (Fig. 64F). Pleopod 2: peduncle's

medial margin with 4 coupling spines and 2 plumose setae, lateral

margin with I long and many short setae; endopod width 0.7 times

width of exopod; appendix masculina widest basally, tapering me-

dially and widening again near apex, length 0.8 times endopod

length (Fig. 64G). Pleopod 3: peduncle's medial margin with 4

coupling spines and 2 plumose setae; endopod width 0.74 times

exopod width (Fig. 64H). Pleopod 4: peduncle's medial margin

with 4 coupling spines and 2 plumose setae, lateral margin with 1

large spine and 1 short spine; endopod width 0.80 times exopod

width (Fig. 641). Pleopod 5: peduncle's lateral margin with 1 simple

spine; endopod width 0.82 times exopod width ( Fig. 64J ). Pleopodal

exopods 3-5 with complete or nearly complete medial transverse

incision.

Pleolelson subtriangular. lateral margins slightly concave near

truncate apex; dorsum with median longitudinal carina, fianked by

74

R. C. Brusca. R. Wetzer, and S. C. France

Figure 60. Meiacimlana of the tropical eastern Pacific, dorsal views: A, M. cosiaricensis (LACM 80-60. 1 ). holotype. male. B, M. cosuiricensis (LACM

80-60.2), paratype, female. C, M. calypsn n. sp. (LACM 84-287.1, GAL 84-10), holotype, male.

subniedian longitudinal carinue, carinae usually with tubercles;

pleotelson apex with 2 marginal spines and several simple setae

(Fig. 60A). Uropods with small apical notch on each ramus, 6 or 7

PMS arising from each notch. Uropodal exopod does not extend to

pleotelson apex, 0.66 width of endopod, medial margin with 2 large

spines interspersed with PMS. lateral margin with 2 small spines,

simple setae, and PMS. Uropodal endopod extends to pleotelson

apex; medial margin with 3 spines interspersed with PMS and some

simple setae; lateral margin with I spine and PMS (Fig. 64E).

Female. — Similar to male, except dorsal tuberculation on the

pleon and pleotelson is reduced or absent, the pleotelson's lateral

margins are slraighler and not as concave as in males, coxal plates

II-Vl are less visible in dorsal view, and pleonite 1 is not necessar-

ily hidden by pereonite VII (Fig. 6()B).

Size. — Small, to maximum length of 4.0 mm.

Distiihutiim. — Thus far recorded from Pacific Costa Rica,

Panama, and the Galapagos Islands. Material examined is, with the

exception of the wo Anton Bninn lots, all littoral. The Anion Brinin

Galapagos records suggest that the specimens were taken in deep

water, but the station numbers on label data do not occur in the

station lists of Chin et al. (1972).

Rentark.s. — An intertidal and shallow subtidal species, found in

rocky littoral areas with lurf-algac anil in dark sandy/rocky habitats.

Brusca and Iverson ( U)iS.S)did not atlcquately figure this species, so

we illustrate it completely here

Nohuoliuia Bruce, 1981

Type .Ki>ecie.\. — Ciroluno hiriipes Milne Edwards, I S40, by sub-

sequent designation (Bruce 1981a). Type specimens at the Museum

National d'Histoire Naturelle, Paris.

Synonymy. — Emended and subsequent to Bruce ( 1986a:52).

Natatolcma. Bruce 1985: 708. Brusca and Iverson 1985: 37.

Botosaneanu et al. 1986: 412. Kensley and Schotte 1989: 139.

Descripikm. — Body length approximately 2.5-3.0 times width;

dorsum smooth, without ornamentation. Eyes usually well devel-

oped, but absent or with reduced ommatidia in some species. Ros-

tral process minute or absent. Frontal lamina elongate, narrow, flat,

not projecting, length 3-4 times width; clypeus broad, wider than

long, flat (sessile); labrum narrower than clypeus. Antennular pe-

duncle and flagellum short, flagellum does not extend beyond

anterior region of pereonite I; second article not articulated at right

angle to first article (as in Eiinr/ice): peduncular article 3 longest;

flagellar articles compressed, basal anicles often fused. Antennae

much longer than antennules; peduncular articles 1-2 short, 3^

subequal, 5 longest. Mandible with broad tridentate incisor, medial

cusp often reduced on left mandible; spine row well developed as a

rounded lobe with stout spines. Ma.xillule"s medial lobe with 3 or 4

stout circumplumose spines and often a few small simple spines;

lateral lobe with 9-12 stout apical spines, often with minute subapi-

cal lateral spines or barbs. Maxilla with medial lobe short and

broad. Maxillipedal palp 5-articulate; endite short, with 1-3 cou-

pling spines.

Pereonite 1 longest. Posterior angles of coxae U-VII become

more acute posteriorly. Pereopodal dactyli often with small spine at

base of ungui; superior margins of ischium and merus of pereopods

l-lll strongly produced; pereopods IV-Vll longer than pereopods

Tropical Eastern Pacific Cirolanidae

75

Figure 61. Melacirolami calypso n. sp. (LACM 84-287.1, GAL 84-10). holotype. male: A,

antennule (left). B. antenna (left). C, frontal lamina, clypeus, and labrum. D. mandible (right). E,

maxillule (nght). F, maxilla (right). G, maxilliped (nght).

I-III and with abundant long setae; pereopods VI-VII with is-

chium-propodus flattened and provided with long setae; pereopod

VII with dense medial row of long setae along flat anterior surface.

Penes indistinct or represented by small flattened lobes.

Pleon of 5 free pleonites; pleonite I often patlially concealed by

pereonite VII; pleonite 5 completely encompassed by lateral mar-

gins of pleonite 4. Pleopod I "s exopod almost twice as wide as

endopod; pleopodal peduncles broader than long, lateral margins

without lobes or with weak lobes; rami similar; all pleopodal rami

with PMS, except endopod of pleopod 5, which has reduced or no

PMS; appendix masculina inserted basally or subbasally on

endopod of male's pleopod 2. Pleopod 3: peduncle's medial margin

without coupling spines or plumose setae; endopod with

proximomedial lobe. Pleotelson usually with abundant marginal

setae and spines. Uropodal peduncle's inner angle produced and

subacute; rami with PMS and usually spines; endopod usually

without notch on distal medial angle, except in Nanilnla/ui

variguberna.

Remarks. — Bruce (1981a) split Clrolana into seven different

genera, erecting three new genera, including Natatolana. He did not

describe or figure CiroUma hirtipes Milne Edwards, 1840, the type

species. Characters diagnostic of Ncitauikiiw include the glabrous

appearance and absence of sculpturing of the dorsum, short anten-

nules, flattened articles on the posterior pereopods, and the medial

row of long setae on the flat anterior surface of pereopod VII.

Similar-appearing genera are Dotichotana and Polholana.

Dolichokma has similar pereopods but differs in the form of the

frontal lamina, which has the ventral surface excavated and the

posterior margin produced into a venlrally projecting lobe (the

ventral surface of the frontal lamina is flat in Nalalolana). and in the

lack of PMS on pleopodal endopods (only the endopod of pleopod 5

is naked in NatatoUiiui). Polilohimi differs from Nalalolana in the

following ways: the bases of the posterior pereopods are less ex-

panded; the appendix inasculina arise subbasally, rather than ba-

sally as in Nalalolana (although in some species of Natatolana the

appendix masculina arises slightly above the basal position — see N.

76

R. C. Brusca, R. Wetzer. and S. C. France

Figure 62. Meracirolana calypso n. sp. (LACM 84-287.1, GAL 84-10), holotype. male: A, pereopod I (right). B. pereopod IV (right). C, pereopod Vll

(right). D, penes. E, dorsal view of uropod (right). F, pleopod 1 (right). G, pleopod 2 (right). H, pleopod 3 (right). I, pleopod 4 (right). J, pleopod 5 (right).

Tropical Eastern Pacific Cirolanidae

77

Figure 63. Metacirokma costaricensis (LACM 80-60.1), holotype, male: A. antennule (left). B.

antenna (left). C. frontal lamina, clypeus, and labrum. D, mandible (right). E, maxillule (right). F,

maxilla (right). G. maxilliped (right).

bowmani Bruce, 1986); pleopod Ts peduncle is subquadrate (it i,s

wider than long in Nalatolaiui); and the uropodal endopod has a

distolateral notch (absent in Natatolana. except in N. variguberna).

Natatolana, Politolana. and Dolicholana are part of Bruce's

(1986a) "Conilera genus-group," along with Conilera. OrpheUmu.

and Conilorpheus. Wetzer et al. ( 1987) discussed this group, pro-

vided a key to the genera, and removed Oncilorpheiis from it.

Natatolanii. with 38 described species, is the second largest

genus in the family, and it has the widest distribution of any cirolanid

genus, with more species known from temperate and cold waters

than in any other genus. Natatolana is primarily a shelf and slope

taxon, ranging from the shallow subtidal to about 2000 m, although

occasional specimens have been collected intertidally. Bruce ( 1 986a)

divided the genus into four "species-groups," the N. pellucida group.

the N. valida group, the A', albicaudata group, and the N. woodjonesi

group. The groups are distinguished froin one another by the basis of

pereopod VII, the posterolateral margin of pleonite 4, the pleotelson

dorsum, and the posterior margin of the pleotelson.

In all the species we have examined, it appears that the medial

lobe of the maxillule in species with the 3-spine configuration also

possesses a moderately to well developed protuberance on the

lateral margin, whereas species with the 4-spine configuration lack

this protuberance.

There are four Natatolana species in the eastern Pacific: N.

(7j//f;i.v/.v (Menzies, 1962a), M «aM//.v(Menzies and George, 1972),

A', californiensis (Schultz, 1966), and N. carlenae n. sp. Only the

last two are tropical.

World list of species. —

78

R. C. Brusca. R. Welzer, and S. C. France

Figure 64. Meu.nrotcma , »w„nr,-«vn (LACM 80-60.1 ), holotype. male: A. pereopod 1 (left). B. pereopod IV (left). C, pereopod VII (left). I), penes. K,

ventral view of uropod (right). K ple(.p..d I . G, pleopod 2. H, pleopod 3. 1. pleopod 4. J, pleopod 5.

Tropical Eastern Pacific Cirolanidae

79

Figure 65. Nululolwui of the tropical eastern Pacific, dorsal views: A, N. ciilifornieiisis (AHF 6048). holotype. male. B, N. carleiiiie n. sp. (USNM

252731), holotype, male.

1. N. alhicaiithiki (Stebbing. 1900). Australia. Philippines, and Islands, Indian Ocean.

Japan. 5. /V. arrirai/ffa (Holdich. Harrison, and Bruce. 1981 ). Queens-

2. N. amplociila Bruce. 1986. Kai Islands and the Arafura Sea. land, Australia.

Indonesia. 6. A'. (;/v(»()(/ Bruce, 1986. Victoria. Australia.

3. /v. (»i,?i(/(V Bruce. 1986. Queensland, Australia. 7. N. hokoUxxiizc. 1986. Queensland, Australia.

4. N. anopthatma (Kussakin and Vasina. 1982). Kerguelen 8. M /)r»fo/(.v (Lilljeborg, 1851 ). Europe and South Africa.

80

R C. Brusca. R. Wetzer, and S. C. France

Figure 66. NaiaUilana ciilifiiniien.\i\ (AHF 6048). holotype. male: A. antennule (left), only pro.ximal

region of aethelascs shown. B. antenna (left). C. frontal lamina, clypeus. and labrum. D. mandible (left). E,

maxillule (left). F, maxilla (left). G. maxilliped (left).

9.

10.

11.

12.

13.

14.

13.

16.

17.

18.

19.

20.

21.

N. hdwnuini Bruce, 1986. New South Wales. Australia. 23

N. biiiha Bruce, 1986. Queensland, Australia. 24

N. caecii iDoWfus. 1903). Europe. 25

N. californiensis (Schuhz, 1 966). California to Gulf of Califor- 26,

nia. 27,

N. caiienae n. sp. Pacific Mexico to Panama. 28

N. chilensis (Menzies. 1962). Chile. 29,

A', corpidenta (Hale. 1923). South Australia. 30,

A', ciirta (Richardson. 1910). Philippines. 31

A', endota Bruce, 1986. New South Wales, Australia.

N. galalhea Bruce, 1986. Gulf of Carpenteria, Australia. 32

A', gallica (Hansen, 1905). Atlantic coast of Europe. 33

A', goning Bruce, 1986. Victoria, Australia.

A', gracilis (Hansen, 1890). West Indies to Brazil.

A', hirtipes (Milne Edwards, 1840). South Africa.

N. insignis Hobbins and Jones, 1993. Red Sea.

N. iiuennedia (Vanhoffen, 1914). Antarctica.

N. jciponensis (Richardson, 1904). Japan.

N. kahiba Bruce. 1986. New South Wales, Australia.

A', karkiiriiok Bruce, 1986. Queensland, Australia.

A', laewilla Bruce, 1986. New South Wales, Australia.

N. longispiiui Bruce. 1986. Victoria, Australia.

N. Iiiriir Bruce. 1986. Western Australia.

A', luticola (Holdich, Harrison, and Bruce, 1981 ). Queensland,

Australia.

A', matong Bruce, 1986. Tasmania, Australia.

N. meridioncdis (Hodgson, 1910). Antarctica. [Bruce (1986a)

synonymized A', idbinota (Vanhoffen, 1914) with N. meridio-

ncdis: Brandt (1988) apparently was unaware of this syn-

onymy].

Tropical Kaslern Pacific Cirolanidae

Figure 67. Natatolana culiforniensis (AHP 604S),ho\otype, male: A. pereopod I (left). B. pereopod IV (left). C, pereopod Vll (left). D, penes. E, dorsal

view of uropod (left). F, pleopod I (left). G, pleopod 2 (left). H, pleopod ."! (left). I. pleopod 4 (left). J, pleopod 5 (left).

82

R. C. Brusca. R. Wetzer, and S. C. France

Figure 68. Natalolana carlenae n. sp. (USNM 252731), holotype. male: A, antennule (right), only

proximal region of aethetascs shown. B. antenna (right). C. frontal lamina, clypeus. and labrum. D, mandible

(right). E, maxillule (right). F, ma.xilla (right). G. maxilliped (right).

34. N. namimddi Bruce. 1986. Victoria, Australia.

35. N. narica (Bowman. 1971 ). New Zealand.

36. N. tiatalensis (Barnard. 1940). South Africa.

37. N. natcilis (Menzies and George, 1972). Peru.

38. N. neglecta (Hansen. 1890). Mediterranean.

39. N. nilida (Hale, 1952). Kerguelen and Crozet Islands, Indian

Ocean.

40. N. obrusata (Vanhoffen, 1914). Antarctica.

41. N. ocu lata {Vanhoffen. 1914). Antarctica.

42. N. pallidocula (Kussakin and Vasina. 1982). Kerguelen

Islands, Indian Ocean.

43. A', pasuirei (Gianibiagi, 1925). Tierra del Fuego.

44. /V. /)e'//i/c7</(( (Tattersall. 1921). New Zealand and Australia.

45. N. pihda ( Barnard. 1 955 ). South Africa.

46. M pro//.v(7 Bruce, 1986. Queensland, Australia.

47. A', rossi (Miers, 1876). New Zealand.

48. N. schmidti {Hunsen. 1905). Faroes, northeast Atlantic.

49. A', teiudsnlis {Miers. 1884). Australia.

50. N. ihaline Bruce. 1986. Queensland. Australia.

51. A', thiinir Bruce. 1 986. Bass Strait. Australia.

52. A', ralida (Hale, 1940). Bass Strait. Australia.

53. N. varigiihenui (Holdich. Harrison, and Bruce, I98I).

Australia.

54. A. v;>to(Hale. 1925). South Australia.

55. N. virilis (Barnard, 1940). South Africa.

56. N. woodjonesi (Hale, 1924). South Australia.

57. N. wowine Bruce. 1986. Victoria. Australia.

58. N. widlunya Bruce. 1986. New South Wales, Australia.

Key to Tropical Eastern Pacific Natauilana Species

1. Without eyes; pleotelson with 6-10 apical spines; mandibular

palp with simple setae only; appendi,\ masculina broadly curv-

ing, basally wide Nahitolaua californiensis

T?iipiciil F.iislcrn Pacific Ciriilaiiidae

83

Figure 69. Naiatokmu carlenae n. sp. ( AHF 959- 1 ). scanning electron micrographs: A, frontal lamina, clypeus, and labrum, 40x. B, frontal lamina, 80x.

C, pereopod I, dactylus, 40x. D, pereopod 1, merus and carpus, 80x.

84

R. C. Brusca, R. Wetzer, and S. C. France

— With eyes (sometimes unpigmented but always with distinct

ommatidia): pleotelson with 10-13 apical spines; mandibular

palp with simple and comb setae; appendix masculina sublinear

N. carlenae n. sp.

Natatukma Lalifonnensis (Schultz, 1966)

Figs. 65A, 66, 67

Namwlana callforniensis Bruce 1981a: ?8; 1986a: 222. Brusca 1980;

228. Brusca and Iverson 1985: 37.

Cimlaiw ailifoniieiuis Schuhz 1966: 14; 1969: 178. Brusca and Ninos

1978: 379.

Ciroluim demiinita Menzies and George 1972: 9. (Not Cirokimi

dimimita of Menzies, 1962b, and other authors).

Type material examined.— (\) Male holotype (LACM 60.88.4,

AHFType No. 6048); U.S.A., California, San Diego Co., Coronado

Canyon, 32° 30.70' N, 117° 21.62' W, on green mud, 794 m; R/V

Ve/ero/VSta. 685 1-60; 1 Feb. 1960. (2) Paratypes (LACM 60.76.3,

AHF Cat. No. 952-1): U.S.A., California, Los Angeles Co.. San

Clemente Island, Tanner Canyon, 54.8 mi., 250° T from China

Point Light, 32° 37.87' N, 1 1 8° 58.70' 'W, 792 m; RA' Velero /VSta.

6833-60; 29 Jan. I960; 2 specimens.

Other material examined. — California specimens; (3) Santa

Catalina Island. 6.75 mi. 92° T from Long Point Light; 33°24' N,

118° 13' 'W; RA' Velero IV Sta. 2228-53, AHF Cat. No. 752-1; 28

Feb. 1953; I male and I juvenile. (4) Santa Catalina Island, 33° 22'

30" N, 1 18° 36' 38" W, Campbell grab on sandy gray-green mud; R/

V Velero IV Sta. 2847-.54; 23 June 1954; I specimen. (5) San

Clemente Island; R/V Velero IV Sta. 635 1 -59, AHF Cat. No. 933- 1 .

LACM; 2 specimens. (6) San Clemente Island. 30.5 mi. 62° T to

ChinaPoint;Sla. 24607-76, AHF Cat. No. 774-01, LACM; I male.

(7) San Clemente Island, 63' T to China Point, 2-30 m; Sta. 24606,

AHF Cat. No. 772-01, LACM; 2 mancas. (8) San Clemente Island,

29.7 mi. 66° T to China Point; AHF Cat. No. 773-0 1 . LACM; 1 male.

(9) No exact locality. 900-1250 m; SDNHM; 28 Aug. 1978; RA

Calafia, Calif. Dept. Fish and Game. coll. R Gregory; 12 specimens.

Gulf of California specimen: ( 10) Angel de la Guarda Island. 7

mi. 253° T from south end. 1135-1138 m; RA Velero IV Sta.

11827-67. LACM; I Dec. 1967; 1 nonovigerous female.

Description o/^H!((/t'.— Cephalon width 1.8 times length. Eyes

absent (Fig. 65A). Antennules extend barely to posterior region of

cephalon; tlagellum of 8-12 free anicles. basal articles fused, each

article with 1-5 long unjointed aesthetascs (only the most basal

portion of aesthetascs figured) (Fig. 66A). Antenna reaching middle

of pereonite II; nagelluin of 10-22 articles (Fig. 66B). Frontal

lamina not expanded anteriorly, narrowing and rounded (Fig. 66C).

Mandibular spine row with about 11-14 stout spines; 2 lateral

(outer) cusps of incisor weakly developed; middle and distal ar-

ticles of palp with simple setae only (Fig. 66D). Maxillule"s medial

lobe with lateral protuberance, deep notch, and 3 stout strongly

tapering circumplumose spines; lateral lobe with about 12 large

spines, the largest with barbs, followed by 9 small subapical mar-

ginal spines (Fig. 66E). Maxilla medial lobe with 9 plumose setae

and 4 simple setae; lateral lobes with 1 1 and 5 simple setae, respec-

tively (Fig. 66F). Left maxillipedal endite with 2 or 3 coupling

spines, right maxillipedal endite with I or 2 coupling spines and 2

apical and 2 subapical plumose setae; palp articles with simple

marginal setae, most distal article also with comb setae as figured

(Fig. 66G).

Pereon widest at pereonites IV and V. Coxae IV-VII produced

beyond posterior margins of their segments; most posterior coxae

visible in dorsal aspect (Fig. 65 A). Pereopod I: inferior margin of

ischium with long setae but no spines; inferior margins of merus,

carpus, and propodus with stout spines as figured; superior lobe of

ischium forms spoonlike depression into which merus collapses;

carpus short (Fig. 67A). Pereopod IV not much longer pereopod I;

distal superior margins of ischium and merus not produced; articles ^

with setae and stout simple spines as figured (Fig. 67B). Pereopod

VII not much longer than pereopod IV; basis and ischium with long

plumose setae as figured; basis 2.5 times longer than wide; ischium,

merus, caipus, and propodus with spines and simple setae as fig-

ured (Fig. 46C). Pereonite VII with small penile lobes (Fig. 67D).

Pleopodal rami with PMS as figured (Figs. 67F-J). Pleopod 1;

peduncle's medial margin with 5 coupling spines and 3 plumose

setae; lateral margin with 1 small spine; endopod 0.78 limes width

of exopod (Fig. 67F). Pleopod 2: peduncle's medial margin with 5

coupling spines and 3 plumose setae, lateral margin with 1 small

spine; endopod 0.88 times width of exopod; appendix masculina

scythelike, narrowing from base to apex, length 1.15 times exopod

length (Fig. 670). Pleopod 3; peduncle's medial margin with 4

coupling spines and 4 plumose setae, lateral margin with I short

spine; endopod 0.95 times as wide as exopod, with short incision on

lateral margin; exopod with short incision on medial margin (Fig.

67H). Pleopod 4: peduncle's medial margin with 4 coupling spines

and 3 plumose setae, lateral margin with 1 spine; endopod 0.95

times as wide as exopod, with short incision on lateral margin;

exopod with short incisions on medial and lateral margins (Fig.

671). Pleopod 5: peduncle with 1 short spine on lateral margin,

endopod width subequal to exopod width; exopod with short inci-

sions on medial and lateral margins (Fig. 67J).

Pleotelson subtriangular, lateral margins slightly convex (not

straight); distal quarter of pleotelson with marginal serrations and

6-10 spines interspersed with long PMS; dorsum with shallow

paired submedian depressions near base (Fig. 65A). Uropods ex-

tend barely beyond apex of pleotelson, narrowed apically; rami

without apical notches, margins slightly serrate, fringed with long

PMS and short spines. Uropodal exopod 0.5 width of endopod;

medial margin of exopod with 2-4 stout spines, lateral margin with

5-8 stout spines. Uropodal endopod with 4-6 stout spines on me-

dial margin, 3^ spines on lateral margin. Uropodal peduncle's

inner angle with distal PMS; distolateral angle with 3 long spines

(Fig. 67E).

female. — Similar to male. Females not bearing oostegites have

variously developed maxillipedal epipods.

Size — To maximum length of 13.4 mm.

Di.strilmtiim. — Natatotana californiensis is primarily a south-

ern California animal; we have examined a single specimen from

the Gulf of California. Collection depths range from 792 to 1250 m.

Remarks. — The holotype is a male, not a female as reported by

Schultz (1966: 15). Our description is based on the holotype and

other California material. The single specimen from the Gulf of

California differs from the California specimens in one regard only;

the clypeus is notched anteriorly, below the posterior tip of the

frontal lamina. Depth data for several of the Velero IV collection

stations could not be found.

Nalatolana carlenae n. sp.

Figs. 65B. 68-70

T\pe material examined.— ( 1 ) Male holotype (USNM 252731 )

and 5 male and female paratypes (USNM 252732): Mexico, Sonora

(Gulf of California), Tiburon Island, on muddy sand, 73-101 m;

Sta. No. 563-36, USNM Ace. No. 139772; 10 Mar. 1936.

Additional paratvpes. Pacific Baja California specimens; (2)

Cedros Island, 28° ()4' N, 115° 20' W, 29 m; RA' Velero IV Sta.

1703-49. LACM; 5 Mar 1949; 6 specimens. (3) 2 mi. S.E. of

Cedros Island Light; 28° 20' N. 115° 10' "W. 101 m; RA Velero III

Sta 1265-41. LACM; 28 Feb. 1941; 5 specimens. (4) Dewey

Channel. San Eugenio Point. 27° 49' 32" N. 115° 06'15" W; R/V

Velero III Sta. 1260-41. LACM; 27 Feb. 1941; 1 male. (5) Thurloe

Head. 27° 34' N. 114° 50' W, 70 m; RA Velero IV Sta. 1 1841-67,

LACM; 7 Dec. 1967; I male and I female.

Tropical Eastern Pacific Cirolanidac

85

Figure 70. Naiulolami carlenae n. sp. (USNM 252731). holotype, male; A, pereopod I (right). B, pereopod IV (right), C, pereopod VII (right). D,

uropod (right). E, pleopod I (right). F, pleopod 2 (right). G, pleopod 3 (nght). H, pleopod 4 (right). I, pleopod 5 (nght).

86

R. C. Brusca, R. Wetzer. and S. C. France

Figure 71. Oncilorpheus jenyhiirnardi n. sp.

( LACM 39-5 1,17, AHF Cat, No, 896-04), holotype.

male.

Gulf of California specimens (all Baja California): (6) North of

Angel de la Guarda Island, 73-128 m; RA' Velero III Sta, 546-36,

USNM Ace, No. 139772; 5 Mar. 1936; 2 specimens. (7) North of

Angel de la Guarda Island, 1 5~ 1 8 m; RA' Velero III Sta No. 55 1 -36,

USNMAcc.No. 139772; 1 specimen. (8) Angel de la Guarda Island,

Puerto Refugio, 165 m; R/V Velero III Sta. 709-37. USNM Ace. No.

144492; 2 1 Mar. 1937; 3 specimens. (9) Angel de la Guarda Island,

Puerto Refugio, on sand, 119 m; R/V Velero III Sta. 544-36, USNM

Ace. No. 139772;4Mar. 1936;4specimens.(10)Angelde la Guarda

Island, Puerto Refugio, with oysters, 28-55 m; R/V Velero III Sta.

No. 542-36, USNM Ace. No. 1 39772; 4 Mar. 1 936; 1 specimen. (11)

Los Angeles Bay, on sand, 46-73 m; R/V Velero III Sta. 535-36,

USNMAcc.No. 139772;2Mar. 1936; I specimen. (12) LosAngeles

Bay, 33 m; R/V Velero III Sta. 702-37, USNM Ace. No. 144492; 20

Mar. 1937; 2 specimens. (13) LosAngeles Bay, 59 m; R/V Velero IH

Sta. 701-37, USNM Ace. No. 144492; 20 Mar. 1937; 3 specimens.

(14) North of Coyote Point, 25° 49' N, 111° 11' W, dredge, sand,

mud, and shell, 51 m; R/V Velero IVSVd. 1753-49, LACM; 20 Mar.

1949; 2 specimens. (15) Between Cenalvo Island and La Paz,

benthic trawl, 247 m; Sta. 3, AHF Cat. No. 959-1, LACM; 7 Sept.

1969; coll. R. Schaffer;4 specimens. (16) Ensenada de los Muertos,

with shell fragments, 73 m; R/V Velero III Sta. 629-37, USNM Ace.

No. 144492; 5 Mar. 1937; I specimen.

Central Eastern Pacific specimens; ( 1 7) Costa Rica, Port Parker;

10° 57' N, 85° 49' W, sandy mud, 9-18 m; R/V Velero III Sta. 936-

39, LACM; 25 Mar 1939; 2 specimens. (18) Panama, Secas Island,

mud and shells, 46 ni; USNM Ace. No. 128938; 22 Feb. 1934; I

male, damaged, probably dried and rehydrated.

Description of male. — Cephalon width 1.9 times length. Eyes

elongate, well-developed (Fig. 65B), unpigmenled or golden (in

ethanol), visible in ventral aspect. Antennular peduncle articles

with simple and palmate setae; flagellum of 10 or II articles, each

article with 1-3 long unjointed aesthetascs (only most basal por-

tions of aesthetases figured) (Fig. 68A). Antenna reaching pereonite

III; flagellum of 23 to 24 articles, with simple and palmate setae as

figured (Fig. 68B). Frontal lamina with anterior margin expanded

and rounded (Fig. 68C, 69A, B). Mandibular spine row with about

13 stout spines; inner and middle cusps of incisor acute, outer cusp

rounded, all 3 cusps with elevated ridges; middle and distal palp

articles with simple and comb setae, apical seta of distal article very

long as figured (Fig. 68D). Maxillule's medial lobe with lateral

protuberance, 3 stout cireumplumose spines, and 3 small simple

spines; lateral lobe with about 12 large spines, largest spines with

barbs (Fig. 68E). Maxilla's medial lobe with about 1 1 plumose and

8 simple setae; lateral lobes with 15 and 5 simple setae, respectively

(Fig. 68F). Left and right maxillipedal endites short, each with 2

coupling spines and plumose setae; palp articles with simple setae,

most distal article also with comb setae (Fig. 68G).

Pereon widest at pereonites IV and V. Coxae IV-VII visible in

dorsal view, produced well beyond the posterior margins of their

respective pereonites; coxa VII produced almost to posterior mar-

gin of pieonite 2 (Fig. 65B). Pereopod I: merus, carpus, and

propodus with stout spines as figured; lobe of ischium and merus

with very large distal spines; lobe of ischium forms distal spoonlike

depression into which merus collapses; caipus very short (Figs.

69C, D, 70A). Pereopod IV with distal margins of ischium and

merus not produced as on pereopod I; inferior margins of articles

with very long simple setae and spines as figured (Fig. 70B).

Pereopod VII long, very setose; basis 1.6 times longer than wide;

ischium with simple and plumose setae, simple spines, and about 6

serrate spines; merus and carpus with simple setae and simple and

serrate spines (Fig. 70C). Pereonite VII without penes.

Pleopodal rami with PMS as figured; all pleopodal peduncles

with 1 lateral spine and cluster of sublateral simple setae (Figs.

70E-I). Pleopod 1: peduncle's medial margin with 5 coupling

spines and 4 plumose setae; I spine near lateral margin; endopod

width 0.66 times width of exopod (Fig. 70E). Pleopod 2; peduncle's

Tropical Eastern Pacific Cirolanidae

87

Figure 72. Onalorpheus jerrybarnanU n. sp. (LACM 39-51.17, AHF Cat. No. 896-04): A. antennuie (right). B. antenna (right). C. frontal lamina,

clypeus, and labrum. D. mandible (left). E, maxillule (left). F, maxilla (left). G, maxilliped (left).

medial margin with 4 coupling spines and 5 plumose setae; endopod

width 0.98 times width of exopod; appendix masculina simple:

length 0.97 times endopod length (Fig. 70F). Pleopod 3: peduncle's

medial margin with 4 coupling spines and 7 plumose setae; endopod

subequal to exopod in width, with short incision on lateral margin;

exopod with short incision on tnedial margin (Fig. TOG). Pleopod 4:

peduncle's medial margin with 5 coupling spines and 7 plumose

setae; endopod subequal to exopod in w idth. with short incisions on

medial and lateral margins; exopod with short incision on medial

margin (Fig. 70H). Pleopod 5: peduncle somewhat irregularly

88

R. C. Brusca. R. Wetzer. and S. C. France

Figure 73. Oncilorpheus jernhamardi n. sp. (LACM 39-51.17, AHF Cat. No. 896-04|: A. pereopod I (left). B, pereopod IV (left). C. pereopod VII

(left). D, dorsal view ofpereonite III. E. ventral view of penes on stemite VII, and pleopod I. Oncilorpheus stehhingi (holotype): F, frontal lamina, clypeus

and labrum. G, dorsal view ofpereonite III.

Tropical Eastern Pacific Ciroianidae

Figure 74. Omilorpheiis jenyhaniardi n. sp. A, uropod {left) from paratype, female (USNM 252738). B-F. pleopods (left) from holotype, male

(LACM 39-5 1 . 1 7, AHF Cat. No. 896-04): B. pleopod 1 . C, pleopod 2. D, pleopod 3. E, pleopod 4. F, pleopod 5.

90

R. C. Brusca. R. Wetzer. and S. C. France

shaped, lateral margin produced into elongate lobe; endopod

subequal to exopod in width; proximolaleral angle of exopod

slightly produced, rounded, lobelike (Fig. 701).

Pleotelson subtriangular. lateral margins slightly convex (not

straight); distal quarter ot'pleotelson with marginal serrations. PMS.

and 10-14 stout spines; dorsum devoid of tubercles or carinae, but

with a pair of shallow submedian depressions near base. Uropods

extend beyond apex of pleotelson. margins slightly serrate, fringed

with long PMS and stout spines, narrowed apically. w ithout apical

notches on rami (Fig. 65B). Uropodal exopod 0.58 times width of

endopod; medial margin of exopod with about 3 spines, lateral

margin with 7-9 spines. Uropodal endopod with 4-7 spines on

medial margin, spines on lateral margin, and a single apical spine.

Uropodal peduncle inner angle with PMS; dorsal surface of lateral

(outer) angle with ?< stout spines and PMS as figured (Fig. 70D).

Female. — Similar to male.

Ultrastntctiinil Feulures. — When viewed with an SEM. most

of the body and appendages show a scalelike cuticular structure.

The dorsal regions of the antennules, antennae, and body have

minute cuticular sensillae (Figs. 69A, B).

Size. — To maximum length of 16.8 mm.

Distrihution. — Known from northwestern Baja California (Pa-

cific), the Gulf of California. Costa Rica, and Panama, at depths

ranging from 9 to 1 168 m. Although the depth range of this species

is remarkably broad, we have not observed any differences between

shallow- and deep-water specimens. This species is generally found

in waters shallower than is N. calijomiensis. Of 18 specimen lots

with depth data. 16 were from depths of 9 to I6.'> nV. and only 2 were

from depths of 247 to 1 168 m; most records are from depths of 25 to

170 m. Recorded substrates include sand. mud. and shell fragments;

1 specimen was found "with oysters." Nulalolana cailenae appears

to be fairly common in soft-bottom subtidal habitats, particularly in

the Gulf of California.

Remarks. — This species is very similar to Nalalohma

californiensis. The main characters separating the two are the pres-

ence of eyes in N. carlemie. the number of apical pleotelson spines

(10-14 in N. caiienae. 6-10 in N. californiensis). and uropod

spination. Several small, barely postmanca specimens of

Natatolana carlenae have definite pigmented eyes and 12

pleotelson spines, indicating that these are not age- or size-related

variations but good species differences.

Etxmology. — This common Gulf of California species is named

for the senior author's daughter. Carlene. in appreciation of the

many collecting trips she participated on in the Sea of Cortez, many

before she was old enough to know what an isopod was.

Oncilorpheus Paul and Menzies, 1 97 1

7\7J<' species. — Oncilorpheus slehhingi Paul and Menzies, 1 97 1 .

by original designation. Type specimens at USNM.

Oncilorpheus Paul and Menzies 1971: 29. Kensley and Schotte 1989:

139.

Description. — Body elongate, 4.0-5.0 times longer than broad;

dorsum of pereon with numerous pits or scalloped depressions.

Eyes moderate in size. Cephalon lacking rostrum, moderately im-

mersed in pereonite I; posterior region of cephalon with nearly

complete incision line. Frontal lamina robust, projecting

anteroventrally. 1.8-2.4 times longer than broad, apex subacute or

rounded; clypeus flat (sessile), short, broad, wider than long; la-

bium narrower than clypeus. Antennular peduncle 3-articulate; sec-

ond article not articulated at right angles to first article (as in

Eurydice); peduncular basal articles generally longer than wide;

article 3 longest. Antenna short, only slightly longer than antennule;

peduncle 5-arliculate; articles 4 and 5 subequal in length and longer

than others. Mandible with tridentate incisor; palp 3-articulate,

middle article longest; spine row a small tapered lobe with stout

spines. Maxillule's medial lobe with 3 stout circumplumose spines;

lateral lobe with about 10 stout spines. Maxilla reduced, short, a

single lobe, apex with simple setae. Maxillipedal palp 5-articulate;

endite small, with 1 or 2 coupling spines and plumose setae.

Pereopodal dactyli with short stout accessory spine at base of

unguis; distal superior margin of ischium of pereopods I-III not

greatly produced; distal superior margin of merus produced. Pereo-

pods V-VIl with basis not markedly flattened or enlarged; pereo-

pod VII long and slender; ischium, merus. carpus, and propodus

subequal in length, each one successively narrower. Penes well

developed on sternite of pereonite VII.

Pleon of 5 somites, which may be fused medially; lateral mar-

gins of pleonite 5 largely encompassed by pleonite 4. Pleopod I's

exopod large, indurate, and operculate; endopods of pleopods 2-5

lack PMS. Pleopodal peduncles 2-5 barely wider than long, with-

out accessory lobes; appendix masculina of male arises subbasally

from medial margin of endopod of pleopod 2. Pleotelson tapering

strongly posteriorly, apex subacute or bluntly rounded, never in-

dented. Uropodal peduncles longer than rami; inner angle strongly

produced, with bluntly rounded apex; exopod triangular, distally

acute.

Remarks. — Several emendations to the original description of

this genus (Paul and Menzies 1971) are necessary. In both

Oncilorpheus slehhingi and O. jerryharnardi n. sp., the cephalon is

moderately immersed in pereonite 1. Figure 2 of Paul and Menzies'

paper fails to show this degree of immersion in O. slehhingi. The

frontal lamina, clypeus. and labrum of O. slehhingi are refigured in

our Figure 73F. Paul and Menzies did not describe the uniquely

reduced maxillae of this genus; in both O. slehhingi and O.

jerrxbarnardi n. sp. the lateral and medial lobes are reduced to a

single simple lobe (indicated by a faint cuticular ridge in C.

jerryharnardi n. sp.). Their Fig. 3C identifies this appendage as "the

inner plate of the second maxilla." They also did not describe or

figure the unique body sculpturing typical of the genus (Figs. 71, 73

D, G).

World lisl of species. —

1. O. slehhingi Paul and Menzies. 1971. Venezuela.

2. O. jernharnardi n. sp. Pacific Costa Rica and Panama.

Oncilorpheus jerryharnardi n. sp.

Figs. 71-74

Type maierial examined. — (I) Male holotype (LACM 39-

51.17): Panama (Pacific). Medidor Island, Honda Bay; RA' Velero

III Sta. 948-39, AHF Cat. No. 896-04. Paratypes: (2) USNM

252737. Costa Rica (Pacific). Playa Blancas. along north point,

with mud. sand, and algae, 28 m; R/V Velero HI Sta. 461-35.

USNM Ace. No. 131571; 8 Feb. 1935; I male. (3) LACM 39-48.8.

Panama (Pacific), Secas Island, 50-52 m; KIV Velero III Sta. 945-

39. AHF Cat. No. 2094; I female. (4) USNM 252738, Panama

(Pacific), Secas Island; No. 458; 6 Feb. 1935; 3 specimens. (5)

USNM 252739. Panama (Pacific), Secas Island, south of group,

with mud and shells. 46 m; No. 250, USNM Ace. No. 128938; 22

Feb. 1934; 2 specimens.

Descriplion of male. — Cephalon wider than long; frontal mar-

gin convex, evenly rounded. Eyes situated posterolateral ly (Fig.

7 1 ). Antennule short, reaching posterior margin of cephalon; flagel-

lum of 6-8 articles (Fig. 72A). Antenna reaching middle of

pereonite I; fiagellum of about 15 articles (Fig. 72B). Frontal lamina

about 1.5 times as long as broad; apex forms blunt angle (Fig. Fig.

72C). Mandible with broad tridentate incisor; spine row with about

5 large stout spines; molar process with about 26 small acute spines;

middle palp article with about 7 comb setae and 1 simple seta, distal

article with about 14 simple and comb setae (Fig. 72D). Maxillule's

Tropical Eastern Pacific Cirolanidae

91

medial lobe with 3 eircumpluniose spines and 1 simple seta: lateral

lobe with 10 stout spines, some armed with small apical barbs (Fig.

72E). Maxilla with about 9 simple apical setae and numerous fine

lateral setae (Fig. 72F). Maxillipedal palp articles 2 and ?< with

plumose setae on lateral margins; all articles with simple setae; left

endite with I coupling spine, right endite with 2 coupling spines,

each with ."i apical plumose setae (Fig. 72G).

Body very straight-sided; pereonite I longest. IV-Vll subequal.

longer than II and III; all pereonites subequal in width (Fig. 71).

Coxae well developed but not projecting postenorly beyond their

respective pereonites; not visible in dorsal view. Pereonites heavily

calcified, dorsum with numerous ridges and scalloped pits (Figs.

71. 73D). Pereopod I stout, with simple and plumose setae and

spines as figured; inferior margin of merus with 6 distinct blunt

molariform spines (Fig. 7.^A). Pereopod IV with carpus and

propodus slender, slightly longer than merus. all articles with many

plumose and simple setae and spines (Fig. 7.^B). Pereopod VII with

carpus and propodus slender, longer than merus; all articles with

many plumose and simple setae and spines as figured (Fig. 7.^C).

Penes small, set close together in middle of sternite VII (Fig. 73G).

Pleon widest and longest at pleonite 4. Pleon with strong me-

dian dorsal carina, terminating at pleotelson apex. All pleonites

(except occasionally 1) fused medially (Fig. 71). Pleopodal rami

with PMS as figured (Figs. 74B-F). Exopod of pleopod I highly

calcified, longer than endopod. 3.2 times wider than endopod.

fringed with short, close-set PMS; endopod slender. 3 limes longer

than wide, fringed with short PMS; peduncle subquadrate. with 6

coupling spines on medial margin and 2 plumose setae near

distolateral margin (Fig. 74B). Pleopod 2: peduncle's medial mar-

gin with 4 coupling spines and 6 plumose setae; exopod slightly

longer and wider than endopod; appendix masculina with rounded

spinose apex; length 1 .26 times endopod length (Fig. 74C). Pleopod

3: peduncle with 2 coupling spines and 3 plumose setae on medial

margin; exopod slightly wider than endopod. subequal in length,

exopod with incomplete transverse incision (Fig. 74D). Pleopod 4:

peduncle w ith I or 2 coupling spines and 2 plumose setae on incdial

margin and 1 plumose seta on lateral margin; exopod wider than

endopod but subequal in length, exopod with incomplete transverse

incision; acute protuberance on endopod subapical margin (Fig.

74E). Pleopod 5: peduncle without coupling spines, with I plumo.se

.setae on lateral margin; exopod subequal in length and width to

endopod. with incomplete transverse incision; acute protuberance

on endopod"s subapical margin (Fig. 74F).

Pleotelson subtriangular. lateral margins sinuate, apex narrowly

truncate. Uropodal peduncle broadly expanded, about twice as wide

as endopod. apex of distal-medial angle bluntly rounded (Fig.

74A). Exopod half as wide as endopod; both endopod and exopod

extend beyond pleotelson apex. Pleotelson and uropods with abun-

dant marginal setae, but apparently lacking spines (Fig. 71).

Female. — Similar to male.

Size. — To a maximum length of 15 mm.

Distribulion. — A sublidal species, found at depths of 28 to 52 m

on mud. sand, and shell bottoms. So far known only from three

localities in the Pacific: Medidor and Secas Islands. Panama, and

Playas Blancas. Costa Rica.

Remarks. — Oncilorpheus stehhinfii is known from only 3 speci-

mens collected off Venezuela in the Atlantic (11° 57' N. 64° 37' W).

at 73 m depth (Paul and Menzies 1971). This distribution suggests

that the only two species of the genus may be vicariant descendants

of the transislhmian "Tertiary Caribbean Province" described by

Woodring (1957. 1966). Croizat et al. (1974). Rosen (1975). Brusca

(1980). and others.

Oncilorpheus jcrnharnanli n. sp. differs from O. slehhiiii^i by

the convex frontal margin of the cephalon. the restriction of cuticu-

lar sculpturing to marginal regions of the pereonites. the middorsal

longitudinal carina of the pleon. pleoniles 2-5 being fused medi-

ally, and the sinuate lateral margins of the pleotelson.

Bruce (1986a) and Botosaneanu et al. (1986) placed

Oncilorpheus in an informal genus-group closely corresponding to

the ■■C(;/(/7<'/((-group" of Monod ( 1 930). Wetzer et al. ( 1 987 ) recom-

mended removal of Oncilorpheus from this group.

Eninology. — This species is named in memory of the great

amphipod taxonomist J. Laurens Barnard, an inspiration, often a

sage, and always a kind and gentle man. The patronym stands in

appropriate company with the only other described species in this

genus.

ACKNOWLEDGMENTS

We thank Angelika Brandt. Tom Bowman. Bryan Kensley,

Marilyn Schotte. and especially Niel Bruce for helpful comments

on the manuscript. Thanks also go to Paul Delaney, who, thanks to a

National Science Foundation Grant, worked as an assistant during

the early stages of the research. Most of the drawings were done by

Frances Runyan and John Simpson. We are also indebted to the

rutilant Philip Unitt for his masterful editing and sustained humor in

the face of considerable odds. This work was primarily supported

by grants to Brusca from the National Science Foundation (BSR

87-96330 and 89-18770). Funding for additional field work was

pixnided by two grants to Brusca from the Weiler Foundation, and

special thanks go to William Bullis and Bartlett Bumap for their

support. This is contribution 1 19 from the Grice Marine Biological

Laboratory.

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/loM/cn.w.v (Flabellifera; Cirolanidae) from an exposed sandy beach.

Comparative Biochemistry and Physiology 7.'iA(4):625-629.

Barnard. J. L.. and G. S. Karaman. 1991. The families and genera of

marine gammandean Amphipoda (except marine gammaroids).

Records of the Australian Museum. .Supplement 13 (parts 1/2).

Barnard. K. H. 1914. Contributions to the crustacean fauna of South

Africa. No. 3. Additions to the marine Isopoda. with notes on some

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Note added in proof. The following new generic names in the family Cirolanidae were published while this paper was in press: Aaiokmu Bruce, 1993;

Dodecaluna Carpenter. 1994; Plaki>Uimi Bruce. 1993; SeychelUma Kensley and Schotte, 1994; Ziilialana Botosaneanii and Viloria, 1993.

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