Part 2 - Sphaeromatidae::“Cute As Buttons”

166 FLABELLIFERA • CORALLANIDAE 

Figure 76. Excorallana quadricornis: A, cephalon and pereonite 1 6; B, ventral 

cephalon; C, frontal lamina enlarged. Excorallana sexticornis: D, cephalon and 

o 

pereonite 1 8; E, cephalon 9; F, ventral cephalon. 

RECORDS Turks and Caicos Islands, 1 m; St. Thomas and St. Croix, U.S. 

Virgin Islands, 48—55 m; Cuba; Puerto Rico, on gills of rays Aetobatus narinari 

and Dasyatis americana, and on squirrel fish; Belize, intertidal to 15.2 m, in 

tertid bble sediments in Syringodium and Thalassia 

g r a s s b e d s , o n b r o w n a l g a T u r b i n a r i a , o n M a d r a c i s s p . s p o n g e , o n A g a r i c i a s p 

c o r a l ; G u l f o f M e x i c o .

Excorallana warmingii 167 

F i g u r e 7 7 . Excorallana tricornis tricornis: A , c e p h a l o n a n d p e r e o n i t e 1 6 ; B , v e n t r a l 

c e p h a l o n ; C, p l e o t e l s o n a n d u r o p o d s . 

R E M A R K S T h e s u b s p e c i e s E x c o r a l l a n a t r i c o r n i s o c c i d e n t a l i s R i c h a r d s o n , 1 9 0 5 a , 

f r o m s o u t h e r n C a l i f o r n i a , d i f f e r s f r o m t h e G u l f a n d C a r i b b e a n s u b s p e c i e s i n 

l a c k i n g a g a p b e t w e e n t h e m a r g i n s o f t h e p l e o t e l s o n i c i n c i s i o n , a n d i n h a v i n g 

a r e l a t i v e l y w i d e r u r o p o d a l e x o p o d w h i c h s h o w s a d i s t i n c t l y a s y m m e t r i c a l 

a p i c a l n o t c h . 

E x c o r a l l a n a w a r m i n g i i ( H a n s e n , 1 8 9 0 ) 

F i g u r e 7 5 1 , J 

D I A G N O S I S ( J 9 . 7 m m , 9 1 2 . 0 m m . C e p h a l o n u n o r n a m e n t e d . E y e s c o n 

t i g u o u s , o c c u p y i n g m o s t o f d o r s a l s u r f a c e o f h e a d . P o s t e r i o r m a r g i n s o f 

p l e o n i t e s v e r y f a i n t l y t u b e r c u l a t e . F r o n t a l l a m i n a , l e n g t h s l i g h t l y m o r e t h a n 

t w i c e b a s a l w i d t h , t a p e r i n g a n t e r i o r l y t o r o u n d e d a p e x . P l e o t e l s o n u n o r n a 

m e n t e d e x c e p t f o r t w o f a i n t s u b m e d i a n t u b e r c l e s b a s a l l y ; l a t e r a l i n c i s i o n s 

l a c k i n g ; a p e x b r o a d l y r o u n d e d , w i t h f i v e l o w b u t d i s t i n c t m a r g i n a l t e e t h . 

*

168 FLABELLIFERA • CORALLAN1DAE 

Figure 78. Nalicora rapax: A, 9; B> maxilla 1; C> maxilla 2; D, maxilliped. 

RECORDS Bahamas; between Cuba and the Yucatan Pensinsula; Puerto 

Rico. 

Off Brazil near Rio de Janeiro. 

Nalicora Moore, 1901 

DIAGNOSIS Maxilla 1 exopod a single strongly falcate distal spine with 

knoblike mesial process, and basal caplike convex papilla-bearing structure. 

Maxilla 2 of four articles, distal article slender. Maxillipedal palp of five 

articles; endite lacking.

Nalicora rapax Moore, 1901 

Figure 78 

FLABELL1FERA • CYMOTHOIDAE 169 

DIAGNOSIS 6 6.9 mm, ovigerous 9 10.0 mm. Eyes well developed. Frontal 

lamina basally slender, widening anteriorly, apex subacute. Posterior half of 

body bearing numerous scattered stiff setae. Pereonites 4-7 with row of low 

rounded tubercles near posterior margin. Posterior margins of pleonites 3—5 

faintly tuberculate, more noticeable in 6. Pleotelson wider than long; lateral 

margins faintly sinuous; apex rounded. 

RECORDS Florida Keys, 55 m; Puerto Rico, 50-150 m; Gulf of Mexico off 

Florida, 37-73 m. 

Family Cymothoidae Leach, 1818 

DIAGNOSIS Antennules and antennae reduced, no clear distinction between 

peduncles and flagella. Mandibular palp of three articles. Maxilla 1 with four 

terminal spines. Maxilla 2 apically bilobed, armed with several spines. Max- 

illipedal palp of two articles, terminal article bearing hooks. All seven pairs of 

pereopods prehensile, ending in strongly hooked dactyli. Pleopods lacking 

marginal setae in adults. 

REMARKS The cymothoids are exclusively ectoparasites on marine, fresh 

water, and brackish-water fishes. Most cymothoids occur in shallow water, 

mainly in tropical and subtropical areas. The position of attachment on the 

host (externally, in the buccal cavity, or in the gill chamber) is usually genus- 

or species-specific. The body of gill parasites is often asymmetrical, being 

slightly twisted, perhaps an effect of the position on the host. The mouthparts 

are highly adapted for the parasitic mode of life, while all seven pairs of 

pereopods are strongly prehensile. The posterior pereopods of some genera 

have the basal article expanded and carinate, allowing for increased mus 

culature. The secretion of anticoagulants in the juvenile stages further aids 

the blood-feeding habit. The surface area of the pleopods is often increased 

by the development of lobes on the bases or the lamellae, providing an in 

creased respiratory ability. 

The post-mancal juvenile stages (sometimes referred to as the aegathoid 

stage) have large eyes, and highly setose pleopods for active swimming. The 

juveniles will attach themselves indiscriminantly to any convenient fish host, 

but eventually attach to the preferred host-species. The juvenile then de 

velops into a functional male, losing the swimming setae of the pleopods. 

Both juveniles and males feed actively, drawing blood from the host fish. The

170 FLABELLIFERA • GYMOTHOIDAE 

Key to genera of Cymothoidae 

1. Antennule broader and usually longer than antenna; cephalon very 

weakly sunk into pcreonite 1 2 

Antennule not broader or longer than antenna; cephalon distinctly 

immersed in, or not at all immersed in pereonite 1 4 

2. Bases of antennules widely separated 3 

Bases of antennules contiguous Glossobius 

3. Body curved to one side; pleonitc 1 extended laterally more on one side 

than on other Mothocya 

Body rarely curved to one side; pleonitc 1 extended equally on each 

side Renocila 

4. Pereonites and coxal plates 4-7 strongly expanded on one side only 

Agarna 

No pereonites or coxal plates strongly expanded 5 

5. Cephalon not immersed in pereonite 1; posterior margin of cephalon 

trisinuate 6 

Cephalon to some degree immersed in pereonite 1; posterior margin of 

cephalon not trisinuate 7 

6. Posterolateral angles of pereonites 2-6 not produced; coxal plates short, 

rarely reaching posterior margin of their pereonites Anilocra 

Posterolateral angles of pereonites 2-6 posteriorly increasingly 

produced; coxal plates usually reaching to posterior margin of their 

pereonites Nerocila 

7. Basal antennular articles expanded and contiguous Ceratothoa 

Basal antennular articles expanded but not contiguous, or basal 

antennular articles neither expanded nor contiguous 8 

8. Basal antennular articles expanded but not contiguous Kuna 

Basal antennular articles neither expanded nor contiguous 9 

9. Plconal margins continuous with pereonal margins, pleon not abruptly 

narrowed, only weakly immersed in pereonite 7 Lironeca 

Pleon to some degree narrower than pereon; pleon usually deeply 

immersed in pereonite 7 Cymothoa

FLABELLIFERA • CYMOTIIOIDAE 171 

male eventually becomes a female (all cymothoids are protandrous) should a 

female not already be present. In some species, the female is nonfeeding. In 

those species which settle either in the mouth cavity or gill chamber of the 

host, integumental pigment is frequently lost, and the eyes become reduced. 

Given the highly variable morphology of the cymothoids, in part imposed 

by the parasitic mode of life, and the existence of polymorphism and possible 

sibling species, the taxonomy of this family demands the examination of large 

numbers of specimens. As a further aid to identification, Table 3 is provided, 

giving host species, parasite, and site of attachment. 

TABLE 3. GYMOTHOID PARASITES FROM THE CARIBBEAN AREA, LISTED BY FISH 

HOST SPECIES 

Fish host Cymothoid parasite Site of attachment 

Abudefduf saxatilis 

Acanthurus bahianus 

Acanthurus chirurgus 

Alutera schoepji 

Anchoa lamprotaenia 

Apogon lachneri 

Apogon maculatus 

Apogon townsendi 

Ariusfelis 

Astrapogon stellatus 

Batrachoides surinamensis 

Caranx hippos 

Caranx latus 

Caranx ruber 

Caranx sp. 

Chaetodipterus faber 

Chaetodon capistratus 

Chaetodon ocellatus 

Chaetodon sedentarius 

Chaetodon striatus 

Anilocra abudefduji 

Kuna insularis 

Anilocra acanthuri 

Anilocra acanthuri 

Nerocila acuminata 

Lironeca tenuistylis 

Mothocya bohlkeorum 

Renocila colini 

Renocila colini 




Cymothoa oestrum 





Anilocra chaetodontis 




beneath eye 

gill chamber 

9 at base of pectoral fin; 

immature on or near 

pectoral or pelvic fin 

9 at base of pectoral fin; 

immature on or near 

pectoral or pelvic fin 

on or at base of fin 

posterior to pectoral fin 

in gill chamber 

next to dorsal fin 





inside mouth 

inside mouth 

inside mouth 

inside mouth 


beneath eye 

beneath eye 

beneath eye 

beneath eye 

(continued)

172 

TABLE 3. {Continued) 


Chilomycterus schoepfi 

Chromis cyaneus 

Chromis multilineatus 

Cynoscion nebulosus 

Cynoscion sp. 

Epinephelus cruentatus 

Epinephelus Julvus 

Epinephelus guttatus 

Epinephelus itajara 

Epinephelus sp. 

Exocoetus spp. 

Genes rhombeus 

Haemulon aurolineatum 

Haemulon carbonarium 

Haemulon chrysargyreum 

Haemulon Jlavolineatum 

Haemulon macrostomum 

Haemulon plumieri 

Haemulon sciurus 


Anilocra chromis 

Anilocra chromis 

Cymothoa excisa 


Anilocra haemuli 





Glossobius impressus 

Lironeca redmanni 








Hemirhamphus brasiliensis Glossobius hemirhamphi 

Hirundichthys speculifer 

Holacanthus tricolor 

Holocentrus ascensionis 

Glossobius impressus 

Anilocra holacanthi 

Anilocra holocentri 

Hyporhamphus unifasciatus Mothocya nana 

Leiostomus xanthurus 

Lepiosteus spatula 

Lutjanus analis 

Lutjanus mahogoni 

Lutjanus synagris 

Megalops atlanticus 

Monacanthus ciliatus 

Mugil cephalus 

Myripristis jacobus 

Ocyurus chrysurus 

Orthopristis chrysoptera 











Anilocra myripristi 




beneath eye 

beneath eye 

inside mouth 

inside mouth 

beneath eye 

beneath eye 

beneath eye 



inside mouth 


beneath eye 

beneath eye 

beneath eye 

beneath eye 

beneath eye 

beneath eye 

beneath eye 

inside mouth 

inside mouth 

beneath eye 

? between eyes, 8 and 

immature beneath 

eye 



inside mouth 


on at base of fin 

inside mouth 

inside mouth 

inside mouth 

inside mouth 



9 between eyes, imma 

ture beneath eye 

inside mouth 

inside mouth

Agarna cumulus 173 


Orthopristis ruber 

Paranthias Jurcifer 

Phaeoptyx conklini 

Phaeoptyx pigmentaria 

Pogonias cromis 

Pomacentrus partitas 

Priacanthus arenatus 

Scomberomorus cavalla 

Scomberomorus maculatus 

Scomberomorus regalis 

Selar crumenophthalmus 

Serranus tigrinus 

Sphoeroides maculatus 

Synodus foetens 






Anilocra partiti 






Renocila bowmani 

Renocila waldneri 



Agarna Schioedte and Meinert, 1883 

beneath eye 

beneath eye 




beneath eye 

inside nouth 




inside mouth 




inside mouth 

DIAGNOSIS Cephalon with posterior margin not trilobed; immersed in per- 

eonite 1. Antennular bases contiguous. Pereonites 4-7 on one side flattened 

and expanded; coxal plates of pereopods 4-7 also expanded and flattened 

but generally hidden by lateral expansion of pereonites. Bases of posterior 

three pereopods with well-formed carinae. Pleonites 1 and 2 immersed in 

pereonite 7; pleonites 2-5 with free fingerlike lateral margins. 

Agarna cumulus (Haller, 1880) 

Figure 79 

DIAGNOSIS 9 18 mm. Eyes present, indistinct. Pereon strongly "humped" 

dorsally. Uropod about { fo length of pleotelson; uropodal exopod slightly 

longer, and twice width of endopod. Pleotelson triangular, length 3 A basal 

width, apex rounded. 

RECORDS No host recorded: Key West, Florida.


Figure 79. Agarna cumulus: A, 9, dorsal view; By 9, ventral view, coxal plates 

stippled. 

Anilocra Leach, 1818 

DIAGNOSIS Cephalon usually narrowed anteriorly to triangular apex folded 

ventrally between bases of antennules; posterior margin trilobed; not im 

mersed, or only weakly immersed in pereonite 1. Coxal plates small, com 

pact, not reaching level of posterior margin of their respective pereonites. 

Pereopods increasing in length posteriorly, pereopod 7 often markedly longer 

than 6. Pleon not immersed or only slightly immersed in pereonite 7. 

Pleopods 3—5 often formed into deep pockets or pleats. Uropods often ex 

tending beyond pleotelsonic apex. 

REMARKS Williams and Williams (1981) have provided a comprehensive 

treatment of this genus and nine of its species in the West Indies. Table 1 in 

this latter paper provides characters for separating these nine species. This 

table also indicates that for each species, the site of attachment of the adult to 

the host fish is specific, with six species attaching under the eye of the host.

Key to species of Anilocra 

Anilocra abudefdufi 175 

1. Pereopods 2-4 with swelling on outer margin of dactylus 2 

Pereopods 2-4 lacking swelling on outer margin of dactylus 5 

2. Body axis distorted by more than 10° holacanthi 

Body axis distorted by less than 5° 3 

3. Dactylus of pereopod 7 longer than propodus partiti 

Dactylus of pereopod 7 shorter than propodus 4 

4. Posteroventral angle of pereonite 7 overlapping pleonite 1 only 

Posteroventral angle of pereonite 7 overlapping pleonites 1 and 2 

abudefdufi 

chaetodontis 

5. Posteroventral angle of pereonite 7 produced 6 

Posteroventral angle of pereonite 7 not produced 7 

6. Uropod reaching posterior margin of pleotelson myripristis 

Uropod not reaching posterior margin of pleotelson haemuli 

7. Posteroventral angle of pereonite 7 overlapping pleonite 1 . . . . holocentri 

Posteroventral angle of pereonite 7 not overlapping pleonite 1 8 

8. Uropod reaching posterior margin of pleotelson acanthuri 

Uropod not reaching posterior margin of pleotelson chromis 

Anilocra abudefdufi Williams and Williams, 1981 

Figure 80A-C 

DIAGNOSIS Ovigerous 9 19.0-31.0 mm, 6 7.0-8.5 mm. Pereopods 2-4 

with swelling on outer margin of dactylus. Posteroventral angle of pereonite 6 

slightly produced, of pereonite 7 more produced, overlapping pleonite 1. 

Uropodal endopod variable, not reaching, to extending well beyond, apex of 

exopod. Color: upper lateral half to three-fourths of dorsal surface of 9 when 

attached to host is dark brown; rest of dorsal surface light brown to yellow. 

Attaching beneath eye of host.


Figure 80. Anilocra abudefdufi: A, 9, lateral view; B, pereopod 3; C, pereopod 7. 

Anilocra acanthuri: D, pleotelson and uropods. Anilocra chaetodontis: E, 9, lateral 

view. Anilocra chromis: F, pleotelson and uropods. 

Anilocra acanthuri Williams and Williams, 1981 

Figure 80D 

DIAGNOSIS Ovigerous 9 29.0-40.0 mm, 6 4.0-8.0 mm. Pereopods 2-4 

without swelling on outer margin of dactylus. Posteroventral angles of per- 

eonites not produced. Uropod not reaching posterior margin of pleotelson. 

Endopod of uropod variable, not reaching, to extending well beyond, apex of 

exopod. Color: dorsal surface of 9 black to lead gray, ventral surface gray. 

Attaching under pectoral fin of host. 

RECORDS Doctorfish Acanthurus chirurgus: Florida Keys; Bahamas; Puerto 

Rico; U.S. Virgin Islands. Ocean surgeon Acanthurus bahianus: Florida Keys; 

Bahamas; Cuba; Jamaica; Dominican Republic; Puerto Rico; U.S. Virgin 

Islands.

Anilocra chaetodontis Williams and Williams, 1981 

Figure 80E 

Anilocra haemuli 177 

DIAGNOSIS Ovigerous 9 18-28 mm, 6 4-5 mm. Pereopods 2-4 with swell 

ing on outer margin of dactylus, Posteroventral angles of pereonites 4—7 be 

coming progressively produced, that of pereonite 7 overlapping pleonite 2. 

Uropod not reaching posterior margin of pleotelson; uropodal endopod ex 

tending beyond apex of exopod. Pleotelson as wide as long to slightly wider 

than long. Color: dorsal surface of 9 black to lead gray, ventral surface gray. 

Attaching beneath eye of host. 

RECORDS Foureye butterflyfish Chaetodon capistratus: Bahamas; Puerto Rico; 

British and U.S. Virgin Islands. Banded butterflyfish Chaetodon striatus: 

Bahamas; Puerto Rico; British Virgin Islands. Spotfin butterflyfish Chaetodon 

ocellatus: Bahamas; Puerto Rico; U.S. Virgin Islands. Reef butterflyfish 

Chaetodon sedentarius: Puerto Rico. 

Anilocra chromis Williams and Williams, 1981 

Figure 80F 

DIAGNOSIS Ovigerous 9 16-28 mm, 8 4-9 mm. Pereopods 2-4 lacking 

swelling on outer margin of dactylus. Posteroventral angles of pereonites not 

produced. Uropod extending beyond posterior margin of pleotelson; uropo 

dal endopod not reaching beyond exopod. Color: upper lateral one-fourth to 

two-thirds of dorsal surface of 9 when attached is dark gray, shading to off- 

white lower lateral area. Attaching beneath eye of host. 

RECORDS Brown chromis Chromis multilineatus: Puerto Rico; British and 

U.S. Virgin Islands. Blue chromis Chromis cyaneus: Bahamas; Dominican Re 

public. No host recorded: Anguilla. 

Anilocra haemuli Williams and Williams, 1981 

Figure 81A,B 

DIAGNOSIS Ovigerous 9 21-40 mm, 6 7 mm. Body axis distorted less than 

5°. Pereopods 2—4 lacking swelling on outer margin of dactylus. Postero 

ventral angle of pereonites 6 and 7 produced, latter overlapping pleonite 1. 

Uropod not reaching posterior margin of pleotelson; uropodal endopod 

reaching beyond apex of exopod. Color: dorsal surface of 9 yellow to light 

brown. Attaching beneath eye of host.


Figure 81. Anilocra haemuli: A, 9, dorsal view; B, 9, lateral view. Anilocra 

holacanthi: C, 9. Anilocra holocentri: D, 9. Anilocra myripristis: E, pleotelson and 

uropods. Anilocra partiti: F, 9; G, pereopod 7. 

RECORDS French grunt Haemulon Jlavolineatum: Florida Keys; Puerto Rico; 

British and U.S. Virgin Islands. Tomtate Haemulon aurolineatum: Jamaica; 

Puerto Rico. Smallmouth grunt Haemulon chrysargyreum: Puerto Rico; U.S. 

Virgin Islands. Caesar grunt Haemulon carbonarium: Puerto Rico; U.S. Virgin

Anilocra holocentri 179 

Islands. Spanish grunt Haemulon macrostomum: Puerto Rico. White grunt 

Haemulon plumieri: Florida Keys; Yucatan Peninsula. Bluestriped grunt 

Haemulon sciurus: Florida Keys. Cora cora Orthopristis ruber: Margarita Island, 

Venezuela. Coney Epinephelus Julvus: Bahamas; Dominican Republic; Puerto 

Rico; U.S. Virgin Islands; Guadeloupe. Red hind Epinephelus guttatus: Puerto 

Rico; British and U.S. Virgin Islands. Graysby Epinephelus cruentatus: 

Bahamas; Dominican Republic; U.S. Virgin Islands. Creole-fish Paranthias 

jurcifer: Dominican Republic; Puerto Rico; Colombia. No host recorded: 

Cuba; Jamaica; Dominica; Barbados; Venezuela; Brazil. 

Anilocra holacanthi Williams and Williams, 1981 

Figure 81C 

DIAGNOSIS Ovigerous 9 21-33 mm, 6 4-7 mm. Body axis distorted by 

more than 10°. Pereopods 2-4 with swelling on outer margin of dactylus. 

Posteroventral angles of pereonites 5—7 progressively more produced, that of 

pereonite 7 overlapping pleonite 1. Uropod not reaching posterior margin of 

pleotelson; uropodal endopod reaching beyond apex of exopod. Color: dorsal 

surface of 9 black to lead gray. Attaching beneath eye of host. 

RECORDS Rock beauty Holacanthus tricolor: Bahamas; Jamaica; Dominican 

Republic; Puerto Rico; British and U.S. Virgin Islands. 

Anilocra holocentri Williams and Williams, 1981 

Figure 81D 

DIAGNOSIS Ovigerous 9 32-46 mm, 6 5-9 mm. Body axis distorted less 

than 5°. Pereopods 2—4 lacking swelling on outer margin of dactylus. 

Posteroventral angle of pereonite 7 produced, overlapping pleonite 1. 

Uropod not reaching posterior margin of pleotelson; uropodal endopod 

reaching beyond apex of exopod. Color: dorsal surface of 9 dark brown, 

ventral surface light brown. 9 attaching between eyes of host; 6 or transi 

tional stage beneath eye. 

RECORDS Squirrelfish Holocentrus ascensionis: Puerto Rico; U.S. Virgin 

Islands. 

No host recorded: Patagonia, Straits of Magellan.

180 FLABELLIFERA • CYMOTHOIDAE 

Anilocra myripristis Williams and Williams, 1981 

Figure 81E 

DIAGNOSIS Ovigerous 9 29-40 mm, 6 6-7 mm. Body axis distorted less 

than 5°. Pereopods 2-4 lacking swellings on outer margin of dactylus. 

Posteroventral angle of pereonites 6 and 7 produced, latter overlapping 

pleonite 1. Uropod reaching beyond posterior margin of pleotelson; uropodal 

endopod reaching beyond apex of exopod. Color: dorsal surface of 9 light 

reddish brown, ventral surface yellow. 9 attaching between eyes of host; 

immature or transitional forms sometimes beneath eve. 

4 

RECORDS Blackbar soldierfish Myripristis jacobus: Bahamas; Dominican Re 

public; Puerto Rico. 

Anilocra partiti Williams and Williams, 1981 

Figure 81F,G 

DIAGNOSIS Ovigerous 9 12-16 mm, transitional 7.6-9.0 mm. Body axis 

distorted less than 5°. Pereopods 2-4 with swelling on outer margin of dac 

tylus. Pereopod 7 with dactylus longer than propodus. Posteroventral angle 

of pereonite 7 produced, overlapping pleonite 1. Uropod not reaching pos 

terior margin of pleotelson; uropodal endopod not reaching apex of exopod. 

Color: dorsal surface black to slate gray. Attaching beneath eye of host. 

RECORDS Bicolor damselfish Pomacentrus partitas: Jamaica. 

Ceratothoa Dana, 1852 

DIAGNOSIS Cephalon more or less immersed in pereonite 1, posterior mar 

gin not trisinuate. Bases of antennules expanded, contiguous. Coxal plates 

compact; anterior plates not extending beyond posterior margins of their 

respective pereonites; posterior coxal plates may or may not be produced 

beyond the posterior margins of the pereonites. Anterior pleonites narrowed, 

immersed in pereonite 7, Copulatory stylet lacking on pleopod 2 of 6 of some 

species. 

Ceratothoa deplanata Bovallius, 1885 

Figure 82A 

DIAGNOSIS 9 18 mm. Cephalon subtriangular, anterior margin rounded. 

Pereopods 4—7 with strongly carinate bases. Uropod reaching or extending

Ceratothoa deplanata 181 

Figure 82. A, Ceratothoa deplanata (from Bovallius, 1885); B, Cymothoa caraibica; C, 

Cymothoa excisa; D, Cymothoa oestrum. 

slightly beyond posterior margin of pleotelson; rami subequal in length and 

width. Pleotelson basally wider than long, posterior margin broadly 

rounded. Color: bright yellow. 

RECORDS Haiti, host not recorded.

182 FLABELUFERA • GYMOTHOIDAE 

Cymothoa Fabricius, 1793 

DIAGNOSIS Body usually not distorted. Cephalon with posterior margin not 

trilobed; more or less immersed in pereonite 1; latter with anterolateral cor 

ners produced to embrace cephalon. Bases of antennules not expanded, well 

separated. Anterior coxal plates not reaching posterior borders of their re 

spective pereonites, posterior coxal plates nearly reaching or extending be 

yond posterior borders of pereonites. Pleon narrower than, and immersed in 

pereonite 7. Pleonites increasing in length and width posteriorly. 

Key to species of Cymothoa 

1. Anterolateral angles of pereonite 1 reaching to half length of cephalon 

or less; eyes or traces of eyes present 2 

Anterolateral angles of pereonite 1 broad, reaching to anterior margin 

of cephalon; eyes absent oestrum 

2. Anterolateral angles of pereonite 1 narrow, subacute excisa 

Anterolateral angles of pereonite 1 broad, rounded caraibica 

Cymothoa caraibica Bovallius, 1885 

Figure 82B 

DIAGNOSIS 9 17 mm, 6 12-16 mm. Anterior margin of cephalon broadly 

rounded. Eyes large, distinct. Broadly rounded anterolateral angles of per 

eonite 1 reaching to about midlength of cephalon. Bases of pereopods 4—7 

with strong, rounded carina. Uropodal rami subequal in length, equal to 

peduncle in length. Pleotelson width about twice length, posterolateral mar 

gin broadly rounded. 

RECORDS Puerto Rico; Gulf of Mexico. 

Cymothoa excisa Perty, 1833 

Figure 82C 

DIAGNOSIS Ovigerous 9 20-24 mm. Anterior margin of cephalon in dorsal 

view truncate to slightly excavate; eyes small, indistinct. Anterolateral angles 

of pereonite 1 narrowly rounded to subacute, reaching anteriorly to about 

midlength of cephalon. Pereopods 4—7 with high rounded carina on basis.

Glossobius 183 

Uropods hardly reaching halfway along lateral margin of pleotelson; exopod 

slightly longer than endopod. Pleotelson about twice wider than long; 

broadly rounded and somewhat bilobed. 

RECORDS Yellowtail snapper Ocyurus chrysurus: Yucatan Peninsula, Mexico; 

Carrie Bow Cay, Belize; Margarita Island, Venezuela; Panama. Mutton 

snapper Lutjanus analis: Yucatan Peninsula, Mexico; Panama. Lane snapper 

Lutjanus synagris: Panama. Mahogany snapper Lutjanus mahogoni: Panama. 

Pigfish Orthopristis chrysoptera: Florida, Gulf of Mexico. Spot Leiostomus 

xanthurus: Texas, Gulf of Mexico. Spotted seatrout Cynoscion nebulosus: Texas, 

Gulf of Mexico. Inshore lizardfish Synodusfoetens: Texas, Gulf of Mexico. No 

host recorded: Massachusetts; South Carolina; Georgia; Florida Keys; 

Bahamas; Cuba; Trinidad; Brazil. 

Cymothoa oestrum (Linnaeus, 1793) 

Figure 82D 

DIAGNOSIS Ovigerous 9 38 mm. Cephalon in dorsal view with ante 

rolateral angles rounded, anterior margin slightly excavate; eyes absent. An 

terolateral angles of pereonite 1 expanded, broadly rounded, reaching to 

level of anterior margin of cephalon. Pereonites 4—7 with high rounded 

carina on basis. Uropod reaching posteriorly beyond midlength of 

pleotelson; exopod slightly longer than endopod. Pleotelson length slightly 

more than half basal width. 

RECORDS Bigeye scad Selar crumenophthalmus: Bermuda; U.S. Virgin Is 

lands. Bigeye Priacanthus arenatus: Bermuda. Bar jack Caranx ruber: Florida 

Keys; Carrie Bow Cay, Belize. Horse-eye jack Caranx latus: Bahamas; Bar 

bados. Crevalle jack Caranx hippos: Venezuela. Jack Caranx sp.: Jamaica; Cu 

rasao. Hind Epinephelus sp.: Grenada. Parrotfish: Jamaica. Seatrout Cynoscion 

sp.: Panama. Tarpon Megalops atlantica: Texas, Gulf of Mexico. No host re 

corded: Honduras; Haiti. 

Glossobius Schioedte and Meinert, 1883 

DIAGNOSIS Cephalon not immersed in pereonite 1; excavate on either side 

in anterior half, forming broad and anteriorly rounded median area; anten 

nae fitting into excavate areas. Bases of antennules contiguous, expanded. 

Antennules broader and longer than antennae. Bases of pereopods 4—7 with 

posterior margin expanded and flattened. Pleonites 1-3 immersed in per 

eonite 7.


Key to species of Glossobius 

1. Coxal plates of perconites 1 and 2 anteroventrally protruding impressus 

Coxal plates of pereonites 1 and 2 close to body, not protruding 

Glossobius hemiramphi Williams and Williams, 1985a 

Figure 83A 

hemirhamphi 

DIAGNOSIS Ovigerous 9 27 mm. Eyes small but distinct. Cephalon pointed 

anteriorly. Fused coxa of pereonite 1 and free coxa of pereonite 2 carinate but 

not protruding. Coxa of pereonite 7 semicircular in dorsal view. Pleotelson 

with middorsal length more than half basal width; lateral margins somewhat 

tapered; posterior margin variable, sinuate or excavate. Uropods reaching to 

or slightly beyond posterior pleotelsonic margin; rami subequal in length, 

exopod slightly broader than endopod. 

RECORDS Ballyhoo Hemiramphus brasiliensis; Puerto Rico. 

Glossobius impressus (Say, 1818) 

Figure 83B 

DIAGNOSIS Ovigerous 9 33 mm. Eyes small but distinct. Cephalon 

rounded anteriorly. Fused coxal plate of pereonite 1 and distinct coxal plate 

of pereonite 2 protruding strongly in oblique anteroventral direction. Uropod 

reaching to posterior half of pleotelson; exopod shorter and narrower than 

endopod. Pleotelson basal width twice length, posteriorly broadly bilobed. 

Attaching inside mouth of host. 

RECORDS Flyingfish Exocoetus spp.: Rio de Janeiro, Brazil; North Atlantic, 

especially in the Gulf Stream. 

Mirrorwing flyingfish Hirundichthys speculifer: North Atlantic. No host rec 

ord: Senegal, West Africa. 

Kuna Williams and Williams, 1986 

DIAGNOSIS Cephalon somewhat immersed in pereonite 1. Anterior margin 

of pereonite 1 not trisinuate. Number of articles in antennules and antennae

Kuna 185 

Figure 83. A, Glossobius hemiramphi; B, Glossobius impressus; C, Kuna insularis; D, 

Lironeca redmani; E, Lironeca tenuistylis. 

reduced. Antennule somewhat expanded; basal article expanded but not 

contiguous. Copulatory stylet present on pleopods 1-3 in 6. Pleonites dor- 

sally strongly convex, not immersed in pereonite 7.


Kuna insularis (Williams and Williams, 1985b) 

Figure 83C 

DIAGNOSIS Ovigerous 9 11.1-17.2 mm, 6 4.2-8.7 mm, transitional 9.6- 

9.8 mm. Antennules and antennae consisting of four articles each. Uropods 

short, not reaching posterior margin of pleotelson. Clavate eopulatory stylet 

present on pleopods 1—3 in 6. Pleotelson basally broader than long, pos 

terior margin broadly rounded. 

RECORDS Sergeant major Abudefdufsaxatilis: Carrie Bow Cay, Belize; Cura 

sao; Panama. 

Lironeca Leach, 1818 

DIAGNOSIS Cephalon weakly to deeply immersed in pereonite 1; posterior 

border rarely trisinuate. Bases of antennules not expanded, well separated. 

Posterior pereopods with carinae on bases in 6, carinae present or absent in 

9. Pleonites subequal in width; pleonites 1 and 2 rarely narrowed and 

weakly to moderately immersed in pereonite 7. Pleopods highly folded, and 

with lamellar or digitiform accessory gills in some species. 

Key to species of Lironeca 

1. Uropodal endopod about twice longer than wide; pleon somewhat 

immersed in pereon redmanni 

Uropodal endopod about three times longer than wide; pleon barely 

immersed in pereon tenuistylis 

Lironeca redmanni Leach, 1818 

Figure 83D 

DIAGNOSIS Ovigerous 9 19.5-25.0 mm. Cephalon barely immersed in per 

eonite 1. Pleon somewhat immersed in pereon, but lateral margins of pleonite 

1 free. Pleotelson basally wider than long. Uropodal rami reaching well be 

yond posterior margin of pleotelson; exopod longer than endopod, both rami 

somewhat broad, endopod about twice longer than wide. Attaching to gills of 

host.

Mothocya 187 

RECORDS New Jersey to Florida; gills of kingfish, Jamaica; Cuba; St. 

Christopher; Spanish mackerel Scomberomorus maculatus and cero Scomberomorus 

regalisy Puerto Rico; king mackerel Scomberomorus cavalla, Colombia; Genes rho- 

mbeus, Panama; spot Leiostomus xanthurus, Gulf of Mexico. 

Brazil. 

Lironeca tenuistylis (Richardson, 1912b) 

Figure 83E 

DIAGNOSIS 9 13 mm. Cephalon barely immersed in pereonite 1. Uropodal 

rami reaching beyond rounded posterior margin of pleotelson; exopod longer 

than endopod; endopod slender, about three times longer than wide. Pleonite 

1 barely immersed in pereonite 7. Pleotelson basally wider than long. Attach 

ing to host between pectoral and anal fin. 

RECORDS Longnose anchovy Anchoa lamprotaenia: Panama. 

Mothocya Costa, 1851 

DIAGNOSIS Cephalon more or less immersed in pereonite 1. Bases of anten- 

nules widely separated; antennules longer and more robust than antennae. 

Coxae nearly reaching or extending beyond posterior margin of respective 

pereonites. Pleon somewhat immersed in pereonite 7. Uropodal exopod 

longer than endopod. 

REMARKS Bruce (1986b) revised the genus Mothocya, The species of Moth 

ocya are almost entirely gill parasites on the fish families Hemiramphidae, 

Apogonidae, Belonidae, and Atherinidae. 

Key to species of Mothocya 

1. Cephalon anteriorly narrowed, slightly immersed in pereonite 1; 

pleotelson subrectangular bohlkeorum 

Cephalon anteriorly broad, deeply immersed in pereonite 1; pleotelson 

subtriangular nana


Mothocya bohlkeorum Williams and Williams, 1982 

Figure 84B 

DIAGNOSIS Ovigerous 9 7.6-8.5 mm, 6 3.7 mm. Cephalon anteriorly nar 

rowed in dorsal view, ventrally flexed, broadly rounded; slightly immersed in 

pereonite 1. Pleotelson subrectangular. Uropods extending slightly beyond 

posterior margin of pleotelson; exopod only slightly longer than endopod. 9 

lateral lobes of pleopodal peduncles not developed. Endopods of pleopods 3- 

5 with small proximomedial lobe. 

RECORDS Whitestar cardinalfish Apogon lachneri: Puerto Rico. Dusky car- 

dinalfish Phaeoptyx pigmentaria: Bahamas. Freckled cardinalfish Phaeoptyx con- 

klini: Florida Keys; Bahamas. Conchfish Astrapogon stellatus: Leeward 

Islands. 

Mothocya nana (Schioedte and Meinert, 1884) 

Figure 84A 

DIAGNOSIS Ovigerous 9 11.0-17.0 mm, 6 7.9-8.3 mm. Cephalon deeply 

immersed in pereonite 1; rostrum anteroventrally narrowly rounded. Uropo- 

dal exopod markedly longer than endopod. Pleotelson broad, with posterior 

margin rounded sufficiently to give appearance of being subtriangular. 

RECORDS Halfbeak Hyporhamphus unifasciatus: Chesapeake Bay, Maryland; 

Georgia; Florida; Colon, Panama. Halfbeak Hemiramphus bermudensis: 

Bermuda. 

Nerocila Leach, 1818 

DIAGNOSIS Body generally more depressed than in most cymothoid genera, 

rarely curved. Cephalon with anterior margin convex, narrowly rounded, or 

concave; not, or only slightly, immersed in pereonite 1. Pereonite 1 anterior 

margin trisinuate. Posterolateral angles of pereonites weakly to strongly pro 

duced, increasing in length posteriorly. Coxal plates prominent, usually al 

most reaching or extending to posterior margin of their respective pereonites. 

Juveniles and 6 usually with spines on posterior pereopods; 9 lacking these 

spines. Pleon not immersed in pereonite 7. Pleonites subequal in length; 

pleonites 1 and 2 usually produced posterolateral^. Pleopods typically with 

small lamellar accessory gills; pleopods 3-5 folded into deep pockets or 

pleats. Uropods usually extending beyond pleotelsonic apex.

Figure 84. A, Mothocya nana; By Mothocy 

acuminata; D, Nerocila acuminata f. aster. 

B


Nerocila acuminata Schioedte and Meinert, 1881 

DIAGNOSIS Ovigerous 9 16.2-19.0 mm. Cephalon with anterior margin 

convex. Posterolateral angles of all, or of posterior pereonites only, produced 

into acute or subacute angles. 

RECORDS Striped burrfish Chilomycterus schoepfi: Texas, Gulf of Mexico. 

Northern puffer Sphoeroides maculatus: New York. Striped mullet Mugil 

cephalus: Texas, Gulf of Mexico. Jewfish Epinephelus itajara: Texas, Gulf of 

Mexico. Hogfish: Bermuda. Alligator gar Lepisosteus spatula: Louisiana, Gulf 

of Mexico. Hardhead catfish Ariusfelis: Texas, Gulf of Mexico. Sawfish: Flor 

ida (Atlantic). Black drum Pogonias cromis: Texas, Gulf of Mexico. Orange 

filefish Alutera schoepfi: Texas, Gulf of Mexico. Toadfish Batrachoides sur- 

inamensis: Colon, Panama. Spot Leiostomus xanthurus: Florida, Gulf of Mexico. 

Spadefish Chaetodipterus faber: Florida, Gulf of Mexico; Virginia. Fringed fil 

efish Monacanthus ciliatus: Florida, Gulf of Mexico. No host recorded: Mas 

sachusetts; Florida Keys; Florida, Gulf of Mexico. Louisiana, Gulf of Mex 

ico. Texas, Gulf of Mexico. 

REMARKS Brusca (1981) has shown that this highly variable species occurs 

on both sides of the Isthmus of Panama, in two relatively distinct forms. 

Intergrades between the two forms do occur but are uncommon. Brusca 

(1981:159) also lists all the host-records for this species in the eastern Pacific. 

Nerocila acuminata Schioedte and Meinert, 1881, forma acuminata 

Figure 84C 

DIAGNOSIS Cephalon width equal to or greater than length; frontal margin 

narrowly rounded. Posterolateral angles of anterior pereonites weakly pro 

duced, rounded to subacute; of posterior pereonites more strongly produced, 

subacute to acute. Coxal plates 3—7, 4—7, or 5—7 with acute posterolateral 

angles; coxae rarely reaching beyond posterior margins of their respective 

pereonites. 

Nerocila acuminata Schioedte and Meinert, 1881, forma aster 

Figure 84D 

DIAGNOSIS Cephalon always wider than long; anterior margin broadly 

rounded. Posterolateral angles of all pereonites strongly produced, acute, all 

reaching well beyond posterior margins of their respective pereonites. Coxal 

plates 2—7 strongly produced with acute posterior angles.

Renocila Miers, 1880 

Renocila colini 191 

DIAGNOSIS Body rarely curved. Cephalon anteriorly weakly to distinctly 

truncate. Antennular bases well separated. Antennules and antennae some 

what flattened, antennules usually broader and longer than antennae. Per- 

eonites 5—7 with posterolateral corners more or less strongly produced. 

Pleonites not laterally incised. 

REMARKS Williams and Williams (1980) provide a key to nine species of 

Renocila. 

Key to species of Renocila 

1. Posteroventral angle of pereonite 7 reaching pleonite 1 colini 

Posteroventral angle of pereonite 7 reaching beyond pleonite 1 2 

2. Dorsal surface of body brown; posteroventral angle of pereonite 7 

reaching pleonite 2 waldneri 

Dorsal surface of body black; posteroventral angle of pereonite 7 

reaching pleonite 3 bowmani 

Renocila bowmani Williams and Williams, 1980 

Figure 85A 

DIAGNOSIS 9 18.0 mm, 6 11.5 mm. Posteroventral angles of pereonites 5— 

7 produced, that of pereonite 7 overlapping pleonites 1-3. Pereopods 1—3 

lacking swelling on dactylus. Pereopods 6-7 subequal in length. Uropodal 

exopod longer than endopod. Pleotelson length 3 A basal width. Color: dorsal 

surface of body and appendages uniform black. Attached to dorsum of body 

close to dorsal fin. 

RECORDS Harlequin bass Serranus tigrinus: Dominican Republic. 

Renocila colini Williams and Williams, 1980 

Figure 85B,C 

DIAGNOSIS Ovigerous 9 12.0-17.5 mm, 6 7.5-13.0 mm. Pereonites 5-7 

with posteroventral angle produced, that of pereonite 7 overlapping pleonite 

1 only. Pereopods 1—3 lacking swelling on dactyli; pereopods 6—7 subequal in 

length. Uropod reaching beyond pleotelson, endopod more than half length


c 

Figure 85. A, Renocila bowmani. Renocila colini: B, 9; C, d. D> Renocila waldneri. 

of exopod. Pleotelson l /i to 72 wider than long, with slight rounded apex 

Color: dorsal surface of body and appendages uniformly yellowish brown 

Attached to dorsum of body, close to dorsal fin.

FLABELLIFERA • L1MNOR11DAE 193 

RECORDS Flamefish Apogon maculatus: Puerto Rico. Belted cardinalfish Ap- 

ogon townsendi: Puerto Rico. 

Renocila waldneri Williams and Williams, 1980 

Figure 85D 

DIAGNOSIS Ovigerous 9 15.3-19.3 mm, 6 5.0-10.8 mm. Posteroventral 

angle of pereonite 5 moderately produced, of pereonites 6-7 more strongly 

produced, that of pereonite 7 overlapping pleonites 1 and 2. Pereopods 1-3 

without swelling on dactyli. Pereopods 6 and 7 subequal in length. Uropodal 

exopod slightly longer than endopod. Pleotelson basally wider than long; 

posterior margin broadly and evenly rounded. Color: dorsal surface of body 

uniform brown; appendages yellowish brown. Attached to dorsum of body 

close to dorsal fin. 

RECORDS Harlequin bass Serranus tigrinus: Dominican Republic. 

Family Limnoriidae Harger, 1879 

DIAGNOSIS Body ovate in cross section, often becoming more setose posteri 

orly. Cephalon subspherical, freely articulating with pereonite 1; eyes lateral. 

Antennules and antennae well separated at bases. Mandible with strong in 

cisor; lacking molar and well-defined lacinia mobilis, but with species- 

distinctive lacinioid bristle or seta; palp usually of three articles. Maxillipe- 

dal palp of five articles; endite well developed. Coxae present on pereonites 

2-7. Pleon consisting of five free pleonites plus pleotelson; latter subcircular, 

set obliquely to axis of body, usually with anterolateral crests. Uropod with 

strong protopod inserted ventrolaterally. 

Key to genera of Limnoriidae 

1. Uropodal rami very unequal 2 

Uropodal rami subequal Paralimnoria 

2. Mandibular incisors possessing rasp and file Limnoria 

Mandibular incisors lacking rasp and file Phycolimnoria

194 FLABELLIFERA • LIMNORIIDAE 

REMARKS This family includes a number of species that are of considerable 

economic importance. Given that species of Limnoria are wood borers, 

wooden structures such as wharf pilings that are immersed in sea water and 

even in water of reduced salinity are vulnerable to attack by these gribbles. 

Prolonged exposure can lead to weakening and eventual collapse of these 

structures (see Ray, 1959). Even creosote-treated wood is not fully protected; 

Limnoria tuberculata will bore into such wood to where the creosote has not 

penetrated. 

The isopods rasp at the wood fibres with the rasp and file structures of the 

mandibles, usually following the grain of the wood. With this boring activity, 

saprophytic fungi and bacteria invade the wood and assist in the breakdown 

process. Limnoria lack cellulase-secreting microflora in their gut, but proba 

bly secrete a cellulase themselves (Boyle and Mitchell, 1978). It is also prob 

able that the fungi and bacteria, the latter often densely aggregated on the 

setae of the isopod, form part of the animals' diet. In the natural environ 

ment, Limnoria perform an important role in the breakdown of dead wood, 

especially in mangrove areas. 

Sexual dimorphism of the pleotelson does occur in some species. This as 

pect of the morphology, however, has hardly been investigated. 

Limnoria Leach, 1814 

DIAGNOSIS Antennular flagellum of four articles. Antennal flagellum of 

three to five articles. Incisor of right mandible equipped with filelike struc 

ture on upper surface; incisor of left mandible with rasplike structure. Rami 

of pleopod 5 lacking marginal setae. Uropodal exopod much shorter than 

endopod, bearing terminal claw. Pleotelson smooth, or variously ornamented 

with tubercles and ridges. 

Limnoria indica Becker and Kampf, 1958 

Figure 86A,B 

DIAGNOSIS 6 3.0 mm, ovigerous 9 3.0 mm. Pleonite 5 with submedian pair 

of strong rounded ridges, converging slightly posteriorly. Pleotelson basally 

with two pairs of submedian tubercles and pair of lateral tubercles. 

RECORDS Cozumel, Mexico; Man o'War Cay, Belize. 

India; Hong Kong; Philippines; east coast of Australia.

Key to species of Limnoria 

Limnoria insulae 195 

1. Dorsal surface of pleotelson lacking prominent tubercles, ridges, or 

carinae (L. simulata may appear to lack ornamentation; in this 

species the tubercles are very small) 2 

Dorsal surface of pleotelson bearing tubercles, ridges, or carinae .... 3 

2. Pleotelson flat; pleonite 5 with broadly rounded middorsal ridge 

Pleotelson cup shaped; pleonite 5 with strong narrowly rounded 

platycauda 

middorsal ridge insulae 

3. Pleotelson with basal tubercles but lacking ridges 4 

Pleotelson with ridges but lacking freestanding tubercles 7 

4. 6 pleotelson with single strong middorsal tubercle unicornis 

Pleotelson with more than one basal tubercle 5 

5. Pleotelson with three basal tubercles tuberculata 

Pleotelson with more than three basal tubercles 6 

6. Pleotelson with four basal tubercles in line (difficult to detect) simulata 

6 pleotelson with six basal tubercles indica 

7. Pleotelson with single middorsal longitudinal ridge multipunctata 

Pleotelson with two rounded basal ridges 8 

8. Pleonite 5 with strong Y-shaped ridge Pfeffe 

Pleonite 5 with two posteriorly converging ridges saseboensis 

Limnoria insulae Menzies, 1957 

Figure 86C 

DIAGNOSIS 6 3.0 mm, ovigerous 9 3.4 mm. Pleonite 5 with strong middor 

sal ridge. Pleotelson cup shaped, lateral crests extended anteromesially, sep 

arated basally by distinct gap; posterior margin and lateral crests not 

tuberculate. 

RECORDS Twin Cays, Belize.


200HM 

Figure 86. Limnoria indica: A, pi 6;B, pi 9. Limnoria insulae: C 

pleotelson. Limnoria multipunctata: D, pleotelson in oblique-lateral view 

L i m n o r i a m u l t i p u n c t a t a M e n z i e s , 1 9 5 7 

F i g u r e s 8 6 D ; 8 7 A 

D I A G N O S I S 6 2 . 8 m m , o v i g e r o u s 9 3 . 0 m m . P l e o n i t e 5 d o r s a l l y s m o o t h . 

P l e o t e l s o n w i t h m i d d o r s a l l o n g i t u d i n a l r o u n d e d r i d g e b e a r i n g s e v e r a l

Limnoria multipunctata 197 

Figure 87. Limnoria multipunctata: A, pleotelson; Limnoria pfefferi: B} pleotelson 

Limnoria platycauda: C, pleotelson; Limnoria saseboensis: D> pleotelson; Limnoria 

simulata: E, pleotelson; Limnoria tuberculata: F, pleotelson. 

b u t t o n - s h a p e d t u b e r c l e s i n p o s t e r i o r h a l f ; p o s t e r i o r m a r g i n a n d l a t e r a l c r e s t s 

t u b e r c u l a t e . 

R E C O R D S P u e r t o R i c o ; J a m a i c a ; T w i n C a y s , B e l i z e 

J a p a n ; K a i I s l a n d s , S o u t h P a c i f i c .


Limnoria pfefferi Stebbing, 1904 

Figure 87B 

DIAGNOSIS 8 3.8 mm, ovigerous 9 4.0 mm. Pleonite 5 with conspicuous 

middorsal Y-shaped carina. Pleotelson basally with pair of submedian 

rounded ridges; lateral crests lacking tubercles. 

RECORDS Florida Keys; Bahamas; Puerto Rico; U.S. Virgin Islands; Twin 

Cays and Man o'War Cay, Belize; Yucatan Peninsula, Mexico. 

Minikoi Atoll and Aldabra Atoll, Indian Ocean; Philippines; New Guinea; 

Panama. 

Limnoria platycauda Menzies, 1957 

Figure 87C 

DIAGNOSIS 6 2.5 mm, ovigerous 9 2.6 mm. Pleonite 5 with broad middor 

sal longitudinal rounded ridge. Pleotelson lacking dorsal ornamentation; 

posterior margin and lateral crests bearing tubercles. 

RECORDS Cuba; Puerto Rico to Curasao; Cozumel, Mexico; Twin Cays 

and Man o'War Cay, Belize. 

Aldabra Atoll, Indian Ocean. 

Limnoria saseboensis Menzies, 1957 

Figure 87D 

DIAGNOSIS 6 3.5 mm. Pleonite 5 with submedian pair of ridges, converging 

slightly posteriorly. Pleotelson basally with submedian pair of anteriorly tu- 

berculate ridges; posterior margin and lateral crests tuberculate. 

RECORDS Miami, Florida. 

Japan; Fiji. 

Limnoria simulata Menzies, 1957 

Figure 87E 

DIAGNOSIS 6 3.8 mm, ovigerous 9 4.0 mm. Pleonite 5 with obscure me 

dian longitudinal groove. Pleotelson basally with submedian pair of tubercles 

and small lateral tubercles, latter often difficult to detect; lateral crests 

tuberculate. 

RECORDS Florida Keys; U.S. Virgin Islands; Gulf of Mexico.

Limnoria tuberculata Sowinsky, 1884 

Figure 87F 

Paralimnoria andrewsi 199 

DIAGNOSIS 6 2.8 mm, ovigerous 9 3.0 mm. Pleonite 5 with two anterior 

tubercles, one middorsal posterior tubercle, area between tubercles de 

pressed. Pleotelson basally with middorsal tubercle, followed by pair of sub- 

median tubercles, all three tubercles having short obscure carina; posterior 

margin and lateral crests tuberculate. 

RECORDS Rhode Island to Venezuela; Cuba; Man o'War Cay, Belize; Gulf 

of Mexico. 

Uruguay; West Africa; Mediterranean; Black Sea; India; Hong Kong; 

Hawaii; Australia; California. 

REMARKS This species has frequently been recorded under the name Lim 

noria tripunctata Menzies, 1951a. 

Limnoria unicornis Menzies, 1957 

Figure 88A,B 

DIAGNOSIS 6 2.6 mm, ovigerous 9 2.6 mm. Mandibular palp of one arti 

cle. Pleonite 5 with somewhat obscure Y-shaped ridge middorsally. 

Pleotelson in 6 with strong basal slightly curved middorsal tubercle; lateral 

crests lacking tubercles. 

RECORDS Bahamas; Man o'War Cay and Twin Cays, Belize. 

Caroline Islands; Palau; Society Islands. 

Paralimnoria Menzies, 1957 

DIAGNOSIS Antennular flagellum of five articles. Antennal flagellum of five 

or six articles. Mandibular incisor with rasp and file. Pleopod 5, rami bearing 

marginal setae. Uropodal rami subequal in length, each with clawlike apex. 

Paralimnoria andrewsi (Caiman, 1910) 

Figure 88C,D 

DIAGNOSIS 6 2.6 mm, 9 2.6 mm. Pleonite 5 with or without triangular 

middorsal depressed area. Pleotelson with basal submedian pair of tubercles 

either obscurely or strongly carinate; lateral crest tubercles of variable 

strength.


Figure 88. Limnoria unicornis: A, pleotelson, 8; B, pleon, 6*, in lateral view. 

Paralimnoria andrewsi: C, p l e o n i t e 5 a n d p l e o t e l s o n ; D , u r o p o d . Phycolimnoria clarkae: 

E , p l e o n i t e 5 a n d p l e o t e l s o n ; F , u r o p o d a n d p l e o t e l s o n i n l a t e r a l v i e w . 

R E C O R D S F l o r i d a K e y s ; P u e r t o R i c o ; T w i n C a y s , B e l i z e ; C u r a g a o 

C h r i s t m a s I s l a n d s , I n d i a n O c e a n ; S a m o a ; H a w a i i ; J a p a n . 

R E M A R K S M e n z i e s ( 1 9 5 7 ) d i s c u s s e s t h r e e f o r m s o f t h i s s p e c i e s : F o r m a typ- 

i c a , w h i c h l a c k s a c e n t r a l d e p r e s s e d a r e a d o r s a l l y o n p l e o n i t e 5 a n d h a s a p a i r 

o f s u b m e d i a n o b s c u r e l y c a r i n a t e t u b e r c l e s o n t h e p l e o t e l s o n ; F o r m a A , w h i c h 

h a s a t r i a n g u l a r d e p r e s s e d a r e a d o r s a l l y o n p l e o n i t e 5 a n d a p a i r o f s u b m e -

FLABELLIFERA • SEROLIDAE 201 

dian tubercles supported by strong carinae on the pleotelson; Forma B5 hav 

ing a triangular depressed area dorsally on pleonite 5 and an obscurely cari- 

nate pair of tubercles on the pleotelson. Given that at least two of these forms 

have been recorded occurring together, it would seem that this is merely a 

highly variable species. 

Phycolimnoria Menzies, 1957 

DIAGNOSIS Mandibular incisor lacking rasp and file. Uropodal rami une 

qual, exopod longer than endopod, latter usually with clawlike apex. 

REMARKS Most species of Phycolimnoria are algal borers, frequently encoun 

tered in the holdfasts of brown algae such as Macrocystis, Laminaria, and 

Sargassum, The one species recorded from the Caribbean, P. clarkae, however, 

has only been taken from decaying wood. 

Phycolimnoria clarkae Kensley and Schotte, 1987 

Figure 88E,F 

DIAGNOSIS 6 4.3 mm, ovigerous 9 3.3-4.4 mm. Uropodal exopod less 

than half length of endopod, straight, tipped with short squat claw. Pleonite 

5 with broad raised middorsal region having irregular bumps. Pleotelson 

wider than long, with two rounded submedian ridges basally, becoming ob 

solete posteriorly. 

RECORDS Bahamas; Twin Cays, Belize. 

Aldabra Atoll, Indian Ocean. 

Family Serolidae Dana, 1852 

DIAGNOSIS Body dorsoventrally depressed. Eyes present or absent. 

Cephalon fused with pereonite 1 dorsally. Mandible bearing palp. Maxillipe- 

dal palp of one to four articles. Pereonites 2—4 with coxae demarked; per- 

eonites 5 and 6 with coxae not demarked; pereonite 7 narrow, lacking free 

lateral margins. Pereopod 1 in 6 and 9 subchelate, pereopod 2 subchelate or 

ambulatory in 6, ambulatory in 9. Pleonites 1 and 2 free, articulated, re 

mainder of pleonites fused with telson. Pleopods 1—3 small, natatory; 

pleopods 4 and 5 large, operculate. Uropods lateral, biramous. 

REMARKS The serolids reach their greatest diversity (and their greatest size 

of up to 80 mm in length) in the southern oceans, with few species extending

202 FLABELLIFERA • SPHAEROMATIDAE 

into the subtropics and tropics. The deep- and abyssal-dwelling species usu 

ally lack eyes. The animals are epibenthic, living in the upper few centime 

ters of the bottom sediment, where they are scavengers and carnivores. 

Serolis Leach, 1818 

DIAGNOSIS Body markedly dorsoventrally flattened. Coxal plates produced 

laterally. Mandible having lacinia mobilis and single spine. Maxillipedal 

palp of three articles (rarely two to four). Pereopod 2 exhibiting sexual di 

morphism, subchelate in cJ, ambulatory in 9. Pleopods 1-3, peduncles elon 

gate, rami subelliptical. Pleopod 3, exopod uniarticulate. 

Serolis mgrayi Menzies and Frankenberg, 1966 

Figure 89 

DIAGNOSIS S 4.5 mm, ovigerous 9 4.7 mm. Eyes present. Cephalon with 

two middorsal tubercles. Pereonites 2—4 each with faint rounded tubercle 

just mesial to coxal suture. Pereon and pleon with faint middorsal longitudi 

nal carina bearing small blunt tubercle on posterior margin of each segment. 

Pleonites 1 and 2 with lateral margins not contributing to body outline, over 

lapped by pereonite 6. Pleotelson broadly triangular, with lateral carina in 

anterior half; apex truncate. Uropodal rami reaching to or slightly beyond 

pleotelsonic apex. 

RECORDS Off North Carolina, 18-34 m; off South Carolina, 22 m; off 

Georgia, 18-47 m; Florida Keys, 18-88 m; Trinidad; Venezuela, 95 m; Flor 

ida, Gulf of Mexico, 11-88 m. 

Family Sphaeromatidae H. Milne Edwards, 1840 

DIAGNOSIS Antennular peduncle of three articles, antennal peduncle of five 

articles. Mandible stout, lacinia mobilis and molar usually well developed, 

palp of three articles. Maxillipedal palp of five articles. Mouthparts in some 

genera metamorphosed and somewhat reduced in ovigerous 9 . Pleon of five 

partially or completely fused pleonites, often indicated by lateral sutures, 

plus dorsally convex and sometimes inflated pleotelson. Uropods lateral, ex 

opod free if present, endopod fused with sympod. Sexual dimorphism often 

marked, especially in pleotelsonal structure. Animal often capable of con 

globating or folding over. Young brooded in internal pouches or anterior or 

posterior pockets; oostegites variable in number, if present.

FLABELLIFERA • SPHAEROMATIDAE 203 

Figure 89. Serolis mgrayi: A, 6; B, pereopod 1; C, pereopod 2, 8. 

REMARKS Right into the 1980s this family was routinely divided into three 

groups, based on the structure of the two posterior pairs of pleopods: 

Platybranchiatae—pleopods 4 and 5 with both rami membranous and lack 

ing branchial pleats; Hemibranchiatae—pleopods 4 and 5 with branchial 

pleats on endopods only; Eubranchiatae—pleopods 4 and 5 with branchial 

pleats on both rami. These three "groups" were recognized formally as sub 

families by Hurley and Jansen (1977) but the names were not based on con-

204 FLABELL1FERA • SPHAEROMATIDAE 

tained genera and were replaced with current subfamily names by Bowman 

(1981) and Iverson (1982), the latter providing diagnoses for all five sub 

families. Four of these are represented in the Caribbean area; the fifth, the 

Tecticipitinac, contains only the single primarily Pacific genus Tecticeps. 

While the subfamilial status now appears to be resolved, many of the gen 

era still require unambiguous diagnoses. The work of Harrison (1984) on the 

structure of the female broodpouch, with its various components of 

oostegites, internal pouches, and anterior and posterior pockets (Figure 90), 

along with the metamorphosis of the female mouthparts (see Figure 96) has 

helped enormously to standardize the genera. Nevertheless, these features of 

the female remain unknown in several genera. Further, with this stabilization 

based on females, many problems of incorrect generic designation have been 

uncovered. In this work, Harrison's generic diagnoses are followed as far as 

possible. Where uncertainty exists, this is indicated. In some cases, we may 

still be unaware of existing problems: future work will without doubt result in 

the shifting of species to different genera, as well as in the creation of new 

genera. 

Key to subfamilies of Sphaeromatidae 

1. Pereopod 1 prehensile in both sexes; pereopod 2 prehensile only in 6 

Ancininae 

Pereopods 1 and 2 ambulatory 2 

2. Pleopods 4 and 5 lacking branchial pleats Cassidininae 

Pleopods 4 and 5 with branchial pleats on endopods 3 

3. Pleopods 4 and 5 with branchial pleats on both rami . . . Dynameninae 

Pleopods 4 and 5 with branchial pleats on endopods only 

Subfamily Ancininae Tattersall, 1905 

Sphaeromatinae 

DIAGNOSIS Body markedly dorsoventrally depressed. Cephalon fused me 

dially with pereonite 1. Pereopod 1 prehensile in 8 and 9. Pereopod 2 pre 

hensile in 6 only. Pleopods 4 and 5 similar, lacking branchial pleats. 

Uropods uniramous.

internal pouch 

Ancinus belizensis 205 

Figure 90. Diagrammatic representation of 9 sphaeromatid, showing marsupial 

structures (adapted from Harrison, 1984). 

Ancinus H. Milne Edwards, 1840 

DIAGNOSIS Eyes dorsal. 9 mouthparts not metamorphosed. Mandibular 

molar absent; palp of three articles. Maxilla 1 of single ramus, endite rudi 

mentary. Maxilla 2 of two rami. 9 with oostegites absent; brood held in two 

opposing pockets, opening as narrow ventral slit between pereopods 4. Pleon 

consisting of short anterior pleonite with free lateral margin, plus broadly 

triangular pleotelson. Pleopod 1 uniramous, endopod absent. Pleopod 2 op- 

erculiform. Pleopod 3, exopod of single article. Uropod lacking exopod, sym- 

pod not laterally expanded. 

Key to species of Ancinus 

1. Pleotelson as long as basal width, apex narrowly rounded . . . brasiliensis 

Pleotelson with basal width greater than length, apex subtruncate 

Ancinus belizensis Kensley and Schotte, 1987 

Figure 91A-C 

belizensis 

DIAGNOSIS 8 4.1 mm, 9 2.8 mm. Body oval, about twice longer than wide. 

Dorsal integument strongly pitted. Antennular flagellum of 12 articles; an-


Figure 91. Ancinus belizensis: A, 9; B, pereopod 1 cJ; C, pereopod 2 8. Ancinus 

braziliensis: D, adult (from Glynn and Glynn, 1974). 

tennal flagellum of 10 articles. 8 pereopod 2, dactylus strongly curved, 

reaching to proximal lobe of propodus. Pleopod 2 about 2.5 times longer than 

basal width. 

RECORDS Carlson Point, Belize, in seagrass flats, 0.5 m. 

Ancinus brasiliensis Lemos de Castro, 1959 

Figure 91D 

DIAGNOSIS 8 7.0 mm, 9 6.0 mm. Body about twice longer than wide. Dor 

sal integument smooth. Antennular flagellum of 17 articles; antennal 

flagellum of 10 articles. 8 pereopod 2, dactylus strongly curved, reaching to 

midlength of posterior margin of carpus. Pleopod 2 almost three times longer 

than basal width.

Cass idin idea 207 

RECORDS Brazilian coast from Rio de Janeiro northward, 1.5 m; Costa 

Rica, Panama; shallow infratidal below sandy beaches. 

REMARKS Glynn and Glynn (1974) discussed color polymorphism in this 

species. 

Subfamily Cassidininae Iverson, 1982 

DIAGNOSIS Cephalon not medially fused with pereonite 1. Pereopod 1 am 

bulatory. Pleopods 4 and 5, both rami lacking transverse pleats, outer rami 

unsegmented. Pleopod 5, outer ramus with low subapical squamiferous pro 

tuberances. Pleotelsonic apex entire. Uropods with exopods reduced. 

REMARKS The genus Dies has twice been recorded from the Caribbean: D. 

arndti Ortiz and Lalana, 1980, from Cuba, and D. barnardi Carvacho, 1977, 

from Guadeloupe. This genus is distinguished from Cassidinidea solely on the 

basis of the penial structure: biramous in Cassidinidea, uniramous in Dies. 

Harrison (1984) has pointed out that the separation of these two genera has 

not been satisfactorily resolved. The penis of neither the Cuban nor the 

Guadeloupan species has been illustrated, but the whole-animal illustrations 

of both look suspiciously like Cassidinidea ovalis. Examination of material of D. 

barnardi from the Paris Museum supports the view that this species was based 

on immature material of C. ovalis. Neither of the so-called species oi Dies are 

dealt with in this work, both being regarded as junior synonyms of C. ovalis. 

Key to genera of Cassidininae 

1. Frontal lamina visible dorsally between antennular bases; two basal 

articles of antennular peduncle not expanded Cassinidinea 

Frontal lamina not visible between antennular bases; two basal articles 

of antennular peduncle broadly expanded Paraleptosphaeroma 

Cassidinidea Hansen, 1905b 

DIAGNOSIS Body strongly dorsoventrally depressed. Eyes dorsal, situated 

at posterolateral corners of cephalon. Latter somewhat sunken into pereonite 

1. Frontal lamina expanded, visible dorsally between antennular bases. An 

tenna directed laterally. Pleon consisting of one free pleonite having short 

free lateral margin, plus broadly triangular pleotelson. Uropodal endopod


well developed, fused with sympod; exopod markedly reduced. Penial rami 

elongate, separate. 9 mouthparts not metamorphosed. Oostegites absent. 

Brood housed in pouch formed by opposing pockets overhanging ventrum, 

opening by slit between fourth pereopods. 

Key to species of Cassidinidea 

1. Posterior margin of pleotelson truncate ovalis 

Posterior margin of pleotelson rounded mosaica 

Cassidinidea mosaica Kensley and Schotte, 1987 

Figure 92A 

DIAGNOSIS 6 1.8 mm, ovigerous 9 1.6 mm. Body twice longer than wide. 

Dorsal integument bearing close-packed flattened tubercles. Pleotelson tri 

angular, with posterior margin narrowly rounded, dorsally convex, basally 

inflated. 

RECORDS Carrie Bow Cay, Belize, 1.5-10 m; in silty sand and rubble be 

tween patch reefs and coral buttresses. 

Cassidinidea ovalis (Say, 1818) 

Figure 92B-E 

DIAGNOSIS 6 and 9 3.6 mm. Body width slightly less than half length. 

Dorsal integument smooth. Pleotelson with raised anteromesial area, but 

lacking sculpture; posterior margin truncate. 

RECORDS New Jersey to Florida, in marsh mud and among dead leaves, 0— 

1 m; Trinidad; Belize; Panama; Dominica; Louisiana and Vera Cruz, Gulf of 

Mexico. Known from waters of less than l%o to 35%o. 

Paraleptosphaeroma Buss and Iverson, 1981 

DIAGNOSIS Body oval in outline, entire circumference with transparent 

flange of fused setae on two expanded basal articles of antennule, on per- 

conites, pleonite 1, and uropods. Expanded basal articles of antennules con-

Paraleptosphaeroma 209 

Figure 92. Cassidinidea mosaica: A, 6. Cassidinidea ovalis: B, 6] C, pereopod 1; Dy 

pleopod 4; E, pleopod 5. Paraleptosphaeroma glynni: Fy 6. 

tiguous in midline. Single articulated pleonite with short free lateral margin. 

Uropodal sympod and endopod fused; exopod articulated, much shorter 

than fused endopod.


Paraleptosphaeroma glynni Buss and Iverson, 1981 

Figure 92F 

DIAGNOSIS 6 2.58 mm, ovigerous 9 2.38 mm. Pleotelson basally broad, 

tapering to notched posterior margin. Fused uropodal endopod and sympod 

of each side almost touching posterior to pleotelsonic apex. 

RECORDS Portsmouth, Dominica, intertidal rock pools. 

Punta Paitilla, Pacific Panama. 

REMARKS Buss and Iverson (1981) demonstrated that this species displays 

sequential protogynous hermaphroditism, and that the change from female 

to male seems to be mediated by social conditions, especially the proportion 

of males to females. The principal food source for this species was shown to 

be abascan bryozoans. 

Key to genera of Dynameninae 

1. Pleotelson very similar in both sexes 2 

Pleotelson showing marked sexual dimorphism 3 

2. Cephalon and pleotelson smooth, lacking ridges Ischyromene 

Pleotelson and cephalon with ridges Cerceis 

3. Uropods lamellar in both sexes 4 

Uropods lamellar in 9, endopod reduced, exopod elongate-cylindrical 

in 6 5 

4. Ovigerous 9 lacking oostegites;

Subfamily Dynameninae Bowman, 1981 

Discerceis 211 

DIAGNOSIS Cephalon not fused with pereonite 1. Pereopods 1 and 2 am 

bulatory. Pleopods 4 and 5, both rami having branchial pleats. Pleopod 4, 

exopod unjointed, usually lacking setae, endopod with few setae at most. 

Pleotelsonic apex often with terminal notch or foramen, especially in


Figure 93. "Cerceis" carinata: A, 6. Discerceis linguicauda: B, d. Dynamenella 

acutitelson: C, 9; D, pleon 8. Dynamenella angulata: E, 9. 

internal pouches (number unknown). Pockets absent. 6 with uropodal endo- 

pod and sympod fused, very short; exopod elongate, cylindrical.

Discerceis linguicauda (Richardson, 1901) 

Figure 93B 

Dynamenella 213 

DIAGNOSIS 6 7.2 mm. Dorsal integument, especially of posterior half, with 

numerous scattered granular tubercles. Uropodal endopod and sympod 

fused, very short, exopod elongate, subcylindrical and slightly bowed. Ante 

rior half of pleotelson inflated, with three elongate rounded ridges (each com 

posed of two contiguous tubercles) ending posteriorly in subacute tubercle; 

posterior margin trilobed, median lobe broadly rounded, with subacute tu 

bercle at base, lateral lobe truncate, well separated from median lobe. Head 

and pereonite 1 not fused. Frontal lamina visible dorsally between antennal 

bases. Penes short, separate. Copulatory stylet basally relatively broad, dis- 

tally broadly rounded. 

RECORDS Cape Catoche, Yucatan, Mexico, 48-50 m. 

REMARKS This species is known only from the four male syntypes. 

Dynamenella Hansen, 1905b 

DIAGNOSIS Species exhibiting obvious sexual dimorphism. Both sexes lack 

ing processes on pereon and pleon, Uropodal rami lamellar. Exopod of 

pleopod 3 with or without articulation. 9: Mouthparts not metamorphosed. 

Broodpouch lacking oostegites, but formed by two opposing ventral pockets 

opening in midline between fourth pereopods. Apex of pleotelson with notch, 

Key to species of Dynamenella 

1. 6 with pleotelsonic foramen 2 

6 lacking foramen but with notch, or appearing entire; 9 pleotelson 

with faint notch visible acutitelson 

2. 6 with four strong pleotelsonic ridges; 9 with subcircular pleotelsonic 

foramen quadrilirata 

6 lacking pleotelsonic ridges; 9 with posterior margin of pleotelson 

entire perforata


groove, or foramen. 6: Penes basally fused, rami long, tapering. Copulatory 

stylet proximally broad, tapering to acute tip, reaching to or just beyond 

apex of endopod. Uropods broader than in $. Posterior pleotelson with dor- 

sally directed foramen connected to apex by narrow slit. 

REMARKS The species described by Richardson (1901) as Dynamene angulata 

from No Name Key, Florida, and referred to by some authors as a Dyna- 

menella, while figured here (Figure 93E), is not included in the present key. 

The species is known only from immature females; correct generic placement 

is thus not possible. 

Dynamenella acutitelson Menzies and Glynn, 1968 

Figure 93C,D 

DIAGNOSIS 6 3.5 mm, 9 2.3 mm. 6: Pereonites 4-6 with transverse ridge 

over dorsum, ridge interrupted to form short median section. Pleotelson with 

two submedian and two lateral rounded tubercles basally, two submedian, 

poorly defined ridges in central area; posterior margin tapering in dorsal 

view, with slit eitherjust visible or appearing entire. In lateral view, posterior 

pleotelson seen to be laterally compressed, forming narrow groove. 

RECORDS Puerto Rico, intertidal rocks and algae. 

REMARKS Menzies and Glynn (1968) described this species with two vari 

eties, the holotype as D. acutitelson var. typica, and 11 paratypes as D. acu 

titelson var. glabrothorax. The major difference between these varieties lay in 

the presence of transverse ridges on pereonites 4—6 in typica and their absence 

xnglabrothorax. The holotype, however, at 3.5 mm, would seem to be a mature 

male, while all the paratypes are smaller. The differences described by Men 

zies and Glynn (1968) may thus be due to immaturity. As further compara 

tive material is lacking, these varieties (or whatever their true status) are not 

recognized here. 

Menzies and Glynn (1968, fig. 30a) illustrate D. acutitelson var. glabrothorax 

as having scattered tiny granules over the dorsal integument. These were not 

seen when the type material was reexamined. 

Harrison and Holdich (1982) placed this species in Paradella, based on the 

literature. However, the penes for both varieties are shown as short and sepa 

rate, as in Ischyromene. Again, until further mature males and ovigerous 

females are seen, the generic placement of this species must remain in doubt.

Dynamenella perforata (Moore, 1901) 

Figure 94A,B 

Geocerceis barbarae 215 

DIAGNOSIS 6 3.2 mm, 9 2.6 mm. 6: Pleon bearing two low rounded sub- 

median "mounds." Pleotelson with strongly convex anterior two-thirds, with 

T-shaped foramen. Pleon and pleotelson with numerous scattered small tu 

bercles. Uropodal rami broadly ovate, outer margins crenulate. 9: 

Pleotelson broadly rounded in dorsal view, posterior margin entire. Inner 

uropodal ramus distally subacute. 

RECORDS Bermuda to Puerto Rico, intertidal coral rubble and algae, and 

under chiton Acanthopleura granulate; Dominican Republic; Cuba. 

Dynamenella quadrilirata Kensley, 1984 

Figure 94C-H 

DIAGNOSIS 6 2.6 mm, 9 2.5 mm. 6: Two low rounded submedian tuber 

cles on last pleonite. Anterior half of pleotelson inflated, with four rounded 

longitudinal ridges; posterior half tapered, somewhat dorsally flexed, with 

cordate foramen. Uropodal rami distally rounded, outer margins crenulate 

to dentate. 9: Lacking pleonal tubercles. Pleotelson inflated, unornamented, 

posterior margin forming subcircular foramen. 

RECORDS Carrie Bow Cay, and Twin Cays, Belize; intertidal to 3 m. 

Geocerceis Menzies and Glynn, 1968 

DIAGNOSIS Ovigerous 9 with mouthparts metamorphosed. Broodpouch 

with three pairs of oostegites, on pereonites 2—4, just overlapping in midline. 

Brood held in internal pouches (number unknown). Pockets absent. Uropo 

dal rami lamellar, shorter than pleotelson. 6 uropodal endopod fused with 

sympod, very short; exopod elongate, club shaped. Pleopod 2 with copula- 

tory stylet articulating distally on endopod. 

Geocerceis barbarae Menzies and Glynn, 1968 

Figure 95A-C 

DIAGNOSIS 8 3.3 mm, 9 2.5 mm. Pleopod 3 exopod of single article. Pleon 

with two elongate sutures reaching lateral pleon margin. 6: Frontal lamina 

expanded into ventrally directed beaklike process. Penes separate, relatively


Figure 94. Dynamenella perforata: A, 6; B> pleon 9. Dynamenella quadrilirata: C, 6; 

D, pleon 9; E, pleon

Ischyromene 217 

Figure 95. Geocerceis barbarae: A, 8; B, pleon 9; C,


nounced. Uropodal rami lamellar. 8 pleopod 2 with copulatory stylet 

basally narrow, reaching to or just beyond distal margin of endopod. 

Ischyromene barnardi (Menzies and Glynn, 1968) 

Figure 95D 

DIAGNOSIS 6 4.5 mm, 9 3.7 mm. Both sexes lacking processes on pereon 

and pleon. Accessory unguis of pereopods often bifid. Pleopod 3, exopod of 

single article. Uropodal rami lamellar. 6: Pereonite 7, posterior margin 

bilobed. Penes short, separate to base. 

RECORDS Puerto Rico, intertidal. 

Paracerceis Hansen, 1905b 

DIAGNOSIS Pleopod 3 exopod with transverse suture in distal half. Pleon 

with two long sutures reaching to posterolateral margin. 6: Penial rami 

short, separate. Pleotelson with basal area strongly vaulted; deep posterior 

notch sometimes having denticles on inner margins, and/or median tooth at 

base of notch. Uropodal endopod short, fused with sympod; exopod elongate, 

club shaped. 9: Mouthparts metamorphosed. Mandible fused with 

cephalon. Broodpouch of four pairs of oostegites, three posterior pairs over 

lapping. Brood retained in internal pouches. Uropodal rami subequal, lamel 

lar. Pleon usually less ornamented than in

Paracerceis caudata 219 

3. 6, pleotelsonic notch deep, margins usually with two teeth on each 

side; strong median tubercle on anterior pleotelson bluntly bifid; 9, 

pleotelson with one or two rounded median tubercles and 2 smaller 

tubercles on each side caudata 

6, pleotelsonic notch shallow, with tiny lateral denticles; median 

tubercle of pleotelson conical, acute; 9, pleotelson with three large 

conical acute tubercles and several smaller scattered tubercles in 

anterior half cohenae 

Paracerceis caudata (Say, 1818) 

Figure 96 

DIAGNOSIS 6 8.1 mm, 9 6.4 mm. 8: Pleotelson with blunt median bifid 

tubercle, with two smaller tubercles on each side. Pleotelsonic notch usually 

with two strong denticles on each margin, basal median tooth lacking. 

Uropodal exopod reaching well beyond pleotelson, slightly bowed, with 2—4 

setose bumps on outer margin. 9: Pleonite 5 with three low tubercles. 

Pleotelsonic apex broadly rounded in dorsal view, with two rounded median 

tubercles and two smaller tubercles on each side. Uropodal rami subequal, 

lamellar, outer distal angle of each acute. 

RECORDS Bermuda; New Jersey to Florida Keys; Yucatan to Venezuela; 

Turks and Caicos Islands; Cuba; Puerto Rico; Bahamas; Jamaica; Haiti; St. 

Maartens, 0.2-127 m; St. Lucia; Gulf of Mexico. Found in the following 

algae: Caulerpa, Halimeda, Turbinaria> Amphiroa, Laurencia, Dictyota; between 

sponges and tunicates on red mangrove roots; in coral rubble; in spur and 

groove zone of reefs, lagoon, back reef, seagrass flats, and fringing 

mangroves. 

REMARKS Menzies and Glynn (1968:55, fig. 22f) named and figured P. 

caudata var. brevipes from Puerto Rico. This variant was characterized as hav 

ing the margins of the pleotelsonic notch lacking denticles. Given the con 

siderable variation in ornamentation in this species, we feel that no validity 

can be given to the name "brevipes." 

This is the commonest sphaeromatid in the Caribbean, and it has very 

broad ecological requirements, being found in a wide range of habitats and 

depths.


Figure 96. Paracerceis caudata: A, 8,B, pleon 9; C, mandible 6] D, maxilla 1 6; E, 

maxilla 2 6; F, maxilliped 6; G> mandible 9; Hy maxilla 19;/, maxilla 1 9; J, 

maxilliped 9. 

Paracerceis cohenae Kensley, 1984 

Figure 97A,B 

DIAGNOSIS 6 10.0 mm, 9 7.9 mm. 6: Pereonites each with median tuber 

cle and several smaller lateral tubercles near posterior margin of somite. 

Pleonite 5 with large median conical tubercle. Anterior two-thirds of

Paracerceis glynni 221 

pleotelson inflated, faintly tripartite, with strong median conical tubercle; 

notch in posterior margin shallow, with low median tooth and tiny lateral 

denticles; posterolateral margins finely dentate. Uropodal exopod cylindri 

cal, distally denticulate, six to seven times longer than basal width. 9: Per- 

eon and pleon much as in 6 y but pleotelsonic notch shallower and 

posterolateral margins not denticulate. Uropodal rami subequal, lamellar, 

exopod with distolateral angle acute. 

RECORDS Carrie Bow Cay, Belize, 15-16 m. Only known from sponge 

Callispongia plicifera growing on outer reef slope. 

Paracerceis edithae Boone, 1930 

Figure 97C-E 

DIAGNOSIS 6 4.0 mm, 9 3.1 mm. 6: Posterior three pereonites and 

pleonites each with irregular row of small tubercles near posterior margin, 

densely setulose tubercles becoming spinose more posteriorly. Pleotelson 

with strong median conical tooth in anterior half, flanked by convex spinose 

mound. Pleotelsonic notch deep, with elongate median basal tooth bearing 

strong acute tooth at its base. Lobes of posterior pleotelsonic margin broad, 

flattened, margins denticulate. Uropodal exopod tuberculate, tapering, api- 

cally acute. 9: Integument much less tuberculate-spinose than in 6. Imma 

ture 9, posterior margin of pleotelson with faintly rounded median lobe. In 

mature 9, posterior margin distinctly trilobed. Uropodal rami subequal, 

lamellar, distally rounded, with tiny distolateral spine on exopod. 

RECORDS Bahamas, 60-66 m, in vase sponge; Haiti; Puerto Rico, 20—25 m. 

Paracerceis glynni Kensley, 1984 

Figure 97F,G 

DIAGNOSIS 6 6.4 mm, 9 5.2 mm. 6: Integument becoming strongly setose 

and tuberculate posteriorly from about pereonite 5. Posterior margin of inf 

lated anterior area of pleotelson bearing strong median conical tubercle and 

smaller acute lateral tubercle, with low swelling beneath each lateral tuber 

cle. Posterior notch deep, narrow, with small basal median tooth, lobes form 

ing notch tricuspid, outer cusps recurved dorsally. Uropodal exopod fairly 

straight, cylindrical, apically acute. 9: Body far less setose and tuberculate 

than 6. Pleotelson with strongly inflated anterior area having very faint mid- 

dorsal tubercle; notch well marked, formed by triangular lobes of

222 

FLABELLIFERA • SPHAEROMATIDAE 

E 

Figure 97. Paracerceis cohenae: A, 6\ B, pleotelson, 9. Paracerceis edithae: C, 

pleotelson, 9; D, mature 6] E, immature S. Paracerceis glynni: F, 8\ G} pleotelson, 

9. Paracerceis nuttingi: H, 9. 

pleotelsonic margin. Uropodal rami subequal, flattened, endopod with distal 

margin faintly trituberculate; exopod with few distal tubercles. 

RECORDS Alligator Light, Florida, 11 m; Carrie Bow Cay, Belize, 11-15.2 

B 

G

Paradella 223 

m, from green alga Halimeda sp. on forereef, and from sponge Aphysina 

jistularis. 

Paracerceis nuttingi (Boone, 1921) 

Figure 97H 

RECORDS Barbados; Puerto Rico, 1.5 m, from Cymodocea seagrass, and coral 

rubble and sponges. 

REMARKS The types of this species from Barbados consist only of females 

(total length 4.1 mm). Menzies and Glynn (1968) record an immature male 

i 

from Puerto Rico with an incipient pleotelsonic notch. This specimen, how 

ever, still has the subequal lamellar uropodal rami. The mature male, with 

the characteristically reduced uropodal endopod and cylindrical exopod, is 

unknown. The possibility exists that this is not a true Paracerceis. 

Paradella Harrison and Holdich, 1982 

DIAGNOSIS Marked 

ual dimorphism. Accessory unguis of pereopods simple, not bifid. Pleopod 3 

Key to species of Paradella 

1. Pereonite 7 with projecting bilobed flange; pleon and pleotelson finely 

but distinctly granulate 2 

Pleon and pleotelson smooth 3 

2. 8 with pleotelsonic foramen distinctly heart shaped, with median 

point; four submedian tubercles of pleotelson in 8 and 9 somewhat 

elongate; 9 pleotelson posteriorly narrowed, slit visible dorsally 

8 with pleotelsonic foramen wider than long, but lacking median 

point; four submedian tubercles of pleotelson in 8 small, rounded, 

obscure in 9; 9 pleotelson posteriorly truncate, slit not visible 

dianae 

dorsally plicatura 

3. Tubercles on pleotelson in 8 and 9 small to obscure; 8 with 

pleotelsonic foramen subcircular quadripunctata 

Tubercles on pleotelson broadly rounded mounds in 8 and 9; 8 with 

pleotelsonic foramen wider than long tumidicauda

224 FLABELL1FERA • SPHAEROMAT1DAE 

exopod with articulation. Uropodal rami lamellar. 9: Mouthparts not meta 

morphosed. One pair of oostegites arising from pereonite 4, short, not reach 

ing midline. Brood held in pouch formed by two opposing pockets covering 

entire ventrum, opening by transverse slit between 4th pereopods.

Paradella quadripunctata 225 

Figure 98. Paradella dianae: A, 8; B, pleopod 2 8; C, pleon 9. Paradella plicatura: 

Dy 8; E, pleon 9. Paradella quadripunctata: F, 8; G, pleon 9. Paradella tumidicauda; 

H, pleon 9 (from Glynn, 1970); /,


Paradella tumidicauda (Glynn, 1970) 

Figure 98H,I 

DIAGNOSIS 8 6.7 mm, 9 6.5 mm. 6: Last pleonite with two submedian 

swellings. Pleotelson with four submedian broadly rounded swellinglike tu 

bercles and two pairs of lateral tubercles. Foramen wider than long, posterior 

contiguous borders of foramen each bearing rounded swelling. 9: Pleotelson 

with four submedian swellinglike tubercles, sometimes with two obscure lat 

eral tubercles; posterior slit not visible dorsally, area surrounding slit 

swollen, horseshoe shaped. 

RECORDS Margarita Island, Venezuela, from among intertidal barnacles. 

Subfamily Sphaeromatinae H. Milne Edwards, 1840 

DIAGNOSIS Cephalon not fused with pereonite 1. Pereopods 1 and 2 am 

bulatory. Pleopods 4 and 5, endopods having branchial pleats, exopods un- 

pleated, membranous, of two articles. Uropods biramous. 

Key to genera of Sphaeromatinae 

1. Uropodal exopod with outer margin serrate Sphaeroma 

Uropodal exopod with outer margin entire or faintly crenulate 2 

2.

Cymodoce ruetzleri 227 

Pleopod 5, exopod of two articles, distal article with apex and internal mar 

gin covered with fine teeth, anterior surface with long distally toothed boss; 

proximal article with two small toothed bosses at internodistal angle. 6: 

Maxillipedal palp articles 2—4 bearing setigerous lobes. Penial rami elongate, 

separate. Pleon usually more tuberculate than in 9. Uropodal exopod lamel 

lar, shorter than endopod. 9: Mouthparts metamorphosed. Broodpouch 

formed by four pairs of oostegites arising from pereonites 1-4, overlapping in 

midline. Brood housed in five pairs of internal pouches. 

"Cymodoce" barrerae (Boone, 1918) 

Figure 99A,B 

DIAGNOSIS 9 7.5 mm. 9: Body dorsally strongly vaulted, unornamented. 

Frontal lamina distally broadly rounded, lateral shoulders rounded. 

Mouthparts not metamorphosed. Pleotelson anteriorly strongly inflated with 

barest indication of two submedian swellings; posterior margin trilobed, with 

median lobe strong, narrowly rounded, outer lobes much smaller and ventral 

to median lobe. Uropodal endopod distally obliquely truncate; exopod dis 

tally acute. 

RECORDS Cabanas, Cuba. 

REMARKS This species is known only from the nonovigerous female 

holotype. Loyola e Silva (1960) placed the species in Cymodoce, based on a 

female specimen from Brazil. As the mouthparts are not metamorphosed, 

this does not agree with the present concept of Cymodoce, but with neither 

ovigerous females nor males available, the correct generic placement cannot 

be determined. 

Cymodoce ruetzleri Kensley, 1984 

Figure 99C-G 

DIAGNOSIS 6 5.0 mm, 9 4.2 mm. 6: Integument with numerous small 

tubercles, becoming densely setose posteriorly. Pleonite 4, posterior margin 

broadly bilobed. Pleotelson bearing pair of strong conical tubercles with 

acute tips, each tubercle flanked by low rounded tubercle; apex trilobed, 

outer lobes triangular, acute, sharp spine at base of incision, median lobe 

apically blunt. Uropodal exopod apically acute, oval in cross section, endo 

pod and sympod fused, somewhat flattened, apex triangular with strong 

tooth. 9: Pleotelson with two conical apically acute tubercles, apex barely 

notched, with short rounded lobe slightly offset from posterior margin. Both


A 

Figure 99. "Qymodoce" barrerae: A, $; B, frontal lamina. Cymodoce ruetzleri: C, pleopod 4; G, pleopod 5. 

uropodal rami flattened; exopod with tiny apical tooth, endopod distally 

truncate-rounded, with small mediodistal tooth. 

RECORDS Carrie Bow Cay, Belize, 0.5-13 m; in algal clumps, reef crest 

rubble, and seagrass flats. 

E

Exosphaeroma Stebbing, 1900 

Exosphaeroma alba 229 

DIAGNOSIS Maxillipedal palp articles 2-4 produced medially into lobes. 

Pereonites 6 and 7 dorsally unarmed. Pleopod 3, exopod biarticulate. 6: 

Penes short, separate. Copulatory stylet of pleopod 2 elongate, slender. 

Pleotclson lacking strong apical notch. 9: Mouthparts not metamorphosed. 

Broodpouch of three pairs of oostegites on pereonites 2-4; oostegites short, 

not reaching midline. Brood held in four pairs of internal pouches. 

Key to species of Exosphaeroma 

1. Pleotelson with posterior margin entire, evenly convex 2 

Pleotelson with posterior margin faintly notched or trilobed 3 

2. Frontal lamina with length less than 1.5 times greatest width diminuta 

Frontal lamina with length almost two times greatest width 

productatelson 

3. Pleotelson with posterior margin faintly trilobed, and with three low 

rounded tubercles anteriorly yucatanum 

Pleotelson with posterior margin faintly notched 4 

4. Pleotelson posteriorly broadly notched; two rounded submedian 

tubercles on inflated midregion antillense 

Pleotelson with faint narrow notch posteriorly; lacking dorsal tubercles 

Exosphaeroma alba Menzies and Glynn, 1968 

Figure 100A-C 

DIAGNOSIS 8 2.0 mm, 9 2.3 mm. Frontal lamina anteriorly broadly 

rounded, basally slightly wider than midlength. Pleotelson similar in 8 and 

9; anterodorsally inflated and unornamented, posteriorly tapering to slight 

median notch, seen in dorsal view. Uropodal rami distally shallowly serrate, 

exopod 2.5 times longer than wide. 

RECORDS Puerto Rico, intertidal to 0.5 m; in algae on rocks, and under 

Chiton tuberculatus and C. marmoratus. 

alba


Figure 100. Exosphaeroma alba: A; B, frontal lamina; Cy uropod. Exosphaeroma 

antillense: D; E, frontal lamina; F, uropod. Exosphaeroma diminuta: G; H, frontal 

lamina; /, uropod. Exosphaeroma productatelson: J; K> frontal lamina; L, uropod. 

Exosphaeroma yucatanum: My pleon (from Richardson, 1905). 

Exosphaeroma antillense Richardson, 1912d 

Figure 100D,F 

DIAGNOSIS 9 5.0 mm. Frontal lamina anteriorly tapering to subacute apex. 

Pleotelson with two broadly subconical submedian tubercles on inflated an-

Exosphaeroma yucatanum 231 

terior area; posterior margin subtruncate to very faintly emarginate. Uropo 

dal exopod distally crenulate, length slightly more than twice greatest width; 

endopod with faint distal notch. 

RECORDS Montego Bay, Jamaica. 

REMARKS The single ovigerous female holotype is the only known specimen 

of this species. The overlapping oostegites suggest that this may not be an 

Exosphaeroma. 

Exosphaeroma diminuta Menzies and Frankenberg, 1966 

Figure 100G-I 

DIAGNOSIS 8 2.2 mm. Frontal lamina widest at midlength, anteriorly 

truncate-rounded. Pleotelson with posterior margin broadly rounded. 

Uropodal rami not quite reaching pleotelsonic apex; exopod margin distally 

crenulate. 

RECORDS Chesapeake Bay to Florida; Venezuela; sand dwelling, intertidal 

and shallow subtidal. 

Exosphaeroma productatelson Menzies and Glynn, 1968 

Figure 100J-L 

DIAGNOSIS 6 2.5 mm, 9 1.5 mm. Sexes essentially similar. Frontal lamina 

widest at midlength, where slight shoulder apparent, anteriorly broadly 

rounded, 1.6 times longer than wide. Pleotelson unornamented, anteriorly 

inflated, posterior margin entire, evenly convex. Uropodal exopod distally 

shallowly serrate, almost four times longer than wide; endopod wider than 

exopod. Broad lateral patches of pigment on pleotelson in both sexes. 

RECORDS Puerto Rico, intertidal to 0,5 m, in algae on rocks; Texas, Gulf of 

Mexico. 

Exosphaeroma yucatanum (Richardson, 1901) 

Figure 100M 

DIAGNOSIS Frontal lamina anteriorly tapering from widest point to sub 

acute apex, proximally narrower than at midlength. Pleotelson posteriorly 

obscurely trilobed, median lobe narrowly rounded, longest; three low 

rounded tubercles on pleotelson in anterior region.


RECORDS Cape Catoche, Yucatan, Mexico, 48 m. 

REMARKS This species was described from a single specimen which has 

since been lost. The true generic placement of this species is thus undeter 

mined and full description awaits the finding of more material. 

Harrieta Kensley, 1987c 

DIAGNOSIS 9 with mouthparts metamorphosed. Broodpouch of three pairs 

of oostegites on pereonites 2—4, overlapping in midline; brood held in five 

pairs of internal pouches. Uropodal rami subequal, lamellar in 9, exopod 

twice length of endopod and oval in cross section in 8. Pleopod 2 in 6 with 

copulatory stylet articulating basally on endopod, curved, barely reaching 

apex of endopod. Penes basally fused, rami slender, elongate, tapering. 

Harrieta faxoni (Richardson, 1905) 

Figure 101 A,B 

DIAGNOSIS 6 6.0 mm, 9 6.5 mm. 6: Frontal lamina with broad slightly 

convex anterior margin. Two low rounded submedian tubercles on cephalon 

near posterior margin. Two rounded submedian tubercles on last pleonite. 

Pleotelson anteriorly inflated with two submedian tubercles; posterior mar 

gin trilobed. 9: Essentially similar to 69 but posterior margin of pleotelson 

less markedly trilobed, with median lobe longer, and uropodal rami subequal 

in length. 

RECORDS Florida to Texas, Gulf of Mexico, intertidal and subtidal in 

Thalassia, Halodule, and Syringodium seagrass beds, in salinities of 7%o to 36%o. 

Sphaeroma Bosc, 1802 

DIAGNOSIS Maxillipedal palp with three distal articles poorly developed, 

lacking lobes; fringe of robust plumose setae with swollen bases on internal 

margin of endite; distal margin of endite with simple setae. Pereopods 1—3 

with plumose setae on ischium and merus. Posterior margin of pleotelson 

entire, similar in 6 and 9. Pleopod 3, exopod uniarticulate. Uropodal ex 

opod with outer margin serrate. Able to conglobate. 6: Penes short, 

rounded, separate. Pleopod 2, copulatory stylet articulating basally on endo 

pod, slender, reaching well beyond rami. 9: Mouthparts not meta-

Figure 101. Harrieta faxoni: A, 6; B, pie 

Sphaeroma terebrans; E, Sphaeroma walkeri. 

B 

E


morphosed. Three pairs of overlapping oostegites arising from pereonites 2- 

4 (but S. terebrans has anterior pair rudimentary). 

REMARKS The genus Sphaeroma is one of the few sphaeromatids in which the 

number of oostegites varies, from the diagnostic three pairs, through two 

normal pairs (as in S. terebrans), to having the oostegites completely absent 

(as in S. annandalei). 

Jacobs (1987) has provided a useful reevaluation of the European, Medi 

terranean, and northwest African species of Sphaeroma and related genera. 

Key to species of Sphaeroma 

1. Pleotelson posteriorly bluntly triangular, with 4 strong anterior 

tubercles 

terebrans 

Pleotelson posteriorly broadly rounded 2 

2. Pleotelson dorsally smooth or with few low tubercles quadridentata 

Pleotelson dorsally with numerous strong tubercules walkeri 

Sphaeroma quadridentata Say, 1818 

Figure 101C 

DIAGNOSIS 6 11.0 mm, 9 8.0 mm. Pleotelson anteriorly inflated, some 

times with few low rounded tubercles, posteriorly flattened to concave; pos 

terior margin entire, broadly rounded. 

RECORDS New England to Florida; Gulf of Mexico, intertidal to 1 m, often 

in pilings and partially submerged dead tree trunks, and commonly associ 

ated with barnacles. 

Sphaeroma terebrans Bate, 1866 

Figure 10 ID 

DIAGNOSIS 6 10.0 mm, 9 11.5 mm. Pereonite 7 with pair of submedian 

and pair of lateral tubercles. Dorsal pleon densely tuberculate. Posterior 

pleonite with pair of submedian acute tubercles. Pleotelson anteriorly with 

submedian pair and lateral pair of tubercles, posteriorly rounded-triangular.

FLABELLIFERA • TRIDENTELLIDAE 235 

RECORDS Virginia to Florida; Belize; Cuba; Venezuela to Brazil; Gulf of 

Mexico. 

Nigeria, east coast of southern Africa, India, Sri Lanka, Thailand, Indo 

nesia, Philippines, Australia. 

REMARKS There is no agreement on whether this species is synonymous 

with S. destructor Richardson, 1897. This latter (if distinct) bores into wood 

pilings in estuarine waters, while S. terebrans is found in the prop roots of the 

red mangrove tree, Rhizophora mangle. In this habitat, the isopods are inter 

preted either as being destructive agents (e.g., Rehm and Humm, 1973) or as 

promoting increased root growth (Simberloff et al., 1978). It is unlikely that 

the bored wood itself is a source of food for the isopods; rather, as with the 

genus Limnoria, the food is probably detritus or fungi and bacteria growing on 

the wood fragments in the burrows or on the setae of the appendages. 

Sphaeroma walkeri Stebbing, 1905 

Figure 101E 

DIAGNOSIS 6 9.5 mm, 9 10.0 mm. Pereonites 3—7 with transverse row of 

large rounded tubercles. Last pleonite with row of prominent tubercles and 

smaller scattered tubercles laterally. Pleotelson anteriorly inflated, posteri 

orly concave and cuplike, with four irregular longitudinal rows of large tu 

bercles plus many small scattered tubercles. Posterior margin rounded, en 

tire to irregularly crenulate. Uropodal endopod with several rounded 

tubercles on dorsal surface; exopod with row of smaller tubercles on ventral 

surface. 

RECORDS Probably pan-tropical. Florida to Puerto Rico, intertidal. 

Family Tridentellidae Bruce, 1984 

DIAGNOSIS Eyes well developed. Pereonites 2—7 with distinct coxae. Pleon 

consisting of five free pleonites plus pleotelson. Mandible with acute incisor; 

lacinia mobilis absent; molar present; palp of three articles. Maxilla 1, outer 

ramus styliform with three to five strong terminal spines, and several short 

recurved subapical spines. Maxilla 2 uniramous, biarticulate, bearing small 

sometimes tridentate spines or scales distally. Maxillipedal palp of five arti 

cles; endite slender, lamellar, usually lacking coupling hooks.

236 FLABELLIFERA • TRIDENTELLIDAE 

Tridentella Richardson, 1905 

DIAGNOSIS Body dorsally often bearing spines, tubercles, or carinae, more 

developed in 8 than in 9. Frontal lamina narrow, pentagonal. Antcnnular 

peduncle of three articles; antennal peduncle of five articles. Mandibular 

molar weakly sclerotized. Pereopods 1-3 weakly prehensile; pereopods 4-7 

ambulatory. Copulatory stylet of pleopod 2 rodlike, arising proximally on 

mesial margin of endopod. Pleopod 5 endopod lacking marginal setae. 

REMARKS Delaney and Brusca (1985) provide useful taxonomic and dis 

tributional comments on the family Tridentellidae. 

Tridentella virginiana (Richardson, 1900b) 

Figure 102 

DIAGNOSIS 8 9.5 mm, ovigerous 9 9.5-11.0 mm. Cephalon and pereon 

dorsally smooth, pleon minutely granular. Uropodal rami with distal mar 

gins faintly dentate, apically narrowly rounded, endopod wider and slightly 

longer than exopod. Pleotelson basally wider than middorsal length; pos 

terior margin broadly rounded to subtruncate. 

RECORDS Nova Scotia to Florida; off Georgia, 550 m; Gulf Stream off Key 

West, 220 m. 

Suborder Gnathiidea Leach, 1814 

DIAGNOSIS Eyes usually well developed, rarely on short lateral processes, 

occasionally absent. Mandibles in 8 greatly enlarged, projecting anteriorly 

from cephalon, not used in feeding. Mandibles lacking in 9. Mouthparts of 

praniza larva styliform, with acute mandibles projecting anteriorly (see Fig 

ure 103D). Pereopod 1 modified, forming second pair of broad opercular 

maxillipeds covering mouthparts, referred to as pylopods. Pereopods 2—6 

ambulatory. Pereonite 7 reduced, lacking pereopod. Pleonites separate, nar 

rower than pereon. Uropods lateral, rami lamellar, forming tailfan with 

telson. Praniza larva with pereonites 4-6 enlarged, sometimes inflated. 9 

with pereonites 4-6 greatly inflated, forming broodpouch for internally 

brooded eggs (see Figure 103E). 

REMARKS The gnathiideans are entirely marine, most described species 

being from shallow waters. The males and females are frequently found in 

association with sponges and do not feed. The praniza larva is an efficient 

swimmer and has been recorded from shallow-water plankton, but is more

GNATHIIDEA 237 

Figure 102. Tridentella virginiana: A, 9; B, pereopod 1; C, maxilla 1; D, mandible; 

Ey maxilliped; F, maxilla 2. 

frequently encountered as a fish parasite, the favored site for sucking the 

host's blood being in the nares. Upton (1987a, 1987b) has shed light on the 

unusual life history of at least one gnathiid genus, Paragnathia. 

The taxonomy of the Gnathiidae is based almost entirely on males, the 

praniza and females of most species being remarkably similar.

238 GNATHIIDEA • GNATHIIDAE 

Family Gnathiidae Harger, 1879 

DIAGNOSIS As for the suborder Gnathiidea. 

Gnathia Leach, 1814 

DIAGNOSIS In addition to features mentioned in diagnosis of suborder: Eyes 

present in most species. Pylopod with two small articles distal to broad oper 

cular article 2, terminal article minute. 

Key to species of Gnathia (6 only) 

1. Anterior margin of cephalon with medial process or slightly convex . . 2 

Anterior margin of cephalon concave or lacking medial process 9 

2. Anterior margin of cephalon broadly triangular, projecting, with small 

lateral teeth triospathiona 

Anterior margin of cephalon not triangular and projecting 3 

3. Cephalon and two free anterior pereonites dorsally granular 4 

Cephalon and two free anterior pereonites smooth 5 

4. Lobe of outer margin of mandible notched; pereonite 5 twice wider 

than middorsal length velosa 

Lobe of outer margin of mandible rounded; pereonite 5 1.5 times wider 

than middorsal length 6 

5. Anterior margin of cephalon with distinct medial process virginalis 

Anterior margin of cephalon barely convex, lacking medial process 

rat hi 

6. Inner proximal lobe of mandible distinct 7 

Inner proximal lobe of mandible indistinct samariensis 

7. Inner proximal lobe of mandible entire 8 

Inner proximal lobe of mandible with rounded toothlike marginal 

structures Johanna 

8. Pereonites 3-5 poorly defined puertoricensis 

Pereonites 3-5 clearly defined magdalenensis 

9. Anterior margin of cephalon concave, lacking projections gonzalezi 

Anterior margin of cephalon with four projections beethoveni

Gnathia beethoveni Paul and Menzies, 1971 

Figure 103 A 

Gnathia magdalenensis 239 

DIAGNOSIS 6 3.0 mm. Anterior margin of cephalon with two low tubercles 

flanking shallow medial notch plus slightly larger pair of lateral tubercles. 

Cephalon lacking dorsal tubercles. Pereonite 5 1.5 times wider than middor- 

sal length. Uropodal endopods reaching beyond telsonic apex. 

RECORDS Off Venezuela, 95 m. Colombia. 

Gnathia gonzalezi Miiller, 1988 

Figure 103B 

DIAGNOSIS 6 2.0 mm. Body smooth. Anterior margin of cephalon concave. 

Pereonites 3—5 distinct; pereonite 5 2.5 times wider than middorsal length. 

Cutting margin of mandible with four or five low rounded teeth. 

RECORDS Colombia, 30 m. 

Gnathia Johanna Monod, 1926 

Figure 103C 

DIAGNOSIS 8 2.1 mm. Anterior margin of cephalon medially convex be 

tween pair of submedian tubercles. Pereonites 4 and 5 poorly separated. 

Proximomedial lobe of mandible having four or five rounded crenulations, 

with tiny seta between adjacent crenulations. 

RECORDS U.S. Virgin Islands, 29-46 m; Colombia. 

Gnathia magdalenensis Miiller, 1988 

Figure 103D 

DIAGNOSIS 6 3.0 mm. Anterior margin of cephalon with three tubercles, 

median tubercle slightly shorter than submedian pair. Cephalon with few 

scattered low granulations dorsally. Pereonite 5 about 1.5 times wider than 

middorsal length. Proximomedial lobe of mandible entire. 

RECORDS Carrie Bow Cay, Belize, intertidal; Colombia, 18 m.

Figure 103. Gnathia beethhoveni: A, 6. Gnathia gonzalezi: B, 6 (after Miiller, 1988). 

Gnathia Johanna: C,

Gnathia puertoricensis Menzies and Glynn, 1968 

Figure 103E-G 

Gnathia triospathiona 241 

DIAGNOSIS 6 3.0 mm, ovigerous $ 1.8 mm. Anterior margin of cephalon 

having three tubercles between mandibular bases, medial tubercle narrower 

than submedian pair. Dorsal integument finely granular, with coarser gran 

ules mediodorsal to eye. Pereonites 4 and 5 indistinctly separated. Mandible 

lacking proximomedial lobe. 

RECORDS Carrie Bow Cay, Belize, intertidal to 2 m; Puerto Rico, intertidal; 

Cuba. 

Gnathia rathi Kensley, 1984 

Figure 104 A 

DIAGNOSIS 6 1.9 mm, ovigerous 9 2.2 mm. Frontal margin faintly convex 

to straight between single low lateral tubercle mesial to mandibular bases. 

Dorsal integument of cephalon and anterior two free pereonites coarsely 

granular. Lateral margins of telson faintly denticulate. Pereonites 4 and 5 

poorly separated. 

RECORDS Carrie Bow Cay, Belize, 1-36 m. 

Gnathia samariensis Miiller, 1988 

Figure 104B 


median tubercle slightly shorter than submedian pair; dorsal integument 

smooth. Pereonites 4 and 5 well differentiated; pereonite 5 about 2.2 times 

wider than middorsal length. Mandible lacking proximomedial lobe. 

RECORDS Colombia. 

Gnathia triospathiona Boone, 1918 

Figure 104C 

DIAGNOSIS 6 8.8 mm. Anterior margin of cephalon with broad-based tri 

angular projection bearing three low teeth; deep V-shaped depression pos 

terior to anterior margin, with low flanking granulations. 

RECORDS Off Key West, in Gulf Stream, 218 m.

Figure 104. Gnathia rathi: A, 6; Gnathia samariensis: B, 6 (after Miiller, 1988); 

Gnathia triospathiona; C, 6; Gnathia virginalis: D, 6; Gnathia velosa: E, 6 (after 

Miiller, 1988). 

E

Gnathia velosa Muller, 1988 

Figure 104E 

MIGROGERBERIDEA • MIGROGERBERIDAE 243 


median tubercles slightly shorter and narrower than submedian pair. Dorsal 

integument of cephalon and anterior three pereonites granular. Pereonite 5 

about 2.5 times wider than middorsal length. Lateral lobe of mandible 

notched. 

RECORDS Colombia. 

Gnathia virginalis Monod, 1926 

Figure 104D 


median tubercles slightly longer than submedian pair. Dorsal integument of 

cephalon and anterior three pereonites granular. Pereonite 5 about 1.7 times 

wider than middorsal length. Lateral lobe of mandible rounded. 

RECORDS U.S. Virgin Islands, 29 m; Colombia. 

Suborder Microcerberidea Lang, 1961 

DIAGNOSIS Cephalon free. Mandibles with reduced palp, or lacking palp. 

Maxillipedal palp of five articles. Pereon of seven free segments. Pereopod 1 

subchelate; pereopods 2—7 ambulatory. Pleon of two free pleonites plus 

pleotelson. Pleopod 1 in 6 usually absent. Pleopod 2 modified for copulation. 

Pleopod 3 uniramous, opercular. Pleopod 4 biramous. Pleopod 5 reduced. 

Uropods usually uniramous or biramous. 

Family Microcerberidae Karaman, 1933b 

DIAGNOSIS Eyes absent. Body elongate, slender. Antennular peduncle of 

three articles; antennal peduncle of six to eight articles. Mandibular palp of 

single article; molar reduced to single stout fringed spine. Maxilla 2 reduced 

to single ramus bearing two distal fringed lobes. Pereopods 2—7 ambulatory, 

dactyli biunguiculate. 

REMARKS The species of the Microcerberidae are all very small (less than 2 

mm total length) and are most often found in interstitial habitats. They have 

been recorded from marine, brackish, and freshwater environments.

244 MICROCERBERIDEA • MICROCERBERIDAE 

The microcerbcrideans were often classified with the Anthuridea, mainly 

because of the similarity in body shape. Wagele (1983) however, has con 

vincingly demonstrated the asellotan affinities of the group. 

Key to genera of Microcerberidae 

1. MaxilHpedal palp articles 2 and 3 enlarged; basis of pereopods lacking 

spinous process Yvesia 

MaxilHpedal palp articles 2 and 3 not markedly enlarged; basis of 

pereopods with spinous process Microcerberus 

Microcerberus Karaman, 1933b 

DIAGNOSIS MaxilHpedal palp articles 2 and 3 not enlarged. Articles 2 and 3 

of antennal peduncle with spinous process. Basis of pereopods with spinous 

process. Propodus of pereopod 2 with two denticulate proximal spines. 

Microcerberus syrticus Kensley, 1984 

Figure 105A-E 

DIAGNOSIS (5 1.1 mm, 9 1.1 mm. Tergal lobes of pereonites 2—4 rounded. 

Apical lobe of 6 pleopod 2 acute. 

RECORDS Carrie Bow Cay, Belize, interstitial in intertidal sand bar. 

REMARKS In addition to M. syrticus, six species of Microcerberus have been 

recorded from the Caribbean area: M. littoralis Chappuis and Delamare De- 

boutteville, 1956, from the Bahamas; M. minutus Coineau and Botosaneanu, 

1973, from Cuba; M. mirabilis Chappuis and Delamare Deboutteville, 1956, 

from the Bahamas; M. nunezi Coineau and Botosaneanu, 1973, from Cuba; 

M. renaudi Chappuis and Delamare Deboutteville, 1956, from the Bahamas; 

M. simplex Coineau and Botosaneanu, 1973, from Cuba. The reader is re 

ferred to the original descriptions for separation of the species.

\ 

Figure 105. Microcerberus syrticus: A, 8; B, pereopod 1; C, maxilliped; D, pereopod 

2; Ey pleopod 2 8. Yvesia striata (from Coineau and Botosaneanu, 1973): F9 

maxilliped; G, pereopod 1; H, pereopod 2. 

G

246 MICROCERBER1DEA • ONISCIDEA 

Yvesia Coineau and Botosaneanu, 1973 

Yvesia striata Coineau and Botosaneanu, 1973 

Figure 105F-H 

DIAGNOSIS 9 1.6 mm. Antennal peduncular articles 2 and 3 smooth, lack 

ing spinous processes. Maxillipedal palp articles 2 and 3 enlarged. Bases of 

pereopods unarmed, lacking spinous processes. Propodus of percopod 1 with 

single smooth proximal spine. Body having longitudinal ventrolateral striae. 

RECORDS Oriente, Cuba, interstitial on beach. 

Suborder Oniscidea Latreille, 1803 

DIAGNOSIS Compound eyes usually present. Antennules usually very short. 

Antennae with 4- or 5-articulate peduncle; flagellum varying from few arti 

cles to multiarticulate. Mandibular palp present. Distal articles of maxillipe 

dal palp often reduced. Coxae of pereonites 1—7 usually distinct, expanded. 

Pleopods respiratory, often with pseudotrachea; 6 with pleopod 2, and 

sometimes pleopod 1 as well, modified for copulation. Uropods terminal or 

subterminal with terete rami, or ventral and opercular, with reduced rami. 

REMARKS The Oniscidea includes all the isopods that have successfully in 

vaded the terrestrial environment. While still in some degree reliant on exter 

nal moisture, their morphological and behavioral adaptations have allowed 

them to live in almost all terrestrial habitats, from hot, dry deserts, through 

tropical rainforests and grasslands, to cold-temperate niches. Several forms 

have successfully inveigled themselves into termite or ant colonies, where 

with varying degrees of morphological adaptations they take advantage of 

the security of these habitats. A small number of species have evolved to live 

in more constantly wet habitats. Several species may be found in the marine 

intertidal, either living in and under piles of decomposing litter along the 

high-tide line, digging into beach sand, or sheltering in the damp cracks and 

crevices of rocky shores. A few may also be found in mangrove swamps. 

A breakdown of families, genera, and species is not provided for this sub 

order, but those few species that are commonly encountered in intertidal 

habitats are dealt with individually. Schultz (1974, 1984) records several 

oniscidean isopods from the Caribbean area.

Key to genera and species of littoral Oniscidea 

ONISCIDEA 247 

1. At least one uropodal ramus reaching well beyond outline of body ... 5 

Uropodal rami very short, not reaching beyond outline of body 2 

2. Uropods ventral, not visible in dorsal view 3 (Tylos) 

Uropods visible in dorsal view Armadilloniscus ninae 

3. Ventral extensions of pleonite 5 meeting in midline Tylos niveus 

Ventral extensions of pleonite 5 not meeting in midline 4 

4. Ventral extensions of pleonite 5 very short, obsolete Tylos wegeneri 

Ventral extensions of pleonite 5 well separated Tylos marcuzzii 

Ventral extensions of pleonite 5 just falling short of meeting in midline 

Tylos latreillei 

5. Uropodal rami both elongate, subequal 6 (Ligia) 

Uropodal rami very unequal in length 8 

6. Propodus of 6 pereopod 1 with distal rounded lobe Ligia exotica 

Propodus of 6 pereopod 1 lacking rounded lobe 7 

7. Apex of 6 pleopod 2 club shaped Ligia olfersii 

Apex of 6 pleopod 2 bifid Ligia baudiniana 

8. Antennal flagellum of two articles Rhyscotus texensis 

Antennal flagellum of three articles 9 (Vandeloscia) 

9. Endopod of 6 pleopod 1 with large scalelike subapical process 

Endopod of 6 pleopod 1 with small scalelike subapical process 

Armadilloniscus Ul'yanin, 1875 

Armadilloniscus ninae Schultz, 1984 

Figure 106A 

Vandeloscia riedli 

Vandeloscia culebrae 

- - - - - , 

DIAGNOSIS 8 3.2 mm, 9 4.1 mm. Uropodal sympod expanded to form part 

of body outline; rami set mesial to expanded base, with exopod half length of 

endopod. 

RECORDS Ambergris Cay, Belize; under damp objects along beach drift 

line.

Figure 106. A, Armadilloniscus ninae. Ligia baudiniana: B; C} 6 pleopod 2 endopod. 

Ligia exotica: D, dactylus and propodus of pereopod 1; E, 6 pleopod 2 endopod. 

Ligia olfersii: F, 6 pleopod 2 endopod. G, Rhyscotus texensis. Tylos latreillei: H, 

ventral pleon. Tylos marcuzzi: I, ventral pleon (from Schultz, 1984). Tylos niveus: J, 

lateral view; K} ventral pleon. Tylos wegeneri: L} ventral pleon. Vandeloscia culebrae: 

M, apex of pleopod 1 endopod. N, Vandeloscia riedli.

Ligia Fabricius, 1798 

Ligia baudiniana H. Milne Edwards, 1840 

Figure 106B,C 

Rhyscotus texensis 249 

DIAGNOSIS 6 and 9 up to 22 mm. Antennal flagellum elongate, multiar- 

ticulate. Apex of 6 pleopod 2 bifid, with lateral lobe longer and more slender 

than mesial lobe. Uropods inserted terminally on pleotelson; sympods 

elongate-cylindrical; rami slender, elongate, subequal. 

RECORDS Bermuda; Bahamas; U.S. Virgin Islands; Antigua; Carrie Bow 

Cay, Belize; Bonaire; Aruba; Trinidad; Tobago; Gulf of Mexico. 

REMARKS As is typical in the genus Ligia, this species may be seen on rocks 

i 

and sea walls, as well as piles of drift debris at low tide. When disturbed, they 

run rapidly, to shelter in damp crevices and hollows. 

Ligia exotica Roux, 1828 

Figure 106D,E 

DIAGNOSIS 6 28.5 mm, ovigerous 9 32.0 mm. Propodus of 8 pereopod 1 

with rounded lobe on inner distal surface. Apex of 6 pleopod 2 club shaped, 

convoluted. 

RECORDS New Jersey to Uruguay; Indo-Pacific. 

Ligia olfersii Brandt, 1833 

Figure 106F 

DIAGNOSIS 6 20.0 mm, ovigerous 9 24.0 mm. Apex of 6 pleopod 2 simple, 

club shaped. 

RECORDS South Florida to Rio de Janeiro, Brazil; Texas, Gulf of Mexico. 

Rhyscotus Budde-Lund, 1885 

Rhyscotus texensis (Richardson, 1905) 

Figure 106G 

DIAGNOSIS 6 and 9 6.0 mm. Antennal flagellum of two unequal articles. 

Uropodal endopod at least twice length of exopod, inserted distally on base, 

exopod inserted distally on base. Pleotelson broadly triangular. 

RECORDS Carrie Bow Cay, Belize; Texas, Gulf of Mexico.

250 ONISCIDEA 

Tylos Latreille, 1826 

Tylos latreillei Audouin, 1826 

Figure 106H 

DIAGNOSIS 6 12.8 mm, 9 13.0 mm. Ventral extensions of pleonite 5 not 

meeting in midline. 

RECORDS Bermuda; Cuba; Puerto Rico; Honduras. 

Mediterranean. 

Tylos marcuzzii Soika, 1954 

Figure 1061 

DIAGNOSIS 6 6.6 mm. Antennal flagellum of four articles. Ventral exten 

sions of pleonite 5 well separated. 

RECORDS Florida Keys; Bahamas; Leeward Islands; Ambergris Cay, Be 

lize; under debris on sand beach drift line. 

Tylos niveus Budde-Lund, 1885 

Figure 106J,K 

DIAGNOSIS 8 11.0 mm., 9 12.0 mm. Antennal flagellum of four articles. 

Ventral extensions of pleonite 5 expanded, medially contiguous. 

RECORDS Bahamas; Florida Keys; Cuba; Dominica; Lesser Antilles; Bo 

naire; Curasao, under piles of decaying mangrove leaves at beach drift line; 

Carrie Bow Cay, Ambergris Cay, Belize, under deep piles of dead plant ma 

terial on beach drift line; Tobago; Panama. 

Rio de Janeiro, Brazil. 

Tylos wegeneri Vandel, 1952 

Figure 106L 

DIAGNOSIS 6 10.5 mm, 9 15 mm. Antennal flagellum of three articles. 

Ventral extensions of pleonites short or nearly absent. Pleonite 5 lacking free 

lateral margins. 

RECORDS Tobago; Venezuela, under decaying beach debris on drift line; 

Trinidad.

Vandeloscia Roman, 1977 

Vandeloscia culebrae (Moore, 1901) 

Figure 106M 

VALVIFERA 251 

DIAGNOSIS 8 5.0 mm, 9 6.1 mm. Tiny lateral tubercles present on per 

eonites. Endopod of pleopod 1 in 8 with small scalelike subapical process on 

laterally folded tip. 

RECORDS Florida Keys; U.S. Virgin Islands; Puerto Rico; under decaying 

plant material, especially Thalassia testudinea accumulated along beach drift 

line, 

Vandeloscia riedli (Strouhal, 1966) 

Figure 106N 

DIAGNOSIS 8 5.9 mm, 9 6.0 mm. Tiny obsolete tubercles present on all 

pereonites. Endopod of 8 pleopod 1 with large scalelike subapical process on 

laterally folded tip. 

RECORDS Yucatan Peninsula, Mexico; Ambergris Cay, Belize; Barbuda; 

Venezuela; Brazil. 

Gulf of Aqaba; Red Sea; northeastern coast of Africa; Madagascar; Bay of 

Bengal; St. Helena Is. 

Suborder Valvifera Sars, 1882 

DIAGNOSIS Pereopodal coxae, in addition to usual dorsal coxal plates, ex 

panded ventrally to form plates. Penes situated ventrally on articulation be 

tween pereon and pleon, or on pleonite 1. Pleonites and pleotelson variously 

fused. Uropods forming operculum covering over pleopods. 

Key to families of Valvifera 

1. Body often geniculate, flexed between pereonites 4 and 5; anterior 

pereopods setose for feeding, posterior pereopods ambulatory 

Arcturidae 

Body never geniculate; all pereopods ambulatory Idoteidae

252 VALV1FERA • ARGTUR1DAE 

REMARKS Of the six families in the suborder, only two have been recorded 

in the Caribbean area, the Idoteidae and the Arcturidae. 

Family Arcturidae Sars, 1897 

DIAGNOSIS Pereonite 1 either distinct, or completely or imcompletely fused 

with cephalon. Anterior four pairs of pereopods directed anteriorly, usually 

strongly setose; posterior three pairs of pereopods ambulatory, used for cling 

ing to substrate. Body often bent between pereonites 4 and 5. Uropods usu 

ally biramous, with minute endopod concealed by larger exopod. Pleonites 

variously fused with pleotelson. Sexual dimorphism often marked. 

REMARKS Menzies and Kruczynski (1983) described three species of 

arcturids from the west coast of Florida, in depths of 55—73 m: Arcturella 

spinata, Arcturella bispinata, and Edwinjoycea horologium. These species are not 

covered here. 

Key to genera of Arcturidae 

1. Pereonite 1 not fused with cephalon; at least one free pleonite 

Pereonite 1 fused with cephalon; pleonites fused with pleotelson 

Astacilla Cordiner, 1793 

Thermarcturus 

Astacilla 

DIAGNOSIS Antennae at least half length of body. Pereopod 1 with strong 

terminal claw on dactylus. Pereopods 2—4 lacking dactyli. Endopod of 

Key to species of Astacilla 

1. Body integument lacking ornamentation cymodocea 

Body integument with spines or tubercles 2 

2. Pereonite 4 in 6 and 9 with strong middorsal tubercle; pairs of spines 

lacking on pereonites regina 

Pereonite 4 lacking strong middorsal tubercle; pairs of spines on all 

pereonites lasallae

Astacilla regina 253 

pleopod 1 6 with median notch and three specialized setae; pleopod 2 copu- 

latory stylet apically trifid. Pereonite 4 considerably longer than preceeding 

or following pereonite. 

Astacilla cymodocea Menzies and Glynn, 1968 

Figure 107A,B 

DIAGNOSIS 6 6.4 mm, ovigerous 9 9.0 mm. Body cylindrical, ovigerous 9 

with pereonite 4 somewhat bulged, 6 with pereonite 4 elongate-cylindrical. 

Shallow groove marking fusion between cephalon and pereonite 1. Pleonites 

fused with pleotelson, with two incomplete shallow dorsal grooves marking 

lines of fusion anteriorly. Pleotelson lacking any shoulders, posteriorly tape 

red to narrowly rounded apex. 

RECORDS Florida Keys; Puerto Rico, 1.5 m, on Cymodocea sp. seagrass; Car 

rie Bow Cay, Belize, 1—2 m, on Syringodium filiforme seagrass. 

REMARKS In life, A. cymodocea is bright green, blending in with its preferred 

substrate of seagrasses. 

Astacilla lasallae Paul and Menzies, 1971 

Figure 107C 

DIAGNOSIS 9 3.5 mm. Cephalon with large rounded area bearing pair of 

spines; all pereonites and two anterior fused pleonites bearing pair of short 

submedian spines. Pleotelson with strong anterior shoulder, posteriorly tri 

angular, tapering sharply to narrowly rounded apex. 

RECORD Off Venezuela, 95 m. 

REMARKS This species is known only from the small female holotype, and 

until a mature male and ovigerous female are found, it cannot be confidently 

diagnosed. 

Astacilla regina Kensley, 1984 

Figure 107D-G 

DIAGNOSIS 6 6.5 mm, ovigerous 9 7.1 mm. Body strongly tuberculate, 

many tubercles acute. Cephalon with two submedian pairs of acute tuber 

cles; fused pereonite 1 and pereonites 2 and 3 each with single middorsal 

acute tubercle. Pereonite 4 with strong middorsal tubercle situated in ante-

254 VALVIFERA • ARGTURIDAE 

Figure 107. Astacilla cymodocea: A, 8; By 9. Astacilla lasallae: C, 6. Astacilla regina: 

D, 6\ E, 9; F, pereopod 4; G, pereopod 1. Thermarcturus venezuelensis: H, 9 (from 

Paul and Menzies, 1971). 

rior half. Pleotelson with strong lateral shoulder in anterior half, second 

shoulder in posterior half, then tapering to rounded apex. 

RECORDS Carrie Bow Cay, Belize, on forereef slope, 27—36 m; Barbados, 

100—400 m; St. Lucia, 2—3 m, associated with crinoids.

Thermarcturus Paul and Menzies, 1971 

VALVIFERA • IDOTEIDAE 255 

DIAGNOSIS Pereonite 1 not fused with cephalon. Pereonite 4 subequal in 

length to pereonite 3, not markedly elongate. Pereopods 2—4 having dactyli 

but lacking elongate setae. Body cylindrical, flexed between pereonites 4 and 

5. Pleon consisting of two free pleonites plus pleotelson. 

Thermarcturus venezuelensis Paul and Menzies, 1971 

Figure 107H 

DIAGNOSIS 9 4.5 mm. Cephalon, all pereonites, and anterior two pleonites 

each with submedian pair of dorsal tubercles, those on pereonites 2 and 3 

broad and expanded. Pleonite 2 with pair of bulbous lateral swellings, pos 

terior margin triangular. Pleotelson with lateral shoulder anteriorly, posteri 

orly triangular. 

RECORDS Off Venezuela, 95 m. 

REMARKS Only the holotype (which seems to be lost) is known of this spe 

cies. Considerable uncertainty exists regarding some of the features. 

Family Idoteidae Fabricius, 1798 

Subfamily Idoteinae Dana, 1852 

DIAGNOSIS Flagellum of antenna either multiarticulate; clavate, i.e., with 

large basal articles and with or without one to four reduced distal articles; or 

Key to genera of Idoteinae 

1. Antennal flagellum multiarticulate Idotea 

Antennal flagellum clavate 2 

2. Pereopod 4 reduced, considerably smaller than pereopods 3 or 5 .... 3 

Pereopod 4 not reduced, of similar size to pereopods 3 and 5 

Erichsonella 

3. Pleon consisting of three complete and one incomplete pleonites plus 

pleotelson Cleantioides 

Pleon consisting of two complete and two incomplete pleonites plus 

pleotelson Miratidotea

256 VALVIFERA • IDOTEIDAE 

vestigial. Maxillipedal palp consisting of five or fewer articles. Uropods uni- 

ramous or biramous, rami usually much smaller than sympod. Pleonites 

variously fused with pleotelson; number of fused pleonites often indicated by 

lateral sutures or furrows. 

REMARKS Brusca (1984) has reviewed the phylogeny, evolution, and bio- 

geography of the subfamily Idoteinae, the only one of the five subfamilies 

recorded from the Caribbean. 

Cleantioides Kensley and Kaufman, 1978 

DIAGNOSIS Antennal flagellum a single clavate article. Maxillipedal palp of 

four or five articles. Pereopod 4 somewhat reduced. Uropod uniramous. 

Pleon consisting of three complete and one incomplete pleonites plus 

pleotelson. 

Cleantioides planicauda (Benedict, 1899) 

Figure 108A 

DIAGNOSIS Ovigerous 9 5.5 mm. Body parallel sided. Maxillipedal palp of 

five articles. Pleotelson posteriorly broadly rounded, with obliquely truncate 

subcircular dorsal area in posterior half. 

RECORDS Maryland to Florida; Puerto Rico; Panama; Louisiana, Gulf of 

Mexico, intertidal to 44 m; often in hollow stems and roots of seagrasses, and 

tubes of the polychaete Diopatra cuprea. 

Oaxaca, Pacific Mexico. 

REMARKS Cleantioides planicauda has been recorded only once in the eastern 

Pacific, where it occurs with the more common C. occidentalis (Richardson, 

1899). 

Key to species of Erichsonella 

1. Pereonites with dorsal spines 2 

Pereonites lacking dorsal spines attenuata 

2. Pereonites 1—4 with middorsal and lateral spines floridana 

Pereonites 1-4 with middorsal spines only Jiliformis

Erichsonella 257 

Figure 108. Ay Cleantioides planicauda; B, Erichsonella attenuata 8\ C, Erichsonella 

filiformis 6\ Dy Erichsonella Jloridana 9; E, Idotea balthica 6] F, Idotea metallica $; G, 

Miratidotea bruscai $. 

Erichsonella Richardson, 1901 

DIAGNOSIS Antennal flagellum clavate. Maxillipedal palp of four articles. 

Uropod uniramous. Pleonites completely fused with pleotelson. 

REMARKS Pires (1984) reviewed the genus Erichsonella and did not recog 

nize the subspecies E. filiformis tropicalis Menzies and Glynn, 1968.


Erichsonella attenuata (Harger, 1873) 

Figure 108B 

DIAGNOSIS 6 11.4 mm, ovigerous 9 12.0 mm. Body dorsally smooth. 

Cephalon lacking middorsal elevation. Antcnnulc reaching only slightly be 

yond antennal peduncular article 2. Pleotelson with slight marginal bulge in 

anterior half, indicating ventrolateral articulation of uropod. 

RECORDS Connecticut to Miami; Florida, Mississippi, Texas, Gulf of Mex 

ico; intertidal to 2 m, usually associated with submerged seagrass and algal 

beds. 

REMARKS While not recorded in the Florida Keys, this species does reach 

Miami, and continues into the Gulf of Mexico. 

Erichsonella filifor mis (Say, 1818) 

Figure 108C 

DIAGNOSIS 6 10.5 mm, ovigerous 9 8.2 mm. Body dorsally with bifid tu 

bercle on cephalon, and low rounded middorsal tubercle on pereonites. An- 

tennule reaching midlength of antennal peduncular article 3. Basis of per- 

eopods 2—7 with larges tubercles. Pleotelson with distinct lateral shoulder in 

anterior half. 

RECORDS Connecticut to Florida, shallow infratidal to 55 m; Bahamas; 

Turks and Caicos Islands; Puerto Rico; Quintana Roo, Yucatan Peninsula, 

Mexico, 60-109 m; Florida and Texas, Gulf of Mexico. 

Brazil. 

Erichsonella jloridana Richardson, 1901 

Figure 108D 

DIAGNOSIS Ovigerous 9 10.0 mm. Antennule reaching distal end of anten 

nal peduncular article 3. Cephalon with strong trifid tubercle. Pereonites 1—7 

each with posteriorly directed spine near posterior margin; pereonites 1—4 

each with lateral spine. Basis of pereopods 2-7 smooth. 

RECORDS Florida Keys, intertidal to 2 m; Florida, Gulf of Mexico, interti 

dal mud flats.

Idotea Fabricius, 1798 

Miratidotea 259 

DIAGNOSIS Antennal flagellum multiarticulate. Maxillipedal palp of four or 

five articles. Uropod uniramous. Pleon consisting of two complete and one 

incomplete pleonites plus pleotelson. 

Key to species of Idotea 

1. Posterior margin of pleotelson truncate metallica 

Posterior margin of pleotelson with distinct median lobe balthica 

Idotea balthica (Pallas, 1772) 

Figure 108E 

DIAGNOSIS 6 24.5 mm, ovigerous 9 13.2-23.5 mm. Anterior margin of 

cephalon concave. Cephalon dorsally smooth. Pereonites evenly convex, 

smooth. Posterior margin of pleotelson with rounded median lobe. 

RECORDS Worldwide in tropical to cold-temperate waters, often on floating 

seaweed, from surface to 357 m. 

Idotea metallica Bosc, 1802 

Figure 108F 

DIAGNOSIS 6 30.0 mm, ovigerous 9 22.2 mm. Cephalon with sinuous fur 

row in posterior half. Pereonites 2—4 laterally with rounded convex area close 

to coxae. Posterior margin of pleotelson truncate. 

RECORDS Worldwide in tropical to cold-temperate waters, often on floating 

seaweed, from surface to 200 m. 

Miratidotea Kensley, 1987a 

DIAGNOSIS Antennal flagellum of single clavate article. Maxillipedal palp 

of four articles. Uropod uniramous. Pleon consisting of two complete and two 

incomplete pleonites plus pleotelson.


Miratidotea bruscai Kensley, 1987a 

Figure 108G 

DIAGNOSIS Ovigerous 9 13.0 mm. Body parallel sided. Maxillipedal palp 

of four articles, terminal article very short. Pereopods 1—3 increasing in 

length posteriorly, pereopod 4 reduced, shorter than pereopod 5, and with 

dactylus spinelike, pereopods 5—7 increasing in length. Pleotelson consisting 

of two complete and two incomplete pleonites plus pleotelson; latter with 

broadly rounded posterior margin, and with bifid median process situated 

dorsal to posterior oblique-concave area. 

RECORDS Carrie Bow Cay, Belize, 1.5 m, in hollow root-internodes of sea- 

grass Syringodium filiforme.

Zoogeography 

FAUNAL PROVINCES 

The area under discussion has been divided into several faunal regions or 

provinces, of which the Caribbean, West Indian, and Brazilian are the major 

ones (Briggs, 1974). The extent and boundaries of the provinces have been 

variously defined depending on the group of organisms under discussion. 

Inevitably, zones of overlap exist, but for the purposes of this discussion, the 

following rough limits have been used. 

Brazilian Province: This province stretches from Cape Frio near Rio de 

Janeiro in Brazil to the mouth of the Orinoco River in Venezuela. The out 

flow of freshwater from the major rivers of this region has probably contrib 

uted to the isolation of the Brazilian coral reefs and their associated fauna 

from those of the Caribbean. This isolation is demonstrated by the consider 

able endemism of the Brazilian reef fauna and that of the Caribbean reef 

fauna, with very few species being common to both. 

Caribbean Province: This province has two components, a northern part 

in peninsular Florida, that stretches from around Cape Kennedy on the east 

coast to Tampa or Sanibel Island on the west coast, and a southern compo 

nent that runs from the mouth of the Orinoco River to around Cabo Rojo or 

Tampico on the gulf coast of Mexico. The northern Gulf of Mexico is ex 

cluded from this province and is characterized as being warm-temperate, 

rather than subtropical (Briggs, 1974:66). 

West Indian Province: This includes all the islands of the West Indian 

chain, the Bahamas, and the isolated outrider, Bermuda. The West Indian 

Province closely approaches the Caribbean Province in the Yucatan Penin 

sula to the north, and between Grenada and Trinidad in the south. There is 

also some indication of the isolating effect on the Bahamas of the Florida 

Current through the Straits of Florida. 

It has been suggested, on the basis of the molluscan fauna, that a relict of 

the Neogene Gatunian Province exists around northern Venezuela and Col 

ombia (Petuch, 1982). While several isopod species have been recorded only 

from this area, these are all described in a single paper that covers a very 

small part of this region (Paul and Menzies, 1971). There is as yet too little 

evidence to explore the idea of this relict fauna further. 

261

262 ZOOGEOGRAPHY 

ANALYSIS OF THE ISOPOD FAUNA 

In the following discussion, the West Indian and Caribbean provinces are 

treated as one, the isopod faunas offering little evidence to warrant a separate 

treatment of each. 

It is a truism that for any discussion of the zoogeography of an area to have 

meaning, the true extent of the fauna must be known. With the area under 

review, collecting effort has been uneven, and the true faunal composition of 

many regions is still incompletely known. Obviously, any conclusions based 

on such incomplete data are approximate and subject to revision. Neverthe 

less, certain general patterns or trends emerge when the present isopod fauna 

is broken down into its components. 

The deepwater isopod fauna of the Caribbean (i.e., from deeper than 200 

m) has barely been explored, and little is to be gained from discussing the 

relatively few species known. A list of these deeper dwelling species is in 

cluded (Table 4). 

Although about 280 shallow-water species are covered by this work, cer 

tain categories of species must be excluded, for various reasons, before anal 

ysis can be attempted. Such excluded groups include the species of Oniscidea 

(being essentially terrestrial forms and not part of the marine regime); the 

cymothoid species and the species of Aegidae (being fish parasites for at least 

part of their life history, and whose distribution is complicated by the dis 

tribution and mobility of the hosts); the limnoriids (being wood-borers whose 

distribution is more a function of the distribution of floating wood); and the 

true cave species (which have a history more reflective of the geological his 

tory of the area than of the marine regime). The epicarideans have a distribu 

tion somewhat complicated by the distribution of their crustacean hosts and 

their pelagic epicaridean and cryptoniscan larvae. Nevertheless, the decapod 

hosts of the great majority of species covered here are Caribbean endemics, 

and inclusion of the epicarideans changes very little the overall patterns of 

distribution, as demonstrated by the two figures provided (Figure 109). After 

making these exclusions there remain about 166 species (218 with the epi 

carideans) that can be broken down into the following components (figures 

in brackets include epicarideans): 

1. True Caribbean/Bahamian species—124 species, 74.8% [147, 67.5%]. 

These are the species recorded only from the Caribbean and the Bahamas. 

The term endemic is avoided, as too little is known of the actual distribution 

of many species. Of these species, 86 [87] have been recorded from a single 

locality. 

2. Species occurring south of the discussion area, and extending into 

Brazil—5 species, 3.0% [9, 4.1%]. These low numbers indicate that the

TABLE 4. CARIBBEAN ISOPODS RECORDED FROM DEPTHS GREATER THAN 200 M 

SUBORDER ANTHURIDEA 

Family Paranthuridae 

Neoanthura coeca Menzies, 1956b. South of Jamaica, 1244 m 

SUBORDER ASELLOTA 

Family Dendrotiidae 

Dendrotion hanseni Menzies, 1956b. South of Jamaica, 1244 m 

Family Desmosomatidae 

Desmosoma magnispina Menzies, 1962a. Bay of Panama, 1906 m 

Family Echinothambematidae 

Echinothambema ophiuroides Menzies, 1956a. North of Puerto Rico Trench, 

5104-5122 m 

Family Eurycopidae 

Acanthocope spinosissima Menzies, 1956b. South of Jamaica, 1224 m 

Storthyngura pulchra caribbea (Benedict, 1901). Off Windward Islands, 1256 m 

Storthyngura snanoi Menzies, 1962a. Colombia abyssal plain, 4071 m 

Family Haploniscidae 

Antennuloniscus dirneroceras (Barnard, 1920). North of Puerto Rico Trench, 

5440-5410 m; South Atlantic off South and West Africa, 1400-3921 m; 

off Argentina, 5843 m 

Haploniscus unicornis Menzies, 1956a. North of Puerto Rico Trench, 5104- 

5122 m 

Hydroniscus quadrifrons Menzies, 1962a. North of Puerto Rico Trench, 5271- 

5684 m 

Family Ischnomesidae 

Haplomesus tropicalis Menzies, 1962a. Colombia abyssal plain, 4071 m; off 

South Africa, 2526 m; Mediterranean 

Heteromesus bifurcatus Menzies, 1962a. Colombia abyssal plain, 4071 m 

Ischnomesus armatus Hansen, 1916. North of Puerto Rico Trench, 5494-5477 

m; Davis Straits, 2702 m 

Ischnomesus caribbicus Menzies, 1962a. Off Panama, 1714 m 

Ischnomesus multispinis Menzies, 1962a. Off Panama, 975 m 

Family Janiridae 

Abyssianira dentifrons Menzies, 1956a. North of Puerto Rico Trench, 5104- 

5122 m; off Argentina, 5024-5293 m; off southwest Africa, 4588 m 

Ianirella caribbica Menzies, 1956b. South of Jamaica, 1244 m 

Ianirella vemae Menzies, 1956a. Near Puerto Rico Trench, 5104-5122 m 

Spinianirella serrata Kensley and Heard, 1985. Off Puerto Rico, 350 m 

Family Macrostylidae 

Macrostylis caribbicus Menzies, 1962a. Off Colombia, 2875-2941 m 

Macrostylis minutus Menzies, 1962a. North of Puerto Rico Trench, 5163- 

5494 

{continued)


TABLE 4. {Continued) 

Macrostylis setifer Menzies, 1962a. North of Puerto Rico Trench, 5477- 

5494 m 

Macrostylis vemae Menzies, 1962a. North of Puerto Rico Trench, 5410— 

5684 m 

Family Mesosignidae 

Mesosignum kohleri Menzies, 1962a. Colombia abyssal plain, 2868-4076 m 

Family Nannoniscidae 

Nannoniscus camayae Menzies, 1962a. Off Panama, 1714 m 

SUBORDER GNATH11DEA 

Family Gnathiidae 

Akidognathia poteriophora Monod, 1926. OffU. S. Virgin Islands, 914 m. 

SUBORDER VALVIFERA 

Family Arcturidae 

Antarcturus annaoides Menzies, 1956b. South of Jamaica, 1244 m 

Arcturus caribbaeus Richardson, 1901. OfTAves Island, 1360 m 

Arcturus purpureus Beddard, 1886. Off Leeward Islands, 900 m 

Note: Records from deep water around Bermuda are not included. 

great area of mixed-salinity waters resulting from the outflow of the Orinoco, 

Amazon, Tocantins, and Parnaiba rivers form an effective barrier to the 

movement of shallow-water isopod species. 

3. Species having an amphi-Panamic distribution—7 species, 4.2% [8, 

3.7%] (Table 5). In spite of the history of immergence and emergence of the 

Isthmus of Panama, this very small amphi-Panamic component in the Carib 

bean isopod fauna suggests that most of this fauna has evolved since the last 

emergence of the late Pliocene. Given the limited mobility of most isopod 

species, the Panama Canal seems to have played a minimal role in contribut 

ing to this component. 

4. Species occurring outside of the western Atlantic (but excluding the 

amphi-Panamic species)—3 species, 1.8% [7, 3.2%]. 

5. The role of the Gulf of Mexico isopod fauna (see Clark and Robertson, 

1982) in the composition of the Caribbean/Bahamian is complex and diffi 

cult to analyze. One hundred and thirteen species of shallow-water isopods 

have been recorded from the Gulf of Mexico (Table 6). This number would 

indicate that many species remain to be recorded in this region. Of these 113 

species, 61 (54%) have also been reported from the Caribbean region. It is 

therefore possible that there exists a true Gulf of Mexico fauna, whose evolu 

tion was perhaps spurred by the relative isolation and reduction of the Gulf

Car 

Car 

Sout 

South 3.0% % 

Panam 4.2% 

Epicaridea excluded 

Panam 3.7% 

Out 3.2% 

North 

Epicaridea included 

5.5% 

ZOOGEOGRAPHY 265 

GoM/N/Car 7.2% 

oM/Car 5.4% 

3.6% 

GoM/N/Car 9.6% 

/Car 6.4% 

Figure 109. Relative proportions of the zoogeographic components of the 

Caribbean isopod fauna, with and without the parasitic Epicaridea. Car, 

Caribbean; Out, extra-western Atlantic; GoM/Car, Gulf of Mexico-Caribbean; 

GoM/N/Car, Gulf of Mexico-Northern-Caribbean; North, northern; Panam, amphi- 

Panamic; South, southern. 

during a low-water stand (100 m below present sea level) during the 

Pleistocene. A significant proportion (about 27 species, 28%) of the Gulf of 

Mexico isopods are known from the eastern coast of the United States north


TABLE 5. SPECIES OF ISOPODS OCCURRING ON BOTH SIDES OF THE ISTHMUS 

OF PANAMA 

*Aega deshaysiana (H. Milne Edwards, 

1840) 

Anopsilana browni (Van Name, 1936) 

Cleantioides planicauda (Benedict, 

1899) 

Excirolana braiiliensis Richardson, 

1912 

Excorallana tricornis (Hansen, 1890) 

*Nerocila acuminata Schioedte and 

Meinert, 1881 

* fish parasite or fish predator 

of Cape Kennedy, which would indicate a significant cooler-water compo 

nent. What proportion of originally Gulf species have spread into the Carib 

bean, and what proportion of Caribbean and temperate east coast species 

have entered the Gulf, cannot yet be assessed, given our incomplete knowl 

edge of the Gulf fauna. Because of this unresolved situation, three categories 

of species have been separated: species ranging from north of Cape Kennedy 

into the Caribbean—6, 3.6% [12, 5.5%]; species occurring in the Gulf of 

Mexico and the Caribbean—9, 5.4% [14, 6.4%]; species occurring north of 

Cape Kennedy, in the Gulf, and in the Caribbean—12, 7.2% [21, 9.6%] The 

conclusion that the fauna of the Gulf of Mexico contains an endemic compo 

nent, a Caribbean component, and a warm-temperate component was also 

reached by Topp and HofT (1972), in an analysis of the pleuronectiform 

fishes of the Gulf. 

THE BAHAMAS 

The Florida Current flowing through the Straits of Florida has been sug 

gested as a factor in reducing the movement of shallow-water fauna between 

peninsular Florida and the Florida Keys on the west and the Bahamas on the 

east (Briggs, 1974). Comparison of the number of isopod species on either 

side of the Straits of Florida (13 from the Bahamas, 50 from southern penin 

sular Florida and the Florida Keys) supports this view. Of the 13 species 

from the Bahamas, only four arc "endemic," three of these being interstitial 

microcerberideans. 

Paradella dianae (Menzies, 1962b) 

Paraleptosphaeroma glynni Buss and 

Iverson, 1981 

Probopyrus pandalicola (Packard, 

1879) 

*Rocinela oculata Harger, 1883 

*Rocinela signata Schioedte and 

Meinert, 1879 

Uromunna reynoldsi Frankenberg and 

Menzies, 1966

TABLE 6. ISOPOD SPECIES OCCURRING IN THE GULF OF MEXICO 

SUBORDER ANTHURIDEA 

*Accalathura crenulata (Richardson, 

1901) 

*Amakusantkura magnified (Menzies 

and Frankenberg, 1966) 

Cyathura polita (Stimpson, 1855) 

Horoloanthura irpex Menzies and 

Frankenberg, 1966 

Kupellonura formosa (Menzies and 

Frankenberg, 1966) 

*Mesanthura Jloridensis Menzies and 

Kruczynski, 1983 

Mesanthura hopkinsi Hooker, 1985 

Mesanthura pulchra Barnard, 

1925 

Paranthura Jloridensis Menzies and 


Ptilanthura tricarina Menzies and 


Skuphonura lindae Menzies and 


*Xenanthura brevitelson Barnard, 

1925 

SUBORDER ASELLOTA 

Carpias Jloridensis Menzies and 


Gnathostenetrioides pugio Hooker, 

1985 

*Joeropsis coralicola Schultz and 

McCloskey, 1967 

*Joeropsis rathbunae Richardson, 

1902 

Mexicope kensleyi Hooker, 1985 

Munnogonium wilsoni Hooker, 1985 

*Pleurocope Jloridensis Hooker, 

1985 

*Santia milleri (Menzies and Glynn, 

1968) 

*Stenetrium stebbingi Richardson, 

1902 


Uromunna hayesi Robertson, 1978 

*Uromunna reynoldsi Frankenberg 

and Menzies, 1966 

SUBORDER EPICARIDEA 

Allodiplophryxus Jloridanus Markham, 

1985 

*Aporobopyrina anomala Markham, 

1973 

*Azygopleon schmitti (Pearse, 1932) 

*Bopyrina abbreviata Richardson, 

1904 

*Bopyrione synalphei Bourdon and 

Markham, 1980 

*Cancricepon choprae (Nierstrasz and 

Brender a Brandis, 1925) 

Dactylokepon sulcipes Adkison, 1982 

Eophryxus subcaudalis (Hay, 1917) 

*Gigantione mortenseni Adkison, 

1984b 

Gigantione uberlackerae Adkison, 

1984b 

*Hemiarthrus synalphei (Pearse, 

1950) 

Hyperphrixus castrensis Markham, 

1985 

*Munidion longipedis Markham, 

1975a 

Ovobopyrus alphezemiotes Markham, 

1985 

Parabopyrella mortenseni (Nierstrasz 

and Brender a Brandis, 1929) 

*Parabopyrella richardsonae 

(Nierstrasz and Brender a 

Brandis, 1929) 

Parabopyriscus stellatus Markham, 

1985 

*Probopyria alphei (Richardson, 

1900b) 

Probopyrinella heardi Adkison, 

1984a 

{continued)

268 

TABLE 6. (Continued) 

*Probopyrinella latreuticola (Gissler, 

1882) 

Prodajus cf. bigelowiensis Schultz and 

Allen, 1982 

Pseudione cognata Markham, 1985 

Pseudione upogebiae Hay, 1917 

*Schizobopyrina urocaridis (Richardson, 

1904) 

*Stegophryxus hyptius Thompson, 1902 

*Synsynella choprae (Pearse, 1932) 

*Synsynella deformans Hay, 1917 

Synsynella Integra Bourdon, 1981 

* Urobopyrus processae Richardson, 1904 

SUBORDER FLABELLIFERA 

*Aega deshaysiana (H. Milne Edwards, 

1840) 

*Aega ecarinata Richardson, 1898 

Aega incisa Schioedte and Meinert, 

1879 

Alcirona krebsii Hansen, 1890 

Ancinus depressus (Say, 1818) 

Anilocra acuta Richardson, 1910 

Anilocra laticauda H. Milne Edwards, 

1840 

*Bathynomus giganteus A. Milne 

Edwards, 1879 

*Cassidinidea ovalis (Say, 1818) 

Ceratothoa transversa (Richardson, 

1900b) 

Cirolana borealis Lilljeborg, 1851 

*Cirolana obtruncata Richardson, 1901 

* Cirolana parva Hansen, 1890 

Conilera cylindracea (Montagu, 1804) 

*Cymothoa caraibica Bovallius, 1885 

*Cymothoa excisa Perty, 1833 

*Cymothoa oestrum (Linnaeus, 1793) 

*Cerceis carinata Glynn, 1970 

*Eurydice convexa Richardson, 1900b 

Eurydice littoralis (Moore, 1901) 

*Eurydice piperata Menzies and 


*Excirolana braziliensis Richardson, 

1912a 

*Excirolana mayana (Ives, 1891) 

Êxcorallana antillensis (Hansen, 

1890) 

Excorallana mexicana Richardson, 

1905a 

*Excorallana tricornis (Hansen, 

1890) 

*Harrieta faxoni (Richardson, 

1905a) 

*Limnoria tuberculata Sowinsky, 

1884 

Lironeca ovalis (Say, 1818) 

* Lironeca redmanni Leach, 1818 

Lironeca texana Pearse, 1952 

Lironeca tropicalis Menzies and 


*Nalicora rapax Moore, 1901 

*Nerocila acuminata Schioedte and 

Meinert, 1881 

Olencira praegustator (Latrobe, 1802) 

*Paracerceis caudata (Say, 1818) 

*Paradella dianae (Menzies, 1962b) 

Paradynamene benjamensis 

Richardson, 1905 

*Politolana polita (Stimpson, 1853) 

*Rocinela insularis Schioedte and 

Meinert, 1879 

*Rocinela oculata Harger, 1883 

*Rocinela signata Schioedte and 

Meinert, 1879 

*Serolis mgrayi Menzies and 


*Sphaeroma quadridentata Say, 1818 

*Sphaeroma terebrans Bate, 1866 

SUBORDER GNATHIIDEA 

Gnathia Jloridensis Menzies and 


SUBORDER MICROCERBERIDEA 

Microcerberus mexicanus Pennak, 1958

SUBORDER VALVIFERA 

Antarcturus jloridanus (Richardson, 

1900b) 

Arcturella bispinata Menzies and 


Arcturella spinata Menzies and 


Astacilla lauffi Menzies and 


Chiridotea excavata Harper, 

1974 

Cleantioides planicauda (Benedict, 

1899) 

species also occurring in the Caribbean 


Edotea lyonsi (Menzies and 

Kruczynski, 1983) 

Edotea montosa (Stimpson, 1853) 

Edwinjoycea horologium Menzies and 


*Erichsonella attenuata (Harger, 

1873) 

* Erichsonella filiformis (Say, 1818) 

* Erichsonella Jloridana Benedict, 

1901 

Erichsonella isabelensis Menzies, 

1951b 

*Idotea metallica Bosc, 1802 

Note: Records for the Gulf of Mexico have been assembled from published 

BERMUDA 

literature; in most cases, actual material has not been examined. 

Twenty-nine species of isopods have been recorded from Bermuda (Table 7). 

Of these, nine are endemics (three being cave forms). The remaining 20 

species have all been recorded from the Caribbean region, indicating a strong 

subtropical connection, in spite of the relatively high latitude (32°15'N). Al 

though Bermuda is of Eocene or Oligocene age, the tropical fauna was prob 

ably decimated by the low temperatures of the last Pleistocene glaciation 

(Briggs, 1974:76). 

GAVE ISOPODS 

With the expanding efforts of cave divers, more and more true stygobiont 

forms are being found. Concurrently, discussion of the origin of cave fauna 

has spurred several theories, all invoking the geological history of the Carib 

bean area. 

Among the isopods, cave forms have been found in four suborders, the 

Asellota, Anthuridea, Flabellifera, and Microcerberidea. Two valuable dis 

cussions on the origin of cave crustaceans may be found in Stock (1986) and 

Wagele (1985). 

The only true cave asellote, Atlantasellus cavernicolus Sket, was collected 

from Bermuda.


TABLE 7. ISOPOD SPECIES OCCURRING AT BERMUDA 

Alcirona krebsi Hansen, 1890 

*Anthomuda stenotelson Schultz, 1979 

*Apanthura harringtortiensis Wagele, 

1981 

*Arubolana aruboides (Bowman and 

Iliffc, 1983) 

*Atlantasellus cavernicolus Sket, 1979 

Bopyrissa wolffi Markham, 1978 

Cancricepon choprae (Nierstrasz and 

Brender a Brandis, 1925) 

Carpias bermudensis Richardson, 1902 

*Carpias minutus (Richardson, 1902) 

*Colanthura tenuis Richardson, 1902 

Colopisthus parvus Richardson, 1902 

* Curassanthura bermudensis Wagele, 

1985 

Dynamenella perforata (Moore, 1901) 

Eurydice per sonata Kensley, 1987b 

* recorded only from Bermuda 

Excorallana quadricornis (Hansen, 1890) 

Joeropsis rathbunae Richardson, 1902 

Leidya bimini Pearse, 1951 

Paracerceis caudata (Say, 1818) 

Paranthura infundibulata Richardson, 

1902 

Pendanthura tanaiformis Menzies and 

Glynn, 1968 

Para theIges piriformis Markham, 1972b 

Parathelges tumidipes Markham, 1972b 

Probopyrinella latreuticola (Gissler, 

1882) 

Pseudione affinis (Sars, 1882) 

*Stegias clibanarii Richardson, 1904 

Stenetrium stebbingi Richardson, 1902 

Stenobermuda acutirostrata Schultz, 1979 

Synsynella choprae (Pearse, 1932) 

Synsynella deformans Hay, 1917 

The anthuridean cave representatives are found in two families: the genus 

Curassanthura Kensley in the Paranthuridae, and the genus Cyathura subgenus 

Stygocyathura Botosaneanu and Stock in the Anthuridae (see Figure 110). 

Three species of Curassanthura are known, one each from Curagao, Ber 

muda, and Lanzarote in the Canary Islands. Curassanthura halma Kensley, 

from Curagao, is an interstitial form found in hypersaline waters. Curas 

santhura bermudensis Wagele was found in water of about 26%o salinity. The 

Lanzarote species, C. canadensis Wagele, came from seawater in a lava cave. 

Wagele (1985) suggests that this amphi-Atlantic distribution of Curassanthura 

is the result of plate tectonics separating an ancestral hypogean progenitor 

that had a Tethyan distribution. 

The genus Cyathura has representatives in the sea, in estuarine-brackish 

habitats, and in freshwater caves, and is found in the Atlantic, Indian, and 

Pacific oceans. This widespread distribution suggests a very long history for 

the genus. Using the morphology of the male copulatory stylet, Wagele 

(1985) suggests that marine ancestors, having a Tethyan distribution, en 

tered freshwater interstitial habitats. The series of regressions of sea level 

forms. 

serv

Figure 110. Map showing distribution of cave anthurideans.

* cave cirolanids 

Figure 111. Map showing distrib


The flabelliferan family Cirolanidae contains five stygobiont genera in the 

Caribbean: Anopsilana, Arubolana, Bahalana, Creaseriella, and Haptolana (Figure 

111). Six other genera are known from the North American continent: 

Antrolana, Cirolanidesy Mexilana, Speocirolana, Sphaerolana, and Troglocirolana, all 

of which, except Antrolana from the Appalachian Valley of Virginia, occur in 

Mexico and Texas (see Notenboom, 1981). A few of these forms occur in 

brackish water, but most are found in freshwater of caves. Cave cirolanids 

are also known from Palau, North and East Africa, Madagascar, Bulgaria, 

Greece, Jugoslavia, Israel, France, and Spain. This widespread distribution 

again suggests a Tethyan marine origin, with dispersal and isolation due to 

sea regressions. 

The suborder Microcerberidea and the asellotan family Microparasellidae 

contain almost entirely interstitial forms, although few occur in caves. At 

least two genera, Microcerberus and Angliera, have very widespread distribu 

tions and are known from marine, brackish-water, and freshwater habitats, 

and may well have a history similar to that of Cyathura.

Appendix 

Since the manuscript of this work was completed and sent to press, a few 

papers have appeared either describing new species, mentioning new 

records for the Caribbean and associated areas, or instituting a major 

new taxon. It was thought useful to include these, if only in an appendix, 

to make the work as current as possible. The relevant taxa are listed 

alphabetically, with the full citation given below. 

Antheluridae Poore and Lew Ton, 1988 

Poore, G. C. B., and H. M. Lew Ton. 1988. Antheluridae, a new 

family of Crustacea (Isopoda: Anthuridea) with new species from 

Australia. Journal of Natural History 22:489—506. 

Within the geographical area covered by this work, only Anthomuda 

belongs to this new family. 

Aporobopyrus collardi Adkison, 1988 

Adkison, D. L. 1988. Pseudione parviramus and Aporobopyrus collardi, 

two new species of Bopyridae (Isopoda: Epicaridea) from the Gulf 

of Mexico. Proceedings of the Biological Society of Washington 

101(3):576-584. 

Booralana tricarinata Camp and Heard, 1988 

Camp, D. K., and R. W. Heard. 1988. Booralana tricarinata, a new 

species of isopod from the western Atlantic Ocean (Crustacea: Iso 

poda: Cirolanidae). Proceedings of the Biological Society of Washington 

101 (3):603-613. 

Originally recorded from the outer shelf and upper slope off the 

Little Bahama Bank and the Antilles Islands in 110-610 m, this 

species has since been recorded off Haiti in 620 m. 

Bythognathia yucatanensis Camp, 1988 

Camp, D. K. 1988. Bythognathia yucatanensis, new genus, new species, 

from abyssal depths in the Caribbean Sea, with a list of gnathiid 

275

276 APPENDIX 

species described since 1926 (Isopoda: Gnathiidae). Journal of Crust 

acean Biology 8(4):668-678. 

Edotea samariensis Muller, 1988 

Muller, H. G. 1988. Idoteidae aus N-Kolumbien mit Beschreibung 

von Edotea samariensis n. sp. (Crustacea: Isopoda: Valvifera). 

Senckenbergiana biologica 68(4/6):407-412. 

Gnathia Johanna Monod, 1926 

Miiller, H. G. 1988. Redescription of Gnathia Johanna, 1926 (Iso 

poda) from St. John, Virgin Islands. Bulletin Zoologisch Museum, Uni- 

versiteit van Amsterdam 11(15): 129—133. 

Phycolimnoria bacescui Ortiz and Lalana, 1988 

Ortiz, M., and R. Lalana. 1988. Una nueva especie del genero Phy 

colimnoria (Isopoda, Limnoriidae) de aguas cubanas. Revista de Inves 

tigations Marinas, La Habana 9(2): 37—42. 

Pseudione parviramus Adkison, 1988 

Adkison, D. L. 1988. Pseudione parviramus and Aporobopyrus collardi, 

two new species of Bopyridae (Isopoda: Epicaridea) from the Gulf 

of Mexico. Proceedings of the Biological Society of Washington 

101(3):576-584.

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Index 

abbreviata, Bopyrina, 110, 267 

Abudefduf saxatilis, 171, 175, 186 

abudefdufi, Anilocra, 171, 175 

Abyssianira dentifrons, 263 

Acanthocope spinosissima, 263 

acanthopkora, Domecia, 111 

Acanthopleura granulata, 215 

acanthura, Anopsilana, 124, 125 

acanthuri, Anilocra, 171, 175, 176 

Acanthurus 

bahianus, 171, 176 

chirurgus, 171, 176 

acanthurus, Macrobrachium, 112 

crenulata, 64, 65, 267 

setosa, 64, 65 

Achelion occidentalism 110 

acuminata, Nerocila, 171, 172, 173, 190, 266 

268 

forma acuminata, 190 

forma &yter, 190 

acuta, Anilocra, 268 

acutirostrata, Stenobermuda, 106, 270 

acutirostris, Processa, 113 

acutitelson, Dynamenella, 213, 214 

ilagfl, 116, 117 

antillensiSy 117 

incisa, 268 

-4i?£fl (Aega), 116, 117 

deshaysiana, 117, 266, 268 

ecarinata, 117, 268 

-dtfgfl (Rhamphion), 116, 117 

^wtata, 117, 119 

tenuipes, 117, 119 

Aegidae, 115, 116, 262 

Aetobatus narinari, 166 

Pseudione, 113, 270 

Upogebia, 112 

Agaricia, 88, 166 

agaricicola, Metacirolana, 153, 154 

A£

294 INDEX 

angulata 

Dynamene, 214 

Dynamenella, 214 

angustifrons, Hexapanopeus, 111 

Anilocra, 170, 174, 175 

abudefdufi, 171, 175 

acanthuri, 171, 175, 176 

acuta, 268 

chaetodontis, 171, 175, 177 

fAnwiw, 172, 175, 177 

haemuli, 172, 173, 175, 177 

kolacantki, 172, 175, 179 

holocentri, 175, 179 

laticauda, 268 

myripristis, 172, 175, 180 

/wr/i/i, 173, 175, 180 

annandalei, Sphaeroma, 234 

annaoides, Antarcturus, 264 

annulicornis, Pandalus, 113 

annulipes, Pagurus, 113 

anomala, Aporobopyrina, 110, 267 

anops, Creaseriella, 137 

Anopsilana, 124, 273 

acanthura, 124, 125 

ArMWH, 124, 125, 266 

owiata, 124, 125 

cubensis, 124, 126 

;m, 124, 127 

radicicola, 124, 127 

annaoides, 264 

jloridanus, 269 

Antennuloniscus dimeroceras, 263 

Antheluridae, 275 

Anthomuda, 17, 23, 291 

stenotehon, 23, 270 

Anthuridae, 16, 270 

Anthuridea, 2, 14, 15, 16, 244, 263, 269 

antiguai, Plesionika, 113 

antillense, Exosphaeroma, 229, 230 

antillensis 

Aega, 117 

Dromidia, 111 

Excorallana, 161, 162, 268 

Paranthura, 69 

Antrolana, 273 

Apanthura? 17, 25 

cracenta, 25 

cram, 25, 26 

harringtoniensis3 25, 26, 270 

Apanthuroides? 17, 26 

millae, 27 

Aphysina fistulariSy 223 

lachneri, 171, 188 

maculatus, 171, 193 

townsendi, 171, 193 

Apogonidae, 187 

Aporobopyrina, 110 

anomala, 110, 267 

Aporobopyrus, 110 

collardi, 275 

curtatus, 110 

arbuscula, Oculina, 88 

Archosargus probatocephalus, 122 

Arcturella, 252 

bispinata, 252, 269 

#wiflte, 252, 269 

Arcturidae, 251, 252, 264 

caribbaeus, 264 

purpureus, 264 

arenatus, Priacanthus, 173, 183 

Argeia, 110 

atlantica, 110 

Argeiinae, 107 

Ariusfelis, 171, 190 

Armadilloniscus; 247 

m'/zai, 247 

Iscfinomesus, 263 

Petrolisthes, 110 

armillatuSy Alpheus, 112 

arndti, Dies, 207 

aruboidesy Arubolana, 144, 145, 270 

Arubolana, 144, 273 

aruboides, 144, 145, 270 

/ma/a, 144, 145 

parvioculata, 144, 145 

ascensionis, Holocentrus, 172, 179 

Aselloidea, 75 

Asellota, 15, 15, 73, 263, 267, 269 

Astacilla, 252 

cymodocea, 252, 253 

/ajtf//

Atlantasellidae, 75 

Atlantasellus, 75 

cavernicolus, 75, 269, 270 

atlantica 

Argeia, 110 

Megalops, 172, 183 

attenuata, Erichsonella, 256, 258, 269 

aurolineatum, Haemulon, 172, 178 

Azygopleon, 110 

schmitti, 110, 267 

bacescui, Phycolimnoria, 276 

Bagatus, 83 

Bahalana, 124, 127, 128, 273 

cardiopus, 128 

geracei, 128 

mayana, 128 

bahianus, Acanthurus, 171, 176 

bajonado, Calamus, 122 

Balanopleon, 110 

tortuganus, 110 

Balistes vetula, 122 

balthica, Idotea, 259 

barbadensis, Micropanope, 111 

barbarae, Geogerceis, 215 

6anzar

296 INDEX 

Cancrion carolinus, 111 

capistratus, Chaetodon, 171, 177 

caraibica, Cymothoa, 182, 268 

Caranx, 122, 171 

hippos, 171, 183 

latus, 171, 183 

ruber, 171, 183 

carbonarium, Haemulon, 172, 178 

carcinus, Macrobrachium, 112 

cardiopus, Bahalana, 128 

caribaeus, Dactylokepon, 111 

caribbaeus, Arcturus, 264 

caribbea, Storthyngura pulchra, 263 

Ianirella, 263 

Malacanthura, 45 

Ischnomesus, 263 

Macrostylis, 263 

caribea, Uromunna, 94 

carinata, 'Cerceis,' 211, 268 

carolii, Metaphrixus, 111 

carolinense, Tozeuma, 112 

carolinus, Cancrion, 111 

Gzr/u

Colopisthus, 144, 146 

parvus, 147, 270 

confixa, Cortezura, 31 

Conilera cylindracea, 268 

Conilerinae, 123, 139 

conklini, Phaeoptyx, 173, 188 

constricta, Munida, 112 

convexa, Eurydice, 147, 148, 149, 268 

coralicolay Joeropsis, 88, 267 

Corallanidae, 115, 157 

corallicola, Minyanthura, 53 

corallinus, Pylopagurus, 112 

Cortezura, 17, 29 

confixa, 31 

penascoensis, 31 

cracenta, Apanthura, 25 

crassa, Virganthura, 73 

Creaseriella, 124, 137, 273 

anops, 137 

crenata, Anopsilana, 124, 125 

crenulata, Accalathura, 64, 65, 267 

crenulitelson, Cirolana, 132, 133 

cromis, Pogonias, 173, 190 

cruciaria, Astalione, 110 

cruris, Apanthura, 25, 26 

cruentatus, Epinephelus, 172, 179 

crumenophthalmus, Selar, 173, 183 

Cryptoniscoidea, 107 

Crytoniscidae, 107, 109 

cubana, Cyathura (Cyathura), 33 

cubense, Munidion, 111 


tfocme/a, 119, 120 

cuborientalis, Cyathura (Stygocyathura), 34, 35 

culebrae, Vandeloscia, 247, 251 

cumanensis, Malacanthura, 45 

cumulus, Agarna, 173 

cuprea, Diopatra, 256 

curacaoensis, Hippolyte, 110 

Curassanthura, 64, 67, 270 

bermudensis, 67, 270 

canariensis, 270 

Afl/mfl, 67, 68, 270 

curassavica, Cyathura (Stygocyathura), 35 

o/rn, Haliophasma, 41 

curtatus, Aporobopyrus, 110 

cuvieri, Galeocerdo, 122 

cyaneus, Chromis, 172, 177 

Cyathura, 17, 31, 270, 273 

/>o/z7a? 267 

(Cyathura), 31 

cubana, 33 

(Stygocyathura), 31, 33, 35, 270 

cuborientalis, 34, 35 

curassavica, 35 

hummelincki, 35 

motasi, 35, 36 

orghidani, 35, 36 

parapotamica, 35, 36 

salpiscinalis, 35, 38 

sbordonii, 35, 38 

specus, 35, 38 

univam, 35, 38 

Cyclograpsus interger, 111 

cylindracea, Conilera, 268 

Cymodoce, 226, 227 

barrerae, 227 

ruetzleri, 227 

Cymodocea, 223, 253 

cymododea, Astacilla, 252, 253 

Cymothoa, 170, 172, 182 

caraibica, 182, 268 

«OM, 172, 173, 182, 268 

INDEX 297 

wrfrem, 171, 172, 173, 182, 183, 268 

Cymothoidae, 115, 169, 170 

Cynoscion, 172, 183 

nebulosus, 172, 183 

Dactylokepon 

caribaeus, 111 

sulcipes, 267 

Dajidae, 107 

Dardanus jucosus, 1122 

Dasyatis americana, 122, 166 

decorata, Mesanthura, 53 

deformans, Synsynella, 110, 113, 268, 270 

delaneyi, Excorallana, 160, 161 

Dendrotiidae, 263 

Dendrotion hanseni, 263 

dentata, Aega, 117, 119 

dentifrons, Abyssianira, 263 

deplanata, Ceratothoa, 180 

depressa, Panoplax, 111 

depressus, Ancinus, 268 

deshaysiana, Aega, 117, 266, 268 

Desmosoma magnispina, 263 

Desmosomatidae, 263 

destructor, Sphaeroma, 235 

desultor, Asymmetrione, 110 

rfifl/M*, Paradella, 224, 266, 268 

Dicropleon 

periclimenis, 111 

Dictyota, 219 

Z)r>j, 207 

flnw/rf, 207 

barnardi, 207

298 INDEX 

dimeroceras, Antennuloniscus, 263 

diminuta, Exosphaeroma, 229, 231 

diogenes, Petrochirus, 110 

Diopatra cuprea, 256 

Diplophryxus, 111 

Discerceis, 210, 211 

linguicauda, 213 

dispar, Paraliomera, 111 

distorta, Leidya, 111 

Domecia 

acanthophora, 111 

hispida, 111 

Dromidia antillensis, 111 

dubitans, Angliera, 91 

Dynamene angulata, 214 

Dynamenella, 210, 213, 214, 224 

acutitelson, 213, 214 

var. glabrothorax, 214 

var. typica, 214 

angulata, 214 

perforata, 213, 215, 270 

quadrilirata, 213, 215 

Dynameninae, 204, 210, 211 

ecarinata, Aega, 117, 268 

Echinothambema ophiuroides, 263 

Echinothambematidae, 263 

edithae, Paracerceis, 218, 221 

Edo tea 

lyonsi, 269 

montosa, 269 

samariensis, 276 

edulis, Processa, 113 

edwardsi, Plesionika, 113 

Edwinjoycea horologium, 252, 269 

eglanteria, Raja, 122 

Eisothistos, 16, 38, 39 

petrensis, 39 

ten, 39 

mm, Plesionika, 113 

Entoniscidae, 107, 109 

Entophilinae, 107 

Eophrixus subcaudalis, 111, 267 

Epicaridea, 4, 14, 107, 267 

Epinephelus, 122, 172, 183 

cruentatus, 172, 179 

>/z^, 172, 179 

guttatus, 172, 179 

*7, 122, 172, 190 

morio, 122 

Erichsonella, 255, 256, 257 

attenuata, 256, 258, 269 

Jiliformis, 256, 258, 269 

tropicalis, 257 

floridana, 256, 258, 269 

isabelensis, 269 

Eriphia gonagra, 111 

Eubranchiatae, 203 

Eurycopidae, 263 

Eurydice, 143, 147 

con^xfl, 147, 148, 149, 268 

littoralis, 148, 149, 268 

personata, 147, 149, 270 

piperata, 147, 149, 268 

Eurydicinae, 123, 139, 143 

excavata, Chiridotea, 269 

Excirolana, 144, 149, 150 

braziliensis, 150, 266, 268 

mayana, 150, 153, 268 

Atrial, Cymothoa, 172, 173, 182, 268 

Excorallana, 157, 159, 161 

antillensis, 161, 162, 268 

berbicensis, 161, 162 

delaneyi, 160, 161 

fissicauda, 161, 162 

mexicana, 160, 161, 268 

oculata, 161, 163 

quadricornis, 161, 165, 270 

sexticornis, 161, 165 

subtilis, 160 

tricornis, 266, 268 

occidentalis, 167 


warmingii, 161, 167 

exilipes, Palaemonetes, 113 

Exocoettus, 172, 184 

Exosphaeroma, 226, 229, 231 

fl/Afl, 229 

antillense, 229, 230 

diminuta, 229, 231 

productatelson, 229, 231 

yucatanum, 229, 231 

exotica, Ligia, 247, 249 

faber, Chaetodipterus, 171, 190 

fasciata, Mesanthura, 47, 49 

faustinum, Macrobrachium, 112 

/flxwii, Harrieta, 232, 268 

felis,Arius, 171, 190 

filiforme, Syringodium, 253, 260 

Jiliformis, Erichsonella, 256, 258, 269 

fimbriata 

Pleurocryptella, 112 

Processa, 113 

fissicauda, Excorallana, 161, 162 

fistularis, Aphysina, 223 

Flabellifera, 2, 14, 114, 115, 268, 269 

Jlavolineatum, Haemulon, 122, 172, 178

Jlinti, Munida, 111 

jloridana 

Erichsonella, 256, 258, 269 

Pleurocrypta, 112 

florid anus 

Allodiplophryxus, 267 

Antarcturus, 269 

Thor, 110, 111 

jloridensis 

Carpias, 267 

Gnathic, 268 

Mesanthura, 53, 267 

Paranthura, 69, 71, 267 

Pleurocope, 98, 267 

joetens, Synodus, 173, 183 

foliatus, Parathelges, 112 

formosa, Kupellonura, 267 

formosus, Alpheus, 111, 112 

fritzmuelleri, Synalpheus, 110, 111 

jucorum, Latreutes, 112 

Jucosus, Dardanus, 112 

Julvus, Epinephelus, 172, 179 

Jurcifer, Paranthias, 173, 179 

Galathea rostrata, 112 

galathinus, Petrolisthes, 110 

Galeocerdo cuvieri, 122 

geminsula, Amakusanthura, 18 

Geocerceis, 210, 215 

barbarae, 215 

geracei, Bahalana, 128 

Genes rhombeus, 172, 187 

giardi, Synalpheion, 113 

giganteus, Bathynomus, 131, 268 

Gigantione 

mortenseni, 111, 267 

uberlackerae, 267 

Ginglymostoma cirratum, 122, 158 

glabrothorax, var., Dynamenella acutitelson, 214 

Glossobius, 170, 172, 183, 184 

hemiramphi, 172, 184 

impressus, 172, 184 

Paracerceis, 218, 221 

Paraleptosphaeroma, 210, 266 

Cwato, 238 

beethoveni, 238, 239 

Jloridensis, 268 

gonzalezi, 238, 239 

Johanna, 238, 239, 276 

magdalenensis, 238, 239 

puertoricensis, 238, 239 

ra^z, 238, 241 

samariensis, 238, 241 

triospathiona, 238, 241 

ye/

300 INDEX 

hemiramphi3 Glossobius, 172, 184 

Hemiramphidae, 187 

Hemiramphus 

bermudensis, 188 

brasiliensis, 172, 184 

hemphilli, Synalpheus, 110, 111 

hendleri, Pendanthura, 56 

herbstii, Panopeus, 111 

herrerai, Microcharon, 93 

heterocarpus, Plesionika, 113 

heterochaelis, A Ipheus, 112 

Heteromesus bijurcatus, 263 

Hexapanopeus angustifrons, 111 

Hippolyte 

curacao ensis 3 110 

pleutacanthus, 110, 111 

Zostericola, 110 

hippos, Caranx, 171, 183 

Hirundichthys speculifer, 172, 184 

hispida, Domecia, 111 

holacanthi, Anilocra, 172, 175, 179 

Holacanthus tricolor, 172, 179 

holocentri, Anilocra, 175, 179 

Holocentrus ascensionis, 172, 179 

hopkinsi, Mesanthura, 47, 51, 267 

Horoloanthura irpex, 267 

horologium, Edwinjoycea, 252, 269 

hummelincki, Cyathura (Stygocyathura), 35 

Hydroniscus quadrifrons, 263 

Hyperphrixus castrensis, 267 

Hypoconcha 

sabulosa, 111 

spinosissima, 111 

Hyporhamphus unifasciatus, 172, 188 

hyptius, Stegophryxus, 113, 268 

Hyssuridae, 16, 58, 60 

fanirella 

caribbica3 263 

vemae, 263 

Afotai, 255, 259 

balthica, 259 

metallicay 259, 269 

Idoteidae, 251, 252, 255 

Idoteinae, 255, 256 

Iliacantha 

liodactyla, 111 

subglobosa3 111 

imbricata, Parapagurion, 112 

impressa, Politolana, 140 

impressus, Glossobius, 172, 184 

imswe, Kupellonura, 60 

imw/a, Arubolana, 144, 145 

/nriftZj 4*£A, 268 

indica, Limnoria, 194, 195 

infundibulata, Paranthura, 69, 71, 270 

insulae, Limnoria, 195 

insular is 

Alcirona, 158 

A'ttrtfl, 170 

Rocinela, 119, 120, 268 

integra, Synsynella, 268 

interger, Cyclograpsus, 111 

intermedius, Palaemonetes, 113 

loninae, 107 

Iridopagurus, 112, 113 

zm, 113 

z'm, Iridopagurus, 113 

irmae, Haliophasma, 43 

fr/fcx, Horoloanthura, 267 

irrasa, Munida, 111 

irritans, Munidion, 111 

isabelensis, Erichsonella, 269 

Ischnomesidae, 263 

Ischnomesus 

armatus, 263 

caribbicus, 263 

multispinis, 263 

Ischyromene, 210, 214, 217 

bamardi, 218 

iVfljflrfl, Epinephelus, 122, 172, 190 

jacobus, Myripristis, 172, 180 

jacqueti, Sclerocrangon, 110 

Janiridae, 80, 81, 263 

Janiroidea, 75, 79, 80, 81 

Joeropsidae, 80, 87 

Joeropsis, 87, 88 

bifasciatus, 88 

coralicola, 88, 267 

personatus, 88, 90 

rathbunae, 88, 90, 267, 270 

JOAHHM, G/ifltfifl, 238, 239, 292 

jonesi, Anopsilana, 124, 127 

kadiakensis, Palaemonetes, 113 

kensleyi, Mexicope, 81, 267 

kohleri, Mesosignum, 264 

AWAJI, Alcirona, 158, 268, 270 

tfiwifl, 170, 184 

insularis, 171, 186 

Kupellonura, 60 

formosa, 267 

imswe, 60 

lachneri, Apogon, 171, 188 

Lachnolaimus maximus, 122 

Laminaria, 201 

lamprotaenia, Anchoa, 171, 187 

laodicense, Gnathostenetroides, 78

lasallae, Astacilla, 252, 253 

lata, Parabopyrella, 112 

lathridia, Amakusanthura, 18, 20 

laticauda, Anilocra, 268 

laticeps, Skuphonura, 58 

latreillei, Tylos, 247, 250 

Latreutes Jucorum, 112 

latreuticola, Probopyrinella, 112, 268, 270 

latus, Caranx, 171, 183 

/flw#?, Astacilla, 269 

Laurencia, 219 

Aimini, 111, 270 

distorta, 111 

Leiostomus xanthurus, 172, 183, 187, 

190 

Lepisosteus spatula, 172, 190 

leptorhynchus, Pandalus, 113 

lewisi, Chalixanthura, 27, 29 

Licranthura, 16, 43 

amyle, 43 

Lfcifl, 247, 249 

baudiniana, 247, 249 

wrrtiai, 247, 249 

302 INDEX 

mayana 

Bahalana, 128 

Excirolana, 150, 153, 268 

mcclendoni, Synalpheus, 110, 111 

Megalops atlantica, 172, 183 

menziesi, Metacirolana, 153, 154 

Mesantkura, 17, 45 

bivittata, 47 

decorata, 53 

fasciata, 47, 49 

JloridensiSy 53, 267 

hopkinsit479 51, 267 

looensis, 47, 51 

paucidens, 47, 51 

pulchra, 47, 52, 267 

punctillata, 47, 53 

reticulata, 47, 53 

Mesosignidae, 264 

Mesosignum kohleri, 264 

Metacirolana, 144, 153 

agaricicola, 153, 154 

Afl/ifl, 153, 154 

menziesi, 153, 154 

sphaeromiformis, 153, 154 

metallica, Idotea, 259, 269 

Metaphrixus carolii, 111 

mexicana, Excorallana, 160, 161, 268 

mexicanus, Microcerberus t 269 

Mexicope, 80, 81 

kensleyiy 81, 267 

Mexilana, 273 

mgrayiy Serolis, 202, 268 

Microcerberidae, 243, 244 

Microcerbendea, 14, 243, 269, 273 

Microcerberus, 244, 273 

littoralis, 244 

mexicanuSy 269 

minutuSy 244 

mirabiliSy 244 

nunezi, 244 

renaudiy 244 

simplex, 244 

syrtiaiSy 244 

Microcharon, 90, 91 

herrerai, 93 

phreaticus, 93 

sabulurrty 91 

Micropanope barbadensis, 111 

Microparasellidae, 80, 90, 273 

Microphrys bicornutus, 110 

//nV^r, Munida} 112 

millae, Apanthuroides, 27 

milleri, Santiay 99, 267 

minocule, Stenetrium, 100, 102 

minusy Synalpheus, 110, 113 

minuta, Cirolana, 132, 135 

Cfl#wj, 82, 84, 270 

MacrostyliSy 263 

Microcerberusy 244 

Minyanthura, 16, 53 

corallicola, 53 

mirabilis 

MadraciSy 29, 41 

Microcerberusy 244 

Miratidoteay 255, 259 

bruscai> 260 

Monacanthus ciliatuSy 172, 190 

montaguiy Pandalus, 113 

montosa, Edoteat 269 

morioy Epinephelus, 122 

Gigantione, 111, 267 

Parabopyrellay 112, 267 

mosaica, Cassidinidea, 208 

motasiy Cyathura (Stygocyathura), 35, 36 

Mothocya, 170, 187 

bohlkeorum, 171, 173, 187, 188 

wflwfl, 172, 187, 188 

Mugil cephalus, 172, 190 

multilineatus, Chromis, 172, 177 

multipunctata, Limnoria, 195, 196 

multispinis, Ischnomesus, 263 

constricta, 112 

TKnrt, 111 

irrasas 111 

longipeSy 112 

772 £/w, 112 

schroederiy 112 

simplex, 110 

stimpsoni, 111 

valida, 110 

Munidion 

cubensCy 111 

irritans, 111 

longipedisy 112, 267 

Munna, 93 

petronastes, 94 

Munnidae, 80, 93 

Munnogoniumy 96 

wilsoni, 96 

Mycteroperca 

bonaci, 122 

venenosa, 122 

Myripristis jacobus, 172, 180

myripristis, Anilocra, 172, 175, 180 

Nalicora, 157, 168 

rapax, 169, 268 

nana, Mothocya, 172, 187, 188 

Nannoniscidae, 264 

Nannoniscus camayae, 264 

narinari, Aetobatus, 166 

Natatolana, 139 

gracilis, 140 

nebulosus, Cynoscion, 172, 183 

Nemanthura, 43 

Neoanthura coeca, 263 

Neopanope 

packardii, 111 

ttxana sayi, 111 

Neostenetroides, 77, 78 

stocki, 78 

Nerocila, 170, 172, 188 

acuminata, 171, 172, 173, 190, 266, 268 

forma acuminata, 190 

forma aj/er, 190 

ninae, Armidilloniscus, 246, 247 

nfzwtf, T^/ar, 247, 250 

normanni, Alpheus, 112 

northropi, Palaemon, 113 

nunezi, Microcerberus, 244 

nuttingi, Paracerceis, 218, 223 

obtruncata, Cirolana, 132, 135, 268 

occiden talis 

Achelion, 110 

Cleantioides, 256 

Excorallana tricornis, 167 

Parathelges, 112 

ocellatus, Chaetodon, 171, 177 I 

Excorallana, 161, 163 

Rocinela, 119, 120, 266, 268 

Oculina arbuscula, 88 

Ocyurus chrysurus, 172, 183 

owfram, Cymothoa, 171, 172, 173, 182, 183, 

268 

ohione, Macrobrachium, 112 

Olencira praegustator, 268 

Zigia, 247, 249 

Macrobrachium, 112 

Ontilorpheus, 124, 139 

stebbingi, 139 

Oniscidea, 1, 4, 14, 15, 246, 247, 262 

ophiuroides, Echinothambema, 263 

Orbioninae, 107 

orghidani, Cyathura (Stygocyathura), 35, 

36 

Orthopristis 

chrysoptera, 172, 183 

raA^r, 122, 173, 179 

ovalis 

Cassidinidea, 207, 208, 268 

Lironeca, 268 

Ovobopyrus alphezemiotes, 267 

oxyophthalmus, Paguristes, 112 

Pachygrapsus transversus, 111 

packardii, Neopanope, 111 

Paguristes 

oxyophthalmus, 112 

tortugae, 112 

Pagurus 

annulipes, 113 

bonairensis, 110, 113 

brevidactylus, 112, 113 

longicarpus, 110, 113 

provenzanoi, 110, 112, 113 

Palaemon 

northropi, 113 

pandaliformis, 113 

Palaemonetes 

exilipes, 113 

intermedius, 113 

kadiakensis, 113 

paludosus, 113 

pugio, 113 

vulgaris, 113 

paludosus, Palaemonetes, 113 

pandalicola, Probopyrus, 112, 266 

pandaliformis, Palaemon, 113 

Pandalus 

annul ico rni s, 113 

Ari, 113 

leptorhynchus, 113 

montagui, 113 

pandionis, Synalpheus, 111, 113 

Panopeus herbstii, 111 

Panoplax depressa, 111 

Parabopyrella 

lata, 112 

mortenseni, 112, 267 

richardsonae, 112, 267 

thomasi, 112 

Parabopyriscus stellatus, 267 

Paracerceis, 210, 218, 223 

cflurfate, 219, 268, 270 

var. brevipes, 219 

roA«ifl*, 219, 220 

wfifAae, 218, 221 

£/>wii, 218, 221 

Hatting, 218, 223

304 INDEX 

Paradella, 210, 214, 223 

dianae, 224, 266, 268 

plicatura, 223, 224 

quadripunctata, 223, 224 

tumidicauda, 223, 226 

Paradynamene benjamensis, 268 

Paragnathia, 237 

Paraleptosphaeroma, 207, 208 

îww, 210, 266 

Paralimnoria, 193, 199 

andrewsi, 199 

forma A, 200 

forma B, 201 

forma typica, 200 

Paraliomera dispar, 111 

Paramunnidae, 80, 96 

Paranthias farcifer, 173, 179 

Paranthura, 64, 69 

antillensis, 69 

barnardiy 69, 71 

floridensis, 69, 71, 267 

infundibulatay 69, 71, 270 

Paranthuridae, 16, 64, 263, 270 

Parapagurion imbricata, 112 

Parapagurus, 112 

parapotamica, Cyathura (Stygocyathura), 35, 

Para theIges 

foliatus, 112 

occidentalism 112 

piriformis, 112, 270 

tumidipes, 112, 270 

partiti, Anilocra, 173, 175, 180 

partitus, Pomacentrus3 173, 180 

parva, Cirolana, 132, 135, 268 

parvioculata, Arubolana, 144, 145 

parviramus, Pseudione, 276 

parvus, Colopisthus, 147, 270 

patulipalma, Stenetrium, 100, 102 

paucidens, Mesantkura, 47, 51 

pectiniger, Synalpheus, 110, 111, 113 

penascoensis; Cortezura, 31 

Pendanthura, 17, 56 

hendleri, 56 

tanaiformis, 56, 270 

penna, Calamus, 122 

perforata, Dynamenella, 213, 215, 270 

Periclimenes 

americanus, 111, 113 

longicaudatus, 113 

periclimenis, Dicropleon, 111 

personata, Eurydice, 147, 149, 270 

personatus, Joeropsis, 88, 90 

petrensis, Eisotkistos, 39 

Petrochirus diogenes, 110 

Petrolisthes 

armatus, 110 

galathinus, 110 

marginatus, 110 

petronastesy Munna, 94 

pfefferi, Limnoria, 195, 198 

Phaeoptyx 

conklini, 173, 188 

pigmentaria, 173, 188 

phreaticuSy Microcharoriy 93 

Phreatocoidea, 14 

Phycolimnoria, 193, 201 

bacescuiy 276 

clarkae, 201 

Phyllodurinae, 107 

pigmentariay Phaeoptyx, 173, 188 

piperata, Eurydice, 147, 149, 268 

piriformis, Parathelges, 112, 270 

planicauda, Cleantioides, 256, 266, 268 

Platybranchiatae, 203 

platycauda, Limnoria, 195, 198 

Plesionika 

antiguai, 113 

edwardsiy 113 

mw, 113 

heterocarpus, 113 

martia, 113 

p leuracanthus, Hip poly te, 110, 111 

Pleurocope, 97 

floridensisy 98, 267 

Pleurocopidae, 80, 81, 96 

Pleurocryptella fimbriata, 112 

plicatura, Paradella, 223, 224 

plicifera, Callispongia, 221 

plumieri, Haemulon, 172, 179 

Pogonias cromis, 173, 190 

polita 

Cyathura, 267 

Politolana, 140, 143, 268 

Politolana, 139, 140 

impressa, 140 

ô/ito, 140, 143, 268 

Pomacentrus partitus, 173, 180 

Pontonia margarita, 113 

Porce liana say ana, 110 

A>n7^, 88, 90 

poteriophora, Akidognathia, 264 

praegustator, Olencira, 268 

Priacanthus arenatus, 173, 183 

probatocephalus, Archosargus, 122 

Probopyria alphei, 112, 267

Probopyrinella 

heardiy 267 

latreuticola, 112, 268, 270 

Probopyrus pandalicola, 112, 266 

Processa 

acutirostris y 113 

canaliculata y 113 

eduliSy 113 

Jimbriatay 113 

tenuipes, 113 

processaey Urobopyrusy 113, 268 

Prodajus cf. bigelowiensis, 268 

productatelson, Exosphaeroma3 229, 231 

provenzanoiy Pagurus> 110, 112, 113 

psamathus, Anglieray 91 

Pseudasymmetrione, 113 

Pseudione 

qffinis, 113, 270 

cognata, 268 

parviramuSy 276 

upogebiae, 268 

Pseudioninae, 107 

Ptilanthura tricarina, 267 

puertoricensis, Gnathia} 238, 241 

pugilatoty Uca, 111 

GnathostenetroideSy 77', 267 

PalaemoneteSy 113 

pulchra 

caribbedy Storthyngura, 263 

Mesanthura, 47, 52, 267 

punctatuSy Carpias, 82, 85 

punctillatdy Mesanthura, 47, 53 

purpureus, Arcturus3 264 

PylopaguruSy 110 

corallinus, 112 

quadricornisy Excorallana, 161, 165, 270 

quadridentata, Sphaeroma, 234, 268 

quadrifrons, Hydroniscus, 263 

quadriliratdy Dynamenella, 213, 215 

quadripunctata, Paradella, 223, 223 

racovitzai, Angliera, 91 

radicicola 


Xylolana, 157 

i?a/a eglanteridy 122 

rapaXy Nalicora, 169, 268 

rathbunae 

JoeropsiSy 88, 90, 267, 270 

Lysmata, 112 

rafAi, Gnathic, 238, 241 

redmanni, Lironecay 172, 173, 186, 268 

regalis, Scomberomorus, 173, 187 

regina, Astacilla, 252, 253 

renaudiy Microcerberus, 244 

Renocila, 170, 191 

bowmani, 173, 191 

«/!/!!, 171, 191 

waldneriy 173, 191, 193 

reticulatas Mesanthura, 47, 53 

INDEX 305 

reynoldsiy Uromunnay 94, 95, 266, 267 

RhamphioUy see i4«ga (Ramphion) 

Rhizophora mangle, 235 

rhombeus, GerreSy 172, 187 

Rhyscotus, 247, 249 

fexwuij, 247, 249 

richardsonae, Parabopyrella, 112, 267 

ricordiy Sesarma, 111 

nVrf/z, Vandeloscia, 247, 251 

Ritkropanopeus karrisiiy 111 

Rocinela, 116, 119 

cubensisy 119, 120 

insularis, 119, 120, 268 

oca/ate, 119, 120, 266, 268 

jignafa, 119, 120, 266, 268 

rostratay Galathea, 112 

Caranx, 171, 183 

Orthopristisy 122, 173, 179 

ruetzleri, Cymodoce, 227 

sabulosa, Hypoconcha, 111 

sabulum, Microcharony 91 

salpiscinalis, Cyathura (Stygocyathura), 35, 38 

samariensis 

Edotea, 276 

GnafAia, 238, 251 

Santiay 98 

ffiiV/m, 99, 267 

Santiidae, 80, 98 

Sargassum, 84, 201 

saseboensiSy Limnoria, 195, 198 

saxatilis, Abudefdufy 171, 175, 186 

sayana, Porcellana, 110 

ja>^", Neopanope texana, 111 

sbordoniiy Cyathura (Stygocyathura), 35, 

38 

Schizobopyrina urocaridis, 113, 268 

schmittiy Azygopleon, 110, 267 

schoepfi 

Alutera, 171, 190 

Chilomycterusy 172, 190 

schroederi, Munida, 112 

sciuruSy Haemulon, 172, 179 

Sclerocrangon jacqueti, 110 

Scomberomorus 

cavalla, 173, 187

306 INDEX 

Scomberomorus (cont.) 

maculatus, 173, 187 

regain, 173, 187 

scopulosa, Chalixanthura, 27, 29 

sedentarius, Chaetodon, 171, 177 

Selar crumenophthalmuSy 173, 183 

Serolidae, 114, 115, 201 

Serolis, 202 

mgrayi, 202, 268 

Serranus tigrinus, 173, 191, 193 

serrata, Spinianirella, 263 

serratum, Stenetrium, 100, 102 

serricaudus, Carpias, 82, 87 

Sesarma ricordi, 111 

seticornis, Stenorhynchus, 110 

setifer, Macrostylis, 264 

setosa, Accalathura, 64, 65 

sexticorniSy Excorallana, 161, 165 

Amakusanthura, 18, 21 

Rocinela, 119, 120, 266, 268 

signijica, Amakusanthura, 18, 23 

Microcerberus, 244 

Munida, 110 

simulata, Limnoria, 195, 198 

Skuphonura, 17, 58 

laticeps, 58 

lindae, 267 

snanoi, Storthyngura, 263 

somala, Haptolana, 138 

Sparisoma viride, 122 

spathulicarpusy Stenetrium, 100, 104 

spatula, Lepisosteus, 172, 190 

speculifer, Hirundichthys, 172, 184 

specus, Cyathura (Stygocyathura), 35, 38 

Speocirolana, 273 

Sphaerolana> 273 

Sphaeroma, 226, 232, 234 

annandalei, 234 

destructor, 235 

quadridentata, 234, 268 

terebrans, 234, 235, 268 

ttWfen, 234, 235 

Sphaeromatidae, 114, 115, 202, 204 

Sphaeromatinae, 204, 226 

sphaeromiformis, Metacirolana, 153, 

154 

Sphoeroides maculatus, 173, 190 

Sphyraena barracuda, 122 

j/>i»ata, Arcturella, 252, 269 

Spinianirella serrata, 263 

spinosissima 

Acanthocope, 263 

Hypoconcha, 111 

Oncilorpheus, 139 

Stenetrium, 100, 104, 267, 270 

Stegias clibanarii, 113, 270 

Stegophryxus hyptius, 113, 268 

steindachneri, Haemulon, 122 

Astrapogon, 171, 188 

Parabopyriscus, 267 

Stenetriidae, 99 

Stenetrioidea, 99 

Stenetrium, 99, 100 

bowmani, 100 

minocule, 100, 102 

patulipalma, 100, 102 

serratum, 100, 102 

spathulicarpus, 100, 104 

stebbingi, 100, 104, 267, 270 

Stenobermuda, 99, 106 

acutirostrata, 106, 270 

Stenorhynchus seticornis, 110 

stenotelson, Anthomuda, 23, 270 

stimpsoni, Munida, 111 

stocki, Neostenetroides, 78 

pulchra caribbea, 263 

snanoi, 263 

striata, Yvesia, 246 

striatus, Chaetodon, 171, 177 

Stygocyathura, see Cyathura (Stygocyathura) 

subcaudalis, Eophrixus, 111, 267 

subglobosa, Iliacantha, 111 

subtilis, Excorallana, 160 

sulcipes, Dactylokepon, 267 

surinamensis, Batrachoides, 171, 190 

surinamicum, Macrobrachium, 112 

symmetricus, Ambidexter, 113 

synagris, Lutjanus, 172, 183 

synalphei 

Bopyrione, 110, 267 

Hemiarthrus, 111, 267 

Synalpheion giardi, 113 

Synalpheus 

bousfieldi, 110 

brevicarpus, I ] 0 

AiwwW, 110, 111, 113 

fritzmuelleri, 110, 111 

goodei, 110, 111 

hemphilli, 110, 111

longicarpus, 110, 111, 113 

mcclendoni, 110, 111 

minus, 110, 113 

pandionis, 111, 113 

pectiniger, 110, 111, 113 

Synodus foe tens, 173, 183 

Synsynella, 110, 113 

choprae, 113, 268, 270 

deformans, 110, 113, 268, 270 

Integra, 268 

Syringodium, 166, 232 

filiforme, 253, 260 

syrticus, Microcerberus, 244 

tanaiformis, Pendanthura, 56, 270 

Tecticeps, 204 

Tecticipitinae, 204 

/l^, 117, 119 

Processa, 113 

tenuis, Colanthura, 65, 270 

tenuistylis, Lironeca, 171, 186, 187 

terebrans, Sphaeroma, 234, 235, 268 

ten, Eisothistos, 39 

testudinea, Thalassia, 251 

texana, Lironeca, 268 

texensis, Rhyscotus, 247, 249 

Thalassia, 166, 232 

testudinea, 251 

Thermarcturus, 252, 255 

venezuelensis, 255 

thomasi, Parabopyrella, 112 

floridanus, 110, 111 

manningi, 111 

f/Wn, Bopyrinella, 110 

tigrinus, Serranus, 173, 191, 193 

tortugae, Paguristes, 112 

tortuganus, Balanopleon, 110 

Tozeuma carolinense, 112 

transversa, Ceratothoa, 268 

transversus, Pachygrapsus, 111 

tricarina, Ptilanthura, 267 

tricarinata, Booralana, 275 

trichostoma, Haptolana, 138 

Clibanarius, 110, 112, 113 

Holacanthus, 172, 179 

tricornis, Excorallana, 161, 165, 266, 268 

occidentalis, 167 


Tridentella, 236 

virginiana, 236 

Tridentellidae, 115, 235, 236 

triospathiona, Gnathia, 238, 241 

tripunctata, Limnoria, 199 

/n/ow, Carpias, 82, 87 

Troglocirolana, 273 

tropicalis 

Erichsonella filiformis, 2bl 

Haplomesus, 263 

Lironeca, 268 

INDEX 307 

tuberculata, Limnoria, 194, 195, 199, 268 

tuberculatus, Chiton, 229 

tumidicauda, Paradella, 223, 226 

tumidipes, Parathelges, 112, 270 

Turbinaria, 166, 219 

7>/Wj 247, 250 

/

308 INDEX 

Virganthura, 64, 73 

crassa, 73 

virginalis, Gnathia, 238, 243 

virginiana, Tridentella, 236 

viridari, Alpheus, 112 

viride, Sparisoma, 122 

vittatus, Clibanarius, 110, 112 

vulgaris, Palaemonetes, 113 

waldneri, Renocila, 173, 191, 193 

walkeri, Sphaeroma, 234, 235 

warmingii, Excorallana, 161, 167 

wegeneri, Tylos, 247, 250 

wilsoni, Munnogonium, 96 

wolffi, Bopyrissa, 110 

wurdemanni, Lysmata, 112 

xanthurus, Leiostomus, 172, 183, 187, 190 

Xenanthura, 60 

brevitelson, 62, 267 

A>/

Part 2 - Sphaeromatidae::“Cute As Buttons”

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