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On the Distribution of Greater Antillean Bats1 Thomas A. Griffiths

Department of Biology, Illinois Wesleyan University, Bloomington, Illinois 61702, U.S.A.

and

David Klingener

Department of Zoology, University of Massachusetts, Amherst, Massachusetts 01003, U.S.A.

ABSTRACT Compilation and analysis of geographic distributions of Antillean bat species reveal two overall distribution patterns in Greater Antillean bats. In the first, found among most of the earliest known bat colonizers of the Antilles, Cuban and Hispaniolan bats have a close taxonomic relationship, whereas Jamaican and Puerto Rican bats are more distantly related. This pattern supports a limited vicariance model for early Antillean colonizers, or possibly a dispersal model modified to take into account mid-Tertiary geography. In the second, found among more recent bat colonizers, species accumulate in the western Antillean islands of Cuba and Jamaica, with limited dispersal eastward. The second pattern supports a dispersal model modified to take into account Pleistocene climatic cycles.

IN RECENT YEARS, three approaches to explain the present- day distributions of Antillean organisms have become prevalent: the dispersal theory, the vicariance theory, and the ecological determinism theory. Proponents of the dispersal theory (Simpson 1956, Koopman 1958b, Terborgh 1973; Pregill 1981, to name a few) argue that overwater dispersal can account completely for observed present-day distributions, although habitat of the island in question also will affect whether a potential colonizer is successful (i.e., ecological determinism). With the increasing accep- tance of plate tectonics in the last 20 yr, explanations for present-day distributions have been formulated that take into account the complex geological history of the Carib- bean basin (vicariant theories of zoogeography; see Rosen 1975, MacFadden 1980). Unfortunately, adherents of one or the other approach often reject summarily the possible involvement of other factors.

We have compiled herein an extensive and exhaustive list of distributions of bat species of the Antillean islands. The purpose of this paper is to test the relative importance of dispersal and vicariance as they may apply to Greater Antillean bat species. To accomplish this, we examine the distributions, attempting to detect generalized patterns of distribution. We then attempt to evaluate the importance of vicariance and dispersal in producing the patterns.

GEOLOGIC HISTORY OF THE CARIBBEAN.-Several theories account for the formation of the Greater Antilles (see Molnar & Sykes 1969, Malfait & Dinkleman 1972, Mey- erhoff & Meyerhoff 1972). One view, widely accepted by vicariance biogeographers, states that an island archipelago termed the "proto-Antilles" formed between Central and

South America in pre-Cretaceous times (Freeland & Dietz 1971, Meyerhoff & Meyerhoff 1972, Burke et al. 1984). This island cluster included parts of what are now Jamaica, Hispaniola, Puerto Rico, the Oriente Province of Cuba (though not the rest of Cuba), and various other islands now submerged. The proto-Antilles remained stationary while the North American and South American plates moved westward (Jordan 1975), thus bringing the Ca- ribbean plate to a position just east of proto-Central Amer- ica.

On the other hand, Perfit and Heezen (1978) suggest that during the Cretaceous the North American plate moved southward and was subducted under the Caribbean plate. This subduction raised the oceanic lithosphere of the northern Caribbean plate, forming an ancestral Cay- man Ridge/Nicaraguan Plateau, which in turn led to the formation of a chain of volcanic islands. By the late Eocene, volcanism had mostly stopped, and increased uplift had formed the ancestral Cayman Ridge/Nicaraguan Plateau into an island arc stretching from proto-Central America through the Oriente Province (still a separate island) to Hispaniola and Cuba (see Fig. 1). Jamaica did not then exist as a single island, but may have existed as a cluster of small islands. Hispaniola and Puerto Rico were large islands at this time.

At the end of the Eocene, the Cayman Ridge broke away from the Nicaraguan Plateau and began moving north and east relative to North America (Perfit & Heezen 1978), carrying with it the Oriente Province, the Jamaica island group, much of Hispaniola, and other islands. The Oriente Province moved northeast to eastern Cuba, was sutured onto that end, and began to be uplifted, forming the Sierra Maestra. Hispaniola moved to its present position. Jamaica became fully emergent sometime in the Miocene (Perfit & Heezen 1978). Much of the remainder

I Received 16 September 1986, revision accepted 15 December 1986.

240 BIOTROPICA 20(3): 240-251 1988

of the island arc disappeared via subsidence, including the Cayman Ridge and Nicaraguan Plateau. Hispaniola, Ja- maica, and Puerto Rico are still moving eastward relative to North America and Cuba, although the rate of move- ment is disputed (Hedges 1982).

After the island arc subsided, the Greater Antilles attained much of the degree of isolation they possess today. The Lesser Antillean island arc developed since the late Eocene via volcanism caused by the subsidence of the Aves Ridge (Nagle 1971, Meyerhoff & Meyerhoff 1972). By post-Eocene times, this island arc was presumably open to invasion and may have served as a pathway for South American bats to invade the Greater Antilles. The only other known events that reduced the isolation of both the Greater and Lesser Antilles were the lowered sea levels during the extensive glaciations of the Pleistocene. Sea levels were lowered 100 to 150 m (Donn et al. 1962, Gascoyne et ail. 1979, Cronin et al. 1981), exposing a series of "stepping stone" islands to Cuba and especially Jamaica. Florida and Cuba were less isolated because of the emergence of the Great Bahama and Cay Sal Banks (Fig. 2). The distance between South America and the Lesser Antillean island of Grenada was reduced from about 125 km to about 30 km.

Burke et al. (1984) provided a comprehensive review of the geological literature on plate tectonics of the Ca- ribbean basin, and Hedges (1982) discussed the biogeographic implications of many current geological hypotheses. Hedges observed that virtually all geologists cited agreed that tectonic motion has occurred along a fault extending from Guatemala through the Cayman Trough to northern Hispaniola and the Puerto Rico Trench. Cuba and Hispaniola are thought to have been in close proximity to one another, or actually connected, some time between mid-Miocene (17 MYA) and the late Pliocene (5 MYA), the time depending on which estimate of the rate of motion one accepts.

METHODS The distributional data on living Antillean bat species are summarized in Tables 1 and 2 and in the Appendix. Though our analysis is directed at distributions of Greater Antillean bats (from Cuba, Hispaniola, Jamaica, and Puer- to Rico), we also tabulated distributions of bats on islands of the Lesser Antilles. Some chiropteran species are known only from fossils in the Greater Antilles, and some are locally extinct on certain islands. Table 3 contains distributional data on fossils that either extend the ranges of species listed in Table 2 or provide information on all Greater Antillean species not listed in Table 2 (in other words, those known only from fossil specimens). Data were collected from the excellent distributional summary by Baker and Genoways (1978), and from our own examination of museum specimens at the American Museum

K)~~~~~~~~~~~~~~~~~~~P

CR/NP ORIENTE

proto - CENTRAL AMERICA

SOUTH AMERI CA

FIGURE 1. Map showing the relationship between proto- Central America, South America, and the Greater Antilles as it might have appeared in the Oligocene if Perfit and Heezen's subduction hypothesis is correct (after Perfit & Heezen 1978, with data from Ladd 1976). Areas outlined off the tip of proto- Central America were probably not single islands, but clusters of small islands. Other hypotheses discussed (see text) would place Hispaniola, Jamaica, and Puerto Rico further west in the Oligocene, with contact between Hispaniola and Cuba occurring as late as the Pliocene. CR/NP = Cayman Ridge/Nicaraguan Plateau complex.

of Natural History in New York, the National Museum of Natural History in Washington, the Field Museum of Natural History in Chicago, and the University of Mas- sachusetts Museum of Zoology in Amherst. Additional confirmations, corrections, and new additions to the An- tillean faunal lists were taken from: Anderson and Nelson (1965), Anthony (1917, 1919), Baker and Genoways (1976), Baker et at. (1978), Buden (1975a, b), Choate and Birney (1968), Davis (1973), Eger (1977), Genoways and Baker (1972, 1975), Genoways and Jones (1975), Genoways et al. (1973, 1981), G. G. Goodwin (1959), R. E. Goodwin (1970), Hall (1981), Hall and Bee (1960), Hall and Jones (1961), Hill and Evans (1985), Husson (1960), Jones and Baker (1979), Jones and Phillips (1970, 1976), Jones and Schwartz (1967), Klingener et al. (1978), Kock and Stephan (1986), Koopman (pers. comm., 1958a, b, 1968, 1975, 1976, 1981), Koopman and Williams (1951), Koopman et al. (1957), LaVal (1973), Martin and Schmidly (1982), Martin (1972), Miller (1900, 1904,

Greater Antillean Bat Distribution 241

TABLE 1. Geographical and faunal data for the 21 Antillean islands considered in this paper. (Forfossil occurrences, see Table 3.)

Maxi- mum eleva- No. No. No.

Area tion spe- Ant. isl. Island (kiM2) (m) cies end.- end.b

1. Cuba 111,463 1,973 26 12 4 2. Hispaniola 76,193 3,170 18 10 1 3. Jamaica 11,424 2,256 21 11 4 4. Puerto Rico 8,865 1,338 13 7 0 5. Guadeloupe 1,510 1,484 11 6 2 6. Martinique 1,100 1,388 10 4 0 7. Dominica 751 1,447 12 4 1 8. St. Lucia 616 959 8 3 0 9. Barbados 430 337 6 3 0

10. St. Vincent 344 1,225 9 3 0 11. Antigua 280 405 7 2 0 12. Grenada 277 841 12 0 0 13. St. Croix 207 355 4 1 0 14. Grand Cayman 184 c. 25 8 3 0 15. St. Kitts 168 1,156 4 0 0 16. Barbuda 161 61 6 2 0 17. Montserrat 98 914 9 4 0 18. Anguilla 91 59 5 2 0 19. St. Martin 88 406 5 1 0 20. St. Eustatius 31 600 5 2 0 21. Saba 13 860 3 1 0

a Ant. End. = species of bats endemic to 2 or more Antillean islands. b Is. End. = species of bats endemic to a single Antillean island.

1918), Gary Morgan (pers. comm.), Morgan and Woods (1986), Orr and Silva Taboada (1960), Ottenwalder and Genoways (1982), Pierson et al. (1986), Schwartz and Jones (1967), Silva Taboada (1974, 1979), Silva Ta- boada and Koopman (1964), Simpson (1956), Smith (1972), Starrett and Rolle (1963), Swanepoel and Gen- oways (1978), Varona (1974), and Woloszyn and Silva Taboada (1977). On the basis of personal examination of specimens, we do not follow Buden (1976) in placing all Antillean Erophylla in the species Erophylla sezekorni; rather, we follow Klingener et al. (1978) and recognize E. sezekorni of Cuba and Jamaica as a distinct species from E. bombifrons of Hispaniola and Puerto Rico. In some instances, we follow the most recent reviewer of a taxon; for example, we follow Hall (1981) in recognizing three distinct species of Lasiurus (L. pfeifferi, L. degelida, and L. minor) in the Antilles, rather than prior convention that treated them as three subspecies of L. borealis. As will be evident in the following, whenever we have had to choose between two (or more) ways of treating a taxon, it has not made any difference to our analyses, except for our decision to follow Klingener et al. (1978) rather than Buden (1976) on Antillean Erophylla.

RESULTS The distributions of the 38 living species of bats currently known to inhabit the islands of the Greater Antilles are listed in Appendix I. In Table 1, we have summarized the numerical data on an island-by-island basis. We list geographic data about each island, followed by a listing of total number of bat species per island, total number of Antillean endemic species (bat species found only in the Greater and/or Lesser Antilles), and the number of bat species endemic to the single island in question. In Tables 2 and 3, we have listed the 38 living species plus 9 species known only from fossils in the Greater Antilles, with distributions in three columns. In column 1 are the 16 living (and 3 additional fossil) Antillean species that are conspecific with mainland species. In column 2 are the 11 living (and 3 additional fossil) species that are congeneric with mainland species. In column 3 are the 11 living (and 3 additional fossil) Greater Antillean bat species that are neither conspecific nor congeneric with mainland species.

The third column of each table shows an interesting arrangement of distributions within every taxon but Ero- phylla. Within the species Monophyllus redmani, which is found throughout the Greater Antilles, Cuba and His- paniola share a subspecies, M. r. clinedaphus, whereas different subspecies are found on Jamaica (redmani) and on Puerto Rico (portoricensis). A different species, Mono- phyllusplethodon, is known from fossil specimens in Puerto Rico (Table 3) and from live specimens taken in the Lesser Antilles. Within the subfamily Stenodermatinae, four endemic species are found on the Antilles. Cuba and His- paniola share a genus: Cuba has Phyllops falcatus [and the fossil species P. vetusl and Hispaniola has P. haitiensis. Jamaica and Puerto Rico have distinct genera of their own (as do the Lesser Antilles with the endemic Antillean stenodermatine species Ardops nichollsi). Within the subfamily Brachyphyllinae, Cuba and Hispaniola share a single species within the genera Phyllonycteris and Brachy- phylla, but not Erophylla. In the case of the genus Phyl- lonycteris, Jamaica has its own species, P. aphylla, and Puerto Rico had its own species, P. major, now extinct (Table 3). In the case of Brachyphylla, Puerto Rico and the Lesser Antilles share a distinct species (B. cavernarum), but apparently Brachyphylla is extinct on Jamaica (B. pumila, Koopman & Williams 195 1, considered B. nana by Swanepoel & Genoways 1978, though the Jamaican population showed some morphological distinctiveness).

An analysis of the distributions of species in the first and second columns reveals that only one of the taxa found on more than one Greater Antillean island has a "special Cuba-Hispaniola taxonomic relationship" similar to that found for the bats in the third column. Mormoops megalophylla is known from fossil specimens in Cuba and Hispaniola. Of the remaining 32 species found in the first

242 Griffiths and Klingener

and second columns of Tables 2 and 3, five taxa have distributional patterns that do not reveal much about their biogeographic history. Noctilio leporinis and Mormoops blainvillii are distributed over all four Greater Antilles without any special relationships between islands. Mac- rotus waterhousii has the same distributional pattern, tak- ing into consideration its fossil occurence on Puerto Rico (Table 3). Tadarida brasiliensis and Molossus molossus are thought to have endemic subspecies on each island, but relationships between them are in need of study. Of the remaining 27 species, 5 are clearly relatively recent arrivals on Cuba from the mainland: Nycticeius humeralis, Tadar- ida laticaudata, Eumopsperotis, Lasiurus intermedius, and Antrozous pallidus. In Table 3, Desmodus rotundus is also probably a relatively recent arrival on Cuba. Two living species are clearly relatively recent arrivals on Jamaica, Glossophaga soricina and Eumops auripendulus. Of the fossil species, Tonatia bidens also probably belongs with this group. Of the remaining 18 species, no taxon has a Cuba-Puerto Rico or a Jamaica-Puerto Rico pattern of special relationship. Three taxa show a Cuba-Jamaica pattern (Eumops glaucinus, Pteronotus macleayi, and Pterono- tusparnellii), and one shows aJamaica-Hispaniola pattern (Natalus micropus). The species L. pfeifferi, L. degelida, and L. minor were, until recently, considered subspecies of the mainland species L. borealis. If they are considered sister species, they would together form the sole example of a Hispaniola-Puerto Rico pattern (L. minor is found on Hispaniola and Puerto Rico). Arnold et al. (1980) demonstrated that Eptesicus lynni from Jamaica was closely related to E. fuscus (though not necessarily most closely related to Antillean E. fuscus). Whatever relationship E. lynni has to other Antillean Eptesicus, E. fuscus is the sole example of a taxon with a Cuba-Hispaniola-Puerto Rico pattern, and E. lynni is either a more distantly related Antillean relative or a separate arrival from the mainland. Two species show a Cuba-Jamaica-Hispaniola pattern, Tadarida macrotus and Natalus major (the latter known only from fossils on Cuba; Goodwin 1959). Two show the final possible pattern, a Jamaica-Hispaniola-Puerto Rico pattern, Artibeus jamaicensis and Pteronotus fuliginosus.

Systematic relationships are not presently known of the living species Natalus lepidus, nor of the two large fossil mormoopids (Pteronotus pristinus and Mormoops magna) and the endemic fossil stenodermatine (Artibeus anthonyi) of Cuba. They may represent speciation events in Cuba, derived either from other Cuban or Antillean species or from a mainland progenitor. The final Antillean species, Mormopterus minutus of Cuba, is intriguing in that it is apparently most closely related to two Peruvian species of Mormopterus (see Freeman 1981) and may be closely related to African molossid species. We cannot make state- ments about patterns of distribution until its systematics

are better understood, but we would not be surprised if fossil Mormopterus are someday discovered on Hispaniola.

Among congeneric and conspecific species, there is no single dominant pattern of distribution such as appears in the "neither" species. However, analyzing the 32 conspecific-congeneric species in a slightly different fashion, 12 of the 32 are found only on Cuba (L. intermedius, L. pfeifferi, N. humeralis, T. laticaudata, E. perotis, A. pallidus, M. minutus, N. lepidus, D. rotundus, P. pristinus, M. magna, and A. anthonyi). Four of the 32 are found only on Jamaica (G. soricina, T. bidens, E. lynii, and E. auripendulus). Two of the 32 are found only on Cuba and Jamaica (E. glaucinus and P. macleayi). One species is found on Cuba and Hispaniola (M. megalophylla). Three more species are found on Cuba, Jamaica, and Hispaniola (M. waterhousii, N. micropus, and T. macrotis). Thus a total of 18 of 32 species of bats have a distribution restricted to one or more of the western-most Greater An- tillean islands, and 22 of the 32 are not found farther east than Hispaniola. Interestingly, all of these except M. minutus are conspecific or at least congeneric with species found to the west or north in Central America, North America, or both.

We have also produced three species-area curves (cf MacArthur & Wilson 1963, 1967) for the islands listed in Table 1 (see Fig. 3). One curve ("L" in Fig. 3) incorporates data from the 17 Lesser Antillean islands alone. The slope (z) of curve L is 0.25 (r = 0.74). Curve L is extended in Figure 3 (dotted line) up to islands the size of the Greater Antilles. A second curve ("G" in Fig. 3) was plotted using data from the Greater Antilles alone. The slope for curve G is 0.14 (r = 0.64). The third curve ("B") incorporates data from both the Greater and Lesser Antilles; the slope z is 0.21 (r = 0.88).

DISCUSSION We suggest two overall distributional patterns for Greater Antillean bats. One of these patterns, which we shall term the "Cuba-Hispaniola (C/H) pattern," can be found among the group of bats that includes the earliest bat colonizers of the Greater Antilles (column 3 of Tables 2 and 3). In every taxon discussed, except within the genus Erophylla, there is a peculiar "special taxonomic relationship" between the bats of Cuba and Hispaniola. The C/H pattern exists only in M. megalophylla of all the species listed in columns 1 or 2 of Tables 2 and 3. No single overall distributional pattern was found in bats of columns 1 and 2; rather, the pattern of distribution found shows bats to be largely restricted to the western Greater An- tilles either to Jamaica alone, Cuba alone, or to Jamaica and Cuba together. We call this the Western Antilles (WA) pattern.

Neither the C/H pattern nor the WA pattern supports


TABLE 2. Summary of living Greater Antillean bat species showing distributions of species conspecific, congeneric, and neither conspecific nor congeneric with mainland species.

Conspecific Congeneric Neither

Family Noctilionidae Noctilio leporinus

mastivus C, H, J, P

Family Mormoopidae Pteronotus parnellii Pteronotus fuliginosus parnellii C, J torrei C gonavensis H (part) fuliginosus H, J, P pusillus H (part) Pteronotus macleayi portoricensis P macleayi C

griseus J Mormoops blainvillii C, H, J, P

Family Phyllostomidae Subfamily Phyllostominae

Macrotus waterhousii minor C waterhousii H jamaicensis J

Subfamily Glossophaginae Glossophaga soricina Monophyllus redmani

antillarum J clinedaphus C, H redmani J portoricensis P

Subfamily Stenodermatinae Artibeus jamaicensis Phyllops falcatus C

jamaicensis H, J, P Phyllops haitiensis H parvipes C Ariteus flavescens J

Stenoderma rufum P Subfamily Brachyphyllinae

Phyllonycteris poeyi C, H Phyllonycteris aphylla J Brachyphylla nana C, H Brachyphylla cavernarum P Erophylla sezekorni C, J Erophylla bombi frons H, P

Family Natalidae Natalus major H, J Natalus lepidus C Natalus micropus

macer C micropus H, J

Family Vesperitilionidae Eptesicus fuscus Eptesicus lynii J

dutertreus C hispaniolae H wetmorei P

Lasiurus intermedius Lasiurus pfeifferi C insularis C Lasiurus degelida J

Lasiurus minor H, P Antrozous pallidus

koopmani C Nycticeius humeralis

cubanus C


TABLE 2. Continued.


Family Molossidae Tadarida brasiliensis Mormopterus minutus C

muscula C constanzae H murina J anti'llularum P

Tadarida laticaudata C Tadarida macrotis C, H, J Eumops auripendulus J Eumops glaucinus C, J Eumops perotis C Molossus molossus

tropidorhynchus C verrilli H miller J fortis P

a straight vicariance hypothesis for bat colonization of the Greater Antilles, although the C/H pattern might support a limited vicariance hypothesis for the more ancient An- tillean bat species. If vicariance accounted for all presently known Antillean bat species, we would expect to see a single taxonomic pattern among all (or at least most)

Antillean bat species today. This pattern should reflect the order of breakup between Greater Antillean islands: for example, if Puerto Rico were isolated first in time, with Cuba and Hispaniola connected until relatively recently, we would expect that the bats of Cuba and Hispaniola would be most closely related and those of Puerto Rico

TABLE 3. Known fossil species of the Greater Antilles, not including islands where a species is known from both living forms and fossil forms. (See Table 2 for living forms.) Bat species are listed in three columns: species that are conspecific, congeneric, and neither conspecific nor congeneric with living mainland species.


Family Mormoopidae Pteronotus pristinus C

Mormoops megalophylla C, H Mormoops magna C

Family Phyllostomidae Subfamily Phyllostominae

Tonatia bidens J Macrotus waterhousii pa

Subfamily Glossophaginae Monophyllus plethodon pb

Subfamily Stenodermatinae Artibeus anthonyi C Phyllops vetus C

Subfamily Brachyphyllinae Brachyphylla nana(?) Ja Phyllonycteris major P

Subfamily Desmodontinae Desmodus rotundus C

Family Natalidae Natulus major ca

a Extinct on island indicated, but alive elsewhere in the Greater Antilles. b Extinct on island indicated, but alive in the Lesser Antilles.


most taxonomically distant. This is simply a reflection of the fact that, although bats are quite vagile over land and short water barriers, they seem not to cross larger water barriers easily (Koopman 1970). The problem with the straight vicariance theory is that it cannot explain the two patterns (C/H and WA) simultaneously.

On the other hand, the patterns of bat distribution in the Greater Antilles do not support a simple waif dispersal model either, at least not without some quali- fication. If chance dispersal (for example, a bat being blown off course by a tropical storm) is postulated to account for present-day distributions, then the patterns of distribution of bat species in the Antilles should be more random than is evident. Certainly those islands that are closest to the source (Jamaica and Cuba to Central Amer- ica, for example) would be expected to have the largest number of source-related species. This accounts for part of the WA pattern. However, unless some additional factor at work "directs" species only to Cuba and Jamaica and limits species to those two islands, the WA pattern should break down relatively quickly. For example, if a waif arrives on Jamaica and founds a population that grows and thrives, unless some outside "Cuba factor" is working, it is just as likely that a Jamaican bat will get to Hispaniola and found a population as it is that it will get to Cuba under the waif dispersal model. More im- portantly, we would expect that species would be con- stantly arriving on the western islands and founding populations that then disperse to more easterly islands. A species capable of crossing the wide ocean barrier between Central America and Cuba should be able to cross the comparatively narrow Cuba-Hispaniola barrier, for example. This doesn't seem to have happened much in the WA species. The waif dispersal model poses one other problem: unless some factor allows species easy access to only Cuba and Jamaica, then certainly some bat species should arrive and colonize only Hispaniola or only Puerto Rico under the waif dispersal model. This also seems not to have happened.

We suggest that neither a straight vicariance nor a simple waif dispersal model can account for the present day distributions of Antillean bats. We propose an alter- nate two-event model that not only accounts for all the observed characteristics of bat distributions, but also agrees with the geological history of the Caribbean basin as it is currently understood. MacFadden (1980) interpreted the distribution of insectovorans in the Greater Antilles as the result of a three-event scenario, with extinction of some populations being the third event. He explained the presence of solenodontids on Cuba and Hispaniola by dispersal from the latter island to the former. We suggest that the presence of solenodontids on both islands could be due to the same event that we believe accounts for distribution patterns of "old" Antillean bats.

The C/H pattern could support a limited vicariance model, or it could support a much-modified dispersal

model. There has been speculation recently in the geological literature (see Hedges 1982) that Cuba and Hispaniola were connected, or nearly so, between 5 and 17 MYA. If this were the case, then the breakup of this connection would be a vicariant event that split the faunas of the two islands, and we would expect to see a pattern similar to the C/H pattern we have observed herein. The more distantly related species on Jamaica and Puerto Rico would either be disperser-founded populations from Cuba/His- paniola or (less likely) remnants of the original vicariant breakup of the Greater Antilles.

Alternatively, if at some point in the late Tertiary, a dispersal route existed to Cuba and Hispaniola that made it possible for large populations of Central American bats to colonize Cuba and Hispaniola simultaneously, then stabilizing selection would tend to keep the Cuban and Hispaniolan populations morphologically similar. Such would not be the case for the much smaller, disperser- founded populations of Jamaica and Puerto Rico, resulting in the C/H pattern. For this explanation to work, we must explain why Jamaica, which is closer to Central America than Hispaniola, would not have been colonized.

Figure 1 shows the Caribbean basin as it may have appeared in the Oligocene (after Ladd 1976, and Perfit & Heezen 1978). Note that Jamaica was mostly submerged, that the "toe" of Central America extended much further out toward the Antilles than today, and that there was a chain of islands (the most distal being the Oriente, now the eastern-most province of Cuba). Such an island chain certainly could have provided a pathway for large populations of bats to colonize both Cuba and Hispaniola simultaneously, bypassing Jamaica. However, if Perfit and Heezen (1978) are correct about the age of the chain of islands, then the original colonization would have had to occur in the Oligocene or early Miocene (cf Koopman 1981) to utilize the chain. It seems most unlikely that a "special taxonomic relationship" between bats of Cuba and Hispaniola could still be recognized at the genus, species, or subspecies level after all that time. If future study shows that the age of the island chain was younger than suggested by Perfit and Heezen, then this hypothesis might explain the C/H pattern. Present evidence suggests that, although the original colonizers of the Greater An- tilles might have utilized such a chain of islands initially to colonize the Greater Antilles, the C/H pattern is best explained by a vicariant event between Cuba and His- paniola that occurred more recently, perhaps slightly over 5 MYA.

The second pattern of distribution, the WA pattern, can be explained only by dispersal of species from North and Central America, but with one very important qual- ification: such dispersal was not random, but "directed" by geological events specifically to Cuba and Jamaica. Figure 2 shows the Caribbean basin as it might have appeared in the Pleistocene during a lowering of the sea level. Note that Hispaniola (and Puerto Rico, not shown)


90 85 80 70

FLORIA

25 Cay Sal B N G. B. Bank

%> \ v Pe droll Bank t

4 /

\Ni~~~~~~~~~cerga Pteu ( ; / d1

go es so SourH ~~~~~~~~ ~~~~ANEI OL s

90 85 80 75

FIGURE 2. Map of Central America and the western Greater Antilles. Thin line: how the land masses appear today. Thick line: how the land masses would appear if ocean levels dropped to the lowest levels reached during the Pleistocene glaciarions. G. B. Bank-Great Bahamas Bank. Bar in lower left equals 100 km.

are relatively isolated islands, much as they are today. Cuba is much more accessible from North America via a much-enlarged Florida and several undersea banks that have become islands. Cuba is also slightly more accessible from Central America. Jamaica is much more accessible from Central America via the exposed Nicaraguan Plateau and a number of exposed banks.

We suggest that this second event, the exposure of a 'stepping stone" set of islands out to the western Antilles, would have resulted in exactly the WA pattern we have observed. In addition, the relative recentness of the drops in sea level during the Pleistocene explains why such a large number of species on both Cuba and Jamaica have not yet dispersed eastward. Note that in Figure 3, both Cuba (1) and Jamaica (3) are above their respective equilibrium numbers of species for the B and G curves, and

Jamaica is above for all three curves. Both Hispaniola (2) and Puerto Rico (4) are well below their equilibrium numbers. This would be expected if our model is correct.

Vicariance cannot account for the WA pattern observed, for two reasons. First, we are aware of no geological evidence that supports a Cuba-Jamaica connection. Sec- ond, if such a connection did exist at a time more recent than the hypothetical Cuba-Hispaniola connection (see above), the resulting Cuba-Jamaica taxonomic pattern would both obliterate the more ancient C/H pattern and result in a much higher number of Cuban-Jamaican shared species than the 3 species currently known.

In summary, we suggest that the two distributional patterns we have observed among species of Antillean bats (C/H and WA) indicate that the colonization of the Antilles was a two-stage process, dominated by two geo-


100

w *

loo -

a-

t) ~ ~ ~ ~ ~ 0 1000 0-0 100,00

cn _

o ; 5 = J 7 S ;~~~~~~

* 1 3

~ 21

10 100 1,000 10,000 100,000

AREA (km2)

FIGURE 3. Species-area relationships for the 21 islands listed in Table 1. G = curve for the Greater Antillean islands, L = curve for the 17 Lesser Antillean and selected other islands (dotted line is an extrapolation to islands the size of the Greater Antilles), B = all islands.

logical events. The first event might have begun with the appearance sometime in the Tertiary of a chain of islands leading out to Cuba and Hispaniola, but bypassing Ja- maica. Early colonizers would have utilized the chain to reach the Greater Antilles. More recently, it appears that the connection or close proximity of Cuba and Hispaniola allowed free migration between these two now separated islands. The decoupling of Cuba and Hispaniola would have been a vicariant event for the land-based faunas of the two islands, and it would have resulted in the C/H pattern we have observed among the oldest Antillean bat species. The second event, much later, would have been the multiple sea-level lowerings that occurred during the Pleistocene. This event would have resulted in a number of species, some now extinct, colonizing primarily Jamaica and Cuba (but Hispaniola and Puerto Rico to a much

lesser extent). The resulting pattern of species would cor- respond well with the WA pattern we have observed here.

ACKNOWLEDGMENTS

We thank Drs. Margery Coombs, Donald Kroodsma, Charles Pitrat, Carroll Schloyer, and Dana Snyder of the University of Massachusetts; Dr. William Wall of Georgia College; Dr. J. Knox Jones, Jr., and Robert Owen of Texas Tech University; and several anonymous reviewers for reviewing various versions of the manuscript. We are particularly grateful to Douglas Smith, University of Massachusetts, Gary Morgan, Florida State Mu- seum, and Dr. Karl F. Koopman, American Museum of Natural History, all of whom contributed unpublished data to our study and reviewed earlier versions of the manuscript. Special thanks to Dr. Charles 0. Handley, Jr., Dr. Koopman, Dr. Bruce D. Patterson, and Dr. Robert M. Timm for allowing us access to specimens in their care.

LITERATURE CITED ANDERSON, S., AND C. E. NELSON. 1965. A systematic revision of Macrotus (Chiroptera). Am. Mus. Novit. 2212: 1-39. ANTHONY, H. E. 1917. Two new fossil bats from Porto Rico. Bull. Amer. Mus. Nat. Hist. 37: 565-568 (pl. 56).

1919. Mammals collected in eastern Cuba in 1917, with descriptions of two new species. Bull. Amer. Mus. Nat. Hist. 41: 625-643.

ARNOLD, M. L., R. J. BAKER, AND H. H. GENOWAYS. 1980. Evolutionary origins of Eptesicus lynni. J. Mammal. 61: 319-322. BAKER, R. J., AND H. H. GENOWAYS. 1976. A new species of Chiroderma from Guadeloupe, West Indies (Chiroptera, Phyllo-

stomatidae). Occas. Pap. Mus. Tex. Tech Univ. 39: 1-9. , AND . 1978. Zoogeography of Antillean bats. Spec. Publ. Acad. Nat. Sci. Phila. 13: 53-97.


BAKER, R. J., H. H. GENOWAYS, AND J. C. PATrON. 1978. Bats of Guadeloupe. Occas. Pap. Mus. Tex. Tech Univ. 50: 1-16. BUDEN, D. W. 1975a. Monophyllus redmani Leach (Chiroptera) from the Bahamas, with notes on variation in the species. J.

Mammal. 56: 369-377. 1975b. A taxonomic and zoogeographic appraisal of the big-eared bats (Macrotus waterhousii Gray) in the West Indies.

J. Mammal. 56: 758-769. 1976. A review of the bats of the endemic West Indian genus Erophylla. Proc. Biol. Soc. Wash. 89: 1-16.

BURKE, K., C. COOPER, J. F. DEWEY, P. MANN, AND J. L. PINDELL. 1984. Caribbean tectonics and relative plate motions. In W. E. Bonini, R. B. Hargraves, and R. Shagram (Eds.). The Caribbean-South American plate boundary and regional tectonics, pp. 31-63. Geol. Soc. Am. Mem. No. 162.

CHOATE, J. R., AND E. C. BIRNEY. 1968. Sub-recent Insectivora and Chiroptera from Puerto Rico, with the description of a new bat of the genus Stenoderma. J. Mammal. 49: 400-412.

CRONIN, T. M., B. J. SZABO, T. A. AGER, J. E. HAZEL, AND J. P. OWENS. 1981. Quaternary climates and sea levels of the U.S. Atlantic coastal plain. Science 211: 233-240.

DAVIS, W. B. 1973. Geographic variation in the fishing bat, Noctilio leporinus. J. Mammal. 54: 862-874. DONN, W. L., W. R. FARRAND, AND M. EWING. 1962. Pleistocene ice volumes and sea level lowering. J. Geol. 70: 206-214. EGER, J. L. 1977. Systematics of the genus Eumops (Chiroptera: Molossidae). R. Ont. Mus. Life Sci. Contrib. 110: 1-69. FREELAND, G. L., AND R. S. DIETZ. 1971. Plate tectonic evolution of Caribbean-Gulf of Mexico region. Nature 232: 20-30. FREEMAN, P. W. 1981. A multivariate study of the family Molossidae (Mammalia, Chiroptera): morphology, ecology, evolution.

Fieldiana-Zool., Chic. Nat. Hist. Mus. n.s. 7: vii + 173 pp. GASCOYNE, M., G. J. BENJAMIN, AND H. P. SCHWARTZ. 1979. Sea-level lowering during the Illinoian glaciation: evidence from

a Bahama "blue hole." Science 205: 806-808. GENOWAYS, H. H., AND R. J. BAKER. 1972. Stenoderma rufum. Mamm. Species 18: 1-4.

, AND . 1975. A new species of Eptesicus from Guadeloupe, Lesser Antilles (Chiroptera: Vespertilionidae). Occas. Pap. Mus. Tex. Tech Univ. 34: 1-7. , AND J. K. JONES, JR. 1975. Additional records of the stenodermine bat, Sturnira thomasi, from the Lesser Antillean island of Guadeloupe. J. Mammal. 56: 924-925. , R. J. BAKER, AND R. S. LOREGNARD. 1973. Two species of bats new to the fauna of Trinidad. Mammalia 37: 362- 363. , R. C. DOWLER, AND C. H. CARTER. 1981. Intraisland and interisland variation in Antillean populations of Molossus molossus (Mammalia: Molossidae). Ann. Carnegie Mus. 50: 475-492.

GOODWIN, G. G. 1959. Bats of the subgenus Natalus. Am. Mus. Novit. 1977: 1-22. GOODWIN, R. E. 1970. The ecology of Jamaican bats. J. Mammal. 51: 571-579. HALL, E. R. 1981. The mammals of North America, Vol. 1. John Wiley and Sons, New York.

, AND J. W. BEE. 1960. The red fig-eating bat Stenoderma rufum Desmarest found alive in the West Indies. Mammalia 24: 67-75. , AND J. K. JONES, JR. 1961. North American yellow bats. "Dasypterus" and a list of the named kinds of the genus Lasiurus Gray. Univ. Kans. Publ. Mus. Nat. Hist. 14: 73-98.

HEDGES, S. B. 1982. Caribbean biogeography: implications of recent plate tectonic studies. Syst. Zool. 31: 518-522. HILL, J. E., AND P. G. H. EVANS. 1985. A record of Eptesicusfuscus (Chiroptera: Vespertilionidae) from Dominica, West Indies.

Mammalia 49: 133-136. HUSSON, A. M. 1960. De Zoogdieren van de Nederlandse Antillen. Natuurwetenschappelijke Werkgroep Nederlandse Antillen,

Curacao. JONES, J. K., JR., AND R. J. BAKER. 1979. Notes on a collection of bats from Montserrat, Lesser Antilles. Occas. Pap. Mus. Tex.

Tech. Univ. 60: 1-6. , AND C. J. PHILLIPS. 1970. Comments on systematics and zoogeography of bats in the Lesser Antilles. Stud. Fauna Curacao Other Caribb. Isl. 32: 131-145. , AND . 1976. Bats of the genus Sturnira in the Lesser Antilles. Occas. Pap. Mus. Tex. Tech. Univ. 40: 1-16. , AND A. SCHWARTZ. 1967. Bredin-Archbold-Smithsonian biological survey of Dominica. 6. Synopsis of bats of the Antillean genus Ardops. Proc. U.S. Nat. Mus. 124: 1-13.

JORDAN, T. H. 1975. The present-day motions of the Caribbean plate. J. Geophys. Res. 80: 4433-4439. KLINGENER, D., H. H. GENOWAYS, AND R. J. BAKER. 1978. Bats from southern Haiti. Ann. Carnegie Mus. 47: 81-99. KOCK, D., AND H. STEPHAN. 1986. Une chauve-souris nouvelle pour la Martinique, Antilles francaises: Monophyllus plethodon

luciae. Mammalia, 50: 268. KOOPMAN, K. F. 1958a. A fossil vampire bat from Cuba. Breviora M. C. Z. 90: 1-4.

1958b. Land bridges and ecology in bat distribution on islands off the northern coast of South America. Evolution 12: 429-439.

1968. Taxonomic and distributional notes on Lesser Antillean bats. Am. Mus. Novit. 2333: 1-13. 1970. Zoogeography of bats. In B. H. Slaughter and D. W. Walton (Eds.). About bats, pp. 29-50. Southern Methodist

Univ. Press, Dallas. 1975. Bats of the Virgin Islands in relation to those of the Greater and Lesser Antilles. Am. Mus. Novit. 2581: 1-7. 1976. Zoogeography. In R. J. Baker, J. K. Jones, Jr., and D. C. Carter (Eds.). Biology of bats of the New World

family Phyllostomatidae, Spec. Publ. Mus. Tex. Tech Univ. 10: 1-218. 1981. The distributional patterns of New World nectar-feeding bats. Ann. Mo. Bot. Gard. 68: 352-369.

AND E. E. WILLIAMS. 1951. Fossil Chiroptera collected by H. E. Anthony in Jamaica, 1919-1920. Am. Mus. Novit. 1519: 1-29. M. K. HECHT, AND E. LEDECKY-JANACEK. 1957. Notes on the mammals of the Bahamas with special reference to the

bats. J. Mammal. 38: 164-174.


LADD, J. W. 1976. Relative motion of South America with respect to North America and Caribbean tectonics. Bull. Geol. Soc. Am. 87: 969-976.

LAVAL, R. K. 1973. A revision of the neotropical bats of the genus Myotis. Nat. Hist. Mus. Los Ang. Cty. Sci. Bull. 15: 1-54. MAcARTHUR, R. H., AND E. 0. WILSON. 1963. An equilibrium theory of insular zoogeography. Evolution 17: 373-387.

, AND . 1967. The Theory of island biogeography. Princeton Univ. Press, Princeton, New Jersey. MAcFADDEN, B. J. 1980. Rafting mammals or drifting islands: biogeography of the Greater Antillean insectivores Nesophontes

and Solenodon. J. Biogeogr. 7: 11-22. MALFAIT, B. T., AND M. G. DINKLEMAN. 1972. Circum-Caribbean tectonic and igneous activity and the evolution of the Caribbean

plate. Bull. Geol. Soc. Am. 83: 251-272. MARTIN, C. O., AND D. J. SCHMIDLY. 1982. Taxonomic review of the pallid bat, Antrozous pallidus (LeConte). Spec. Publ. Mus.

Tex. Tech Univ. 18: 1-48. MARTIN, R. A. 1972. Synopsis of Late Pliocene and Pleistocene bats of North America and the Antilles. Am. Midl. Nat. 87:

326-335. MEYERHOFF, A. A., AND H. A. MEYERHOFF. 1972. Continental drift, IV: The Caribbean "plate." J. Geol. 80: 34-60. MILLER, G. S. 1900. A second collection of bats from the island of Curacao. Proc. Biol. Soc. Wash. 13: 159-162.

1904. Notes on the bats collected by William Palmer in Cuba. Proc. U.S. Nat. Mus. 27: 337-348 (pl. IX). 1918. Three new bats from Haiti and Santo Domingo. Proc. Biol. Soc. Wash. 31: 39-40.

MOLNAR, P., AND L. R. SYKES. 1969. Tectonics of the Caribbean and Middle American regions from focal mechanisms and seismicity. Bull. Geol. Soc. Am. 80: 1639-1684.

MORGAN, G. S., AND C. A. WOODS. 1986. Extinction and the zoogeography of West Indian land mammals. Biol. J. Linn. Soc. 28: 167-203.

NAGLE, F. 1971. Caribbean geology, 1970. Mar. Sci. Bull. 21: 375-439. ORR, R. T., AND G. SILVA TABOADA. 1960. A new species of bat of the genus Antrozous from Cuba. Proc. Biol. Soc. Wash. 73:

83-86. OTTENWALDER, J. A., AND H. H. GENOWAYS. 1982. Systematic review of the Antillean bats of the Natalus micropus-complex

(Chiroptera: Natalidae). Ann. Carnegie Mus. 51: 1-38. PERFIT, M. R., AND B. C. HEEZEN. 1978. The geology and evolution of the Cayman Trench. Bull. Geol. Soc. Am. 89: 1155-

1174. PIERSON, E. D., W. E. RAINEY, R. M. WARNER, AND C. C. WHITE-WARNER. 1986. First record of Monophyllus from Montserrat,

West Indies. Mammalia 50: 269-270. PREGILL, G. K. 1981. An appraisal of the vicariance hypothesis of Caribbean biogeography and its application to West Indian

terrestrial vertebrates. Syst. Zool. 30: 147-155. ROSEN, D. E. 1975. A vicariance model of Caribbean biogeography. Syst. Zool. 24: 431-464. SCHWARTZ, A., AND J. K. JONES, JR. 1967. Bredin-Archbold-Smithsonian biological survey of Dominica. 7. Review of bats of

the endemic Antillean genus Monophyllus. Proc. U.S. Nat. Mus. 124: 1-20. SILVA TABOADA, G. 1974. Fossil Chiroptera from cave deposits in central Cuba, with descriptions of two new species (genera

Pteronotus and Mormoops) and the first West Indian record of Mormoops megalophylla. Acta Zool. Cracov. 19: 33-73. 1979. Los Murcielagos de Cuba. Editorial Academia, La Habana, Cuba.

AND K. F. KOOPMAN. 1964. Notes on the occurrence and ecology of Tadarida laticaudata yucatanica in eastern Cuba. Am. Mus. Novit. 2174: 1-6.

SIMPSON, G.G. 1956. Zoogeography of West Indian land mammals. Am. Mus. Novit. 1759: 1-28. SMITH, J.D. 1972. Systematics of the chiropteran family Mormoopidae. Univ. Kans. Mus. Nat. Hist. Misc. Publ. 56: 1-132. STARRETT, A., AND F. J. ROLLE. 1963. A record of the genus Lasiurus from Puerto Rico. J. Mammal. 44: 264. SWANEPOEL, P., AND H. H. GENOWAYS. 1978. Revision of the Antillean bats of the genus Brachyphylla (Mammalia: Phyllosto-

matidae). Bull. Carnegie Mus. Nat. Hist. 12: 1-53. TERBORGH, J. 1973. Chance, habitat, and dispersal in the distribution of birds in the West Indies. Evolution 27: 338-349. VARONA, L. S. 1974. Catalogo de los Mamiferos Vivientes y Extinguidos de las Antillas. Academia de Ciencias de Cuba, Habana. WOLOSZYN, B. W., AND G. SILVA TABOADA. 1977. Nueva especie f6sil de Artibeus (Mammalia: Chiroptera) de Cuba, y tipificaci6n

preliminar de los dep6sitos fosiliferous cubanos contentivos de mamiferos terrestres. Inst. Zool. Acad. Cienc. Cuba (Poeyana) 161: 1-17.

APPENDIX I. Distribution of Living Species of Antillean Bats Used in this Paper.

Cuba: Noctilio leporinus (NI), Pteronotusparnellii (Pp), Pteronotusfuliginosus (Pf, Pteronotus macleayi (Pm), Mormoops blainvillii (Mb), Macrotus waterhousii (Mw), Monophyllus redmani (Mr), Artibeusjamaicensis (Aj), Phyllopsfalcatus, Brachyphylla nana (Bn), Erophylla sezekorni, Phyllonycterispoeyi, Natalus lepidus, Natalus micropus, Eptesicusfuscus (Ef), Lasiuruspfeifferi, Lasiurus intermedius, Nycticeius humeralis, Antrozous pallidus, Tadarida brasiliensis (Tb), Tadarida laticaudata, Tadarida macrotis (Tm), Mormopterus minutus, Eumops glaucinus, Eumops perotis, Molossus molossus (Mm).

Hispaniola: NI, Pp, Pf, Mb, Mw, Mr, Aj, Phyllops haitiensis, Bn, Erophylla bombifrons, Phyllonycteris poeyi, Natalus major (Nm), Natalus micropus, Ef, Lasiurus minor, Tb, Tm, Mm.

Jamaica: NI, Pp, Pf, Pm, Mb, Mw, Glossophaga soricina, Mr, Aj, Ariteusfiavescens, Erophylla sezekorni, Phyllonycteris aphylla, Nm, Natalus micropus, Eptesicus lynii, Lasiurus degelida, Tb, Tm, Eumops auripendulus, Eumops glaucinus, Mm.

Puerto Rico: NI, Pp, Pf, Mb, Mr, Aj, Stenoderma rufum, Brachyphylla cavernarum (Bc), Erophylla bombifrons, Ef, Lasiurus minor, Tb, Mm.


Guadeloupe: NI, Monophyllus plethodon (Mp), Sturnira thomasi, Chiroderma improvisum, Aj, Ardops nichollsi (An), Bc, Natalus stramineus (Ns), Eptesicus guadeloupensis, Tb, Mm.

Martinique: NI, Pteronotus davyi, Mp, Sturnira lilium, Aj, An, Bc, Myotis martiniquensis, Tb, Mn. Dominica: NI, Pteronotus davyi, Mp, Sturnira lilium, Aj, An, Bc, Ns, Ef, Myotis dominicensis, Tb, Mm. St. Lucia: NI, Mp, Sturnira lilium, Aj, An, Bc, Tb, Mm. Barbados: NI, Mp, Aj, Bc, Myotis martiniquensis, Mm. St. Vincent: NI, Glossophaga longirostris, Mp, Sturnira lilium, Aj, Artibeus lituratus, An, Bc, Mm. Antigua: NI, Mp, Aj, Bc, Ns, Tb, Mm. Grenada: Peropteryx macrotis, NI, Pteronotus davyi, Micronycteris megalotis, Glossophaga longirostris, Anoura geoffroyi, Carollia

perspicillata, Artibeus cinereus, Aj, Artibeus lituratus, Myotis nigricans, Mm. St. Croix: NI, Aj, Bc, Mm. Grand Cayman: Mw, Aj, Phyllops falcatus, Bn, Erophylla sezekorni, Ef, Tb, Mm. St. Kitts: NI, Aj, Tb, Mm. Barbuda: NI, Mp, Aj, Bc, Tb, Mm. Montserrat: NI, Mp, Chiroderma improvisum, Aj, An, Bc, Ns, Tb, Mm. Anguilla: Mp, Aj, Bc, Ns, Mm. St. Martin: NI, Aj, Bc, Tb, Mm. St. Eustatius: Aj, An, Bc, Tb, Mm. Saba: Aj, Bc, Ns.

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