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Arboreality in Tree Squirrels (Sciuridae)

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Arboreality in Tree Squirrels (Sciuridae) ARBOREALITY IN TREE SQUIRRELS (SCIURIDAE) RICHARD W. THORIN.GTON, JR., AMY M.'L MILLER, AND CHARLES G. ANDERSON Department of Vertebrate Zoology, Smithsonian Institution, Washington, DC. 20560 (RWT) Department of Ecology, Evolution, and Marine Biology, University of California, Santa Barbara, CA 93106 (AMLM) Department of Zoology, University of Tennessee, Knoxville, TN 37916 (CGA) ABSTRACT.•The purpose of this paper is to examine the hypothesis that arboreality evolved more than once among extant tree squirrels. The North American fossil, Douglossia (formerly Protosciurus), from the late Eocene is considered to have been an arboreal squirrel (Emry and Thorington, 1982). Thus, arboreality may be primitive for squirrels, although the European Palaeoscinrus from the early Oligocene was probably terrestrial (Vianey-Liaud, 1974). It is un- clear how these are related to the Recent tree squirrels, the Sciurini of the Americas and Northern Eurasia, the Ratuflni and Callosciurini of Southern Asia, and the Protoxerini and Funambulini, mostly of Africa. Three hypotheses of relationships among Recent squirrels im- ply a terrestrial origin for some group of tree squirrels: Moore (1959) hypothesized that Sciwotamias and Tamiasciurus are closely related and that the North American red squirrel is derived from a terrestrial ancestor, Callahan and Davis (1982) hypothesized that Sciwotamias and Ratufa are closely related and suggested a terrestrial origin for the Asian giant tree squir- rels, and Moore (1959) also suggested that the African tree squirrels, Protoxerini and Funambulini, may have evolved from xerine ground squirrels. A review of morphological evi- dence suggests that Sciwotamias is more closely related to chipmunks, the Tamiina, than to either Tamiasciurus or Ratufa. This contradicts the first two hypotheses cited above and the suggestion that arboreality evolved independently in these two tree squirrels. We cite reasons for doubting the third hypothesis, that African tree squirrels have a terrestrial origin, but do not critically examine it in this paper. INTRODUCTION The squirrel family, Sciuridae, includes two (Black, 1963; Bruijn, et al., 1980). Thus, it is pos- radiations of ground squirrels (Marmotini, Xerini) sible that arboreality is primitive for squirrels and and a larger number of radiations of tree squirrels that extant terrestrial squirrels have evolved from in South America, North America, Eurasia, and tree squirrels once among the Marmotini of North Africa. Formerly, it was presumed that arboreal America and Eurasia, independently in the Xerini squirrels evolved from terrestrial squirrels. For of Africa and Southern Asia, and also indepen- example, Black (1963) considered the chipmunks dently in most tribes of tree squirrels. We here (tribe Tamiini) the basal group from which all other examine several hypotheses that arboreality has squirrels evolved. Similarly, Moore (1959) thought evolved more than once in the family Sciuridae. that the African tree squirrels were derived from Moore (1959) proposed that Sciurotamias, the the African ground squirrels. However, the earli- Chinese rock squirrel, and Tamiasciurus, the North est fossil squirrel, Douglossia jeffersoni formerly American red squirrel, are closely related, and that Protosciurus c.f. jeffersoni), from the late Eocene, the red squirrel evolved from a terrestrial rock 35 Ma, was a tree squirrel (Emry and Thorington, squirrel ancestor, similar to S. davidianus. 1982; Emry and Korth, 1996). The earliest chip- Callahan and Davis (1982) presented an alterna- munks date from the early Miocene, ca. 25 Ma tive hypothesis about Sciurotamias, arguing that In M.A. Steele, J. F. Merritt, and D. A. Zegers (eds.). 1998. 119 Ecology and Evolutionary Biology of Tree Squirrels. Special Publication, Virginia Museum of Natural History, 6: 320 pp V VIRGINIA MUSEUM OF NATURAL HISTORY it is closely related to Ratufa They explicitly sug- two proposed ground squirrel-tree squirrel hypoth- gested that Ratufa evolved from a terrestrial an- eses, each with an alternative hypothesis. cestor. Thus, the unresolved phylogeny of Sciurotamias, the Chinese rock squirrel, is central THE STATUS OF THE TAMIASCIURINI to the question of whether arboreality evolved more Moore (1959) created a tribe, the Tamiasciurini, than once among tree squirrels. for Tamiasciurus and Sciurotamias because both Moore (1959) also suggested that the African have three transbullar septa, a rare condition tree squirrels evolved from the xerine ground squir- among squirrels. Both Moore (1959) and Black rels. This hypothesis has not received careful con- (1963) noted that Tamiasciurus is polymorphic for sideration from subsequent investigators. Moore two and three transbullar septa. Corbet and Hill listed four characters shared by the Xerini and the (1992) list two transbullar septa for Sciurotamias Protoxerini. One of these is the separation of the forresti and three for Sciurotamias davidianus. buccinator and masticatory foramina. This condi- Thus, both genera are polymorphic for number of tion is primitive for rodents (Wahlert, 1991) and transbullar septa. Callahan and Davis (1982) therefore does not serve as a good phylogenetic showed that Sciurotamias lacks the specialized indicator. A second is the prominence of the mas- reproductive tract of Tamiasciurus. These obser- seteric tubercle, but this is found in only one ge- vations detract from the likelihood that Moore's nus in the Protoxerini. Therefore, it is likely that it hypothesis is a good one, but do not test it. The evolved independently in the Protoxerini and the hypothesis would be falsified by finding many de- Xerini. This leaves two characters, long orbits and rived features shared by Sciurus and Tamiasciurus, short interorbital width, purportedly supporting but not by Sciurotamias. It is not falsified by show- Moore's hypothesis. In details the orbits of the ing that proposed synapomorphies of Tamiasciurus Xerines and the Protoxerines are very different, and Sciurotamias are invalid. Bryant (1945) de- however, especially in the shapes of the lacrimal, scribed many osteological and myological similari- jugal, and maxillary bones. On the basis of cranial ties between Sciurus and Tamiasciurus, conclud- evidence, Lavocat (1973) considered the earliest ing that Tamiasciurus should not be considered a fossil squirrels in Africa to be tree squirrels de- distinct genus from Sciurus. He was very careful rived from southwest Asia. This also weakens to distinguish between primitive and derived char- Moore's speculation; but we will not critically ex- acters, but he did not study Sciurotamias. amine it in this paper. Similarity, the immunological study by Hight et In this contribution, we examine and test four al. (1974) and the protein electrophoresis study hypothesized sister groups: Tarniasciurus-Sciurus, by Hafner et al. (1994) suggested that Sciurus and Tamiasciurus-Sciurotamias, Tam.ias-Sciurotam.ias, Tamiasciurus are closely related, but neither study and Ratufa-Sciurotamias (Fig. 1), challenging the Sciurotamias Tamiasciurus Sciurus Sciurus Tamiasciurus Sciurotamias Ratufa Tamias Sciurotamias Tamias Ratufa Sciurotamias C/ D, Fig. 1.•Sister-group hypotheses examined: A. Commonly accepted hypothesis that Sciurus and Tamiasciurus are more closely related to each other than either is to Sciurotamias, B. Moore's (1959) hypothesis that Tamiasciurus and Sciurotamias are more closely related to each other, C. Milne-Edwards' (1871) hypothesis that Sciurotamias is closely related to chipmunks, and D. Callahan and Davis (1982) hypothesis that Sciurotamias is more closely related to Ratufa. 120 TH0BINGTONETAL_ARBOREAUTY|NTRE included Sciurotamias <=n «, ^ SO thev do nnt ^•.__J. 'HMtLb Moores hypothe^Tfn spi elf 1° T COntrad-t nesses of these "tests" of Mn ! l0gicaI Weak" concept of the Tal °f M°°res hypothesis, his £ts^ and Thorington P1982 oL SCiUrids (E•y ment f th doned (Hoffmann et a" f^ COmP^Y aban- Primitive state is listed as fof^f ° ^ states are listed as (la) and (ibi f S^ deriVed THE STATUS OF THE RATUFiMi - M1asc^^nus^-f the two genera arcloselyS5 ^ SUggGSted tha* be included in the tribe R^/r ^ Should b°th -any ways these t£ 1^^ ^ ****** In ferent squirrels. To^s fif* remarkably dif- UJNo septa: ifota/a (1954:479) showing the bac•hgUr^4 °f Prasad glans penis with rnany ridge"" °c "^ ^ the and noted ^^£2T^ C°^ cus, looks very different frnSr & VCntral sul" half transbullar seSa ThT," ^ ^ and one- gUre 3 ofCall e and Davis (1982-45) ^h ahan modal number for sXJsj!f - the primitive SciurotamicL X?^*° *«*» Penis of and COuId be two or three transbXslT^f'iar se Question the basis forfh ^ Thus' We derived. Pta. Lack of septa is Sciurotamias in the RatufiniTh P,laceme"t of Y St the chee Pouch and the hoof-Kketl ' as " k dence. However &L•S ^ Sporting evi- ^ngTon aSrT ^'^ »»"*. 8 P UCh Peri0 edge; porti «d Kata/a la^ teTuXmo * *"* ° of bulla anterior a^sZrior t i °n served that Rata/a's-hoonn;,?1"6, We have <*- Pollen: Scftinxs. TamiS ? thC meatus •t by a modified firs" £* - supported Rat br ad (1) Lacks flange^S ;/"*' "f«- terminal phalanx but th^ n * ° " flattened -Penor to it sw^tC^ ~ -nd not. Agai^ the* crit cTsm's ^ ^oto•- is The primitive conditinr, their h Pothesis, which has genSv h ' **' ^ unswollen bulla (WahlertZou ^dents is a" 9 11 Black s Corbet and Hill, 199| "X^"?*** (e.g., lustration of Pr0toJctimZ* ^ ' ^963) test it, we compare it with Th' ? &1' 1993^ T« & e and unswollen ^ZTerZeT^ ^ that ScturofamL is ^ * ^ hypothesis Scurotamios is consideredf
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