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Hesperornis (Aves) from Ellesmere Island and palynological correlation of known Canadian localities L.V. Hills, E.L. Nicholls, L. (Koldo) M. Núñez-Betelu, and D.J. McIntyre

Abstract: A tarsometatarsus of Hesperornis from the Kanguk Formation on Ellesmere Island is described and illustrated and constitutes the northernmost occurrence of the genus. Palynological evidence and stratigraphic position indicate that Canadian occurrences of Hesperornis are Campanian in age except the specimen from the Mason River Formation which might be as young as early Maastrichtian. Paleogeographic reconstructions indicate that all specimens are from a seaway extending along the western interior of North America. Résumé : Un tarso-métatarse de Hesperornis trouvé dans la Formation de Kanguk, sur l’île d’Ellesmere, est décrit et illustré, et il constitue la découverte la plus éloignée au nord de ce genre. Les enseignements palynologiques et la position stratigraphique indiquent que les découvertes au Canada de Hesperornis sont d’âge du Campanien, à l’exception du spécimen livré par la Formation de Mason River, dont l’âge pourrait être aussi jeune que Maastrichtien précoce. Les reconstructions paléogéographiques indiquent que tous les spécimens proviennent d’un détroit de mer s’étendant le long de l’intérieur occidental de l’Amérique du Nord. [Traduit par la Rédaction]

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Introduction During a regional stratigraphic and palynological study of the Kanguk Formation in the northeastern Sverdrup Basin in July 1992, vertebrate remains assignable to dinosaurs, mosasaurs, sharks, a bony fish (Xiphactinus sp.) and a bird (Hesperornis sp.) were recovered. The purpose of this paper is to record and describe this occurrence of Hesperornis from Ellesmere Island and to review the distribution and age of Hesperornis in North America. The specimen has been accessioned by the Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta (TMP).

Location and stratigraphy The specimen (TMP 97.4.1) of Hesperornis was recovered 155 m above the base of the Kanguk Formation and approximately 30 m below the contact with the Eureka Sound Group on southwestern Fosheim Peninsula, Ellesmere Island, Arctic Canada (79°45′N, 85°36′W; Fig. 1). The Kanguk Formation is a widespread marine unit in the Received January 7, 1999. Accepted May 14, 1999. L.V. Hills.1 Geology and Geophysics, The University of Calgary, 2500 University Drive NW, Calgary, AB T2N 1N4, Canada. E.L. Nicholls. Royal Tyrrell Museum of Palaeontology, Box 7500, Drumheller, AB T0J 0Y0, Canada. L.K.M. Núñez-Betelu. Department of Stratigraphy and Paleontology, University of the Basque Country, P.O. Box 644, 48080 Bilbao, Basque Country, Spain. D.J. McIntyre. 3503 Underhill Drive NW, Calgary, AB T2N 4E9, Canada. 1

Corresponding author (e-mail: hills@geo.ucalgary.ca).

Can. J. Earth Sci. 36: 1583–1588 (1999)

Sverdrup Basin (Embry 1991) and is composed of shales and sandstones with bentonite beds throughout. It ranges from latest Cenomanian–Turonian to Campanian (probably mid-Campanian) in age according to studies of foraminifera by Wall (1983), palynomorphs by Núñez-Betelu (1994) and Núñez-Betelu and Hills (1992a, 1992b), an ammonite by Hills et al. (1994).

Systematics Hesperornithiformes Family Hesperornithidae Marsh, 1872 Hesperornis Marsh, 1872 Hesperornis sp.

Distal end of a right tarsometatarsus of a small hesperornithid (Fig. 2). The bone is not well preserved and shows some signs of post-depositional abrasion. Metatarsals II and III have been laterally crushed, and metatarsal III is broken at the base of the distal trochlea. A deep, polished groove marks the proximal edge of the vascular foramen between metatarsals III and IV; the rest of metatarsal III is missing. The anterior face of the tarsometatarsus is laterally compressed. There is a deep groove between metatarsals III and IV, terminating in the vascular foramen. A lesser groove is present between metatarsals II and III. As is typical of hesperornithids, the distal end of metatarsal IV is the largest, the transverse diameter (at the level of the vascular foramen) being exactly twice that of metatarsal III. Metatarsals II and III are considerably smaller, and offset along the medial side of the shaft. Although the exact length of metatarsal III cannot be determined, its narrow base suggests that it did not extend as far as metatarsal IV. © 1999 NRC Canada

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Fig. 1. Location of the Hesperornis specimen TMP 97.4.1 (ĺ¤š) in the Kanguk Formation on southwestern Fosheim Peninsula, Ellesmere Island, Canadian Arctic Archipelago in relation to northern North America.

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The trochlear surface of metatarsal IV is asymmetric, with the medial condyle being more strongly developed than the lateral. As in both Hesperornis and Parahesperornis, a sickle-shaped diagonal groove is present proximal to the enlarged inner condyle (Marsh 1880, Pls. XVI, XVII; Martin 1984, Fig. 1B). The predominant size of the outer condyle indicates that TMP 97.4.1 is referable to the Hesperornithidae. In the Baptornithidae, the condyles of metatarsals III and IV are subequal in size (Marsh 1880). It resembles Hesperornis and differs from Parahesperornis in that the outer condyle is at least twice the size of the middle condyle and is more distally situated. In Parahesperornis the outer condyle is only about 25% larger than the middle condyle, and the two are approximately equal in length (Martin 1984). The vascular foramen between the condyles of metatarsals III and IV is present in both Hesperornis and Parahesperornis (Marsh 1880, p. 96; Martin 1984, Fig. 1). The presence of this vascular foramen in Baptornis is uncertain. Specimen TMP 97.4.1 is small for Hesperornis. The transverse diameter of the distal edge of trochlea IV is only 10 mm. In the six specimens of Hesperornis described by Marsh (1880), the comparable measurement is 14–15 mm. This suggests that TMP 97.4.1. may represent a juvenile. There are four recognized species of Hesperornis (Martin 1984). Unfortunately, the present specimen is too incomplete to be identified to the species level and is left as Hesperornis sp. Hesperornis has long been recognized as a boreal form (Martin and Stewart 1982; Bryant 1983; Nicholls 1989; Nicholls and Russell 1990). Its most southerly occurrence is in the Niobrara Formation of Kansas (Nicholls and Russell 1990), whereas TMP 97.4.1 is the most northerly known occurrence. Like many of Russell’s (1967) Anderson River specimens, TMP 97.4.1 appears to be a juvenile. This supports the hypothesis that the birds were breeding at northern latitudes (Russell 1967).

Geological age Samples from Hesperornis-bearing strata from the Kanguk Formation (Fosheim Peninsula, Ellesmere Island), the Smoking Hills Formation (Anderson River, District of Mackenzie, N.W.T.), the Mason River Formation (Anderson River, District of Mackenzie, N.W.T.), and the Pierre Shale (Morden, Manitoba) were examined palynologically to establish their age. All of these assemblages correspond to approximately the same time interval. Fosheim Peninsula, Ellesmere Island, Northwest Territories (specimen TMP 97.4.1) The Hesperornis-bearing interval from the Kanguk Formation at Fosheim Peninsula, Ellesmere Island, contains abundant organic matter, including abundant and wellpreserved marine and terrestrial palynomorph taxa (Table 1). This interval can be correlated with the middle part of Biozone 4 (Núñez-Betelu 1994), upper part of Division H-1 (McIntyre 1974), lower part of Biozone CVIa (Doerenkamp et al. 1976), and upper part of Interval I (Ioannides 1986) and is, therefore, early to middle Campanian in age.

1585 Fig. 2. Illustration of the Hesperornis tarsometatarsus (specimen TMP 97.4.1) from the Kanguk Formation. Scale bar = 1 cm.

Anderson River, Northwest Territories Sample C-132693 (shale from bones of associated fauna) from the Smoking Hills Formation at Husky Bend on the Anderson River contains abundant organic material, much of it amorphous and a rich dinoflagellate assemblage with numerous species (Table 1). This dinoflagellate assemblage is similar to that described from Division H-1 in the Smoking Hills Formation at Horton River by McIntyre (1974). Chatangiella ditissima, abundant in C-132693, is also abundant in Division H-1 and was assigned an early Campanian age. Chatangiella hexacalpis, which is common in C-132693, was described from the early Campanian of the Western Interior by Harker et al. (1990). The occurrence of Chatangiella spectabilis, Dorocysta litotes, Isabelidinium acuminatum, and Rhiptocorys veligera also indicates correlation with the middle part of Division H-1 where Dorocysta litotes last occurs and Isabelidinium acuminatum makes its first appearance. The dinoflagellate species present in C-132693 clearly indicate correlation with Division H-1 of McIntyre (1974) and also indicate an early Campanian age. Mason River Formation, Northwest Territories Sample C-132694 (shale from bones of associated fauna) from the Mason River Formation, also at Husky Bend on the Anderson River, contains abundant organic material but has a limited dinoflagellate assemblage with few species (Table 1). The presence of Laciniadinium firmum (Harland) Morgan and abundant Chatangiella biapertura (McIntyre) Lentin and Williams indicates correlation with Divison H-3 of McIntyre (1974). The sample is probably late Campanian, but an early Maastrichtian age is possible. Morden, Manitoba, Pembina Member, Pierre Shale Three samples (F74.04.06, M84.12,18, and P80.06.14) from the Pembina Member of the Pierre Shale at Morden, Manitoba, contain dinoflagellate assemblages which have many species in common with assemblages from Horton River, Anderson Plain, and the Kanguk Formation on Ellesmere Is© 1999 NRC Canada

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Fig. 3. Paleogeographic reconstruction of North America during the Campanian (modified after Williams and Stelck 1975; Kauffman 1984; Nicholls and Russell 1990; and Hills et al. 1994). Stars indicate locations where Hesperornis has been recovered. 1, Echooka River, Alaska (Bryant 1983); 2, Anderson River, Northwest Territories (Russell 1967); 3, Pembina Member, Pierre Shale, Morden, Manitoba (Bardack 1968; Nicholls and Russell 1990); 4, Foremost Formation, South Saskatchewan River, Alberta (Fox 1974); 5, Judith River Formation, Montana (Gilmore 1915), and Clagget Formation, Dog Creek, Fergus County, Montana (Shufeldt 1915); 6, Pierre Shale, South Dakota (MacDonald 1951); 7, Pierre Shale, Nebraska (Marsh 1880; Martin and Tate 1967); 8, Mesaverde Formation, Wyoming (Case 1978); 9, Niobrara Chalk, Kansas (Marsh 1880); 10, Niobrara Formation, South Dakota (Martin and Varner 1992); 11, Pierre Shale, Saskatchewan (Tokaryk 1999).

land. The abundance of C. ditissima and the occurrence of Chatangiella decorosa, C. hexacalpis, I. acuminatum, Laciniadinium biconiculum, and R. veligera indicate that the samples can be correlated with Division H-1 of McIntyre

(1974) from the Anderson Plain. Chatangiella hexacalpis was recorded from the Pembina Member by Harker et al. (1990). The dinoflagellate floras indicate an early Campanian age for the samples. ÂŠ 1999 NRC Canada

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Table 1. Dinoflagellate taxa from the Hesperornis sp. localities in Canada.

Dinoflagellates Alisogymnium euclaense Alterbidinium minus Chatangiella biapertura Chatangiella decorosa Chatangiella ditissima Chatangiella granulifera Chatangiella hexacalpis Chatangiella spectabilis Chamydophorella discreta Chamydophorella nyei Dinogymnium sibiricum Dorocysta litotes Endosorinium campanula Fromea chytra Fromea fragilis Heterosphaeridium difficile Isabelidinium acuminatum Isabelidinium cooksoniae Kallosphaeridium ringnesiorum Laciniadinium arcticum Laciniadinium biconiculum Laciniadinium firmum Laciniadinium williamsi Rhiptocorys veligera Scriniodinium crystallinum Spinidinium clavus Spinidinium uncinatum Spongodinium delitiense Trithyrodinium suspectum

Fosheim Peninsula

Anderson River

Kanguk Formation

Smoking Hills Formation

Morden Mason Formation

Pierre Shale

× × × × × × × ×

× × ×

× × × × ×

× × ×

× × × × ×

× × × ×

× × ×

× ×

× × × ×

Paleogeography

Conclusions

The North American distribution and paleogeography of Hesperornis are summarized in Fig. 3. Many papers have examined the paleogeography of the formations involved (Williams and Stelck 1975; Kauffman 1984; Nicholls and Russell 1990; Hills et al. 1994). These reconstructions indicate that Hesperornis inhabited a Campanian seaway extending north–south along the Western Interior of North America. The specimen from Alaska (Coniacian to Campanian, Bryant 1983) may pose a problem if it is as old as Coniacian. The Campanian age, however, fits with the occurrences reported by Russell (1967) for the Anderson River and the currently reported occurrence on Fosheim Peninsula, Ellesmere Island. All occurrences are in marine strata except possibly the specimen recorded by Fox (1974). In Alberta, the Foremost Formation contains marginal marine, coastal sediments and interbeds of terrestrial sediments (Ogunyomi and Hills 1977; McNeil et al. 1995), and therefore the specimen collected by Fox could also have been recovered from marginal marine sediments. However, at least one genus of Hesperornithiformes inhabited coastal if not terrestrial margins (Tokaryk and Harington 1992). Feduccia (1996) summarizes known occurrences of Hesperornis species and notes also that all were from estuarine to marine strata with some records from many miles offshore.

The new occurrence, a juvenile, the northernmost known occurrence of Hesperornis, along with research by Russell (1967) suggest that these birds were reproducing at these high latitudes. Paleogeographic reconstructions (Fig. 3) indicate that the northern and southern occurrences were connected by a seaway and that further studies should yield specimens in the intervening area.

Acknowledgments The authors would like to acknowledge the support of the Natural Sciences and Engineering Research Council of Canada (L.V. Hills), Polar Continental Shelf Project, the Geological Survey of Canada, the Royal Tyrrell Museum of Palaeontology, University of Calgary, and University of Basque Country for financial support to L.K.M. NúñezBetelu. The authors appreciate the reviews by C.R. Harington, P. McNeil, and T.T. Tokaryk.

1588 Bryant, L.J. 1983. Hesperornis in Alaska. PaleoBios, 40: 1–8. Case, G. 1978. News from members, Eastern Region, Jersey City. News Bulletin, Society of Vertebrate Paleontology, 114: 16–17. Doerenkamp, A., Jardine, S., and Moreau, P. 1976. Cretaceous and Tertiary palynomorph assemblages from Banks Island and adjacent areas (N.W.T.). Bulletin of Canadian Petroleum Geology, 24(3): 272–417. Embry, A.F. 1991. Mesozoic history of the Arctic Islands. In Geology of the Inuitian Orogen and Arctic Platform of Canada and Greenland. Edited by H.P. Trettin. Geological Survey of Canada, Geology of Canada, No. 3, Chapt. 14, pp. 371–433. Feduccia, A. 1996. The origin and evolution of birds. Yale University Press, New Haven and London. Fox, R.C. 1974. A middle Campanian nonmarine occurrence of the toothed bird Hesperornis Marsh. Canadian Journal of Earth Sciences, 11: 1335–1338. Gilmore, C.W. 1915. Hunting vertebrate fossils in Montana. Smithsonian Institute, Miscellaneous Collections, 65(6): 15. Harker, S.D., Sarjeant, W.A.S., and Caldwell, W.G.E. 1990. Late Cretaceous (Campanian) organic-walled microplankton from the Interior Plains of Canada, Wyoming and Texas: biostratigraphy, paleontology and paleoenvironment interpretation. Palaeontographica, Abteilung B, 219: 1–243. Hills, L.V., Braunberger, W.F., Núñez-Betelu, L.K., and Hall, R.L. 1994. Paleogeographic significance of Scaphites depressus in the Kanguk Formation (Upper Cretaceous), Axel Heiberg Island, Canadian Arctic. Canadian Journal of Earth Sciences, 31: 733–736. Ioannides, N.S. 1986. Dinoflagellate cysts from Upper Cretaceous – Lower Tertiary sections, Bylot and Devon islands, Arctic Archipelago. Geological Survey of Canada, Bulletin 371. Kauffman, E.G. 1984. Paleobiogeography and evolutionary response dynamics in the Cretaceous Western Interior Seaway of North America. In Jurassic–Cretaceous biochronology and paleogeography of North America. Edited by G.E.G. Westermann. Geological Association of Canada, Special Paper 27, pp. 273–306. MacDonald, J.R. 1951. The fossil vertebrata of south Dakota. In Guidebook, Fifth Field Conference, Society of Vertebrate Paleontology, West South Dakota, pp. 63–74. Marsh, O.C. 1880. Odontornithes: a monograph on the extinct toothed birds of North America. Government Printing Office, Washington, D.C. Martin, J.E., and Varner, D.W. 1992. The occurrence of Hesperornis in the Late Cretaceous Niobrara Formation of South Dakota. Proceedings of the South Dakota Academy of Science, 71: 95–97. Martin, L.D. 1984. A new hesperornithid and the relationships of Mesozoic birds. Transactions of the Kansas Academy of Science, 87(3–4): 141–150. Martin, L.D., and Stewart, J.D. 1982. An ichthyornithiform bird from the Campanian of Canada. Canadian Journal of Earth Sciences, 19: 324–327. Martin, L.D., and Tate, J., Jr. 1967. A Hesperornis from the Pierre Shale. Nebraska Academy of Science Proceedings, 77: 40. McIntyre, D.J. 1974. Palynology of an Upper Cretaceous section, Horton River, District of MacKenzie, N.W.T. Geological Survey of Canada, Paper 74-14.

Can. J. Earth Sci. Vol. 36, 1999 McNeil, D.H., Wall, J.H., and Eberth, D. 1995. Parasequences and foraminiferal distributions in the Campanian Foremost Formation of southern Alberta. Geological Survey of Canada, Open File 3058, pp. 93–97. Nicholls, E.L. 1989. Marine vertebrate of the Pembina member of the Pierre Shale (Campanian, Upper Cretaceous) of Manitoba and their significance to the biogeography of the Western Interior Seaway. Ph.D. thesis, University of Calgary, Calgary, Alta. Nicholls, E.L., and Russell, A.P. 1990. Paleobiogeography of the Cretaceous Western Interior Seaway of North America: the vertebrate evidence. Palaeogeography, Palaeoclimatology, Palaeoecology, 79: 149–169. Núñez-Betelu, L.K. 1994. Sequence stratigraphy of a coastal to offshore transition, Upper Cretaceous Kanguk Formation: a palynological, sedimentological, and rock-eval characterization of a depositional sequence, northeastern Sverdrup Basin, Canadian Arctic. Ph.D. thesis, University of Calgary, Calgary, Alta. Núñez-Betelu, L.K., and Hills, L.V. 1992a. Preliminary paleopalynology of the Kanguk Formation (Upper Cretaceous), Remus Creek, Ellesmere Island, Canadian Arctic Archipelago: 1. Marine palynomorphs. Revista Española de Paleontographica, 7(2): 185–196. Núñez-Betelu, L.K., and Hills, L.V. 1992b. Preliminary paleopalynology of the Kanguk Formation (Upper Cretaceous) Remus Creek, Ellesmere Island, Canadian Arctic Archipelago: 2. Terrestrial palynomorphs. Revista Española de Paleontographica, 7(2): 197–206. Ogunyomi, O., and Hills, L.V. 1977. Depositional environments, Foremost Formation (Late Cretaceous), Milk River area, southern Alberta. Bulletin of Canadian Petroleum Geology, 25: 929– 968. Olson-Storrs, L., and Parvis, D.C. 1987. The Cretaceous birds of New Jersey. Smithsonian Contributions to Paleobiology, 63: 1– 22. Russell, D.A. 1967. Cretaceous vertebrates from the Anderson River, N.W.T. Canadian Journal of Earth Sciences, 4(1): 21–38. Shufeldt, R.W. 1915. The fossil remains of a species of Hesperornis found in Montana. Auk, 32(3): 290–294. Tokaryk, T.T. 1999. The toothed bird Hesperornis (Hesperornithiformes) from the Pierre Shale (Late Cretaceous) of Saskatchewan. Canadian Field Naturalist, 113(4). Tokaryk, T.T., and Harington, C.R. 1992. Baptornis sp. (Aves: Hesperornithiformes) from the Judith River Formation (Campanian) of Saskatchewan, Canada. Journal of Paleontology, 66(6): 1010–1012. Wall, J.H. 1983. Jurassic and Cretaceous foraminiferal biostratigraphy in the eastern Sverdrup Basin, Canadian Arctic Archipelago. Bulletin of Canadian Petroleum Geology, 31: 246– 281. Williams, G.D., and Stelck, C.R. 1975. Speculations on the Cretaceous paleogeography of North America. In The Cretaceous System in the Western Interior of North America. Edited by G.E. Caldwell. Geological Association of Canada, Special Paper 13, pp. 1–20.