Journal of Vertebrate Paleontology

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Journal of Vertebrate Paleontology 21(1):119-131. 2001
doi: 10.1671/0272-4634(2001)021[0119:SAOPCM]2.0.CO;2

STRATOCLADISTIC ANALYSIS OF PALEOCENE CARPOLESTIDAE (MAMMALIA, PLESIADAPIFORMES) WITH DESCRIPTION OF A NEW LATE TIFFANIAN GENUS

JONATHAN I. BLOCHa, DANIEL C. FISHER, KENNETH D. ROSE, and PHILIP D. GINGERICH

aDepartment of Geological Sciences and Museum of Paleontology, University of Michigan, Ann Arbor, Michigan 48109-1079

bDepartment of Cell Biology and Anatomy, Johns Hopkins University School of Medicine, Baltimore, Maryland 21205

Abstract

“Carpodaptes” jepseni is a morphologic intermediate between Carpodaptes and Carpolestes, with the number and position of cusps on p4 more consistent with placement in Carpodaptes but relative size of p4 more like Carpolestes. The type and only previously known specimen of “C.” jepseni, a partial dentary with p4–m2, is from Divide Quarry (Tiffanian Land-Mammal Age) in the Fort Union Formation of the Bighorn Basin, Wyoming. New specimens of “C.” jepseni from Divide Quarry include a nearly complete dentary with p4–m3 and alveoli for all anterior teeth, and the first known upper dentitions with P1–M2 and an alveolus for C1. Specimens of Carpodaptes cygneus are also described from Divide Quarry, demonstrating the occurrence of two distinct carpolestid species at the same locality.

Stratocladistic analysis of the thirteen known carpolestid species, using thirty-two morphologic characters and stratigraphic order, produced eight most-parsimonious phylogenetic trees associated with a single cladogram. The topology of the cladogram generated using stratocladistics is identical to that of the single most-parsimonious cladogram from cladistic analysis of the same morphologic data, but stratocladistics allows greater resolution than cladistics at the level of phylogenetic trees. New specimens demonstrate extreme shortening of the anterior jaw of C. jepseni, a derived state not present in other carpolestids. This suggests that C. jepseni occupies a side-branch close to, but not at, the ancestry of the Carpolestes clade (an ancestor-descendant lineage composed of the sequence C. dubius, C. nigridens, and C. simpsoni, in that order). “C.” jepseni is here placed in a new genus, Carpomegodon.

Received: February 13, 2000; Accepted: August 25, 2000



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APPENDIX 1


Description of characters used in phylogenetic analyses. Outgroup taxa were Purgatorius and Pronothodectes. The cladistic analysis used Purgatorius to root the preferred cladogram. All characters except character 33 (the stratigraphic character, not used in the cladistic analysis) were unordered. Those characters that were variable within a species were coded as polymorphic. Autapomorphic characters (characters 1, 5, and 15) were included in the stratocladistic analysis, as they can affect hypotheses of ancestry; cladistic results are reported with and without them.

Mandible anterior to p4—not foreshortened, with substantial to moderate space both anterior and posterior to the mental foramen (0), or foreshortened, with barely enough room for a mental foramen between the anterior root of p4 and the alveolus of i1 (1).

i1—subequal in size to c1 and not strongly procumbent (0), or enlarged relative to c1 and strongly procumbent (1).

i1 crown—short and lanceolate (0), moderately long and lanceolate (1), or long and slender (2).

Basal cusp on lingual cingulum of i1—absent (0), or present (1).

i2—present and aligned with other anterior teeth (0), or vestigial and displaced buccally in relation to other anterior teeth (1).

i3—present (0), or absent (1).

c1—larger than i2 (0), slightly smaller than i2 (1), or distinctly smaller than i2 (2).

c1 anterior projection—weak (0), or strong and touching the crown of i2 (1).

p2—present with two roots (0), present with one root (1), or absent (2).

p3—subequal in size relative to p4 (0), reduced in size relative to p4 (1), or absent (2).

p3—double-rooted, occupying two alveoli (0), or with partially or wholly fused roots, occupying a single alveolus (1).

Crown of p3—premolariform and unreduced(0), or button-like and reduced (1).

Crown of p4—premolariform, not hypertrophied and not exodaenodont (0), plagiaulacoid, hypertrophied and exodaenodont (1).

Metaconid on p4—absent (0), present and posterolingual to protoconid (1), or present and in line with other cusp(s) in the trigonid (2).

Paraconid on p4—present (0), or absent (1).

Number of accessory apical cusps on p4—none (0), one which, together with the three trigonid cusps, forms a blade with four longitudinally arranged cusps (1), two which, together with the three trigonid cusps, form a blade with five longitudinally arranged cusps (2), three which, together with the three trigonid cusps, form a blade with six longitudinally arranged cusps (3), four which, together with the three trigonid cusps, form a blade with seven longitudinally arranged cusps (4), or five to six which, together with the three trigonid cusps, form a blade with eight to nine longitudinally arranged cusps (5).

Labial height of p4 relative to that of m1—low (0), intermediate (1) or high (2).

Outline of plagiaulacoid p4 in labial view—gently rounded (0), somewhat pointed (1), or very pointed (2).

Posterior apical cusp on crest uniting main shearing blade with talonid cusp of plagiaulacoid p4—near penultimate apical cusp (0), or roughly equidistant between penultimate apical cusp and talonid cusp (1).

Posterior apical cusp of plagiaulacoid p4—cuspate and above a strong vertical rib on lingual face (0), or weak, with slight to no expression of a vertical rib on lingual face(1).

Lower molars—not exoedaenodont (0), or somewhat exoedaenodont (1).

Trigonid of m1—triangular, with paraconid in a lingual position (0), subtriangular, with paraconid in a more anterior position (1), anteroposteriorly aligned, with the paraconid directly anterior to the protoconid (2), or anteroposteriorly aligned, with the paraconid directly anterior to the protoconid and trigonid more elongate anteroposteriorly (3).

P3—smaller than P4 (0), subequal to or slightly larger than P4 (1), or much larger than P4 (2).

P3 protocone—a single, distinct cusp (0), a single weak cusp (1), or accompanied by a hypocone to form two lingual cusps (2).

Lingual basin on P3—absent (0), weakly present (1), well developed (2), or well developed and anteroposteriorly enlarged (3).

Number of median crests on P3—None (0), one, represented by pre- and postconule cristae (1), two or three, represented by pre- and postconule cristae and one or two more posterolingual crests not continuous across the tooth crown (2).

Buccal side of P3—with two cusps, a major cusp followed by a smaller posterior cusp (0), three cusps with anterior cusp separated somewhat from two posterior cusps (1), four cusps with an anterior cusp separated somewhat from three posterior cusps (2), five cusps with an anterior cusp separated somewhat from three posterior cusps and an anterior fifth cuspule forming a short anteroexternal spur (3), or five cusps with anterior one fully as developed as others, resulting in an elongate anteroexternal spur (4).

Occlusal view of P4—triangular (0), subtrapezoidal (1), trapezoidal (2), or square (3).

Number of cusps on buccal row of P4—two (0), four (1), or five to six (2).

Number of median crests on P4—none (0), one with a distinct central conule (1), two, with the more buccal crest showing a distinct central conule and the more lingual crest represented by pre- and postconule cristae (2), or two, with as many as five nearly equal cusps on the more buccal crest and the more lingual crest represented by pre- and postconule cristae (3).

Number of lingual cusps on P4—one (0), or three (1).

Preprotocrista on P4—present (0), or absent (1).

Stratigraphic Range—Puercan (0), Torrejonian (1), Tiffanian1 (2), Tiffanian2 (3), Tiffanian3 (4), Tiffanian4 (5), Tiffanian5 (6), Tiffanian6 to Clarkforkian1 (7), Clarkforkian2–3 (8).



APPENDIX 2


table

Character matrix used for phylogenetic analyses. Characters that are either not known for a particular taxon, due to incomplete fossil material, or that are not applicable, are treated as missing (“?”)

FIGURE 1. A, Locality map and B, stratigraphic section of the northern Bighorn Basin with Paleocene quarries indicated. MQ, Mantua Quarry; RBQ, Rock Bench Quarry; CPQ, Cedar Point Quarry; DQ, Divide Quarry; PQ, Princeton Quarry. Figures modified from Bartels (1987). Biostratigraphic zonation after Gingerich (1983, 2000), Archibald et al. (1987), Williamson (1996), and Eberle and Lillegraven (1998)

FIGURE 2. A, Lingual; B, buccal; and C, occlusal views of the right dentary of Carpomegodon jepseni (UM 80575) with p4–m3 and the alveoli of all anterior teeth. Note that p4 is very large and high crowned and that there are six distinct apical cusps on p4, the first five closely spaced and the last one set farther back midway between the fifth cusp and the talonid heel. An additional weak cuspule is situated between the fifth and sixth cusp of p4. Scale in mm

FIGURE 3. Enlarged view of the dentary of Carpomegodon jepseni (UM 80575) anterior to p4. Note that the anterior part of the dentary is very short, with two small alveoli between p4 and a relatively large alveolus for the medial incisor. A very small excavation at the posterolateral margin of the large alveolus probably housed a vestigial tooth. We interpret these alveoli to have held i1–2, c1, and p3; hence the lower dental formula of Carpomegodon jepseni is 2.1.2.3. Scale in mm

FIGURE 4. Upper dentition of Carpomegodon jepseni from Divide Quarry in occlusal view: A, P1(C1?)-M2 (UM 85918) and B, P3–M2 (UM 86241). Note the presence of three alveoli anterior to P3 in the maxilla, with no premaxilla-maxilla suture evident around the anteriormost alveolus. P1(C1?) and P2 are single-rooted. Scale in mm

FIGURE 5. Enlarged view of the maxilla (UM 86241) anterior to P3. Note the presence of three alveoli anterior to P3. We interpret these alveoli to have held P2, P1, and C1. Scale in mm

FIGURE 6. Camera lucida drawings of carpolestid p4s. A–B, 3 Carpolestes and 1 Carpomegodon species and C–D, 4 Carpodaptes species. Outlines are in buccal view with anterior to the right. Shading represents interpretations of p4 outlines with ranges of species variation indicated. The p4s were oriented about a vertical line passing through the highest apical cusp and the ventral notch and a perpendicular horizontal line that touches the base of the posterior lobe. Note that the p4 of Carpomegodon jepseni, while mostly higher crowned than that of any of the other species, overlaps somewhat with Carpolestes dubius in height and length. The p4 profile of Carpomegodon jepseni is very distinct in size from that of any of the Carpodaptes species. Note, however, that p4 of Carpodaptes aulacodon is similar in shape to that of C. jepseni

FIGURE 7. Plots of carpolestid p4 size. A, p4 height as a function of m1 height; B, p4 length as a function of m1 length; and C, p4 height as a function of p4 length. The points represent mean sample measurements (in mm). Dashed parallel lines are isoclines of constant difference. Solid line is a least-squares regression through the mean sample measurements. Note that the p4 of Carpomegodon jepseni is higher relative to its own length, than is the case for other carpolestids. The length of the Carpomegodon p4 relative to that of its m1 is not larger than for other carpolestids, with the exception of Elphidotarsius shotgunensis and Elphidotarsius florencae, which seem to deviate from the relationship demonstrated in other species of carpolestids in having a shorter p4 relative to m1 length

FIGURE 8. A, occlusal views of upper dentitions for Carpolestes dubius (YPM-PU 19349); B, Carpomegodon jepseni (UM 86241) and C, Carpodaptes hazelae (AMNH 33980, type). Note that Carpomegodon jepseni is similar to Carpolestes dubius, and differs from Carpodaptes hazelae, in having an anteroexternal extension on P3, and in having P4 smaller than P3. Scale in mm

FIGURE 9. A, buccal view of right dentary (UM 80669); B, buccal view of left dentary (UM 85286); and C, occlusal view of left maxilla (UM 91324) of Carpodaptes cygneus from Divide Quarry. The i1 is similar to those illustrated by Krause (1978) from the Roche Percée local fauna, Saskatchewan, in having a crown that is long and slender, laterally compressed, and apically tapering. Scale in mm

FIGURE 10. Hypotheses of phylogenetic relationship among North American carpolestids based on dental characters (Appendices 1 and 2). A, cladistic analysis yielded a single most-parsimonious cladogram generated by a branch-and-bound algorithm (Swofford, 1993) and rooted with Purgatorius: tree length = 65 (68, with autapomorphies), consistency index = 0.92 (0.93, with autapomorphies), retention index = 0.94. All characters were unordered. Unambiguous synapomorphies supporting each node are as follows (change is from 0 to 1 for binary characters; state indicated for multistate characters) node 1–2, 6, 7(1), 9(1), 30(1); node 2 (Carpolestidae)-10(1), 11, 13, 14(1), 16(1), 21, 24(1), 25(1), 26(1), 28(1), 29(1); node 3–3(1), 22(1), 23(1), 25(2), 27(1); node 4–24(2); node 5–14(2); node 6–9(2), 12, 16(2), 17(1), 22(2). 27(2), 28(2), 29(2), 30(2), 31, 32; node 7–19(1); node 8–27(3); node 9–20; node 10–18(2), 22(3); node 11–17(2), 23(2), 25(3); node 12 (Carpolestes)-3(2), 7(2), 8, 16(5), 18(1). 20, 28(3). B, stratocladistic analysis yielded eight overall most-parsimonious trees associated with a single topology, identical to the most-parsimonious cladogram generated by PAUP. Terminal taxa are shown in bold type with heavy lines, while ancestral taxa are shown in light type with arrows pointing to their correct position on the tree. E. shotgunensis, E. russelli, and C. cygneus are equivocal in their placement as terminal taxa or at ancestral nodes

FIGURE 11. A, stratigraphic occurrence and B, ranges for the thirteen carpolestids. Purgatorius was used to root the trees in the cladistic analysis and Pronothodectes was included as an outgroup taxon. Stratigraphic position of samples is shown at the greatest resolution allowed by interbasinal correlations. Samples known from a faunal zone are plotted (arbitrarily) as being from the middle of that zone. In the plot of stratigraphic ranges (B) the stratigraphic character is coded in 9 states, all of which are subdivisions of the Paleocene. The first occurrence of Elphidotarsius wightoni is in either Ti-1 or Ti-2 (Fox, 1990). To allow for the possibility that E. wightoni is known from Ti-1, it was coded as occurring in that interval. Resolution of this issue could affect the conclusions of this analysis, as a later first occurrence of E. wightoni would provide support for different hypotheses of ancestry. Stratigraphic occurrence of C. oriens is here plotted as mid-to-late Clarkforkian, but it is possibly early Eocene in age (see text). Resolution of this issue will not affect the results of this analysis

FIGURE 12. One of the phylogenetic trees (with equivocal ancestors as sisters only) resulting from the stratocladistic analysis (Fig. 10B). Divergences of sister taxa are shown schematically, with no implied knowledge of the exact timing of cladogenesis. Note that Carpomegodon jepseni (Ti-4) is the sister taxon to the clade that includes Carpolestes dubius, C. nigridens, and C. simpsoni, which represent an ancestor-descendant lineage spanning the late Tiffanian (Ti-5) through Clarkforkian NALMAs

table

TABLE 1. Summary of dental measurements of Carpomegodon jepseni from Ti-4 Divide Quarry. Total sample includes 12 specimens. Abbreviations: N, samples size; , mean; s, standard deviation; V, coefficient of variation; L, crown length; W, crown width; and MD, mandibular depth. Measurements in mm

table

TABLE 2. Summary of dental measurements of Carpodaptes cygneus from Ti-4 Divide Quarry. Total sample includes 8 specimens. Abbreviations as in Table 1. Measurements in mm

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