The Phylogenetic Relationships of Adapidae (Primates, Lemuriformes)

THE PHYLOGENETIC RELATIONSHIPS OF ADAPIDAE (PRIMATES, LEMURIFORMES)

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JEFFREY H. SCHWARTZ AND IAN TATTERSALL

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JEFFREY H. SCHWARTZ Assistant Professor, Department of Anthropology University of Pittsburgh

IAN TATTERSALL Associate Curator, Department of Anthropology The American Museum of Natural History

VOLUME 55: PART 4 ANTHROPOLOGICAL PAPERS OF THE AMERICAN MUSEUM OF NATURAL HISTORY NEW YORK : 1979 ANTHROPOLOGICAL PAPERS OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 55: part 4, pages 271-283, figures 1, 2

Issued January 23, 1979 Price. $1.15 a copy

ISSN 0065-9452

Copyright ©) The American Museum of Natural History 1979 ABSTRACT The Eocene Holarctic primates of the family are to Lemur and Varecia. This phylogenetic rela- Adapidae have generally been considered to be close tionship implies that a split prior to earliest Eocene to the ancestry of the lemurs of Madagascar, them- occurred between the common ancestor of Lep- selves conceived of as a homogeneous group. Re- ilemur-Megaladapis-Hapalemur-Adapidae and that of examination of available evidence reveals, however, Lemur-Varecia and, in tum, implies earlier dates yet that the adapids appear to be more closely related to for the divergences establishing the major taxa of the Lepilemur, Megaladapis, and Hapalemur than they modern Malagasy primate fauna.

INTRODUCTION The consensus regarding the phylogenetic adapid relationships has been in the direction of relationships of the Eocene primate family emphasizing the affinity of these Eocene forms Adapidae has changed remarkably little in the to non-Malagasy primate groups, or of denying more than half-century since Gregory (1920) any special relationship whatever to other pri- published his exhaustive monograph on mate taxa. In contrast, however, our own pre- Notharctus, the best-known North American liminary investigation of relationships among representative of the group. Although Gregory's the so-called lower primates has suggested that contention that the adapid subfamily Notharc- the adapids are not "basal" lemuriforms, and tinae contained the ancestor of the New World that in fact they form a sister-group with only monkeys has not survived into the recent litera- Hapalemur and Lepilemur among the extant ture, his basic concept of the adapids as primi- Malagasy lemurs.1 It is this relationship, and its tive lemuroids has been almost universally implications, that we wish tentatively to ex- adopted. plore here. This basic assessment has remained largely unchanged even while suggestions as to the ACKNOWLEDGMENTS affinity of the adapids with other primate groups have multiplied. Thus Gingerich (1973, We thank Ms. S. F. Peters, Dr. H. B. 1975a, 1975b), whilst proposing Rollins, and especially Dr. L. Krishtalka for that the origin discussion and criticism of various aspects of of the "higher" primates is to be sought within this work, and Mr. N. Amorosi for the figures. Adapidae, nonetheless emphasized the relation- Drs. M. C. McKenna and S. Anderson (the ship of this family with the Malagasy le- American Museum of Natural History), C. L. muroids. Szalay (1974), however, has Gazin, expressed his belief that "The Eocene lemurs R. Emry, and R. Thorington (Smithso- have a more primitive dentition, with less spe- nian Institution), P. Andrews, Ms. T. Molleson cialized and Ms. P. Napier (British Museum [Natural teeth and the tritubercular molars of History]), Drs. R. Saban (Museum National early Tertiary primates, than any of the d'Histoire Madagascan species . . . which . . . puts them Naturelle) and M. R. Dawson (Car- closer negie Museum of Natural History) kindly to the ancestor of the catarrhines than allowed access to specimens in their charge. any Madagascan lemur" (p. 53). Cartmill and This Kay (1978) have departed yet further from research was supported in part by Univer- orthodoxy in arguing that no characters can be sity of Pittsburgh administered NIMH BSSG distinguished which indicate any particular rela- funds and a John G. Bowman Faculty Grant (to tionship between the adapids and any other JHS). group of primates, haplorhine or strepsirhine. Evidently, then, such shifting as there has 1We use the vernacular term "lemur" solely to mean been away from the classical viewpoint of "Malagasy primate." 273 274 ANTHROPOLOGICAL PAPERS AMERICAN MUSEUM NATURAL HISTORY VOL. 55

ABBREVIATIONS American Museum of Natural History; USNM, The following abbreviations are used in the the Smithsonian Institution; BM (NH), British text for names of institutions: AMNH: the Museum (Natural History).

THE MORPHOLOGICAL EVIDENCE

THE DENTITION: MOLAR MORPHOLOGY acterized by: (1) cuspal compression with sharp, almost continuous (except for the region In support of his contention that the adapids, of the entoconid) intercusp cresting; and (2) a or at least the adapines, were near the ancestry cristid obliqua confluent with the paracristid. of the lemurs and lorises, Gregory (1915, 1920, All of these derived states are also seen in 1922) repeatedly called attention to similarities the adapids (cf. Gingerich, 1975a; Gregory, in molar morphology between Adapis parisien- 1915, 1920; Stehlin, 1912). In the notharctines sis and Lepilemur; he thus regarded the latter as they are less pronounced than they are in the representing the primitive strepsirhine condi- adapines, amongst which their degree of ex- tion. More recently Gingerich (1975a) has pression is very close to that found in the Lep- chosen to compare A. parisiensis instead with ilemur-Megaladapis-Hapalemur group. Hapalemur. Gingerich chose Hapalemur for Some specimens of Adapis parisiensis comparison because it is supposedly "interme- (Stehlin, 1912) and of Notharctus (Gregory, diate" in the range of variation exhibited by 1920) yield information on patterns of dental other extant lemurs, and "because of its rela- eruption and some Notharctus also contribute tively generalized nature . . . which . . . may data on dental development. From Stehlin's and closely approximate the condition of the an- Gregory's descriptions and illustrations and our cestral lemur stock" (p. 70). We have, study of juvenile specimens, it appears that in however, recently determined that Lepilemur both of these adapids, M1 and then M2 erupt and Hapalemur exhibit a number of derived first; M3 comes into place later, somewhat after character states in their dental morphology the anterior dentition. In Adapis and which are not shared with other extant strep- Notharctus, the anteriormost premolar (which is sirhines and which are otherwise seen among most probably a retained deciduous tooth the lemurs only in the subfossil Megaladapis [Schwartz, MS]), erupts well ahead of the poste- (Tattersall and Schwartz, 1974). Derived fea- rior three premolars which, in turn, replace tures of upper molar morphology of Hapale- their predecessors in the sequence P4 -* P3 -- mur, Lepilemur and Megaladapis include: (1) a P2. Study of maxillary (AMNH 13025) and reduction of lingual, especially cingular, devel- mandibular (AMNH 13029) remains of No- opment; (2) a protocone which, although large, tharctus tyrannus, in which the posterior three does not dominate the trigon; (3) buc- deciduous premolars have not been shed, re- colingually compressed (and hence with sub- veals a developmental pattern for the permanent crescentic-crescentic cross sections) paracones premolars of P2 --p4pl P3. The sequence of and metacones which are connected by marked eruption of the permanent premolars thus dif- para- and metacristae; and (4) a strikingly de- fers from the developmental sequence. veloped preprotocrista on MV-3 which passes Amongst strepsirhines, the only taxa which anterior to the paraconule and connects with a also convert a P2 P4 -> P3 developmental crest or shelf extending from the parastylar re- pattern to P4 -> P3 P2 in eruption are Lep- gion. ilemur and Megaladapis (Lamberton, 1938; The lower molars also display derived char- Schwartz, 1974a, 1975a). This is in marked acters not shared with other Malagasy forms or contrast to the majority of lemurs and lorises with primates in general. These teeth are char- which possess the primitive sequence of premo- 1979 SCHWARTZ AND TATTERSALL: ADAPIDAE 275 lar appearance (P2 -- P4 -* P3) in both devel- toothcomb is invariably associated with an opment and eruption. Hapalemur, Lemur catta, unfused mandibular symphysis, it is likely and the archaeolemurines are similar to Lep- that these two character states were present to- ilemur, Megaladapis, and adapids in that their gether in the ancestral form (cf. Tattersall and premolars erupt in a posteroanterior sequence Schwartz, 1974). but differ in that this is also the order in which Many authors have suggested that the tooth- these teeth develop (Schwartz, 1974a, 1975a). comb (and, by implication, an unfused sym- Other aspects of craniodental morphology indi- physis) was derived from a condition similar to cate that the sequence P4 -- P3 -> P2 in devel- that seen in most adapids, in particular Adapis, opment and eruption arose independently in L. wherein the lower anterior teeth are orthally catta, the archaeolemurines and Hapalemur (cf. emplaced and associated with a fused mandibu- Tattersall and Schwartz, 1974). lar symphysis (e.g., Stehlin, 1912; Gregory, The early appearance of Ml followed by M2 1920, 1922; Hill, 1953; Clark, 1962; Martin, also represents a derived character state (Sch- 1972; Cartmill, 1975; Gingerich, 1975a). Si- wartz, 1974a, 1975b); the primitive state is mons (1972), however, has correctly remarked marked by the eruption first of the anterior that Adapis, at least, could not have given rise dentition. The only other strepsirhines to dis- to modern strepsirhines precisely because its play early eruption of MI, followed by M2, are fused symphysis is a specialization. In the light Megaladapis, Lepilemur, Galago demidovii, of this observation, it would be extremely ques- and, to some extent, Microcebus murinus. tionable to view the adapid condition as predat- Again, wider comparisons indicate the acquisi- ing the acquisition of an unfused symphysis in tion independently of this feature in the Mega- association with a toothcomb, as this would ladapis-Lepilemur-(and presumably also, with imply that the unfused symphysis, at least, was subsequent loss)-Hapalemur group. secondarily derived, and not a primitive reten- The evidence of both molar and premolar tion. eruption sequences thus rules out the possibility Although most known adapids possessed a that adapids (or at least those in which the fused symphysis in association with orthally characters can be discerned) were ancestral to emplaced lower teeth, the well-represented the lemurs as a whole, and strongly suggests a early Eocene adapid Pelycodus, found in both sister relationship between Megaladapis- North America Ind Europe (Russell, Louis and Lepilemur-Hapalemur and Adapidae. Savage, 1967), possessed an unfused symphysis (Gregory, 1920) and (possibly) procumbent THE DENTITION: EVOLUTION OF THE lower anterior teeth (Simons, 1972). As Gre- TOOTHCOMB gory (1920) pointed out, the unfused sym- With the exception of Daubentonia, all ex- physis, at least, is primitive for Adapidae. tant strepsirhines possess a toothcomb. Com- In arguing for Adapis as ancestral to lemurs parative morphological, as well as develop- and lorises, Gingerich (1975a) pointed out that ment/eruption studies have shown that, while the lower canines of A. parisiensis are func- there are fewer teeth in the indriine toothcomb tionally incisors, i.e., that the anterior dentition than in that of other lemurs and lorises, the functioned as a unit, as is the case with the lateral procumbent teeth (canines) of all these toothcomb (or "dental scraper"). Gingerich forms are homologous as are the teeth of the suggested that, with the modification of this central set (incisors) (Martin, 1972; Schwartz, dental complex to form a toothcomb, sym- 1974b). It thus seems virtually certain that the physeal fusion would no longer be necessary toothcomb was not acquired independently, but and, thus, would not occur during ontogeny. was present prior to the major strepsirhine di- This argument, however, could equally well vergences (Martin, 1972; Schwartz, 1974b; Tat- apply the other way around to explain the tran- tersall and Schwartz, 1974). Similarly, since sition from a toothcomb to an orthal anterior except among the larger subfossil lemurs the dentition (as happened, for instance, in the evo- 276 ANTHROPOLOGICAL PAPERS AMERICAN MUSEUM NATURAL HISTORY VOL. 55 lution of the Malagasy subfossil Hadropithecus BASICRANIAL STRUCTURE: THE BONY [Tattersall, 1973b; Tattersall and Schwartz, EAR 1974]). Indeed, such a transition would make better sense functionally; certainly, when other The character which is perhaps most com- characters are considered, indications are that monly adduced to indicate for the adapids not the possession of a toothcomb, in association only affinities with, but also a position near the with an unfused symphysis (as suggested in ancestry of, the Malagasy lemurs, is the struc- Pelycodus and possibly Pronycticebus [Simons, ture of its bony ear. In essence, this argument 1972]), was primitive for Adapidae. involves the presence of an enlarged auditory bulla, of petrosal formation, which extends laterally beyond the inferior border of the tym- MANDIBULAR MORPHOLOGY panic ring. This configuration is characteristic of the adapids and also of most of the lemurs In a review of relationships among the Mal- with the notable exception of the highly modi- agasy lemurs (Tattersall and Schwartz, 1974), fied Megaladapis and the palaeopropithecines; we proposed that a relatively long and slender in the latter, this may well be a size-related mandibular corpus, in association with a modification (Tattersall, 1973a). hooked gonial region, represents the primitive Although this type of bullar conformation condition for strepsirhines. Pelycodus and has been argued by many to represent the prim- Notharctus retain this primitive condition, itive primate condition (e.g., McDowell, 1958; whereas the other adapids tend to have a rela- Clark, 1962; Szalay and Katz, 1973; Tattersall tively shorter and deeper mandibular corpus and Schwartz, 1974; Cartmill, 1975) recent (cf. Gazin, 1958; Gregory, 1920; Stehlin, 1912). work on the ontogeny of the bulla (Cartmill, With regard to the mandibular condyle, we 1975; MacPhee, 1977) indicates otherwise. distinguished three discrete morphologies Rather it appears that possession of an extrabul- among the lemurs: (1) the primitive condition, lar tympanic ring-as in earlier ontogenetic as seen in Lemur, in which the condyle is stages-is primitive and that the condition broad transversely, and the somewhat posteri- wherein the lateral edge of the bulla has grown orly directed articular facet is distinct from, laterally beyond the ectotympanic is derived rather than confluent with, the posterior surface (Schwartz, Tattersall and Eldredge, in press). of the condylar neck; (2) a derived condition Common possession of this apomorphy by seen in the indriines, in which the condyle may adapids, Lemur, Varecia, Lepilemur, Hapale- be broad transversely, but is also more or less mur, Daubentonia, and indriids most certainly strongly curved in the coronal plane, and where bespeaks their common ancestry but indicates the articular facet is confluent with the posterior nothing about relationships among these taxa. aspect of the condylar neck; and (3) a different derived condition, seen among the lemurs only BASICRANIAL STRUCTURE: THE in Lepilemur, Megaladapis, and some Hapale- CAROTID CIRCULATION mur (e.g., USNM 63355) in which a distinct articular facet descends, medial to the condylar Considerable emphasis has been placed on neck, from the posterior aspect of the trans- the pattern of carotid circulation in the deter- versely broad condyle. This last configuration mination of the phyletic relationships of mam- is elsewhere found only in the adapids, mals. The diversity of taxa characterized by an amongst which the posteromedial extension of internal carotid artery which sends off three the articular facet is most pronounced in major branches (medial, promontory, and sta- Notharctus (e.g., AMNH 21960, 21864 and pedial), with the promontory larger than the 13230), Smilodectes (e.g., USNM 17995 and stapedial, suggests strongly that this condition 21815) and Adapis (e.g., BM(NH) 1633 and is primitive for Eutheria (cf. McDowell, 1958; 7506); it is present, but slightly less pro- McKenna, 1966; Szalay, 1975). Since this con- nounced, in Pelycodus (e.g., AMNH 15019). dition is also characteristic of the microsyopid 1979 SCHWARTZ AND TATTERSALL: ADAPIDAE 277 primates (McKenna, 1966), it is reasonable to In contrast to each of these configurations is regard it as having been present in the ancestral that seen in Lemur and Varecia: the palatine primate. Within Primates, deviations from this broadly contacts the frontal and lacrimal and condition thus represent derived states. thus separates the maxilla from both of these One such derived state is where the medial bones (Clark, 1962; Kollman, 1925); when branch of the internal carotid is absent, and the Phaner, Hapalemur, and Lepilemur do not dis- stapedial is larger than the promontory artery. play the configuration discussed above, they This occurs in adapids, Lemur, Varecia, Hapa- have this one. This pattern also characterizes lemur, the indriines, the archaeolemurines, and, tupaiids, marsupials, Dermoptera and Mac- to some extent, in the palaeopropithecines roscelidea, although some specimens are seen (Saban, 1963; Szalay, 1975; Szalay and Katz, to have part of the ethmoid exposed between 1973; Tattersall, 1973b). Lepilemur and Mega- the palatine and the lacrimal (Evans, 1942; ladapis possess different conformations: in the Muller, 1934; Saban, 1956). Tarsius has a simi- former the stapedial artery is feebly developed, lar disposition of the ethmoid vis-a-vis the pal- whilst in the latter it is entirely absent. atine and lacrimal; lorisids, cheirogaleids and Anthropoidea display variants of this configura- THE ORBITAL FOSSA tion (see Cartmill, 1975, for further discussion). STRUCTURE OF Whilst not explicitly stating this, Clark Another character which has received much (1962) implied that the palatine-lacrimal config- attention in discussions of the relationships uration, as seen in Lemur and Tupaia, is primi- among "lower" primates is the structure of the tive for primates. This interpretation is tied to medial orbital wall. In both North American the multifaceted belief that "insectivores" are and European adapids in which this feature is generalized, primitive mammals, Tupaia repre- preserved, contact between the frontal and sents the ancestral primate condition, and maxilla separates the palatine from the lacri- Lemur is the most primitive primate; therefore, mal, while contact between the palatine and whatever Tupaia and Lemur share must have frontal separates the orbitosphenoid from the been present in the ancestral primate. Adapids maxilla (Clark, 1962; Gregory, 1920; Piveteau, do not have this configuration of the medial 1957). This configuration is also seen in the orbital wall. On the other hand, Cartmill (1975) indriines, Palaeopropithecus, Daubentonia, has argued that, since shrews and hedgehogs Plesiadapis, and occasionally in Phaner, Hapa- are probably closely related to primates, the lemur, and Lepilemur (Cartmill, 1975; Genet- pattern shared by these taxa reflects the primi- Varcin, 1963; Kollman, 1925; Tattersall and tive primate state, and that the pattern seen in Schwartz, 1974); it also characterizes many ex- adapids is primitive. However, while shrews, tant Lipotyphla-soricids, talpids, tenrecids, hedgehogs, and some primates show contact and chrysochlorids (Butler, 1956). In Archae- between the frontal and maxilla, soricids and olemur, while the maxilla separates the palatine erinaceids themselves do not share the same and lacrimal, the fronto-maxillary suture is total configuration of the medial orbital wall. appreciably longer-presumably as a result of Furthermore, the "diversely adapted Mad- secondary loss of the prenasopalatine portion of agascar lemurs," which Cartmill (1975, p. 340) the palatine (Kollman, 1925)-and the absence cited in support of his argument, include Pha- of this portion of the palatine permits a broad ner, Lepilemur, and Hapalemur, which varia- contact between the maxilla and orbito- bly develop the pattern seen in Lemur. Thus sphenoid, thus separating the palatine from the those taxa which consistently display the same frontal. In non-primates, this particular pattern conformation of the entire medial orbital wall is seen, for example, in Leptictis (representa- as do the adapids are the indriines, Pal- tive of the late Cretaceous-early Tertiary er- aeopropithecus, Plesiadapis, and non-erinaceid- notherian leptictids [McKenna, 1975]), as well solenodontid lipotyphlan insectivores; the pal- as erinaceid and solenodontid lipotyphlan insec- atine is much reduced in Daubentonia (Koll- tivores (Butler, 1956). man, 1925). Erinaceids, solenodontids, and 278 ANTHROPOLOGICAL PAPERS AMERICAN MUSEUM NATURAL HISTORY VOL. 55 leptictids share the same pattern with Archae- imply. The basis for the formation of this the- olemur. ory of relationships, and the uncertainties in- From the available evidence it is obviously volved, are perhaps best discussed by not easy to decide which of the above config- characterization of the branching-points on the urations of the medial orbital wall is primitive diagram. for Primates. In addition to the claims of Clark The common ancestor of the entire assem- (1962) and Cartmill (1975), Martin (1968) rea- blage under consideration (node 1 in fig. 1) was soned that the ancestral primate had the eth- highly derived in many ways relative to the moid exposed, as in Tarsius, Anthropoidea, ancestor it shared with the Lemur-Varecia most lorisids and cheirogaleids, and sometimes group. Its molars were characterized by buc- in marsupials, macroscelidids, and Dermoptera. colingual compression of the cusps and by Whether Clark's hypothesis or Martin's is cor- marked intercuspal cresting; in the lower mo- rect, the pattern seen in the adapids emerges as lars this cresting was interrupted lingually be- an autapomorphic condition. Cartmill's argu- tween the metaconid and the entoconid. The ment rests heavily on the assumption that talonid heel of M3 was elongate. The premolars shrews and hedgehogs are closely related to appeared in a posteroanterior sequence, at least primates. While this may be a viable hypothe- in eruption, and the anterior molars came in sis (Krishtalka, 1976 and personal commun.), early. The posteromedial aspect of the man- we believe it unwise to use preconceived no- dibular condyle was prolonged inferiorly to tions of phylogeny to determine the derived- provide a secondary articulation with the primitive polarity of the morphologies which postglenoid process. There was a marked poste- are then supposed to indicate relationships. rior convergence of the temporal lines. If wide distribution-"communality of pos- But if it is relatively easy to characterize the session" (e.g., Hennig, 1966; Schaeffer, Hecht ancestor represented by node 1 (a reflection of and Eldredge, 1972)-of a character among the the distinctiveness of the assemblage), there ex- taxa under consideration as well as in an array ist considerable problems in determining rela- of taxa of varied affinities bespeaks primitive- tionships within the group. The common ness, we must agree with Martin (1968) that the ancestor possessed a toothcomb as a primitive ancestral primate had the ethmoid exposed. On retention from an earlier ancestor (see fig. 2). the other hand, development of different con- Known adapines uniformly possessed orthally figurations of the medial orbital wall in mem- implanted anterior lower teeth, as did all the bers of the same genus (e.g., Phaner, notharctines with the possible exception of Lepilemur, Hapalemur) may reflect more accu- Pelycodus. If this derived trait is shared be- rately the futility of using this character com- cause of ancestral retention in the adapines and plex in phylogeny reconstruction. notharctines, then the relationships of these two groups are as expressed by node 2b (fig. 1). If Pelycodus indeed possessed procumbent ante- PHYLOGENETIC RELATIONSHIPS rior lower teeth, this would strongly suggest It is apparent from the preceding survey that that the primitive conformation had been lost the closest affinities of the adapids lie with the independently in the two groups. Such an inter- Lepilemur-Megaladapis-Hapalemur group. As pretation accords better with molar morphol- we have seen, some of the classical features ogy, which favors the phylogeny represented which have traditionally been focused upon by by node 2a (fig. 1). This common ancestor primate paleontologists do not help to clarify would have possessed further buccolingual the issue of adapid relationships; but it does compression of the molar cusps, and sharper seem to us that the dental evidence, in particu- intercuspal crests; it would also have shown the lar, is compelling. At this stage, unfortunately, development of a small hypocone on the upper the details of the relationships within this entire molars. It is the relationship expressed by node assemblage of fossil and living forms are not 2a (fig. 1) which we find to be the more plausi- entirely clear, as the dotted lines in figure 1 ble. 1979 SCHWARTZ AND TATTERSALL: ADAPIDAE 279

FIG. 1. Hypothesis of evolutionary relationships within Lemuroidea.

Node 3 (fig. 1) is the hardest of all to locate some adapines than to others. For instance, the within the scheme because Hapalemur is char- loss of the anteriormost premolar in Lepilemur, acterized principally by its retention of features Megaladapis, and Hapalemur may more specif- primitive for the clade (although, of course, it ically align them with Mahgarita, Caeno- is highly derived relative to its common ances- pithecus and Protoadapis, as opposed to Adapis tor with the Lemur- Varecia group). A striking and its four-premolared allies. This question autapomorphy resides in the loss in most Hapa- will be explored in a future, more detailed, lemur of the derived condition of the temporo- study. mandibular joint. But despite the fact that relationships within The common ancestor of Lepilemur and this assemblage of primates have yet to be Megaladapis (node 4 in fig. 1) had lost the worked out, there seems to be little question upper permanent incisor teeth, and possessed a about the alignment of the adapids with the highly unusual pattern of development and Lepilemur-Megaladapis-Hapalemur group, to eruption of the premolars whereby the develop- the exclusion of the other genera conventionally mental sequence P2 -* P4 P3 yielded to a assigned to Lemuridae. That this relationship sequence of eruption of P4 P3 -b P2. has not been recognized before is presumably We fully realize that comparison of genera largely due to the pervasive notion that the with higher taxa is an unsatisfactory process, Malagasy lemurs constitute a single homogene- although the reconstruction of ancestral mor- ous, monophyletic grouping-an assumption photypes helps avoid the problem. However, it which has only very recently begun to be chal- is possible that careful evaluation of relation- lenged. This realignment of relationships, al- ships within the Adapinae may reveal that the though apparently well-grounded, is thus Malagasy forms are more closely related to somewhat radical, and carries profound im- 280 ANTHROPOLOGICAL PAPERS AMERICAN MUSEUM NATURAL HISTORY VOL. 55 plications not only for the classification of the tors) which we know must have been extant at lemuriforms, but also for the times of phyletic that time. This is certainly more compelling divergence within the strepsirhines and for the than citing the absence of adapids in Africa as history of the geographical dispersion of these evidence for a southerly migration of strep- primates. sirhine ancestors; there exists at present no ap- While a popular view of mammalian, and in propriate African (or Malagasy) fossil primate particular primate, dispersal would have waves record. of northern forms migrating southward (e.g., Since Africa emerges as the only credible Van Couvering and Van Couvering, 1975; source of such invading forms, adapids must Walker, 1972), the implications of our sug- have been present on that continent at a time gested relationships among strepsirhines (fig. 2) when faunal interchange was still possible with instead favor the hypothesis that Africa was the Laurasia. While such interchange prior to the source of forms which subsequently invaded closure of the Tethys seaway in the early Neo- Madagascar and Holarctica. The importance of gene seems to have been possible during the Africa as a center of early primate diversifica- mid-late Eocene and mid-late Oligocene tion and dispersal is thrown into even more (Adrover and Hugueney, 1975; McKenzie, dramatic focus precisely because the well- 1970; Van Couvering, 1972; Van Couvering known Holarctic Paleogene primate fauna lacks and Van Couvering, 1975), the presence of many elements (particularly indriiform ances- adapids in the earliest Eocene of Euramerica

FIG. 2. Hypothesis of evolutionary relationships within Strepsirhini. 1979 SCHWARTZ AND TATTFERSALL: ADAPIDAE 281 had to have resulted from a northward invasion liest appearance of any adapid, which lends during the latest Cretaceous-early Paleocene added weight to our suggestion (Tattersall and regression (cf. Van Couvering and Van Cou- Schwartz, 1974) that the last common ancestor vering, 1975). If our theory of relationships of all the lemurs was very early in date, and is correct, the split between the Lemur- reinforces the hypothesis that the Malagasy pri- Varecia ancestor and the ancestor represented mate fauna is the result of multiple early invas- by node 1 in figure 1 occurred before the ear- ions, not of in situ diversification.

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In Vlth Congress Regional Committee on primates in relation to continental config- Mediterranean Neogene Stratigraphy, Pro- uration. In Bishop, W. W., and J. A. ceedings. Bratislava, Slovak Acad. Sci., Miller (eds.), Calibration of hominoid ev- pp. 363-367. olution. Edinburgh, Scottish Academic Walker, A. Press, pp. 195-218. 1972. The dissemination and segregation of early

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