Distribution, Probable Evolution, and Fossil Record of West Indian Woodpeckers (Family Picidae)
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DISTRIBUTION, PROBABLE EVOLUTION, AND FOSSIL RECORD OF WEST INDIAN WOODPECKERS (FAMILY PICIDAE) ALEXANDER CRUZ Department of Biology University of Colorado Boulder, Colorado 80302 R ESUMEN : La familia Picidae (carpinteros) esta representada en la fauna de las Antillas por dote especies vivientes y dos especies fosiles. Las primeras estan comprendidas en dos generos endemicos y seis generos de distribution mas amplia. Las segundas constan de un genero conocido y otro especimen de afinidad desconocida. Los carpinteros estan mejor representados en Cuba, donde hay cinco especies residentes en comparacion con las Antillas Menores, a excepcion de la Guadalupe, donde no hay especies residences. Durante la epoca glacial del Pleistocene el nivel del agua era inferior al actual y muchas zonas fueron expuestas. Durante esta epoca y posiblemente antes (Plioceno), la mayoria de la avifauna de las Antillas se derivo de las regiones continentals cercanas. Los carpinteros de las Antillas probablemente se originaron en tres diferentes regiones: Norte America, Centro America, y Sur America. HE family Picidae, whose fossil history Cayman, Gran Bahama, Abaco, and T dates back to the Lower Pliocene of Watling’s Island), Centurus radiolatus North America (Brodkorb, 1970) is re- (Jamaica), Centurus striatus (Hispaniola), presented in the West Indian faunal region Melanerpes portoricensis (Puerto Rico and (Fig. 1) by twelve living species, eleven Vieques), Melanerpes herminiero (Guada- resident and one migratory. These com- loupe), Colaptes auratus (Cuba and Grand prise two endemic genera (Nesoctites and Cayman), Colaptes (Nesoceleus) fernandi- Xiphiodiopicus) and six genera of a greater nae (Cuba), Xiphiodiopicus percussus (Cuba distribution (Colaptes, Melanerpes, Centu- and the Isle of Pines), Dendrocopos villosus rus, Sphyrapicus, Dendrocopos, and Cam- (New Providence, Andros, Grand Bahama pephilus). The woodpeckers are best Mores, and Abaco), Sphyrapicus varius represented in Cuba which has five resident (winter resident on many West Indian species in comparison to the Lesser Antilles islands), Campephilus principals (Cuba), which, with the exception of Guadaloupe, and Nesoctites micromegas (Hispaniola and do not have any resident species. Gonave). The species present and their distribution are as follows (Bond, 1956): Centurus PROBABLE EVOLUTION AND FOSSIL RECORD superciliaris (Cuba, Isle of Pines, Grand The West Indies are oceanic islands, which (as fa as is known) have never been Carib. J. Sci., 14 (3-4). Dec. 1974. connected to the surrounding mainland 183 184 A. CRUZ (Myers, 1938; Bond, 1948, 1963; Simpson, (Cuban Parrot), and Centurus superciliaris 1956; Darlington, 1957) and this is also (West Indian Red-bellied Woodpecker). It abundantly demonstrated by the nature of is surprising, as Short (op. cit.) has also the West Indian fauna itself (Myers, op. cit., noted, that Colaptes auratus has extended Bond, op. cit., Simpson, op. cit., and Dar- its range from Cuba to the small island of lington, op cit.). They are not connected Grand Cayman (71 square miles) but to any continental shelf, but are close to apparently has not successfully invaded the banks off Central America, Florida, and other larger and nearby islands (Isle of the Yucatan Peninsula. During the glacial Pines, Jamaica, and Hispaniola) which periods of the Pleistocene the water level have suitable habitats. was lower than that of the present level Dendrocopos villosus (Hairy Woodpecker) and many land areas were exposed, present on some of the Bahama Islands increasing the size of many islands and belongs to a species which is widely reducing the width of many water gaps distributed in North America, ranging (Fig. 1). It was during this time and from Alaska to Panama. The Bahamian probably earlier (Pliocene) that much of races (two subspecies) were probably the West Indian avifauna was derived from derived from Floridian stock sometime in the surrounding mainland area. The West the Pleistocene. Dendrocopos villosus is Indian woodpeckers were probably derived known from the Pleistocene of California from three different sources: North America (Miller and Demay, 1942) and from a Post- (Florida), Middle America, and South Colombian site in Abaco Island, Bahamas America. (Conklin, 1970). The Bahamas also shares The three non-endemic species of wood- a number of other birds in common with peckers of the Antillean faunal region Florida, namely Sitta pusilla (Brown- (Colaptes auratus, Dendrocopus villosus, headed Nuthatch) and Dendroica dominica and Campephilus principals) appear to (Yellow-throated Warbler). have been derived from North America Campephilus principals bairdii (Cuban (Bond, 1963). The genus Colaptes probably Ivory-billed Woodpecker) is considered by dates back to the Lower Pliocene of North Bond (1960) as a race of C. p. principals America. Wetmore (1931) examined a (North American Ivory-billed) which was Pliocene picid ulna from Nebraska and formerly found in the southeastern United determined that it was related to the States. The North American form is present day species of Colaptes. Short probably extinct and the Cuban form is (1965) states: “It (the picid ulna) is extremely rare. In 1956 it was reported colaptine and indistinguishable from those that twelve or thirteen occurred in Oriente of Colaptes melanochloros and Colaptes Province, Cuba (Lamb, 1957). In 1967 two auratus mexicanoides.” The earliest record were reported just to the south of Cupeyal of Colaptes auratus from the West Indian (Oriente); in 1968 a female was seen in the region is from the Pleistocene of Great mountains northwest of Cupeyal (Bond, Exuma Island (Wetmore, 1937). Inter- 1968). It is thought that no more than six estingly, its modern West Indian range birds currently survive in the woods of Moa extends only to Cuba and Grand Cayman. and Barrocoa, both in Oriente (Garrido, Possibly Colaptes first reach the Baha- 1970). The genus Campephilus dates back mas (Great Exuma Island) from Florida to the Upper Pliocene of Texas (Brodkorb, when the Bahama Bank was exposed 1970). There is no fossil record for the (Fig. 1) and from there flew to Cuba which Cuban Ivory-billed, but the North American was very close at this time. The Grand form has been reported from prehistoric Cayman form of Colaptes auratus probably sites in Ohio, Illinois, West Virginia, and reached the island from Cuba during the Georgia (Brodkorb, 1971). As with the late Pleistocene. Grand Cayman also two preceding species, the Cuban Ivory- shares a number of other birds in common billed was probably derived from North with Cuba, namely Melopyrrha nigra American stock (Florida). Other birds (Cuban Bullfinch), Amazona leucocephala shared by Cuba with North America are DISTRIBUTION, PROBABLE EVOLUTION 185 Grus canadensis, Agelaius phoeniceus (also of the centurine woodpeckers in the on some of the Bahama Islands), and West Indies, especially on Cuba little more Sturnella magna (which occurs to northern can be said about its fossil history and South America). evolution. Two endemic species of woodpeckers also It is interesting to note that the avifauna appear to have been derived from North of the Bahamas was richer during the America. There are Colaptes (Nesoceleus) Pleistocene than it is at present. Brodkorb fernandinae (Fernandina’s Flicker) and (op. cit.) notes that nine of the twenty-one Centurus superciliaris (West Indian Red- resident Pleistocene birds identified from bellied Woodpecker). Short (op. cit) feels New Providence, Bahamas, are extinct. that pre-fernandinae flicker stock may Among those recorded are Centurus su- have reached Cuba. from the ‘north (Flo- perciliaris, where no living species now rida) in the middle or late Pliocene. Until occur. Also reported is a paleospecies of recently Colaptes (Nesoceleus) fernandinae woodpecker, Bathoceleus hyphalus, which was placed in its own monotypic genus differs in generic characters from other Nesoceleus, but Short (op. cit.) notes that West Indian woodpeckers. Brodkorb the morphological differences from Co- (op. cit.) notes that during the early laptes are not of sufficient magnitude to portion of the Wisconsin (fourth) glacial warrant maintaining it in the monotypic state the sea dropped to the ten fathom genus Nesoceleus and he places it in the mark exposing the Great Bahama Bank genus Colaptes. (Fig. 1) and uniting Andros, Bimini, New The members of the genus Centurus in Providence, Exuma, Cat Island, Eleuthera, the West Indies probably originated from and Long Island into one large island and North America and Central America. The providing many suitable habitats. The Centurus superciliaris group is similar to rising of the sea during the interglacial Centurus carolinus of the eastern United period, decreasing the size of the island States. Selander and Giller (1963) examined and destroying many suitable habitats, a large series of specimens of both species could possibly have been the cause of the and found many similarities between them. extinction of these species. My own field experience with the Grand Cayman form (C. s. caymanerzsis) and the The fossil record of Centurus radiolatus Florida C. carolinus tends to support this (Jamaican Woodpecker) is unkown. It conclusion. The probable origin of the closely resembles the Centurus aurifrons Centurus super ciliaris group is North group, especially C. aurifrons dubius of the America. There are two Pleistocene records Yucatan Peninsula, Tabasco, northern for Centurus superciliaris, both from the Chiapas, northern