9 



1995; Mouchaty et al. 1995; Perry et al. 1995). However, a primitive or ancestral status 

 for either Pagophilus or Pusa within the Phocini has been suggested by some authors 

 [Burns & Fay (1970) and Shaughnessy & Fay (1977), and McLaren (1966, 1975) 

 respectively]. 



Altogether, these systematic difficulties within the Phocini likely stem from the relatively 

 recent major radiation of the group [latest Miocene (Arnason et al. 1995) or post-early 

 Pliocene and/or Pleistocene (Ray 1976a)], so that its members are not clearly differentiated 

 from one another. This is especially true for Phoca vitulina, a species that has been 

 described as being in the midst of a rapidly evolving "species swarm" (Ray 1976a: 402). 

 Individuals of this species from the Atlantic and Pacific Oceans are readily distinguishable 

 from one another (Allen 1902; Doutt 1942; Chapskii 1955a, 1967; Davies 1958b; Arnason 

 et al. 1995), but there is much debate as to the exact subspecific make-up of the Pacific 

 subgroup [for summaries, see Shaughnessy & Fay (1977) or Bigg (1981)]. This impacts 

 here primarily on the larga seal, a taxon distinguishable from other Pacific P. vitulina 

 based on geographical, ecological, behavioural, and morphological grounds (Shaughnessy 

 1975), but of uncertain taxonomic status. It has variously been regarded as an unnatural, 

 "garbage" taxon (Allen 1902), as a subspecies of P. vitulina (Scheffer 1958; Burns 1970; 

 Shaughnessy 1975; Baram et al. 1991), as the species Phoca largha (Chapskii 1955a, 

 1967; McLaren 1966, 1975; Shaughnessy & Fay 1977; Bigg 1981; King 1983; Arnason 

 et al. 1995), or of uncertain status (Allen 1880). As well, the population boundaries of the 

 larga seal are highly contentious, ranging from encompassing all Pacific harbour seals 

 (Chapskii 1955a), to only those inhabiting the western Cis-Asiatic region (roughly from 

 the Chukchi Sea to the coast of China) (Scheffer 1958; Chapskii 1967), to only those in 

 this latter region that breed on the pack ice (Shaughnessy & Fay 1977). For our purposes, 

 we will accept the larga seal as the species Phoca largha [species description given in 

 Chapskii (1967)] in order to establish its systematic relationship with Phoca vitulina. As 

 well, we will recognize this species as inhabiting the entire western Cis-Asiatic region, a 

 distribution that has become increasingly accepted. 



The systematic status of the Monachinae 



Since its inception, membership of the subfamily Monachinae has fluctuated from 

 including only Monachus spp., to its present status of encompassing all southern 

 hemisphere seals (lobodontines plus Mirounga spp.) plus Monachus spp. This instability 

 appears to be due simply to an increasing refinement of phocid taxonomy with time, 

 although it may relate to the suggestion of Wyss (1988a) that the subfamily is paraphyletic. 

 This novel suggestion is apparently connected with the paraphyly of Monachus, and with 

 the recognition of M. schauinslandi as the sister taxon of the remaining phocids in 

 particular (Berta & Wyss 1994; see above). Although a strongly divergent adaptive 

 radiation has been noted for the monachines (Ray 1976b), paraphyly of the subfamily 

 would contradict a number of apparent synapomorphies, particularly among postcranial 

 elements (see King 1966; Hendey & Repenning 1972; de Muizon 1982a), which would 

 have to be re-interpreted as phocid symplesiomorphies. It also clashes with biomolecular 

 evidence (Sarich 1975, 1976), and the finding that the monachines and phocines are 

 equally ancient lineages with, distinct representatives of each being found among the first 



