It has been more than ten years since Becker (1987a:25) briefly reviewed the fossil birds of the "Agate Fossil Quarries." The specimens described herein were collected in 1908 by field crews from the University of Nebraska State Museum, but have never been identified or reported upon until now. This collection includes the first record of a crane, Gruiformes, and additional specimens of the fossil hawk, Buteo ales Wetmore. Buteo typhoius Wetmore should be removed from the list of species from Agate.
For historical accounts, the most comprehensive
geologic data and the best paleoenvironmental interpretation for
the Monument and surrounding fossil localities are discussed
in the works of Robert M. Hunt, Jr. (1972, 1978, 1981,
1985, 1990, and 1995) and Hunt and Skolnick (1996).
[An asterisk (*) indicates genera or species described from the Agate Fossil Beds. The classification sequence followed below is that of the AOU Check-list, 6th edition. The following acronyms are used: American Museum of Natural History, Department of Vertebrate Paleontology (AMNH); Carnegie Museum of Natural History (CMNH); Harold J. Cook collection (HC); Museum of Comparative Zoology (MCZ); Princeton University Geological Museum (PUGM) now at Yale University, Peabody Museum; University of Nebraska State Museum (UNSM)].
(hawks, eagles, falcons, and allies)
(kites, hawks, eagles, and allies)
(kites, hawks, and eagles)
Genus Promilio Wetmore 1958
Promilio efferus (Wetmore) 1923
Holotype.AMNH 6299, left tarsometatarsus missing internal half of the shaft; Agate Fossil Beds National Monument, Upper Harrison (late Arikareean), Sioux Co., Nebraska.
Promilio efferus is the earliest record for a kite in North America (Brodkorb, 1964:274). Wetmore (1923:504) had tentatively placed efferus in the genus Proictinia Shufeldt (1915: 301) from the late Miocene [latest Clarendonian or earliest Hemphillian, Long Island local fauna, Phillips County, Kansas; see Steadman (1981:171) for comments on age of Long Island local fauna]. Later, Wetmore (1958:2) decided that Proictinia was more closely related to the Everglade Kite, Rostrhamus sociabilis, and that P. efferus was more like the Old World carrion eating kites in the genus Milvus. Promilio efferus is equivalent in size and is similar osteologically to species of Milvus. Wetmore (1958:3) placed efferus in the subfamily Milvinae, but in a new genus, Promilio, based on perceived differences.
Genus Buteo Lacepede, 1799 (hawks)
Buteo typhoius Wetmore, 1923
Holotype.AMNH 1754, distal two-thirds of the right tarsometatarsus missing the trochlea for Digit II; from the Lower Snake Creek, Olcott Formation (early Barstovian), 23 miles south of Agate, Sioux Co., Nebraska.
Referred material.HC 477, distal one half of the right tibiotarsus, Agate Fossil Beds National Monument, Stenomylus Quarry 1.5 miles east of Carnegie Hill and University Hill quarries (referred by Wetmore, 1928:149).
Discussion.Buteo typhoius is osteologically very similar to its living congeners. Wetmore compared B. typhoius exclusively to the eastern Red-tailed Hawk, B. jamaicensis borealis, from which it can be distinguished by subtle osteological differences and its much larger size (approximately 50 percent; see Wetmore, 1923:485-492; 1928:149-150).
None the less, because of the disparity in age of the type locality of B. typhoius, which is the Lower Snake Creek, early Barstovian, (see Becker, 1987b for the interesting history and insight into the age of the locality) and the age of the Monument, late Arikareean, I do not think B. typhoius occurs at Agate. The tibiotarsus (HC 477) that Wetmore (1928:149) referred to B. typhoius most likely belongs to the next species B. ales (Wetmore). Therefore, B. typhoius should be stricken from the Monument species list.
*Buteo ales (Wetmore) 1926
Holotype.CMNH 1828, complete right tarsometatarsus; Agate Fossil Beds National Monument, Carnegie Hill (Quarry No. 2).
Referred material.UNSM 3001, hallux; UNSM 3002, right humeral shaft; UNSM 3004, left ulna missing the olecranon; UNSM 3006, right femur, proximal end; UNSM 5782, left ulna, distal one/quarter; UNSM 5783, right tarsometatarsus, distal end missing trochlea of Digit 4; UNSM 5784, left tibiotarsus, proximal two/thirds; UNSM 5785, right tarsometatarsus, distal one/quarter missing anterior half of trochlea of Digit 3; UNSM 5786, left humerus, distal end missing the entepicondyle.
Discussion.Buteo ales (Wetmore, 1926:403) was a large hawk. The length of the holotype tarsometatarsus, CMNH 1828, is 90.2mm with a distal width of 16.2mm. A hawk of this size is at the top of the size range for the eastern Red-tailed Hawk, B. jamaicensis borealis, given by Friedmann (1950:239). With the identification and referral of the UNSM fossils to B. ales, the assignment of this species to Buteo is affirmed. The ulna, UNSM 3003, may appear to be too long for this species (164.0mm), but is within reason when sexual size dimorphism is taken into consideration for North American raptors (Snyder and Wiley, 1976).
Measurements of selected fossils.hallux (UNSM 3001) length of cord = 24.0mm; ulna (UNSM 3004) length missing the olecranon = [164.0mm], proximal width = 15.2mm, distal width = 11.0mm, depth of external condyle = 11.3mm, mid-shaft width/depth = 7.1/7.2mm; ulna (UNSM 5782) distal width/depth of external condyle = 10.8/11.2mm; tibiotarsus (UNSM 5784) width/depth across proximal articular surface = 13.5/17.5mm, length of fibular crest = 28.5mm, width/depth of shaft below fibular crest = 7.9/6.9mm; tarsometatarsus (UNSM 5785) distal width = 17.1, depth of trochlea Digit 4 = 8.3mm, width of trochlea Digit 2 = 7.2mm; humerus (UNSM 5786) distal width missing entepicondyle = [21.0mm].
Wetmore identified three other fossils only to family rank with the following comments: HC 466, right ulna, proximal end about the size of a caracara, Polyborus (Wetmore, 1923:506); PUGM 12157, right tarsometatarsus, distal end which resembles a Marsh Hawk, Circus hudsonius (Wetmore, 1923:507); and CMNH 2207, large claw somewhat smaller than a Golden Eagle, Aquila chrysaetos (Wetmore, 1926:406). I have not seen any of these fossils, but the right ulna (HC 466) which is the size of a caracara should be re-examined with the UNSM ulnae referred to B. ales.
*Species Palaeastur atavus Wetmore 1943
Holotype.HC 693, right tarsometatarsus, distal one/third; Agate Fossil Beds National Monument, Stenomylus Quarry.
Discussion.Wetmore (1943:230) likened this new
genus and species of extinct hawk to the monotypic
white Hawk-eagle (Spizastur melanoleucus). Spizastur melanoleucus is the smallest of the booted eagles, which are considered by Brown and Amadon (1968:22) to be "the most highly evolved members of the family and indeed of all birds of prey." They live in dense, humid evergreen, and semideciduous forests in Central and South America (Blake, 1977; Howell and Webb, 1995).
(grouse, quail, turkeys, and allies)
(curassows, guans, and chachalacas)
*Species Boreortalis tantala (Wetmore) 1933
Holotype.HC 498, right tibiotarsus, distal end; Agate Fossil Beds National Monument, Carnegie Hill. (This specimen is now in the AMNH collections).
Discussion.In the Neogene there are five species of boreal chachalacas in the genus Boreortalis Brodkorb (1964:304-305). Boreortalis (Brodkorb, 1954:180) is closely related to Ortalis, the genus of living chachalacas found in southern Texas through Central America and into South America (Blake, 1977). It was Brodkorb's impression that chachalacas had a Nearctic origin and only during the Great American Biotic Interchange did they expand their range into South America. Although boreal today, the chachalacas from these North American fossil localities would have been much more tropical to subtropical in nature.
(grouse, quail, and turkeys)
*Species Palaealectoris incertus Wetmore 1930
Holotype.MCZ 2190, left humerus, proximal one/half and distal end with part of shaft missing; Agate Fossil Beds National Monument.
Discussion.This fossil grouse is between a Northern Bobwhite, Colinus virginianus, and the Spruce Grouse, Dendragapus canadensis, in size. Wetmore (1930:152-153) thought it similar to the Spruce Grouse, but only distantly related. Today, the Spruce Grouse is a member of the boreal species community. Palaealectoris may represent an ancestral grouse living in a subtropical environment before the dichotomy we see today between boreal species like the Spruce Grouse and the prairie grouse like the Greater Prairie-Chicken, Tympanuchus cupido.
(cranes, rails, limpkins, and their allies)
Genus and species indeterminate
Referred material.UNSM 3003, right ulna mid-shaft; UNSM 3005, right humerus, distal end; both are from Agate Fossil Beds National Monument, University Hill, University Quarry.
Discussion.This right ulna (UNSM 3003) and humerus (3005) are the first crane fossils to be identified from the Monument. Although fragmentary, the fossils are from a species of crane approximately the size of the modern Sandhill Crane, Grus canadensis (Linnaeus). It should be kept in mind, however, that in the late Clarendonian there is an Old World crane (Balearica) in Nebraska (Feduccia and Voorhies, 1992:241), and the fossils from Agate may represent either of these cranes.
(thick-knees, plovers, sandpipers, and their allies)
(coursers and pratincoles)
* Species Paractiornis perpusillus (Wetmore) 1930
Holotype.MCZ 2191, left tarsometatarsus missing the intercotylar prominence; Agate Fossil Beds National Monument, Carnegie Hill.
Discussion.Wetmore (1930:153) originally named
and described this new genus and species as the earliest
representative of the modern oystercatchers, Haematopodidae.
However, Olson and Steadman (1978:972-976) have shown
this species to be the first New World record for the Old
World pratincoles (Glareolinae). Pratincoles are short-legged
insect feeders, which feed on the wing much like swallows.
Once again, we have a member of the paleoavifauna from
Agate which is representative of a greater, worldwide avifauna
still in existence at the end of the Oligocene and into the
The fossil birds of the Agate Fossil Beds National
Monument are very interesting at several levels. First, they are
interesting at an alpha taxonomic level as new genera and
species. Secondly, birds are frequently used as
environmental indicators of certain habitats and to evaluate the health of
the environment. The birds from the Monument support
Hunt's interpretation of the paleoenvironment as an area with
ephemeral stream channels, open plains, but with riparian areas
along the streams. Thirdly, it is significant that the
paleoavifauna represents a worldwide fauna, which only recently has
been recognized at other important localities like Quercy,
France; Messel, Germany; Green River, Wyoming; and the Naze
in England. Olson (1989:2023-2029) has dubbed this the
global avifauna, of which our world today has only relictual
distribution patterns of that once cosmopolitan avifauna.
I would like to thank Robert M. Hunt, Jr. for giving
me the opportunity and encouragment to work on these very
interesting fossils. I am also indebted to Vincent L. Santucci
for allowing me the time and opportunity to submit this
contribution on the fossil birds of Agate. Much thanks goes out to
following curators, collection managers, and their institutions for helping me with loans of fossils and/or modern osteological specimens for this study: Robert M. Hunt, Jr. and R. George Corner, Nebraska State Museum, University of Nebraska; Larry D. Martin and M. Desui, Vertebrate Paleontology Department and the late Robert M. Mengel, the late Marion Jenkinson, Richard Prum, and Mark Robbins, Museum of Natural History, The University of Kansas; S. David Webb and Marc Frank, Division of Vertebrate Paleontology and David Steadman and Tom Webber, Division of Ornithology, Florida Museum of Natural History, The University of Florida. I would like to thank Linda D. Chandler, Dennis Parmley, William P. Wall, and the editors for critically reading one or more manuscripts and improving this paper.
Becker, J. J. 1987a. Neogene avian localities of North America. Smithsonian Institution Press, Washington, D.C.
. 1987b. Revision of "Falco" ramenta Wetmore and the Neogene evolution of the Falconidae. The Auk 104:270-276.
Blake, E. 1977. Manual of Neotropical Birds, vol. 1 (Spheniscidae (Penguins) to Laridae (Gulls and Allies). The University of Chicago Press, Chicago and London. 674pp.
Brown, L. and D. Amadon. 1968. Eagles, Hawks and Falcons of the World. New York, McGraw-Hill, 2 vols., 945pp.
Brodkorb, P. 1954. A chachalaca from the Miocene of Florida. Wilson Bulletin 66(3):180-183.
. 1964. Catalogue of fossil birds: Part 2 (Anseriformes through Galliformes). Bulletin of the Florida State Museum, Biological Sciences 8(3):195-335.
Feduccia, A. and M.R. Voorhies. 1992. Crowned Cranes (Gruidae: Balearica) in the Miocene of Nebraska, in Papers in Avian Paleontology (K.E. Campbell, Jr., ed.). Natural History Museum of Los Angeles County, Science Series No. 36:239-248.
Friedmann,H. 1950. The Birds of North and Middle America. Part 11. United States National Museum, Bulletin 50. 793pp.
Howell, S.N.G. and S. Webb. 1995. A guide to the birds of Mexico and Northern Central America. Oxford University Press. New York. 851pp.
Hunt, R.M., Jr. 1972. Miocene Amphicyonids (Mammalia, Carnivora) from the Agate Spring Quarries, Sioux County, Nebraska. American Museum Novitates No. 2506.
. 1978. Depositional setting of a Miocene mammal assemblage, Sioux County, Nebraska (U.S.A.). Palaeogeography, Palaeoclimatology, Palaeoecology, 24:1-52.
. 1981. Geology and Vertebrate Paleontology of the Agate Fossil National Monument and surrounding region, Sioux County, Nebraska (1972-1978). National Geographic Society Research Reports, v. 13:263-285.
. 1985. Faunal succession, lithofacies, and depositional environments in Arikaree rocks (Lower Miocene) of the Hartville Table, Nebraska and Wyoming, in Fossiliferous Cenozoic deposits of western South Dakota and northwestern Nebraska. Dakoterra v. 2(2):155-204.
. 1990. Taphonomy and sedimentology of Arikaree (Lower Miocene) fluvial, eolian, and lacustrine paleoenvironments, Nebraska and Wyoming; A paleobiota entombed in fine-grained volcaniclastic rock, pp. 69-111, in Lockley, M.G. and A. Rice, eds., Volcanism and fossil biotas: Boulder, Colorado, Geological Society of America, Special Paper 244.
. 1995. The Miocene carnivore dens of Agate Fossil Beds National Monument, Nebraska: oldest known denning behavior of large mammalian carnivores, pp.3-7, in V.L. Santucci and L. McClelland, eds. National Park Service Paleontological Research No.16.
, and R. Skolnick. 1996. The giant mustelid Megalictis from the early Miocene carnivore dens at Agate Fossil Beds National Monument, Nebraska: earliest evidence of dimorphism in New World Mustelidae (Carnivora, Mammalia). Contributions to Geology, University of Wyoming, v. 31(1):35-48.
Olson, S.L. 1989. Aspects of Global Avifaunal Dynamics during the Cenozoic. Acta 19 Congressus Internationalis Ornithologici, v. 2. Henri Ouellet, ed. "1988" Ottawa: University of Ottawa Press.
, and D.W. Steadman. 1978. The fossil record of the Glareolidae and Haematopodidae (Aves: Charadriiformes). Proceedings of the Biological Society of Washington 91(4):972-981.
Snyder, N.F.R. and J.W. Wiley. 1976. Sexual size dimorphism in hawks and owls of North America. Ornithological Monographs No. 20. American Ornithologists' Union. Washington, D.C.
Steadman, D.W. 1981. A re-examination of Palaeostruthus hatcheri (Shufeldt), a late Miocene sparrow from Kansas. Journal of Vertebrate Paleontology 1(2):171-173.
Wetmore, A. 1923. Avian fossils from the Miocene and Pliocene of Nebraska. Bulletin of The American Museum of Natural History 48(12):483-507.
. 1926. Description of a fossil hawk from the Miocene of Nebraska. Annals of the Carnegie Museum 16(3-4):403-408.
. 1928. The tibio-tarsus of the fossil hawk Buteo typhoius. Condor 30(2):149-150.
. 1930. Two fossil birds from the Miocene of Nebraska. Condor 32(3):152-154.
. 1933. A fossil gallinaceous bird from the Lower Miocene of Nebraska. Condor 35(2):64-65.
. 1943. Two more fossil hawks from the Miocene of Nebraska. Condor 45(6):229-231.
. 1958. Miscellaneous notes on fossil birds.
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