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286 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FIGURE 1.—Distribution of localities, or groups of localities if situated close to each other, of enantiornithine birds (solid circles). later Turgai Strait (Figure 3). Later on, there was no land connection between Laurasia and Gondwana until the Tithonian, when the present Gibraltar Strait was already very narrow. It seems, however, that the present-day Iberian Peninsula was colonized from other parts of Europe in the Valanginian-Berriasian because previously it had been an island surrounded by ^ > FIGURE 2.—Coastlines in the Valanginian-Berriasian, 138 Ma BP (modified from Smith et al., 1995, reprinted with the permission of Cambridge University Press). Squares indicate Early Cretaceous enantiornithine localities in Spain and northeastern China, which at that time were situated on continents divided by the Turgai Strait. (E=Europe, IP=Indian Peninsula, TS=Turgai Strait.) more or less wide seas. After the Tithonian, the part of the Tethys dividing Laurasia and Gondwana was wide again until the Tertiary. Probably the enantiornithine birds inhabiting these parts of the earth evolved independently during that time. The colonization of Gondwana took place in the Bajocian via the eastern part of North America and Africa (although we do not have any evidence for the occurrence of the Enantiornithes in Africa), in view of its land connection with South America C=> FIGURE 3.—Coastlines in the Bajocian, 170 Ma BP (modified from Smith et al., 1995, reprinted with the permission of Cambridge University Press). Arrows indicate hypothetical directions of dispersal of Enantiornithes.
NUMBER 89 287 and Australia by way of the Antarctic (Figure 4). A second wave of colonization of North America was possible later, independent of the colonization of Gondwana, in the Early Cretaceous (Valanginian). In Chiappe's (1991) opinion, the Enantiornithes evolved in Gondwana, whereas Zhou (1995a) claims that Laurasia was their cradle. In Chiappe's (1991) conception, the whole process of colonization of the earth ran in the opposite direction, which, from the viewpoint of the history of continents, also is possible. If the Enantiornithes separated in the Bajocian, they theoretically could have originated anywhere on the earth. Vorona berivotrensis Forster et al. (1996), discovered in Madagascar and considered to be a sister group of the Enantiornithes, speaks in favor of Chiappe's conception. On the other hand, the age of the remains and the differentiation of the forms from Laurasia seem to support Zhou's (1995a) opinion, and this is the reason for adopting my present course of reasoning. No matter which of these two theories is right, the history of continents indicates that the subclass Enantiornithes evolved in the Middle Jurassic, more than 25 million years before Archaeopteryx. The genera Nanantius and Enantiornis were first described from Gondwana in the Albian of Australia (Molnar, 1986) and the Maastrichtian of South America (Walker, 1981), respectively. The acceptance of land dispersal for the Enantiornithes against a background of the history of continents raises doubts that the Late Cretaceous remains mentioned from Uzbekistan and the Gobi Desert (Nesov and Panteleev, 1993; Kurochkin, 1995a, 1996) could actually belong to these genera. Even if their flight abilities were considerably greater than in the Early Cretaceous Enantiornithes, at that time the oceans between all Bocheriski, Z. 1997("1996"). Enantiornithes—dominuj^ca grupa ladowych ptakow kredowych [Enantiornithes—A Dominant Group of the Cretaceaous Terrestrial Birds]. Przegl^d Zoologiczny, 40(3^1): 175-184, 4 figures. [In Polish, with English summary. Date on title pages is 1996; actually published in 1997.] Chatterjee, S. 1991. Cranial Anatomy and Relationships of a New Triassic Bird from Texas. Philosophical Transactions of the Royal Society of London, Biological Sciences, 332(1265):277-342, 40 figures. London: The Royal Society. 1994. Protoavis from the Triassic of Texas: The Oldest Bird. [Abstract.] Journal fur Ornithologie, 135(3):330. Chiappe, L.M. 1991. Cretaceous Birds of Latin America. Cretaceous Research, 12:55-63. 1993. Enantiornithine (Aves) Tarsometatarsi from the Cretaceous Lecho Formation of Northwestern Argentina. American Museum Novitates, 3083: 27 pages, 13 figures. Dong, Zhi-Ming 1993. A Lower Cretaceous Enantiornithine Bird from the Ordos Basin of Inner Mongolia, People's Republic of China. Canadian Journal of Literature Cited FIGURE 4.—Coastlines in the Albian, 105 Ma BP (modified from Smith et al., 1995, reprinted with the permission of Cambridge University Press). The star indicates the Albian locality of Nanantius in Queensland. Arrow indicates the latest possibility of colonization of Australia, assuming that the Antarctic was colonized earlier (not later than in the Tithonian). (IP=Indian Peninsula.) the places mentioned above were too wide to permit crossing (see Figure 4 and Rich, 1976). It also is doubtful that the genus Nanantius would have survived for 25 million years (i.e., from the Albian to the Campanian) or even longer. Earth Sciences, 30:2177-2179. Forster, C.A., L.M. Chiappe, D.W. Krause, and S.D. Sampson 1996. The First Cretaceous Bird from Madagascar. Nature, 382:532-534, 4 figures. Haq, B.W., and F.W.B. Van Eysinga 1987. Geological Time Table. Fourth edition. Amsterdam: Elsevier Science Publishers. [Wall plate.] Hou, L. 1995. Morphological Comparisons between Confuciusomis and Archaeopteryx. In A. Sun and Y. Wang, editors, Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota; Short Papers, pages 193-202. Beijing: China Ocean Press. Kurochkin, E.N. 1995a. The Assemblage of the Cretaceous Birds in Asia. In A. Sun and Y. Wang, editors, Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota; Short Papers, pages 203-208, 3 figures. Beijing: China Ocean Press. 1995b. Synopsis of Mesozoic Birds and Early Evolution of Class Aves. Archaeopteryx, 13:47-66. 1996. A New Enantiornithid of the Mongolian Late Cretaceous, and a General Appraisal of the Infraclass Enantiornithes (Aves). 50 pages, 13 figures, 3 plates. Moscow: Russian Academy of Sciences,
- Page 245 and 246: NUMBER 89 235 mal was found under t
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- Page 309 and 310: NUMBER 89 299 would seem unlikely a
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286<br />
SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY<br />
FIGURE 1.—Distribution of localities, or groups of localities if situated close to each other, of enantiornithine<br />
birds (solid circles).<br />
later Turgai Strait (Figure 3). Later on, there was no land connection<br />
between Laurasia and Gondwana until the Tithonian,<br />
when the present Gibraltar Strait was already very narrow. It<br />
seems, however, that the present-day Iberian Peninsula was<br />
colonized from other parts of Europe in the Valanginian-Berriasian<br />
because previously it had been an island surrounded by<br />
^ ><br />
FIGURE 2.—Coastlines in the Valanginian-Berriasian, 138 Ma BP (modified<br />
from Smith et al., 1995, reprinted with the permission of Cambridge University<br />
Press). Squares indicate Early Cretaceous enantiornithine localities in Spain<br />
and northeastern China, which at that time were situated on continents divided<br />
by the Turgai Strait. (E=Europe, IP=Indian Peninsula, TS=Turgai Strait.)<br />
more or less wide seas. After the Tithonian, the part of the<br />
Tethys dividing Laurasia and Gondwana was wide again until<br />
the Tertiary. Probably the enantiornithine birds inhabiting these<br />
parts of the earth evolved independently during that time.<br />
The colonization of Gondwana took place in the Bajocian via<br />
the eastern part of North America and Africa (although we do<br />
not have any evidence for the occurrence of the Enantiornithes<br />
in Africa), in view of its land connection with South America<br />
C=><br />
FIGURE 3.—Coastlines in the Bajocian, 170 Ma BP (modified from Smith et al.,<br />
1995, reprinted with the permission of Cambridge University Press). Arrows<br />
indicate hypothetical directions of dispersal of Enantiornithes.