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Phocinae and Phylogenetic Affinities of the Monk Seals

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Phocinae and Phylogenetic Affinities of the Monk Seals AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2924, pp. 1-38, figs. 1-10, tables 1, 2 October 3, 1988 On "Retrogression" in the Evolution of the Phocinae and Phylogenetic Affinities of the Monk Seals ANDRFE R. WYSS' CONTENTS Abstract ...................................................................... 1 Introduction .................................................................. 2 Acknowledgments. 3 Phocid Monophyly and Historical Considerations ................................. 3 Comparative Morphology and Phylogenetics ...................................... 6 "Monachus" Affinities ....................................................... 6 Characters Pertinent to the Placement of "Monachus" ........................... 6 Affinities of the "Monachinae" ................................................ 15 "Monachine" Characters and the "Monachine"/Phocine Division ................. 16 Systematic Conclusions and Implied Patterns of Character Evolution ................ 30 References .................................................................... 36 ABSTRACT The origin of the phocine seals appears to have corroborated hypothesis of the monophyly of the been associated with a pattern of massive char- Phocinae, it is concluded that the monophyly of acter reversal, particularly as concerns features of the "Monachinae" remains significantly more the postcranial skeleton. To establish that the doubtful. Monk seals have been placed within this primitive features occurring in phocines do in fact latter subfamily in all recent phylogenetic treat- represent reversals and not simply retentions from ments of the Phocidae. Analysis here reveals that an ancestral phocid condition, a reexamination of this arrangement is less strongly supported than is the two currently recognized subdivisions of the generally assumed. Moreover, the genus compris- Phocidae is undertaken. In contrast to the strongly ing monk seals, "Monachus," does not itself ap- Department of Vertebrate Paleontology, American Museum of Natural History. Copyright American Museum of Natural History 1988 ISSN 0003-0082 / Price $3.75 2 AMERICAN MUSEUM NOVITATES NO. 2924 pear to be monophyletic. Thus, in the preferred Phocinae must therefore be postulated. This pat- branching arrangement presented here both the tern of character transformation is corroborated "Monachinae" and "Monachus" are portrayed as whether or not "monachine" paraphyly is accept- paraphyletic, with the monk seals placed as se- ed. Moreover, these reversals can be substantiated quential sister-taxa to the remaining phocids. whether or not one ascribes to the multiple origin Certain aspects of the phocine postcranium, of pinnipeds. Acceptance of either "monachine" which may be considered primitive, are not prim- paraphyly or pinniped monophyly, however, ac- itive for phocids as a whole, and an exceptional centuates the pattern of reversal seen among pho- series ofreversals within the clade comprising the cines. INTRODUCTION The so-called "true" or "earless" seals of focus ofthis paper. If other characters can be the family Phocidae are undoubtedly among recruited to support the placement of pho- the most highly specialized members of the cines as the sister-group of other phocids, it eutherian order Carnivora. It is well known might then become judicious to suggest that that in many aspects of the postcranial skel- the postcranial features ofphocines are prim- eton members of the phocid subfamily Pho- itive for phocids. Alternatively, if phocines cinae differ markedly from other phocids. can be shown to be removed from a sister- Hitherto ignored is the observation that many group position, then the unusual primitive ofthe features in which phocines depart from phocine postcr-anial attributes may reason- the conditions typical of other phocids are ably be interpreted as reversals. The appar- decidedly primitive. Also overlooked is the ently conservative aspects of phocine mor- fact that these primitive traits not only set phology include features associated with the phocines apart from other phocids, but serve architecture of the fore- and hind-flippers, as well to distinguish phocines from all other and the more proximal appendicular skele- pinnipeds. From a phylogenetic standpoint, ton, as well as a lesser number of soft ana- then, the situation seen in phocines is com- tomical and cranial traits. patible with two interpretations; either (1) Phocids are currently divided into the these characters represent primitive hold- postcranially plesiomorphic Phocinae and the overs from the ancestral phocid condition, or "Monachinae,"2 which are more typically (2) they represent reversals to more primitive pinniped in many postcranial aspects. One states that have occurred subsequent to the means of establishing that phocines (assum- early diversification of phocids. Preference ing for the moment their monophyly) are not for one ofthese transformational hypotheses the sister-group of other phocids is a dem- necessarily depends on the acceptance of a onstration of the paraphyly of the presumed particular notion of phocid phylogeny and phocine sister-group the "monachines." thus the remarks presented below are pri- Therefore I first examine the presumed basal marily aimed at an examination of some of ("monachine"/phocine) phocid dichotomy, the more general aspects ofthe interrelation- dealing specifically with the monophyly of ships of these highly aquatic carnivorans. "Monachus"-a taxon widely recognized as The greatly divergent morphology of pho- representing an early offshoot in the evolu- cids, coupled with their scant fossil record tion of phocids-and a consideration of the and a general historical lack of detailed ex- affinities of its three Recent species to other amination, has resulted in both a poor un- phocids. This leads to an evaluation of the derstanding of the origin of the group as a evidence bearing on the monophyly of the whole and considerable uncertainty regard- higher taxonomic group of which "Mona- of its ing the mutual phylogenetic affinites 2 Because the monophyly of the "Monachinae" and members. Higher-level intra-phocid rela- its type genus "Monachus" has yet to be securely dem- tionships are crucial to deciding between the onstrated and is in fact called into question by this anal- two alternative sequences of transformation ysis, these nomena are included in quotes throughout outlined above, and thus form the principal this paper. 1988 WYSS: PHOCINE SEALS 3 chus" is widely held as being a member, the are monophyletic. Advanced features com- "Monachinae." This phylogenetic frame- mon to members ofthis highly modified group work then permits the central objective of are many, including: the inability to draw the this paper, a more detailed consideration of hind limbs forward during terrestrial loco- the occurrence ofwide-scale character rever- motion, the lack of a lesser trochanter on the sal in the evolution of phocine seals, a pos- femur, the lack of an alisphenoid canal, a sibility suggested above. transversely directed basal cochlear whorl, an The present account is not intended as an opening of the cochlear fenestra outside the exhaustive treatment ofintraphocid relation- tympanic cavity to form a cochlear foramen, ships; it merely represents a preliminary ex- the lack of an internal auditory meatus and amination of these affinities as they pertain prefacial commissure, strongly everted ilia, a specifically to the questions outlined above. massively developed posterior astragalar I make no claim to either the originality of process and greatly reduced calcaneal tuber, any ofthe observations made or to any ofthe heavily pachyostotic mastoid region, ptery- characters considered. My efforts are aimed goids reaching the auditory bulla, pit for in- simply at putting into a phylogenetic context sertion oftympanohyal anterior to stylomas- one particular-and highly peculiar-char- toid foramen, greatly inflated heavily ossified acter distribution. caudal entotympanics, and an ilial psoas ma- jor insertion (Weber, 1904; Gray, 1905; Faw- ACKNOWLEDGMENTS cett, 1918; Howell, 1929; Kummer and NeiB3, Chester Tarka is gratefully acknowledged 1957; Burns and Fay, 1970; Repenning, 1972; for sharing with a most recalcitrant pupil what Hunt, 1974; deMuizon, 1981a, 1981b; King, he could ofhis prodigious photographic skills. 1983). Features common to phocids that also I thank Lorraine Meeker for her ever patient are approached or matched in Odobenus, its tutelage with the figures and for her pains- close fossil allies, and/or the Miocene allo- taking "heroics" with figure 10. I have ben- desmine and dematophocine "otarioids" in- efited greatly from discussions with, and the clude: greatly expanded cochlear fenestra, continued encouragement of, Dr. Richard H. greatly expanded basal whorl of the scala Tedford. Drs. Tedford, Sydney Anderson, tympani, separation of the entrances of cra- Annalisa Berta, Francis H. Fay, John J. Flynn, nial nerves VII and VIII, inguinal testes, dark Malcolm C. McKenna, Michael J. Novacek, epidermis, hind limb propulsion, greatly en- and Randall T. Schuh read and provided larged incus, malleus and epitympanic recess, valuable comments on the manuscript. I prof- mortised jugal-squamosal contact, nasals ited enormously from a particularly detailed wedging deeply between the frontals, short and incisive critique by Dr. Fay; I am deeply premaxillary-nasal contact, absence of pin- appreciative
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