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DSA10 Heissig Kurt Heissig Bayerische Staatssammlung für Paläontologie und Geologie, München Origin and early dispersal of the squirrels and their relatives Heissig, K., 2003 - Origin and early dispersal of the squirrels and their relatives - in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 277-286 [ISSN 0923-9308] Published 1 December 2003 The first appearances of squirrels and aplodontids in both Europe and North America point to an unknown origin in between both continents. More recent analyses of the tectonic history of the North Atlantic and Arctic ocean basins are consistent with the hypothesis, that an exchange between both sides of the Atlantic has been possible at least until the early Oligocene, the time of origin of both groups. The long lasting isolation of Northern Europe from both the southern parts of the continent and Asia gives enough room for the assumption of a separate evolutionary centre. Correspondence: Kurt Heissig, Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Str. 10, D-80333 München, e-mail: [email protected] Keywords: Sciuridae, Aplodontidae, Paleobiogeography, North Atlantic, Paleogene INTRODUCTION known origin. To get an idea in which of the Squirrels (Sciuridae GRAY, 1821) are a empty spots on our paleobiogeographic map holarctic rodent group, appearing first in we should look for the origin of squirrels and Europe as immigrants of the Grande Coupure aplodontids, we have first to analyse the at the beginning of the Lower Oligocene relationships of both families to one another together with the probably related and with more or less similar isolated genera. Aplodontidae TROUESSART, 1897. In North America however, there appear related gene- THE DATA ra, partly with a protrogomorphous skull, partly with a superficial masseter halfway Characters and clades from the protrogomorphous to the sciuro- To get an idea of the interrelationship of the morphous or even a myomorphous structure. various genera that have been assigned to the All possible ancestors of squirrels or aplo- sciurid-aplodontid group I undertook several dontids in any Eocene fauna are too far from attempts of a character analysis, using the these groups to establish a near relationship. following taxa: Emry & Korth (1996) suggest "some Reithroparamys like ischyromyid" as an early As outgroup some rather primitive ancestor of both families. But most of the paramyines: Paramys delicatus LEIDY, 1873; shared dental characters could also be suspec- Paramys copei LOOMIS, 1907; and ted to be mere parallelisms within the vast Leptotomus leptodus (COPE, 1873). group of Ischyromyidae. As our knowledge of Eocene mammal faunas As species of probable but unknown is restricted to some well known areas, it is relation to squirrels or aplodontids: not astonishing to find immigrants without a Ailuravus macrurus WEITZEL, 1949 (A. picteti 277 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003 as a later member of the same lineage could Douglassia jeffersoni. These two species are be omitted); Mytonomys robustus (PETERSON, grouped nearly together in all obtained trees, 1919); Paracitellus cingulatus HEISSIG, 1979; sometimes forming a separate clade, some- P. marmoreus HEISSIG, 1979; P. eminens times forming a sistergroup of the true squir- DEHM, 1950; Cedromus wilsoni KORTH & rels. The last species is clearly set off from EMRY, 1991; and Douglassia jeffersoni all true squirrels by the separate entoconid (DOUGLASS, 1901). with a long labial spur (entolophid). So it may belong to some ancestral stock, separa- As representants of early aplodontids ted by a protrogomorphous masseter and this including Prosciurinae and Allomyinae: plesiomorphic tooth detail. The squirrel-like Spurimus scotti BLACK, 1971; Prosciurus skull and skeleton demonstrated by Emry & relictus (COPE, 1873); Campestrallomys Thorington (1982) may be due to convergence. dawsonae (MACDONALD, 1963); Campestral- The clade of the species of undefined lomys annectens KORTH, 1989; Campestral- affinities may form a sistergroup of one of lomys siouxensis KORTH, 1989; Ephemeromys the two other main clades, but these never hospes WANG & HEISSIG, 1984; and Plesi- can be grouped with one another. In the pure spermophilus angustidens FILHOL, 1882, as a pattern cladistic cladogram this group is atta- primitive Allomyinae. ched to the aplodontid clade and the squirrels are isolated, without the sistergroup of As earliest representants of true squirrels: Cedromus wilsoni. If the characters are Palaeosciurus goti VIANEY-LIAUD, 1974; weighted in three levels and the form of the Oligopetes radialis HEISSIG, 1979; and protoconule, the anterior connection of the Oligopetes lophulus HEISSIG, 1979. mesostyle and the reduction of the labial crest of the entoconid are considered to be irrever- These 20 species have been analysed accor- sible, this clade of doubtful species appears ding to 27 dental characters with different as the sistergroup of the squirrel clade. In this cladistic methods. The results yielded, if any, case the pair of Mytonomys and Douglassia is very similar pictures. Generally there is an excluded from the relationship of squirrels. outgroup comprising Leptotomus leptodus, With all these manipulations it was impos- Paramys copei and Spurimus scotti. So this sible to rearrange the tree into two main cla- last species cannot be comprised within the des, the intermediate forms being sorted in Aplodontidae and this family therefore arri- both of them. This result has consequences ves in North America not before the begin- for the formation of a biogeographical hypo- ning of the Oligocene. The arrangement of thesis. The results of the analysis can not be these three species is varying according to the summarized in one or more cladograms, first weighting of different characters. because of the subjectivity of the character There are three main clades of rather con- selection, second because of the unknown stant composition. One comprises the three degree of character variability in most invol- species of true squirrels, sometimes with ved species. Even characters traditionally Cedromus wilsoni as a sistergroup at the used are highly variable in some species. On base, a result obtained also by Korth & Emry the other hand the interior configuration of (1991: 993) on the basis of skull characters. the three main clades is rather stable against The second one comprises the early manipulations, so that there may be some Aplodontidae, including Campestrallomys reason to accept them as real units. So the annectens in a rather basal position. The third idea of any closer relationship between squir- one comprises the group of unclear affinities, rels and aplodontids depends on the subjecti- in most solutions including Paramys delica- ve high value of the elongation of the proto- tus, but excepting Mytonomys robustus and cone at the cost of the hypocone. 278 HEISSIG: Origin and dispersal of squirrels The paleobiogeographical distribu- Campestrallomys dawsonae and C. sioux- tion ensis, which show up always within this pattern clade and not with Campestrallomys annec- There is only one species in the Eocene of tens, appear not earlier than in the middle or Europe which shows undoubted affinities to late Oligocene. one of the three main clades: Ailuravus macrurus from Messel. Mytonomys from The paleogeographic situation of North America, which was considered by possible land bridges Wood (1976) and Korth (1984) to be a mem- According to the rapidly decreasing faunal ber of the Ailuravinae, never shows a group- similarity of North America and Europe from ing together with this species but is always the base of Middle Eocene onwards, the ope- near the protrogomorphous but otherwise ning of the Atlantic was assumed by paleon- squirrel-like Douglassia jeffersoni. On the tologists to begin at that date (e.g. Simpson other hand Ailuravus macrurus is by far too 1947, Kurtén 1966, McKenna 1972, big to be ancestral to any of the Oligocene Springhorn 1984). More recent tectonic species. investigations have revealed, however, that The bulk of the Aplodontidae and Sciuridae the earliest spreading zone extended west of appear without doubt at or near the base of Greenland before the spreading activity was the Oligocene on both continents. The sciu- restricted to the eastern branch of the Atlantic rids are restricted to Europe in this first time, around 36 Ma BP (Eldholm 1990). The final but members of both subfamilies, Sciurinae separation of the Barents shelf from the nort- and Petauristinae occur probably at the same hern edge of Greenland must therefore have time with the Grand Coupure. Members of occurred some time later, probably during the the other clades appear on both sides of the Oligocene (Fig. 1). This assumption is corro- Atlantic. As there is still some uncertainty borated by the transpressive folding of where the Eocene-Oligocene boundary has to Eocene strata on Spitsbergen (Geyssant & be placed within the Chadronian of North Lepvrier 1982), indication of a direct neigh- America, the results remain a little doubtful. bourhood of Greenland during most of the There is no comparable immigration event as Eocene. in Europe. The undoubted Aplodontidae On the other hand, there was a broad nearly appear in Europe with the subfamilies continuous epicontinental seaway along the Prosciurinae (Ephemeromys) and Allomyinae Polish lowlands from the Middle Eocene till (Plesispermophilus
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