Aves: Enantiornithes): New Data on Biostratigraphy and Biogeography

Home , Avisauridae, Avisaurus, Lecho Formation, Neuquenornis, Patagonykus

Asociación Paleontológica Argentina. Publicación Especial 7 ISSN 0328-347X VII International Symposium on Mesozoic Terrestrial Ecosystems: 175-177.Buenos Aires, 30-6-2001

The North American avisaurids (Aves: Enantiornithes): new data on biostratigraphy and biogeography

1 2 Thomas A. STIDHAM ,2 and J. Howard HUTCHISON

Abstraet. In addition to the two described Avisaurus species (A. archibaldi Brett-Surman and Paul and A. gloriae Varrichio and Chiappe), undescribed material from the Maastrichtian contributes important morphologic, biostratigraphic, and biogeographic data on North American avisaurids. A juvenile Avisaurus specimen of an undescribed species from the Maastrichtian Hell Creek Formation of Montana reveals the presence of distal tarsals in enantiornithines and an ontogenetic stage shared with non-avian dinosaurs. All known Avisaurus fossils were deposited in a variety of habitats along the Western Interior Seaway of North America and areas more interior. The presence of other clades of birds in the Hell Creek Formation indicate that avisaurids formed only part of a diverse avifauna at the close of the Cretaceous in North America. Phylogenetic analysis of the new Maastrichtian material places these new specimens as the sister to the other known Avisaurus species within Avisauridae. This hypothesis of avisaurid relationships points to a South American origin of this clade with a dispersal or vicariant event to North America before or early in the Campanian. This event was penecontemporaneous with the Laurasian movement of alvarezsaurids and sauropods. These Montanan enantiornithine specimens represent the northern-most record of this bird clade in North America. These Maastrichtian species and specimens are from very late in the Maastrichtian and are probably the youngest known enantiornithines. The stratigraphic position of these birds with respect to the rest of the fauna indicate that their extinction was nearly synchronous with the extinction of other toothed bird clades and other vertebrates at the end of the Cretaceous.

Key words. Aves. Enantiornithes. Avisaurus. biogeography

Introduction American record is the type specimen Avisaurus gloriae. The fauna of the Two Medicine Formation, in- Enantiornithes is the most diverse Mesozoic cluding A. gloriae, is part of the Campanian Judithian avian clade known, and their fossilized remains have North American Land Mammal Age and is thought been collected from all continents except Antarctica. to be time equivalent with the Judith River Avisauridae, currently known only from the New Formation deposited about 78 million years ago World (North and South America), is among the best (Goodwin and Deino, 1989).Therefore, the age of the known clades of enantiornithines. Several studies earliest record of North American avisaurids (middle support its monophyly (Chiappe, 1993;Chiappe and Campanian) appears to be at least slightly younger Calvo, 1994;Sanz et al., 1995),although this conclu- than the early Campanian or older age of oldest sion was not reached by Kurochkin (1996). South American avisaurid species (Dingus et al., Avisauridae comprises two Argentine species 2000). This stratigraphic and geographic distribution (Neuquenornis volans Chiappe and Calvo and of Avisaurus parallels that of the water ferns, Soroaoisaurus ausiralis Chiappe) and two named Hydropteris (Erwin, 1998), and other vertebrates North American species (Avisaurus archibaldi and A. (McCord, 1998)in North America. gloriae), as well as undescribed Avisaurus material (Stidham, 1998). N ew material The origin of Avisauridae appears to have been before the middle Campanian in South America. The Undescribed avisaurid fossils from the Hell Creek North American record begins in the Campanian of Formation of Montana includes tarsometatarsal frag- Montana and continues into the Maastrichtian of ments representing one or two undescribed taxa, as Montana. The sole described Campanian North well as a large distal tibiotarsus attributable to A. archibaldi. Additional currently indeterminate Hell Creek enantiornithine material (Stidham, 1998)may 'Department of Integrative Biology. represent additional avisaurid material. 'Museum of Paleontology, and lMuseum of Vertebrate Zoology, The presence of three distal tarsals in enantior- University of California, Berkeley, CA 94720, USA. T.A.s. E-mail: [email protected] nithines is confirmed by analysis of x-rays of A. ©Asociación Paleontológica Argentina 0328-347X/01$00.00+50 176 T.A. Stidham and J.H. Hutchison archibaldi and the other Hell Creek material. The [u- are found to belong to other clades as their phyloge- venile avisaurid material demonstrates an ontogenetic positions with respect to North American taxa netic pattern of tarsometatarsal fusions quite differ- become better known. Overall, these groups and ent from ornithuran birds, but similar to many non- many others support the presence of a land bridge avian dinosaurs. The avisaurid tarsometatarsus did (Gayet et al., 1992) between North and South America not pass through a stage in which it had a free distal starting before the middle Campanian, since volant, tarsal cap as in ornithuran birds, and therefore ground-dwelling, and aquatic taxa of various sizes avisaurid (and possibly all enantiornithine) tar- dispersed in both directions within the same interval sometatarsi likely represent a condition not homolo- of time before the late Campanian. This hypothe- gous with the ornithuran tarsometatarsus. sized land bridge coincides with a drop in global sea Phylogenetic analysis of the new avisaurid mate- level after a late Turonian highstand and before a rial indicates monophyly of the North American ma- highstand in the middle Campanian (Haq et al., terial within a monophyletic Avisauridae. These re- 1987), possibly exposing previously submerged por- sults do not support Kurochkin's (1996) phylogenet- tions of Central America. Even with these data, the ic hypothesis of enantiornithine interrelationships. apparent absence of dispersal of other clades, notably The Hell Creek material appears to be the sister multituberculates, eutherians, and most theropod group to A. archibaldi and A. gloriae. Soroavisaurus is clades, during this same interval raises questions the sister taxon to the North American avisaurids, about the selectivity of dispersal between these con- with Neuquenornis as the sister to all other avisaurids. tinents. These are the same results reported by Chiappe (1993) and Varrichio and Chiappe (1995). At this Environment point, we cannot determine if this new Hell Creek material represents two or three undescribed species All currently described Avisaurus material is from of different sizes or is a growth series of one species. the Hell Creek Formation and the Two Medicine Formation in Montana. The Hell Creek Formation was deposited along the western coastal plain of the Biogeography Western Interior Seaway in North America. This The paraphyly of the Argentine sister taxa with coastal plain was forested with rivers and streams respect to the monophyletic North American dissecting it. Other birds known from the Hell Creek Avisaurus species points to a dispersal or vicariant Formation include hesperornithiforms and neor- event from South to North America, resolving the nithines (Stidham, 1998), and they indicate that biogeographic ambiguity cited by Gayet et al. (1992). avisaurids were only a minor part of a very diverse The stratigraphic data from these avian fossils sug- avifauna. In contrast, the Two Medicine Formation gests that this event likely occurred before the middle was deposited further interior, away from the coast, Campanian. This phylogeographic pattem is seen in with Avisaurus gloriae the only bird known from the other vertebrate clades. Within Alvarezsauridae, the formation (Varrichio and Chiappe, 1995). Argentine Alvarezsaurus and Patagonykus are the sister taxa to the Mononykinae in North America and Extinction Asia (Chiappe et al., 1996). In both of these cases, the North American members of these clades appear to The Hell Creek enantiornithine material is latest have dispersed to North America (or the northern Cretaceous in age, and is probably the youngest hemisphere) before the middle Campanian. The known enantiornithine material. Contrary to the presence of mononykines in North America idea of a gradual extinction of toothed bird clades (Hutchinson and Chiappe, 1998), possibly even be- through the Maastrichtian (Tokaryk, 1992), the pres- longing to the same species as described Asian taxa ence of enantiornithines and other toothed bird may indicate that alvarezsaurids dispersed to Asia clades (Stidham, 1998) near the K-T boundary in from South America by way of North America North America supports the possibility that they per- (Novas, 1996). In addition, South American ti- ished along with most dinosaurs in the Cretaceous tanosaurids are probably the sister taxa to the mass extinction. No enantiornithines or other Campanian and Maastrichtian North American toothed bird clades are known from the overlying sauropods (Salgado et al., 1997). On the other hand, Paleocene Tullock Formation or even the Paleocene hadrosaurs, ceratopsians, and ankylosaurs apparent- parts of the Hell Creek Formation. The presence of a ly took the opposite route (North to South America) relatively modern flight mechanism in avisaurids during the Campanian (Gayet et al., 1992; Salgado (Chiappe and Calvo, 1994) seems to eliminate poor and Coria, 1996). This conclusion will need to be re- flight capability as a possible reason for their extinc- vised if the South American members of these clades tion. However, as noted above, differences between

A.P.A. Publicación Especial 7, 2001 The North American avisaurids (Aves:Enantiorntithes) 177

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A.P.A.Publicación Especial 7, 2001