A Checklist of the Beetles of Cuba with Data on Distributions and ...
A Checklist of the Beetles of Cuba with Data on Distributions and ...
A Checklist of the Beetles of Cuba with Data on Distributions and ...
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22<br />
Arthropods <str<strong>on</strong>g>of</str<strong>on</strong>g> Florida <strong>and</strong> Neighboring L<strong>and</strong> Areas, Vol. 18<br />
Hispaniola, Jamaica, Caymans) <strong>and</strong> near c<strong>on</strong>tinental<br />
l<strong>and</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> North America (Florida) <strong>and</strong> Central<br />
America (Yucatan peninsula ). Most study <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g><br />
patterns <strong>and</strong> processes in Caribbean biogeography<br />
have been based <strong>on</strong> vertebrates <strong>and</strong> plants (Woods<br />
1989). Only recently have insects been used to help<br />
underst<strong>and</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> history <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Caribbean, <strong>and</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g><br />
impact <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Caribbean <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> evoluti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> insects.<br />
Examples <str<strong>on</strong>g>of</str<strong>on</strong>g> biogeographic analyses <str<strong>on</strong>g>of</str<strong>on</strong>g> Caribbean<br />
insect groups are butterflies (Millere <strong>and</strong> Miller<br />
1989; Scott 1972), lygaeid bugs (Slater 1988),<br />
auchenorhynchous bugs (Ramos 1988), scaritine<br />
<strong>and</strong> platynine carabid beetles (Nichols 1988,<br />
Liebherr 1988), caddis flies (Hamilt<strong>on</strong> 1988),<br />
drosophilid flies (Grimaldi 1988), ants (Wils<strong>on</strong> 1988)<br />
<strong>and</strong> sweat bees (Eichwort 1988). Genaro <strong>and</strong> Tejuca<br />
(2001) provide a discussi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> patterns in <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Cuba</str<strong>on</strong>g>n<br />
insect fauna in general.<br />
A variety <str<strong>on</strong>g>of</str<strong>on</strong>g> biogeographic patterns exists bey<strong>on</strong>d<br />
those noted by Darlingt<strong>on</strong> (1957) for vertebrates.<br />
There are no fossil beetles to help in a rec<strong>on</strong>structi<strong>on</strong><br />
<str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>ir history in <str<strong>on</strong>g>Cuba</str<strong>on</strong>g>. Only a recent<br />
dytiscid <strong>and</strong> a scarab have been recovered from<br />
asphalt tar seeps in Matanzas Province (Iturralde-<br />
Vinent et al. 2000; Valdés 1999).<br />
Biogeographic History <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>Cuba</str<strong>on</strong>g><br />
The biogeographic history <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>Cuba</str<strong>on</strong>g> has l<strong>on</strong>g<br />
stimulated interest, speculati<strong>on</strong>, <strong>and</strong> investigati<strong>on</strong>.<br />
The basics <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>Cuba</str<strong>on</strong>g>n zoogeography, <str<strong>on</strong>g>with</str<strong>on</strong>g> data drawn<br />
from <str<strong>on</strong>g>the</str<strong>on</strong>g> vertebrate animals, were summarized by<br />
Darlingt<strong>on</strong> (1957), who categorized <str<strong>on</strong>g>the</str<strong>on</strong>g> West Indies<br />
as a “fringing archipelago.” The vertebrate fauna<br />
is unbalanced in its diversity. The absence <str<strong>on</strong>g>of</str<strong>on</strong>g> strictly<br />
fresh-water fishes, <str<strong>on</strong>g>the</str<strong>on</strong>g> limited diversity <str<strong>on</strong>g>of</str<strong>on</strong>g> amphibians<br />
<strong>and</strong> reptiles, <strong>and</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> virtual absence <str<strong>on</strong>g>of</str<strong>on</strong>g> l<strong>and</strong><br />
mammals is evidence that <str<strong>on</strong>g>Cuba</str<strong>on</strong>g> has always been<br />
isolated by oceanic water gaps, just as is <str<strong>on</strong>g>the</str<strong>on</strong>g> case<br />
for <str<strong>on</strong>g>the</str<strong>on</strong>g> rest <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Greater Antilles (Hedges 2001).<br />
There is also a distributi<strong>on</strong>al structure or pattern<br />
in <str<strong>on</strong>g>the</str<strong>on</strong>g> vertebrate fauna. There is a primary<br />
pattern in relict-endemic vertebrate groups (<str<strong>on</strong>g>with</str<strong>on</strong>g><br />
small distributi<strong>on</strong>al ranges) which shows an eastward<br />
diminuti<strong>on</strong> in overall species diversity, suggesting<br />
col<strong>on</strong>izati<strong>on</strong> primarily from <str<strong>on</strong>g>the</str<strong>on</strong>g> Yucatan<br />
Peninsula <str<strong>on</strong>g>of</str<strong>on</strong>g> Mexico directly into <str<strong>on</strong>g>Cuba</str<strong>on</strong>g>. There is a<br />
sec<strong>on</strong>dary pattern suggesting dispersal to <str<strong>on</strong>g>Cuba</str<strong>on</strong>g><br />
through Jamaica. Some <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>se groups <str<strong>on</strong>g>the</str<strong>on</strong>g>n moved<br />
from <str<strong>on</strong>g>Cuba</str<strong>on</strong>g> into <str<strong>on</strong>g>the</str<strong>on</strong>g> isl<strong>and</strong>s fur<str<strong>on</strong>g>the</str<strong>on</strong>g>r to <str<strong>on</strong>g>the</str<strong>on</strong>g> east<br />
(Hispaniola <strong>and</strong> Puerto Rico). A full underst<strong>and</strong>ing<br />
<str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> historical development <str<strong>on</strong>g>of</str<strong>on</strong>g> distributi<strong>on</strong>al patterns<br />
requires some knowledge <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> geological<br />
history <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>Cuba</str<strong>on</strong>g>.<br />
Geological History <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>Cuba</str<strong>on</strong>g><br />
The Greater Antilles (<strong>and</strong> <str<strong>on</strong>g>Cuba</str<strong>on</strong>g>) were formerly<br />
thought to be composed <str<strong>on</strong>g>of</str<strong>on</strong>g> blocks <str<strong>on</strong>g>of</str<strong>on</strong>g> c<strong>on</strong>tinental rock<br />
<strong>and</strong> to be relatively fixed in <str<strong>on</strong>g>the</str<strong>on</strong>g>ir locati<strong>on</strong> through<br />
geological time. The isl<strong>and</strong>s are composed <str<strong>on</strong>g>of</str<strong>on</strong>g> a complex<br />
<str<strong>on</strong>g>of</str<strong>on</strong>g> volcanic rocks <strong>and</strong> marine limest<strong>on</strong>es. It is<br />
now known that <str<strong>on</strong>g>the</str<strong>on</strong>g> isl<strong>and</strong>s have had a complex<br />
history <str<strong>on</strong>g>of</str<strong>on</strong>g> locati<strong>on</strong>, submergence, <strong>and</strong> emergence.<br />
D<strong>on</strong>nelly (1988, 1989) has summarized <str<strong>on</strong>g>the</str<strong>on</strong>g> modern<br />
underst<strong>and</strong>ing <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> geological history <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g><br />
Caribbean, but <str<strong>on</strong>g>the</str<strong>on</strong>g> Caribbean Basin remains <strong>on</strong>e<br />
<str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> most c<strong>on</strong>troversial areas in <str<strong>on</strong>g>the</str<strong>on</strong>g> world for a<br />
rec<strong>on</strong>structi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> its past history. The best modern<br />
overview, from <str<strong>on</strong>g>the</str<strong>on</strong>g> point <str<strong>on</strong>g>of</str<strong>on</strong>g> biogeography, is that <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
Iturralde-Vinent <strong>and</strong> MacPhee (1999). These references<br />
have been used to c<strong>on</strong>struct <str<strong>on</strong>g>the</str<strong>on</strong>g> following<br />
summary.<br />
<str<strong>on</strong>g>Cuba</str<strong>on</strong>g> seems to be composed mainly <str<strong>on</strong>g>of</str<strong>on</strong>g> a terrane<br />
<str<strong>on</strong>g>of</str<strong>on</strong>g> slivers <str<strong>on</strong>g>of</str<strong>on</strong>g> oceanic material stacked by compressive<br />
deformati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> linear masses <str<strong>on</strong>g>of</str<strong>on</strong>g> sufficient<br />
thickness to achieve emergence. This seems to be<br />
<str<strong>on</strong>g>the</str<strong>on</strong>g> result <str<strong>on</strong>g>of</str<strong>on</strong>g> lateral compressi<strong>on</strong> at <str<strong>on</strong>g>the</str<strong>on</strong>g> edge <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g><br />
Caribbean tect<strong>on</strong>ic plate as it moved from an eastern<br />
Pacific locati<strong>on</strong> into a locati<strong>on</strong> between <str<strong>on</strong>g>the</str<strong>on</strong>g> North<br />
<strong>and</strong> South American plates in late Cretaceous <strong>and</strong><br />
early Tertiary (Eocene) times. <str<strong>on</strong>g>Cuba</str<strong>on</strong>g> also seems to<br />
include parts <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Florida-Bahamas c<strong>on</strong>tinental<br />
margin, <strong>and</strong> possible fragments <str<strong>on</strong>g>of</str<strong>on</strong>g> c<strong>on</strong>tinental margin<br />
derived from <str<strong>on</strong>g>the</str<strong>on</strong>g> south (<str<strong>on</strong>g>the</str<strong>on</strong>g> Chortis block).<br />
All <str<strong>on</strong>g>the</str<strong>on</strong>g>se fragments, isl<strong>and</strong> arc sediments, <strong>and</strong><br />
volcanics seem to have been c<strong>on</strong>solidated in <str<strong>on</strong>g>the</str<strong>on</strong>g> late<br />
Cretaceous <str<strong>on</strong>g>with</str<strong>on</strong>g> renewed compressive deformati<strong>on</strong><br />
in <str<strong>on</strong>g>the</str<strong>on</strong>g> Eocene. Thus, <str<strong>on</strong>g>Cuba</str<strong>on</strong>g> possibly provides a small<br />
excepti<strong>on</strong> to <str<strong>on</strong>g>the</str<strong>on</strong>g> strictly n<strong>on</strong>-c<strong>on</strong>tinental origin <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
<str<strong>on</strong>g>the</str<strong>on</strong>g> Greater Antilles. But <str<strong>on</strong>g>the</str<strong>on</strong>g>re is no evidence for<br />
any direct, emergent, c<strong>on</strong>tinental l<strong>and</strong> c<strong>on</strong>necti<strong>on</strong><br />
at any time. However, <str<strong>on</strong>g>the</str<strong>on</strong>g>re is evidence that <str<strong>on</strong>g>the</str<strong>on</strong>g>re<br />
were very early but limited l<strong>and</strong> c<strong>on</strong>necti<strong>on</strong>s to<br />
what is now Hispaniola <strong>and</strong> Puerto Rico, but not<br />
Jamaica.<br />
After its origin by c<strong>on</strong>solidati<strong>on</strong>, <str<strong>on</strong>g>the</str<strong>on</strong>g> history <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
<str<strong>on</strong>g>Cuba</str<strong>on</strong>g> was probably <strong>on</strong>e <str<strong>on</strong>g>of</str<strong>on</strong>g> isolati<strong>on</strong> from <str<strong>on</strong>g>the</str<strong>on</strong>g> rest <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
<str<strong>on</strong>g>the</str<strong>on</strong>g> Greater Antilles. From 40 Ma (milli<strong>on</strong> years<br />
ago) to 10 Ma it lay north <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> east-west strikeslip<br />
faults that c<strong>on</strong>tinued to juxtapose different slivers<br />
<str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> o<str<strong>on</strong>g>the</str<strong>on</strong>g>r isl<strong>and</strong>s (especially Hispaniola).<br />
<str<strong>on</strong>g>Cuba</str<strong>on</strong>g> is presently composed <str<strong>on</strong>g>of</str<strong>on</strong>g> three c<strong>on</strong>nected<br />
blocks: Western <str<strong>on</strong>g>Cuba</str<strong>on</strong>g>, West Central <str<strong>on</strong>g>Cuba</str<strong>on</strong>g>, <strong>and</strong><br />
Eastern <str<strong>on</strong>g>Cuba</str<strong>on</strong>g>. Available evidence suggests that<br />
<str<strong>on</strong>g>the</str<strong>on</strong>g>se have existed separately as subaerial l<strong>and</strong> since<br />
<str<strong>on</strong>g>the</str<strong>on</strong>g> Eocene-Oligocene boundary, at about 35 Ma,<br />
<strong>and</strong> that <str<strong>on</strong>g>the</str<strong>on</strong>g> three blocks have been joined into <str<strong>on</strong>g>the</str<strong>on</strong>g><br />
present isl<strong>and</strong> since <str<strong>on</strong>g>the</str<strong>on</strong>g> mid-Miocene (after 8 Ma).