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A late Pleistocene bird community at the northern edge of the tropics in Sonora, Mexico.

INTRODUCTION

For decades, biogeographers have recognized that the highly varied plant and animal communities of northern Mexico ate produced by gradients in temperature and precipitation, which in turn are a consequence of the complex interplay of latitude, longitude, and elevation (White, 1949; Marshall, 1957; Martin, 1958; Martin et al., 1998). In Sonora, for example, warm-temperate desertscrub grades into tropical thornscrub and deciduous forest at elevations from 400 to 1100 m (Wiseman, 1980; Brown, 1982; Felger et al., 2001). Along the major rivers, and especially their perennial tributaries, are rich plant communities that combine floral elements of Sonoran desertscrub, desert grassland, foothills thornscrub, oak woodland and tropical deciduous forest (Figs. 1, 2; Van Devender et al., 2005). The floral diversity in these valleys and canyons is enhanced further by trees characteristic of riparian areas, such as cottonwoods (Populus), willows (Salix), ashes (Fraxinus), walnuts (Juglans) and sycamores (Platanus). Not surprisingly, such mesic areas often teem with birdlife in Sonora today (Russell and Monson, 1998; Flesch, 2008; DWS, pers. obs.).

It is in this general setting that the late Pleistocene fossil site of Terapa was discovered in 1999. Positioned adjacent to the Rio Moctezuma in a region dominated by foothills thornscrub but within 10 km of desert grassland and oak woodland, the Terapa fossil locality was first reported by Mead et al. (2006), who conservatively noted 58 vertebrate taxa (3 fishes, 3 amphibians, 3 reptiles, 24 non-passerine birds and 25 mammals). In this paper we provide osteological, ecological and biogeographic details about the fossil avifauna from Terapa, which through subsequent field and laboratory research now consists of 31 nonpasserine taxa.

Terapa is only the second late Pleistocenc site in northern Mexico where abundant bird fossils have been studied; the other is San Josecito Cave, which is >1100 km southeast in the Sierra Madre Oriental of Nuevo Leon (23[degrees]57'21"N, 99[degrees]54'45"W, elev. 2,250 m; Steadman et al., 1994).

MATERIALS AND METHODS

The fossils were identified by direct comparison with modern bird skeletons in the Florida Museum of Natural History, University of Florida (UF), supplemented with specimens from the Louisiana State University Museum of Natural Sciences (LSUMNS) and the National Museum of Natural History, Smithsonian Institution (USNM). Fossils from Terapa ate curated and catalogued in a temporary archive (with the prefix "Teta-") at the Neogene Vertebrate Paleontology Laboratory, East Tennessee State University. Osteological details for species that are extinct or otherwise distinctive ate provided in the species accounts, with terminology that follows Baumel et al. (1993), supplemented by Howard (1929). MI fossil specimens ate adult (fully ossified) unless designated otherwise. Field methods for excavations at Terapa were described in Mead et al. (2006). A dagger ([dagger]) designates an extinct species. Unless cited otherwise, modern distributions of birds are from Russell and Monson (1998), and late Pleistocene distributions ate from Lundelius et al. (1983).

SITE SETTING, FAUNA, AND CHRONOLOGY

The fossil locality of Terapa (29[degrees]41'N, 109[degrees]39'W, elev. 605 m) lies on a terrace just above the Rio Moctezuma, which is the major northwestern tributary of the Rio Yaqui (Fig. 2). The subtropical climate of this region features hot summers with monsoonal rains, and mild, generally frost-free winters (Fig. 3).

[FIGURE 1 OMITTED]

The geographic and geologic setting, stratigraphy, chronology and overall fossil fauna (plants, invertebrates, vertebrates) of Terapa were described in Mead et al. (2006), from which we briefly summarize some highlights. A Pleistocene lava flow along the Rio Moctezuma impounded a basin that filled with fossiliferous sediments to a depth of 11 m. The impoundment produced a lake and marsh environment adjacent to woodland and perhaps grassland communities. A tropical to subtropical riparian corridor may have persisted from the Gulf of California (350 km downstream from Terapa) up the Rio Yaqui to the Rio Moctezuma including the marsh at Terapa. The occurrences of cf. Crocodylus acutus (American crocodile), Pampatherium (extinct pampathere, or giant armadillo), Glyptotherium (extinct glyptodont) and a capybara (Hydrochoerinae) at Terapa are unique in interior northern Sonora (Mead et al., 2006, 2007).

[FIGURE 2 OMITTED]

Teeth and post-cranial remains of Bison from throughout the deposits indicate that the Terapa local fauna represents the Rancholabrean North American Land Mammal Age of the late Pleistocene. The Rancholabrean is defined to begin with the arrival of Bison sometime between 210 and 160 ka and to terminate at the end of the Pleistocene at 14 to 12 ka (Bell et al., 2004). Although the age of the Terapa local fauna originally was believed to be 440 [+ or -] 130 ka based on a single, low-resolution Ar/Ar date on the basalt dam (Mead et al., 2006), subsequent chronological analyses place the age of the deposit and its fossils at ca. 40 ka.

[FIGURE 3 OMITTED]

RESULTS

The fossil material includes 19 extant species of birds that ave widespread today in or near freshwater of grassland habitats in Sonora, if not most of temperate North America. This group is dominated by species that occur in Sonora as migrants and winter residents (Table 1). Most of these species also ate recorded widely in the late Pleistocene in the United States and, to a much lesser extent, Mexico (Lundelius et al., 1983; Webb et al., 2004). The remaining 12 species ate sufficiently unusual, or out of place, that they warrant special notice.

Plegadis cf. P. chihi. White-faced Ibis. Distal end of tibiotarsus (Teta-111). This fossil agrees with the tibiotarsus of Plegadis chihi in size, being much smaller than in Eudocimus of especially Ajaia (=Platalea). The fossil agrees further with Plegadis and not Eudocimus in its smooth, elongate and concave supratendinal bridge, the narrow internal condyle, and the wide anterior intercondylar fossa. Plegadis chihi is found in Sonora today an uncommon migrant or wintering bird (Fig. 4). The osteologically similar Glossy Ibis, P. falcinellis, occurs in eastern North America. The only previous late Pleistocene report of P. chihi is from Rancho La Brea, California (Howard, 1962). Compared to complete, modern skeletons of various juvenile ibises, the Terapa specimen belongs to a bird that probably was volant, but perhaps too young to migrate, therefore suggesting local breeding. The nearest modern breeding records of P. chihi are from southwestern Arizona and northern Durango, ca. 600 km NW and 600 km SE of Terapa, respectively (Ryder and Manry, 1994; Zaun et al., 2003).

[dagger] Ciconiidae sp. Large stork. Shaft of ulna (Tera-112). This very fragmentary specimen is much larger than the ulna of the living Mycteria americana, the sole species of stork recorded today in Sonora (rare; no breeding records) or anywhere else in northern Mexico. The only storks recorded in North America in the late Pleistocene ate M. wetmorei (Olson, 1991) and Ciconia maltha (Lundelius et al., 1983). Both of these extinct species ate larger than M. americana, as ave the two other extant species of Neotropical storks (Jabiru mycteria, southern Mexico to Argentina; C. maguari, Colombia to Argentina). Other late Pleistocene records of storks from Mexico are Ciconia? sp. from the Tequixquiac local fauna, State of Mexico, and M. wetmorei from the San Marcos local fauna, Jalisco (Howard, 1969). While we classify the unknown stork from Terapa as aquatic (Table 1), it might have been a grassland dweller that fed mainly on carrion as some African storks do today (Steadman and Martin, 1984).

Branta canadensis Canada Goose. Sternal end of coracoid, distal end of ulna, shaft of radius, distal end of tibiotarsus (Tera-113-116). These specimens are referred to Branta rather than to Anser (including Chen) because of these characters: coracoid - medial half of Facies articularis sternalis more excavated (concave); ulna - Tuberculum carpale wider, less excavated on ventral surface; radius - similar placement and development of intermuscular lines; tibiotarsus - general agreement in size and overall morphology. Branta canadensis is a rare wintering species in Sonora today, with only four inland records (Fig. 5). Late Pleistocene records of B. canadensis ave numerous and widespread from ca. 28[degrees] to 43[degrees]N and 82[degrees] to 120[degrees]W.

[dagger] Anabernicula cf. A. oregonensis. Extinct shelduck. Complete coracoid, scapulo-humeral end of coracoid, distal end of tibiotarsus (Teta-6, 57, 117). We refer these specimens to the extinct tadornine genus Anabernicula rather than to Anas because of these characters: coracoid - Sulcus musculo supracoracoidei deep, Cotyla scapularis deep, Processus procoracoideus rugose in dorsal aspect with subparallel lineations, Facies articularis sternalis deep and distinct in dorsal aspect; tibiotarsus - Condylus medialis narrow; in distal aspect, intercondylar region relatively shallow. We agree with Howard (1964) that Anabernicula is similar osteologically to the extant Old World genus Tadorna. The coracoid from Terapa measures 46.1 mm (total length) and 8.4 mm (width across triosseal canal). These measurements in the type series (n = 4) of A. oregonensis from the Fossil Lake local fauna, Oregon (Rancholabrean) are 44.7-52.0 mm and 8.4-9.1 mm (Howard, 1964). The only other Mexican record of Anabernicula is that of "cf. Anabernicula medium-large species" from the early Blancan NALMA (late Pliocene) GTO 26 site, Guanajuato (Steadman and Carranza, 2006). The coracoid from Guanajuato (IGCU 4816-7) is indistinguishable qualitatively from the one from Terapa, and also is similar in size (total length 47.4 mm, width across triosseal canal 8.5 mm). We now refer IGCU 4816-7 (and its associated wing elements) to the A. oregonensis lineage as well, although the species remains uncertain. See Bickart (1990) and Olson and Rasmussen (2001) for a discussion of Anabernicula, a genus badly in need of revision. Aside from Fossil Lake, the only other late Pleistocene record of A. oregonensis is from Dark Canyon Cave, New Mexico, ca. 560 km NE of Terapa.

Ictinia sp. Mississippi/Plumbeous-type kite. Distal half of tarsometatarsus (Teta-119). This specimen is referred to Ictinia, rather than to other genera of small New World accipitrids (Chondrohierax, Elanoides, Gampsonyx, Elanus, Harpagus and Accipiter) by having this unique combination of characters: Foramen vasculare distale wide and long relative to width of Corpus tarsometatarsi (small in Chondrohierax) ; Trochlea metatarsi II extends distad to level of Trochlea metatarsi III (does not extend this far in Elanoides, Elanus, Harpagus or Accipiter) ; Trochlea metatarsi III as wide as long in acrotarsial aspect (narrower in Gampsonyx and Accipiter); Trochlea metatarsi II wide (narrower in Elanus, Harpagus and Accipiter); in medial aspect, plantar extension of Trochlea metatarsi II moderate (greater in Gampsonyx, Elanus, Harpagus and Accipiter. Tera-119 is larger than available specimens of either living species of Ictinia (Table 2), which do not seem distinguishable from each other based on the distal tarsometatarsus. This is the first fossil record of any age for Ictinia. Neither living species is recorded from Sonora in modern times, although I. mississippiensis nests locally southern Arizona and New Mexico (Bolen and Flores, 1993; Parker, 1999). At its western limit in Arizona, the Mississippi Kite prefers riparian woodlands dominated by cottonwoods (Populus fremontii) that are >15 na tall (Glinski and Ohmart, 1983); this riparian tree grows commonly along the Rio Moctezuma at Terapa today. The exclusively Neotropical I. plumbea (Plumbeous Kite) does not nest today north of ca. 23[degrees]N in southern Tamaulipas, Mexico (Howell and Webb, 1995).

[FIGURE 4 OMITTED]

[dagger] Buteogallus (Wetmoregyps) daggetti. Daggett's Eagle or "Walking Eagle." Shaft of tarsometatarsus, pedal phalanx (Tera-122, 136). These fossils are qualitatively very similar to, but much larger than, modern specimens of Buteogallus (Heterospizias) meridionalis. Olson (2007) proposed that the extinct, late Pleistocene accipitrid known as Wetmoregyps daggetti (originally described as Morphnus daggetti Miller 1915) is a "scaled-up version" of the living Savanna Hawk B. meridionalis, which today inhabits tropical savannas and grasslands from Panama to Argentina. A tarsometatarsus of B. daggetti from its type locality (Rancho La Brea, California) is 49% wider than the mean of seven modern USNM specimens of B. meridionalis (Olson, 2007). The width of the tarsometatarsal shaft from Terapa (Tera-122) is 9.91 mm, which is 46% and 60% larger, respectively, than in two modern specimens of B. meridionalis--6.78 mm in UF 33555 ([female]) and 6.18 mm in UF 38903 ([male]). The shaft of the pedal phalanx from Terapa (Tera-136) is 4.86 mm wide, which is 18% and 25% larger, respectively, than in modern specimens of B. meridionalis--4.10 mm in UF 33555 ([female]) and 3.88 mm in UF 38903 ([male]). The extinct Buteogallus (Wetmoregyps) daggetti is known previously from only three other late Pleistocene localities: Rancho La Brea, California (Miller, 1915, 1925); Carpinteria, California (Miller, 1928, 1931); and San Josecito Cave, Nuevo Leon, Mexico (Miller, 1943; Steadman et al., 1994).

[FIGURE 5 OMITTED]

[FIGURE 6 OMITTED]

Aquila chrysaetos Golden Eagle. Scapular/humeral facet of coracoid, distal end of radius (Tera-120, 121). These specimens agree with the coracoid and radius in modern A. chrysaetos in all details. The Golden Eagle is a rare permanent resident of Sonora, mainly from north of Terapa (Fig. 6). Late Pleistocene records of A. chrysaetos are widespread in the western United States; the only other Pleistocene record from Mexico is from Tequixquiac, state of Mexico (Howard, 1969).

Caracara cheriway Crested Caracara. Nearly complete coracoid (Tera-123). This specimen is referred to Caracara cheriway (=Polyborus plancus) rather than to any species of Falco, Herpetotheres, or Milvago because of this combination of characters: in medial aspect, Processus acrocoracoideus and Facies articularis clavicularis elongate rather than wide and blunt; Sulcus musculo suproacoracoidei smooth (not with irregular surface) and slightly concave (not flat); humero-ventral margin of Corpus coracoideus rounded (not sharp). The Crested Caracara is a fairly common resident in Sonora below 1000 m elevation, including a rather isolated modern record near Terapa (Fig. 7). Late Pleistocene records of C. cheriway (often reported as C. prelutosa) are numerous across the southern United States; previous late Pleistocene Mexican records are from Tequixquiac, state of Mexico (Howard, 1969).

[FIGURE 7 OMITTED]

Callipepla (Lophortyx) douglasii Elegant Quail. Distal two-thirds of humerus (Tera-3). This specimen is referred to C. douglasii rather than to similarly sized species of Odontophorinae (Colinus virginianus, Callipepla gambelii, Callipepla squamata) because of these characters: Tuberculum supracondylare ventrale large and ventrally protrudent; Sulcus musculo humerotricipitis deep; Processus flexorius large. The osteological similarity of Callipepla and "Lophortyx" has long been recognized (Holman, 1961:190). Terapa is near today's northern limit (Fig. 8) of C. douglasii, an inhabitant of tropical thornscrub, tropical deciduous forest, mesquite grassland and oak woodland. The northernmost records are both in tropical thornscrub at ca. 30[degrees]20'N; the first is near Cucurpe in the Rio San Miguel drainage (T. R. Van Devender and P. D. Jenkins, pers. comm.), and the second is 3 km south of Arizpe where the Rio Agua Caliente flows into the Rio Sonora (DWS and G. K. Pregill, 3, 4 Feb. 2007). There is no previous fossil history for C. douglasii.

[FIGURE 8 OMITTED]

Gallinula chloropus Common Moorhen. Complete femur (Tera-2). This specimen is referred to G. chloropus using characters in Steadman (1988) and Kirchman and Steadman (2006). There are no modern records of G. chloropus from the upper Rio Yaqui drainage (Fig. 9). The nearest records are from Mututicachi on the Rio Sonora (Russell and Monson, 1998:86) and Cienega Saracachi (Locality 4 in Fig. 1) in the drainage of Rio San Miguel, a major tributary of the Rio Sonora.

Porphyrio martinicus [Porphyrula martinica] Purple Gallinule. Distal end of carpometacarpus (Tera-145). Inclusion of Porphyrula in Porphyrio follows Steadman (1988). This specimen is referred to P. martinicus rather than to North American rallids of roughly similar size (Rallus longirostris, R. elegans, Gallinula chloropus) because the distal metacarpal symphysis is more concave, and metacarpal III is relatively stouter in medial aspect. The only modern record of P. martinicus in Sonora is from Guirocoba (23 Jul. 1933), ca. 340 km south of Terapa (Fig. 9). A widespread neotropical resident, including the Pacific slope of Mexico southward from Sinaloa (Howell and Webb, 1995:243; Russell and Monson, 1998:86), P. martinicus has been recorded as a vagrant hundreds or even thousands of kilometers outside of its normal range (Remsen and Parker, 1990). There are a number of reports of P. martinicus in the past two decades from southern Arizona, a region teeming with birdwatchers, unlike Sonora. The only other late Pleistocene record of P. martinicus is from Ichetucknee River, Florida (Campbell, 1980).

[FIGURE 9 OMITTED]

[dagger] Strix brea La Brea Owl. Proximal end anda shaft of tarsometatarsus, pedal phalanx (Tera-124-126). The tarsometatarsus (section of distal shaft) agrees with that in Strix [varia, occidentalis, virgata] rather than in Bubo [virginianus] in its sharper plantar and acrotarsial margins of shaft. The least width of this shaft is 6.64 mm; this is the only measurement directly comparable to those given by Howard (1933) for the holotype of S. brea (least width of shaft 7.1 mm, proximal width 14.3 mm, distal width 16.4 mm, and total length 67.2 mm). In five modern UF specimens of S. (Ciccaba) virgata from Mexico (2 females, 3 unsexed), this measurement ranges from 5.16 to 5.56 mm, with total length ranging from 43.4 to 50.0 mm. In 10 modern UF specimens of S. varia from Florida, South Carolina, and Pennsylvania, these measurements vary from 5.96 to 6.62 mm and from 53.2 to 60.5 mm. The pedal phalanx (lacking distal end) agrees with that in Strix [varia, occidentalis, virgata] rather than in Bubo [virginianus] in its less angular and less excavated distal margins of the articular surface. The extinct Strix brea is known previously only from the late Pleistocene of Rancho La Brea, California (type locality; Howard, 1933), Carpinteria, California (Van Devender et al., 1985), Dry Cave, New Mexico (Harris, 1970), and Rancho La Brisca, Sonora (Van Devender et al., 1985).

Finally, we note the recovery at Terapa of 62 specimens of small- to large-sized passerine birds (Passeriformes) that represent at least eight species. These fossils will be the subject of a future paper.

DISCUSSION

Paleoclimate, Paleovegetation.--We are just beginning to learn about long-term changes in the vertebrate fauna of northern Mexico. Other than Terapa, the only Rancholabrean local fauna in Sonora that has been studied in detail is Rancho La Brisca (Van Devender et al., 1985; locality 3 of Fig. 1 herein). The absolute ages of these local faunas are poorly understood; neither can be assigned confidently even to a glacial vs. interglacial interval, although Van Devender et al. (1985) suggested that the Rancho La Brisca local fauna was deposited during the last interglacial (ca. 125 ka), because of the presence of subtropical anurans (Bufo cf. B. kelloggi, B. mazatlanensis, Leptodactylus melanonotus) that live today no closer than ca. 200 km south of Rancho La Brisca in the lower Rio Yaqui drainage.

We note, however, that diatoms from a deep sea core in the Guaymas Basin, off the coast from the mouth of the Rio Yaqui, represented subtropical taxa during the most recent full glacial (Byrne, 1982). If winter temperatures in the region were moderated relative to modern ones (see Fig. 3) during glacial intervals, then organisms with northern range limits controlled by winter temperatures may have occurred farther north in the latest Pleistocene than in the Holocene, even if summer temperatures were cooler during glacial than interglacial intervals.

Other evidence of late Quaternary climate change in the southwestern U.S. and northern Mexico is based on plant macrofossils from arid mountain ranges, and pollen or diatom sequences from inland lake basins (Betancourt et al., 1990; Bradbury, 1997; Bradbury et al., 2000). In assessing the past 65,000 y of climate change in northwestern Chihuahua, Metcalfe et al. (2002) stated that "it is clear that the aridity which typifies the area today is the exception rather than the norm ..." The same trend appears to apply to southwestern U.S. (California, Nevada, Utah, Arizona, New Mexico), which was characterized by abundant late Pleistocene pluvial lakes and other wetlands of all sizes (e.g., Oviatt et al., 2003; Menking et al., 2004). Such bodies of freshwater would have sustained breeding waterfowl, many species of which probably were migrants or winter residents at Terapa.

Radiocarbon dated plant macrofossils from late Pleistocene packrat (Neotoma) middens from southern Arizona, southern New Mexico, trans-Pecos Texas, Coahuila, Durango and Baja California all indicate the presence of pinyon pine (Pinus edulis, P. monosperma, etc.) before 12 ka, and in some cases extending back to >30 ka, at elevations much lower than where these conifers live today (Van Devender, 1990a, b; Holmgren et al., 2003). This implies that reduced summer temperatures and dominant winter precipitation characterized northern Mexico prior to 12 ka.

Connin et al. (1998) analyzed the oxygen and carbon isotopes from tooth enamel of megamammals (Mammuthus, Equus, Bison, Camelops, among others) from the southwestern U.S., including many specimens from the border region near Mexico. For the period from 40 to 10 ka, they found C4 plants (arid and tropical grasses, some herbs and shrubs) to have been dominant south of 35[degrees]N. Summer rains, with a tropical/subtropical source, probably were necessary to sustain C4 productivity at levels consistent with the nearly pure C4 feeding by the megaherbivores. This idea is compatible with the vertebrate fauna from Terapa, which overall suggests a tree-lined wetland in a subtropical woodland. That grasses were not in short supply is supported by the occurrence of Bison and two species of horses (Equus excelsus, E. conversidens) (Carranza-Castaneda and Roldan-Quintana, 2007).

Quaternary Biogeography of Birds of Northwestern Mexico.--While the numerous fossil eggshell fragments from Terapa remain unidentified, fossil bones from juvenile birds provide evidence that the following species nested at of very near Terapa in the late Pleistocene--Pied-billed Grebe Podilymbus podiceps, Great Blue Heron Ardea herodias, White-faced Ibis Plegadis cf. P. chihi and American Coot Fulica cf. F. americana. Among these species, only the ibis no longer nests in Sonora.

Of the three non-passerine birds identified to species from fossils at Rancho La Brisca (Van Devender et al., 1985), the Green-winged Teal Arias crecca, and extinct owl Strix brea, were recorded as well at Terapa. Missing thus far at Terapa is the widespread, living species of turkey Meleagris gallopavo, which has been recorded from a Rancholabrean fauna at Arizpe in the Rio Sonora drainage (Cracraft, 1968), and is likely to be discovered at Terapa with additional fossil collecting.

There is no overlap at all among the 31 non-passerine species recorded as fossils at Terapa and the 10 species of non-passerine birds that DWS and G. K. Pregill observed alive there on 22-23 Mar. 2005. Although both the modern and fossil samples are small, and the sampling methods are very different (bone deposition in a wetland vs. intense, systematic birdwatching), these data suggest substantial turnover in the regional bird community since the late Pleistocene.

Acknowledgments.--We thank Arturo Baez, Hector Ruiz Durazo, Santiago Garcia Lopez, Elsa Maria Aruna Moore, and the citizens of San Clemente de Terapa and El Llano. Special thanks go to Sandra L. Swift for countless hours of screen-washing, sorting and curation of fossils. We thank Matthew Reetz and Jessica Oswald for preparing the figures, and Monica Arakaki for kindly translating the abstract into Spanish. For dedicated field and laboratory work, we are grateful to Joaquin Arroyo-Cabrales, Arturo Baez, Eric Baez, Jordan Bright, Mary Carpenter, Fred Croxen, Nicholas Czaplewski, Marci Hollenshead, Margaret Imhof, Darrell Kaufman, Aimee Mead, Gary Morgan, Gregory Pregill, Christopher Shaw, Sandra Swift, Richard White and many students from Northern Arizona University. The manuscript was improved by comments from Thomas Van Devender and two anonymous reviewers.

SUBMITTED 29 OCTOBER 2008

ACCEPTED 17 AUGUST 2009

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DAVID W. STEADMAN (1)

Florida Museum of Natural History, University of Florida, Gainesville 32611

AND

JIM I. MEAD

Department of Geosciences and Don Sundquist Center for Excellence in Paleontology, East Tennessee State University,

Johnson City 37614

(1) Corresponding author: e-mail: dws@flmnh.ufl.edu
TABLE 1.--Birds identified from the Pleistocene (Rancholabrean
NALMA) Terapa local fauna, Sonora, Mexico. ([dagger)], extinct
species. Taxa in brackets [xxx] are not necessarily different from
those identified more specifically. Current status categories: E,
extinct; M, migrant; R, resident; W, widespread, common aquatic
species (not included in Results). Habitat categories (very
generalized): A, aquatic; T, terrestrial (non-aquatic)

                                                     Current
FAMILY                                   Number     status in
Species                                of fossils    Sonora

PODICIPEDIDAE
  Podilyrnbus podiceps                     1        M, R, W
    (Pied-billed Grebe)
  Podiceps nigricollis (Eared              2        M, W
    Grebe)

PELECANIDAE
  Pelecanus erylhrorhynchus (White         1        M, R?, W
    Pelican)

PHALACROCORACIDAE
  Phalaoocorax auritus (Double-            1        M, R, W
    crested Cormorant)

ARDEIDAE
  Ardea herodias (Great Blue               4        M, R, W
    Heron)
  Ardea alba (Great Egret)                 1        M, R, W

THRESKIORNITHIDAE
  Plegadis cf. P. chihi (White-            1        M
    faced Ibis)

CICONIIDAE
  ([dagger]) Ciconiidae sp. (large         1        E
    extinct stork)

ANATIDAE
  Branta canaderu[degrees]is (Canada               4        M
    Goose)
  Anser sp. (Snow or White-fronted         1        M, W
    Goose)
  [Anserinae sp. (goose) ]                [1]       [M]
  ([dagger]) Anabernicula cf. A.           3        E
    aregonensis (extinct shelduck)
  Anas crecca (Green-winged Teal)          7        M, W
  Anas plat yrhynchos/diazi                16       M, R, W
    (Mallard or Mexican Mallard)
  Anas acula (Northern Pintail)            6        M, W
  Anas strepera (Gadwall)                  1        M, W
  Anas clypeata (Northern                  2        M, W
    Shoveler)
  Anas discors/cyanoptera                  7        M, W
    (Blue-winged or Cinnamon Teal)
  [Anas sp. (small; size of A.            [6]       [M]
    crecca, discors, (yanopter-a)]
  Aythya collarzs (Ring-necked             3        M, W
    Duck)
  Oxyura jamaicensis (Ruddy Duck)          1        M, R, W

ACCIPITRIDAE
  Circus ryaneu.s (Northern                1        M
    Harrier)
  Ictinia sp.                              1        --
    (Mississippi/Plumeous-type kite)
  ([dagger]) Buleogallus                   2        E
    (Wetrnoregyps) daggetti
    (Daggett"s Eagle)
  Arluila chiysaetos (Golden               2        R
    Eagle)

FALCONIDAE
  Caracara cheriway (Crested               1        R
    Caracara)

ODONTOPHORIDAE
  Callipepla douglasii (Elegant            1        R
    Quail)

RALLIDAE
  Parzana carolina (Sora)                  1        M, W
  Gallinula chloropus (Common              1        R
    Moorhen)
  Porphyrio martinicus (Purple             1        M?
    Gallinule)
  Fulica cf. americana (American           2        M, R, W
    Coot)

CHARADRIIDAE
  Charadrius vociferus (Killdeer)          1        M, R, W

STRIGIDAE
  ([dagger]) Strix brea                    3        E
    (La Brea Owl)

PASSERIFORMES
  At least 8 species of songbirds          61       ?

                                          Modern
FAMILY                                 record within   Habitat
Species                                   100 km

PODICIPEDIDAE
  Podilyrnbus podiceps                       X            A
    (Pied-billed Grebe)
  Podiceps nigricollis (Eared                X            A
    Grebe)

PELECANIDAE
  Pelecanus erylhrorhynchus (White           X            A
    Pelican)

PHALACROCORACIDAE
  Phalaoocorax auritus (Double-              X            A
    crested Cormorant)

ARDEIDAE
  Ardea herodias (Great Blue                 X            A
    Heron)
  Ardea alba (Great Egret)                   X            A

THRESKIORNITHIDAE
  Plegadis cf. P. chihi (White-             --            A
    faced Ibis)

CICONIIDAE
  ([dagger]) Ciconiidae sp. (large          --            A
    extinct stork)

ANATIDAE
  Branta canaderu[degrees]is (Canada                --            A
    Goose)
  Anser sp. (Snow or White-fronted           X            A
    Goose)
  [Anserinae sp. (goose) ]                  [X]          [A]
  ([dagger]) Anabernicula cf. A.            --            A
    aregonensis (extinct shelduck)
  Anas crecca (Green-winged Teal)            X            A
  Anas plat yrhynchos/diazi                  X            A
    (Mallard or Mexican Mallard)
  Anas acula (Northern Pintail)              X            A
  Anas strepera (Gadwall)                    X            A
  Anas clypeata (Northern                    X            A
    Shoveler)
  Anas discors/cyanoptera                    X            A
    (Blue-winged or Cinnamon Teal)
  [Anas sp. (small; size of A.              [X]           A
    crecca, discors, (yanopter-a)]
  Aythya collarzs (Ring-necked               X            A
    Duck)
  Oxyura jamaicensis (Ruddy Duck)            X            A

ACCIPITRIDAE
  Circus ryaneu.s (Northern                  X            T
    Harrier)
  Ictinia sp.                               --            T
    (Mississippi/Plumeous-type kite)
  ([dagger]) Buleogallus                    --            T
    (Wetrnoregyps) daggetti
    (Daggett"s Eagle)
  Arluila chiysaetos (Golden                 X            T
    Eagle)

FALCONIDAE
  Caracara cheriway (Crested                 X            T
    Caracara)

ODONTOPHORIDAE
  Callipepla douglasii (Elegant              X            T
    Quail)

RALLIDAE
  Parzana carolina (Sora)                    X            A
  Gallinula chloropus (Common               --            A
    Moorhen)
  Porphyrio martinicus (Purple              --            A
    Gallinule)
  Fulica cf. americana (American             X            A
    Coot)

CHARADRIIDAE
  Charadrius vociferus (Killdeer)            X            T

STRIGIDAE
  ([dagger]) Strix brea                     --            T
    (La Brea Owl)

PASSERIFORMES
  At least 8 species of songbirds            ?            ?

TABLE 2.--Measurements (in mm) of the distal tarsometatarsus in Ictinia

                                                     Minimum
                                                   width shaft
                                                   immediately
                                                   proximal to
                      Catalogue                       Fossa
   Species            number     Sex  Locality     metatarsi I

Fossil                Tera-119   --   Sonora          3.78
I. mississippiensis   OF 44986   M    New Mexico      3.30
I. mississippiensis   OF 23824   M    Texas           3.55
I. mississippiensis   OF 21849   --   Florida         3.17
I. mississippiensis   OF 45499   F    Florida         3.39
I. mississippiensis   OF 23825   F    Oklahoma        3.49
I. plumbea            OF 38882   M    Captive         3.32
I. plumbea            OF 23826   F    Colombia        3.60

                         Length from
                         proximal end        Distal width
                      of Fossa metatarsi      excluding
                      I to distal end of    lateral flange
                           Trochlea          of Trochlea
   Species               metatarsi II        metatarsi II

Fossil                      11.55                7.03
I. mississippiensis          9.38                6.85
I. mississippiensis         10.13                7.02
I. mississippiensis         10.23                6.84
I. mississippiensis         10.68                7.15
I. mississippiensis         10.28                6.99
I. plumbea                  10.41                6.68
I. plumbea                  10.54                7.04
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Author:Steadman, David W.; Mead, Jim I.
Publication:The American Midland Naturalist
Article Type:Report
Geographic Code:1MEX
Date:Apr 1, 2010
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