Notvtates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

AMERICAN MUSEUM Notvtates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2762, pp. 1-16, figs. 1-9, table 1 May31, 1983

A Revision of the European Eocene Primate Genus Protoadapis and Some Allied Forms

IAN TATTERSALL' AND JEFFREY H. SCHWARTZ2

ABSTRACT The European Eocene primate genus Protoad- Pronycticebus, is represented by material formerly apis, as generally conceived of, is long-lived and allocated to "Protoadapis" (or Europolemur) highly speciose, but nonetheless homogeneous. klatti, as well as by Pronycticebus gaudryi and by Reappraisal of the material involved shows, how- a new species we describe here. Pronycticebus may ever, that among the 10 species allocated by Gin- possibly bear affinities with certain Pelycodus. The gerich (1977) to Protoadapis and its closest rela- third major morph in the assemblage is comprised tives Cercamonius and Pronycticebus, three of specimens allocated to "Protoadapis" angusti- separate groups are discernible, together with three dens (now filholi), "Protoadapis" (or Cercamoni- individual species that are clearly misattributed. us) brachyrhynchus, and possibly "Protoadapis" Within the genus Protoadapis we recognize for weigelti; the earliest available name for the single the moment two species, P. curvicuspidens and P. species represented by this material is Cercamoni- recticuspidens, both originally described (as us brachyrhynchus, and its affinities lie with the species of Plesiadapis) by Lemoine in 1878. These larger species of Notharctus. The species "Pro- two differ somewhat in trigonid morphology, but toadapis" ulmensis, based on isolated teeth, ap- both are quite distinct from all other material sub- pears to be related to Adapis, the genus in which sequently referred to Protoadapis. As thus consti- it was originally described, and the affinities ofthe tuted, Protoadapis may be most closely compa- two species "Protoadapis" russelli and "P." louisi rable to Agerinia. A second genus, by priority are unclear. INTRODUCTION Victor Lemoine, the French physician and by Lemoine. Lemoine attributed the first paleontologist, described in 1878 a collection group of material (from Cernay) to the type of teeth and fragmentary jaws, from various species ofthe genus, Plesiadapis tricuspidens; early Tertiary deposits near Reims, as be- the rest, consisting of younger specimens longing to four species of the primate genus from near Epemay (probably Ay), he divided Plesiadapis, which Paul Gervais had estab- among Plesiadapis curvicuspidens, P. crassi- lished the year before on specimens furnished cuspidens, and P. recticuspidens, listed in that

' Curator, Department of Anthropology, American Museum of Natural History. 2 Research Associate, Department of Anthropology, American Museum of Natural History; Associate Professor, Department of Anthropology, University of Pittsburgh.

Copyright © American Museum of Natural History 1983 ISSN 0003-0082 / Price $1.75 2 AMERICAN MUSEUM NOVITATES NO. 2762 order. Two years later Lemoine (1880) trans- Ulm in southern Germany. In 1977 he transferred these last three species to his new genus ferred this species to Protoadapis, an action Protoadapis, listing them in similar order af- shortly endorsed by Gingerich (1977) and ter his (apparently Paleocene) species Pro- Szalay and Delson (1979). More recently yet, toadapis copei, a form he neither described Gingerich (1975) removed P. brachyrhyn- nor illustrated, and whose material basis is chus from Protoadapis to its own genus, Cer- uncertain. A decade later, in a review of the camonius, but two years later created three entire assemblage offossils he had recovered new species ofProtoadapis: weigelti from the from the neighborhood of Reims, Lemoine Geiseltal, and russelli and louisi from the (189 1) reduced to two the number of species French earliest Eocene site of Avenay (Gin- he recognized within Protoadapis: P. curvi- gerich, 1977). He also proposed the new cuspidens and P. recticuspidens. He empha- name filholi to replace Filhol's angustidens, sized the resemblances of these to Plesiada- on the grounds that angustidens had origi- pis, and regarded both genera as "formes plus nally been described as a species of Adapis, lemuriennes" than other genera included in and that the species name was therefore his fossil collection from the area of Reims. preoccupied by the trinomen Adapis pari- Despite the rapid exit of the species Pro- siensis angustidens that had been created for toadapis copei and P. crassicuspidens, and the other material by Filhol himselfin 1883. Sza- fact that most recent authors have regarded lay and Delson (1979) recently returned P. curvicuspidens and P. recticuspidens as brachyrhynchus to Protoadapis, while reject- synonymous, the species content of Protoad- ing Gingerich's three new species (but not the apis has nonetheless expanded enormously new name, filholi); they further reinstated since Lemoine's time. For example, Ginger- Weigelt's genus Europolemur. ich's (1977) recent summary list ofEuropean From this brief taxonomic history it is adapid species contains eight species of Pro- quite evident that, while agreement among toadapis, almost a third ofthat author's total recent authorities is far from complete, Pro- for the family. toadapis would on the face of it appear to be Stehlin (1912) was the first paleontologist not only the longest-lived (earliest Eocene to after Lemoine to contribute a new species to the latest Eocene) of the primates of the Eu- Protoadapis, in describing as Protoadapis (his ropean Paleogene, but also by far the most Protadapis) brachyrhynchus a mandibular prolific of species. Among other potential specimen from the French late Eocene Quer- contenders Adapis has been the only other cy locality of Prajous. Teilhard de Chardin serious claimant for this distinction. Ginger- (1922) subsequently added Filhol's (1888) ich (1977), for instance, lists eight species of "Adapis" angustidens, represented by an- the genus. However, Gingerich's list of Ada- other dentary from the Quercy phospho- pis species contains a heterogeneous assort- rites, to the roster ofProtoadapis species; and ment of taxa of diverse affinities, and the as- in 1932 Cooper described Protoadapis eppsi sessment by Szalay and Delson (1979) ofthe on the basis of a partial mandible and other genus as monotypic, containing only the material recovered at Abbey Wood, in the species A. parisiensis, seems more reason- English earliest Eocene. Simons (1962) trans- able. Similarly, our recent reappraisal of the ferred eppsi to its own new genus, Cantius material that has at one time or another been (whence some authorities would nowadays assigned to the genus Protoadapis has led us remove it to Pelycodus); but at the same time to the conclusion that over the years Pro- he synonymized Weigelt's (1933) Europole- toadapis, like Adapis, has become a "catch- mur klatti (and its own synonym Megatarsius all" taxon that casts a distorting shadow of abeli) with Protoadapis, as P. klatti. Weigelt's illusory uniformity over what is, in fact, material came from the middle Eocene brown another diverse assemblage of taxa. We pre- coal deposits ofthe Geiseltal, near Halle, east sent our conclusions in detail below; and, since Germany (DDR). sweeping systematic statements about the In 1971 Schmidt-Kittler described a new fossils at issue have been made regularly over species of Adapis, A. ulmensis, from the late the past century without much more than Eocene fissure fill deposits ofEhrenstein near passing reference to the actual morphologies 1983 TATTERSALL AND SCHWARTZ: PROTOADAPIS 3 involved, we preface them with descriptions gually emplaced. In both preserved molars of the material. the metaconid and protoconid flow together at their bases, and in general the cusps seem to have been rather puffy. ACKNOWLEDGMENTS Two referred dentaries also seem to belong We thank Drs. Burkart Engesser and Jo- to this species. AL-5180 (fig. 1) is a right hannes Hiirzeler ofthe BNM, Dr. D. E. Rus- partial mandible with three premolars and sell of the MNHN, Dr. Hartmut Haubold of the first two molars, all very heavily worn. the GPIH, and Drs. P. M. Vandermeersch The most anterior premolar is small, single- and Wim Van Neer ofthe PLV, for their help rooted, and premolariform. That behind it is and for their permission to study material in a rather tall premolariform tooth, with a their care. We are also grateful to Dr. Todd strongly truncated talonid. The last premolar Olson for his comments on an earlier draft is more submolariform, with a small para- of the manuscript, and to Ms. Joan Whelan conid and a larger metaconid. Both cusps lie for the scanning electron micrographs. Sup- low on the lingual face of the tooth. The last port for this work was provided through the two premolars as well as both molars possess generosity of the Richard Lounsbery Foun- strong, quite complete buccal cingulids. Ml dation. This is contribution no. 4 of the has a small but distinct paraconid that was Lounsbery Laboratory, Department of An- closely approximated to the metaconid; in thropology, American Museum of Natural M2 this small cusp is more medially placed. History. The entoconid on M2 would have been small and apparently rather conical. A referred left ABBREVIATIONS dentary, FSL 1991, contains the last pre- BNM, Naturhistorisches Museum, Basel molar and MI, and is a shade less worn than CMNH, Carnegie Museum of Natural History, the other specimens. The preserved teeth are Pittsburgh similar to their counterparts in AL-5 180; in FSL, Faculte des Sciences, Universite de Lyon M1 it appears that the cristid obliqua did not GPIH, Geiseltalmuseum (formerly Geologisches- meet the metaconid, and that the paraconid Palaontologisches Institut), Martin-Luther- was separated from the paracristid by a Universitait, Halle-Wittenberg, DDR groove (as in some large species ofNotharctus MNHN, Institut de Paleontologie, Museum Na- which, however, possess narrower talonids). tional d'Histoire Naturelle, Paris The holotype ofProtoadapis recticuspidens PLV, Instituut voor Aardwetenschappen, Katho- is MNHN AL-5 182, a right dentary fragment lieke Universiteit te Leuven, Belgium containing M2-3 (fig. 1). This specimen is highly worn, and only M3 preserves any sig- MORPHOLOGY nificant morphology. The trigonid of this The type of Protoadapis curvicuspidens is tooth is extremely compressed, and lacks a MNHN AL-5 179 (fig. 1), a right partial den- paraconid. The low paracristid arcs between tary with M2-3, very greatly worn. The para- the protoconid and metaconid; the cristid ob- conid shelf on both lower molars preserved liqua is very buccally situated and terminates is quite broad; that on M3 is situated more at the buccal side of the base of the proto- inferiorly than that on M2. On both lower conid. The bases ofthe protoconid and meta- molars the cristid obliqua terminates at the conid are confluent and form a steep wall base ofthe protoconid; the buccal cingulid is facing the talonid. The talonid cusps are rel- complete on M2 and partial on M3. The tal- atively indistinct and peripheral, and the talonid seems to have been low, and the basin onid heel is rather short and lingually em- shallow, at least on M2; and the talonid is placed. Two other dentaries, one with an only slightly broader than the trigonid. A excellent dentition, have been referred to this small enamel swelling in the middle of the species, with which in fact they do not appear paraconid shelfmay suggest a tiny paraconid, to belong (see below). especially on M2; there is a similar swelling The type specimen of Filhol's "Adapis" at the buccal termination of the shelf. M3 angustidens (Protoadapisfilholi ofGingerich, bears a moderately elongate heel, quite lin- 1977) is a left dentary (fig. 2) that has been 4 AMERICAN MUSEUM NOVITATES NO. 2762

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FIG. 1. Scanning electron micrographs of casts of: (left) MNHN AL-5 179, type of Protoadapis curvicuspidens; (center) MNHN AL-5182, type of Protoadapis recticuspidens; (right) MNHN AL-5180, referred to Protoadapis curvicuspidens. All scales represent 1 mm. lost since it was redescribed, illustrated, and containing M2-3, exists, however, in the col- assigned to Protoadapis by Teilhard de Char- lections of the Katholieke Universiteit te din (1922). Another Quercy specimen prop- Leuven, as PLV-35 (fig. 3). The "puffy" mo- erly referable to the species, a left dentary lars of this specimen have a small paraconid 1983 TATTERSALL AND SCHWARTZ: PROTOADAPIS 5

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FIG. 2. Above: occlusal and lateral views, from Stehlin (1912), of BNM QV 619, type of "Pro- toadapis" brachyrhynchus. Below: occlusal and internal views, from Teilhard de Chardin (1922) of the type of "Protoadapis angustidens," now lost. Line drawing is approximately natural size; other illustrations are somewhat enlarged. FIG. 3. Occlusal scanning electron micrograph of a cast of two teeth preserved in Leuven PLV- 35, referred specimen of "Protoadapis.filholi." Scale represents 1 mm. closely appressed in the midline to the bases ofthe metaconid and protoconid. This small cusp is the terminus of a small, arcuate paracristid. In both molars the cristid obliqua terminates medially, between the major trigonid jaw with the last premolar and M,-2. In com- cusps, creating a fairly deep hypoflexid notch. parable parts the dentition of this specimen Both teeth show well-developed buccal cin- closely matches both Teilhard's illustration gulids. The paraconid on M3 is lower on the of the type of Protoadapis angustidens, and face of the trigonid than is its counterpart on PLV-35; in preserved areas it also appears to M2. In each tooth a small thickening of the display a similar alveolar count to the latter. enamel exists just lingual to the paraconid, Again, the cheek teeth are characterized by separated from it by a narrow groove; this their "bulkiness." The last premolar is dom- "pseudo-twinning" of the paraconid occurs inated by a low, broad protoconid that bears further inferiorly on M3 than on M2. a stout metaconid closely approximated to it Stehlin's (1912) Protoadapis brachyrhyn- low down on its lingual face. The talonid is chus, transferred by Gingerich (1975) to its somewhat compressed, and the enclosed tal- own genus Cercamonius, is based on BNM onid basin is displaced lingually. The molar QV 619 (figs. 2, 4), a robust partial left lower trigonids lack distinct basins, are somewhat 6 AMERICAN MUSEUM NOVITATES NO. 2762

molars are rather straight and terminate at the juncture between the closely approxi- 1. 4 1,v., mated paraconid/metaconid pairs. The talonid basins are quite shallow and lack dis- lit. lk.'.IL - . "k -"A .. tinct hypoconulids. All teeth preserved display strong buccal cingulids. The primate usually known as Europole- mur or Protoadapis klatti is represented by a series of Geiseltal specimens in the collec-

-. ; tions ofthe GPIH, and has also been reported ; from elsewhere. The holotype is a crushed cranium and mandible, GPIH 232/3656 (fig. 6), and notable referred specimens include an unnumbered mandible with the two posterior 4v premolars and M, 3 (fig. 5), and the anterior part of a face, GPIH 4258 (fig. 6), with in- terlocked upper and lower teeth. Two further specimens are highly worn: GPIH 4234, an upper dentition, and GPIH 549, a lower dentition. The lower canines of klatti, as represented by this tooth in GPIH 549, are classically caniniform: simple, conical, and trenchant, with no elaboration of the base. In the lower molars the paraconid is small but twinned, notably on M1, and lies at the terminus ofan inferiorly trending paracristid. The paraconids lie a bit medial to the metaconid, and decrease in size from Ml to M3. There is a complete rounded hypocristid between the hypoconid and the entoconid in M1-2, un- interrupted by any notch or by the development ofa hypoconulid. On M2-3 the cristid obliqua is arcuate but terminates midway between the metaconid and protoconid; on M, this crest probably contacted the metaconid, but wear makes this unverifiable. The last two premolars are laterally compressed; the posterior bears a small, medially placed "hypoconid," which is joined by a weak cristid obliqua that runs up the posterior face ofthe protoconid. The talonid basin is further de- FIG. 4. Occlusal scanning electron micrograph lineated by a stronger crest emanating lin- of a cast of teeth remaining in BNM QV 619, type of "Protoadapis brachyrhynchus." Scale represents gually from the protoconid and running in- 1 mm. feriorly to join the hypoconid, creating an enclosed basin. The talonid is more weakly defined on the penultimate premolar. Both premolars possess a sharp, steeply sloping compressed, and possess distinct paraconid paracristid that turns lingually at the base of shelves that end in small paraconid swellings, the tooth, forming a moderate depression lin- lower down on the trigonid in M2 than in M,. gually. Distinct cingulids are present on the The cristids obliquae of the two preserved posterior halves of the posterior premolars. 1983 TATTERSALL AND SCHWARTZ: PROTOADAPIS 7

Ml bears a virtually complete buccal cingulid; this structure is less complete posteriorly on M2 and still less so on M3. From the crushed type jaw the antepenultimate premolar appears rudimentary (it may be a re- tained deciduous tooth), but possesses two roots that seem disproportionately large compared to crown size. The canines are slightly flattened and at their apex curve lingually; they bear a deep anterior vertical groove. The lateral incisor is larger than the medial but both are similar in being spatulate and small crowned. In the upper dentition, the crown of the central incisor is approximately half as large again as that of the lateral; both teeth are elongate and somewhat spatulate although lingually there is a marked margocrista. The upper canine seems to have been laterally compressed, bearing an anterior and a larger posterior vertical groove; the impression is of a pointed, daggerlike tooth. The antepenultimate upper premolar is small-crowned and of simple premolariform shape; the only elaboration is a tiny posterior swelling bearing slight medial and lateral cingula. The penultimate premolar of the type can be seen undamaged only laterally; in outline it appears as a roughly equilateral triangle. It bears cingular swellings in the para- and metastylar regions, which are joined by a weak cingulum. There appears to have been a lingual distension of the protocone region, which may have borne a small cusp. The last premolar is similar to the foregoing one in buccal Ij outline, but is slightly smaller and bears more distinctly developed stylar regions and cingula. This tooth is quite transverse, and bears a relatively large protocone from which em- anate a weak and truncated postprotocrista and a stronger preprotocrista and protocone fold. The postprotocrista becomes confluent with a small precingulum while the postcingulum reaches beyond the postprotocrista. The upper molars are characterized by very prominent trigon cusps and a very small tal- L, on. Of the crests of the talon the pre- and postprotocrista are the most emphasized, and FIG. 5. Occlusal scanning electron micrograph are more V- than U-shaped in configuration. of a cast of "Protoadapis klatti," unnumbered On Ml-2 the preprotocrista bears a strong specimen in GPIH. Scale represents 1 mm. parastyle which is confluent with the precingulum, and also a paraconule. On all molars 8 AMERICAN MUSEUM NOVITATES NO. 2762

FIG. 6. Above: upper dentition of GPIH 232/3656, type specimen of "Protoadapis klatti." Below: lateral view of GPIH 4258, referred specimen of same. Each scale represents 3 mm.

the precingulum also extends down and pocone occupies the posterolingual corner of around the face of the protocone and be- the tooth, thereby giving these teeth a squar- comes confluent with an equally strong post- ish appearance, although they are still highly cingulum. On M'-2 a distinct and pointed hy- transverse, dominated by the broad, short tri- 1983 TATTERSALL AND SCHWARTZ: PROTOADAPIS 9 gon. Buccal cingula are complete and well the cristid obliqua seems to have been developed on all upper molars, and MI ad- straight, terminating medially and producing ditionally bears a small mesostyle. On the deep hypoflexid notches and rather narrow reduced M3 the trigon enamel is somewhat talonids. wrinkled. In the same contribution, Gingerich cre- Protoadapis ulmensis is represented by the ated two additional new species of Protoad- few isolated teeth described by Schmidt-Kit- apis, P. russelli and P. louisi. Each new tler (1971, 1977) from the late Eocene Ger- species was based on an isolated lower molar, man site of Ehrenstein. As illustrated by and a largely unspecified variety of other Schmidt-Kittler, the upper molars of ulmen- specimens, also isolated teeth. The type of sis are characterized in preserved parts by Protoadapis russelli, Louis coll. Av- 183 from broad, arcuate protocristae; moderately com- the earliest Eocene site of Avenay, a pre- pressed buccal cusps interconnected and sumed right M1, has trigonid cusps that are bounded laterally by distinct crests; and well- very distinct and well-separated; the talonid defined pre- and postcingula that typically is low, with a shallow basin. The metaconid become confluent around the base of the and entoconid are separated by a steep slope, large, somewhat compressed protocone. In and the posterior face of the talonid is rather all upper molars the posterior face of the straight, bearing a small and medially posi- tooth seems to be somewhat swollen, and a tioned hypoconulid. The cristid obliqua is moderate ledge appears in the hypocone re- weak, and terminates medial to the base of gion. In all known upper molars the strong the metaconid. preprotocrista seems to arc past the paracone, The type of Protoadapis louisi, Louis coll. from which it is separated by a narrow Av- 118, is also a presumed right MI from groove. Schmidt-Kittler's illustration of one Avenay, but is larger than Av- 183. The tri- ofthe referred upper molars of ulmensis, Eh- gonid cusps ofthis specimen are tall, pointed, renstein 3, 1971, XXIV-4, shows an elabo- well-separated, and set in a large trigonid ration ofthe enamel fringing the lingual mar- with a deep basin. The talonid cusps are lower gin of the protocone to a degree that is also but still pointed, with a small and medially characteristic of Lepilemur. Another partic- placed hypoconulid. A distinct cristid obli- ular resemblance to the Lepilemur is found qua terminates in the midline between the in a low, broad enamel ridge that diverges bases of the protoconid and metaconid, cre- from the postprotocrista toward the post- ating a narrow but deep hypoflexid notch. A cingulum. Although these teeth as illustrated deep groove separates the metaconid from bear a remarkable resemblance to their coun- the entoconid. terparts in Lepilemur, they also differ in being Two specimens that have generally been slightly more transverse. But while the com- referred to Protoadapis recticuspidens (or P. parison with Lepilemur is reinforced on the curvicuspidens) have clearly been wrongly at- one hand by the detailed similarity between tributed. These are: MNHN-Louis- 15-Ma the single known upper premolar of ulmensis (fig. 7), a left dentary with a large caniniform and the middle upper premolar of the Mal- tooth followed by alveoli, the two posterior agasy form, on the other hand the isolated premolars, and M1 3, from the French Cui- M3 attributed to ulmensis is clearly dissimilar sian site ofMancy; and MNHN AL-5 181 (fig. from that of Lepilemur. The fossil tooth is 8), a left dentary from Epernay with the last badly damaged, however, and its attribution premolar and M2, illustrated as Protoadapis can only be conjectural. but not ascribed to species, by Lemoine in The Geiseltal specimen, GPIH 1209, on 1891. Descriptions of these specimens fol- which Gingerich (1977) based his new species low. Protoadapis weigelti, is totally inadequate for Louis- 1 5-Ma. The caniniform tooth in this such a purpose. A right dentary fragment with jaw is genuinely caniniform: simple, pointed, M1 2, this fossil preserves virtually no surface tall, slightly recurved, and without basal elab- detail of the teeth. From what little remains, oration. The two preserved premolars are rel- the trigonid of M2 seems to have been more atively trenchant, especially the penultimate, compressed than that of M. In both teeth which has a markedly truncated talonid. The 10 AMERICAN MUSEUM NOVITATES NO. 2762

the paracristid, bearing a small and dimin- ishing paraconid, moves inferiorly. In M3 the paracristid terminates at the base ofthe metaconid. Both M2 and M3 would have been larger than the corresponding teeth in the unworn type of recticuspidens. The cristid obliqua of M3 terminates low between the bases of the protoconid and metaconid, creating a rela- il, tively deep hypoflexid notch. The talonid heel of M3 is rounded, not very elongate, and 1 somewhat displaced lingually. There are two small alveoli in front ofthe caniniform tooth; behind this is an alveolus for a small, single- rooted tooth, and between this and the penultimate premolar, two roots for another tooth. AL 5181. At the front of the jaw there is an enormous backwardly sloping alveolus with apparently no teeth in front of it (the symphyseal surface seems to be present in this specimen); possibly this alveolus is ar- tificially enlarged and has "captured" smaller alveoli anterior to it. Behind the large alveolus is a small diastema followed by a small compressed alveolus followed in turn by two associated alveoli succeeded by a single alveolus of moderate size. The last premolar has a moderately sized metaconid low on the FIG. 7. Occlusal view of the dentition of face of the protoconid; it also had a distinct MNHN-Louis-15-Ma, type of new species de- and slightly buccal hypoconid and an en- scribed in this paper. Scale represents 3 mm. closed, lingually displaced talonid basin. The last premolar also had a complete buccal cingulid, and a distinct paracristid descended the face of the protoconid to arc lingually and last premolar lacks a paraconid but possesses ascend to meet the metaconid. M2 lacks a a distinct metaconid on the lingual face of distinct paraconid; the paracristid slopes the protoconid. It also has a distinct but down the face of the protoconid and termi- somewhat truncated talonid whose basin is nates at the base of the metaconid, enclosing displaced lingually, and, like the molars, a tiny fovea. The talonid cusps, especially the bears a relatively complete buccal cingulid. entoconid, are relatively small and relegated A sharp paracristid descends steeply down to the periphery of the tooth where they are the face of the protoconid, at the base of more or less confluent with the cristids that which it arcs lingually and rises to become enclose a relatively shallow talonid basin. confluent with the metaconid. The cristid The cristid obliqua terminates at the midline obliqua on Ml meets the metaconid, but this of the back of the trigonid, and there is a crest terminates centrally on M2 and M3. The distinct although not robust buccal cingulid. talonid basins are shallow and somewhat nar- The two preserved teeth in this jaw are very row; they are rounded distally and lack hy- close in morphology to those of the L- 15 poconulids on M 12. The trigonids become Mancy jaw, which increases the probability more compressed in the sequence Ml3, and that, originally, small teeth were indeed pres- the small swelling at the terminus ofthe para- ent at the front of the jaw. cristid becomes smaller and shifts medially Finally, it is with this group that Grandi- in the same sequence. Also in this sequence dier's (1904) species Pronycticebus gaudryi is 1 983 TATTERSALL AND SCHWARTZ: PROTOADAPIS I1I most appropriately described. This is unique- ly known from the holotype, MNHN 1893- 11 (fig. 9), an exquisitely preserved cranium and mandible from the French late Eocene (Headonian) site ofMemerlein. Both the cranium and the mandible preserve the last two premolars and M1I3, with alveoli anteriorly. The teeth of the mandible seem to be less worn than those ofthe upperjaw, which have lost a certain amount of detail. The penultimate upper premolar is dominated by a large buccal paracone, with a small protocone lingually. There is a distinct parastyle and a larger metastyle. In the last upper premolar these stylar buildups are subequal in size and are connected buccally by a cingulum. The paracone is broader than in the penultimate premolar, and the protocone is distinctly larger. A well-developed preprotocrista con- nects the protocone and parastyle; a postcingulum runs from the metastylar region and terminates distally in a small swelling at the base of the protocone. The upper molars are characterized by their transverseness and are relatively short. M' is smaller than M2-3, which are subequal in size. The anterior molar is convex on its anterior surface and con- cave posteriorly; this posterior waisting is emphasized by the distension of the hypocone and metacone regions. The metacone distension is caused in part by a distinct and prominent metastylar crest. The upper molar protocristae seem to have been neither broad nor parabolic in configuration. A crest that emanates from the paracone region and skirts the paracone to meet the precingulum ap- FIG. 8. Occlusal scanning electron micrograph pears not to be the preprotocrista, since the of a cast of teeth preserved in MNHN AL-5 181, latter crest can be discerned, faintly, running referred specimen ofnew species described in this to the apex ofthe paracone. M2 is only slightly paper. Scale represents 1 mm. convex anteriorly, and posteriorly it is essen- tially straight sided. The paracone is much larger than the metacone, but as in M' the centrocristae are well emphasized. The hy- The penultimate lower premolar is domi- pocone is confluent with the postcingulum, nated by the protoconid, which bears a small and appears to have been more shelflike than truncated talonid and lacks the posterior ex- cusplike. M3 is roughly triangular, with a cavation characteristic of Adapis and its al- shelflike ridge posterior to the base of the lies. The last premolar is also dominated by protocone. As in M2, the buccal crests are the protoconid; this bears anteriorly, at its emphasized and the paracone is dominant. base, a small paraconid shelf, and posteriorly, All molars bear a distinct precingulum that well down on its lingual surface, a small meta- reaches partway around the base of the pro- conid. The talonid is somewhat better ex- tocone; in M2 this structure terminates in a pressed than is that of the penultimate lower tiny style, and in M3 in a distinct pericone. premolar, and bears a minute, medially 12 AMERICAN MUSEUM NOVITATES NO. 2762

L J

lqpp- 2,, . . i.

FIG. 9. Occlusal scanning electron micrographs of casts of the upper (left) and lower preserved dentitions of MNHN 1893-1 1, type of Pronycticebus gaudryi. Scale represents 0.5 mm. placed hypoconid from which emanates a development increases in the sequence M1I3. cristid obliqua that runs down to the meta- The trigonid cusps of the lower molars are conid, and a hypocristid that encloses the tal- somewhat compressed and the trigonids have onid basin. In both preserved premolars the sheer posterior walls. Except for the enlarged paracristid turns lingually to enclose a talonid heel of M3, the molars lack any sign depression. Lingual to the hypoconid is a of hypoconulids. The cristid obliqua on Ml tiny, compressed cusp that may represent the kinks round to the metaconid, but this crest entoconid. This weak buccal cingulid is more terminates at the base of the protoconid on complete than on the penultimate premolar. M2-3. The trigonid of Ml is somewhat com- Buccal cingulids on the molars occur pri- pressed, that of M2 more so, and that of M3 marily around the trigonids; such cingular more compressed yet. The molar trigonids 1983 TATTERSALL AND SCHWARTZ: PROTOADAPIS 13 are all similar, however, in that a weak para- preciably broader than the trigonid. On the cristid runs down from the protoconid and basis of such poor material it is perhaps un- arcs lingually, where it becomes more ex- wise to speculate in unduly precise terms panded and supports twinned small cuspules about the affinities of Protoadapis; neverthe- that may be identified as twinned paraconids. less, one might note that the buccal emplace- On M1_2 the talonid basin is enclosed by the ment of the cristid obliqua, the peripherali- rather arcuate cristid obliqua and hypocris- zation ofthe cusps that leads to the complete tid; from front to back in the molar sequence enclosure of the talonid, at least of M2, and the cristid obliqua becomes less well defined, the truncated nature of the better-preserved and the paracristid becomes lower. The tal- M3 of recticuspidens, may invite comparison onid heel on M3 is displaced lingually. ofProtoadapis with Agerinia, as was done by Szalay and Delson (1979). DISCUSSION The second major group represented in the assemblage consists of Protoadapis klatti, As we have already noted, Protoadapis as Pronycticebus gaudryi, and a previously un- generally conceived is perhaps the most high- recognized species represented by the ly speciose and long-lived primate genus of MNHN specimens Louis- 1 5-Ma and AL- the European Paleogene. However, within 5181. Although a substantial case could be the material that has traditionally been al- made for recognizing three genera within this located to this genus and its close allies, there assemblage, it is clear that the forms com- are in fact three major groups discernible, in prising it are very closely related, and for the addition to three described species that are purposes of taxonomic simplicity we regard not only clearly not Protoadapis, but are also them here as belonging to a single genus, by referable to none of these major groups. In- priority Pronycticebus. We should note that sofar as can be told from the morphology Szalay and Delson (1979) have already em- preserved, the types ofLemoine's two species phasized the affinities between their concept curvicuspidens and recticuspidens are not of Protoadapis and Pronycticebus; this was only somewhat dissimilar to each other, but presumably on the basis of the better Mancy both are distinctly different from all other specimen rather than on the type material. specimens subsequently referred to Protoad- Before discussing further the enlarged genus apis. Because of the heavily worn condition Pronycticebus, we will formally name the new of the specimens involved, it would be ill- species, of which the morphology has been advised to attempt to separate curvicuspidens given at some length above. and recticuspidens at other than the species level, at least at present. It may be noted, however, that curvicuspidens differs strongly ORDER PRIMATES from recticuspidens in having a much better PRONYCTICEBUS GRANDIDIER, 1904 developed trigonid, with a broad paracristid Pronycticebus mancyi, new species and the tips ofthe two main cusps divergent; the protoconids and metaconids are much Protoadapis curvicuspidens (in part), Russell, more closely approximated in the somewhat Louis, and Savage, 1967. smaller recticuspidens, and the paracristid is TYPE: MNHN-Louis- 15-Ma, left dentary both smaller and higher on the anterior molar preserving a large caniniform tooth, the last face. In any event, at present it seems most two premolars, and M1-3 (fig. 7). prudent to regard these two species as con- TYPE LOCALITY: Mancy, France: early generic, and as the only known representa- Eocene. tives of Protoadapis. In this view, the dis- HYPODIGM: Type plus MNHN AL-5181, tinctive characteristics ofthe restricted genus left dentary with last premolar and M2 (fig. Protoadapis include the following: a broad 8), from Epernay. paraconid shelf, a buccally emplaced cristid ETYMOLOGY: To reflect the provenance of obliqua that terminates at the base of the the type specimen. protoconid, a low and shallow talonid basin, DIAGNOSIS: Larger than Pronycticebus at least on M2, and a talonid that is not ap- gaudryi and comparable in size with Pro- 14 AMERICAN MUSEUM NOVITATES NO. 2762 nycticebus klatti. Cheek teeth relatively much that bears some anterior and posterior stylar broader than in gaudryi; cusps more inflated. development and buccal cingula. The pei- Differs from klatti in the greater height of its ultimate premolar bears a small protocone penultimate premolar, which has less distinct swelling that is more discernible as a cusp on buccal cingulids but more lingual cingulid de- the last premolar, and a distinct protocone velopment, and which shows a well-defined fold. To judge from the preserved roots in internal depression. Bears one more (tiny) gaudryi, both species had a large, stout upper premolar than does klatti. The last premolar canine that may have been rather more com- of mancyi is relatively larger and more in- pressed in klatti, at least at the alveolar mar- flated than that of klatti, with a more distinct gin. Distinctions between gaudryi and klatti metaconid and a broader talonid basin. All also reside in the greater development ofpre- molars of mancyi bear stronger and more and postcingula in klatti; these become con- complete buccal cingulids than those of fluent around the base of the upper molar klatti, and show relatively larger (although protocones. Pronycticebus gaudryi also re- still absolutely tiny) paraconids. tains an additional (fourth) upper premolar that is absent in klatti; this disparity may also Pronycticebus gaudryi has traditionally have been true ofthe lower dentition. In this been regarded as quite distinct among "ada- regard klatti is also distinguished from man- pid" primates (see, for example, Gingerich cyi, at least in the lower jaw. [1977]). Szalay and Delson (1979), however, At the present state of our knowledge it is have suggested that Pronycticebus may be de- difficult to postulate precise affinities for scended from their concept of Protoadapis Pronycticebus. If, however, there is a genuine (which includes filholi and related forms, but phylogenetic signal in characters such as the excludes klatti). We concur with neitherjudg- arcuateness of the cristid obliqua, the encir- ment; for if one allows for dental wear it is clement of the talonid basin by crests, the clear that morphologically gaudryi is very transverseness of the upper molars, the close to "Protoadapis" klatti as known from V-shaped configuration of the protocristae, the Geiseltal. Distinguishing features in the and an apparent morphocline from front to lower dentition (the upper is known only in back in the reduction of the paraconid and gaudryi and klatti) of a genus Pronycticebus its transformation into a shelf in close prox- that embraces the species gaudryi, klatti, and imity to the metaconid, then perhaps the mancyi include the following: the presence in most appropriate comparisons might lie the lower molars oftiny twinned paraconids, among species of Pelycodus such as P. ral- most evident on M, but also discernible on stoni and P. trigonodus. the posterior molars; arcuate cristids obli- The third group within the assemblage is quae on M2 that are more internally directed the most distinctive and thereby the easiest than in Protoadapis; a cristid obliqua on M3 to delineate. This is not least because it con- that terminates on the lingual side ofthe base sists of a single species which contains the of the protoconid; and a more elaborate tal- fossils that have been variously allocated to onid heel than in Protoadapis. Protoadapis filholi (or angustidens), Cerca- The upper dentitions of Pronycticebus monius (or Protoadapis) brachyrhynchus, gaudryi and Pronycticebus klatti are known and possibly Protoadapis weigelti. Szalay and from decent specimens, the unique represen- Delson (1979) have also recently suggested tative ofthe former (fig. 9) more heavily worn the synonymy of weigelti with filholi, and in than the best specimens ofthe latter (e.g., fig. the same contribution proposed that the lat- 6). In both species the upper molars are dis- ter might bear an ancestral relationship to tinguished by being rather transverse, with brachyrhynchus. We should note that since distinct protocristae forming a V, and a the holotype ofProtoadapis weigelti is totally somewhat ledgelike postcingulum that ter- inadequate to support analysis, we hazard minates lingually in a broad hypocone. The that it belongs here largely on the basis of its last two premolars are each dominated by a size, almost the only character it retains. rather compressed and trenchant paracone The specimens referred to the species fil- 1983 TATTERSALL AND SCHWARTZ: PROTOADAPIS 15

TABLE 1 Summary of Species Recognized or Discussed in This Paper, and Their Synonymies This Paper Synonyms Reference GROUP 1 Protoadapis curvicuspidens (Lemoine, 1878) Plesiadapis curvicuspidens Lemoine, 1878 Protoadapis recticuspidens (Lemoine, 1878) Plesiadapis recticuspidens Lemoine, 1878 GROUP 2 Pronycticebus gaudryi Grandidier, 1904 Pronycticebus klatti (Weigelt, 1933) Europolemur klatti Weigelt, 1933 Megatarsius abeli Weigelt, 1933 Protoadapis klatti Simons, 1962 Pronycticebus mancyi, new species Protadapis sp. Lemoine, 1891 Protoadapis curvicuspidens Russell, Louis, and Savage, 1967 GROUP 3 Cercamonius brachyrhynchus (Stehlin, 1912) Adapis angustidens Filhol, 1888 Protadapis brachyrhynchus Stehlin, 1912 Protoadapis angustidens Teilhard de Chardin, 1922 Protoadapis weigelti Gingerich, 1977 Protadapisfilholi Gingerich, 1977 Other Taxa Notes Protoadapis eppsi Cooper, 1932 Transferred to new genus as Cantius eppsi by Simons (1962); often regarded as Pelycodus Protoadapis ulmensis (Schmidt-Kittler, 1971) Not Protoadapis; appears to be an adapine sensu stricto. Protoadapis russelli Gingerich, 1977 Not Protoadapis; primate incertae sedis. Protoadapis louisi Gingerich, 1977 Not Protoadapis; primate incertae sedis. holi (or angustidens) and brachyrhynchus pared since Stehlin's time with North consist of lower jaws which differ strikingly American Eocene primates such as Pelycodus from other material that has in the past been and Notharctus. In our view, Teilhard's referred to Protoadapis. All have broad, (1922) older, and Gingerich's more recent "swollen" cheek teeth that show considerable (1975) comparison with the larger species of lingual development and do not incorporate Notharctus is highly appropriate. In the the cusps into shearing crest systems. The "puffiness" ofthe teeth, in the closed-up and molar paraconids are appressed to the meta- lingually shifted trigonids, in the wrinkling conids, further down the tooth in the se- and posterior lowering of the paraconid re- quence M13. The "pseudo-twinning" of the gion, in the relative straightness ofthe cristids paraconid is in fact no more than a wrinkling obliquae, and in the occurrence of a narrow ofthe enamel, rather than a clear delineation and shallow entoconid notch, Cercamonius of two cusps of the kind seen in Pronyctice- brachyrhynchus provides a remarkably close bus. The cristids obliquae are straight, and match for forms such as Notharctus venti- the talonid basins are very shallow. The ento- colus (as represented by CMNH 37159). It conid is bounded internally by a narrow differs from the largest Notharctus, N. cras- groove which appears posteriorly as a small sus, in lacking robust cresting ofthe posterior notch. premolar. Unquestionably, however, it is with this group of Eocene primates that Cerca- Cercamonius brachyrhynchus is thus high- monius is most closely allied. ly distinctive relative to other European pri- This division ofthe total assemblage leaves mates, and has, indeed, regularly been com- the isolated teeth assigned by Schmidt-Kit- 16 AMERICAN MUSEUM NOVITATES NO. 2762 tler (1977) to Protoadapis ulmensis, and by Pal. Univ. Michigan, vol. 24, no. 15, Gingerich (1977) to Protoadapis russelli and pp. 163-170. louisi. The ulmensis material seems referable 1977. New species ofEocene primates and the to the clade of Adapis sensu stricto, as phylogeny ofEuropean Adapidae. Folia Schmidt-Kittler (1971) had originally con- Primat., vol. 28, pp. 60-80. Lemoine, V. cluded; it does not belong with any of the 1878. Communication sur les ossements fos- other material discussed here. The species siles des terrains tertiaires inferieurs des russelli and louisi are strikingly characterized environs de Reims. Bull. Soc. Hist. Nat. by distinct and well-separated trigonid cusps, Reims, for 1878, pp. 90-113. a feature which by itself distinguishes them 1880. Communication sur les ossements fos- from all of the other primates under consid- siles des terrains tertiaires inferieurs des eration. Indeed, the remarkably large trigo- environs de Reims. Assoc. Fran9aise nids ofthese specimens demarcate them even Avanc. Sci., Cong. de Montpellier, Ses- from the most primitive species ofPelycodus. sion 8, pp. 585-594. Both appear to belong to the same but 1891. Etude d'ensemble sur les dents des group, mammiferes fossiles des environs de their broader affinities are obscure to us. Cer- Reims. Bull. Soc. Geol. France, ser. 3, tainly the suggestion by Szalay and Delson vol. 19, pp. 263-290. (1979) that they may belong with filholi Russell, D. E., Pierre Louis, and D. E. Savage seems inappropriate. 1967. Primates of the French early Eocene. Univ. Calif. Publ. Geol. Sci., vol. 73, pp. 1-46. SUMMARY Schmidt-Kittler, N. Our review of the material that has been 1971. Eine unteroligozan Primatenfauna von assigned at one time or another to the Eu- Ehrenstein bei Ulm. Mitt. Bayer. Staats- genus Protoadapis samml. Palaont. Hist. Geol., vol. 1, pp. ropean Eocene primate 171-204. shows that the genus has in fact become an 1977. Neue Primatenfunde aus unteroligoza- umbrella for a heterogeneous assortment of nen Karstspaltenfullungen Suddeutsch- taxa. We find that among this material three lands. Mitt. Bayer. Staatssamml. Pa- major groups are represented, together with lIont. Hist. Geol., vol. 17, pp. 177-195. three described species that belong to none Simons, E. L. of these. For convenience of reference we 1962. A new Eocene primate genus, Cantius, summarize our conclusions in table 1 by list- and a revision of some allied European ing those species we recognize together with lemuroids. Bull. Brit. Mus. (Nat. Hist.), their synonyms. Geol., vol. 7, no. 1, pp. 1-36. Stehlin, H. G. 1912. Die Sliugetiere des schweitzerischen LITERATURE CITED Eocaens. Critischer Catalog der Mate- rialen. Abh. Schweiz. Pal. Ges., vol. 38, Cooper, C. F. pp. 1165-1298. 1932. On some mammalian remains from the Szalay, F. S., and E. Delson lower Eocene ofthe London Clay. Ann. 1979. Evolutionary history of the primates. Mag. Nat. Hist., Dec. 10, vol. 9, pp. New York, Academic Press, 580 pp. 458-467. Teilhard de Chardin, P. Filhol, H. 1922. Les mammiferes de l'Eocene inferieur 1888. Description d'une nouvelle espece francais et leurs gisements. Ann. Pale- d'Adapis. Bull. Soc. Philomath. Paris, ont., vol. I1, pp. 1-108. ser. 7, vol. 12, pp. 10-12. Weigelt, J. Gingerich, P. D. 1933. Neue Primaten aus der mitteleozanen 1975. A new genus of Adapidae (Mammalia, (oberlutetischen) Braunkohle des Gei- Primates) from the late Eocene ofsouth- seltals. Nova Acta Leopoldina, n.f., vol. ern France, and its significance for the 1, pp. 97-156. origin ofhigher primates. Contrib. Mus.