Bijdragen tot de Dierkunde, 54 (1): 25-50 — 1984

Amsterdam Expeditions to the West Indian Islands, Report 38.

The distribution and zoogeography of freshwater

Ostracoda (Crustacea) in the West Indies,

with emphasis on species inhabiting wells

by

Nico W. Broodbakker

Institute of Taxonomie Zoology, University of Amsterdam,

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

Summary Résumé

The distribution of Ostracoda in islands of discute freshwater the On la distribution des Ostracodes dulcicoles sur les the West Indies and part of the Venezuelan mainland is îles des Indes Occidentales et dans certaines zones de discussed. The ostracod fauna of wells and epigean Venezuela. La faune d’Ostracodes des puits et des habitats habitats Some Certaines des is compared. species, e.g. Cypretta spp. are épigés est comparée. espèces (par exemple found often in others significantly more wells, are only espèces de Cypretta) sont trouvées nettement plus souvent found in and dans des d’autres epigean habitats, e.g. Hemicypris spp. Cypris puits, (par exemples Hemicypris spp. et subglobosa. Some species are found significantly more often Cypris subglobosa) sont rencontrées seulement dans des in others to habitats il epigean habitats, e.g. Stenocypris major, seem épigés. Ensuite, y a des espèces nettement plus have and trouvées dans les habitats no preference, e.g. Physocypria affinis Cypridopsis souvent épigés (Stenocypris major

tandis spp. par exemple), que d’autres (exemples: Physocypria

occurrence of Ostracoda and other faunal semblent avoir de Joint groups affinis et Cypridopsis spp.) ne pas préfé- in wells is studied, especially for hadziid amphipods, which rence. are to on small crustaceans. There On examine la dans les à supposed predate may présence puits, conjointement be some influence ofthe ofhadziid on celle des d’autres presence amphipods Ostracodes, groupes d’animaux, surtout the distribution of some of but this des dont species Ostracoda, Amphipodes Hadziides, on suppose que ce sont could not be tested for most islands and des de Crustacés. statistically prédateurs au dépens petits Il est possi-

ble ait certaine de la de species. qu’il y une influence présence ces

Some models are discussed. Most biogeographical Amphipodes sur la distribution de certaines espèces ostracods have arrived the islands la des îles seem to at by dispersal. d’Ostracodes, mais, pour plupart et des espèces,

No ostracods were found evidence for the ceci n’a être mis évidence bases statisti- representing pas pu en sur des

modelof distribution and regression speciation. Practically ques. all seem to be stable and did not evolve species genetically Des modèles biogéographiques différents sont discutés. into true hypogean species. Only in the presumably older La plupart des Ostracodes semblent avoir abordé les îles islands and are Des Hispaniola Cuba, true hypogean species par dispersion. Ostracodes représentant des preuves

which seem to have evolved from relict freshwater le “modèle de des found, pour régression” ne sont pas connus ancestors. Caraïbes. Pratiquement toutes les espèces semblent être

The results the that Cuba and stables évolué donner de strengthen hypothesis génétiquement et n’ont pas pour

been which vraies formes Des Hispaniola have part of a proto-Antillean arc, hypogées. espèces effectivement hypo- has been connected with Yucatan. The fact that no n’ont îles gées été découvertes que sur Hispaniola et Cuba,

were found in the Lesser Antilles hypogean species supposées plus anciennes; ces espèces semblent avoir évo- strengthens the hypothesis that these islands have emerged lué à partir d’ancêtres dulcicoles relictes. above the sea level independentlyof the Proto-Antilles and Les résultats obtenus viennent à l’appui de l’hypothèse much later in time. Cuba fait d’un que Hispaniola et ont partie arc proto-

Antillais été relié Yucatan. Le fait les Petites ayant au que

Antilles n’ont fourni pas des espèces hypogées, est un

de de argument en faveur l’hypothèse que l’émergence ces

îles été a indépendente et de beaucoup postérieure à celle

* Report 37 is published in the same issue of this journal. des Proto-Antilles.

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INTRODUCTION vironment and. adapt to real subterranean

conditions.

the As In period 1973-1982 the Amsterdam Expe- (2) a place where typical groundwater

in the surround- ditions have made groundwater explorations organisms can emerge from

and of the and 43 West Indian islands part ing groundwater system, perhaps even

Venezuelan mainland. Most of the stations survive.

since these form the of To like the in- sampled were wells, way (3) study ecological processes,

the subterranean in fluence of restricted access to fauna, especially space, water volume,

A number karstified sediments. great of light conditions and other environmental

hypogean animals were encountered, but also factors, on the animals, and on the com-

animals. The between them. many epigean predominant petition

and four As observations what groups were Oligochaeta, Gastropoda (4) a place allowing on

when in crustacean taxa: Ostracoda, cyclopoid happens epigean species come con-

Copepoda, Amphipoda, and Thermosbaenacea tact with groundwater species.

(Stock, 1983a). The ostracods were considered Wells are like islands which can be colonized,

to be important because of their abundance and often by epigean, cosmopolitan species. The

in whether colonization is constancy hypogean habitats, being present questions are common,

in more than halfof the stations sampled, often whether many species succeed, which species

in large numbers. They can also be part of the are better colonizers, and whether it is possible

food for for the hypogean chain, being a possible prey some species to adapt to groundwater

larger hypogean amphipods (Stock, 1983a). environment. Furthermore, the differences in

Furthermore, the ostracods were thought to be composition and diversity of populations can be

usable for biogeographic modelling, since their studied.

distribution It became clear was thought to be influenced that only in the islands

and predominantly by dispersal, though some Hispaniola Cuba some species (belonging

and Strandesia species could have originated by regression of to the genera Neocypridopsis i) are

marine in ancestors or vicariant events. The dif- present, which are possibly some way

models and their ferent biogeographic use- adapted to live in groundwater environments,

fulness discussed for ostracods in this arti- of their small reduced are on account very size, or cle. The islands harbour other eye pigment. same

this it became clear that of all the Caribe- During study most species, belonging to new genus the ostracods encountered belong to epigean candona of the subfamily Candoninae, which

is caused the fact in In groups. This by that are stygobiont (Broodbakker, 1983c). a well

encountered in in of karstified sediments, as tropical the mainland Venezuela another hypo- and wells with to the new subtropical regions, a large gean species was found, belonging diameter habitat from the Danielocandona represent a differing genus (Broodbakker, 1983c).

subterranean surrounding area (Danielopol, Therefore it became inevitable to study most-

These wells and 1981). play an important role for ly the aspects (3) (4). Since most species

because of the it aquatic life; dry seasons or dry were supposed to be epigean, was necessary climats in these regions, and the lack of other to obtain samples from epigean habitats for aquatic habitats, they provide some ecological comparison. Dr. P. Wagenaar Hummelinck

shallow wells about 600 ostracod stability. Only open dry out kindly provided samples

of the like during part year, most shallow pools from mostly epigean habitats, sampled and ponds. predominantly in the Lesser Antilles, between

A well be in several 1930 and He stations can regarded ways: 1973. sampled many

As a where faunal times in different months (1) place some epigean ele- several or even years,

ments can survive and the islands perhaps even pene- especially on Curaçao, Bonaire,

in the and trate deeper true groundwater en- Aruba St. Martin.

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I received Dr. D. L. of the Strandesia Furthermore, through genus (Broodbakker, 1983d,

Danielopol 37 ostracod samples from Cuba, 1984a), of the subfamily Candoninae (Brood-

from but also few from for the mostly caves, a pools bakker, 1983c), and species Tanycypris

and the underflow of running waters, sampled meridana (Furtos, 1936) (Broodbakker, 1984b).

Drs. L. Botosaneanu, St. and T. Other like and by Negrea genera, Cypretta Cypridopsis,

Orghidan, members of the Cubano-Romanian could not be studied because of the lack of good

Biospeleological Expeditions to Cuba, in 1969, descriptions and of time.

1970 and 1973. In the the distribution of all present paper

few have been freshwater all islands Only papers published con- ostracods, on sampled, is

cerning the freshwater ostracods of the West In- recorded and discussed. The ostracod fauna of

dies. Brady (1902) described Cypretta sarsi from wells is discussed and compared with that of

St. Thomas, Sharpe (1910) described Chlamydo- epigean habitats. This is done since most of the

barbadensis and Brehm wells theca from Barbados, habitats sampled were either or stagnant

(1949) described some species, viz. Cypretta epigean habitats. Roughly 50% of the samples

taken in both margalefi, Cytheridella ilosvayi Von Daday, 1905 types of habitat, being 360 wells,

and Stenocypris major (Baird, 1859), from Cuba. and 182 stagnant epigean habitats, yielded

Most other have been described Klie Ostracoda. 37 of which 24 in species by Only caves, Cuba,

stations (1933) and Triebel (1961, 1962) from yielded Ostracoda, while only few caves have

sampled by Dr. P. Wagenaar Hummelinck. been sampled in the other islands.

described some from the effect of the of Margalef (1961) species Furthermore, presence

the Venezuelan island of Margarita, which hypogean amphipods, in particular Metani-

proved to be already described by Klie (1933) phargus, is studied. Finally, the different

from Bonaire and Aruba. Van den Bold discussed in (1958) zoogeographical models are con-

mentioned some species from rivers in nection with the distribution of the freshwater

Trinidad, which he thought to be identical with ostracods in the West Indian islands.

European species. Harrison & Rankin (1976),

in a hydrobiological study of St. Vincent,

number this reported a of species from island, MATERIAL AND METHODS

which were in part wrongly identified.

Information concerning the samples is provided in tableI, During my research it became clear that in Stock (1979), Wagenaar Hummelinck (1940a-b, 1953 & had been which many species not yet described, as S and WH in the and in 1981) (abbreviated sequel) my meant that of research had to a great part my the the West In- previous papers concerning Ostracoda of

consist of a taxonomie analysis of a few dies.

freshwater ostracod and of the sorted data on habitat, conditions, genera, partly Computer light water table, water depth and the abundance of hypogean subfamily Candoninae. It appeared accompa- with the nying fauna, were statistically compared presence describe both the the necessary to carapace and of the different ostracod of G-tests. species by way In order describe limbs as precise as possible. to

the Dr. and I The G-test. — Instead of chaetotaxy, Danielopol designed chi-square tests, G-tests were

because the G-test has a theoretical a descriptive system (Broodbakker & Danielo- performed, advantage the and is over former, simpler (Sokal & pol, 1982). Furthermore, the ecology of species computationally 1981: for the Rohlf, 704). Critical values G-test can be the chosen dif- belonging to groups was studied, found in a since the distribution of G chi-square table, ap- ferences between of different populations the distribution. The proximates chi-square tests are one-

habitats and in islands, especially carapace tailed since only large values of G are used for rejecting

the that and there is no influence of the factor length, were examined, the geographical hypothesis in

question. G was computed for frequency distributions distribution of these and related species was arranged by a single criterion of classification. Frequency the of discussed. This was done for species the distributions for all dominant species were calculated for

genus Heterocypris (Broodbakker, 1982, 1983a), islands and of environmental grouped classes factors, or of the of genus Hemicypris (Broodbakker, 1983b), types accompanying fauna.

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the were calculated multiplying Venezuela is described and Expected frequencies by discussed separate- ostracods in each number of samples containing type or ly. The criteria for grouping the islands were class (re,) with the total number of samples yielding the either short mutual distance, proximity to the ostracod in question («]), divided by the total number of the bank. in the island mainland, or a position on same samples yielding ostracods respective group

which is in table III. So: All islands are listed in table I and or island («2), illustrated sampled

shown in 1. In the table of the n fig. many species F t_ni.

re2 encountered in the West Indies are listed, ex-

for which in Only when all expected frequencies are larger than 5, it is cept species were only found one or

to calculate a G- value. Whenever a G-value was permitted two islands, for species of Darwinula, and for significantly large, it was adjusted by applying Williams' Perissocytheridea cribrosa (Klie, 1933). The species correction (q) (Sokal & Rohlf, 1981: 705): that are not listed are discussed in the text. In

= — of the of table I the total number of q=\+ -7 (v degrees freedom) right part

fresh- and brackish-water stations that have G r been sampled by the Amsterdam Expeditions 1 and the expeditions of Dr. Wagenaar Humme- Data were sorted with the CDC computer of SARA linck listed The combined (Stichting Academisch Rekencentrum Amsterdam). are separately.

with Sinclair home G-tests were performed a ZX-spectrum number of samples yielding ostracods is given,

computer. well the number of from each of as as samples

the listed habitats that yielded ostracods.

DISTRIBUTION OF THE SPECIES MAINLAND OF VENEZUELA

I: (Table nrs. 1, 2)

In fauna of this chapter the ostracod every Part of the Estado Falcon was sampled exten-

the Amsterdam sampled island group and the mainland of sively in 1982 by Expeditions,

of the Fig. 1. Map Caribbean, showing (dotted) the 200 m line being the edge of the continental shelf. The numbers of

the islands and regions correspond with those given in table I.

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the Sierra de San calcareous the in especially Luis, a belonging to Cypridopsini was found an

chain of mountains, and the Peninsula de interstitial sample from the Estado Sucre. This

a karst with cenotes. will be described in Morocoy, region many species a forthcoming One Strandesia venezolana Broodbakker, species, paper.

is in the 1983d, very common, especially

cenotes, but it was also found in the underflow

in this of running waters state and in a river 50 THE VENEZUELAN ISLANDS

km south of It Caracas (Broodbakker, 1983d). (Table I: nrs. 7 to 12)

encountered in Pseudocandona Isla was once a cave. The Margarita, which is situated on the

aff. antilliana, Ps. and hum- geratsi Candonopsis continental shelf of Venezuela, presents twelve in melincki were found the underflow of running species of ostracods. One species, Rudjakoviella

waters in the Estado Falcon, but also in other prolongata (Triebel, 1962) being the only known

states of Venezuela, as Sucre and Aragua. All representative of the tribe Rudjakoviellini of the

these candonid described and assumed be species are subfamily Cypridinae, was to

discussed in Broodbakker C. hum- endemic (1983c). to Margarita. However, it was also

melincki also found in the of in the of was underflow run- found a pool on island Iguana Los

ning waters in the Distrito Federal. Testigos (WH 30). Triebel (1962) described it

Darwinula and from roadside Cypretta sp., sp. Physocypria sp. a pool on Margarita (WH 24). It

found in and of were some cenotes springs the appeared to be present in samples from the

Estado Falcon. In and 18 and one muddy pool a very pools WH S 82/19 as well.

shallow, muddy spring, specimens of Chlamydo- Heterocypris margaritae Margalef, 1961, is the

theca aff. found arcuata (Sars, 1901) were found, once species most commonly on Margarita. It

with also found the together Physocypria sp. was on Venezuelan islands La

The in 1982 Los and La districts of Falcon sampled are Blanquilla, Testigos, Tortuga relatively humid and rich in vegetation. In the (Broodbakker, 1983a). arid Peninsula de which of Triebei hum- Paraguana, is part (1961) described Chlamydotheca

the Estado Falcon, species like Chl. aff. arcuata melincki with two subspecies, hummelincki and and S. venezolana were not found. The ostracod dispar, from St. Eustatius (WH 507). During fauna of this peninsula, which was sampled by my research it became clear that this species has

the Amsterdam in 1978 Expeditions and by Dr. a wide distribution, being common in the

P. Wagenaar Hummelinck in 1937, shows Venezuelan islands, St. Eustatius, St. Martin

more affinities with that of the arid islands off and other Lesser Antilles, including Barbados.

the and with that of the coast, partly arid penin- However, as early as 1910, Sharpe had describ-

sula de coastal district the Isla ed Chl. barbadensis from small Araya, a opposite a pond on

In twelve wells Margarita. only four out of Government Hill, Barbados. Triebel con-

in sampled Paraguana, ostracods were found. sidered this to be a different species, because of

In three of these wells minor differences in the Cypretta sp. was found, description of the and them also maxillula. two of yielded Heterocypris valves and Since Sharpe' s descrip-

margaritae Margalef, 1961, while the fourth well tion is relatively poor, I consider the two species

three of be yielded specimens Chlamydotheca barba- to synonymous, especially since I found this

1910 discussion densis Sharpe, (see on its tax- species on Barbados, St. Eustatius, and in

from other islands. Therefore onomy below). samples many

In well a in Calabozo, a town 150 km south Chl. hummelincki becomes the of junior synonym of Caracas in the Estado Guârico, a new hypo- Chl. barbadensis. The subspecies dispar, describ-

of gean species a new genus of the subfamily ed by Triebel (1962) from St. Martin, and the

Candoninae lieshoutae was found, Danielocandona subspecies nordestina described by Kotzian

Broodbakker, 1983c. Furthermore, another (1974) from Brazil, have to be ascribed to Chl.

of probably hypogean species a new genus barbadensis.

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caves from 1 4 2

waters running 3 1 samples of underflows from

1 3 3 1 of 1).of springs from 8

(coral-rock) karst 24 fig. 2 ostracod 5 3 2 in holes water from in presence of 8 2 2 4 expeditions etc. pools ponds, from 1 16 26 2 11 37 the 5 9 — 6 —3 7 4 the indicated wells from 18 13 27 Number 211 3— 7 2 3 18 2 12 1982, also indicates number total 20 31 47 71 18

to are 3 8—3—2—1 3 52 67 17 13 15 15 samples S of number total 612743 89 46835 4—3—3 15—3—1 31 151174624 1973 asterisk 6— samples 5 2 58—8—8 3 4 2 WH of number total 32 49 64 18 13 11 numbers An 13—3—3 12 Islands, unispinosa Chlamydotheca listed. Neocypridopsis 6 1 (Island inaudita

are 4 1 1 Indian longula Strandesia 1973. 1 1 stocki Strandesia West to genus. 7 decaryi Cypris 1 16 1 the 1969 encountered

subglobosa 1 5 2 3 1 to same Cypris 3

Cuba, species the hummelincki Candonopsis 2 to to I 1 2 antilliana Pseudocandona 1 4 Expeditions 9* 11 2 6 TABLE ostracod belonging antillensis Heterocypris 9 14 11 1 expeditions meridana Tanycypris 1 1 the 12 32 of 7 Amsterdam species exigua Hemicypris 4 2 1

1 Most reticulata Hemicypris 3 the Romanian 111

3 2 4 5 another hummelincki Potamocypris 11 17 and 13 during Chlamydotheca 1 1 indicated. dispar b. 2111

2 1 6 1973, barbadensis Chlamydotheca 3111 631414 are 11 to 1 1 112 1 1 investigated margaritae Heterocypris 15 1454 2211 1930 samples punctata Heterocypris 1 2

1 3 1 1 1 Stenocypris 2 1 regions major 18 17

sp. Physocypria and Hummelinck, 23

1 brackish-water affinis Physocypria 2 7 4 1 2 1 13 11

4 1 sp. Cypridopsis 1 1 islands — 31 20 —— —— 53 251 — 1 and —8 32 312 111 321 —— 21 11 — — 11 — —4 21 sp. 5 2 1 4 3 7 — 2 Cypretta — 2 Wagenaar 11 19 Indian fresh- P. Saintes West Dr. Only Falcón Galante Curaçao Roques Testigos Tortuga Blanquilla Lucia des Désirade Paraguana Curaçao Margarita Trinidad Grenada Barbados Martinique Dominica Guadeloupe Montserrat Antigua Barbuda Estado Aruba Klein Bonaire Los La La Araya Los St. îles Marie La

1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25.

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caves 1 from 2 2 24 1

waters running 4 samples of underflows from

2 5 springs from 1 1 —

(coral-rock) karst —1 4 1 1 ostracod — in holes water from 31—

of 2 1 4 1 etc. pools ponds, from 18 12 94 8 5 1 wells from 11 11 16 17 20 Number 55 85 87 44 44 33 102 93 55 66 2 3 5 number total 10 31 11 11 115 15 1164 532 17 17 888 21 831 2—2—1—1 5 17 15 16 13 17 22 10 15 14 15 23 25 samples of total 43 12211 43 S number 222

5 2 5 2 5 5 7 7 samples WH of number total 15 24

2 unispinosa Chlamydotheca 37

5 1 * * 1 5 4 inaudita Neocypridopsis 3 8 3* 2 2125 2 22

7 2 3 1 6 longula Strandesia 3 33 2241— 11 14

1 1 1 1 stocki Strandesia 39

decaryi Cypris 1 21 subglobosa Cypris 1

1 1 hummelincki Candonopsis 11 * * antilliana Pseudocandona 11 8* 1*

1 1 6 antillensis Heterocypris

meridana Tanycypris

exigua Hemicypris 1122 1

reticulata Hemicypris 1

1 1 6 hummelincki Potamocypris dispar b. Chlamydotheca 412 1 barbadensis Chlamydotheca 2

margaritae Heterocypris 1

4 1 punctata Heterocypris 1

major 1 1 1 Stenocypris 11 18 21 sp. 1 1 4 Physocypria 12 10 11 affinis Physocypria 9 23 sp. 2 2 7 Cypridopsis 1 6 8 8 —— —— — 33 —— 11 —— 2 —— 212 12 ——— 2 112—— — 211 — 1 —— 45 ——— ——— —— 32——— ———— 13 10 345 115 246 276 23 —— 3 14 33 14 —— —— ——— —— ——— ——— 10 ——— 1 —2— 18 ——— 11 21 11 42 sp. 1 2 Cypretta 8 6 10 39 4 5 4

Brae Cayman Island Gorda Rico Caicos Salvador Providence Kitts Eustatius Barths Martin Croix John Thomas Anguilla Anegada Tortola Culebra Vieques Puerto Cayman Providenciales Mayaguana Crooked Eleuthera Nevis St. St. Saba St. St. St. Virgin St. St. Haiti Cuba Jamaica Grand Inagua South San New

26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54.

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Other species which were found on fauna. Bonaire, in which most samples were

in Margarita as well as on La Blanquilla, Los taken, presented most species, but contrast to

Testigos, and Araya, are Physocypria affinis and Margarita it is not the largest of these three

Potamocypris hummelincki, both described by Klie islands, as it comprises only 60% of the size of

(1933) from Bonaire and Aruba. The latter was Curaçao.

continen- erroneously considered by Margalef (1961) as a On Aruba, which is situated on the

P. tal shelf new species, ombrophila, from Margarita. of Venezuela, Heterocypris antillensis

Margalef also described Physocypria sanctaeannae Broodbakker, 1982, and H. margaritae are the

from Margarita, a species different from P. af- most common species. On Bonaire these species

his which are also but on this island finisaccording to drawings, was, common, Cypretta sp.,

encountered this and however, not during study. Stenocypris major, Cypris decaryi, are common

found La Tor- too. On the same are common- Cypretta sp. was on Margarita, Curaçao species

and Los Steno- H. which tuga, as only species on Roques. ly found, except for margaritae, was

in cypris major (Baird, 1859), Hemicypris exigua present one sample only. However, on this

island is the Broodbakker, 1983b, Cypris decaryi Gauthier, Cypridopsis sp. most common

and all found while it found and 1933, Cypridopsis sp., were once species, was not on Bonaire,

but on Margarita, not on the other Venezuelan only once on Aruba. Neocypridopsis inaudita was

islands. meridana found Bonaire Klein Tanycypris (Furtos, 1936) was on only, and once on encountered and Los small island km 2 between on Margarita Testigos Curaçao, a very (1.2 )

(Broodbakker, 1984b). Neocypridopsis inaudita Curaçao and Bonaire.

found in hole with The of in (Furtos, 1936) was a presence Cypridopsis sp. many brackish La and of but not is water on Tortuga, Heterocypris samples Curaçao, on Bonaire, very

in well is almost absence punctata Keyser, 1975, a small dug-out peculiar, as the complete of on La Blanquilla (Broodbakker, 1983a). Heterocypris margaritae on Curaçao. Further-

the absence of barbadensis It seems that the freshwater ostracod fauna of more, Chlamydotheca on

Bonaire and and of antil- the Venezuelan islands is essentially the same Curaçao, Heterocypris

that of the Netherlands the as nearby Antilles, lensis on Venezuelan islands, is striking.

Bonaire and meri- Another is the Aruba, Curaçao, Tanycypris strange phenomenon presence dana the being only species which was not en- of a species resembling Chlamydotheca mexicana countered on one of these three islands. Isla Sharpe, 1903, in pools on Klein Bonaire, a

2 Margarita presents most species, probably small island (7 km ) 750 m west of Bonaire. because This nowhere else in the on this island more stations were species was found

the other but also sampled than on islands, Caribbean. because there much less freshwater habitats In Darwinula cf. are many springs on Curaçao available on the other smaller islands. Fresh stevensoni (Brady & Robertson, 1870), and/or

is absent La which Pseudocandona antilliana water virtually on Tortuga, Broodbakker, 1983c,

Bonaire D. stevensoni presents only some brackish-water habitats, were found. On cf. was and Los few found in and in Roques, which presents only a in a spring, several cave waters,

found in rainwater pools. a sink hole. On Aruba it was once a

The best find which In other Bonaire way to out species are spring. some springs on essentially present on these small islands would another unidentified species of Darwinula was be raise several Ps. antilliana to ostracods from samples of found. was neither found on dried mud and from dried-in pools or borders of Bonaire nor on Aruba.

from each island. In brackish-water habitats of all three pools many

islands cribrosa Perissocytheridea was found, a

described ARUBA, BONAIRE, CURAÇAO species as Leptocythere by Klie (1933)

brackish rede- (Table I: nrs. 3 to 6) from waters on Bonaire, and

have These islands a comparable ostracod scribed by Keyser (1975) from brackish waters

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in southern Florida. This species was also pres- WINDWARD ISLANDS

ent in brackish-water samples from St. Martin (Table I: nrs. 14 to 17)

and Anguilla. From the volcanic islands, Grenada, St. Lucia,

The ostracod fauna of Aruba, Bonaire and and Martinique, only few samples are avail-

the the have been in Curaçao is essentially same as that of able. No stations sampled The

islands and the arid of the and St. Vincent. Venezuelan parts Grenadines

continent, as was mentioned before. Since the St. Vincent has been the subject of intensive

much Netherlands Antilles were sampled more hydrobiological investigations by Harrison &

than intensively the other islands, more species Rankin (1976). They found several species of

The of were encountered. absence or presence ostracods: Cypretta infesta Klie, 1943, Cypretta

some species in one or more of these islands can sarsi Brady, 1902, and Strandesia elliptica (Sars,

be factor in the of not explained by any ecological 1901), stony parts rivers; Cypridopsis

measured. A find if there are dif- vidua and in way to out Stenocypris sp., permanent pools;

ferences in species diversity and if some species moreover, all these species including Darwinula

of dried are really absent, is to collect samples stevensoni, in marshes. In the marginal vegeta-

mud in islands and raise ostracods from tion of the to a trophically rich, slowly flowing river

these mud samples. they found large numbers of Stenocypris major,

and in river pools near the sea Heterocypris

In TRINIDAD symmetricus (G. W. Müller, 1898). one rivulet

I: (Table nr. 13) Chlamydotheca unispinosa (Baird, 1862) was

From Trinidad only few ostracod samples are found.

A the Har- available. few brackish-water samples con- However, species determined by

tained Thalassocypridinae, a subterranean rison & Rankin as Strandesia elliptica is most prob-

in river yielded Darwinula cf. stevensoni and another ably S. stocki, as is explained Broodbakker

and is H. Darwinula species, a fish pond Stenocypris (1983d); Heterocypris symmetricus probably

and mistake also made Furtos major Cypridopsis sp. punctata, a by (1936)

for a from Yucatan (Broodbakker, Van den Bold (1958) studied ostracod specimen Most will have be collected in rivers Trinidad. He 1983a). Cypretta species to carapaces on restudied since the characters of mentions D. stevensoni, S. major, Cypridopsis cf. morphological

of these are known. Sohn & Kor- vidua (O. F. Müller, 1776), and Candona cf. most barely

from the nicker (1973) wrote an excellent monograph on fabaeformis Fischer, 1854, upper ranges but than 15 above level. the redescribed some of the rivers, more m sea genus, they only

Victor & Fernando redescribed Except for Candonathese are the same species as species. (1981) and describedeleven species from Malaysia, In- found in the few stations sampled by Dr. P.

Van donesia and the All from Wagenaar Hummelinck on this island. Philippines. species

the are or described, den Bold found several other species in the Neotropics barely poorly

which is the most reason why the slowly flowing to semi-stagnant waters in flat important Cypretta encountered during this study country, which he thought to be identical with spp. could not be identified. Under these European species, like: Ilyocypris cf. bradyi Sars, conditions,

all found by Harrison & Rankin are 1890; Cypria cf. exsculpta (Fischer, 1855); C. cf. species also found in the other Antillean ophthalmica (Jurine, 1820); Eucypris cf. ornata (O. most probably

~ ~ islands. F. Muller, 1776); and E. cf. lilljeborgi (G. W.

if all of The that are in the few Müller, 1900). In my opinion, most not species present these American samples from Grenada, St. Lucia and Martini- species represent South species also in the with the are found other Antillean which are not identical species men- que tioned above. islands.

In Barbados wells and some The other species discussed by Van den Bold many epigean stations wells marine were and are and brackish-water species. sampled. Only eight

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most of the epigean habitats yielded ostracods. which only one yielded one ostracod, a

of three The species encountered are the same as those specimen Stenocypris major. Only

of which from the other Lesser Antilles, with the excep- epigean stations were sampled, one

S. and tion of Hemicypris barbadensis, a species endemic yielded major, Cypris subglobosa Hemicypris

to Barbados, that strikingly resembles H. aurita exigua.

(Klie, 1939) fromthe high mountains in Kenya.

It is described in Broodbakker (1983b). MONTSERRAT

I: (Table nr. 23) LEEWARD ISLANDS On Montserrat only four epigean stations have (Table I: nrs. 18 to 40) been sampled, of which a pond harboured S.

and streamlet aff. major, a Cypridopsis vidua, DOMINICA Cypretta sp., and Tanycypris meridana. On Dominica only few stations have been

sampled. Only two samples yielded ostracods. ANTIGUA BARBUDA A slowly flowing streamlet yielded Cypridopsis &

I: nrs. 24 and aff. vidua and Strandesia stocki Broodbakker, (Table 25)

a calcareous and and brackish Barbuda, island, Antigua, a 1983d, a ditch yielded a species of partly volcanic and partly calcareous island, are the subfamily Thalassocypridinae.

situated on the same shelf fragment, just north

of Guadeloupe. On Antigua only one pond, one GUADELOUPE, MARIE GALANTE, basin and five wells ostracods. All ÎLES DES SAINTES, LA DÉSIRADE yielded

species, except for Strandesia sphaeroidea, have (Table I: nrs. 19 to 22) been found also in the islands off the On these islands some epigean stations have

Venezuelan coast. been sampled by Dr. Wagenaar Hummelinck, On Barbuda only 2 out of 13 wells yielded which yielded mostly the same species as the because of the of islands off Venezuelan but for few ostracods, perhaps presence the coast, a

the large hypogean amphipod Metaniphargus a exceptions. Chlamydotheca unispinosa (Baird, , situation which will be discussed in the sequel. 1862) was found on Iles des Saintes, Marie In contrast, practically all epigean stations Galante, and Guadeloupe. This species is sampled yielded ostracods. All species en- common in Haiti, and is moreover found in St.

countered, except for Neocypridopsis debilis Klie, Vincent, the Bahamas, Cuba, Jamaica, the 1940, have also been found in the islands off the Cayman Islands, southern U.S.A., and

Venezuelan coast. N. debilis was described from Yucatan, but not in the other Lesser Antilles. It

northeast Brazil, but was rediscovered in seems to be represented by disjunct populations from Barbuda and St. Martin. islands around samples on the Guadeloupe and on St.

Vincent.

Marie Galante presented two species which NEVIS, ST. KITTS, ST. EUSTATIUS, SABA are not, or rarely, found in the other islands: (Fig. 2; table I: nrs. 26 to 29)

Strandesia trispinosa galantis Broodbakker, 1983d, Of these volcanic islands, the first three are

of S. a subspecies trispinosa which was described situated on the same shallow bank (fig. 2). On by Pinto & Purper (1965) from southern Brazil; Nevis and St. Kitts only four epigean stations and S. All for sphaeroidea Broodbakker, 1983d, which have been sampled. species, except found in several was samples, and furthermore Strandesia stocki, are also found in the islands off in only one sample from Antigua. the Venezuelan coast.

meridana Tanycypris was found on three of the On St. Eustatius only 4 out of 17 wells yield- islands of this ed Dr. group (table I). ostracods. Wagenaar Hummelinck

The Amsterdam Expeditions collected 13 sampled 11 uncovered pool-like wells and 4

well of of which All pump- or samples on Guadeloupe, cisterns, 8 yielded ostracods.

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encountered the also in islands species occur on islands off the encountered are found the other

Venezuelan coast as well. of the Leeward Group.

Saba few stations have been stations have been On only sam- On St. Martin many

One harboured the Amsterdam pled. pool Cypridopsis species sampled, by Expeditions as well

and Heter and Dr. In ocypris antillensis, a spring yielded as by Wagenaar Hummelinck. brackish

Cypretta sp. waters either Heterocypris punctata or

Perissocytheridea cribrosa (Klie, 1933) were found.

Stenocypris major, Potamocypris hummelincki, and

found in Cypridopsis sp., were many pools,

wells. cisterns and open pool-like Cypretta sp.

encountered in wells was and pool-like wells,

and Chlamydotheca barbadensis in a few cisterns.

wells Heterocypris antillensis was found in three

in and and three pools, H. margaritae one pool,

debilis in well and Neocypridopsis a a trough.

Strandesia longula was found in a well, a pool-like

rock well and a pool, and S. stocki in a pool. The

latter described in two Strandesia species are

Broodbakker (1983d).

Mr. dried mud J.J. Jongsma sampled some

for in Slob of few km of me Welgelegen, a west

Philipsburg, on St. Martin. From this sample I

have raised five species of ostracods: Stenocypris

major, Strandesia longula, S. stocki, Physocypria af-

and debilis. The fact that finis Neocypridopsis so

could be raised in few many species just a

from in- weeks, one sample of mud, seems to

dicate that this is another of way getting some

better idea of the ostracod faunaof the West In-

dian islands, especially since part of the species

stations raised were present in only few of the

sampled.

The ostracod fauna of St. Martin seems to be

the for greater part the same as that of the

islands off the Venezuelan coast, with a few ex-

H. reticulata Fig. 2. Map of the islands on the shallow banks of St. ceptions: Hemicypris exigua, (Klie,

Martin and island Saba. and St. Kitts, the Depth contour 1930), Cypris decaryi, and C. subglobosa Sowerby,

intervals of 35 m and 180 m are indicated U.S. (after 1840 (all being epigean species, especially com- Hydr. Off. Charts). in the Antilles mon Netherlands off the

Venezuelan coast, but also found in Puerto

Both in ST. BARTHÉLEMY, ST. MARTIN, ANGUILLA Rico). Cypris species were also found

table I: 30 Haiti. neither of these (Fig. 2; nrs. to 32) However, species was

St. Martin and St. which Both Barthélémy are partly found on St. Martin. The only species

and is the islands off the volcanic partly calcareous; Anguilla were not found in

All islands situated Venezuelan Strandesia merely calcareous. three are coast are: stocki, a species on the same shallow bank. common in Haiti, which was also found in

On few from St. Barthélémy only wells, but no samples Martinique, Dominica, St. Kitts epigean stations, yielded ostracods. All species and St. John, and probably St. Vincent (Brood-

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- U.S. Off. Fig. 3. Map ofthe Puerto Rico Virgin Islands area. Depth contour intervalof 180 m is indicated (after Hydr. Charts).

and which Islands bakker, 1983d); Neocypridopsis debilis, the remaining Virgin by a deep trench

was also found in Barbuda, and was described (fig. 3), only one out of the 13 wells sampled

Klie from Brazil. by (1940) yielded ostracods, belonging to Physocypria sp.

On Anguilla only few epigean stations have Of the five pools sampled by Dr. P. Wagenaar

been sampled. A brackish puddle yielded Hummelinck, two yielded the species Hemicypris

while the brackish Heterocypris punctata, mostly reticulata, Stenocypris major and Cypridopsis sp.

wells is the in the yielded Neocypridopsis inaudita, Eucypris sp., Cypretta sp. most common species

Perissocytheridea cribrosa, and Thalassocypri- wells of the Virgin Islands, followed by

dinae. Darwinula was found once. and sp. Physocypria sp. Cypridopsis sp. (table I).

On Dog Island, a small key just north of Other species encountered are: Candonopsis hum-

well Anguilla, one was sampled, which har- melincki and Pseudocandona antilliana, both found

boured Pseudocandona antilliana and Darwinula in shallow wells once very pool-like on Vieques,

sp. (Broodbakker, 1983c). Strandesia longula in a well on Culebra and in

three wells on Tortola, and a specimen of a

Pseudocandona species in a well on Tortola. On

hummelincki VIRGIN ISLANDS, ST. CROIX, PUERTO RICO Virgin Gorda, Candonopsis was

table I: 33 found in antillensis in (Fig. 3; nrs. to 41) an open well, Heterocypris a

Practically all stations sampled on the Virgin shallow open well, and a hitherto undescribed

in Islands were wells and rock crevices karst species of Neocypridopsis, closely related to N.

Hummelinck in with Darwinula areas. Dr. Wagenaar sampled debilis, a spring together sp.

and Puerto of some epigean stations on St. Croix On Anegada the same species Neocypridopsis

in and in in Rico, and one station on St. Thomas. Only few was found a well, three crevices a

the Amsterdam karst wells could be sampled by Ex- area, together with Neocypridopsis inaudita.

in works peditions Puerto Rico, because water Only few ostracod samples were available

have been constructed, and most old wells have from each island. Many species were found

been filled well in Puerto Rico once in the up. Only one only mostly shallow, open pool-like

wells is therefore yielded a damaged specimen of an ostracod. sampled. It highly probable

richer in The few stations sampled by Dr. Wagenaar that these islands are much species

Hummelinck than is known. for St. yielded relatively many species presently Except Croix,

On St. which Puerto (table I). Croix, is separated from all islands, including Rico, are situated

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It is that which will be in the on the same bank. highly probable this species discussed section on

least 7000 bank emerged as a whole, at only Cuba.

After this time the islands were Other which encountered years ago. species were only

of in the separated because a rising sea level (Pregill & once wells, are: Strandesia pistrix Brood-

This could that all these and male Olson, 1981). mean bakker, 1983d, a specimen of an

islands potentially have the same ostracod unknown species of Strandesia (Broodbakker,

fauna. and 1983d), Cypris subglobosa, C. decaryi, an

According to the available data, the ostracod unknown species of Cypris. The first two Cypris

Puerto St. likewise in of the fauna of the Virgin Islands, Rico and species are common some

West Indian which Croix could be essentially the same as that of islands, being species com-

the other Leeward Islands. No Chlamydotheca monly live in epigean waters, like pools and

have been but this be due species found, may to ponds.

the low number of stations in the wells of epigean sampled, as The most common species

is illustrated the found in the Haiti the Strandesia by many species are epigean species longula,

few epigean samples from Puerto Rico and St. S. stocki, Chlamydotheca unispinosa, and Cypretta

Croix. is this island The distribution and of the first Only one species specific to spp. ecology

is discussed in Brood- group, Neocypridopsis sp., a species closely two species extensively

related to N. debilis. bakker (1984a). Chlamydotheca unispinosa was

found in twice only once Cuba, on Grand

and few times in the Bahamas. It is HAITI (HISPANIOLA) Cayman, a

in the southern of the (Table I: nr. 42) a common species parts

In 1978 and 1979 the Amsterdam Expeditions U.S.A., Central America, and South America,

150 and under- found in Hawaii It sampled about wells, 72 springs and is even (Kotzian, 1974). flows of and 6 in Haiti. West In- running waters, caves, is probably much more common in the

In 102 wells, which is 68% of the total number dian islands, but rarely encountered during this

wells ostracods found. 8 since bur- of sampled, were Only study most Chlamydotheca species are springs, 3 underflows of running waters, and 2 rowing species of muddy, epigean habitats.

Strandesia caves, yielded ostracods. In one cave Cypretta spp. are very common in most West In- cavernicola dian in wells. Broodbakker, 1983d, was discovered, islands, especially and in another Darwinula cf. stevensoni, both Other species encountered in a number of

that found in other wells and species were not any sample are: Stenocypris major, Physocypria sp. from Haiti. inaudita found Cypridopsis sp.; Neocypridopsis was

Except for a male specimen of a Pseudocandona twice in covered wells. species, all species which have been found in

and of springs underflows running waters are also common in wells. In the wells some species CUBA were found once: hummelincki, only Candonopsis (Table I: nr. 43) also found in other Antillean and many islands, Dr. L. Botosaneanu, Dr. T. Orghidan and

auricularia and C. Caribecandona ansa, both other members of the Cubano-Romanian Ex-

which species closely related to C. trapezoidea, peditions in 1969, 1970 and 1973, sampled

found in number of wells. The Caribecan- in was a many caves Cuba, but also a few pools and dona which described and discussed species, are underflows of running waters. Danielopol in Broodbakker are the en- described (1983c), only species (1972) a new species, Danielopolina countered in Haiti, and even the West Indian from orghidani, an anchyhaline cave sampled by

that islands, are clearly stygobiont. Practically these expeditions. This species is closely related all other ostracod found in Haiti, Cuba of the species to species genus Thaumatocypris which is and the other islands less are more or epigean only represented by species living in the deep-

with the of few species, possible exception a sea. It is the only species known in the Carib-

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bean which must have originated by a "regres- tween Yucatan and Cuba, since it is practically

sion type" of evolution, from marine ancestors. impossible for hypogean freshwater species to

barrier. Furthermore, Danielopol (1982) described cross a sea

from iden- Candonopsis cubensis another (freshwater) Darwinulids, which were not further

in found in of the cave, Cuba, a species which proved to belong tified, were many cave samples.

described in Other found in the holes in to the new genus Caribecandona, species few pools,

Broodbakker endemic Cuba holes of landcrabs and the underflow (1983c), a genus to limestone,

and Hispaniola. It is also the only represen- of running waters, include: Cytheridella ilosvayi

Von tative of the subgenus Cubacandona, which is Daday, 1905, Physocypria sp., Cypretta sp.,

be endemic Cuba. and supposed to to Cypridopsis sp., Cypris sp., Eucypris sp.,

Dr. I received 24 Through Danielopol Ilyocypris sp.

ostracod from caves in Cuba. Pseudo- Since most from and samples samples are caves, only

candona antilliana Broodbakker, 1983c, was a few from other habitats, it is not possible to

in of but also in ostracod of found part these samples, get an overall view of the fauna

the underflow ostracod samples from a pool and from of Cuba, or to compare the fauna found,

running waters (Broodbakker, 1983c). Ps. with that of the other islands.

in- cubensis Broodbakker, 1983c, probably an

in under- terstitial species, was found twice the

and flow of running waters, could be endemic JAMAICA, CAYMAN ISLANDS

Two I: 44 to Cuba. possibly stygobiont species of (Table nrs. to 46)

Strandesia, S. botosaneanui and S. cavernicola, were In Jamaica only seven wells have been sampled,

also encountered in the which caves (Broodbakker, of three yielded ostracods ( Cypridopsis

The first be endemic 1983d). species seems to to sp.). Of the 36 samples from springs and the

while found in of 5 Cuba, the second has been a underflows running waters, only yielded cave in Haiti as well. S. cavernicola seems to have ostracods. Pseudocandona caribbeana Broodbak-

reduced with otherStrandesia both Steno- a eye, as compared ker, 1983c, was found twice, and

in the Caribbean found species encountered (Brood- cypris major and Cypridopsis sp. were once.

If cavernicola is In brackish karst lakes and karst bakker, 1983d). S. really a stygo- the few holes biont species, this could indicate further proof sampled, representatives of the subfamily of a former connection between Cuba and Thalassocypridinae were found. In the four

Haiti, as proposed by Hedges (1982). stagnant waters and two rivulets sampled by

in Dr. ostracods S. longula and S. stocki were found the few Wagenaar Hummelinck, no were epigean samples, as well as a variety of found.

Hemicypris exigua different from populations in Of the 24 wells sampled in both Grand other islands by the reticulated inner side of its Cayman and Cayman Brae, 14 yielded

ostracods. carapace (Broodbakker, 1983b). Only Cypretta spp., Cypridopsis sp.,

inaudita and N. mexicana and Strandesia were Neocypridopsis (Fur- Physocypria sp., longula tos, 1938) were found in four and six cave found. Of the 8 pools and the single natural samples, respectively. Both species have been hole in coral rock sampled in Grand Cayman, 5 described and the S. (as Cypridopsis ) from cenotes yielded same species, except for longula. caves in Yucatan. N. inaudita is also found in Furthermore, Chlamydotheca unispinosa, Neocypri- epigean samples from Yucatan and other West dopsis inaudita and two undetermined species,

Indian but N. in in islands, mexicana only caves probably belonging to the genus Candonocypris,

Cuba. This, combined with the fact that N. mex- were found. icana is one of the smallest freshwater ostracod The number of stations yielding ostracods, species presently known, could mean that N. sampled in Jamaica and the Cayman Islands, is mexicana is and therefore allow conclusion about a stygobiont species, not sufficient to any the could indicate proof of a former connection be- ostracod fauna of these islands.

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BAHAMAS CAICOS ISLANDS where each of less & the species is more or com-

I: 47 (Table nrs. to 54) mon has been calculated. For most species the

These in ostracod islands are very poor species, Lesser Antilles, except for the Virgin Islands,

probably because they have been inundated by but including the Peninsula de Paraguana,

seawater 65000 used reference some years ago (Pregill & were as area.

As as some 17000 In the the Olson, 1981). recent years Virgin Islands, Haiti, Cayman

the Great wells formed the ago Bahama Bank must have been a Islands and the Bahamas,

which single emergent land mass because of a sea level greatest part of the stations sampled,

of about 120 Later the land makes it the ostracod drop m . on, mass impossible to compare

was split up in five major island complexes, and fauna of different habitats for these islands.

small land in 16 at present only patches remain, Cypris subglobosa was only found epigean

which still further are receding (Pregill & Olson, habitats on Curaçao, Bonaire, Barbados,

Marie and 1981). Galante Guadeloupe, and only once

coral in well. In Puerto Rico it found in Only wells, water holes in rock, one a was two

cave and a bore hole yielded ostracods in the epigean habitats, and in Haiti in a polluted

islands listed in table I under the 47 well. is clear that this is nrs. to 54. open It species truly

The most common species in the wells of these epigean, which could also be the reason why it

islands are Cypretta spp., Cypridopsis sp., Physo- was almost never found in the Virgin Islands,

and Strandesia and the islands of since in these cypria sp. longula. Neocypridopsis in- west these,

audita found in wells In- was on Mayaguana, islands practically no epigean stations have

agua, Crooked Island, Eleuthera, and in a cave been sampled. The same holds true for all

on Inagua. Heterocypris punctata was encountered Hemicypris species (Broodbakker, 1983b) and

twice meridana on South Caicos, Cypris decaryi once on Tanycypris (see: Broodbakker, 1984b).

less Eleuthera, Darwinula sp. once on Mayaguana, Cypris decaryi was found somewhat fre-

and Chlamydotheca unispinosa twice on Providen- quently in the wells of Bonaire and Curaçao

and Crooked in ciales, twice on Island. than epigean habitats, but this difference was

Most of the species mentioned are commonly not statistically significant.

found in pools and other epigean habitats in Heterocypris margaritae was found more often in

West other islands of the Indies, and are wells, but not significantly, while H. antillensis

therefore much in the found in probably more common was significantly more often epigean

Bahamas than the data hand would habitats also: at suggest. (see Broodbakker, 1983a).

in inaudita be The only pool sampled Eleuthera yielded Neocypridopsis seems to found more

and often in wells but this difference Stenocypris major, Cypretta sp. Neocypridopsis (and caves),

inaudita. could not be tested since only few samples

available. yielding this species are However, it

was also encountered in two covered wells in

THE OSTRACODS IN WELLS Haiti, in sixteen wells and water holes in coral

rock and in in the and in one cave, Bahamas,

The relative abundance of the most common caves in Cuba. It seems therefore to have its ostracod species in the Lesser Antilles (except main distribution in hypogean habitats.

for the Virgin Islands) in wells, becomes clear Stenocypris major and Potamocypris hummelincki

in each from table II, which for species the were found significantly more often in epigean abundance in wells and in epigean lentic habitats, nothwithstanding the fact that habitats found in 18 wells in Haiti like pools, ponds, basins, cisterns etc., Stenocypris major was is compared with the expected abundance in (table III). these two of habitats if the would and types species Physocypria affinis, Cypridopsis sp., have As total no preferences. reference the Chlamydotheca barbadensis seem to have no number of habitats for concerned in the islands preference either wells or epigean habitats.

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TABLE II

Number of ostracod samples from lentic habitats and from wells, compared with the total number ofostracod samples in

the respective habitats and islands, with percentages and G-values (ref. = reference).

Number of pools, Number of wells, G-value

ponds, ditches, etc. sink- and bore holes

Cypris decaryi 21 (78%) 6 (22%) 1.34 Réf.: Curaçao, Bonaire 88 (68%) 42 (32%)

Heterocypris margaritae 23 (55%) 19 (45%) 1.85 Réf.: Islands 3 to 12, and Paraguana 126 (65%) 68 (35%)

Heterocypris antillensis 36 (80%) 9 (20%) 4.61* Réf.: Islands 3 to 6, 25, 31 127 (65%) 67 (35%)

Neocypridopsis inaudita 2 (20%) 8 (80%)

Réf.: Islands 5, 6, 28, 30 to 32 69 (54%) 59 (46%)

Stenocypris major 60 (87%) 9 (13%) 23.30** Potamocypris hummelincki 55 (82%) 12 (18%) 14.36**

16 48 33.64** Cypretta sp. (25%) (75%)

Physocypria affinis 31 (65%) 17 (35%) 0.31

25 13 Cypridopsis sp. (66%) (34%) 0.42

Chlamydotheca barbadensis 15 (58%) 11 (42%) 0.10

Réf.: Islands 3 to 33 and Paraguana 176 (61%) 114 (39%)

TABLE III

Number of ostracod samples, with percentages and G- values, from wells in Haiti and the Bahamas

(fur further explanation see text).

HAITI In all wells Cypretta sp. Chlamydotheca unispinosa

Accompanying Ostracoda (ri(2) -98) («(1) - 39) G (n = 37) G

Clamydotheca unispinosa 37 (38%)(«,) 16 (15%) 9.49**

Strandesia stocki 39 (40%) 13 (33%) 0.69 12 (32%) 0.86

Strandesia longula 33 (34%) 22 (56%) 8.42** 6 (16%) 5.72*

Stenocypris major 18 (18%) 8 (21%) 0.12 3 (8%) 3.89*

12 6 0.33 5 — Physocypria sp. (12%) (15%) (14%)

7 3 - 3 — Cypridopsis sp. (7%) (8%) (8%)

In wells BAHAMAS all Physocypria sp. Cypridopsis sp.

= G = Accompanying Ostracoda (* 81) (n = 45) (n 38) G

38 19 0.40 Cypridopsis sp. (47%) (42%)

Strandesia longula 38 (47%) 16 (36%) 2.37 21 (55%) 1.06

10 0.58 10 0.01 Cypretta sp. 22 (27%) (22%) (26%)

Neocypridopsis inaudita 16 (20%) 7 (16%) 0.53 7 (18%) 0.04

* than ** Significant at better 5% level (one-tailed). Significant at better than 1% level (one-tailed).

The first two are also in wells this be abundant in the genera common genus seem to most of the Virgin Islands, Haiti, and the Bahamas. wells of practically all West Indian islands.

was found In the wells of the Islands Cypretta sp. significantly more Virgin Cypretta sp.

in Some often wells. or perhaps all species of is the most common species, being present in 29

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out of 39 wells and water holes in coral rock from their overall distribution in the wells of the

followed 12 sa.npled, by Physocypria sp. (in islands considered. This was already calculated

and 7 Most wells) Cypridopsis sp. (in wells). for Strandesia longula and S. stocki by Broodbak-

other species encountered were found only ker (1984a), S. longula being found significantly

for Strandesia which was often in the of and once, except longula more presence Cypretta sp., found in four and inaudita wells, Neocypridopsis significantly less often in the presence of

and Neocypridopsis sp., wich were found in three Chlamydotheca unispinosa, than expected.

shallow water holes in coral rock in found much less fre- Anegada. Likewise, Cypretta sp. was

In the wells of Haiti and the Bahamas the quently in the presence of Chl. unispinosa than

species listed in table III were most abundant, expected (table III). The other species in the

the Strandesia and including two species already wells of Haiti the Bahamas were not found

discussed Broodbakker in by (1984a). significantly more or significantly less often

is of the Chlamydotheca unispinosa one most each other's presence, than expected.

common species in the wells of Haiti, while it is An attempt was made to compare the abun-

in practically absent the wells of the Bahamas. dance of the most common species in different

in in This species was scarce the epigean samples environmental classes wells, with that ex-

for from the other islands, while it is also known pected, as was done species of the genus

from in the Broodbakker The fac- some states U.S.A., from Yucatan, Heterocypris by (1983a).

from Brazil and and from Hawaii chosen of Argentina tors were: light conditions, position

(Kotzian, 1974). The reason for its scattered the water table, and water depth. For the

distribution and its absence in wells in other species in the wells of the Lesser Antilles not

in the Lesser data since islands, especially Antilles, re- enough were available, most species

It could that there encountered in less 15 wells mains unsolved. be is some were than only (cf.

and form of competition between Chl. unispinosa and table II), because the factors tested were

measured indicated. Chl. barbadensis, this last species being especially not always or Testing was

in of the Lesser Antilles. for in the common some possible only Cypretta sp. Virgin

Cypridopsis sp. is especially common in wells Islands, and for the most abundant species

of the Bahamas, and much less in wells of Haiti. found in Haiti and the Bahamas (cf. table III).

Most of these have the of No differences found in Cypridopsis sp. shape significant were

C. smaller and do distribution of far vidua, but they are not possess these species as as light condi-

the characteristic dark the markings on tions, position of the water table and water

which could be due to fixation in for- concerned. The data and G-values carapace, depth are

malin. It is quite possible that most of the calculated are therefore not listed in this article.

Cypridopsis species encountered in the West In-

dies have to be assigned to C. vidua. Something

similar holds true for Cypretta species, the most ACCOMPANYING FAUNA IN WELLS

common species in wells of the West Indies. It

could of hadziid be that most of them belong to C. sarsi, a Especially the joint occurrence am-

which described and ostracods is studied. The species was poorly by Brady phipods genus

(1902) from St. Thomas (Virgin Islands). Metaniphargus belongs to the larger hypogean

of which I did not the of the hadziid Stock Physocypria sp., study amphipods group. (1983)

is in wells of the made it that be ef- taxonomy, most common likely hadziid amphipods can

but found in other Bahamas, it was also most fective predators of small crustaceans, like

West Indian islands. cyclopid copepods and the small-sized crusta-

for the Bahamas and for of the Only as a whole, ceans hypogean order Thermosbaenacea.

it calculate if On in 45 of Haiti, was possible to some species Curaçao Metaniphargus was found

found often less often than the wells are more or expected 54 sampled. Cypridopsis sp. was found

in each other's in wells than in 9 and in 4 of the 11 wells presence expected Cypretta sp. genuine

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yielding ostracods, Cypridopsis in 5, and Cypretta of these samples Neocypridopsis inaudita was

in 2 wells together with Metaniphargus. This found together with Metaniphargus. In the other

and and could mean that Cypretta especially sample Perissocytheridea cribrosa Metaniphargus

Cypridopsis are better fitted against prédation by were found.

other Un- encountered in Metaniphargus than ostracod species. Hadziid amphipods were also

fortunately, 7 ostracod samples from stations St. Croix, Jamaica and Haiti, but not in the

lost. and yielding Metaniphargus have got Further- Virgin Islands the Bahamas. On St. Croix

is in of twelve wells more, Cypridopsis common not only wells, only one out sampled yielded

but also in pools, ponds and similar habitats on ostracods, belonging to the genus Physocypria.

in Curaçao, while it is absent on Bonaire and most Hadziid amphipods were only found one

It other well. Venezuelan islands. was found only once on

If is Haiti data Aruba and Margarita. Cypridopsis sp. more Only from enough are present to

resistant the the of the most against prédation by Metaniphargus, compare statistically presence

habitats with the of hadziid question of its abundance in epigean common ostracods presence

is done in IV. and of its absence in the neighbouring islands amphipods, as table Neither of

the in remains unsolved. species listed was found significantly less

On Aruba 7 out of 41 wells the of hadziid only genuine presence amphipods. Only

less in sampled yielded ostracods. Fourteen wells Strandesia longula was found significantly

but no ostracods. In one the of hadziid while S. yielded Metaniphargus presence amphipods,

well found with stocki Metaniphargus was together did not seem to be influenced, as was

is that Aruba Cypretta. It strange on Heterocypris calculated in Broodbakker (1984a: table II).

well brackish the which have margaritae, a species adapted to Cypretta spp. are only species

wells (Broodbakker, 1983a), is only found in been found in large enough numbers to com-

brooklets and and their pools, springs, only present pare presence with that of Metaniphargus

in It be its for all one pool on Curaçao. could that sampled Lesser Antilles together. These

absence in of these is less in the wells islands caused by species were not found significantly the

the abundance of there. of than Metaniphargus presence Metaniphargus expected (table

On 5 wells and 4 stations Guadeloupe pump IV). have been found in The of ostracods in the wells sampled. Metaniphargus was presence was

of the wells. also with that of other faunal 4 pump samples and in 2 In only compared groups.

In the one other well a juvenile of Stenocypris major was Lesser Antilles, only Cypretta was found

found. Since only few wells have been sampled in the wells often enough to compare its abun-

of of the on Guadeloupe it is not evident ifthe absence dance with that other faunal groups, but ostracods is caused by the presence of differences found were not statistically signifi-

Metaniphargus. cant.

15 been For On Antigua wells have sampled of Haiti and the Bahamas joint occurrence which only 1 yielded Metaniphargus and 6 others could be statistically tested for the most com-

is shown in table IV. yielded ostracods. mon species, as However,

2 with On Barbuda caves and 13 wells have been no significant differences the expected joint

be sampled. The two caves and six of the wells occurrence could found, except for yielded Metaniphargus. Only two other wells Neocypridopsis inaudita in the Bahamas, which

fact than yielded ostracods. The that Metaniphargus is was found significantly more often ex-

this island and that wells in the of common on only two pected presence Oligochaeta.

could mutual yielded ostracods, indicate some influence. However, on St. Eustatius 4 out only EPIGEAN VERSUS HYPOGEAN of the 17 wells sampled while yielded ostracods, OSTRACODA this island. Metaniphargus is not found on

of 11 natural crevices and of wells in West On Anguilla 9 out Most the the Indies are coloniz- wells sampled yielded ostracods. In all but one ed by epigean animals, especially Cyclopidae,

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TABLE IV

of other faunal with and from wells in most Lesser Number samples (ostracods versus elements), percentages G-values,

Antilles, the Virgin Islands, Haiti and the Bahamas (for further explanation see text).

ISLANDS 3 TO 33 In all wells In wells VIRGIN ISLANDS In all wells In wells

AND yielding yielding yielding yielding

PARAGUANA Ostracoda Ostracoda Cypretta sp. Cypretta sp.

Accompanying fauna (n = 129) (n = 43) G Accompanying fauna (« = 39) (n = 29) G

Hadziid amphipods 17 (13%) 4 (9%) 0.62 Thermosbaenacea 12 (31%) 10 (34%) 0.18 Cyclopidae 55 (43%) 24 (56%) 3.01 Cyclopidae 24 (62%) 16 (55%) 0.49

Mosquito larvae 20 (16%) 9 (21%) 0.39 Mosquito larvae 6 (15%) 5 (17%) —

Oligochaeta 17 (13%) 7 (16%) 0.34 Oligochaeta 10 (26%) 9 (31%) 0.42

HAITI In all wells In wells yielding

yielding Cypretta sp. Chlamydotheca Stenocypris Physocypria sp.

Ostracoda unispinosa major

= Accompanying fauna (n = 98) (n = 39) G (n = 37) G («=18) G (» 12)

Cyclopidae 81 (84%) 29 (74%) 1.68 34 (92%) 2.64 13 (72%) — 10 (83%)

Oligochaeta 55 (57%) 22 (56%) 0.01 22 (59%) 0.17 7 (39%) 2.15 7 (58%)

Cladocera/Phyllopoda 20 (21%) 5 (13%) 1.54 10 (27%) 0.93 — 5 (42%)

Thermosbaenacea 26 (27%) 11 (28%) 0.06 11 (30%) 0.20 7 (39%) 1.30 2 (17%)

Hadziid amphipods 18(19%) 4(10%) 1.97 10 (27%) 1.67 3 (17%) — 2(17%)

BAHAMAS In all wells In wells yielding

yielding Physocypria sp. Cypridopsis sp. Cypretta sp. Neocypridopsis

Ostracoda inaudita

Accompanying fauna (n = 81) (n = 45) G (n = 38) G (n = 22) G (fi-16) G

Cyclopidae 43 (53%) 24 (53%) 0.01 25 (66%) 2.52 13 (59%) 0.32 7 (44%) 0.56 Oligochaeta 24 (30%) 11 (24%) 0.60 13 (34%) 0.37 5 (23%) 0.53 10 (63%) 7.15»*

— — Cladocera/Phyllodopa 10 (12%) 5 (11%) 0.07 5 (13%) — 5 (23%) 2 (13%)

Hyalella sp.

— — (Amphipoda) 12 (15%) 8 (18%) 0.30 6 (16%) — 3 (14%) 1 (6%)

** Significant at better than 1% level (one-tailed).

insect larvae and Gastropoda. Since most of the 1983b), Tanycypris meridana (see Broodbakker, ostracod wells often and in in species colonizing are en- 1984b), Cypris subglobosa (only one well countered in large numbers, it is clear that they Haiti). For species of the genera Heterocypris

in and is in can ecologically adapt to life wells. However, Strandesia, this aspect discussed Brood-

all be of not species seem to capable colonizing bakker (1983a, 1984a).

Ostracod in wells is low. In of wells, or they are competitively excluded by diversity most

all the other ostracod species. Compared with other samples only one or two species are present. ostracod When the well is and it species, Cypretta spp., Cypridopsis spp., more open shallow, and often to resemble habitats and Physocypria spp., are most en- begins epigean two or

three countered in wells. Many epigean species were species can be encountered. In true

found in like six never or almost never genuine wells, epigean habitats, pools, up to species

all be e.g. Hemicypris species (Broodbakker, can present in a single sample. Furthermore,

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in deal different seasons other species combinations on hypogean Crustacea (e.g. Stock, 1977,

can be found. The species found in pools in- 1980).

clude those in wells. This is in Ostracoda with found agreement However, no hypogean

with the of direct less direct marine findings Danielopol (1981), who or ancestors were found

records likewise low diversities in wells of in inland from any groundwater sample any

Euboea (Greece). Danielopol grouped his wells island. Only one epigean brackish-water

and in six types (I to VI) from more epigean ostracod, Perissocytheridea cribrosa, has colonized

wells the polluted to covered, unpolluted sheltering some of more brackish wells, but it did not

evolve into hypogean animals. The epigean animals live in a hypogean species either.

Euboea in the uncovered wells with It polluted seems that there are no ostracod species for

In low number real waters. his case a (two) of which it was possible to overcome the barrier

hypogean ostracod species was found, while in from salt to fresh water. Most of the interstitial

the West Indian islands, except for Cuba and ostracods living in the vicinity of the sea in the

animals no were en- are still in 18°/ or Hispaniola, hypogean Neotropics living 0o chlorinity

countered all. All if in- close that at wells, colonized, were to (see e.g. Hartmann, 1974).

vaded by epigean, euryoecious species. Species As Danielopol (1981) described for Euboea,

to like wells behave like and colonized belonging genera Heterocypris, Hemicypris islands, are by

and in Strandesia can display much variability both hypogean and epigean species. However,

related in the of length, probably to ecological factors, case the ostracods of all smaller West

but be stable Indian seem to genetically (Broodbakker, islands, the balance seems to have turn-

It that in less 1983a-b, 1984a). appears some species ed favour of the more or cosmopolitan

in but the can ecologically adapt to life wells, epigean species.

in populations living wells do not differ mor-

phologically from populations living in epigean

habitats. ZOOGEOGRAPHY

Apparently, the freshwater ostracods living

in the Lesser Caribbean Islands and in the There is much the controversy concerning unable evolve into Bahamas, were to hypogean origin of the West Indian islands and of their

This is due fauna. Most of the with species. probably to their ecologie opinions are concerned

plasticity, combined with their large potentials the "Dispersal" and the "Vicariance Model"

for passive dispersal, especially of their eggs, of origin.

from island to island, and from pool to pool or The Dispersal Model has gained acceptance

well. there is of the works of Probably a large amount gene chiefly through Darlington (e.g.

flow in the sexual species, and a strong genetic 1957). It assumed that the Antillean islands

in stability the parthenogenetic ones. have originated independently and have been

Other which could colonize the of groups populated by transmarine migrants, by way

the marine groundwater are interstitial, active or passive dispersal.

and marine, brackish-water species, especially The most cited propagators of the Vicariance

in Antillean the islands which are slowly but Model are Rosen (1976), for the Caribbean,

since thousands of constantly rising many and Croizat (e.g. 1976) for the general theory.

or even millions of In this Rosen the Antillean islands years, years. way assumes to be

marine animals and may have "stranded", fragments of a proto-Antillean contiguous area

gradually adapted to nonmarine conditions, forming the connection between North and

of especially by way the partly or wholly South America in the Late Mesozoic. This brackish groundwater of the semi-arid islands, proto-Antillean plate has moved in north-

which are in surface This direction and has in poor water. eastern split up eventually

zoogeographic model, the Regression Model, is the presently existing islands and plates, in the advocated Stock in for by many papers, a great Late Tertiary.

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and Hedges (1982) has recently compiled new Sohn & Kornicker (1979) froze freeze-

evidence of vicariant of the geologic a origin dried eggs of Heterocypris incongruens (Ramdohr,

and 0.01 mb Greater Antilles, including the Virgin Islands 1808) at -25 to -30°C at vacuum.

which situated the Puerto Rican bank. still hatched immersion in are on These eggs after

He remains in doubt about the of the for ostracods origin water. In this way it is possible

other Lesser Antilles. with such resistant be distributed eggs to widely

With the Vicariance Model of because of respect to of circumglobal transport eggs by

of the Caribbean data origin islands, many are high altitude winds at low temperature and

derived from flatworms Or- (e.g. Ball, 1971), pressure.

thocladiinae (Chironomidae) (Saether, 1981), Some species, especially some Candoninae,

Trichoptera (Ross, 1967; Botosaneanu, 1979, can enter in a torpid state when the situation

only for the Greater Antilles), freshwater fauna becomes unfavourable (Delorme & Donald,

in general (Harrison & Rankin, 1976), and 1969; McLay, 1978). Especially for species with

freshwater fishes with brood this (Rosen, 1976). Likewise, less resistant eggs or care, seems

Rosen derived data from survive less favourable condi- (1976) epigean a good way to

and be freshwater shrimps of the family Atyidae tions, or to dispersed.

the hypogean Thyphlatya which were There are several of genus , possible ways passive

refuted Stock for ostracods and their Most of by (1981). dispersal eggs.

In favour of the Model of of these listed in Dispersal origin are McKenzie (1971):

the Caribbean fauna indications many are (1) Ovigerous females or nonovigerous par-

derived from the distribution of their be reptiles (e.g. thenogenetic species, or eggs, can car-

bats ried and other in mud Müller, 1973), (Baker & Genoways, 1978), across oceans barriers

insects like butterflies (Brown, 1978), and caked to the feathers and feet of migrant or

freshwater Gastropoda (Baker, 1945). stormdriven birds, especially waterfowl (Sand-

Recent discussions both models concerning berg, 1964; De Deckker, 1977). There is a

of origin of fauna, can be found in Nelson & migration route of wading birds and ducks from

and Nelson Platnick Rosen (1981) & (1981). Venezuela over Trinidad, Grenada, up north,

which and further St. Puerto Freshwater organisms accomplish over Kitts, St. Thomas,

life their entire cycle in fresh water often have Rico, Haiti, Jamaica, to Cuba. This dispersal

limited dispersal abilities, and could therefore route is postulated to be the essential way for

be excellent objects for testing the different gastropod species of the family Planorbidae to

models. Freshwater ostracods with reach the Antillean islands partly agree (Baker, 1945).

these requirements; however, most of them Over shorter distances even insect carriers

have extensive abilities for passive dispersal. could be effective for dispersal of ostracods

Passive dispersal of ostracods can take place by (Fryer, 1953).

of their of adult animals. way eggs, or (2) Dusty, moisty and even dry wind or gales

The of freshwater of desiccation eggs many species, could be a way dispersal of resis-

the resistant and of adults in especially Cypridinae, are very tant eggs, perhaps even a torpid

all of against types physico-mechanical stress. state (McKenzie, 1971).

first of all desiccation resistant They are by way (3) Eggs or even complete specimens could be of a protective double wall (Van Morkhoven, dispersed via the alimentary canal of birds

Sars raised Proctor the 1962). (e.g. 1889, 1901) many (Proctor, 1964; & Malone, 1965), or

in dried species of Cypridinae cultures from guts of fishes in case of marine ostracods

mud in various continents sampled of the (Kornicker & Sohn, 1971).

world. A case is known in which a Introduction of bar- Cypris species (4) by man, by way water

was raised from dried mud sent from the Near rels or ballast tanks on ships, or mud on boots

East Great 24 30 this hard to Britain, some to years or ships. Unfortunately, is to prove in

Morkhoven, most It is of in the previously (Van 1962). cases. a proven way dispersal

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case of rice-introduction in European countries tually encountered, as became evident in the

(e.g. Petkovski, 1964). chapter on distribution of the species. When the

It will be clear that most freshwater number of species is plotted against the number

ostracods, except for the hypogean species, of stations yielding ostracods in each island, it is

limited value for fine scale been have a evident that the more samples have taken,

Most the ostracods the zoogeography. of freshwater more species are found (fig. 4).

effect: One encountered in the West Indies disperse There can be two reasons for this

relatively easily and are therefore widely is the fact that when an island is larger, more

whilst in distributed. Many species are parthenogenetic, stations to be sampled are available, a

habitats which even more facilitates introduction after small island only few freshwater are

passive dispersal. McKenzie (1971) states that available and only few species are present. The

of be that the number of species the genera Heterocypris, Cypridopsis, other reason can species

and encountered is correlated with number Chlamydotheca, Cypretta, are the most com- truly the

in islands. This is of stations and mon statement confirmed by sampled. Curaçao St. Martin

The the results of the present study. genera yielded the same numberof species, Aruba one

and have be St. Martin Strandesia, Hemicypris Physocypria to less and Bonaire one more. occupies

added to this list. This could indicate that these approximately one-fifth of the size of Curaçao

in have the best and one-third of the size of Bonaire. Aruba is genera particular dispersal

St. abilities. However, the genera Heterocypris and about twice the size of Martin. According to

the and should Cypridopsis are common throughout world, their size the theory, Curaçao yield

while the Bonaire and Aruba and St. genera Chlamydotheca, Cypretta, most species, less,

and Martin the least. and Cuba Strandesia, Hemicypris are common circum- Haiti yielded 20

tropical genera. Most of the species en- and 19 species, respectively, whereas in Haiti

be stable and did three times ostracods. countered seem to genetically as many samples yielded

evolve Barbados and not into endemic species on the smaller Margarita yielded the same

islands. number of species, and the number of samples

This that the of the the but Barbados is far from means greatest part was same, away

is situated the freshwater ostracods of the West Indies must the continent and Margarita on

have arrived the of Venezuela. Barbados is also on islands by passive dispersal, continental shelf

for in Haiti and In the except perhaps some species much smaller than Margarita (fig. 1).

which have in these islands where less than 14 Cuba, may originated samples yielded

from 10 islands or in the former Proto-Antilles. ostracods, a variation 1 to species was

Therefore it should be possible to test some of found. the theories regarding the expected number of Furthermore, other factors probably con-

in the number of island. species islands. MacArthur & Wilson (1967) tribute to species in an calculated relations between the number of Volcanic islands have a structure different from species present and island area, and distance to that of calcareous islands, like greater altitudes,

and the mainland, or to the centre of dispersal. thus more precipitation more freshwater

extended Many authors used this model or the habitats, although often of a torrential type, model with other it to and therefore than semi- variables, applying many can yield more species

likewise in West arid This the animal groups, the Indies. islands. could explain large

the small St. However, only some islands were sampled number of species found on island

other islands and in with the number of extensively, barely, many Martin, as compared

and islands only wells underflows of running species on the largely calcareous islands, waters have been sampled. Furthermore, only Aruba, Bonaire, and Curaçao. few species have been found in most of the The conditions in the more extensively

make islands, and it is highly probable that in most sampled islands are too different to a islands those in more species are present than ac- statistical comparison possible. Furthermore,

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Number of ostracod number of in the various islands Fig. 4. species encountered, versus stations sampled, and regions.

and in than indicated Only islands regions which more six species were found are by name.

have underflows have been Cuba mostly caves been sampled, in Haiti of running waters sam-

mostly wells, and in the Netherlands Antilles all pled.

kinds of habitats. The few differences found do No consistent differences could be found

indicate either between volcanic and the not a species/area regression or a rela- the calcareous

tionship to the distance from the mainland. islands. This is partly due to the fact that the

far volcanic islands were sampled less extensive-

ly-

All species in the Lesser Antilles, the

ZOOGEOGRAPHICALCONCLUSIONS and of the from Bahamas, many species Haiti

and Cuba must have arrived there by passive

It appears that the different Lesser Antilles have dispersal. Some of them probably arrived from

the ostracod with minor South others from the southern essentially same fauna, America, part

differences in like the abundance of of North and from some cases, America, maybe some a

Cypridopsis sp. on Curaçao, the occurrence of proto-Antillean centre of origin.

barbadensis the Hemicypris on Barbados, presence In Cuba one species, and in Haiti three

of Gorda and of ostracods have been found in the Neocypridopsis sp. on Virgin species

and the scattered distribution of the Anegada only, groundwater, belonging to new genus

other many species. However, most of these dif- Caribecandona Broodbakker, 1983c. This genus,

ferences be insuffi- which to can probably explained by seems be very old, could have

cient of most islands far in the Proto-Antilles. The sampling as as originated genus is ostracods are concerned. The only islands hav- divided into two subgenera, one endemic to

and ing a clearly different partly specific fauna Cuba, the other to Haiti. This could be due to

are Hispaniola and Cuba, which is probably vicariance, viz. the separation of Cuba and also due which could 5 to the fact that these islands have a Haiti, have taken place between

and extended to 20 million A large ecological diversity physio- years ago (Hedges, 1982).

be bar- graphy. Furthermore, there seems to a relatively recent separation of Cuba and Haiti

rier for some between Puerto Rico and could also the of Strandesia species explain presence

but this Hispaniola, could also be a result of cavernicola, a species which is postulated to be

because of in selective sampling, especially west hypogean, caves in Cuba, and in one cave in

this and Haiti. boundary practically only wells, caves,

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Another is the of Neo- Greater Antilles. This is in contrast with the striking fact presence

in Cuba. This of Rosen and conform the cypridopsis mexicana in many caves opinion (1975) opin-

species has been described by Furtos (1938) ion of Stock (1981).

in Mexico. It is found In which is from a cave Yucatan, Jamaica, postulated by many

from and authors be only in hypogean habitats Cuba, to a former part of a geotectonical therefore probably has limited dispersal plate independent of that of the Proto-Antilles

abilities. This would mean that there must have (see review in Hedges, 1982 and Stock, 1983b), been a connection between Yucatan and Cuba. only few stations yielded ostracods. It remains

According to the review of Hedges (1982), the therefore uncertain whether this island

model of for least the ostracod and if most probable origin at possesses any hypogean fauna,

Greater Antilles, is the one of Dickinson & so, whether it is different from that in the

Coney (1980), in which the Proto-Greater An- Greater Antilles.

between tilles, a magmatic arc that emerged

South and North America, has moved north- eastward, and split to give the Greater Antilles. ACKNOWLEDGEMENTS

have been For a long period this arc must in Dr. D. L. is thanked for his his Danielopol support, contact with Yucatan. However, the time of stimulating enthusiasm, his contributions in the form of from Yucatan is from 40 separation postulated of and and copies interesting papers discussions, especially 120 million B.P. The the I in the Institut Lim- to years genus Cypridopsis hospitality could always enjoy für

Mondsee, of the Osterreichische is believed to be 37 million years old, according nologie, Abteilung Akademie der Wissenschaften. to data of the fossil record (Van Morkhoven, Prof. Dr. J. H. Stock, coordinator of the Amsterdam This that 1962). would mean Neocypridopsis Expeditions and of this project, is thanked for reading the mexicana be old in order to must a very species sometimes drafts of all articles many, hardly readible, my

its in Cuba as well as in onthe explain presence ostracods ofthe West Indies, and for the opportuni-

he for me to write this thesis. Yucatan, by a vicariance event. ty provided Dr. L. Botosaneanu and Drs. F. F. J. M. Pieters are Unfortunately, no other clearly hypogean thanked for scrutinizing my articles. the species belonging to Candoninae were Dr. P. Wagenaar Hummelinck, Dr. L. Botosaneanu found in the caves and cenotes of Yucatan. and Dr. T. Orghidan are thanked for placing their samples

Most of the encountered Furtos species by at my disposal. This study has been financed by the Netherlands Foun- (1936) in Yucatan seem to be epigean. Some of dation for the Advancement of Tropical Research them have also been found in the Caribbean (WOTRO), The Hague. The fieldwork ofthe Amsterdam islands. her only Neocypridopsis Among species been from Expeditions has supported by grants WOTRO, mexicana and N. yucatanensis, both belonging to the Beijerinck-Popping Fonds (Amsterdam), the Treub the smallest freshwater ostracods of the world Maatschappij (Utrecht), the Amsterdamse Universiteits

0.40 Vereniging and the Fonds Landbouw- (0.35 to mm), are possibly hypogean. (Amsterdam), hogeschool (Wageningen). Only few stations in Puerto Rico, which could have been another part of the Proto-

Antilles, yielded ostracods. The Virgin Islands,

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Received: 4 January 1984

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