Bijdragen tot de Dierkunde, 51 (2): 313-331 — 1981

Amsterdam Expeditions to the West Indian Islands, report 12.

Some new hypogean cirolanid isopod from Haiti

and Mayaguana (Bahamas)

by

Jos Notenboom

c/o Institute of Taxonomie Zoology, University of Amsterdam,

P.O. Box 20125, 1000 HC Amsterdam, The Netherlands

Abstract pus, est décrit d’une grotte de Mayaguana, île appartenant

de B. aux Bahamas. Cette espèce est congénérique geracei A of new stygobiont genus , Haitilana, Carpenter, 1981, de l’île de San Salvador (Bahamas), dont with two new species, H. radicicola and H. acanthura, are des elle se les inermes pléopodes described from distingue par endopodites groundwaters in Haiti. These are the first à l’ornementation de l’endite du L’ar- 3 5 et par maxillipède. hypogean cirolanids recorded from the island of Hispaniola. d’accumulation, Bahalana chipel des Bahamas est une aire et The is characterized its new genus mainly by pleon segmen- dans des C’est la a été trouvée eaux meso- et polyhalines. tation, the clearly prehensile first pereiopods and weakly dispersion, à partir d’ancêtres marins, qui semble être l’expli- prehensile second and third pereiopods, and the 2- and 5- l’évolution des Bahalana. cation la plus plausible pour segmented pedunculus of antennae 1 and 2, respectively.

Haitilana shows the greatest morphological resemblance to

Antrolana lira Bowman, 1964, from the Appalachian Valley,

U.S.A. The localities where Haitilana , was found,

have been flooded INTRODUCTION by the sea during the Cenozoic. Regres-

sion seems to be the most plausible explanation for the

evolution of Haitilana. During the Amsterdam Expedition to the West Another new stygobiont cirolanid, Bahalana cardiopus is , Indian Islands of 1979, stygobiont isopods of the described from a cave on Mayaguana, an island of the Baha- Cirolanidae have found several mas. This species is congeneric with Bahalana geracei Car- family been on

penter, 1981, from the island of San Salvador (Bahamas). islands. J. H. Stock and S. Weinberg found them distinctive of B. Clearly cardiopus are the unarmed endo- in two caves in the Bahamas, Lighthouse Cave podites of pleopods 3 to 5, and the armature of the maxilli- endite. Salvador and Mount Cave pedal The Bahama archipelago is an accumulation on San island, Misery Bahalana has been found in and area. meso- polyhaline waters. cirolanid of on Mayaguana. The Lighthouse Cave Dispersal (from marine ancestors) seems to be the most is described and has plausible explanation for the evolution of Bahalana. being by Carpenter (1981) been named Bahalana The cirolanid from geracei. Résumé Mount Misery Cave is a new species, likewise of Haitilana Un genre nouveau d’Isopodes Cirolanides, avec , the and will be genus Bahalana, described here as deux espèces nouvelles (H. radicicola et H. acanthura) sont B. décrits d’eaux souterraines de Haïti. Ce sont les premiers cardiopus.

Cirolanides hypogés découverts à être à Hispaniola. Le nou- L. Botosaneanu and the present author found

veau genre se caractérise surtout par la segmentation de son hypogean cirolanids at three different places on pléon, par ses premiers péréiopodes nettement préhensiles et Haiti: in karst used par des péréiopodes de la 2ème et 3ème paires faiblement a large spring as groundwater

ainsi les antennes 1 et 2 des préhensiles, que par ayant flagel- catchment and in two shallow wells. These are the

les de 2 et de 5 articles respectivement. Du point de vue first stygobiont cirolanids discovered in Hispan- morphologique, Haitilana ressemble le plus à Antrolana lira iola, the island of which Haiti forms the western Bowman, 1964, connue de l’Appalachian Valley, Virginia, U.S.A. localités Haitilana sub- Les où a été capturée ont été A new endemic is erected for part. genus them, la du c’est mergées par mer au cours Cénozoïque, et ce que with two new species: Haitilana radicicola and H. Stock appelle „Regression model” qui semble être l’explica- acanthura. tion la plus plausible pour l’évolution des Haitilana.

Cirolanide Un autre stygobie Bahalana cardio- to cirolanids nouveau, Up now, stygobiont were known

from Cuba, San Salvador island, Aruba, Mexico, *) 11 Report no. appeared in Bijdr. Dierk., 51 (1): 135-144 and the U.S.A. and (1981). (Texas Virginia) (see fig. 9). 314 J. NOTENBOOM - WEST INDIAN HYPOGEAN CIROLANIDS

from and Haitilana radicicola Presently, they are known Mayaguana n. sp. Figs. 1, 2, 3 a-e.

found around Haiti as well. Furthermore, they are Material. — Amsterdam Expeditions to the West Indian in eastern Africa the Mediterranean, (Somalia) Islands, Sta. 79/633, 1 S (holotype), 7 $ $ and 16 9 9

Haiti: Source Débarasse and in Madagascar. (paratypes). near Jérémie (approx- imate position 18°39'08"N 74°11'30"W), 1 Dec. 1979.

Zoölogisch Museum Amsterdam coll. no. Is. 105.164 a-b,

105.167 a-b. TAXONOMIC PART

Haitilana n. gen. — Males to maximum Description. range a

— Cirolanidae. Blind, unpig- of 6.3 females to 6.5 mm. rather Diagnosis. length mm, Body sclerotized blade, mented (except for masticatory wide, 2.8 times as long as wide (fig. la).

of maxilla 1 and the the spines on the exopodite tips All pereionites, except for first, with coxal

of the cannot coxal of the dactyls pereiopods). Body plates; posterior corners of plates 2 and 3

flat. roll into a ball; rather wide, Posterolateral rounded, those of 4 to 7 posteroventrally pointed

margins of head partly surrounded by pereionite 1. (figs, la, lm); lateral margin of pleonite 1 cover-

4 with Pleonites 1 to 5 distinct, pleonites 2 to ed by coxal plate of pereionite 7; pereionite 1 lateral angularly produced posterolateral margins; slightly longer than the others, pereionites 2 to 4

covered those of the size. Pleo- margins of pleonite 1 by pereio- and 5 to 7 mutually of about same

nite B of nites wide 7 (type Bowman, 1975). 1 to 3 almost as as pereionite 7 ; epimera

the anterior Antennae implanted near margin 2 to 4 posteroventrally pointed, elongately trian-

head. than the of the First antenna longer pe- gular (fig. lm). Dorsal surface of pleonite 1

duncle of the second; peduncle of the first antenna parly covered by pereionite 7. Lateral margins of

2-segmented, of the second antenna 5-segmented; pleonite 5 covered by epimera of pleonite 4.

first and second with human flagellum of the antennae Telson shaped like a tongue, its lateral

table numerous segments (see I). margins rounded, posterior margin with several

Mandibles and maxillae clearly cirolanid. Max- short setae and 4 small spines (fig. lb).

illipedal endopodite 5-segmented; endite with 2 Antenna 1 longer than the peduncle of antenna

and coupling spines several plumose setae. 2, reaching to the middle of pereionite 1 (fig. la).

Pereiopods increasing in length in posterior Peduncle 2-segmented, both segments long and

in with into direction; pereiopod 1 clearly, pereiopods 2 and 3 equal size; segment 1 a depression weakly prehensile; remaining pereiopods ambula- which fits peduncle segment 1 of antenna 2. tory, slender. Flagellum 10- to 12-segmented, first segment twice

Rami of pleopods 1 and 2 undivided; exopodites as long as the others; flagellum segments with 1 of pleopods 3 to 5 2-segmented; exopodites of or 2 aesthetes on ventral surface (figs, lc, Id).

pleopods 1 to 5 and endopodites of pleopods 1 and Antenna 2 reaching to the middle of pereionite

armed with 4 4 2 numerous plumose setae. Appendix (fig. la). Peduncle 5-segmented, segments masculina median and others. basal, implanted on the margin 5 about twice as long as the Flagellum of the its medial with about twice the 22- to 27- endopodite, margin pre- as long as peduncle, apical serrations. segmented, without aesthetes (fig. le). Uropodal protopodite wedge-shaped, exopodite Frontal lamina (fig. Il) well developed, lateral and endopodite subequal; exopodite oblong and margins concave, anterior end wider than poste- narrow, endopodite triangular. rior end.

Telson shaped like a human tongue, margins Clypeus and labrum (fig. ll) forming together rounded, caudal end an structure with armed. oval a slight, rounded concavity

on its posterior margin.

— Haitilana Mandibles Type species. radicicola n. sp. asymmetrical (fig. 3 a & b); left

mandible overlapping the right one. Palp slender,

— Haitilana 2 with distal of slender Etymology. (gender feminine), segment a group 11 to 16 from Haiti + which (Ciro)lana. spines (4 of are longer), segment 3 with BIJDRAGEN TOT DE DIERKUNDE, 51 (2) - 1981 315

1. Haitilana radicicola entire dorsal left and A2 Fig. n. sp., a-k, ￿ holotype: a, , (scale A, A1 right only partially dorsal first d, first distal second represented); b, telson, (B); c, antenna (C); antenna, part (D); e, antenna (C); f, first pereiopod (C); g, second pereiopod (C); h, third pereiopod (C); i, fourth pereiopod (C); j, fifth pereiopod (C); k, sixth pereiopod (C).

ventral and l-m, ￿ paratype: 1, head, (B); m, pereion pleon, lateral (B). 316 J. NOTENBOOM - WEST INDIAN HYPOGEAN CIROLANIDS

and 11 or 12 small marginal spines and 3 longer ter- Endopodite with 5 slender spines 3 or 4 minal spines. Incisor right with 9 or 10, left with plumose setae.

11 small spines. Lacinia mobilis elongated, armed Maxilliped (fig. 3d) with 5-segmented endo-

with with 13 to 17 small spines and with fine podite; segment 1 squarish, armed 1 setiform

with cilia. element; segment 2 trapezoidal, armed 6 or

Maxilla Distal of 7 medial and lateral setiform 1 (fig. 3c). armature exopo- 1 element; segment 3 dite consisting of 1 minute spinule and 10 strongly the largest, squarish, armed with 15 to 17 medial sclerotized spines; three innermost spines with 1 and 4 to 5 lateral setiform elements; segment 4

with den- with medial lateral setiform medial denticle, other spines 2 or more 10 to 13 and 2 to 3

with ticles. Endite armed with 3 spines, of which 1 is elements; segment 5 the smallest, about 20 longer than the two others. setiform elements. Endite armed with 5 plumose

Maxilla 2 (fig. 3e) with 4 and 8 to 10 slender setae (3 terminal) and 2 coupling hooks (reti- elements on palp and exopodite, respectively. nacula).

2. Haitilana radicicola first second second Fig. n. sp. a-g, ￿ holotype: a, pleopod (scale C); b, pleopod (C); c, pleopod,

masculina third fourth fifth apex appendix (D); d, pleopod (C); e, pleopod (C); f, pleopod (C); g, uropod (C). h, ￿ paratype: second pleopod (C). - 317 BIJDRAGEN TOT DE DIERKUNDE, 51 (2) 1981

4 with Pereiopod 1 (fig. If) clearly prehensile; basis Pleopods 3 and (figs. 2d, 2e) a large,

with and ischium scarcely armed; merus squarish, poste- 2-segmented exopodite; basal segment a with rior margin armed with 5 blunt spines and 1 lateral row of plumose setae; distal segment

with small little smal- setiform element; carpus 1 or 2 spines numerous plumose setae; endopodite a

4 setiform elements unarmed and ovoid. and ( 1 longer ) ; propodus ler,

from 3 and about twice as long as wide, posterior margin with Pleopod 5 (fig. 2f) differs pleopods

4 with seti- 4 of 3 or widely-spaced spines, distally 2 in having a lower density setae.

the and 4 such elements armed form elements on anterior, Uropodal protopodite wedge-shaped,

the with terminal with its laterodistal and with on posterior margin; dactylus 2 spines on margin,

with and 6 setiform its mediodistal claw, 2 spines a group of 8 to 10 plumose setae on margin;

little than elements. exopodite a shorter the endopodite,

with 6 and Pereiopod 2 (fig. lg) weakly prehensile, slightly oblong and narrow, armed spines

more slender than 1 basis pereiopod ; scarcely numerous plumose setae; endopodite triangular,

armed with 4 armed; ischium armed with a posteroterminal much wider than the exopodite,

of 2 and anteroterminal of and setae group an group 2 or 3 spines numerous plumose (fig. 2g).

spines; merus rectangular but with a concave Intraspecific variation. — A good this bears two posterior margin; margin groups, this number of individuals of new species was each of 3 and 3 anterodistal spines, spines; carpus present in the Source Débarasse sample (Sta. with 4 squarish, a posterodistal group of spines idea of the intra- 79/633), enabling us to get an and 1 setiform element; propodus more than twice of specific variation in a population a hypogean with as long as wide, posterior margin 3 widely- found in the cirolanid. Variation was mainly spaced spines; dactylus with 1 distal spine and number antennae and the mouthparts. In table I a 1 setiform element, in addition to the claw. of the and three counts based on holotype para- Pereiopod 3 weakly prehensile, similar to pereio- number types are enumerated. Variable are the pod 2 (fig. lh). and the of flagellum segments of the antennae Pereiopods 4 to 7 (figs, li, lj, Ik) increasing armature of the mandibular incisor and lacinia in length, slender, and ambulatory. mobilis. Pleopod 1 (fig. 2a): lateral margin of proto- Constant characters are: podite with 4 spines and 1 seta; exopodite and third — the number of terminal spines on the endopodite of equal length, armed with plumose mandible palp segment; setae; exopodite ovoid, with 1 subbasal spine;

—- number the endite the of spines on of max- lateral with endopodite rectangular, margin a row illa 1; of fine cilia.

— the number of slender spines on the palp of Pleopod 2 ( c5 ) (fig. 2b): lateral margin of maxilla 2; protopodite armed with 4 spines and 3 or 4

•—- the the number of plumose setae on endite, plumose setae; exopodite and endopodite of equal and the the first number of setiform spines on length, armed with numerous plumose setae, both endopodite segment of the maxilliped; inserted ovoid; appendix masculina laterally on •—■ the number of spines on the uropodal proto-, the base of the endopodite, and longer than the endo-, and exopodite; latter, straight, its lateral margin with a proximal

— the of 4 small the presence spines on posterior of its medial with row short cilia, margin minute, margin of the telson. blunt, preapical serrations (fig. 2c).

lateral of — Pleopod 2 (?) (fig. 2h): margin Etymology. The proposed specific name,

with 4 and 3 radicicola is of the Latin words incola protopodite spines plumose setae; , composed exopodite and endopodite of equal length, armed (inhabitant) and radix (root), alluding to the with where plumose setae; exopodite ovoid, endopodite place the have been found, under

in of rectangular (as uropod 1); lateral margin of a blanket Ficus roots in a large karst spring

with used endopodite a row of fine cilia. as groundwater catchment. - WEST INDIAN HYPOGEAN CIROLANIDS 318 J. NOTENBOOM

Haitilana left mandible first maxilla Fig. 3. a-e, radicicola n. sp., ￿ holotype: a, mandible (scale F); b, right (F); c,

(F); d, maxilliped (F); e, second maxilla (F).

mandible maxilla second f-i, Haitilana acanthura n. sp., ￿ holotype: f, right (F); g, first (F); h, maxilliped (F); i, maxilla (F). BIJDRAGEN TOT DE DIERKUNDE, 51 (2) - 1981 319

TABLE I

Variation range in certain characters of four individuals of Hait ilana radicicola from

“Source Débarasse”.

Characteristics holotype paratypes S $ 9(a) 9(b)

antennae

numb. 12 12 11 10 segm. flagellum Al

A2 numb. segm. flagellum 27 25 22 23

mandibleright

numb, spines incisor 10 8 8 numb, spines lac. mob. 13 16

numb. dist. 2 17 16 spines palp segm. ca. 12

numb. term, spines palp segm. 3 3 3 3

numb. marg. spines palp segm. 3 11

mandibleleft

numb, spines incisor 11 8 8

numb, spines lac. mob. 15 17 17

numb. dist. 14 spines palp segm. 2 12

numb. 3 3 3 term, spines palp segm.

numb. marg. spines palp segm. 3 ca. 10

maxilla 1

numb, spines endite 3 3 3 3

numb, spines exopodite 10 10 11 10

maxilla 2 right

numb, spines palp 4 4 4 4

numb, spines exopodite 8 8 9

numb, spines endopodite (longer) 9(4) 9(4) 9(4) 8(4)

maxilliped

numb. plum, setae endite 5 5 5 5

numb, spines endop. segm. 1 1 1 1 1

uropod

numb, spines protopodite 2 2 2 2

numb, spines endopodite 4 4 4 4

numb, spines exopodite 6 6 6 6

telson

numb, little spines post, margin 4 4 4 4

Habitat. — Haitilana radicicola found in it has C was is ca. 0.1 m deep, a temperature of 24.7°

which and a large karstic spring (Source Débarasse) a chlorinity of 20.6 mg/1. Ficus roots enter arises from foot of a middle terrace at the a higher the diaclase through the roof, and partially cover

km from the walls terrace (Morne Débarasse), ca. 2.5 and floor. The cirolanids were found in the

Caribbean Sea. This locality is situated some 2 km water between the Ficus roots and the floor; the

from and the airstrip of Jérémie. These middle and water running over through the roots har-

terraces of Eocene and Pa- boured higher are composed many Hyalella (Amphipoda), Gastropoda,

leocene limestones. Since 1951, the spring is used and Elmidae (Coleóptera). One specimen of the

catchment for the drinkwater found as a water supply isopod was also under a stone outside the

small of the town of Jérémie. The water catch- water catchment, in a small auxiliary spring.

entered ment can be through a metal gate; first

is Haitilana acanthura n. 3 f-i, 4, 5. there a small room (cut out in the rock) with sp. Figs.

several which — concrete tanks, are fed by fast run- Material. Amsterdam Expeditions to the West Indian Islands, Sta. 79/560, 1 9 (holotype) and 1$ (paratype). ning water flowing through a partially man-made Haiti: Marigot, well of Jacques Simein (approximate position this cleft of some 10 m long; cleft, or diaclase, 18°13'51"N 72°18'52"W), 16 Nov. 1979. Zoölogisch Mu- is 0.4 0.6 and is to m high totally dark. The water seum Amsterdam coll. no. Is. 105.163 a-b, 105.166 a-b. 320 J. NOTENBOOM - WEST INDIAN HYPOGEAN CIROLANIDS

— Both Description. specimens are 7 mm and 5 about twice as long as the others. Flagellum

long. The body is 2.3 times as long as wide. Pereio- about 19-segmented, without aesthetes.

nites similar to those of the former species. Pleo- Frontal lamina well developed, rectangular,

anterior little wider nites 1 to 3 almost as wide as pereionite 7. Epimera end a than posterior end.

4 and 1 to 3 posteroventrally pointed; epimere obtuse, Clypeus labrum form together an oval

its lateral margins partially covered by epimere 3. structure with a slight rounded concavity at the

Pleonite 1 partially covered by pereionite 7; lateral posterior margin.

margins of pleonite 5 covered by those of pleo- Mandibles (fig. 3f) asymmetrical; left mandible

the nite 4. overlapping right one. Palp slender, segment

Telson human 2 with distal of about 16 which (fig. 4e) shaped as a tongue, a group setae (of

lateral margins rounded, posterior margin with 4 are longer), segment 3 with about 9 smal mar-

plumose setules and 10 to 12 small spines. ginal setae and 2 longer terminal setae. Incisor

Antenna 1 (fig. 4b) longer than the peduncle right and left with 7 small spines. Lacinia mobilis

armed with of antenna 2, reaching beyond the middle of elongate, 15 (right) or 18 (left)

both small anterior with pereionite 1; peduncle 2-segmented, seg- spines, margin basally some

than ments elongate, segment 2 a little longer cilia.

with Maxilla with segment 1; peduncle segment 1 a depression 1 (fig. 3 g): exopodite distally

into which fits peduncle segment 1 of antenna 2. 12 sclerotized spines, one of which is slender; the

Flagellum 9-segmented, first two segments of innermost spine has a small medial denticle, spines

4 unarmed. equal length; segments 3 to 9 with aesthetes on 2 to are Endite with 3 distal elements.

with with and the ventral surface (segments 5 to 8 2 Maxilla 2 (fig. 3i) 4 10 setae on palp

aesthetes, the others with one). and exopodite, respectively; endopodite with 5

Antenna 2 (fig. 4a) reaching to the middle of naked and 4 plumose setae.

pereionite 5; peduncle 5-segmented, segments 4

4. Haitilana acanthura n. basal of second Fig. sp., ￿ holotype: a, part antenna (scale C); b, first antenna (C); c, first

second dorsal pereiopod (C); d, pereiopod (C); e, telson, (B). BIJDRAGEN TOT DE DIERKUNDE, 51 (2) - 1981 321

Maxilliped (fig. 3h) with 5-segmented endo- Pleopod 2 (fig. 5b): protopodite with 3 spines

with setiform and and podite; segment 1 squarish 1 spine; 2 setae on lateral margin; exo- endopodite

segment 2 triangular, armed with 1 lateral and 3 similar to those of pleopod 1.

with to 5 medial setiform elements; segment 3 the Pleopods 3 and 4 (figs. 5c, 5d) large,

with largest, squarish, armed with 4 or 5 lateral and 6 2-segmented exopodite, basal segment a

medial with lateral distal with to 11 setiform elements; segment 4 row of plumose setae, segment

all around its 3 lateral and 6 to 9 medial setiform elements; numerous plumose setae, margin;

than segment 5 the smallest with 9 to 12 setiform endopodite a little smaller exopodite, ovoid,

spines. The armature of the endite is different in practically unarmed. the two specimens available: the holotype has 3 Pleopod 5 (fig. 5e) differing from pleopods

terminal plumose setae on the left side, 3 plumose 3 and 4 in having less setae.

setae (2 terminal) on the right side, and on both Uropodal protopodite wedge-shaped (fig. 5f),

armed with its and sides 2 coupling hooks; the paratype has 3 plumose 3 spines on laterodistal corner,

and with 6 its mediodistal setae (2 terminal) 2 coupling spines on the plumose setae on margin;

left side, and 4 plumose setae (3 terminal) and exopodite less long and less wide than the endo- and armed with 3 coupling spines on the right side. podite, oblong narrow, 7 spines

and Pereiopod 1 (fig. 4c) clearly prehensile; basis (5 lateral, 2 medial), numerous plumose

ischium less armed and scarcely armed; merus squarish, poste- setae; endopodite more or triangular,

rior with of small with 6 and margin a subbasal group 3 spines numerous plumose setae.

and distal seti- spines, a group of 1 spine and 1

form with of 4 element; carpus 1 group spines Distinction. — This species has a signifi- about twice (2 smaller); propodus as long as of the cantly larger number spines on posterior wide, posterior margin with 3 widely spaced border of the telson, and shows certain other dif- spines, 4 distal setiform elements, and 2 antero- ferences as well, compared to H. radicicola (see distal setiform elements; dactylus distally with a table II). 2 and anterodistal of seti- claw, spines an group 3

form elements.

— The Etymology. proposed specific name, Pereiopod 2 (fig. 4d) weakly prehensile, acanthura is composed of the Greek words axavöa strongly built; basis scarcely armed; posterior mar-

= and = the spine, oüpá tail, alluding to arma- gin of ischium with 1 + 1+2 spines, antero-

ture of the telson, an difference between distal with and important corner 1 spine 1 setule; meras H. acanthura and H. radicicola. longer than wide, posterior margin with 4 isolated

and distal of and spines a group 3 spines a setule,

anterodistal corner with 3 Habitat. — spines; carpus squarish, Haitilana acanthura was caught by

with of and of posterodistal group 3 spines 1 seta; means a Cvetkov net in the well of Jacques than twice propodus more as long as wide, poste- Simein, near the East end of the village of Mari-

rior with 2 isolated and distal the is margin spines a got, at some 300 m from seashore. The well

and distal end situated group of 1 spine 1 seta; of dactylus on the lower terrace, dating from Pleisto-

with 1 and the robust spine some setules; ungulus cene age; adjacent middle terrace is of Eocene/

rather slender. Paleocene age. The lower terrace is mainly com-

built river Pereiopod 3 as pereiopod 2. posed of sediments (gravel, sand and clay).

Pereiopods 4 to 7 increasing in length, slender The well, 5 m deep, is regularly cleaned. The and ambulatory. water, 1 m deep, has a temperature of 25° C and

with median of 30.8 Pleopod 1 (fig. 5a) : protopodite a a chlorinity mg/1.

of 4 and of In addition the the well houses group spines; exo- endopodite equal to cirolanids, length, armed with plumose setae; exopodite ovoid, Typhlatya (Decapoda), Amphipoda, Cyclopoida armed and with a subbasal spine; endopodite rectan- (Copepoda), Ostracoda. gular. 322 J. NOTENBOOM - WEST INDIAN HYPOGEAN CIROLANIDS

Haitilana acanthura first second third (C); d, Fig. 5. n. sp., ￿ holotype: a, pleopod (scale C); b, pleopod (C); c, pleopod

fourth pleopod (C); e. fifth pleopod (C); f, uropod (C). - 1981 323 BIJDRAGEN TOT DE DIERKUNDE, 51 (2)

TABLE II

Comparison of meristic characteristics in some specimens of three isolated populations of Haitilana.

Characteristics H. radicicola H. acanthura H. sp. (24 specimens) (2 specimens) (2 specimens)

body

length (mm) to 6.5 to 7 6 and 7.5

length/widthratio 2.8 2.3 3.3

antenna 1

numb. segm. flagellum 10-12 9 8-9

ratio 1.5 length peduncle segm. 3 to 2 1 1.5

length ratio peduncle segm. 1 to 2 2 1 1-2

antenna 2

20 numb. segm. flagellum 22-27 19-25

mandible right

numb, spines incisor 8-10 12 10

numb, spines lac. mob. 13-16 15 15

numb. term, spines palp segm. 3 3 2 I

mandible left

numb, spines incisor 8-11 10 10 numb, spines lac. mob. 15-17 18 17

numb. 3 3 2 1 term, spines palp segm.

maxilla 1

numb, spines endite 3 3 3

numb, spines exopodite 10-11 12 II

maxilla 2

numb, spines palp 4 4 4

numb, spines exopodite 8 9 10

numb, spines endopodite (longer) 9(4) 9(4) 10(3)

maxilliped numb. plum, setae endite 5 3 2

numb, spines endopodite:

segment 1 1 0 1

segment 5 16-20 13 12

uropod numb, spines protopodite 2 + 0 2+1 2 + 1

numb, spines endopodite 4 6 6

numb, spines exopodite 6 7 7

telson

numb, little spines post, margin 4 10-12 10

Haitilana used for this are enumerated. The characters sp. study

found to be constant in H. radicicola table I) Material. —- Amsterdam Expeditions to the West Indian (cf.

Islands, Sta. 79/578, 2 9 9. Haiti: Dumonet (between Débas are considered as the most important ones for and Caiman, west of Thomazeau), well of Mme. Tissé-Corio- taxonomie purposes. lan (approximate position: 18°38'58"N 72°06'58"W), 21

Amsterdam coll. Haitilana resembles H. acanthura on account Nov. 1979. Zoölogisch Museum no. Is. sp.

105.165 a-b. of the armature of the uropods and the telson.

Characters for Haitilana the ■— the avail- are Remarks. Unfortunately, one of unique sp. length/

of them width ratio of the the number of terminal able specimens was damaged (both were body,

number dead when caught). It is hard to decide whether spines on mandible palp segment 3, the

of elements maxillar or not these specimens belong to H. acanthura, on the second endopodite,

which and of the to they resemble most. In table II a number the number spines on maxillipedal of characteristics of the three samples of Haitilana endite (cf. table II). 324 J. NOTENBOOM - WEST INDIAN HYPOGEAN CIROLANIDS

between Habitat. — The present two specimens were As point of fact, the differences the

in the well of first and Haitilana caught by means of a Cvetkov net pereiopods of Troglocirolana

small. Mme. Tissé-Coriolan at Dumonet, in the lowland are Clearly different are the ovate body, the

4 and plain, called Cul de Sac. This plain was still pereionites 5 (which are the longest), the

flooded the the The and telson with by sea during Pleistocene. broad rounded a crenulate lateral

the well is 6 m deep and dug out in sand. The water margin, the 6-segmented peduncle of second

is of C and and the of the second maxilla 1 m deep, has a temperature 27.2° a antenna, palp pro-

422 well lies with chlorinity of mg/1. The some 30 m vided 5 spines. Despite these differences,

level and is Haitilana. above the sea situated 200 m from Troglocirolana appears to be related to

the Trou Caiman, a shallow lake. In addition to

the cirolanids, the sample contained Amphipoda,

Monodella (Thermosbaenacea), Cyclopoida (Co- Genus Bahalana Carpenter, 1981.

pepoda), Ostracoda, Phyllopoda, and Gastro- Thanks to the courtesy of Dr. J. H. Carpenter, I poda. the have of his had privilege to a preview manu-

in the script press, containing formal description

RELATIONSHIPS OF HAITILANA of this genus. The type-species, B. geracei, and so

far col- unique representative of Bahalana, was The combination occurring in Haitilana of a pleon lected in Lighthouse Cave, San Salvador island, segmentation of "type B" (Bowman, 1975) and Bahamas. The collections of the Amsterdam Ex- of clearly prehensile pereiopods 1, has been found the West Indian peditions to Islands contained a to now in the cirolanid Antrolana up only genera number of specimens of B. geracei taken at the (Appalachian Valley, Virginia) and Typhlociro- type-locality. lana (Mediterranean belt). Rioja (1957) has

described the from genus Troglocirolana Cuba, Bahalana characterized the geracei 1981. by same pleon segmentation as Carpenter,

with Haitilana, but semi-prehensile first pereio- — Material. Amsterdam Expeditions to the West Indian

5 pods. Islands, Sta. 79/186, 9 9,1 $ (1 9 and 1 $ are dam-

aged). Bahamas: San Salvador island, Dixon Hill Lighthouse Of the Antrolana is genus only one species Cave (approximate position 24°05'44"N 74°27'07"W), 24 A. lira In known, Bowman, 1964. opinion, Nov. Amsterdam my 1979- Zoölogisch Museum coll. no. Is. a-b. Antrolana appears to be the closest relative of 105.186

Haitilana. differs in its However, it clearly greater

Remarks. — The four undamaged females are body length (up to 12 mm), the produced ante-

rior 6, 12, 12 and 14 mm long. Their length/ margin of the head, the 6-segmented peduncle avarage width ratio is the 2.3. recorded a of the second antenna, narrow frontal lamina, Carpenter length/ ratio the width of 3 for his specimens. In general, the armature, with 7 spines, of the palp of the specimens examined from the West Indian Islands second maxilla, the hooked appendix masculina, conform the seta and Expedition description of B. bearing a on the apex, the less wedge- geracei. shaped uropodal protopodites.

has Typhlocirolana a number of characters Bahalana by cardiopus n. sp. Figs. 6, 7, 8. which it the clearly differs from Haitila na: body

Material. — Amsterdam Expeditions to the West Indian is to 15 length up mm, the body shape (narrow, Islands, Sta. 79/143, 1 9 (holotype) and 2 9 9 (paratypes, and with parallel margins rather loosely articu- one of which smaller and damaged). Bahamas: Mayaguana, Little Bay, Mount Misery Cave (approximate position lated somites, more rounded in cross-sec- being o 22°25'49"N 73 02'34"W), 12 Nov. 1979. Zoölogisch Mu- the of the tion), short, 3-segmented peduncle Amsterdam coll. seum no. Is. 105.161 a-b, 105.162 a-b. first antenna, the 6-segmented peduncle of the second the antenna, produced frontal lamina, and Description. — Holotype and undamaged the first the other longer pleonite, overreaching paratype both 10 mm long; body about 2.5 times pleonites. as long as wide. Unable to roll into a ball. Head - 325 BIJDRAGEN TOT DE DIERKUNDE, 51 (2) 1981

left Fig. 6. 6. Bahalana cardiopus n. sp., ￿ holotype: a, entire animal, dorsal (scale G, A1 and right A2 not shown); b, head,

ventral dorsal and lateral first second (H); c, telson, (H); d, pereion pleon, (H); e, antenna (I); f, peduncle antenna (I).

wider than long, anterior margin straight. All short; segments 1 and 2 with a depression into

which and pereion somites, except the first, with coxal plates; fit segments 1 2 of antenna 2; segments

of coxal and 4 bear distal sensorial posterior corner plates 2 3 rounded, 3 and a few setae. Flagel- those of pereionite 4 rectangular, those of pereio- lum 19- to 22-segmented; segment 1 twice as long nites 5 to 7 pointed (fig. 6d). Pereionite 1 slightly as segment 2 (fig. 6e). longer than the other pereionites, pereionites 4 to Antenna 2 reaching the posterior margin of

in 2 and 7 approximately equal length, pereionites pereionite 7; peduncle 5-segmented; segment 1

3 intermediate (fig. 6a). Pleonites 1 to 5 distinct, with an anteromedial knob; segments 1 to 3 short,

4 and each with angularly produced lateral margins, those of subequal, segments 5 about as long pleonite 5 slightly produced. as segments 1 to 3 combined; segment 5 with

Telson squarish (fig. 6c), a little wider than about 5 sensorial setae. Flagellum of about 32 seg- long, lateral margins rounded, posterior margin ments. with Frontal lamina about 50 crenulations; in each notch a setule (fig. 6b) a poorly developed, is inserted. Lateral margin unarmed. narrow carina, posterior end wider, triangular, and

Antenna 1 longer than the peduncle of antenna partly covered by the clypeus.

2, reaching to the posterior margin of pereionite Posterior margin of labrum with a rounded

2 (fig. 6a); peduncle 4-segmented, segments 1 concavity (fig. 6b). and 2 small, segment 3 a little less than twice as Mandibles asymmetrical (fig. 7a), left man-

as 1 and 2 4 dible the one. 2 long segments together, segment very overlapping right Palp segment 326 J. NOTENBOOM - WEST INDIAN HYPOGEAN CIROLANIDS

Fig. 7. Bahalana cardiopus n. sp., ￿ holotype: a, right mandible (scale K); b, second maxilla (K); c, maxilliped (K) ;

d, first maxilla first second third fourth fifth (K); e, pleopod (I); f, pleopod (I); g, pleopod (I); h, pleopod (I); i, pleopod (I); j, uropod (I). BIJDRAGEN TOT DE DIERKUNDE, 51 (2) - 1981 327

with of with 6 and inner about twice as long as segment 1 a row 3 to distal 5 to 9 spines; pro-

about 18 setiform elements in its distal half; seg- podus with 3 distal and 5 to 9 inner spines; dacty-

lus about of the with 1 terminal ment 3 slightly less than half as long as segment 2, 1/4 propodus,

and terminal of the armed with 15 small marginal spines 3 ones. spine; ungulus about 1/3 dactylus,

Incisor with about 10 spines; lacinia mobilis with 3 to 4 spinules of varying sizes.

with triangular, with about 12 small spines. Pleopod 1 (fig. 7e): Protopodite a me-

dian of about Maxilla 1 (fig. 7d) with 11 sclerotized spines row about 10 spines; exopodite

third inner wide and the distal on the exopodite, first and spines with twice as as long as endopodite;

second inner of endo- and with of 1 medial denticle, spine unarmed; margins exopodite a row

setule the 4 spines and a on endite. plumose setae; median margin of endopodite

with Maxilla 2 (fig. 7b) armed with 2 and 4 seti- straight, a row of fine cilia.

with form elements on palp and exopodite, respectively; Pleopod 2 (fig. If) : Protopodite a median

with 4 endopodite distally 8 naked and 2 plumose row of 8 spines and short plumose setae; endo-

setae, basally with 1 setiform element. podite slightly wider than in pleopod 1.

3 and 4 Maxilliped (fig. 7c) : Endopodite 5-segmented; Pleopods (figs. 7g, 7h): Protopodite

with and segment 1 squarish, armed with 1 setiform ele- medially 5 to 7 spines 4 to 5 short

with lateral and with ment; segment 2 trapezoidal 1 5 plumose setae; exopodite partial transverse

the with to 7 medial setae; segment 3 largest, squarish, suture, basal segment 2 or 3 lateral setae, distal with armed with 4 lateral and 11 to 16 medial setae; segment numerous plumose setae;

segment 4 longer than wide, armed with 3 lateral endopodite smaller than exopodite, ovoid, un-

and 8 or 9 medial setae; segment 5 linear, bearing armed.

about 14 setae. Endite armed with one coupling Pleopod 5 (fig. 7i): Exopodite heart-shaped,

spine and 3 plumose setae (one shorter). with transverse suture, basal segment with about

and lateral distal Pereiopods 1 to 3 prehensile, dactylus pro- 2 setae, segment sparsely armed with

both about 20 in podus very elongate. plumose setae, placed two groups; Inner smaller than Pereiopod 1 (fig. 8a): side of merus endopodite ovoid, unarmed, the exo-

that reaches produced into a slender projection to podite.

of the this is armed 3/4 propodus; projection on Uropodal protopodite strongly produced into an

its posterior margin with 5 small, subdistal spines; acute mediodistal projection, bearing 1 apical seta;

small. short carpus very laterodistally with 2 spines; exopodite about

Pereiopod 2 (fig. 8b): Basis scarcely armed; half as wide as the endopodite, medial margin

side ischium with outer of with a distal projection plumose setae, apically with a tuft of 4 setae

which extends about the side and 2 short halfway merus; outer spines, lateral margin with 4 short of with which little than merus a long, falcate projection spines; endopodite a longer the exo- extends to the base of the dactylus, inner margin podite, medial margin with plumose setae, apically

small and with 2 spines a knob tipped with a with some 4 short spines and a few naked setae,

inner of the with distal lateral dorsal surface of setule; margin carpus a margin unarmed; endo-

projection which extends to about the middle of podite with 2 sensorial setae (fig. 7j).

the propodus.

Pereiopod 3 (fig. 8c) : Projection on ischium as

in P2, inner than in Distinction. — In the spine longer P2; projection on diagnosis of the genus

side of in mentions outer merus as P2, projection on inner Bahalana Carpenter (1981) the , pres-

side half as long as the carpus; inner projection ence of setation on the endopodites of pleopods

the half the this setation in B. On on carpus overreaching propodus. 3 to 5; is absent cardiopus.

Pereiopods 4 to 7 similar (figs. 8 d-g), ambula- account of the correspondence in body shape, in

tory and longer than pereiopods 1 to 3; ischium pleon segmentation, in the curiously shaped pereio-

with 2 to 4 distal and 2 or 3 inner spines; merus pods 1 to 3, in the mouthparts, and in the pedun-

with 3 to 6 distal and 2 to 3 inner culus of the it spines; carpus antennae, appears justifiable to 328 J. NOTENBOOM - WEST INDIAN HYPOGEAN CIROLANIDS

first second third Fig. 8. Bahalana cardiopus n. sp., ￿ holotype: a, pereiopod (scale I); b, pereiopod (I); c, pereiopod (I); fourth detail fifth detail f, sixth detail seventh d, pereiopod (I), (K); e, pereiopod (I), (K); pereiopod (I), (K); g, pereiopod (I), detail (K). BIJDRAGEN TOT DE DIERKUNDE, 51 (2) - 1981 329

B. with B. the of B. anchi- consider cardiopus congeneric geracei, Cave, habitat cardiopus, may be

the of Bahalana. haline since of influence type-species as well, the presence tidal

B. cardiopus is distinguished not only by the cannot be excluded. During the Amsterdam Expe-

of the of dition the West Indian Islands of H. lack armature on endopodities pleopods to 1979, J.

3 to 5, but also by the shape of the pleopodal Stock and S. Weinberg took water samples in both

endo- and exopodites, the armature of the maxilli- the Lighthouse Cave (23 November 1979) and pedal endite, and the shape and armature of the Mount Misery Cave (12 November 1979), the

mandibular lacinia mobilis. former had of 21264 the a chlorinity mg/1, latter

of 9682 mg/1, in both cases much too high to

— The classification "inland Etymology. specific name, cardiopus, allow as waters".

is composed of the Greek words xapSia ( = heart)

and 7roùç (= leg), alluding to the cordiform

exopodite on the fifth pleopod. This character ZOOGEOGRAPHIC REMARKS

forms a clear distinction between B. cardiopus and

B. geracei. Most cirolanid isopods are marine, but some are

found in hypogean habitats. It is assumed that the

latter evolved from marine Colonization Habitat. — Mayaguana is composed of reef ancestors.

occurred limestones and debris from the Cretaceous period of hypogean habitats may possibly have

and In the late of marine younger. Tertiary or Quaternary, by stranding populations (regression)

this part of the Bahama archipelago has raised or by dispersal (Stock, 1977, 1980).

A of characteristics in above sea level. The type-locality, Mount Misery number occur stygobiont

is cirolanids in different of The Cave, a small cave, located some 600 m from stages apomorphy.

the seaboard and 10 level. The characteristics involved touch the ca. m above sea pleon segmen-

the is tation, the setation and entrance of cave situated in the vertical part segmentation of the pleo- of the construction of the and a cliff at the inland side of a high ridge, and pods, pereiopods, the

of leads to a slightly down-sloping gallery. The ciro- number peduncle segments in the second an-

found lanids were in a muddy hole filled with tenna. Racovitza (1912) already pointed out cer-

water the water dark- tain of these trends. Also within dif- (probably table), in total one genus,

Tidal influence is to be ferent characters show different of ness. presumed present. stages apo-

The water had a chlorinity of 9682 mg/1. No morphy. The origin of this mosaic pattern could

fauna rest differences ancestral accompanying was observed. upon among populations,

but also successive upon joint or action of dispersal

and These regression. two processes could, more- RELATIONSHIPS OF BAHALANA over, have been repeated in different geological periods.

As of the other Carpenter (1981) notes, none Regression seems to form a plausible explana-

cirolanid to be related tion for genera appears closely to the origin of Haitilana. The localities

Bahalana. The is because where members of this have been genus especially unique genus found lie of the construction of the first three level and far pereiopods. slightly above sea not from the

The of B. in has seaside. the Cenozoic discovery cardiopus Mayaguana During these localities were

this inundated the not changed morphological notion. by sea (Geol. Map Haiti, 1972).

Mayaguana is separated from the type-locality The Bahama archipelago is, in contrast with the of B. San of West geracei, Salvador island, by deep water rest the Indies, an accumulation area. In

(>200 m). The habitat of B. geracei, in the geologically fairly recent times, the low islands

is anchihaline have Lighthouse Cave, considered by come above sea level as a result of conglo-

he be and reef Carpenter (1981); supposes the species to meration accumulation of debris due to

link in the evolution of wind and the wash the an ecological missing of waves (Doran, 1955).

hypogean cirolanids. The waters of Mount Misery The groundwater of these small and low islands 330 J. NOTENBOOM - WEST INDIAN HYPOGEAN CIROLANIDS

Fig. 9. Distribution of troglobiont cirolanid genera in the New World. 1, Haitilana; 2, Bahalana; 3, Antrolana; 4, Cirolani- des 5, Troglocirolana; 6, Haptolana; 7, Arubolana; 8, Speocirolana; 9, Conilera (= Speocirolana ?); 10, Sphaerolana; 11,

Mexilana; 12, Creaseriella.

Sea Cretaceous has connections with the sea through fissures and or the Caribbean during the late dissolution effects, the latter possibly originating or Cenozoic. Antrolana or its direct ancestor could from sea level drops during the Glacials (cf. have reached its present distribution area by north-

Harmon et al., 1981). Bahalana has been found ward dispersal through subterranean pathways. in meso- and polyhaline (maybe anchihaline) However, such a dispersal is hard to accept be-

almost groundwaters of two Bahamian islands (San Sal- cause the geological barriers were (and are) vador and Mayaguana). Dispersal, from some unsurmountable (Bowman, 1964). marine to be stock, appears a plausible explana- tion for the origin of Bahalana.

Since Antrolana lira has found in the been ACKNOWLEDGEMENTS

Appalachian Valley in Virginia, it forms a unique I like thank the would to numberless, mostly anonymous, the cirolanids New case among stygobiont of the for guides on Haiti and several other West Indian islands

World. This area has not been in connection with the help they had given to find the different hypogean habi- the Permian the Consulate-General of the Netherlands, the sea since the (Bowman, 1964). On tats; Coopéra- tion Haitiano-Néerlandaise (COHAN), the Curaçao Trading account of morphological resemblances, there is Company de Haiti SA., all at Port-au-Prince, Haiti, and the some affinity between Haitilana and Antrolana. clerical authorities on Haiti for assistance in various ways;

This it is Stock, and for discussions affinity, if not based on parallelism, J. H. L. Botosaneanu S. Weinberg and the pleasant collaboration during the fieldwork; and possibly reflects a common ancestor. This common Susan Parren-Gardner for reviewing the manuscript. ancestor have lived in the Gulf of Mexico fieldwork been may The has supported by grants to J. H. Stock, BIJDRAGEN TOT DE DIERKUNDE, 51 (2) - 1981 331

L. Botosaneanu and S. Weinberg of the Netherlands Founda- COLE, G. A. & W. L. MINCKLEY, 1966. Speocirolana ther- for Advancement of tion the Tropical Research (WOTRO), midronis, a new species of cirolanid isopod

The Hague and to myself of the Landbouw Hogeschool from central Coahuila, México. Tulane Stud. Zool., 13:

Fonds, Wageningen and the Beyerinck-Popping Fonds, Am- 17-22, figs. 1-21. sterdam. & 1970. of cirolanid —, Sphaerolana, a new genus isopod from northern Mexico, with description of two

Southwest. new species. Natural., 15 (1): 71-81, figs.

1-40, 1 table. REFERENCES DORAN, E., 1955. Landforms of the southeast Bahamas. Univ

Texas Publ., 5509: 1-38. BOLIVAR Y PIELTAIN, C, 1950. Estudio de una Cirolana GEOLOGICAL MAP HAITI, 1972. Sécretariat général, Organi- cavernícola nueva de la región de Valles, San Luis sation des Etats Américains. Haïti, Mission d'Assistance Potosí, Mexico. Ciencia (Mexico), 10 (11-12): 211-218. technique intégrée: 1 sheet. (Washington). BOTOSANEANU, L. & J. H. STOCK, 1979. Amsterdam Expedi- HARMON, R. S., L. S. LAND, R. M. MITTERER, P. GARRETT, tions to the West Indian Islands, Report 6. Arubolana H. P. SCHWARCZ & G. J. LARSON, 1981. Bermuda sea imula first n. gen., n. sp., the hypogean cirolanid isopod level during the last interglacial. Nature (London), crustacean found in the Lesser Antilles. Bijdr. Dierk., 28S (5797): 481-483. 49 (2): 227-233, figs. 1-25. RACOVITZA, E.-G., 1912. Cirolanides (première série). Archs. T. Antrolana and BOWMAN, E., 1964. lira, a new genus Zool. exp. gén., (5) 10: 203-329, figs, i-viii, pis. XV- species of troglobitic cirolanid isopod from Madison XXVIII. Cave, Virginia. Int. J. Speleol., 1 (1/2): 229-236, pis. A the of RICHARDSON, H., 1905. monograph on isopods L-LVII. North America. Bull. U.S. natn. Mus., 54: 1-727, figs.

1966. a new and , Haptolana trichostoma, genus species 740.

of troglobitic cirolanid isopod from Cuba. Int. J. Spe- RIOJA, E., 1957. Estudos carcinologicos, XXXV. Datos sobre leol., 2: 105-108, 24-27. pis. algunos isópodos cavernícolos de la isla de Cuba. An.

1975. A new genus and of troglobitic ciro- , species Inst. Biol., México, 27 (2): 437-462. lanid isopod from San Luis Potosí, Mexico. Occ. Pap. STOCK, J. H., 1977. The and zoogeography of the Mus. Texas Tech Univ., 27: 1-7, figs. 1-4. hadziid Amphipoda, with emphasis on the West Indian

Bahalana CARPENTER, J. H., 1981. geracei n. gen., n. sp., a taxa. Stud. Fauna Curaçao, 177: 1-130, figs. 1-54. troglobitic marine cirolanid from Lighthouse model evolution isopod , 1980. Regression as exemplified by

Cave, San Salvador Island, Bahamas. Dierk. 51 the Bijdr. genus Pseudoniphargus (Amphipoda). Bijdr. Dierk.,

(2): 259-267. 50 (1): 105-144.

Received: 9 February 1981