DISTRIBUTION, PROBABLE EVOLUTION, AND FOSSIL RECORD OF WEST INDIAN (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 portoricensis ( and (Fig. 1) by twelve living , 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

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(Myers, 1938; Bond, 1948, 1963; Simpson, (Cuban Parrot), and Centurus superciliaris 1956; Darlington, 1957) and this is also (West Indian Red-bellied ). 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 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 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. 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 () 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 Guatemala, and Cozumel Bahamas. Wetmore (1937) recorded this Island. Selander and Giller (1963) note species from Great Exuma Island and that the Jamaican species is similar to Brodkorb (1959) recorded it from New C. aurifrons dubius, “showing in its white Providence. On both of these islands it forehead and sides of the head and in no longer occurs, but it presently occurs its dark under parts ‘exaggeration’ of on Grand Bahama, Abaco, and Watling’s characters present in that race of C. auri- Island (all in the Bahama group), and it frons.” Pre-radiolatus stock may have been is also present on Cuba, Isle of Pines, and derived from this species. The invasion of Grand Cayman. Protosuperciliaris stock pre-radiolatus stock must have been early probably first reached the Bahamas and enough to provide sufficient time for the from there extended its range to Cuba. Of development of a distinct species. There equal probability is that pro-superciliaris are other probable immigrants from Cen- stock first reached Cuba and from there tral America (Honduran-Nicaraguan bulge) extended its range to the Bahamas. The that may have reached Jamaica in Isle of Pines and Grand Cayman forms this fashion. Bond (1963) and Lanyon were most likely derived from Cuban stock. (1967) note that most Jamaican endemics, Until more is known about the fossil history including Platypsaris niger and Myiopa- 186 A. CRUZ gis cotta, are representative of Middle and the other were united American genera. into a single land mass (Fig. 1) which was Centurus striatus (Hispaniolan Wood- approximately twice the area of present- pecker) has been recorded from the Late Puerto Rico (Heatwole and MacKenzie, Pleistocene of Hispaniola (Bernstein, 1965). 1967) . It was probably during this period Its probable origin is hard to determine, that the extended since it differs from living members of the its range eastward reaching present-day genus Centurus in coloration and in certain Vieques and St. Thomas. The rising of the structural aspects. The nearest mainland sea during the interglacial period decreased area is the Guajira peninsula of Colombia the size of Puerto Rico and left about 15 and its ancestors may have arrived from offshore islands of various sizes to the this area as a waif. Of equal and possibly north and south of Puerto Rico. greater probability is that it may have In habits and mannerisms the Puerto been evolved from some proto-striatus Rican Woodpecker (M. portoricensis) re- stock reaching Hispaniola from Central sembles the North American Red-headed America by Way of Jamaica. Lanyon Woodpecker (M. erythrocephalus), but (op. cit.) notes that: geologists are in no possibly because M. portoricensis has no agreement as to what extent or in what serious competitors in its woodpecker- manner Jamaica, Hispaniola, and Cuba feeding niche it shows a greater diversity were connected, if at all. However, in its foraging and feeding behavior than Darlington (1938) has interpreted faunal M. erythrocephalus (personal observations). relationships among these islands as M. erythrocephalus shares its habitat with evidence that the linkage has been six or seven other woodpeckers in North primarily from Jamaica to Hispaniola and America, whereas M. portoricensis is the to Cuba ...... It seems likely that Jamaica only resident woodpecker on Puerto Rico. and Hispaniola were connected, or nearly Bond (1968) notes that it is particularly so, ...... ” He further notes that: “There are close structurally to M. erythrocephalus. other probable immigrants from Central M. portoricensis was probably derived from America that may have used Jamaica as North American melanerpine stock as a their route of entry into the West Indies waif, but further studies should be under- and then spread to other Greater Antillean taken to determine its relationships with islands. It is noteworthy that some of members of this genus in Central and these, including Elaenia fallax and Nycti- South America. M. portoricensis has been bius griseus, are found only on Jamaica recorded from Pre-Colombian sites (Cueva and Hispaniola, which further suggests a Clara and Cueva Catedral) in Puerto Rico closer faunal link between those islands (Wetmore, 1922), and M. erythrocephalus than between Jamaica and Cuba.” is known from the Pleistocene of Florida Another Picidae, genera and species not (Brodkorb, 1957, and Ligon, 1966). known, has been described by Bernstein Melanerpes herminieri is the only (1965) from the Late Pleistocene of His- resident woodpecker of the Lesser Antillean paniola from a tibiotarsus. It differs from region, occurring only on Guadaloupe. Its Centurus striatus in size and structure, and origin are completely obscure, but it could it is similar to C. carolinus in having a possibly have evolved from M. portoricensis blade-like attachment for the oblique stock reaching Guadaloupe from Puerto ligament. Rico during the Pleistocene sea floor The source of the West Indan Mela- lowering, when the latter was twice its nerpes woodpeckers, M. portoricensis and present size and extended further eastward. M. herminieri is now completely obscure. At this time the distance between Puerto M. portoricensis occurs on Puerto Rico and Rico and Guadaloupe was 225 miles Vieques, and it is said to have inhabited (approximately) and would have facilitated St. Thomas (Bond, 1956). At maximum sea over-water dispersal. Bond (1956) observes level lowering during the Pleistocene, that M. herminieri is more closely related Puerto Rico, Culebra, Vieques, St. Thomas, to M. portoricensis than is the latter to the DISTRIBUTION, PROBABLE EVOLUTION 187

North American M. erythrocephalus. Other nus which is well represented in South Lesser Antillean birds which probably have America, where twenty-five species are had their origin in the Greater Antilles found; one of which extends its range into (Puerto Rico) are Myiarchus oberi from Honduras (De Schauensee, 1966). The M. antillarum stock from Puerto Rico ancestral stock of Nesoctites was probably (Lanyon, 1967) and Geotrygon mystacea of a member of the genus Picumnus that the northern Lesser Antilles. Bond (1963) arrived as a waif fauna from either South states: “Geotrygon mystacea was clearly America or Central America by way of derived from, and is often considered Jamaica. This stock, isolated on the island nonspecific with, the Greater Antillean of Hispaniola, underwent speciation and G. chrysia . . . . . it has spread south to evolved differences to place it in a St. Lucia, but has never been reported monotypic genus. Its ancestors probably from the southernmost islands.” Also arrived during the Pliocene since it has had Dendroica adelaidae which occurs on Puerto sufficient time to differentiate at the genus Rico, Vieques, Barbuda, and St. Lucia. level. The earliest and only fossil record The source of Xiphiodiopicus percussus known is from the Pleistocene of Hispaniola () belonging to (Bernstein, 1965). the endemic monotypic genus Xiphiodiopi- cus is even more completely obscure. The ACKNOWLEDGEMENTS invasion of Cuba by proto-Xiphiodiopicus stock must have been early enough to I should like to express my gratitude to provide sufficient time for obliteration of Dr. Pierce Brodkorb, Department of Zoology, close affinities with other West Indian University of Florida, for the use of his woodpeckers and to evolve into a monotypic bibliographical material, especially on fossil genus. Goodge (1972) notes: “The Cuban records. Support during this investigation Green Woodpecker is similar to Dendroco- came from a Ford Foundation Fellowship pos woodpeckers in having a short glandula awarded to the author. picorum, a posterior origin for the dermo- temporalis, poorly developed geniothyroid, interdigitation of the tracheohyoids, origin LITERATURE C ITED of sternotrachaealis from the first rib, and the presence of an interorbital foramen. BERNSTEIN , L. 1965. Fossil birds from the Like Mesopicos (which is considered closely Dominican Republic. Quart. Journ. Fla. Acad. Sci., 28: 271-284. related to Dendrocopos by Goodge), an BOND, J. 1948. Origin of the fauna of accesory semitendinosus is present. As in the West Indies. Wilson Bull., 60: 207-229. melanerpine woodpeckers ventral and — 1956. Check-list of birds of the West Indies. lateral parts of trachealis are absent. Fourth edition. Philadelphia, Academy of A primitive feature is the origin of serratus Natural Sciences of Philadelphia. — 1963. Derivation of the Antillean avifauna. anterior from the last cervical and first Proc. Acad. Nat. Sci. Philadelphia, 115: thoracic ribs. Perhaps Xiphiodiopicus is 79-98. an isolated remnant of the ancestral — 1968. Check-list of birds of the West Indies, stock that gave rise to Dendrocopos and 13th Supp., p. 7, Philadelphia, Academy of its relatives. No morphological evidence Natural Sciences of Philadelphia. BRODKORB , P. 1957. New passerine birds from indicates that Xiphiodiopicus is a recent the Pleistocene of Reddick, Florida. Journ. offshoot from Dendrocopos or the melaner- Paleont., 31: 129-138. pines.” - 1959. Pleistocene birds from New Providence Island, Bull. Florida State Mus., 4: 269-291. Neosctites migromegas (Antillean ) — 1970. The Paleospecies of Woodpeckers. from Hispaniola and Gonave Islands is the Quart. Journ. Fla. Acad. Sci., 33: 132-136. only member of the subfamily Picumninae -- 1971. Catologue of fossil birds, Part 4 found in the West Indian region. Nesoctites (Columbiformes through ). Bull. Florida State Mus., 15: 163-266. is an endemic monotypic genus, but it is CONKLIN, K. 1970 (1071). Post-Columbian closely related to the other members of birds from Abaco Island, Bahamas. Quart. this subfamily, especially the genus Picum- Jour. Fla. Acad. Sci., 33: 237-240. 188 A. CRUZ

D ARLINGTON , P. J. 1938. The origin of the M ILLER, L. AND D EMAY. 1942. A new fossil bird fauna of the Greater Antilles, with discus- locality. Condor, 44: 283-284. sion of dispersal of over water and M YERS, G. S. 1938. Fresh-water fishes and through the air. Quart. Rev. Biol, 13: West Indian zoogeography. Ann. Rept. 274-300. Smithsonian Inst. for 1937, pp. 339-364. — 1957. Zoogeography: the geographical SELANDER, R. K. AND D. R. GILLER. 1963. Species distribution of animals. New York, John limints in the woodpecker genus Centurus Wiley and Sons. (Aves). Bull. Amer. Mus. Nat. Hist., 124: G OODGE. W. R. 1972. Anatomical evidence for 213-274. phylogenetic relationships among wood- S HORT, L. L., JR. 1965. Variation in West peckers. Auk, 89: 115-129. Indian flickers (Aves, Colaptes). Bull. H EATWOLE, H. AND F. MACK ENZIE. 1967. Herpe- Florida Sate Mus., 10: 1-42. togeography of Puerto Rico. IV. Paleogeo- S IMPSON , G. G. 1956. Zoogeography of West graphv. faunal similarity and endemism. Indian land mammals. Amer. Mus. Novita- Evolution, 21: 429-438. tes, no. 1759: 28 pp. L AMB. G. R. 1957. ICBP Research Report, W ETMORE, A. 1922. Bird remains from caves No. 1. of Puerto Rico. Bull. Amer. Mus. Nat. Hist., L ANYON, W. E. 1967. Revision and probable 46: 297-333. evolution of the Myiarchus flycatchers of — 1931. Record of an unknown woodpecker the West Indies. Bull. Amer. Mus. Nat. from the Lower Pliocene. Condor, 33: Hist., 136: 331-370. 255-256. L IGON, J. D. 1966. A Pleistocene avifauna from — 1937. Bird remains from cave deposits on Haile. Florida. Bull. Florida State Mus.. 10: Great Exuma Island in the Bahamas. Bull. 1-31. Mus. Comp. Zool., 80: 427-441.

F IGURE 1. The West Indian Islands and their banks (in black). The submerged portions of the island banks are show out to the 100-fathom contour (in black). During the Pleistocene these banks were exposed. Solid line comprises the West Indies as a faunal region