Bickart, 1984

A Field Experiment in Avian Taphonomy Author(s): K. Jeffrey Bickart Source: Journal of Vertebrate Paleontology, Vol. 4, No. 4 (Dec.
, 1984), pp. 525-535 Published by: Taylor & Francis, Ltd. on behalf of The Society of Vertebrate Paleontology Stable URL: http://www.jstor.org/stable/4523013 . Accessed: 18/09/2013 00:22
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they rested on bare soil, leaf litter, live plants, or twigs, although as disarticulation reached an advanced stage only bones resting on bare soil continued to be glued to the ground. Subsequent floods, which on several occasions washed over the floodplain and submerged the cages, failed in most cases to change the positions of the bones by more than several centimeters. Only the most severe storm of the season had a significant effect, moving some of the bones of three of the specimens to one edge of their cages, but leaving other bones of the same birds, and all bones of the other three, little moved from their original positions (Fig. 4). Because the mesh size of the cages was large, and because deposited mud was found on carcasses after floods, it is reasonable to assume that the cages did not significantly impede water flow around the carcasses. Other Movements of Carcasses and Bones Figure 4 shows the positions of the bones of four of the caged rock doves after five months. Many of the bones moved outward from the original carcass location and changed orientation. Some of these changes in position and orientation occurred with the most severe floods during the wet early summer. Bones not resting on bare soil, and thus not stuck down, were more susceptible to displacement than bones stuck to the ground; plant growth in four of the cages probably moved some bones, but not significantly. After completion of disarticulation, there was little change in bone orientation and distance from the original location of the carcass. During the winter of 198 182, long periods of coverage by snow and ice alternating with briefer periods of partial or complete thaw also had little effect on bone position. Bones were frozen into the positions they had attained prior to the snows. Substantial movement occurred only with major thaws, when the concomitant floods moved some of the bones nearer to the cage edges. Near the end of the one-year observation period, one of the cages of the five adult rock doves was destroyed and the contents disrupted, probably by humans. Some of the bones of two of the other four adult specimens had moved into contact with the cage edges since my examination at the beginning of March, and undoubtedly would have gone further. Nevertheless, most of the bones of these specimens remained in the centers of the cages. The positions of the bones of the other two specimens were as before, and not in contact with the cage edges. Approximately half of the bones of two of these four specimens became partially buried in the soil and also fixed in place by vegetation. One gull, #15, was carried by floodwaters from the bank into the stream, about 1.6 m downstream and 1 m out from the bank. The gull was exposed in the stream for several days. Sediment carried by stream waters after several storms then buried the gull quickly. The carcass was completely articulated and well covered by feathers when it was buried, with the exception
of the right tibiotarsus, tarsometatarsus, and phalanges, which separated from the rest of the carcass and were buried a short distance away. As the water level of the stream dropped during the dry part of the summer, the mud covering the carcass became exposed. I excavated the carcass 3.5 months after burial. I noted the following changes which occurred after burial (see Fig. 5): (1) almost all feathers and all soft tissue disappeared; (2) the left carpometacarpus and the first phalanx of the second digit moved about 10 cm from the distal ends of the left radius and ulna; (3) the phalanx subsequently disarticulated from the carpometacarpus; (4) both humeri disarticulated from the coracoids and scapulae, and those bones from the sternum; (5) the ribs and the cervical vertebrae disarticulated and became separated or transported away; (6) the plunge of the left tarsometatarsus changed from approximately zero to 900; (7) the left femur disarticulated from the pelvis, the plunge became vertical, and it rotated almost 1800 around its vertical axis; (8) a scapula also plunged at 900, and the right humerus, radius, and ulna plunged at about 300. I observed on the mud over the carcass racoon and dog tracks which were as deep as the top level of the bones, and numerous living plant roots in the mud surrounding the bones. Thus the changes noted above could have been caused by trampling or growth of plants; the former has been suggested as a burial process (Behrensmeyer and Dechant-Boaz, 1980; Laporte and Behrensmeyer, 1980). Decay and Disarticulation The timing of decay and disarticulation showed great individual variation. For six individuals time to complete disarticulation ranged from 13 days for the one juvenile rock dove to about six months for one of the adults. The times for the four adult birds between the extremes ranged from 65 to 100 days (Table 2). The disarticulation of the skeletal elements also showed individual variation, within an overall distinct sequence (Table 2, Fig. 6). Figure 6 indicates: (1) early (i.e., with respect to the 75 days over which most of the disarticulation occurred) disarticulation of ribs from the sternum; (2) early disarticulation of hind limb joints; (3) late disarticulation of vertebrae; (4) late disarticulation of the pectoral girdle; (5) a tendency for proximal wing joints to disarticulate before distal wing joints; (6) completion of leg disarticulation before wing disarticulation. For each of the joints or groups of joints shown in Figure 6, I calculated the time at which 75% of the joints in the sample had disarticulated. These are: vertebrae 34 days; sternum-ribs 25 days; stemrnumcoracoid 35 days; coracoid-scapula 55 days; coracoid, scapula-humerus 20 days; humerus, radius, ulna 45 days; manus 65 days; hind limb 45 days. These figures may be useful in estimating length of time before burial of a carcass (but see discussion of the stream-buried gull, below). Observation of the sequence of disarticulation was difficult because feathers obscured the bones
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JVP 4(4), December1984
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FIGURE 4. Drawings of four of the five caged adult rock doves, showing disarticulation and movement of bones outward from the original locations of carcasses (dotted outlines) in centers of cages. Solid lines next to bones indicate contact of bone and cage edge. Dotted lines cutting across bones indicate burial of undrawn part of bone. Drawing made after five months. a, #1. b, #6 -carcass originally in location at left; severe flood moved carcass, with exception of stuck skull, to location at right, from which bones then scattered outward. c, #5. d, #2.
JVP 4(4), December 1984
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TABLE 2. Times of disarticulation of skeletal joints of six caged rock doves. Specimens indicated by numbers 1-6. Where the same number is noted for a joint twice, it signifies a different disarticulation time for the two sides; where only one number is noted for a bilaterally symmetrical joint, it signifies the same disarticulation time for the two sides. Note that those joints of #3 not on diagram by 110 days were still articulated at that time. The times of disarticulation not on the table for other specimens were not noted. Time in days 0- 65 10 1. Skull-lower mandible 2. Skull-atlas 3. Cervical vert-cerv vert 4. Cerv vert-thoracic vert 5. Thoracic vert-pelvis 6. Pelvis-caudal vert 7. Thoracic vert-ribs 8. Synsacrum-ilia and ischia 9. Sternum-ribs 10. Sternum-coracoid 11. Coracoid-scapula 12. Coracoid and scapula-humerus 13. Humerus-radius and ulna 14. Radius proximalulna proximal 15. Radius distalulna distal 16. Radius and ulna-carpometacarpus 17. Carpometacarpusdigit 2 18. Digit 2, phalanx 1-digit 2, phalanx 2 19. Pelvis-femur 20. Femur-tibiotarsus 21. Tibiotarsus-tarsometatarsus 22. Tarsometatarsusphalanges 2 1115 1620 2125 6 3 6 6 2630 3135 1 1 1 1 1 1, 5 1 1 6 2, 6 2, 6 6 1 1 3, 5 3, 5 2, 5 5 3 3 2 1 2 2 2 6 2, 5 5 2, 6 1 1, 6 2 1 2 6 1 1 2 2 5 6 6 2, 5 2 6 3 3 3 2 3 3 2 6 1 1 6 6 2, 5 2 3640 5 5 5 2 2 2 6 6 2, 6 41- 4645 50 5155 6 56- 61- 66- 81- 9660 65 70 85 101 106110
I
2, 4 2 4, 5 4 2, 4, 6 4 4 4 4 4 4 4 4 4 4 1, 2 6 5 6 1, 5 3 6
and made the exact time of disarticulation of a joint uncertain. Thus the patterns shown in Figure 6 should be considered only probable patterns. Damage to Bones by Carnivores I recovered from the site a small number of bones damaged by racoons and foxes. They were similar in appearance to mammal bones damaged by carnivores (Haynes, 1980; Gifford, 1981; Shipman, 1981). The scraps all show green fractures typical of fresh bone
4--
(D. C. Fisher, pers. comm.) (Fig. 7). The fracture surfaces consist of smooth edges interrupted by jags. Fractures of spongy bone (Fig. 7, middle) look on a gross level rougher or "fuzzier" than fractures of compact bone (Fig. 7, top, bottom), but when magnified (Fig. 8, middle) the green fracture pattern is evident. Tooth punctures are present on two specimens (Fig. 7, top; Fig. 8, top). Of the 19 long bones recovered, 16, or 84%, were damaged by removal of one or both (one specimen) of the articular ends. This has been observed
FIGURE 5. Drawings of gull (#15) washed into stream and buried by sediment. Drawings made following excavation 3.5 months after burial. Top: plan view. Bottom: schematic projection of bone positions onto plane perpendicular to land surface; some bones omitted for clarity.
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might also affect the results. I found remains of one out of the six uncaged rock doves, but remains of four out of 16, or 1/4,of the ring-billed and herring gulls, a higher proportion of the larger birds. This suggests that scavengers were more likely to completely carry away from the site or completely consume at the site the smaller birds, thus leaving at the site fewer remains of them. Hill (1979) concluded from both theoretical and empirical considerations that there is an effective limit to the scattering of bones by random processes (e.g., disarticulation, kicking by passing animals) and that these form "the prevailing process upon which the effects of more specific processes my be imposed" (p. 272). One specific process is disturbance by scavengers. I observed an apparent limit to the scattering of bird bones in the absence of vertebrate scavengers; this may, especially when combined with observations of bone damage, facilitate recognition of degree of scavenging and predation. Knowledge of disarticulation sequences may also do this. For example, if a fossil, semi-articulated skeleton is found to be in a state of disarticulation not consistent with modern observations of undisturbed carcasses (e.g., wings disarticulated, legs not), disturbance by scavengers should be considered as one possible explanation. Just as ".... disarticulation pattern ... may illustrate those features that are unique to various hu-
FIGURE 8. Enlargementof bones shown in Figure 7, in same order. Scale bar equals 5 mm.
ably applicable to areas with: (a) enough rainfall to keep the floodplain well vegetated and wet enough for potential scavengers to have their dens elsewhere; (b) large numbers of scavengers; and (c) low sedimentation rates. Changes in any of these variables might lead to different results. If gluing is dependent not only on fluids from the decaying carcass but also on a wet substrate, it might not occur, for example, on the loose sand of a beach or in an area with low precipitation. Gluing does not last if the bones are not in contact with the soil, and so may be seasonally dependent, not happening in autumn, for example, when a dead bird's bones would rest on a bed of fallen leaves. Carcass size
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FIGURE 9. Tarsometatarsus of a juvenile rock dove, showing damage that occurredon articularends of most of the long bones; cause uncertain.Note holes in middle and bottom trochleae.Scale bars equal 5 mm. 533
man butchery patterns" (Hill, 1980:134), it may illustrate those features unique to non-human scavenging patterns. From the changes in articulation and positions of the bones of the buried gull it is clear, however, that such characteristics of a skeleton do not necessarily become permanent after burial. These observations contrast with Hill's (1979:261) statement that disarticulation "provides an estimate of the interval between death and burial," and with Toots' (1965:38) statement that "Partial burial has the effect of fixing the buried part in the position and state of articulation they had at the time of burial ... ." Disarticulation should, then, be used only cautiously as an indication of duration of subaerial exposure and condition of the bones when buried. The characteristics of bones damaged by predators and scavengers have received much attention from workers on mammals (e.g., Brain, 1980; Binford, 198 1; Gifford, 1981). Ability to distinguish damage by predators and scavengers from damage by other processes may enable one to make inference about presence or absence of predators and scavengers, and intensity of predation and scavenging, among other things. The observations presented in this study are, however, based on only a few specimens, and further work should be done. If weathering can be recognized on fossil bones, one may be able to estimate length of time of subaerial exposure of bones. I observed no weathering of the bones of the adult rock doves after one year. In other environments, however, bird bone may break down very quickly. C. G. Spies (pers. comm.) has observed carcasses reduced from fresh to chalky, crumbly bones within three weeks. Those carcasses were found on islands in Oneida Lake, New York, among vegetation four to five feet high, closely packed, with the water table a few inches below the ground surface and humidity near the ground probably near 100%. Clearly there is a need for much more work, including comparative work with birds and mammals in the same environment. The results of such work would have implications for the contention that bird fossils are rarer than mammal fossils because bird bones are more delicate than mammal bones. An actual example from the avian fossil record may illuminate how modern taphonomic observations may help solve problems in avian paleoecology. The lacustrine deposits of the Big Sandy Formation of Arizona, mentioned in the Introduction above and under study by me, contain some of the richest known concentrations of fossil birds. Most of the fossils are of ducks, geese, and swans; other water-associated types such as storks, rails, flamingoes, and shorebirds are present in lesser numbers, and there are a very few land birds, primarily diurnal raptors. Articulated skeletons or parts of skeletons are common, and the material is on the whole densely packed. The mammals known from the same deposits as the birds include carnivores of at least
five families; they include a fox, Vulpes stenognathus, and other canids, and a procyonid, Bassariscus sp. Among the important questions are: How does this fossil assemblage reflect the species composition and relative abundances of species in the original community? Were ducks really the most common group? How does the species diversity at this site in the early Pliocene compare with species diversity in lacustrine habitats today? Knowledge of how modern carnivores can by scavenging change the composition of an assemblage of carcasses on a land surface will help to reconstruct the original bird community. Observations of degree of disarticulation, and of any damage to the bones from the carnivores will also help, as discussed above. The practical problem arises in this densely packed assemblage of determining what bones go to what individuals; modern observations of disarticulation and scattering will help to solve this, too. In addition, as Hill (1980:134) has noted, ".... disarticulation pattern ... may also explain aspects of differential representation of skeletal parts in fossil accumulations"; the majority of the bird fossils from the Big Sandy Formation are wing elements-a very curious feature of the assemblage, which needs to be explained. CONCLUSIONS This study presents some of the first observations on avian taphonomy and provides a base for further studies, with the goal of understanding the nature and evolution of ancient bird communities. The main results and conclusions of this study are: (1) elimination of bird remains from a potential fossil record may be due to primary removal at death rather than to their supposed fragility--the common explanation-and large predators and scavengers are important agents decreasing the probability of preservation of skeletal remains in the environment of death; (2) adherence of carcasses to the ground increases preservation potential in an environment of light scavenging and high depositional rates, and decreases preservation potential in an environment of heavy scavenging and low depositional rates; (3) there is a distinctive pattern of disarticulation for the avian skeleton, and there is a limit to the scattering of bird bones in the absence of scavengers; (4) carnivore damage to bird bones appears, initially, to be similar to that observed for mammal bones; (5) the above observations on disarticulation, scattering, and damage to bones can provide information on presence, absence, and extent of scavenging and predation; (6) disarticulation and scattering of skeletons can continue after burial. Acknowledgements-I am indebted to many people for their help with this study. E. R. Meyer helped me to obtain rock dove specimens and suggested the method for constructing a string-trailing device. C. G. Spies supplied the carcasses of gulls, identified plants at the site, and contributed numerous suggestions. T. A. and
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F. R. Bickart provided financial support without which my work would not have been possible. George Junne generously photographed the damaged bones. The following people read and commented on one or more drafts of the manuscript, and their suggestions have improved it immeasurably: C. Badgley, R. T. Bakker, A. K. Behrensmeyer, P. Dodson, D. C. Fisher, P. D. Gingerich, D. W. Steadman, and M. G. Wolman. I am particularly grateful to Catherine Badgley for many hours of conversation on paleoecology in general and taphonomy in particular, and to Robert T. Bakker for the stimulating discussions which formed such an important part of my undergraduate studies at The Johns Hopkins University. REFERENCES Behrensmeyer, A. K. 1978. Taphonomic and ecologic information from bone weathering. Paleobiology 4:150162. and Dechant-Boaz, D. E. 1980. The Recent bones of Amboseli Park, Kenya, in relation to East African paleoecology; pp. 72-92 in Behrensmeyer, A. K. and Hill, A. P. (eds.), Fossils in the Making: Vertebrate Taphonomy and Paleoecology. The University of Chicago Press, Chicago, xii + 338 pp. Binford, L. R. 1981. Bones: Ancient Man and Modem Myths. Academic Press, New York, xxv + 320 pp. Brain, C. K. 1980. Some criteria for the recognition of bonecollecting agencies in African caves; pp. 107-130 in Behrensmeyer, A. K. and Hill, A. P. (eds.), Fossils in the Making: Vertebrate Taphonomy and Paleoecology. The University of Chicago Press, Chicago, xii + 338 pp. Daniels, M. C. S. 1979. A catalogue of the fossil birds from the Eocene London Clay of England contained in the collection of M. C. S. Daniels. Part I: Text. Privately published, 107 pp. Duff, R. 1952. Pyramid Valley. Pegasus Press, Christchurch, 48 pp. Gifford, D. P. 198 1. Taphonomy and paleoecology: A critical review of archaeology's sister disciplines. Advances in Archaeological Method and Theory 4:365-438. Haynes, G. 1980. Evidence of carnivore gnawing on Pleistocene and Recent mammalian bones. Paleobiology 6: 341-351. Hill, A. P. 1979. Disarticulation and scattering of mammal skeletons. Paleobiology 5:261-274. 1980. Postmortem damage to the remains of some contemporary East African mammals; pp. 131-152 in Behrensmeyer, A. K. and Hill, A. P. (eds.), Fossils in
the Making: Vertebrate Taphonomy and Paleoecology. The University of Chicago Press, Chicago, xii + 338 pp. Laporte, L. F. and Behrensmeyer, A. K. 1980. Tracks and substrate reworking by terrestrial vertebrates in Quaternary sediments of Kenya. Journal of Sedimentary Petrology 50:1337-1346. MacFadden, B. J., Johnson, N. M. and Opdyke, N. D. 1979. Magnetic polarity stratigraphy of the Mio-Pliocene mammal-bearing Big Sandy Formation of western Arizona. Earth and Planetary Science Letters 44:349-364. Matthew, W. D. and Granger, W. 1917. The skeleton of Diatryma, a gigantic bird from the lower Eocene of Wyoming. Bulletin of the American Museum of Natural History 37:307-326. Murie, O. J. 1954. A Field Guide to Animal Tracks. Houghton Mifflin Company, Boston, xvii + 375 pp. Olson, S. L. and Hilgartner, W. B. 1982. Fossil and subfossil birds from the Bahamas; pp. 22-65 in Olson, S. L. (ed.), Fossil Vertebrates from the Bahamas. Smithsonian Contributions to Paleobiology No. 48, Smithsonian Institution Press, Washington, 65 pp. and James, H. F. 1982. Prodromus of the fossil avifauna of the Hawaiian Islands. Smithsonian Contributions to Zoology No. 365, Smithsonian Institution Press, Washington, vi + 59 pp. - and Pregill, G. K. 1982. Introduction to the paleontology of Bahaman vertebrates; pp. 1-7 in Olson, S. L. (ed.), Fossil Vertebrates from the Bahamas. Smithsonian Contributions to Paleobiology No. 48, Smithsonian Institution Press, Washington, 65 pp. Rich, P. V. 1980. Preliminary report on the fossil avian remains from late Tertiary sediments at Langebaanweg (Cape Province), South Africa. South African Journal of Science 76:166-170. Rickleffs, R. E. and Gill, F. B. 1980. Fifty years of American ornithology. Bulletin of the British Ornithological Club 100:118-122. Schafer, W. 1955. Fossilisation-Bedingungen der Meeressiiuger und V6gel. Senckenbergiana lethaea 36:1-25. 1975. Ecology and Paleoecology of Marine Environments. Translated by I. Oertel. The University of Chicago Press, Chicago, xii + 568 pp. Shipman, P. 1981. Life History of a Fossil. An Introduction to Taphonomy and Paleoecology. Harvard University Press, Cambridge, 222 pp. Toots, H. 1965. Sequence of disarticulation in mammalian skeletons. University of Wyoming Contributions to Geology 4:37-39. Van Tyne, J. and Berger, A. J. 1959. Fundamentals of Ornithology. John Wiley and Sons, Inc., New York, xi + 624 pp.
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A Field Experiment in Avian Taphonomy Author(s): K. Jeffrey Bickart Source: Journal of Vertebrate Paleontology, Vol. 4, No. 4 (Dec.

JVP 4(4), December1984

JVP 4(4), December 1984

tarsus femur humerus radius ulna

JVP 4(4), December 1984

JVP 4(4), December 1984

I ;~~ "-, :' I

JVP 4(4), December 1984

JVP 4(4), December 1984

JVP 4(4), December 1984

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