Amia Cf. Pattersoni from the Paleocene Paskapoo Formation of Alberta

Amia cf. pattersoni from the Paleocene Paskapoo Formation of Alberta

Lance Grande, Li Guo-Qing, and Mark V. H. Wilson

Abstract: A well-prepared anterior half of an amiid skull from the Late Paleocene Paskapoo Formation of south-central Alberta is described. The specimen is either very closely related to, or conspecific with, Amia pattersoni Grande and Bemis, 1998, from the Early Eocene Green River Formation of Wyoming. We leave the specimen as Amia cf. pattersoni until additional material is found to further clarify its relationships. Amia cf. pattersoni is the oldest known specimen clearly identifiable as belonging to the genus Amia (sensu Grande and Bemis, 1998), and the Paskapoo species extends the known geographic range of Amia both northward and westward. The fish assemblage of the Paskapoo Formation represents the most diverse freshwater Paleocene fish fauna known from North America. Based on compari- sons of sample size and relative taxonomic diversity to the better known Green River Formation localities of Wyoming, we predict that further collecting will substantially increase the known diversity of the Paskapoo fauna. The Paskapoo Formation, therefore, has great potential to continue adding to the meager knowledge of pre-Eocene freshwater teleost diversity in North America.

Résumé : Ce document décrit une moitié antérieure bien préparée d'un crâne d'un aminoïde de la Formation de Paska- poo du Paléocene tardif. Le spécimen est soit relié très étroitement à Amia cf. pattersoni Grande et Bemis, 1998 de la Formation de Green River de l'Éocène précoce, au Wyoming, soit qu'il lui est congénère. Nous classons le spécimen en tant que Amia cf. pattersoni jusqu'à ce qu'il y ait plus de matériel pour mieux clarifier ses relations. Amia cf. pattersoni est le plus ancien spécimen connu que l'on peut identifier clairement comme faisant partie du genre Amia (sensu Grande et Bemis, 1998) et l'espèce de Paskapoo agrandit l'étendue géographique connue de Amia à la fois vers le nord et vers l'ouest. L'assemblage de poissons de la Formation de Paskapoo représente la faune la plus diversifiée connue de poissons d'eau douce du Paléocène en Amérique du Nord. En comparant des tailles d'échantillons et la diversité taxo- nomique relative aux localités mieux connues de la Formation de Green River au Wyoming, nous pouvons dire avec certitude que la poursuite de l'échantillonnage augmentera de façon substantielle le diversité connue de la faune de Pas- kapoo. La Formation de Paskapoo a donc un grand potentiel contribuer de façon marquée au très peu de connaissances actuelles sur la diversité des poissons téléostéens du pré-Éocène en Amérique du Nord. [Traduit par la Rédaction] Grande et al. 37

Introduction the Paskapoo Formation will be described in a larger study of fossil and living gars.2 Previous descriptions of the other The Late Paleocene Paskapoo Formation of south-central Paskapoo fishes are as follows. Occurrence in Alberta of Alberta contains the most diverse freshwater Paleocene fish amiids in what is now considered to be the genus Cyclurus assemblage known from North America. Although some has been known since 1928 in deposits of both Paleocene taxa in the formation are so far represented only by scales, and Late Cretaceous ages. The nominal taxa Kindleia isolated bones, or parts of the caudal region (i.e., the amiid fragosa Jordan 1927 (Late Cretaceous) and Stylomyleodon Cyclurus, Gonorynchidae, Cyprinoidea, and Asineopidae), lacus Russell 1928a (Paleocene) were both established for others are represented by articulated, nearly complete skele- amiids with flattened, crushing teeth, based on fragments of tons (Lepisosteidae, Osteoglossidae, Esocidae and jaw bones and associated disarticulated material. Contro- Percopsidae), or a partial skull (the amiid Amia cf. versy about the generic (e.g., Russell 1928b) and specific pattersoni described herein). The lepisosteid skeletons from (e.g., Gaudant 1992) distinctiveness of Russell’s Paleocene species has recently been resolved as follows. Jordan’s Late Received April 9, 1999. Accepted October 20, 1999. Cretaceous species is now classified by Grande and Bemis L. Grande.1 Department of Geology, Field Museum of (1998) as Cyclurus fragosus (Jordan), whereas Russell’s Natural History, Roosevelt Road at Lake Shore Drive, Paleocene species is treated as a nomen dubium, Cyclurus Chicago, IL 60605, U.S.A. "lacus" (Russell), by the same authors. Nevertheless, it dem- Li G.-Q. Institute of Vertebrate Paleontology and onstrates the presence of the genus Cyclurus in the Paleoanthropology, Academia Sinica, P. O. Box 643, Beijing Paleocene assemblage. Grande and Bemis (1998) also 100044, China. briefly reported that a specimen assignable to Amia existed Li G.-Q. (present address) and V.H. Wilson. Department of in the collections of the University of Alberta. That speci- Biological Sciences, University of Alberta, Edmonton, AB men is the subject of the present paper. T6G 2E9, Canada. Osteoglossidae were described based on isolated bones by 1Corresponding author (e-mail: [email protected]). Wilson (1980), and the genus Joffrichthys was established 2Work by Grande and Bemis currently in progress. based on articulated skeletons by Li and Wilson (1996).

Hiodontidae were reported by Wilson (1980) on the basis of nasal; pas, parasphenoid; pastp, parasphenoid tooth patch; scales, isolated skull bones, and a partial caudal skeleton, pmx, premaxilla; po, postinfraorbitals; ro, rostral bone; smx, but later discoveries proved that all of these fragments can supramaxilla; so, subinfraorbital. be referred to the osteoglossid Joffrichthys (Li and Wilson 1996). Gonorynchidae were reported based on scales and Systematic paleontology isolated skull bones (including a peculiar clefted subopercle indicative of the genus Notogoneus) by Wilson (1980). Subclass Actinopterygii Cope, 1887 Cyprinoidea (probably catostomids) were recorded based on (sensu Rosen et al., 1981) isolated cleithra by Wilson (1980). Esocidae were described Series Neopterygii Regan, 1923 based on isolated bones and several articulated skeletons by (sensu Rosen et al., 1981) Wilson (1980, 1984). Asineopidae were reported based on Division Halecostomi Regan, 1923 isolated scales and skull bones by Wilson (1980). Percopsidae were briefly reported based on isolated bones (sensu Patterson, 1973) by Wilson (1980); Wilson also mentioned, but did not de- Subdivision Halecomorphi Cope, 1872 scribe, partial skeletons. Murray and Wilson (1996) and (sensu Patterson, 1973) Murray (1996) later described nearly complete skeletons that Order Amiiformes Hay 1929 they assigned to two different genera of Percopsidae. (sensu Grande and Bemis, 1998) Percopsid species make up about 95% of the approximately Superfamily Amioidea Bonaparte, 1838 2000 articulated fish skeletons known from the Paskapoo (sensu Grande and Bemis, 1998) Formation. The University of Alberta specimen assigned by Grande Family Amiidae Bonaparte, 1838 and Bemis (1998) to Amia is the first articulated specimen of (sensu Grande and Bemis, 1998) an amiid yet found in the Paleocene of western Canada. Al- Subfamily Amiinae Bonaparte, 1838 though incomplete, consisting only of the anterior half of the (sensu Grande and Bemis, 1998) skull, it is extremely well preserved. In the present paper, we Genus Amia Linnaeus, 1766 fully describe the specimen, place it in phylogenetic context, (sensu Grande and Bemis, 1998) and review the potential of the Paskapoo Formation to yield increased diversity of Paleocene freshwater fishes. Remarks Grande and Bemis (1998, p. 32), in a comprehensive revi- Materials sion of Amiidae, provided an emended diagnosis for the ge- For a list of amiine specimens examined, see Grande and nus Amia. Characters used to distinguish the genus Amia Bemis (1998). Nominal species from that list reexamined from Cyclurus (another genus in the subfamily Amiinae) in- here include the following: Amia calva Linnaeus, 1766; clude the following: (1) the presence of sharply pointed teeth Amia scutata Cope, 1875; Amia pattersoni Grande and on the anterior coronoids and vomers (versus styliform teeth Bemis, 1998; Amia "robusta" Priem, 1901; "Amia" hesperia on the anterior coronoids and vomers in Cyclurus); (2) long Wilson, 1982; Stylomyleodon lacus Russell, 1928 and narrow tooth patch extending well anterior to the later- (= Cyclurus "lacus" (Russell)); Amia kehreri Andreae, 1893 ally pointing ascending rami of the parasphenoid (versus a (= Cyclurus kehreri (Andreae)); Paramiatus gurleyi Romer shorter tooth patch in Cyclurus; see Grande and Bemis and Fryxell, 1928 (= Cyclurus gurleyi (Romer and Fryxell)); 1998: Fig. 135); and (3) a high number of preural centra, ir- Amia efremovi Sytchevskaya, 1981 (= Cyclurus efremovi relevant with regard to the fossil being described here, which (Sytchevskaya)); Kindleia fragosa Jordan, 1927 (= Cyclurus is missing the vertebrae. fragosus (Jordan)). For an explanation of use of quotation The Paskapoo amiid skull described herein has features marks, see Grande and Bemis (1998, p.19 for genera, (1) and (2) (discussed in description), which identify it as pp. 315, 316 for species). belonging to the genus Amia and as being the oldest and northernmost occurrence of the genus sensu Grande and Abbreviations Bemis, 1998. The specimen has some additional features, strongly resembling those of Amia pattersoni Grande and Institutional Bemis (1998) and discussed after the description below. FMNH, Field Museum of Natural History, Chicago, Ill.; MNHN, Muséum national d’Histoire naturelle (Institut de Amia cf. pattersoni Paléontologie), Paris, France; UALVP, University of Al- (Figs. 1–3) berta, Laboratory for Vertebrate Paleontology, Edmonton, Alta., Canada. Referred specimen UALVP 37150, the anterior part of an articulated skull Anatomical (Figs. 2 and 3). The specimen is preserved as black-colored Terminology follows Grande and Bemis (1998): ang, angu- bones that were originally contained in a soft, light gray lar; ao, antorbital; ct, coronoid teeth; d, dentary; dpl1, sandstone. dermopalatine 1; dpl2, dermopalatine 2; dsp, dermosphenotic; dt, dentary teeth; ecp, ectopterygoid; enp, endopterygoid; g, Remarks gular (median gular or gular plate); l, lacrimal (first or This specimen was briefly mentioned and illustrated in anteriormost infraorbital); le, lateral ethmoid; mx, maxilla; n, Grande and Bemis (1998, pp. 208, 209), where it was stated

Fig. 1. Maps showing the locality in south-central Alberta that produced the specimen of Amia cf. pattersoni described here. The specimen (UALVP 37150) is from the locality labeled as DW-2 on right side map. Right side map is based on Fox (1990).

that its description was “in progress.” The cited description Zingiberopsis and a possible member of the Musaceae follows below. (banana family) are suggestive of year-round, frost-free con- ditions (G. Hoffman, personal communication). Insect fos- Locality and horizon sils from the same layer include dragonflies (Odonata) The specimen was collected by Kenneth L. Soehn in 1991 (Wighton and Wilson 1986), grasshoppers–crickets approximately 1 m below a productive fossil-mammal and (Orthoptera), caddisflies (Trichoptera), larval and adult champsosaur-bone site called "DW-2" (UALVP site number crane flies (Diptera: Tipulidae) and beetles (Coleoptera) 226) on the north (left) bank of the Blindman River, 10 km (Wighton 1982), and aquatic oribatid mites (Acari: NNE of Red Deer, Alberta (Fig. 1). Coordinates of the local- Oribatidae) (Baker and Wighton 1984), an assemblage indic- ity are as follows: 52° 21.5′ N, 114° 45.3′ W; NE 1/4, S 14, ative of quiet freshwater environments. Fragmentary and in- T 39, R 27, W 4; 838 meters above sea level). This is within tact shells of freshwater mollusks are also locally abundant the Lacombe Member of the Paskapoo Formation (Demchuk and often associated with vertebrate fossil remains, as they and Hills 1991). The Lacombe Member includes the Blind- are in the Blindman River outcrops. man River and nearby Burbank outcrops, as well as the Joffre Bridge outcrops of closely similar age (Demchuk and Hills 1991; Fox 1990). Lacombe Member rocks are Description interbedded siltstones, mudstones, shales, and coals, with subordinate sandstones and conglomerates. The rocks of All that remains of this specimen is the anterior half of the DW-2 are of undoubted freshwater origin and within the head, crushed dorsoventrally. Paskapoo Formation (exposures along the Blindman River constitute the type section of the Paskapoo). The Paskapoo Skull roof and dorsal ethmoid region lithologies are believed to represent predominantly low- The posteriormost elements remaining of the skull roof energy fluvial deposition, with associated overbank environ- are the paired frontals (fr, Fig. 2B). Only the anterior halves ments including ponds and oxbow lakes (Hoffman 1995). of the bones remain. The ornamentation on the dorsal sur- According to Fox (1990), the fossil mammals of DW-2 indi- face of these and most other dermal bones is coarse, as is cate an age of Ti3 (Tiffanian, Late Paleocene), similar to that typical for amiidae. Jutting out from below the anterolateral of several other Paleocene sites in the Red Deer area that corner of each frontal is the unornamented lateral ethmoid also yield fossil mammals and fossil fishes. For a description (le, Fig. 2). This element is similar in appearance to that of of other fossil faunas and localities containing amiines, see all amiines. Just anterior to the frontals are the paired nasals Grande and Bemis (1998). (n, Fig. 2B). These elements resemble those of A. pattersoni and "A." hesperia much more closely than they do the nasals Associated nonfish fauna of A. calva or A. scutata, in that they are very elongated an- Fragmentary plant remains are generally abundant, and teriorly. They are sutured to each other along the midline, as well-preserved plant and insect fossils (Wighton and Wilson are the frontals. In A. calva and other amiids, the nasals start 1986) are locally abundant, as they are at the Burbank out- out as autogenous elements that eventually develop connect- crops near the present fossil site on the Blindman River ing sutures to each other in large individuals. Thus, it is (Fig. 1). Plant taxa identified from a layer of montmorillo- likely that smaller specimens of A. cf. pattersoni would also nitic shale cropping out both along the Blindman River and be expected to have autogenous nasals. As in other amiine at Burbank include horsetails (Equisetum), ferns (Osmunda, species, there is an anterior notch on each nasal for the ante- Dennstaedtia, and Onoclea), taxodiaceous conifers rior nares. The dermosphenotic is preserved on the left side (Glyptostrobus), monocots (Zingiberopsis, Alismaphyllites), only (dsp, Fig. 2B). It is firmly sutured into the skull roof, and dicots (Platanus,?Cercidiphyllum). The occurrence of as in other amiines. Its anterior margin forms part of the or-

Fig. 2. Amia cf. pattersoni (UALVP 37150) in dorsal view. Specimen is the anterior half of a head. Anterior facing left in A and B; see section Abbreviations in this paper). (A) Photograph of specimen partly coated with ammonium chloride. (B) Drawing of skull in A with lower jaw bones omitted. (C) Closeup of lower jaw showing long, sharply pointed coronoid teeth (ct). Scale is in millimeters.

bital margin. The posterior part of the bone is broken off, so contact the frontals. Anteriorly, each antorbital closely ap- the relative length and shape of the bone is unknown. proaches a lateral arm of the rostral. Anterior to the nasals is the median rostral (ro, Fig. 2B). This bone is v-shaped and articulates with the anteromedial Lateral bones of the head edges of the nasals. On the anterior surface of each arm of Nearly the complete circumorbital series is preserved on the rostral is a small opening to the rostral canal (= ethmoid the left side, and part of it is also preserved on the right side. commissure). Laterally, the nasals are sutured to elongate Anteriorly, there is a large lacrimal (l, Fig. 2B), which bears antorbitals (ao, Fig. 2B). Posteriorly, the antorbitals do not a large posterior notch for insertion of the first subinfra-

Fig. 3. Amia cf. pattersoni (UALVP 37150) in ventral view. (A) Photograph. (B) Line drawing. Anterior facing left; see section Abbre- viations in this paper.

orbital (so1, Fig. 2B). The subinfraorbitals (so1–3, so1–4, cies of Amia. The dentary teeth are large and recurved, also Fig. 2B) are numerous, with three on one side and four on with sharply pointed tips (dt, Fig. 2C). The dentary and the other. The presence of four or more subinfraorbitals is premaxillary teeth are larger than the maxillary teeth. one of the derived diagnostic characters of Amia pattersoni, and no other amiine is known to have so many (Grande and Ventral skull region and gular Bemis 1998, p. 189). The posterior margin of the orbit is The ventral side of the specimen shows a few additional formed mostly by the postinfraorbitals (po1 and po2, details. The anterior end of the parasphenoid is preserved Fig. 2B). The full sizes and shapes of the postinfraorbitals (pas, Fig. 3B), and shows that the parasphenoid tooth patch are unknown because the posterior portions of these bones (pastp, Fig. 3B) extends extremely far forward, as in are missing on the left side, and these bones are completely A. pattersoni (e.g., see Grande and Bemis 1998, their fig. missing on the right side. 135) and the fragmentary A. "robusta," from the Paleocene of northern France (considered to be a nomen dubium within Jaws Amia by Grande and Bemis 1998, pp. 207, 208). As in all The maxillae (mx, Fig. 2B) are the largest bones of the amiines, the ectopterygoid and endopterygoid (ecp, enp, upper jaw. The posterior ends of both the right and the left Fig. 3) are covered with a fine sandpaper-like shagreen of maxilla are missing. The dorsal margin of each maxilla is tiny teeth over much of their oral surfaces. The palate is deeply notched for articulation with the large supramaxilla missing posterior to the ectopterygoid, and the posterior part (smx, Fig. 2B). The supramaxilla is heavily ornamented on of the ectopterygoid–endopterygoid suture is not clearly pre- its lateral surface. A complete count of maxillary teeth was served on the specimen. Other notable features on the ven- not possible because of the incompleteness of the bone. The tral surface are the large fang-like teeth of the premaxillae (pmx, Fig. 2B) are mostly covered by the rostral dermopalatines (dpl2, Fig. 3) and what appears to have been and nasals. The left side premaxilla bears six teeth, and the an elongate gular, although much of the posterior and some right bears seven. anterior parts of the gular are broken off. Gular shape is ex- The lower jaw is represented in the specimen mostly by trapolated from the portion of gular that is preserved, the the elongate dentaries, coronoids, and parts of the articular center of ornamentation, and comparison to more complete bones. The coronoids bear multiple rows of small, elongate gulars in other species of Amia (e.g., compare Fig. 3 with teeth with sharply pointed tips (ct, Fig. 2C), as do other spe- Grande and Bemis 1998, their figs. 127, 136).

Discussion (1998) were not able to find stomach contents in any of the thousands of Cyclurus specimens examined. However, many Some morphological features of the Paskapoo amiid de- of the Paleocene fossil sites in Alberta that yield amiid scribed here suggest it is either closely related to or scales and bones also contain bedding-plane concentrations conspecific with Amia pattersoni Grande and Bemis, 1998. of broken mollusc shells. It is at least possible that some of These features include the high number of subinfraorbitals these concentrations were produced through feeding on (3–4), the extreme anterior extent of the parasphenoid tooth molluscs by individuals of Cyclurus spp. patch, and the long narrow shape of the gular. The anterior- Discovery of both Amia and Cyclurus in the Paskapoo reaching parasphenoid tooth patch and elongate gular are Formation also emphasizes the difficulty of identifying to characteristics shared with Amia "robusta" Priem (1901), species the many isolated amiid scales, skull elements, and from the late Paleocene (Thanetian) of France (a species postcranial bones in existing and future collections. As ar- placed as nomen dubium in Grande and Bemis 1998). With gued by Grande and Bemis (1998), most of these bones can- no more than the anterior half of the head preserved, we can- not reliably be identified to genus, because bone shapes are not say with much confidence that the Paskapoo amiid is highly variable both in Amia and in Cyclurus species. Pend- conspecific with "A. pattersoni.” One possible difference being discovery of more diagnostic characteristics, most dis- tween the Paskapoo species and A. pattersoni is the presence articulated skeletal elements in the Paskapoo Formation of a posterior notch in the lacrimals for articulation with the should provisionally be classified as Amiinae, gen. et sp. first subinfraorbitals in the Paskapoo specimen. This notch is indet. absent in the three specimens of A. pattersoni examined by The list of fish taxa known from the Paskapoo Formation Grande and Bemis (1998, e.g., their figs. 123, 124, 126) that should continue to grow with continued collecting. Of the had the lacrimals preserved. But even though the lacrimals approximately 2000 articulated fish skeletons known so far are not notched in the few known specimens of from the Paskapoo Formation, the approximate taxonomic A. pattersoni, we choose not to describe this specimen as a proportions are as follows: Percopsidae 95%, Osmeridae new species because of the small sample size of both A. 3.5%, Esocidae 0.8%, Osteoglossidae 0.2%, and pattersoni and the Paskapoo species. The presence or ab- Lepisosteidae 0.1%. Although there are about two thousand sence of this notch could be part of the normal morphologi- articulated fish skeletons known so far from the Paskapoo cal variation within A. pattersoni. Additional specimens will Formation, nearly all of them come from a single mass- clarify this. Thus, we leave the Paskapoo specimen as Amia mortality deposit that has restricted diversity (three species cf. pattersoni. are represented by articulated skeletons: Li and Wilson The Paskapoo specimen is nevertheless important because 1996; Murray 1996; Wilson 1996). Two more species occur it is the oldest known specimen of the genus Amia, sensu among twenty articulated skeletons at a second locality Grande and Bemis (1998). It also extends the geographic (Wilson 1984; Murray and Wilson 1996). The gar species range of Amia both northward and westward within North comes from a third locality. All other occurrences consist of America (Grande and Bemis 1998, their figs. 7, 97, 118, disarticulated bones or isolated finds such as the one de- 133). It also adds to the growing faunal list from the Late scribed in this paper. In the freshwater deposits of the Paleocene of North America. Eocene Green River Formation, where the known sample It is interesting to note that the two amiine genera Amia size is much larger (several hundred thousand fishes), well and Cyclurus are sympatric in the Paskapoo Formation. Sim- over half of the known genera of fishes each occur at a rate ilarly, species of Amia and Cyclurus are sympatric in the of less than one out of a thousand specimens (e.g., see Early Eocene deposits of the Fossil Butte Member of the Grande 1984, their table II.10, and 1999). Consequently, Green River Formation, Wyoming (Grande and Bemis 1998, continued collection and study of Paskapoo fishes has great pp. 185, 244) and in the Late Paleocene deposits of Mont de potential to add to the meager knowledge of pre-Eocene Berra at Cernay, France (Grande and Bemis 1998:207, 317). freshwater teleost diversity in North America. Amia and Cyclurus, although in the same subfamily, appear to have occupied different ecological niches, based on differ- Acknowledgments ent tooth morphology (Grande and Bemis 1998, pp. 643, 644, and their fig. 134). We thank Jim Tyler and Stephen L. Cumbaa for reading Species of the genus Amia, including specimens of and commenting on this manuscript. For collecting the spec- A. pattersoni Grande and Bemis, 1998, and the Albertan imen, we thank Kenneth Soehn. For loan of materials we amiid fossil described in the present paper are likely to have thank Barry Chernoff and Mark Westneat (FMNH); Daniel been primarily piscivorous as adults. The Albertan fossil has Goujet (MNHN); and Eugenia Sytchevskaya (PIN). We also sharply pointed teeth both at its jaw margins and within its thank Elaine Zeiger for help in word processing. mouth, like those in other species of Amia. 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