Scarcity of fossils is only one basis for the century-old controversy over whether the giant panda is really a bear or a "raccoon" (procyonid). Despite the gross anatomical similarity of giant panda to bears, there are more subtle anatomical and behavioral differences (Morris & Morris 1966; Ewer 1973). The panda bear has distinctive reproductive organs and an unusual scent gland, with which it exhibits unique marking behavior (Morris & Morris 1966). The panda's head and forepaws are specialized for its diet of primarily bamboo pith, a very coarse food. The broad zygomatic arches and high sagittal crest provide attachments for long, massive jaw muscles. (Sicher 1944). Unlike Ursinae and Tremarctinae, the panda bear retains the second and third premolars and sometimes the first. These premolars and the molars are highly developed for crushing and grinding food, being even broader than those of other bears--which, including the extinct Pleistocene Ursinae, have lost all but the last premolar. (Ewer 1973). Among other ursids, modification of the last premolar and the molars for herbivory reached their zenith in the cave bears of Europe and North America and in spectacled bear (Kurte'n 1976). Having observed captive or wild bears of all species, Herrero (pers. comm.), concluded that behavior of even the spectacled bear is more similar to Ursinae than either of those subfamilies are to panda. There are, however, sound reasons why some biologists suspect that the giant panda is more closely related to the red panda (Ailurus fulgens), an animal similar to an American raccoon (Procyon lotor). Ewer (1973) was particularly impressed by differences from typical bears in how pandas hold food (bamboo), whereas both giant panda and red panda hold it the same way. Their teeth (premolars and molars), skull and skull muscles (massiveness) are similarly adapted for feeding on bamboo. Thus, like Morris & Morris (1966) and numerous other biologists before them, Ewer (1973) concluded that giant panda and red panda are more closely related to one another than to other animals. This view has been bolstered by recent evidence on haemoglobin structure. The minimum mutation distance, in amino acid units, is 7 between the two pandas, 11 between giant panda and polar bear, and 18 between red panda and raccoon (Tagle et al. 1986).
Nevertheless, evidence that giant pandas are really bears and red pandas procyonids is overwhelming. This evidence involves anatomical details of the paws, skull, skull musculature, and teeth, as well as karyotypic and biochemical data.
There are numerous other anatomical reasons for concluding that the giant panda is a bear and the red panda a procyonid whose dependence on bamboo caused them to evolve convergently (Davis 1964). This analysis of comparative anatomy by Davis is considered without peer in our century (Gould 1986). Davis showed that the massive skull and jaw muscles and enlarged molars and premolars by both pandas are essentially what one would expect for adaptation to feeding on coarse vegetation--traits comparable to those of the spectacled bear, Florida cave bear, and European cave bear (see Kurten 1976). Furthermore, the red panda's dentition is only roughly similar to that of a panda bear; and the red panda lacks a pseudo-thumb, although the radial sesamoid is somewhat enlarged (Ewer 1973).
The convergence hypothesis is further supported by karyotypic and biochemical evidence. The 3 ursid subfamilies differ markedly in diploid chromosome number (74 Ursinae; 52 Tremarctos; 42 panda bear). But, 36 for red panda is comparable to the normal procyonid number of 38 (Wurster 1969; Thenius 1979). Reviewing data on genetics, biochemistry (Leone & Wiens 1956; Sarich 1973), morphology, and behavior, Thenius (1979:Fig. 7) concluded that the giant panda is a true bear and the red panda a procyonid, as argued by Pocock (1921). That is strongly bolstered by the additional studies on karyotype, DNA hybridization, isozyme genetic distance, and immunological distance by O'Brien et al. (1985). Mayr (1986) dismisses the haemoglobin evidence of giant panda-red panda linkage as just another phenotypic trait in which evolution of giant and red pandas converged due to living in the same habitat, eating mainly the same food. But how haemoglobin similarities between giant and red pandas would have been particularly adaptive to their location and diet remains unclear. Studies might fruitfully be done on the hemoglobins of other mammals in that region or which eat bamboo. In particular, the Asian black bear should be tested.
Mayr's hypothesis on convergent evolution of haemoglobin structure is unconfirmed. But rest of the evidence strongly favors the view that the giant panda is a true bear, assigned to subfamily Ailuropodinae.
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In this chapter, we have examined what is known about bear phylogeny, that is about which bears are ancestors or descendants of one another. We have also considered the geographic distribution of each species. Let us now examine in more detail the environmental factors which may have shaped ursid evolution and geographic radiation, and the ways in which bears met these challenges.