PDF (Lo-Res) - Smithsonian Institution Libraries

More documents

Recommendations

Info

166 birds, such as the two species of Lagopus, which abounded in the steppe-tundra environment, also underwent changes upon its demise, such as reduced geographic ranges and body size. Interestingly, the largest subspecies of the genus, L. I. major, lives today on the steppes of Kazakhstan. In support of this idea is the fact that all the fossil populations examined, except that from Rebielice Krolewskie (the late Pliocene site), come from deposits considered to belong to cold phases of the Pleistocene. The oldest members of the genus Lagopus examined after those from Rebielice come from a cold horizon above the interglacial at Westbury-sub-Mendip in England, which is early middle Pleistocene (oxygen isotope stage 12). Unfortunately, the fossils are few in number and are fragmentary, which makes it difficult to assess to which species they belong. They do, however, possess relatively robust tarsometatarsi, so it may be that Lagopus was already adapted to the steppe-tundra and was larger in relation to today's birds. The next youngest assemblage examined in this survey is that from La Fage, which is late middle Pleistocene. Both species are definitely present, although it may be significant that they appear to be less divergent from each other in their tarsometatarsal lengths than are modern birds (Figure 2). This may be support for Mourer-Chauvire's (1993) suggestion that the species had diverged not long before. The greater areas of the crista pectoralis and crista bicipitalis may indicate that both L. lagopus and L. mutus were larger in the Pleistocene. If there were a primary selective force for large body size, so that birds were heavier, they would require greater muscle bulk to fly, which in turn adds further to body weight. Alternatively, the birds may have become larger because of selection for better-developed flight muscles under a different climatic regime when the birds may have been less sedentary. This hypothesis would be bolstered by the findings of Bocheriski (1974, 1985) and Bocheriski and Tomek (1994), who demonstrated that the distal-wing elements of Lagopus lagopus and L. mutus during the Pleistocene of Poland and Austria were relatively longer than in present-day birds, and that their legs were relatively shorter. This conclusion, however, was not confirmed by the samples analyzed in the present study. Bocheriski (1974) claimed that there was a clear, positive correlation between temperature and tarsometatarsus length, although he pointed out that local vegetation type also was influential. The nature of the variability seen in tarsometatarsal length over both time and space implies that, unlike the shaft widths, local factors may have had an influence. This seems more likely than the variation being a reflection of the other thermoregulatory biogeographic rule (Allen's Rule), which would produce more uniform clines across the birds' former geographic ranges. Therefore, it may be that influences such as the local terrain are more important than the influence of temperature because locomotion is generally regarded as important in determining leg length in mammals (Scott, 1985). Due to the small magnitude of the differences involved and the small size SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY of the birds in relation to the ground relief, however, this conjecture is difficult to test. Conclusion Lagopus lagopus and L. mutus differ allometrically between the Pleistocene and the present, a consistent finding even when fossils from widely distributed areas and times are compared with greatly dispersed modern samples from across Europe. This difference is most readily identified in the tarsometatarsalshaft width and implies a change that was due to a general or widespread effect, not a local adaptation similar to that proposed for the Holocene evolution of the red grouse {L. I. scoticus) in the British Isles (Voous, 1960; Tyrberg 1991). The changes in tarsometatarsal length appear to be reactions to variations in local conditions that remain unknown. The uniformity of change in tarsometatarsal width across Europe, however, implies regional rather than local effects. Global events that occurred at the Pleistocene/Holocene boundary seem to be a likely explanation; however, two problems arise. First, untangling cause and effect between climatic and vegetational factors, which are closely linked, and second, the possibility that a change occurred in the birds' vagility. It is perhaps easier to conceive that the two species of Lagopus changed size due to climate change and its direct effect on vegetation rather than to changes in the bird's degree of sedentariness. The birds appear to have reacted in much the same way as did many mammals that survived the Holocene/ Pleistocene boundary by becoming smaller. It is suggested herein that the change in seasonal length and vegetation type was the primary reason for this, and not temperature. The birds in the northern areas of Europe today, and particularly in mountainous regions further south in the case of L. mutus, probably exist at much the same temperatures as they did in the Pleistocene in southern Europe. This would eliminate temperature as the primary causal factor of size decrease in the genus over this period, because birds of comparable size to those of the Pleistocene are not present in northern Europe today. Dietary shifts caused by vegetational changes are proposed herein as the most significant factor leading to size reduction. It therefore seems reasonable to suggest that the birds in the genus Lagopus in the Palearctic today are relictual populations that originally evolved and diversified into Lagopus lagopus and L. mutus on the steppe-tundras of the late-middle and late Pleistocene. Bocheriski (1974) and Mourer-Chauvire (1975a) had previously demonstrated allometric trends in European Pleistocene Lagopus, but the suggestion of a major reduction in size at the Pleistocene/Holocene boundary over most of Europe is new, as is the suggestion that it was due to the vegetational changes described above. This work has implications for the taxonomic use of allometric differences in skeletal elements. Often such differences are given greater significance than are mere size differences. The example described above clearly shows that a change in size
NUMBER 89 167 may not be conferred to all elements equally. Therefore, dimensions thought to be significant should be considered in terms of their relationship to body size before taxonomic decisions are taken. Furthermore, allometric changes have been observed by neontologists to occur over short periods of time, for example, the increased bill size observed in Geospiza fortis Gould on the island of Daphne Major, Galapagos, as a result of strong selection through differential survival of large-seeded Andrews, Peter 1990. Owls, Caves and Fossils: Predation, Preservation and Accumulation of Small Mammal Bones in Caves, with Analysis of the Pleistocene Cave Faunas from Westbury-sub-Mendip, Somerset, U.K. 231 pages. London: The Natural History Museum. Bertholt, P., and S.B. Terrill 1991. Recent Advances in Studies of Bird Migration. Annual Review of Ecology and Systematics, 22:357-378. Bocheriski, Z. 1974. Ptaki mlodszego czwartorzedu Polsk [The Birds of the Late Quaternary of Poland]. 205 pages. Warszswa-Krakow: Paristwowe Wydawnictwo Naukowe. 1985. Osteological Differentiation in Willow Grouse. Fortschritte der Zoologie (Stuttgart), 30:69-72. 1991. Pliocene Grouse of the Genus Lagopus from Poland. Acta Zoologica Cracoviensia, 34(2):563-577. Bocheriski, Z., and T. Tomek 1994. Fossil and Subfossil Bird Remains from Five Austrian Caves. Acta Zoologica Cracoviensia, 37(2):347-358. Bridgland, D.R. 1994. Quarternary of the Thames. 441 pages. London: Chapman and Hall. Chaline, Jean 1975. Les rongeurs, Page et la chronologie climatique du remmplissage de l'aven de la Fage (Correze). Nouvelles Archives du Museum d'Histoire Naturelle de Lyon, 13:113-117. Cramp, S., editor 1980. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic, 2: Hawks to Bustards. viii+693 pages. Oxford: Oxford University Press. Davis, Simon, J.M. 1981. The Effects of Temperature Change and Domestication on the Body Size of Late Pleistocene to Holocene Mammals of Israel. Paleobiology, 7(1):101-114. Dement'ev, G.P, and N.A. Gladkov, editors 1967. Birds of the Soviet Union. Volume 4. Jerusalem: Israel Program for Scientific Translation. Dennison, M.D., and A.J. Barker 1991. Morphometric Variability in Continental and Atlantic Island Populations of Chaffinches (Fringilla coelebs). Evolution, 45(l):29-39. Ericson, P.G.P. 1987. Osteology of the Eider Somateria mollissima (L.):A Study of Sexual, Geographic and Temporal Morphometric Variation in the Eider Skeleton. 142 pages. Stockholm: Statens Historiska Museum, Statens Historiska Museum Studies 5. Fleischer, R.F., and R.F. Johnson 1982. Natural Selection on Body Size and Proportions in House Sparrows. Nature, 298:747-749. Literature Cited plants in response to a drought (Grant, 1986). These allometric changes have no taxonomic significance but simply represent organisms adapting to the complex pattern of natural selective pressures acting on any given skeletal dimension. Thus, caution should be used when interpreting allometric differences seen in fossils because the differences seen between Pleistocene and Holocene Lagopus are of the type that have sometimes been considered to have taxonomic significance. Geist, V. 1986. Bergmann's Rule is Invalid. Canadian Journal of Zoology, 65:1035-1038. Grant, PR. 1971. Variation in the Tarsus Length of Birds in Island and Mainland Regions. Evolution, 25:599-614. 1986. Ecology and Evolution of Darwin's Finches. 458 pages. Princeton: Princeton University Press. Guthrie, R.D. 1984. Alaskan Megabucks, Megabulls, and Megarams; the Issue of Pleistocene Gigantism. In H.H. Genoways and M.R. Dawson, editors, Contributions in Quaternary Vertebrate Paleontology: A Volume in Memorial to John E. Guilday. Carnegie Museum of Natural History Special Publication, 8:482-510. Pittsburgh. 1990. Frozen Fauna of the Mammoth Steppe: The Story of Blue Babe. 323 pages. London: The University of Chicago Press. Harrison, C.J.O. 1980. Pleistocene Bird Remains from Torneton Cave and the Brixham Windmill Hill Cave in South Devon. Bulletin of the British Museum (Natural History), Geology, 33(2):91-100. Hedges, R.E.M., R.A. Housley, C. Bronk Ramsey, and G.J. Klinken 1994. Radiocarbon Dates from the Oxford AMS System. Archaeometry, 36(2):337-374. Housley, Rupert A. 1991. AMS Dates from the Late Glacial and Early Postglacial in Northwest Europe: A Review. In N. Barton, A.J. Roberts, and D.A. Roe, editors, The Late Glacial in North-West Europe: Human Adaptation and Environmental Change at the End of the Pleistocene. Council for British Archaeology Research Report, 77:25-39. James, Frances C. 1983. Environmental Component of Morphological Differentiation in Birds. Science, 144:548-550. Janossy, D. 1974. Upper Pliocene and Lower Pleistocene Bird Remains from Poland. Acta Zoologica Cracoviensia, 19(21):531-566. 1976. Plio-Pleistocene Bird Remains from the Carpathian Basin, 1: Tetraonidae. Aquila, 82:13-34. Johnston, R.F., and R.K.. Selander 1964. House Sparrows: Rapid Evolution of Races in North America. Science, 144:548-550. Kraft, E. 1972. Vergleichend morphologische Untersuchungen an Einzelknochen nord und mitteleuropa'ischer kleinerer Huhnervogel. 194 pages. Miinchen: Ludwig-Maximilians-Universitat. Lister, A.M. 1992. Mammalian Fossils and Quaternary Biostratigraphy. Quaternary Science Review, 11:329-344.
Page 1 and 2:
»'' § "S^ iFjwtM . •- .m-i Avia
Page 3:
SMITHSONIAN CONTRIBUTIONS TO PALEOB
Page 6 and 7:
ABSTRACT Olson, Storrs L., editor.
Page 8 and 9:
EARLY WATERFOWL (ANSERIFORMES) AND
Page 10 and 11:
v m SMITHSONIAN CONTRIBUTIONS TO PA
Page 12 and 13:
localities, all of them situated in
Page 14 and 15:
Bone of Sus scrofa Linnaeus, introd
Page 16 and 17:
duboisi are larger than the largest
Page 18 and 19:
8 iver (1897) but afterward were tr
Page 20 and 21:
10 SMITHSONIAN CONTRIBUTIONS TO PAL
Page 22 and 23:
Page 24 and 25:
Page 26 and 27:
16 Cor. Hum. Uln. Cpm. Fern. Tbt. T
Page 28 and 29:
Page 30 and 31:
20 sometatarsus are proportionally
Page 32 and 33:
22 Cor. Hum. Uln. Cpm. Fern. Tbt. T
Page 34 and 35:
Page 36 and 37:
26 Cor. Hum. Cpm. Fern. Tbt. Tmt. P
Page 38 and 39:
Page 40 and 41:
Page 42 and 43:
Page 44 and 45:
34 ability in such a way that it ca
Page 46 and 47:
Page 48 and 49:
Page 50 and 51:
Page 52 and 53:
42 2km SMITHSONIAN CONTRIBUTIONS TO
Page 54 and 55:
Site 13 Research Base ENTRANCE Lowe
Page 56 and 57:
Page 58 and 59:
48 the last has unusually slender w
Page 60 and 61:
50 Si 5 15i 10- 5- 20 J 15 I 10 f 5
Page 62 and 63:
Page 64 and 65:
Page 66 and 67:
Page 68 and 69:
Page 70 and 71:
Page 72 and 73:
Page 74 and 75:
Page 77 and 78:
Comparison of Paleoecological Patte
Page 79 and 80:
NUMBER 89 69 TABLE 1.—Vertebrate
Page 81 and 82:
NUMBER 89 71 Mallorca, probably in
Page 83:
NUMBER 89 Alcover, J.A. 1989. Les a
Page 86 and 87:
Page 88 and 89:
Page 90 and 91:
Page 92 and 93:
Page 95 and 96:
The History of the Chatham Islands'
Page 97 and 98:
NUMBER 89 87 Species Common Name Pe
Page 99 and 100:
NUMBER 89 89 NZA 797,1930,1934 Unco
Page 101 and 102:
NUMBER 89 91 anoramphus spp.), Chat
Page 103 and 104:
NUMBER 89 93 TABLE 2.—Land snails
Page 105 and 106:
NUMBER 89 95 counterparts, with som
Page 107 and 108:
NUMBER 89 population, if such exist
Page 109 and 110:
NUMBER 89 99 FIGURE 11.—Skulls of
Page 111 and 112:
NUMBER 89 101 FIGURE 13.—Lower ma
Page 113 and 114:
NUMBER 89 the Chatham Island Pied O
Page 115 and 116:
NUMBER 89 Locality 9, Western Maung
Page 117 and 118:
NUMBER 89 107 yellowish sand (site
Page 119:
NUMBER 89 109 ogy. Notornis, supple
Page 122 and 123:
112 SMITHSONIAN CONTRIBUTIONS TO PA
Page 124 and 125:
Page 126 and 127: 116 SMITHSONIAN CONTRIBUTIONS TO PA
Page 135 and 136: The Middle Pleistocene Avifauna of
Page 137: NUMBER 89 Accordi, B. 1962. La grot
Page 140 and 141: 130 FIGURE 1.—Map showing locatio
Page 142 and 143: 132 Cranium Mandibula Scapula Clavi
Page 149 and 150: Seabirds and Late Pleistocene Marin
Page 151 and 152: NUMBER 89 141 METHODS Only strictly
Page 153 and 154: NUMBER 89 143 FIGURE 2.—Area of s
Page 161 and 162: NUMBER 89 151 FIGURE 10.—Area of
Page 163 and 164: NUMBER 89 153 FIGURE 12.—Area of
Page 165 and 166: NUMBER 89 155 the period studied. T
Page 167: NUMBER 89 157 Walker, C.A., G.M. Wr
Page 174 and 175: 164 Vl 620 M 570 £ 520 S 470f •
Page 180 and 181: 170 cional Autonoma de Mexico, for
Page 184 and 185: 174 ated with this specimen, see Mi
Page 187 and 188: The Fossil Record of Condors (Cicon
Page 189 and 190: NUMBER 89 179 FIGURE 2.—Geographi
Page 191 and 192: NUMBER 89 181 FIGURE 5.—Vulturida
Page 193 and 194: NUMBER 89 183 FIGURE 7.—Referred
Page 195 and 196: Two New Fossil Eagles from the Late
Page 197 and 198: NUMBER 89 187 TABLE 1.—Measuremen
Page 199 and 200: NUMBER 89 189 carpal trochlea relat
Page 201 and 202: NUMBER 89 191 FIGURE 4.—Holotypic
Page 203 and 204: NUMBER 89 193 We compared the parat
Page 205 and 206: NUMBER 89 195 FIGURE 6.—Distribut
Page 207 and 208: NUMBER 89 197 the Florida State Mus
Page 209 and 210: A New Genus of Dwarf Megapode (Gall
Page 211 and 212: NUMBER 89 201 lis hypotarsi along t
Page 213 and 214: NUMBER 89 203 The fossil is larger
Page 215 and 216: NUMBER 89 205 Clark, George A., Jr.
Page 217 and 218: A New Genus and Species of the Fami
Page 219 and 220: NUMBER 89 209 son with other known
Page 221 and 222: NUMBER 89 211 FIGURE 1.—Argornis
Page 223 and 224: NUMBER 89 213 AM AL AM AL AM AL AM
Page 225 and 226: NUMBER 89 215 caput humeri perpendi
Page 227 and 228:
Selmes absurdipes, New Genus, New S
Page 229 and 230:
NUMBER 89 219 FIGURE 2.—Selmes ab
Page 231 and 232:
NUMBER 89 221 Costae: Deformed frag
Page 233 and 234:
A Fossil Screamer (Anseriformes: An
Page 235 and 236:
NUMBER 89 FIGURE 3.—Chaunoides an
Page 237 and 238:
NUMBER 89 227 B C D FIGURE 6.—The
Page 239 and 240:
NUMBER 89 229 FIGURE 9.—Right tib
Page 241 and 242:
The Anseriform Relationships of Ana
Page 243 and 244:
NUMBER 89 233 Subfamily ANATALAVINA
Page 245 and 246:
NUMBER 89 235 mal was found under t
Page 247 and 248:
NUMBER 89 237 tion, with retroartic
Page 249 and 250:
NUMBER 89 FIGURE 7.—Sternum and p
Page 251 and 252:
NUMBER 89 241 der. The bone is very
Page 253:
NUMBER 89 243 Eocene records of the
Page 256 and 257:
Page 258 and 259:
Page 260 and 261:
Page 263 and 264:
Presbyornis isoni and Other Late Pa
Page 265 and 266:
NUMBER 89 255 FIGURE 1.—Referred
Page 267 and 268:
NUMBER 89 257 vical vertebrae of th
Page 269:
NUMBER 89 259 (Olson and Parris, 19
Page 272 and 273:
Page 274 and 275:
Page 276 and 277:
Page 278 and 279:
Page 280 and 281:
Page 282 and 283:
Page 284 and 285:
274 America. Science, 214(4526): 12
Page 286 and 287:
276 concerning the amphicoelous str
Page 288 and 289:
278 ment of the radius that are con
Page 290 and 291:
280 The left coracoid is represente
Page 292 and 293:
Page 294 and 295:
284 Kurochkin, E.N. 1982. [New Orde
Page 296 and 297:
Page 298 and 299:
288 Palaeontological Institute. [Sp
Page 300 and 301:
290 1991). In this paper we use a "
Page 302 and 303:
Page 305 and 306:
Implantation and Replacement of Bir
Page 307 and 308:
NUMBER 89 297 saurs (Figure 1E-G).
Page 309 and 310:
NUMBER 89 299 would seem unlikely a
Page 311 and 312:
Humeral Rotation and Wrist Supinati
Page 313 and 314:
NUMBER 89 303 apparent. It is now c
Page 315 and 316:
NUMBER 89 305 FIGURE 3.—Dorsal vi
Page 317 and 318:
NUMBER 89 307 FIGURE 4.—Wing of t
Page 319:
NUMBER 89 309 Jura-Museums, Eichsta
Page 322 and 323:
Page 324 and 325:
Page 326 and 327:
Page 328 and 329:
Page 330 and 331:
Page 332 and 333:
Page 335:
Early Evolution of Birds: Roundtabl
Page 338 and 339:
Page 340 and 341:
Page 342 and 343:
Page 344 and 345:
Page 346 and 347:
Page 348 and 349:
338 evolved independently twice." M
Page 350 and 351:
Page 352 and 353:
Page 354 and 355:
Page 356:
1 i'~L ,i "^ 1 •vw* HBB!/'' . m
show all

PDF (Lo-Res) - Smithsonian Institution Libraries

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?