A JOURNAL OF AVIAN BIOLOGY

Volume 96 Number 3

The Condor96571-589 Q The Cooper Omithologkzd %cietY 1994

NEW INFORMATION ON THE LATE FROM SAN JOSECITO CAVE, NUEVO LEON, ’

DAVID W. STEADMAN New York State Museum, The State Education Department, Albany, NY 12230

JOAQUIN ARROYO-CARRALES Museum of Tech University,Lubbock, TX 79409 and Laboratorio de Paleozoologia,Subdireccion de ServiciosAcademicos, Instituto National de Antropologiae Historia, Mexico

EILEEN JOHNSON Museum of Texas Tech University,Lubbock, TX 79409

A. FARIOLA GUZMAN Laboratorio de Paleozoologta,Subdireccibn de ServiciosAcademicos, Instituto National de Antropologiiae Historia, Mexico

Abstract. We report 90 bones representing 18 speciesfrom recent excavations at San Josecito Gave, Nuevo Le6n, Mexico. The new material increasesthe avifauna of this rich late Pleistocenelocality from 52 to 62 .Eight of the 10 newly recorded taxa are extant; each is either of temperate rather than tropical affinities (such as the American Woodcock Scolopax minor and Pinyon Jay Gymnorhinuscyanocephalus) or is very wide- spreadin its modem distribution. The two extinct taxa are a (Ciconia sp. or Mycteria sp.) and Geococcyxcalifornianus conklingi, a large temporal subspeciesof the Greater Road- runner. In this region of the Sierra Madre Oriental (about lat. 24”N, long. lOO”W, elev. 2,000-2,600 m). the avifauna was a mixture of speciesthat to&y prefer coniferous or pine-oak forests/woodlands,grasslands/savannas, and wetlands. As with var- ious late Pleistoceneplant and communities of the United Statesand Mexico, no clear modem analog exists for the late Pleistoceneavifauna of San JosecitoCave. Key words: Late Pleistoceneavzfaunas; Mexico; historicalbiogeography; extinct species; temperate/tropicaltransition.

INTRODUCTION est” (Muller 1939), which is the eastern equiv- SanJosecito Cave is located on a steep western alent of “Madrean evergreen woodland” in the flank of the Sierra Madre Oriental, southern (Brown 1982). Within Nuevo I&n, Mexico (lat. 23”57’21”N, long. the 2 km radius, which would accomodate the 99”54’45”W, elev. about 2,250 m; Fig. 1). The primary home range of most vertebrate species site lies in the Municipio de General Zaragoza, whose bones have accumulated in the cave, the about 1 km SSW of Ejido San Josecitoand 8 km elevation varies from 2,000 to 2,600 m. Only SW of Aramberri. The primary vegetation type about 20 km south of San JosecitoCave is Cerro within a 2 km radius of the cave is pine-oak forest Pefia , at 3,650 m one of the tallest peaks and woodland, classifiedas “montane mesic for- in the Sierra Madre Oriental. During four trips in 1988-l 990, researchteams from Mexico and the United Statesbegan a pro- I Received 3 January 1994. Accepted 24 April 1994, gram of carefully evaluating, mapping, and

I5771 578 DAVID W. STEADMAN ET AL.

n Potrero de Zamora w Agua Delgada

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012345 23”55’ FIGURE 1. The location of San Josecito Cave, Sierra Madre Oriental, Nuevo Lebn, MCxico, in relation to local ejidos (squares)and towns (circles). excavating the fossiliferous sediments of San Jo- while bones were accumulating in the late Pleis- secito Cave (Arroyo-Cabrales 199 1; Arroyo-Ca- tocene, it seemslikely that vertical entrancespro- brales and Johnson, in press;Arroyo-Cabrales et vided the only accessto the cave. al. 1989, 1993). Their goal was to improve the Stock and his crew excavated and removed understanding of the stratigraphy, chronology, fossiliferous sediments to an average depth of and taphonomy of San Josecito Cave’s impor- about 12 m below the original sediment surface tant late Pleistocenevertebrate fauna, which had of the cave. They recoveredabout 100,000 bones, been known previously only from material ex- now housed in the Natural History Museum of cavated under the direction of Chester Stock, Los Angeles County (LACM), . California Institute of Technology, from 1935 to Approximately 108 speciesof late Pleistocene 1941 (Stock 1943). vertebrates are known from San Josecito Cave San JosecitoCave is developed in folded Late (Arroyo-Cabrales and Johnson, in press).Stock ’s Jurassic or Early Cretaceous limestone. It is a original excavations yielded bones of these 19 single-drop fissure with three natural entrances speciesof extinct characteristic of the that descend vertically 12 to 30 m to a single late Pleistocene: the bats Desmodusstocki, Ple- main room that is 34 m long by 25 m wide. No cotustetralophodon, ground slothsMegalonyx sp., horizontal or walk-in entrancesexist today. Even Nothrotheriopsshastensis, rabbit Sylvilagus leo- PLEISTOCENE BIRDS FROM SAN JOSECITO CAVE 579

nensis,pocket gopher Orthogeomysonerosus, tropologia e Historia (INAH), Mexico City (Ar- canids Canis dirus, Cuon alpinus,bear Trem- royo-Cabrales and Polaco 1992). arctosfloridanus, cats Felis atrox, Smilodonfa- Unless cited otherwise, the information on talis, horse Equusalaskae, peccary Platygonus generalized modem distributions of Mexican sp., camel Camelopssp., cervids Navahoceros birds is from Friedmann et al. (1950), Blake fricki, Odocoileussp., pronghorn Stockoceros (1953), A. Miller et al. (1957), Petersonand Chalif conklingi,and bovids Euceratheriumsp., Ore- (1973), AOU (1983), and A. Phillips (1986). We amnoscf. 0. harringtoni.Among the 10 species have found no studies of modern birds within of mammals identified thus far from the new 50 km of San JosecitoCave. Birds from various excavation are three extinct species:Desmodus elevations and habitats within 175 km of the site stocki, Nothrotheriopsshastensis, and Equus have been reported by J. Phillips (19 1 l), Sutton alaskae(Arroyo-Cabrales et al. 1993). and Burleigh (1939), Burleigh and Lowery (1942), L. Miller (1943) reported 43 speciesof birds Sutton and Pettingill (1942, 1943), Moore (1947), from Stock’s excavations. Subsequent exami- Eaton and Edwards (1948), Sutton et al. (1950), nation of the 2,100+ bird bones available to Robins and Heed (195 l), Van Hoose (1955), Miller has increased the San Josecito Cave avi- Packard (1957), Zimmerman (1957), Martin de1 fauna to 52 species(Howard 1971; Olson 1974, Campo (1959), Ely (1962) Contreras-Balderas 1984; Arroyo-Cabrales and Johnson, in press; (1973, 1978,1988,1992),andF. Webster(1974). see Table 1 herein), dominated by scavengers While each of these papers helps to interpret the (teratoms, , ravens, certain and biogeography and paleoecology of the San Jo- falcons) and predators (, certain hawks and secito Cave avifauna, the five described in the falcons). next paragraph are most pertinent because of similarities in habitat and elevation. MATERIALS AND METHODS From 12 to 25 April 1941, the field party of The new excavations consist of a 1.0 x 0.3 m Burleigh and Lowery (1942) studied birds in the unit (designated 533N 197E, thickness 1.26 m), pinyon-juniper-oak woodlands at Diamante Pass, a 0.5 x 0.5 m unit (534N 197E; thickness 0.42 Zapalinamt Mountain, Sierra Guadalupe, south- m), anda0.3 x 0.4 munit (542N 196E; thickness easternCoahuila (elev. about 2,100-3,000 m, lat. 0.23 m). The first two units are stratigraphically 25”20’N, long. 100”55’W, about 175 km NNW equivalent to the deepest sediments excavated of San JosecitoCave). This region had been vis- by Stock,based on in situcorrelations with Stock’s ited briefly (6 March 1938) by Sutton and Bur- system of vertical control (marks chiseled on the leigh (1939). Ely (1962) also studied birds in this cave’s wall), supplemented by his field corre- region at various times from 24 December 1957 spondenceand photographs(Arroyo-Cabrales et to 13 June 1959, covering a broader geographic al. 1993). Unit 542N 196E corresponds to the and altitudinal range. As summarized by Sutton uppermost strata from the cave’s original sedi- and Pettingill (1943), Robert B. Lea and Dwain ment surface, 12 m above the other two units. W. Warner recorded birds from 7 to 14 March The radiocarbon chronology of the San Jose- 194 1 near Galeana, southern Nuevo Leon (elev. cite Cave bone deposit is only partially under- about 2,000-2,500 m, lat. 24”50’N, long. stood. Humate-based radiocarbon dates suggest 100“05’W, about 100 km NNW of San Josecito that units 533N 197E and 534N 197E are about Cave). Also near Galeana, Contreras-Balderas 27,000 to 45,000 years B.P., while unit 542N (1992) studied birds in a pine-juniper-creosote- 196E is latest Pleistocene, most likely between bush community (Pinus ponderosa-Juniperus about 11,000 and 27,000 years B.P. (Arroyo- monosperma-Larreatridentata) at 2,100 m elev., Cabrales et al., unpubl. data). lat. 24”41’N, long. lOO”12’W (about 80 km NNW Identification of avian (by DWS) is based of San Josecito Cave) and a creosotebush-yucca on comparisons with modem skeletal specimens community (Larrea tridentata-Yucca jilzjera) at in the collections of the New York State Museum 1,990m elev., lat. 24”43’N, long. lOO”14’W (about (NYSM) and the National Museum of Natural 85 km NNW of San Josecito Cave. History, Smithsonian Institution (USNM). The recently excavatedbones from San JosecitoCave RESULTS are catalogued in the collections of the Labora- Of the 90 bird bones from the new excavations torio de Paleozoologia,Instituto National de An- that can be identified at least to the ordinal level 580 DAVID W. STEADMAN ET AL.

(Table l), 73 are from unit 542N 196E, 16 are were recorded by Sutton and Pettingill (1943) or from 534N 197E, and one is from 533N 197E. Contreras-Balderas (1992). The 39 specimens identified to or species Grassland speciesmake up another very strong represent 18 speciesof birds (12 non-, component (24 species, including 14 listed as six passerines),among which are 32 bones from well in another habitat category).Especially good unit 542N 196E, six from 534N 197E, and one indicators of grassland are Buteo cf. B. regalis, from 533N 197E. The speciescompositions of Circus cyaneus, Numenius cf. N. americanus, these three sub-assemblagesmay be similar, al- Burhinus cf. B. bistriatus, Asio jlammeus, and thoughthis cannot be determined with suchsmall Sturnella sp. These species,however, do not nec- samples. The use of fine-mesh screenis reflected essarily suggesttreeless prairies. They also may in the dramatically improved recovery of non- occurin grassyareas with scatteredtrees or shrubs, Corvuspasserine bones, which make up 67% (60 such as alpine savannahs, open pine-oak wood- of 90) of the bones in our sample, contrasting lands, sagebrush steppe, or even grassy desert- with the “very few” non-Corvus passerinebones scrub such as occurs today in much of the Chi- mentioned but not identified by L. Miller (1943). huahuan Desert. Of San Josecito Cave’s 24 Fifty-one of the 60 passerinebones are from small “grassland” species, 12 were recorded from to medium-sized species that are very difficult southeastern (Burleigh and Lower-y to identify. 1942, Ely 1962) and six from near Galeana, Nue- The two most numerous speciesin our sample, vo Leon (Sutton and Pettingill 1943, Contreras- Asio otus and Cyrtonyx montezumae, were also Balderas 1992). common among the 2,100 + bird bones from San The San Josecito Cave assemblage also in- Josecito Cave studied by L. Miller (1943). Both cludes 12 wetland species (, heron, four of these species are recorded regularly in late ducks, three rails, three shorebirds). Except for Pleistocenecave assemblages(Mead et al. 1984). a few small perennial streams, wetlands are ab- Remarkably, however, 10 of the 18 species in sent in the mountainous terrain near San Josecito our sample are not among those identified by L. Cave today. None of the 12 wetland specieswas Miller (1943). Furthermore, our sample does not recorded by Burleigh and Lower-y(1942), where- include many of the most common speciesin the as Sutton and Pettingill (1943) and Contreras- original fauna1 assemblage, such as Coragyps Balderas (1992) noted only two of them. atratus, Aquila chrysaetos, Caracara plancus, There are no previous modem or prehistoric Falco mexicanus, the peculiar extinct records in Mexico for the American Woodcock Meleagris crassipes(see Rea 1980, Steadman Scolopax minor. The modem winter and breed- 1980), Tyto alba, Bubo virginianus, or Strix oc- ing ranges of S. minor do not extend south of cidentalis. Corvus corax represented more than Texas (AOU 1983), about 500 km and 800 km, half the bones reported by L. Miller (1943), but respectively, northeast of southern Nuevo Leon. only a single bone in our sample. The absence The preferred habitats of S. minor, whether nest- or scarcity of these speciesin our sample may be ing or wintering, are moist woodlands with good an artifact of small sample size, or it may reflect soil cover or brushy swamps. a genuine fauna1 change related to differencesin Within Mexico, the Pinyon Jay Gymnorhinus taphonomy (how the cave was sampling birds) cyanocephalusbreeds only in the mountains of or chronology (potential changesin climate and northern , with occasional non- vegetation). Further excavation will test these breeding records for the northern Sierra Madre proposals. Occidental in and (Moore Certain modern speciesof birds are excellent 195 1, Selander 1956). The nearest resident pop- indicators of habitat today, and as such are im- ulations are in the mountains of south-central portant in reconstructing late Quatemary habi- , about 1,000 km NW of southern tats. At least 26 speciesofbirds from San Josecito Nuevo Leon. The occurrence of G. cyanoceph- Cave inhabit pine-oak forests or woodlands of alus at San JosecitoCave undoubtedly is related Mexico or southwestern United States today to the expansion of coniferous woodlands (es- (category MW in Table 1). Of these, 13 species pecially pinyon-juniper) in northern Mexico dur- were recorded from southeastern Coahuila by ing the late Pleistocene (Van Devender 1990). Burleigh and Lowery (1942) and Ely (1962). Near Within Nuevo Leon, the Scrub Jay Aphelo- Galeana, Nuevo Leon, 12 of the 26 MW species coma coerulescens(= A. caltjornica of A. Phillips PLEISTOCENE BIRDS FROM SAN JOSECITO CAVE 581

TABLE 1. Birds from San JosecitoCave, Nuevo Leon, Mexico. The identification and nomenclatureof certain taxa reported by L. Miller (1943) for the original excavations are modified accordingto Howard (197 l), Olson (1974, 1984), AOU (1983), Arroyo-Cabrales and Johnson(in press),specimen labels in LACM, and herein. The status of some other taxa reported by L. Miller (1943) may changewith further study. Bones from unit 542N are similar in age to those from Stock’s original excavations (11,000 to 27,000 years B.P.), while those from units 533N and 534N are 27,000 to 45,000 years B.P. Numbers are NISP (number of identified specimens).# = NISP in LACM collectionsunreported. Bone element categoriesfor the 1990 excavations:c = carpometacarpus,co = coracoid, f = femur, fu = furcula, h = humerus, m = mandible, mp = manus phalanx, p = pedal phalanx, r = radius,ro = rostrum, s = scapula,t = tarsometatarsus, ti = tibiotarsus, u = ulna, v = vertebra. * = extinct species. + = recorded at Diamante Pass, southeasternCoahuila, by Sutton and Burleigh (1939), Burleigh and Lowery (1942) or Ely (1962). & = recorded near Galeana, southern Nuevo Leon, by Sutton and Petingill(1943) and/or Contreras-Balderas(1992). Distribution categories:MT = mostly tropical (current range is mostly south of 28”N and entirely south of 4o”N); TE = temperate (most of current range is north of 28%); TR = tropical (current range is south of 28%); WI = widespread (both temperate and tropical). Habitat categories(very generalized):G = grasslandor grassysavannah, MW = montane woodland or forest (dominated by some combination of oak, pines, juniper, other conifers); TF = tropical forest; WE = wetland. Distribution and habitat are not designatedfor most extinct speciesbecause of the paucity of late Pleistocene records south of San JosecitoCave.

1990 Excavations

542N :::: Distribution Habitat

Podicipedidae + Podilymbuspodiceps # - - WI WE Pied-billed Grebe Ardeidae Nycticorax nycticorax 1 - - WI WE Black-crowned Night-Heron Aix cf. A. sponsa # - TE WE Wood Duck Anas sp. # - WI WE dabbling duck cf. Histrionicussp. “a few” - TE WE ?Harlequin Duck &Oxyura cf. 0. jamaicensis # - WI WE Ruddy Duck Ciconiidae *Ciconia sp. or Mycteria sp. - 1 - - - stork ra Teratomithidae *Teratornis merriami 15 - - - - Merriam’s Teratom Vulturidae Gymnogypscalifornianus 12+ - TE G,MW California +Coragyps stratus “lots” - - - Black Accipitridae Elanus leucurus 1 - MT G White-tailed Rite +Parabuteo unicinctus *20 - MT G Harris’ Buteo nitidus # - MT MW Gray Hawk Buteo cf. B. regalis # - TE G FerruginousHawk 582 DAVID W. STEADMAN ET AL.

TABLE 1. Continued.

1990 Excavations Oligid 533N TaXOIl excavations 542N 534N Distribution Habitat

+&Buteo jamaicensis - 1 - WI G,MW Red-tailed Hawk mp +Aquila chrysaetos *I5 - - TE G Golden *Spizaetusgrinnelli 2 - - - - Grinnell’s Eagle * Wetmoregypsdaggetti 3 - - - - Daggett’s Eagle *Neogypserrans 7 - - - - Large accipitrid vulture *Neophrontopsamericanus 21 - - - - Small accipitrid vulture f Circuscyaneus 4 - - WI G Northern Harrier Falconidae Caracara plancus “fairly - - - - Crested Caracara abundant” +&Falco mexicanus 40 - - TE G Prairie Falcon +&Falco sparverius 17 3 - WI G American Kestrel c,t,ti Falco sp. # - - - - falcon + Cyrtonyx montezumae 80 3 2 MT MW Montezuma 2h,c C>P Dendrortyx (?) sp. 2 - - TR MW Wood-Quail *Meleagris crassipes 90 - - - - Great-footed Turkey Rallidae limicola - 1 1 WI WE Virginia t t Rallus elegansllongirostris 1 - - - WE King/Clapper Rail +&Fulica americana - - WI WE American Coot Charadriidae Pluvialis sp. # - - WI G,WE plover Scolopacidae Scolopax minor - 2 - TE G,WE American Woodcock c,ti Numenius cf. N. americanus 1 - - WI G,WE Long-billed Curlew Burhinidae Burhinus cf. B. bistriatus 1 - - TR G Double-striped Thick-Knee +&Columba fasciata 9 1 - WI MW Band-tailed Pigeon t +&Zenaida macroura 3 - - WI G,MW Mourning Dove PLEISTOCENE BIRDS FROM SAN JOSECITO CAVE 583

TABLE 1. Continued.

1990 Excavations 533N 542N 534N Distribution Habitat

Psittacidae Rhynchopsittapachyrhyncha 9 - - MT MW Thick-billed + Rhynchopsittacf. R. terrisi # - - TR MW Maroon-fronted Parrot Cuculidae +*Geococcyx californianusconklingi +40 1 - - - Conkling’s Greater h Tytonidae Tyto alba 95 - - WI G,MW Common Barn- Strigidae +Otus asiolkennicotti 3 3 1 WI G,MW Eastern/Western Screech-Owl h,p,t co + Otusjlammeolus 12 - - WI MW Flammulated Screech-Owl Otus trichopsis 6 - - MT MW Whiskered Screech-Owl +Bubo virginianus “several - - WI G,MW Great Homed Owl hundred” *Bubo sp. 1 - - - - Large owl + Glaucidiumgnoma (?) 2 - - WI MW Northern Pygmy-Owl Micrathene cf. W. whitneyi # - - MT G,MW Elf Owl Strix occidentalis 43 - - WI MW Spotted Owl Aegoliusacadicus 9 - - WI MW Northern Saw-whet Owl Asio otus +100 6 1 TE MW Long-eared Owl ~c,~P,u V Asiojlammeus - 1 - WI G Short-eared Owl U Ciccabavirgata +20 - - TR TF Mottled Owl Caprimulgidae + Phalaenoptilusnuttallii 1 1 - WI G,MW Common Poorwill h Picidae +&Colaptes auratus ? - - WI G,MW Northern (Red-shafted?)Flicker Hirundinidae +Hirundo pyrrhonotus - 2 juv - WI G,MW Cliff Swallow hs Corvidae Gymnorhinuscyanocephalus - - 1 TE MW Pinyon Jay h +Aphelocomacoerulescens 1 - WI MW Scrub Jay t + Corvuscorax “more than 1 juv - WI G,MW % the bones” ti 584 DAVID W. STEADMAN ET AL..

TABLE 1. Continued.

1990 Excavations original excavations 542N :z:z Distribution Habitat

Muscicapidae + Turdusmigratorius - 2 - WI MW American Robin mp,t Icteridae +&Sturnellasp. - 2 1 WI G meadowlark p,ti ti Passeriformes “very few” 41 10 - - Small-medium passerines mist mist Totals 2,100+ 13 17 7MT 24 G 9 TE 26 MW 4TR 1 TF 28 WI 12WE

1986:46) occurs only in the mountains of the radius in Ciconia episcopusmicroscelis (female westernmost part of the state, and “possibly very from Zimbabwe, USNM 43 1493) and Mycteria locally elsewhere,fideJ. T. Marshall” (A. Phillips ibis (female from Tanzania, USNM 488 125). As 1986:48). The late Pleistocene range of A. coe- noted by Olson (199 l), the North American late rulescensprobably was less disjunct than it is Pleistocenerecord of still does not include today, again becauseof the greatly expanded ex- any living species. Modern New World storks tent of pinyon-juniper woodlands. prefer low elevation wetlands. The radius from Unless bones of juveniles are recovered, such San JosecitoCave is the highest elevational rec- as for the Cliff Swallow Hirundopyrrhonotus and ord, past or present, for any stork from North or Common Raven Corvus corax, it is difficult to . A carrion-feeding association establish whether the bones of any particular mi- with extinct , much as still occurs in gratory species represent breeding versus non- parts of Africa today, may account for the greater breeding populations. Larger samples may re- late Pleistocene range of North American storks solve this problem eventually in speciessuch as and other avian scavengers(Steadman and Mar- the American Robin Turdus migratorius, for tin 1984, Steadman and Miller 1987). which both resident and migratory populations The second extinct taxon in the new fauna1 exist in the region (J. Webster 1959). While we sample is Geococcyxcalifornianus conklingi, rep- were unable to identify the single bone of Stur- resented by the distal half of a humerus. The nella to species,the breeding form in southern width of this specimen (11.4 mm) resemblesthat Nuevo Leon today is the Western Meadowlark in two late Pleistocene specimens of G. c. conk- S. neglectarather than the Eastern Meadowlark lingi (11.2, 11.7 mm) from Conkling Cavern and S. magna (Lanyon 1962). Shelter Cave, New Mexico (Harris and Crews All but two of the 18 speciesin the new fauna1 1983). These measurements are larger than in sample are extant. The first extinct taxon is an the living G. c. californianus undetermined stork (Ciconia sp. or Mycteria sp.), (8.8-10.6 mm, n = 26; Harris and Crews 1983). represented by the distal end of a radius that is The G. velox of western and smaller than in any extant or extinct New World southern Mexico is even smaller than G. c. cal- speciesof stork. It is also less pneumatic than in ifornianus. Geococcyxconklingi was describedas Jabiru. Compared to the radius from four species an extinct full speciesby Howard (193 1) from each of Ciconia and Mycteria, the San Josecito Conkling Cavern. It is not known outside of Cave specimen has a broader (relative to total southern New Mexico and San Josecito Cave. width of distal end) sulcus tendineus. It is most Harris and Crews (1983) regard G. c. conklingi similar in size and qualitative characters to the as a large geographicand temporal subspeciesof PLEISTOCENE BIRDS FROM SAN JOSECITO CAVE 585 the living G. c. californianus,the larger size being region today (Moore 1947), although some au- the result of to the cooler summers thors (e.g., Hardy and Dickerman 1955) would of the late Pleistocene. If this is so, then G. cal- synonymize R. terrisi with R. pachyrhyncha of ifornianus conklingi probably is ancestral to G. the Sierra Madre Occidental. Burleigh and Low- c. californianus. er-y (1942) reported R. pachyrhyncha from Dia- The same situation is true for at least two other mante Pass, Coahuila. These birds were, in fact, speciesfrom San Josecito Cave for which “ex- R. terrisi, which had not yet been described. tinct” temporal forms (“chronospecies”) have Much of the late Pleistoceneextinction of North been described. Pleistocene specimens of the American birds has been related to dependencies Coragypsatratus often are referred (especially carrion feeding) on the vast large to as C. occidentalisor C. o. mexicanus(see Brod- mammal fauna, most speciesof which died out korb 1964:254, Howard 1968, Steadman and about 12,000 to 10,000 years ago (Martin 1984, Martin 1984). Similarly, Pleistocenebones of the Steadman and Martin 1984, Steadman and Mil- Crested Caracara (Polyborus plancus of AOU ler 1987). Even if we discount the various extinct 1983; Caracaraplancusof Banks and Dove 1992) speciesof birds, no exact modem analog exists have been referred to as P. prelutosusor P. pre- for the late Pleistocene avifauna of San Josecito lutosusgrinnelli (but seeOlson 1976). For neither Cave. In other words, there is no place in Mexico the Black Vulture or Crested Caracara is there or the United Statestoday where, within a radius any evidence that the Pleistocene form is not of several kilometers, one could find the same ancestral to the living species,forming an evo- assemblage of forest/woodland, grassland, and lutionary continuum. wetland species. In recording speciesthat are allopatric today, DISCUSSION AND CONCLUSIONS the San Josecito Cave avifauna resembles the The San JosecitoCave avifauna now consistsof many North American late Pleistocene mam- at least 62 species, the richest late Pleistocene malian “disharmonious faunas.” Each of these assemblage from Mexico. That only 39 newly faunas consistsof speciesof small mammals that identified bones would include 10 speciespre- today occur hundreds of kilometers in different viously unrecorded from the site shows that we directions from the site in question (Lun- are nowhere near reaching the point of dimin- delius et al. 1983; Semken 1984, 1988; Graham ishing returns in sampling the San JosecitoCave 1985; Stafford and Semken 1990). The area that avifauna. Further excavations are sure to dis- most closely represents a modem analog for a close additional species.Five of the newly added disharmonious fauna, even though it may not speciesare passerines(Cliff Swallow, Pinyon Jay, include all mammalian speciesin the late Pleis- Scrub Jay, American Robin, meadowlark sp.). tocene assemblage,is generally to the north or As a larger bone sample becomes available, a northwest of the fossil site. At San JosecitoCave, thorough attempt to identify a significant portion however, it is impossible at present to demon- of the small to medium-sized bones strate contemporaneity of taxa. Becauseso much will be essentialto compare the past and present time is represented, the San Josecito Cave avi- avifaunas of the San Josecito region more pre- fauna is not homologous to a modem species cisely. Although unsurveyed, the modem avi- assemblage.Rather, the San Josecito Cave avi- fauna undoubtedly is dominated by passerines. fauna is a time-transgressive seriesof small sam- The only extinct fossil passerineknown thus far ples of local birds. The new excavations, which from highland Mexico is a bunting (cf. Passerina) feature temperate or widespread species rather from the late Pliocene Yepomera fauna in Chi- than tropical taxa, are a first step toward ad- huahua (Steadman and McKitrick 1982). dressingthis problem. Another aspect of future research is to re-ex- The eastern slopes of the Sierra Madre Ori- amine the material collected by Stock, some of ental are generally wetter, lessseasonal in rainfall which has not been studied for 50 years. Among and temperature, more heavily forested, and the unresolved taxonomic and osteologicalprob- support a more tropical biota than the western lems are the Rhynchopsittaparrots, two species slopes.This difference may be part of the reason of which have been reported from San Josecito why tropical species are poorly represented at Cave. Only one species(R. terrisi) occurs in this San JosecitoCave. One of these tropical species, 586 DAVID W. STEADMAN ET AL. the Mottled Owl Ciccaba virgata, occurs today account for the strong temperate component in only at lower elevations at the latitude (23”57’) the San Josecito Cave avifauna. The cave also of San JosecitoCave. To the south, however, at has yielded bones of the yellow-bellied marmot lat. 2 l”25’ in San Luis Potosi, C. virgata has been (MarmotaJlaviventris), an alpine sciurid that to- recorded in pine-oak forest at 1,500 to 2,100 m day lives only north of Mexico (Frase and Hoff- elevation (Davis 1952). mann 1980; Arroyo-Cabrales and Johnson, in Much remains to be learned about the histor- press). ical vertebrate biogeography of the Sierra Madre At Ranch0 la Brisca, in the western foothills Oriental, the adjacent coastal lowlands to the of the Sierra Madre Occidental in northern So- east, and the Chihuahuan desert to the west. As nora (lat. 30”23’, long. 110”33’, elev. 1,000 m), elsewhere in Mexico (Brown 1985), this region Van Devender et al. (1985) hypothesized that was profoundly influenced by late Quaternary the vertebrate fauna during the Sangamon inter- changesin climate and vegetation (Martin et al. glacial (about 125,000 years ago;Clark 1992) was 1954; Martin 1955, 1958; Martin and Harrell even more tropical than that existing today, with 1957; Bryant and Riskind 1980). Glacial cli- the best modem analog at lower elevations about mates, as opposed to the warm interglacial cli- 240 km to the south-southeast. The deep de- mates of the past 10,000 years, have been op- posits in San JosecitoCave, thus far representing erative for about 85-90% of the past 250,000 only the last glacial interval, eventually may be years (Dansgaard et al. 1993). Late glacial cli- found to sample the previous interglacial, there- mates averaged4-5°C cooler than today virtually by providing data on the waxing and waning of throughout the Neotropics, whereas the associ- temperate versustropical components in the ver- ated moisture regimes were much more variable tebrate communities of northeastern Mexico. (Markgraf 1989, Lozano-Garcia et al. 1993). In the Chihuahuan Desert of Mexico, a cooler and ACKNOWLEDGMENTS perhaps moister late Pleistocene climate led to Steadman’smuseum research was sponsoredby Na- the development of vast wetlands in areasunable tional ScienceFoundation grant EAR-9 11863(Co-PI ’s to sustain such wetlands during the warmer Ho- T. W. Stafford,Jr., R. W. Graham,H. A. Semken,Jr., locene climates of the past 10,000 years (Meyer E. Anderson,DWS). Field work by Arroyo-Cabrales 1973, Van Devender 1990). By contrast, evi- and Johnson was sponsoredby The Graduate School dence from more southern parts of highland and Museum of Texas Tech Universitv (as part of on- going late Quatemary paleoecological-researchof the Mexico suggeststhat glacial times averaged drier Lubbock Lake Landmark), National SpeleologicalSo- than the (Watts and Bradbury 1982, ciety, Cave ResearchFoundation, National Geograph- Lozano-Garcia et al. 1993). ic Society, I.N.A.H., Geological Society of America, Areas of northern Mexico now covered with and American Society of Mammalogists. We appre- ciate accessto the skeletal collection in the Division Chihuahuan desertscrub were occupied until of Birds, USNM (J. P. Angle, M. R. Browning, J. P. about 10,000 years ago by a pinyon-juniper Dean, S. L. Olson) and the vertebrate woodland (Van Devender 1990). Just as with collectionsof LACM (L. Barnes, S. McLeod, D. Whis- birds or mammals, the late Pleistocene plant as- tler). L. Wootan assisted in manuscript preparation. semblages were “disharmonious” because in- Comments by P. S. Martin, S. L. Olson: and T. W. Stafford. Jr. imnroved the manuscrint. We dedicate dependent geographic responses of individual this paper to the memory of Pierce Brodkorb (1908- speciesproduced plant communities without ex- 1992), a pioneer in the study of both modem and pre- act modem analogs(Van Devender 1990). Plants historic Mexican birds. characteristic of higher elevation communities (stunted spruce-pine forests, alpine tundra; see LITERATURE CITED McDonald 1990) also were depressedin eleva- AMERICANORNITHOLXX~ISTS UNION’ [AOU]. 1983. tion as much as 1,000 m lower than at present Check-list of North American birds, 6th ed. Allen (McDonald 1993). Under this cooler than mod- Press, Lawrence, KS. em climatic regime, true alpine conditions may Aaao~o-CABRALES,J. 1991. New studieson San Jo- have extended from the high elevation region of secitoCave, Nuevo Le6n, Mexico. 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