A Checklist of the Beetles of Cuba with Data on Distributions and ...

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22 Arthropods <strong>of</strong> Florida and Neighboring Land Areas, Vol. 18 Hispaniola, Jamaica, Caymans) and near continental land <strong>of</strong> North America (Florida) and Central America (Yucatan peninsula ). Most study <strong>of</strong> <strong>the</strong> patterns and processes in Caribbean biogeography have been based on vertebrates and plants (Woods 1989). Only recently have insects been used to help understand <strong>the</strong> history <strong>of</strong> <strong>the</strong> Caribbean, and <strong>the</strong> impact <strong>of</strong> <strong>the</strong> Caribbean on <strong>the</strong> evolution <strong>of</strong> insects. Examples <strong>of</strong> biogeographic analyses <strong>of</strong> Caribbean insect groups are butterflies (Millere and Miller 1989; Scott 1972), lygaeid bugs (Slater 1988), auchenorhynchous bugs (Ramos 1988), scaritine and platynine carabid beetles (Nichols 1988, Liebherr 1988), caddis flies (Hamilton 1988), drosophilid flies (Grimaldi 1988), ants (Wilson 1988) and sweat bees (Eichwort 1988). Genaro and Tejuca (2001) provide a discussion <strong>of</strong> patterns in <strong>the</strong> <strong>Cuba</strong>n insect fauna in general. A variety <strong>of</strong> biogeographic patterns exists beyond those noted by Darlington (1957) for vertebrates. There are no fossil beetles to help in a reconstruction <strong>of</strong> <strong>the</strong>ir history in <strong>Cuba</strong>. Only a recent dytiscid and a scarab have been recovered from asphalt tar seeps in Matanzas Province (Iturralde- Vinent et al. 2000; Valdés 1999). Biogeographic History <strong>of</strong> <strong>Cuba</strong> The biogeographic history <strong>of</strong> <strong>Cuba</strong> has long stimulated interest, speculation, and investigation. The basics <strong>of</strong> <strong>Cuba</strong>n zoogeography, <strong>with</strong> data drawn from <strong>the</strong> vertebrate animals, were summarized by Darlington (1957), who categorized <strong>the</strong> West Indies as a “fringing archipelago.” The vertebrate fauna is unbalanced in its diversity. The absence <strong>of</strong> strictly fresh-water fishes, <strong>the</strong> limited diversity <strong>of</strong> amphibians and reptiles, and <strong>the</strong> virtual absence <strong>of</strong> land mammals is evidence that <strong>Cuba</strong> has always been isolated by oceanic water gaps, just as is <strong>the</strong> case for <strong>the</strong> rest <strong>of</strong> <strong>the</strong> Greater Antilles (Hedges 2001). There is also a distributional structure or pattern in <strong>the</strong> vertebrate fauna. There is a primary pattern in relict-endemic vertebrate groups (<strong>with</strong> small distributional ranges) which shows an eastward diminution in overall species diversity, suggesting colonization primarily from <strong>the</strong> Yucatan Peninsula <strong>of</strong> Mexico directly into <strong>Cuba</strong>. There is a secondary pattern suggesting dispersal to <strong>Cuba</strong> through Jamaica. Some <strong>of</strong> <strong>the</strong>se groups <strong>the</strong>n moved from <strong>Cuba</strong> into <strong>the</strong> islands fur<strong>the</strong>r to <strong>the</strong> east (Hispaniola and Puerto Rico). A full understanding <strong>of</strong> <strong>the</strong> historical development <strong>of</strong> distributional patterns requires some knowledge <strong>of</strong> <strong>the</strong> geological history <strong>of</strong> <strong>Cuba</strong>. Geological History <strong>of</strong> <strong>Cuba</strong> The Greater Antilles (and <strong>Cuba</strong>) were formerly thought to be composed <strong>of</strong> blocks <strong>of</strong> continental rock and to be relatively fixed in <strong>the</strong>ir location through geological time. The islands are composed <strong>of</strong> a complex <strong>of</strong> volcanic rocks and marine limestones. It is now known that <strong>the</strong> islands have had a complex history <strong>of</strong> location, submergence, and emergence. Donnelly (1988, 1989) has summarized <strong>the</strong> modern understanding <strong>of</strong> <strong>the</strong> geological history <strong>of</strong> <strong>the</strong> Caribbean, but <strong>the</strong> Caribbean Basin remains one <strong>of</strong> <strong>the</strong> most controversial areas in <strong>the</strong> world for a reconstruction <strong>of</strong> its past history. The best modern overview, from <strong>the</strong> point <strong>of</strong> biogeography, is that <strong>of</strong> Iturralde-Vinent and MacPhee (1999). These references have been used to construct <strong>the</strong> following summary. <strong>Cuba</strong> seems to be composed mainly <strong>of</strong> a terrane <strong>of</strong> slivers <strong>of</strong> oceanic material stacked by compressive deformation <strong>of</strong> linear masses <strong>of</strong> sufficient thickness to achieve emergence. This seems to be <strong>the</strong> result <strong>of</strong> lateral compression at <strong>the</strong> edge <strong>of</strong> <strong>the</strong> Caribbean tectonic plate as it moved from an eastern Pacific location into a location between <strong>the</strong> North and South American plates in late Cretaceous and early Tertiary (Eocene) times. <strong>Cuba</strong> also seems to include parts <strong>of</strong> <strong>the</strong> Florida-Bahamas continental margin, and possible fragments <strong>of</strong> continental margin derived from <strong>the</strong> south (<strong>the</strong> Chortis block). All <strong>the</strong>se fragments, island arc sediments, and volcanics seem to have been consolidated in <strong>the</strong> late Cretaceous <strong>with</strong> renewed compressive deformation in <strong>the</strong> Eocene. Thus, <strong>Cuba</strong> possibly provides a small exception to <strong>the</strong> strictly non-continental origin <strong>of</strong> <strong>the</strong> Greater Antilles. But <strong>the</strong>re is no evidence for any direct, emergent, continental land connection at any time. However, <strong>the</strong>re is evidence that <strong>the</strong>re were very early but limited land connections to what is now Hispaniola and Puerto Rico, but not Jamaica. After its origin by consolidation, <strong>the</strong> history <strong>of</strong> <strong>Cuba</strong> was probably one <strong>of</strong> isolation from <strong>the</strong> rest <strong>of</strong> <strong>the</strong> Greater Antilles. From 40 Ma (million years ago) to 10 Ma it lay north <strong>of</strong> <strong>the</strong> east-west strikeslip faults that continued to juxtapose different slivers <strong>of</strong> <strong>the</strong> o<strong>the</strong>r islands (especially Hispaniola). <strong>Cuba</strong> is presently composed <strong>of</strong> three connected blocks: Western <strong>Cuba</strong>, West Central <strong>Cuba</strong>, and Eastern <strong>Cuba</strong>. Available evidence suggests that <strong>the</strong>se have existed separately as subaerial land since <strong>the</strong> Eocene-Oligocene boundary, at about 35 Ma, and that <strong>the</strong> three blocks have been joined into <strong>the</strong> present island since <strong>the</strong> mid-Miocene (after 8 Ma).
In summary, <strong>Cuba</strong> has had a complex geological origin and history. It is partly derived from two former continental margins. It has mostly been submerged. It may have had potential terrestrial (or smaller water gap) connections to <strong>the</strong> Yucatan. These geological potentials, in combination <strong>with</strong> <strong>the</strong> zoogeographic data, are consistent <strong>with</strong> a Tertiary emergent history, <strong>with</strong> continued isolation by water gaps, <strong>with</strong> <strong>the</strong> greatest source <strong>of</strong> over-water colonists being Central America (directly <strong>with</strong> Yucatan and via Jamaica from Honduras). The closing <strong>of</strong> <strong>the</strong> Central American (Panamanian) isthmus in <strong>the</strong> Pliocene modified oceanic circulation and formed <strong>the</strong> Gulf Stream, which must have altered overwater colonization processes. Lower glacial sea-levels exposed vastly greater areas <strong>of</strong> land, and diminished <strong>the</strong> size <strong>of</strong> <strong>the</strong> isolating water gaps. The Pleistocene changes <strong>of</strong> climate in <strong>the</strong> Caribbean are poorly understood, but seemingly lower sea levels were coincident <strong>with</strong> increased aridity, and this decreased <strong>the</strong> dispersal and colonization potential <strong>of</strong> mesic terrestrial biotas. References Costa, C. 2000. Estado de conocimineto de los Coleoptera Neotropicales. Pp. 99-114. In: Martín- Piera, F., Morrone, J. J., and Melic, A. (eds). Hacia un proyecto CYTED para el inventario y estimación de la diversidad entomologica en Iberoamérica: PrIBES-2000. M3m: Monografías Tercer Milenio, vol. 1, SEA, Zaragoza, Spain. Darlington, P.J., Jr. 1957. Zoogeography. John Wiley and Sons, New York. Donnelly, T.W. 1988. Geologic constraints on Caribbean biogeography. Pp. 15-37. In: Liebherr, J.K. (ed.). Zoogeography <strong>of</strong> Caribbean Insects. Cornell Univ. Press, Ithaca, New York. Donnelly, T.W. 1989. History <strong>of</strong> marine barriers and terrestrial connections: Caribbean paleogeographic inference from pelagic sediment analysis. Pp 103-118. In: Woods, C.A. (ed.). Biogeography <strong>of</strong> <strong>the</strong> West Indies; Past, Present and Future, Sandhill Crane Press, Gainesville, FL. Eichwort, G.C. 1988. Distribution patterns and biology <strong>of</strong> West Indian sweat bees (Hymenoptera: Halictidae). Pp. 231-253. In: Liebherr, J.K. (ed.). Zoogeography <strong>of</strong> Caribbean Insects. Cornell Univ. Press, Ithaca, New York. Genaro, J.A. Tejuca, A.E. 2001. Patterns <strong>of</strong> endemism and biogeography <strong>of</strong> <strong>Cuba</strong>n insects. Pp. 77-83. In: Woods, C.A. and F.E. Sergile (eds.), A <strong>Checklist</strong> <strong>of</strong> <strong>the</strong> <strong>Beetles</strong> <strong>of</strong> <strong>Cuba</strong> 23 Biogeography <strong>of</strong> <strong>the</strong> West Indies; patterns and perspectives. CRC Press, Boca Raton, FL. Grimaldi, D.A. 1988. Relicts in <strong>the</strong> Drosophilidae (Diptera). Pp. 183-213. In: Liebherr, J.K. (ed.). Zoogeography <strong>of</strong> Caribbean Insects. Cornell Univ. Press, Ithaca, New York. Hamilton, S.W. 1988. Historical biogeography <strong>of</strong> two groups <strong>of</strong> Caribbean Polycentropus (Trichoptera: Polycentropidae). Pp. 153-182. In: Liebherr, J.K. (ed.). Zoogeography <strong>of</strong> Caribbean Insects. Cornell Univ. Press, Ithaca, New York. Hedges, S.B. 2001. Biogeography <strong>of</strong> <strong>the</strong> West Indies: an overview. Pp. 15-33. . In: Woods, C.A. and F.E. Sergile (eds.), Biogeography <strong>of</strong> <strong>the</strong> West Indies; patterns and perspectives. CRC Press, Boca Raton, FL. Iturralde-Vinent, M. A., and MacPhee, R. D. E. 1999. Paleogeography <strong>of</strong> <strong>the</strong> Caribbean Region: implications for Cenozoic biogeography. Bull. American Museum <strong>of</strong> Natural History no. 238, 95 pp. Iturralde-vinent, M. A., MacPhee, R. D. E., Díaz-Franco ,S., Rojas-Consuegra, R., Súarez, W. and Lomba, A.. 2000. Las Breas de San Felipe, a Quaternary fossiliferous asphalt seep near Martí (Matanzas Province, <strong>Cuba</strong>). Caribbean Journal <strong>of</strong> Science 36: 300- 313. Liebherr, J.K. 1988. Biogeographic patterns <strong>of</strong> West Indian Platynus carabid beetles. Pp. 121- 152. In: Liebherr, J.K. (ed.). Zoogeography <strong>of</strong> Caribbean Insects. Cornell Univ. Press, Ithaca, New York. Marske, K.A. and Ivie, M.A. 2003. Beetle fauna <strong>of</strong> <strong>the</strong> United States and Canada. Coleopterists Bulletin 57: 495-503. Miller, L.D. and Miller, J.Y. 1989. The biogeography <strong>of</strong> West Indian butterflies (Lepidoptera: Papilionoidea: Hesperioidea): a vicariance model. Pp. 229-262. In: Woods, C.A. (ed.), Biogeography <strong>of</strong> <strong>the</strong> West Indies: Past, Present and Future. Sandhill Crane Press, Gainesville, FL. Navarrete-Heredia, J. L. and Fierros-López, H. E. 2001. Coleoptera de México: situación actual y perspectivas de estudio. Pp. 1-21. In: Navarete-Heredia, J. L., Fierros-López, J. L., and Burgos-Soloria, A. (eds.). Tópicos sobre Coleoptera de Mexico. Universidad de Guadalajara-Universidad Autónoma del Estado de Morelos, Guadelajara, Mexico. Nichols, S.W. 1988. Kaleidoscopic biogeography <strong>of</strong> West Indian Scaritinae (Coleoptera: Carabidae). Pp. 71-120. In: Liebherr, J.K. (ed.).
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