California condor associated with spruce-jack pine woodland in the late Pleistocene of New York

QUATERNARY

RESEARCH

28, 415-426 (1987)

California Condor Associated with Spruce-Jack Pine Woodland in the Late Pleistocene of New York DAVID W. STEADMANANDNORTON Biological

Survey,

New

York

State

Museum,

The State

Education

G.MILLER Department,

Albany,

New

York

12230

Received January 6, 1987 A humerus, coracoid, and pedal phalanx of the California Condor, Gymnogyps californianus, were recovered from the Hiscock Site in western New York, in an inorganic stratum containing wood that is 11,000 radiocarbon years old. Associated vertebrates include mastodont, wapiti, and caribou. Pollen and plant macrofossils from the sediments indicate a spruce-jack pine woodland and a local, herb-dominated wetland community. Historic records (all from western North America) and previous late Pleistocene fossils of the California Condor are associated mainly with warm-temperate climates and floras. The New York fossils show that this bird was able to live in a colder climate and in a boreal, coniferous setting at a time when appropriate food (large mammal carrion) was available. The California Condor, which survives only in captivity, has suffered a greater reduction in geographical range than previously suspected. Much of this reduction in range probably occurred ca. 11,000 yr B.P. when the extinction of many North American large mammals resulted in severely reduced availability of food for the California Condor and other large scavenging birds. Q 1987 University of Washington.

INTRODUCTION The late Quaternary record of birds in North America is less complete than that for mammals. This is particularly true north of the Wisconsinan glacial boundary where few fossil birds have been found (Lundelius et al., 1983). Recently collected fossils from western New York provide new information about the birds that reoccupied the newly deglaciated terrain. SITE DESCRIPTION AND STRATIGRAPHY The Hiscock Site, located 1.5 km WNW of the Village of Byron, Genesee County, New York (lat. 43”5’N, long. 78”5’W, elev. 187 m; Byron Quad.), is a rich accumulation of plant and animal remains in a seasonally inundated peatland of 0.8 ha (Fig. 1). A preliminary excavation was conducted in 1959 (Heubusch, 1959). In 1982, a long-term program of excavation was begun by the Buffalo Museum of Science. Four major stratigraphic units are present. These are presented in the column

labeled “SEDIMENTS” in Fig. 2. Unit A is a late Pleistocene inorganic deposit of boulders and cobbles in a matrix of pebbles, sand, silt, and clay. Unit B is a late Pleistocene, largely inorganic deposit of clays and silts with varying components of sands, pebbles, and boulders, with abundant small coniferous twigs and rarely other plant macrofossils. Unit C is an early Holocene peat (thickness varies from 0 to 30 cm) with abundant plant macrofossils (cones, seeds, fruits, and especially wood). Unit D is a late Holocene peat (thickness varies from 0 to 70 cm) with abundant plant macrofossils (wood, cones, seeds, fruits). Spring action is indicated in Units B and C by the presence of scattered sandy pockets left from localized, intermittent vertical movement of water. Details of the excavation methods, stratigraphy, and geomorphology of the Hiscock Site have been reported elsewhere (Laub et al., in press; Miller, in press; Muller and Calkin, in press; Steadman et al., 1986; Steadman and Miller, in press). Preservation of pollen is good

415 0033-5894/87 $3.00 Copyright 0 1987 by the University of Washington. All tights of reproduction in any form reserved.

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STEADMAN

throughout the sequence (Miller, in press; see below). Forty-eight species of amphibians, reptiles, birds, and mammals have been recovered from the Holocene peats (Units C, D), whereas only seven species of vertebrates have been recorded from Units A and B (Steadman, in press). Bones of the extinct mastodont Mummut americanum are confined to and dominate Units A and B (Laub et al., in press). Radiocarbon dates range from 250 1 50 to 680 5 80 yr BP for Unit D, From 8610 +- 80 to 8720 + 80 yr B.P. for Unit C, and from 10,930 ? 90 to 11,250 ? 140 yr B.P. for Unit B (Table 1). A radiocarbon date is also available, presumably for Unit B, of 10,450 r 400 yr B.P. on wood (W-1038) from the preliminary excavation in 1959 (Ives et al., 1964). THE CONDOR FOSSILS Three fossil bones (Figs. 3-5) of the California Condor, Gymnogyps californianus, were recovered within an area of 4.8 m2 from Unit B: a humerus (Buffalo Museum of Science catalog No. E25654; collected in 1984 from square HSSW, depth 114 cm), a coracoid (BMS E25655; collected in 1985 from square HSSW, depth 124 cm), and a terminal pedal phalanx (BMS E25921; collected in 1986 from square GSNE, depth 79 cm). The lesser depth of the pedal phalanx is not indicative of a younger age, for the upper limit of Unit B varies greatly in depth from square to square. Compared with the equivalent bones of modern individuals (Table 2), the humerus and phalanx are relatively smaller and the coracoid is within the size range, suggesting that the three fossils may not belong to the same individual. The condor fossils are grayish brown and mineralized. In contrast the numerous bird

AND MILLER

fossils from the Holocene peat are brown to black (stained with humic acids) and little mineralized. The age of the condor fossils is placed at ca. 11,000 yr B.P. by a radiocarbon date on coniferous twigs of 11,250 + 140 yr BP. from square HSSW at a depth of 115-125 cm. A uniform age of ca. 11,000 yr B.P. for the sediments containing the condor fossils is indicated by a radiocarbon date of 10,930 ? 90 yr B.P. at a depth of 14.5-155 cm, which is 30 cm below the condor fossils (Table 1). At 1.5 standard deviations, this date overlaps with that from the same depth as the condor humerus and coracoid. The fossils are referred to Gymnogyps on the basis of size (Table 2) and qualitative characters determined by comparisons with specimens in the skeletal collection of the Division of Birds, National Museum of Natural History, Smithsonian Institution. Within the family Cathartidae, the fossil specimens are much larger than those in Coragyps, Cathartes, or Sarcoramphus. The coracoid differs from that in Vultur gryphus (the living Andean Condor) in its slightly smaller size and in having a narrower, more ovoid glenoid facet. The humerus differs from that in Vultur gryphus in its slightly smaller size and in having a deeper olecranal fossa, a deeper internal side of the brachial depression, and an attachment of the anterior articular ligament that is shorter but more distinctly elevated from the shaft. These fossils agree qualitatively with Gymnogyps rather than with the similarly sized, extinct Breagyps clarki, in the characters stated by Howard (1972). The terminal pedal phalanx agrees qualitatively with that of Gymnogyps. It seems to be slightly smaller than in modern specimens, although abrasion during sedimentation has rounded much of the surface of

FIG. 1. Hiscock Site basin contours (interval 0.2 m) at peat/inorganic sediment interface as determined by probing with steel rods. Inset at left shows grids of excavated squares. The solid circle (HSSW) indicates the position where the pollen samples were taken.

LATE

PLEISTOCENE

CONDOR

417

418

STEADMAN

3

AND MILLER

‘54

t

PERCENT

OF POLLEN

BROWN, HUMIFIED PEAT WITH SOME WOOD

SUM

l

analyst:

CONES

N.

G. Miller,

HS 2

1985-86

WOODY PEAT

FIG. 2. Pollen diagram showing selected pollen taxa at Hiscock Site, Genesee Co., New York. Further explanation in Appendix A.

this fossil, thus precluding surements.

accurate mea-

PALYNOLOGY AND PLANT MACROFOSSILS In order to characterize the vegetation that existed during deposition of the sediments at the Hiscock Site, pollen was extracted from a stratigraphic series of samples (Fig. 2; Appendix A). Unit B, which contains the condor fossils, has nine TABLE Lab number B-15963 B-16054 B-16733 B-16735 B-16734 B-15962 B-16736 B-16175

Identity of wood Quercus

Fraxinus Fruxinus? Picea or Larix Picea or Larix Picea

Conifer twigs Conifer twigs

similar pollen spectra (pollen zone HS 1) with Gramineae, Cyperaceae, and longspine Compositae types dominant (85%) and low values for spruce and pine (10 and 5%, respectively). Pollen assemblages above HS 1 contain high percentages of pollen from trees (HS 2, HS 3) or from herbs associated with agriculture (HS 4) and conform quantitatively with assemblages known at other sites in western New York (Calkin and

1. RADIOCARBONDATESFROMTHEHISCOCK

SITE

Depth 4% 250 530 680 8,660 8,720 8,610 11,250 10,930

* 2 ? k 2 f 2 2

50 50 80 100 80 80 140 90

Square

(cm)

HSSW HSSW GSNW HSSW HSSW HSSW HSSW HSSW

30 48 60 73 85 90 115-125 145-155

Stratigraphic unit D D D C C C B B

Peat Peat Peat Woody Woody Woody Sandy, Sandy,

peat peat peat silty clay silty clay

Nore. Only those from the vicinity of the Condor humerus and coracoid (squares GSNW and HSSW) are given. All determinations are on wood samples and are reported in radiocarbon years BP corrected for 13Ci12C.

LATE PLEISTOCENE

A

6

CONDOR

D

FIG. 3. California Condor humerus. Fossil (BMS E25654) from the Hiscock Site, New York in cranial (A) and caudal (C) aspects, compared with the same element from a modern skeleton from California (B, D; USNM 3369). Scale = 5 cm.

McAndrews, 1980; Miller, 1973a; Spear and Miller, 1976). Zone HS 2 is marked by high total values of Pinus (to 40%) and Quercus (15%), while HS 3 shows decreased percentages of Pinus and increased representation of Fugus, Acer saccharum, and Carya. Zone HS 4 has high perand Gramineae centages of Ambrosia-type and reduced values for various deciduous trees and Pinus. The radiocarbon dates establish that the

age of sediments between ca. 90 cm and the surface is Holocene, whereas that of Units A and B is largely or totally late Pleistocene. The Holocene record is incomplete, with a hiatus of about 8000 yr between 73 and 60 cm indicated by the radiocarbon age of wood samples at these depths (Table 1). A second, older hiatus is suggested by the abrupt increase in pine pollen percentages between 89 and 79 cm, a change without parallel in other pollen profiles from the re-

420

STEADMAN

AND MILLER

FIG. 4. California Condor coracoid. Fossil (BMS E25655) from the Hiscock Site, New York in dorsal (A) and ventral (C) aspects, compared with the same element from a modern skeleton from California (B, D; USNM 3369). Scale = 5 cm.

gion. This interval marks the onset of peat accumulation at the Hiscock Site. As these events are younger than those recorded in Unit B, they will not be discussed further. The nonarboreal to arboreal pollen ratio (NAP:AP) of HS 1 is 9: 1. This is anomalously high for late glacial sediments in the eastern Great Lakes region, where the oldest late glacial pollen assemblages, which are generally interpreted as indicating tundra or a tundra-forest mosaic, have an NAP:AP ratio of 1: 1. Younger late

glacial assemblages have higher percentages of spruce pollen and an NAP:AP ratio of from 1:3 to 1:9 (Anderson, 1982; Calkin and McAndrews, 1980; Davis, 1983; McAndrews, 1981; Miller, 1973a, b; Spear and Miller, 1976; Terasmae and Matthews, 1980; Watts, 1983). Although Cyperaceae values from 18 to 52% (x of HS 1 spectra = 27%) are not unusual in late glacial sediments of small lakes and ponds in glaciated northeastern North America, much lower percentages of Gramineae and long-spine

I 5 cm

[0

A

B

FIG. 5. California Condor terminal pedal phalanx (digit 2, phalanx 3). Fossil (BMS E25921) from the Hiscock Site, New York in dorsal (A) and lateral (C) aspects, compared with the same element from a modern skeleton from California (B, D; USNM 3369). Scale = 5 cm.

LATE PLEISTOCENE

Compositae pollen (i.e., subf. Asteroideae, excluding Ambrosia and Artemisia) are generally the rule. Zone HS 1 also contains numerous subordinate pollen types of herbs (e.g., Campanula, cf. Cornus, Epilobium, Galium, Potentilla culus, Caryophyllaceae,

palustris,

Ranun-

Cichorioideae, Cruciferae, Labiatae, other Rosaceae, Scrophulariaceae, Urticaceae), some of which are recorded infrequently in analyses of late glacial sediments. The HS 1 pollen assemblage indicates boreal conditions at the time the condor fossils were deposited. Spruce and pine were the principal trees near the basin, and the occurrence of jack pine (Pinus banksiana Lam.) cones at depths of 96, 99, and 118 cm indicates the local presence of this boreal tree. The Hiscock site is ca. 300 km south of the continuous range of the species (Critchfield and Little, 1966, map 56). Cones of white spruce (Picea glauca (Moench) Voss) from sediments of pollen zone HS 2 suggest that this tree may have been associated with jack pine, although no cones of white spruce have been found yet in HS 1 sediments. The density of trees in the vegetation reTABLE

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421

corded in HS 1 is difficult to infer, and therefore whether tundra-forest or forest is represented cannot be decided with certainty. One reason for this is the high probability of overrepresentation by herbs growing in or near the basin. Three lines of evidence support a local edaphic cause for the unusually high NAP:AP ratios: (1) Pollen spectra from contemporaneous late glacial sediments at sites with 90 km of the Hiscock basin have NAP:AP ratios from 1:I to 1:6 (Calkin and McAndrews, 1980; Miller, 1973a, b; Spear and Miller, 1976), showing that high percentages of tree pollen types were a regional phenomenon at 11,000 yr B.P. (2) Nearly 25% of the pollen sum is of insect-pollinated Compositae. This suggests that conditions near or in the basin were unusually appropriate for the growth of these and ecologically similar herbs. (3) There is no sedimentary evidence that the basin has held a lake, although small, shallow, intermittent pools of water probably were present. A water table lower than that now existing at the site and an associated reduction in basin area are indicated by the latest Holocene age of the upper 60 cm of peat and the hiatus in depo-

2. MEASUREMENTS (IN MM) OF THE HUMERUS AND CORACOID OF GYMNOGYPS

Humerus:

Distal width

Least width of shaft

Hiscock fossil (BMS E25654) Modern no. of specimens

45.3 48.1-51.2 7

14.6 14.6-16.4

CALIFORNIANUS

Diagonal depth of external condyle

Diagonal length of brachial depression

25.5 26.9-29.4 6

23.6 25.4-31.8 7

Coracoid:

Total length

Length of glenoid facet

Width of glenoid facet

Depth of triosseal region

Hiscock fossil (BMS E25655) Modem no. of specimens

98.5 94.9- 102.5 9

28.7 28.2-30.9 9

12.3 13.0 I

19.6 19.9 1

Length of coraco-humeral surface

17.3 16.0- 18.9 9

Note. Range and sample size are given for the modern comparative specimens (Division of Birds, Smithsonian Institution; USNM 3369, 13823, 345225, 346582, 489406, 489755, 492447).

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STEADMAN

sition from about 8720 to 680 yr B.P. Since excavations have revealed much variation in relief at the bottom of the basin (up to 50 cm of depth in 2.5 m linear distance), a lowered water table would create more area for the establishment of wetland or mesic herbs. Sediments in basins smaller than 1 ha may be dominated by locally produced pollen (Jacobson and Bradshaw, 1981). The absence or low percentages (generally ~1%) of pollen of trees typical of deciduous or hemlock-northern hardwood forest (e.g., Acer, Betula, Carya, Tilia, Tsuga), together with the pollen of Picea, pollen and macrofossils of Pinus banksiana, and high values of various Cyperaceae, Gramineae, and Compositae, indicates the existence of open, local wetland plant communities with spruce and jack pine on adjacent slopes at the time the condor fossils were deposited. Such upland forest associations conform in floristic composition to those of the modern boreal forest region of eastern Canada. DISCUSSION

AND CONCLUSIONS

The fossils from New York represent a major extension of the late Pleistocene range of the California Condor (Fig. 6). East of the Mississippi River, condors are known only from four early and eight late Pleistocene sites in Florida (Lundelius et al., 1983; S. D. Emslie, personal communication), the closest of which is 1600 km south of the Hiscock Site. Other late Pleis-

AND MILLER

tocene records of Gymnogyps are from Nuevo Leon and Texas to California (Fig. 6). (The record of Gymnogyps sp. from Smith Creek Cave, Nevada (Howard, 1952), has been reevaluated as Breagyps clarki (H. Howard, personal communication to S. D. Emslie).) Although the Hiscock fossils do not demonstrate that condors nested in the immediate vicinity of the site, they do imply the regional presence of a resident population. Condors are not known to stray more than 300 km north of their breeding range (Wilbur, 1973). Historic records of the California Condor are confined to the West Coast, from Baja California to British Columbia (Koford, 1953). Today, the condor survives only in captivity. The decline of the California Condor within the past century has been discussed extensively (Koford, 1953; Wilbur, 1978, 1980; Kiff et al., 1979; Rea, 1981; Snyder, 1983; U.S.F.W.S., 1984; Crawford, 3985). Some causes of the condor’s recent decline, such as poisoning from pesticides and lead, are unique to modern times. The stratigraphic and paleobotanical evidence from the Hiscock Site indicates that California Condors lived in the northeastern United States 11,000 radiocarbon years ago under boreal conditions, only 300-400 km south of the continental ice sheets. This habitat contrasts greatly, for example, with that of the late Pleistocene condor fossils from Vulture Cave, Grand

FIG. 6. The late Pleistocene range of the California Condor. (A) Hiscock Site, Genesee Co., New York; (B) Itchetucknee River, Columbia Co., Florida: (C) Seminole Field, Pinellas Co., Florida; (D) Hog Creek, Sarasota Co., Florida; (E) Friesenhahn Cave. Bexar Co., Texas; (F) Gypsum Cave, Clark County, Nevada; (G) Vulture Cave, Mohave Co., Arizona; (H) Stanton’s Cave, Coconino Co., Arizona; (I) Howell’s Ridge Cave, Grant Co., New Mexico; (J) Conkling Cave, Dona Ana Co., New Mexico; (K) Burnet Cave, Eddy Co., New Mexico: (L) Dry Cave, Eddy Co., New Mexico: (M) Dark Canyon Cave, Eddy Co., New Mexico; (N) Rocky Arroyo Cave, Eddy Co., New Mexico: (0) Mule’s Ear Peak Cave, Brewster Co., Texas; (P) San Josecito Cave, Nuevo Leon, Mexico; (Q) Schuiling Cave, San Bernardino Co., California; (R) Ranch0 La Brea, Los Angeles Co., California; (S) Samwel Cave, Shasta Co., California; (T) Potter Creek Cave, Shasta Co., California; (U) Stone Man Cave, Shasta Co., California. Derived from data in Lundelius et al. (1983). Wetmore and Friedmann (1933). and Miller (1943). Other unpublished late Pleistocene records have been compiled by S. D. Emslie. Except for one record from Chihuahua, Mexico, all of Emslie’s new records are from Florida, Texas, New Mexico, Arizona, or California, and thus do not alter the range indicated by the records depicted here.

423

424

STEADMAN

Canyon, Arizona (lat. 35”40’N, elev. 645 m), which, like most or all other late Pleistocene condor records from the Southwest, are associated with an open juniper woodland (Mead and Phillips, 1981). The previous restriction of late Pleistocene fossils of condors mainly to warm-temperate areas (Fig. 6) may be an artifact of the incompleteness of the vertebrate fossil record in the region of continental glaciation. The record from the Hiscock Site suggests much broader climatic and vegetational tolerances for the California Condor in the late Pleistocene. The vertebrate fauna from the stratigraphic unit at the Hiscock Site that contains the condor fossils (Unit B) is dominated by the extinct mastodont, Mummut americanurn, represented by at least four individuals recovered thus far (Laub et al., in press). Other large mammals from Unit B are the wapiti (Cervus elaphus) and caribou (Rangifer sp.). While proboscideans (Mammut and Mammuthus) have been recorded from seven other late Pleistocene sites that have yielded condor fossils (Lundelius et al., 1983), the Hiscock Site is the first condor-bearing deposit with fossils of caribou and only the second such site with fossils of wapiti. Most late Pleistocene vertebrate localities north of the Wisconsinan glacial boundary consist of single individuals of mastodonts, mammoths, or other large mammals. The rich concentration of large animals at the Hiscock Site, as well as the presence of spring-derived sediments, suggests that the basin served as a watering place or wallow 11,000 yr ago. This may explain why scavenging animals such as condors were recovered from the Hiscock Site but have not been found at other sites in the Northeast. After 11,000 yr B.P., the onset of the Holocene in western New York was characterized by a warmer climate, more diverse forest types, and the development of deep gorges along the Niagara and Genesee rivers. Each of these changes would seem

AND

MILLER

to be either beneficial or neutral, rather than detrimental, for the California Condor. For example, modern condors nest either on ledges or in cavities of large trees (Snyder et al., 1986), both of which would have been more accessible in western New York in the Holocene than in the late Pleistocene. The age of the condor fossils from New York coincides with the time of megafauna1 extinction in North America (ca. 11,000 yr B.P.; Meltzer and Mead, 1983; Mead et al., 1986) and with the time that the California Condor suffered a drastic range retraction (Emslie, 1985, 1987). Based upon the wide climatic and vegetational tolerance of the California Condor, we agree with Emslie (1985, 1987) that the major reason for the large loss of range was a shortage of food (large mammal carrion) caused by the extinction of many large mammals. We believe that the same process that killed the large mammals, whatever that process was (Martin and Klein, 1984), probably affected the California Condor and other scavenging birds not directly, but indirectly by depleting their sources of food, in the form of dead large mammals (Steadman and Martin, 1984). Should condor fossils be found in the Northeast that postdate 11,000 yr BP, a cause other than megafaunal extinction should be sought for the loss of the condor from this region. Whereas the California Condor has survived into historic times, many other large North American scavenging birds (e.g., Teratornis, Breagyps, Cathartornis, Neogyps, and Neophrontops) apparently died out at the end of the Pleistocene. The feeding ecology of this diverse assemblage of scavenging birds may have resembled that of historic times in Africa, where up to seven species of large scavenging birds can occur in the same region, with as many as five of these feeding upon the same large mammal carcass (Steadman and Martin, 1984). Considering the great loss of large mammals and scavenging birds in North

LATE PLEISTOCENE

America at the end of the Pleistocene, perhaps we are fortunate that the California Condor survived into the Holocene. APPENDIX A Samples for pollen analysis were cut from a clean wall in square HSSW on 16 August 1985. Subsamples of 1 or 2 cm3 were dispersed in ethyl alcohol, sieved through a 180~p,rn mesh screen, and treated successively with 2N HCl, 35% HF (heated), 2N HCl, and 10% KOH. Each treatment was separated by distilled water washes. All residues were acetolyzed. Residues were mounted in 2000 cST silicone fluid. Counts were made at 500 x magnification. Pollen percentages for all taxa are based on the sum of trees and upland shrubs and herbs. Wetland and aquatic taxa are excluded. The percentages are given with 95% contidence intervals. Pollen assemblage zones are specified using the ZONATION computer program (Gordon and Birks, 1972) and a modified version of SLOTSEQ (Gordon and Birks, 1974; Futyma and Miller, 1986). ACKNOWLEDGMENTS For generous support and cooperation in the field and lab, we thank Richard S. Laub and other staff members of the Buffalo Museum of Science. We thank Charles and the late Charlotte Hiscock for access to the site and other sorts of cooperation. The hard work of the field crews is appreciated as well. Partial tinancial support was provided by the George G. and Elizabeth G. Smith Foundation, Inc., of Buffalo and the National Geographic Society (Grant 3359-86). Storrs L. Olson and J. Phillip Angle allowed access to skeletons in the Smithsonian Institution. For comments on the manuscript, we thank Steven D. Emslie, Richard P. Futyma, Richard S. Laub, Daniel A. Livingstone, and Paul S. Martin. We especially thank Steven D. Emslie for access to much unpublished information, and Donald M. Lewis for assistance in the laboratory. Dominique Pahlavan drafted Fig. 1 and assisted with other figures. The photographs are by Thaddeus Beblowski and Christopher Supkis. This is Contribution Number 515 of the New York State Science Service.

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