45 



felt to be among the fossil sister taxa of the phocids (Wyss 1987; Berta 1991; Wyss & 

 Flynn 1993; Berta & Wyss 1994). 



Instead, Erignathus is now embedded within the Phocini (rendering the latter paraphyletic), 

 forming the sister taxon to the clade of Histriophoca plus Pagophilus. Despite the nearly 

 universal agreement on the primitive, almost monachine, nature of Erignathus with respect 

 to the remaining phocines (Chapskii 1955a; King 1966, 1983; Burns & Fay 1970; McLaren 

 1975; Ray 1976a; Wyss 1988a; Berta & Wyss 1994; Arnason et al. 1995), this clade 

 consistently demonstrates the highest number of synapomorphies within the Phocini: seven 

 under DELTRAN optimization (also the number of unequivocal synapomorphies) and 13 

 under ACCTRAN optimization. 



Paraphyly of the Phocini is an extremely uncommon suggestion, speaking against an 

 apparent host of putative chromosomal and morphological synapomorphies (McLaren 

 1960a, 1966, 1975; King 1966; Burns & Fay 1970; Arnason 1974, 1977; Arnason et al. 

 1995). To our knowledge, it has only previously been suggested by de Muizon (1982a), 

 Mouchaty et al. (1995), Perry et al. (1995). and possibly Arnason et al. (1995), with 

 Cystophora occupying roughly the same position indicated here for Erignathus. Yet, there 

 are hints in the literature that Erignathus might not be quite as primitive as it is commonly 

 held to be. Ray (1976a) dismisses suggestions of Erignathus possessing monachine 

 tendencies, instead preferring to view it as a conservative, partly aberrant phocine. 

 Chapskii (1955a), who supports a basal placement for Erignathus, also notes a number of 

 derived features for this genus (mostly pertaining to the feeding apparatus) with respect 

 to the remaining phocines. As well, Erignathus displays a number of karyotypic peculi- 

 arities that otherwise contradict its plesiomorphic chromosome number (Arnason 1974, 

 1977). Wyss (1988a) is entirely correct in regarding these features as being autapomorphic 

 and thus phylogenetically uninformative. Nor do they necessarily indicate a paraphyletic 

 Phocini; however, they do potentially hint at a more derived position for Erignathus within 

 the phocines. This latter supposition is tentatively supported here by the relatively large 

 number of character state changes (23 to 25 unweighted steps), very few of which indicate 

 a more primitive placement, in the branch immediately leading to Erignathus. Within 

 phocids, five of these changes are autapomorphic, 16 are convergent with other phocids, 

 only three are reversals to the plesiomorphic phocid condition, and one is a reversal 

 convergently found in some other phocids (see Appendix E and Character Analysis). Nor 

 does Erignathus appear to display an inordinate amount of convergence on the monachine 

 pattern. Most of the convergent characters converge on the states found in selected 

 monachines only, and a good number are convergent on the states found in other phocines. 

 On the basis of this evidence, we would suggest that undue attention has been given to 

 the many unusual and prominent attributes of Erignathus [which may stem from an 

 accelerated rate of evolution, as has been postulated at the molecular level for Pagophilus 

 (Arnason et al. 1995)], at the expense of its many other similarities with the remaining 

 phocines. 



Within the Phocini proper, the paraphyly of a number of taxa is indicated. The new position 

 of Erignathus now also disrupts the monophyly of Phoca (sensu Burns & Fay 1970). As 

 mentioned previously, paraphyly of this taxon is not a new idea, but it is usually attributed 

 to an intrusion by Halichoerus [Chapskii 1955a; de Muizon 1982a; Arnason et al. 1995; 



