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1989 Evolution of the Aeluroid Carnivora: Significance of the Ventral Promontorial Process of the Petrosal, and the Origin of Basicranial Patterns in the Living Families Robert M. Hunt Jr. University of Nebraska-Lincoln, [email protected]

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Hunt, Robert M. Jr., "Evolution of the Aeluroid Carnivora: Significance of the Ventral Promontorial Process of the Petrosal, and the Origin of Basicranial Patterns in the Living Families" (1989). Papers in the Earth and Atmospheric Sciences. 553. https://digitalcommons.unl.edu/geosciencefacpub/553

This Article is brought to you for free and open access by the Earth and Atmospheric Sciences, Department of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in the Earth and Atmospheric Sciences by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. NovitatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2930, 32 pp., 13 figs., 2 tables February 6, 1989

Evolution of the Aeluroid Carnivora: Significance of the Ventral Promontorial Process of the Petrosal, and the Origin of Basicranial Patterns in the Living Families

ROBERT M. HUNT, JR.' CONTENTS Abstract ...... 1 Introduction ...... 2 Acknowledgments ...... 3 Abbreviations ...... 3 Petrosal Form in the Living Aeluroid Families ...... 5...... The Auditory Region in the Oldest Known Aeluroid Carnivorans ...... 7 The Stenoplesictine Concept ...... 12 A New Stenoplesictine Basicranium from Quercy ...... 14 Petrosal Form in the Living Aeluroid Nandinia binotata ...... 15 The Proailurine Basicranium and Petrosal ...... 21 Origin of the Living Aeluroid Families ...... 23 Conclusions ...... 29 References ...... 30

ABSTRACT In studies of carnivoran phylogeny, the config- useful in tracing modem lineages to their first ap- uration and ontogenetic development of the au- pearances in the mid-Cenozoic. Pre-Oligocene ditory bulla enclosing the middle ear have been camivorans, however, lack preserved bullae and

I Research Associate, Department of Vertebrate Paleontology, American Museum of Natural History; Curator, State Museum and Department of Geology, University of Nebraska, Lincoln, Neb. 68588.

Copyright © American Museum of Natural History 1989 ISSN 0003-0082 / Price $3.70 2 AMERICAN MUSEUM NOVITATES NO. 2930 have been difficult to relate to later Cenozoic indicates that the form of Nandinia's petrosal groups. The carnivoran petrosal bone, because of closely approximates the petrosals of Oligocene its durability both in intact skulls and as an iso- stenoplesictine aeluroids from the Quercy fissues lated element, can be tracked through Cenozoic ofFrance. Stenoplesictines are the oldest generally time, and can supply new information on lineage acknowledged aeluroids represented by basicra- continuity through its geometry and spatial rela- nial remains. The strong anatomical correspon- tionships. dence shared by the Quercy stenoplesictine basi- Aeluroid carnivorans are united by a petrosal crania with the basicranium of Nandinia reflects of characteristic shape, distinguished by a ventral the plesiomorphic auditory structure of these promontorial process buttressing the lateral mar- groups. However, relative to stenoplesictines, gin of the basioccipital. The configuration of the Nandinia's auditory region is more primitive in process is highly uniform in most living aeluroids the structure ofthe auditory bulla and surrounding (viverrids, herpestids, hyaenids) but has been sup- basicranium; its basicranium is arrested at a pre- pressed in modem felids by encroachment of an Oligocene structural grade, and is representative inflated auditory bulla. Ancestral proailurine fe- of the projected ancestral aeluroid morphotype. lids, however, retain the process. Survey ofthe aeluroid fossil record suggests that Among living aeluroids, the African palm civet the modem aeluroid basicranial and bulla patterns Nandinia binotata is distinguished by a somewhat developed in the mid- to late Miocene, and were differently configured ventral promontorial pro- well established by the Plio-Pleistocene. Prior to cess, which is more posteriorly situated and ro- the mid-Miocene, an array of archaic basicranial bust. Comparison with fossil aeluroid basicrania patterns characterized the aeluroid Carnivora.

INTRODUCTION An improved understanding of the evo- and details of its structure are only conjec- lution ofthe mammalian order Carnivora has tural at the present time. The nature of au- resulted from detailed anatomical study of ditory bullae in early Tertiary carnivorans the auditory region in both living and extinct may be eventually clarified by study of re- lineages (Flower, 1869; Hough, 1948; Ted- cently discovered fully articulated skeletons ford, 1976). Ontogenetic elements forming ofEocene carnivorans from Messel for which the auditory bulla join to create unique mor- little postmortem disturbance is inferred phopatterns diagnostic of particular lineages (Springhorn, 1980, 1982, 1985). (Hunt, 1974a). Such lineages are most com- However, in the interim, a useful insight monly identified at the family level within comes from another quarter: identification of the order, and can be traced in some cases the small Holarctic Oligocene carnivoran Pa- into the Oligocene. laeogale as a relict lineage of the early Ter- The auditory bulla has not been found in tiary viverravids (Hunt, 1974b; see also Flynn pre-Oligocene fossil carnivoran basicrania, and Galiano, 1982) demonstrates the struc- and only rarely occurs in the extinct creodont ture ofthe auditory bulla in a member ofone carnivores (Mellett, 1977). This situation of the principal early Tertiary carnivoran seems to result from (a) a record of very few families (Viverravidae). Although the den- well-preserved skulls ofPaleocene and Eocene tition of Palaeogale exhibits aeluroid affini- Carnivora in paleontological collections; (b) ties (Flynn and Galiano, 1982), its unique probable loose attachment ofthe bulla to the single-chambered fully ossified bulla is not skull; (c) lack of ossification of some or all typically aeluroid, and suggests that early bulla elements. Tertiary viverravids probably did not possess Based on present knowledge of taxonomic the derived bulla configuration of the living distribution and morphology of the ontoge- aeluroid families. This conclusion raises sev- netic elements making up the auditory bulla eral important questions concerning the liv- in post-Eocene camivorans, we might rea- ing aeluroid groups: When did the modern sonably predict that Paleocene and Eocene bulla patterns develop? Can additional de- carnivorans possessed a multipart bulla rived traits be identified that unite the aelu- (formed by an ectotympanic and one or more roid families in addition to bulla configura- entotympanics) as in living forms. However, tion? such a bulla has yet to be discovered intact, Modern aeluroid bulla patterns appear in 1 989 HUNT: AELUROID CARNIVORA 3 the mid- to late Miocene, based on a review plesictine skulls (M1723, M1381) discussed of aeluroid basicranial patterns in the fossil in this report; to Dr. Guy Musser, American record, identified and summarized in this re- Museum of Natural History, New York, for port. From this conclusion it follows that one permission to study aeluroid carnivorans in should not expect to find aeluroid basicranial the Department of Mammalogy; to Drs. patterns typical of the living families in the Richard Tedford and Michael Novacek, early Tertiary. American Museum of Natural History, for Consequently, in the tracing of carnivoran access to fossil carnivorans in the Depart- lineages, it would be useful to identify other ment of Vertebrate Paleontology; and to Dr. features ofthe skeleton that can be employed Patricia Freeman for loan of carnivorans in as reliable anatomical markers in lieu of the the Zoology Division, University of Nebras- dentition and bulla. The basicranium ofCar- ka State Museum, Lincoln. I am grateful to nivora has been a fertile source of data for G. de Beaumont and Qiu Zhanxiang for use- phylogenetic studies in the past, so it is not ful discussions on fossil aeluroid carnivorans. surprising that a fresh examination of this My thanks to Marc Marcuson for preparation part ofthe skull, particularly the auditory re- of figures 7, 8, and 13. For review of the gion, might provide new insights. My recent manuscript, I appreciate the comments of work indicates that the form of the petrosal Harold Bryant, Patricia Freeman, Neil Land- bone has potential as a phyletic marker. In man, Bruce MacFadden, R. T. Schuh, and the Aeluroidea, a characteristic configuration Richard Tedford. of the petrosal promontorium can be iden- tified (Hunt, 1987: 44). It is found in repre- ABBREVIATIONS sentative species of all the major aeluroid families: A alisphenoid viverrids, herpestids, hyaenids, AC anterior carotid foramen primitive felids, and in the living primitive BO basioccipital aeluroid Nandinia. In addition, it appears that BS basisphenoid the petrosal configuration of aeluroids may Ca anterior crus of ectotympanic be responsible for the unusual double-cham- E caudal entotympanic bered arrangement of their auditory bulla, a EO exoccipital structural peculiarity that has been difficult F facet for ectotympanic on petrosal to explain. FR round window To demonstrate this, I first illustrate and G gonial discuss petrosal form in representative mem- H hypoglossal (condyloid) foramen bers of the living aeluroid groups; then de- ICA internal carotid artery L middle lacerate foramen scribe the structural parallels in petrosal and M mastoid basicranium between the oldest generally ac- P petrosal knowledged aeluroids (stenoplesictines) and PC posterior carotid foramen the living African aeluroid Nandinia, with PLF posterior lacerate foramen particular attention given to a recently iden- PP paroccipital process of exoccipital tified stenoplesictine basicranium in the Brit- R rostral entotympanic ish Museum; and, finally, compare these re- S intrabullar septum of hyaenids sults with basicranial structure in the SQ squamosal remaining group ofprimitive fossil aeluroids, T ectotympanic the I a V ventral process of the petrosal promonto- proailurines. conclude with survey of rium (VPP) aeluroid basicranial patterns in the fossil rec- X line of attachment of caudal entotympanic ord, focusing on the time of origin of these to ectotympanic patterns. AMNH Vertebrate Paleontology, American Museum of Natural History, New ACKNOWLEDGMENTS York AMNH-M Mammalogy, American Museum of I wish to express my appreciation to Dr. Natural History, New York Alan Gentry, British Museum of Natural BMNH Palaeontology, British Museum History, London, for loan of the two steno- (Natural History), London 4 AMERICAN MUSEUM NOVITATES NO. 2930

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Fig. 1. A, Basicranium of the viverrid Civettictis civetta (UNSM-ZM 14114), young adult, Kibwezi, Kenya, oblique ventral view, showing auditory bulla in place in right auditory region, and bulla removed from the left auditory region, revealing the ventral petrosal process (V). Dashed line indicates path of internal carotid artery through left auditory region. White scale bar in this figure and all subsequent photographs is one cm in length. For all abbreviations not defined in figure captions, see p. 3. B, Basicranium of same individual as A, but with ectotympanic and rostral entotympanic elements of the auditory bulla in place, forming an anterior chamber of-the bulla in front of the petrosal. This anterior chamber rests against the ventral petrosal process (V). Dashed line indicates zone of fusion between ectotympanic and the extreme anterior tip of caudal entotympanic; rostral entotympanic (R) is fused to 1 989 HUNT: AELUROID CARNIVORA 5

F:AM Frick Collection, Vertebrate Pa- the basioccipital-basisphenoid suture, and is leontology, American Museum of not anteriorly inclined as in the first group, Natural History, New York resulting in a more robust symmetrical pro- SAM South African Museum, Cape Town montorium. UNSM-ZM University of Nebraska State Mu- Lincoln Form ofthe Nandinia petrosal is strikingly seum, Zoology, similar to petrosal shape occurring in the old- PETROSAL FORM IN THE est generally accepted aeluroid skulls yet FAMILIES identified in the fossil record, the stenople- LIVING AELUROID sictines and proailurines from the Oligocene An aeluroid petrosal is characterized by a Quercy fissures of France (Teilhard, 1915; prominent ventral process or extension ofthe Piveteau, 1943; Lavocat, 1952). The simi- promontorium (fig. 1, V), forming the ven- larity in form and basicranial relationships tromedial edge of the petrosal. The process of Nandinia's petrosal to those of stenople- is best seen in ventral view with bulla re- sictines and proailurines is remarkable. It ap- moved. This ventral promontorial process pears that this is not convergence but the (VPP) is applied to the lateral edge of the result ofretention ofa primitive aeluroid pe- basioccipital, and often extends ventrad be- trosal structure in the living African palm yond the basicranial axis. The arctoid and civet. This trait accompanies numerous other cynoid Carnivora lack the process. Hence, plesiomorphic features which suggest that the the presence of VPP is an aeluroid synapo- structure ofthe auditory region ofthis animal morphy, and the common ancestor of the closely approximates the primitive aeluroid living aeluroid groups must have had a pe- morphotype (Hunt, 1987). trosal with VPP present. The evident correspondence among the In living aeluroids, where a broad sample living aeluroid groups in petrosal structure is ofrepresentative species can be surveyed, the demonstrated by the subsequent descriptions VPP takes one of two distinctive configura- and stereophotographs oftheir auditory anat- tions: omy: (1) In living viverrids, herpestids, and Viverridae. The viverrid petrosal (fig. IA, hyaenids, VPP inclines forward from the pro- B) is illustrated using the auditory region of montorium, and is applied to the basioccip- the bush civet Civettictis civetta. The petrosal ital immediately posterior to or at the basi- occupies the anterior auditory region, sepa- occipital-basisphenoid suture. Thus in these rating the middle ear from the cranial cavity. animals, VPP is situated in proximity to the In primitive Camivora, the posterior edge of anterointernal corner ofthe auditory region;2 the petrosal abuts against the exoccipital, but (2) In the primitive aeluroid Nandinia in viverrids like Civettictis, the petrosal is binotata, VPP is applied to the lateral edge separated from the exoccipital by a wide space of the basioccipital as in the other living ae- occupied by the caudal entotympanic. This luroids, but the process is not as anteriorly separation is due to the enormous enlarge- situated. It is placed closer to the midpoint ment and posterior extension of the caudal ofthe basioccipital's lateral edge, well behind entotympanic in living viverrids, relative to the primitive aeluroid condition in which no 2 Living felids lack the VPP. However, in the ancestral separation occurs. stock of the felids (Proailurinae), VPP is present, but more posteriorly situated relative to the living aeluroid Once the bulla has been removed from the groups. In later Cenozoic and living felids, the inflation skull (fig. 1A), the petrosal can be seen in of the auditory bulla via relative growth of the enlarged ventral view. It is a compact rugose bone with caudal entotympanic results in suppression of VPP, so a prominent ventral process, situated directly that loss ofthe process is a derivative trait ofthe younger lateral to the basioccipital-basisphenoid su- Felidae. ture. Anterior to the process is the middle the ectotympanic (T) rim, producing the thictic bulla configuration of viverrids. The petrosal has been slightly displaced to the left in the photograph so that the registration mark upon its anterior promontorial surface for the rostral entotympanic can be seen (to right and slightly below letter P). Stereopairs. 6 AMERICAN MUSEUM NOVITATES NO. 2930

Fig. 2. Basicranium ofthe viverrid Genetta genetta (AMNH-M 169064), female neonate, Molepolole, Bechuanaland, South Africa, basilar length of skull 53.6 mm, dp3-4 nearly fully erupted, medial view of left auditory region to show relations of ventral petrosal process (V) to anterior chamber (T, R) and posterior (E) chamber of bulla in neonatal viverrid. Ventral petrosal process protrudes below the basi- cranial axis (BO) at this early ontogenetic stage. Note lines of recent fusion between ectotympanic (T), rostral (R), and caudal (E) entotympanics, the three bony elements making up the auditory bulla in the adult. Rostral entotympanic is situated on the anterior slope of the ventral petrosal process. Stereopair. lacerate foramen for the entrance of the in- panic before turning mediad to enter the cra- ternal carotid artery into the cranial cavity. nial cavity at the middle lacerate foramen (fig. When the bulla is in place (fig. 1A), the ven- 1A). tral process is the only part of the petrosal This structural relationship of petrosal, that is exposed on the surface of the basicra- VPP, rostral entotympanic, and ectotympan- nium, its tip forming a tonguelike protrusion ic is constant in the viverrids. Of special sig- between bulla and basicranial axis. nificance is the fact that the rostral entotym- The relationship of the bulla's ecto- and panic and ectotympanic ossifications are entotympanic components to the ventral pe- spatially segregated anterior to the petrosal trosal process (VPP) is precise and of signif- promontorium by development of the ven- icance for the final adult form adopted by the trally extended process, resulting in devel- bulla. In figure 1B, the caudal entotympanic opment ofa separate anterior bulla chamber. and the posterior part of the ectotympanic This suggests that the form of the petrosal, have been removed from the auditory region especially during its early ontogenetic histo- to demonstrate the relationship ofthe rostral ry, could have been responsible for the entotympanic to VPP. Rostral entotympanic division of the bulla into two chambers. Pe- fits against the anterior slope ofVPP, and the trosal form seems to be a significant deter- ectotympanic is then fused to the edge ofros- mining factor in the bulla configuration tral entotympanic (this contact between ros- adopted by Viverridae. tral entotympanic and ectotympanic defines In early ontogeny in viverrids (fig. 2), VPP the thictic bulla ofViverridae-Hunt, 1987). is already developed, separating rostral from Together these two elements form the ante- caudal entotympanic, and marking the en- rior chamber of the viverrid auditory bulla. trance point ofthe internal carotid artery into Note that the internal carotid artery enters the auditory region. So it is reasonable on the auditory region immediately posterior to developmental grounds that petrosal form VPP, travels along its lateral face, then enters plays a role in the creation ofadult bulla pat- a partial tube or groove in rostral entotym- tern. 1 989 HUNT: AELUROID CARNIVORA 7

Herpestidae. The herpestid petrosal is sit- terior growth of ectotympanic beneath the uated much as in viverrids. In adult Her- caudal entotympanic chamber has not been pestes auropunctatus (fig. 3A, B), VPP pro- restricted in any way by the petrosal. In early trudes below the level ofthe basioccipital bone ontogeny of Crocuta (Hunt, 1974a, fig. 35), immediately posterior to the basioccipital- VPP protrudes somewhat ventrad below the basisphenoid suture. As in viverrids, the pro- basioccipital as in the adult, and the over- cess is well formed early in ontogeny (fig. 3A), growth of the petrosal by ectotympanic seen maintaining its shape and relationships with in the adult has already occurred. bulla and basicranial bones into the adult Felidae. In living felids, there is no evident stage. VPP. Despite the apparent absence of the The rostral entotympanic rests on the an- ventral process, felids parallel other living ae- terior slope of VPP in herpestids (figs. 3A, luroids in their dorsoventrally deep petrosal 4A) as in viverrids. Fused to the ventral edge dividing the auditory region into anterior and of rostral entotympanic is the ectotympanic posterior sections. In young lions, the pe- (thictic herpestid condition, Hunt, 1987), and trosal promontorium is prolonged ventrad as along the line ofcontact between the two ele- an elongate arcuate crest. However, were it ments runs the internal carotid artery en- not for the fossil record, the fact that felids closed in the adult in a bony tube (figs. 3A, originally had a VPP would not be known. 4A, B). The petrosal acts as a transverse bar- In the earliest true felids, assigned to Proai- rier within the auditory region, dividing an lurus (figs. 11, 12), the petrosal retains a anterior chamber (formed by rostral ento- prominent VPP. As evolution progresses from tympanic and ectotympanic, fig. 4A) from a Proailurus through Pseudaelurus to later fe- posterior chamber (formed by caudal ento- lids, the developing bulla suppresses the ven- tympanic, fig. 4B). tral process of the promontorium, largely by VPP in the neonatal herpestid (fig. 5) is a means of caudal entotympanic growth and prominent tongue of bone tightly pressed encroachment. Hence VPP is primitive for against the basioccipital's lateral edge about Felidae. at its midpoint. By contrast, in the adult her- pestid (fig. 3B), the well-developed process is positioned closer to the anterior end of basi- THE AUDITORY REGION IN THE occipital. A considerable length ofbasioccip- OLDEST KNOWN AELUROID ital occurs posterior to the process, indicating CARNIVORANS that relative growth during ontogeny poste- The oldest generally acknowledged aelu- riorly extended the basicranial axis and cau- roid carnivorans are represented by fossils dal entotympanic chamber of the bulla, par- from the Quercy fissures ofFrance. The Quer- alleling the growth pattern of bulla and cy district has produced the world's largest basicranial axis in viverrids (compare figs. 1 and most diverse sample of early aeluroids and 2 with figs. 3B and 5). from a single geographic area. The fossils can Hyaenidae. The hyaenid petrosal is devel- be placed for purposes ofdiscussion into three oped as in viverrids and herpestids, having groups, based on consensus common to the a prominent VPP extending ventrad to the scientific literature: (a) the stenoplesictine ae- level of, or slightly below, the basicranial axis luroids, Stenoplesictis and Palaeoprionodon, in living Crocuta (fig. 6). In other hyaenids, often considered viverrids (Teilhard, 1915; VPP is present, but the process can be located Piveteau, 1943, 1961; Bonis et al., 1973; deep within the auditory region hidden by Schmidt-Kittler, 1987) but placed by others the auditory bulla (e.g., Hyaena brunnea). within the Felidae (Beaumont, 1964; Gins- VPP is situated slightly posterior to the basi- burg, 1979); (b) the proailurine felids, Proai- occipital-basisphenoid suture in hyaenids, lurus and Haplogale, generally believed to however the strong posterior growth of cau- include the ancestry of the modern felid ra- dal entotympanic found in viverrids and her- diation; (c) the nimravid cats (Ginsburg, pestids is absent, and the distance posterior 1979), Nimravus, Quercylurus, Eofelis, and to the process is not great, remaining nearly Dinailurictis, first Old World representatives constant during ontogeny. The strong pos- of a mid-Cenozoic radiation ofcatlike mam- 8 AMERICAN MUSEUM NOVITATES NO. 2930

Fig. 3. Auditory region ofthe herpestid Herpestes auropunctatus: A, Internal (medial) view ofpetrosal- auditory bulla complex, removed from the skull, showing a developed ventral petrosal process (V) in both juvenile and adult; top, female adult (AMNH-M 189424), St. John, Virgin Islands, anterior to right; bottom, juvenile male (AMNH-M 59926), Nam Fong, Hainan, China, anterior to left. B, Basi- cranium of adult female in oblique ventral view, demonstrating-the protrusion of the ventral petrosal process (V) below the bones (BO, BS) of the basicranial axis (AMNH-M 239641), St. Croix, Virgin Islands. Stereopairs. mals on the northern continents, regarded by long 1 and Soumailles, are the oldest Euro- some workers as caniform Carnivora and not pean aeluroid carnivorans presently recorded aeluroids. Also, even though its skull is not in the paleontological literature. known, one should mention a problematical Geographic distribution ofthese fossils re- taxon, "Viverra" simplicidens, represented in veals that all are exclusively Old World species the Quercy district by dental remains dis- with the exception ofthe nimravid cats. Nim- cussed by Teilhard (1915). It was later syn- ravids occurred both in Eurasia and North onymized with Proailurus (Beaumont, 1964), America in the Oligocene, and extended into but recently has been listed as a distinct early the Miocene of Europe, north Africa, and aeluroid genus, Anictis Kretzoi, present at the North America, becoming extinct worldwide early Oligocene Quercy locality ofAubrelong in the late Miocene. Stenoplesictines oc- 1 (Bonis et al., 1973; Bonis, 1974; Sige et al., curred in the Oligocene of Europe (Teilhard, 1979: 44). Anictis, if in fact it is an aeluroid, 1915; Sige et al., 1979) and Asia (Hsanda and the nimravid Eusmilus, found at Aubre- Gol, Mongolia, Mellett, 1968), and have been 1 989 HUNT: AELUROID CARNIVORA 9

Fig. 4. Dissected auditory region of adult herpestid Herpestes auropunctatus (AMNH-M 239641), female, St. Croix, Virgin Islands, to show relationship of ventral petrosal process (V) to the auditory bulla: A, Basicranium in lateral view, bony floor removed from both anterior and posterior chambers of auditory bulla. Anterior chamber made up of ectotympanic (T) and rostral entotympanic (R) rests on petrosal, and displays contact between T and R (thictic herpestid state). A strongly inflated posterior chamber formed by caudal entotympanic (E) is situated behind the petrosal promontorium. Ventral petrosal process (V) is largely hidden by the bulla except where the process protrudes below the basioc- cipital; B, Basicranium in oblique ventral view, with the anterior chamber of the bulla (T, R) entirely removed, and only part ofthe caudal entotympanic remaining. Note the thin compressed ventral petrosal process (V) buttressing the lateral edge of basioccipital. Dashed line indicates path of internal carotid artery which runs within a bony tube formed by the medial wall of the bulla. Artery enters bulla at PC, exits bulla at AC, and enters cranium at middle lacerate foramen (L). Stereopairs. 10 AMERICAN MUSEUM NOVITATES NO. 2930

Fig. 5. Basicranium of the herpestid Herpestes (Xenogale) naso (AMNH-M 51089), male neonate, Medje, Zaire, basilar length 44.5 mm, showing developed ventral petrosal process (V) early in ontogeny, segregating rostral (R) from caudal (E) entotympanic bulla elements. Note junction of ectotympanic (T) and caudal entotympanic (E) to form incipient septum bullae. This spatial arrangement ofbulla elements (T, E) is believed to have been present in the hypothetical herpestid-hyaenid ancestral stock whose bulla configuration transformed along two ontogenetic pathways, one leading to the modem herpestid, the other to the hyaenid condition. Stereopair. recently reported from the early Miocene of World aeluroids justifies their placement in Africa (Schmidt-Kittler, 1987). They are un- two groups: (a) stenoplesictines and proai- known from the New World. Proailurines in lurines; (b) nimravids. Nimravids have been the strict sense are known only from the Old discussed in detail elsewhere (Hunt, 1987); World Oligocene and early Miocene, and in they appear to represent an early aeluroid fact are unknown outside Europe (Ginsburg, branch identified by a unique auditory bulla 1 979).3 pattern in which a true septum bullae is ab- Identification of only two fundamental sent, and the anterior wall of the bulla is basicranial patterns among these early Old formed by a nimravid autapomorphy: the overlap of an elaborate bony process of the 3 A unique felid skull found in 1935 in mid-Heming- fordian rocks of North America (Ginn Quarry, Nebras- ectotympanic and an anterior bony lamina ka, FAM 61847) approaches the proailurine grade of of the caudal entotympanic. Although a de- European authors; P2 is present, reduced but still double- veloped VPP is not present, the nimravid rooted. This cat is the oldest true felid known from the promontorium is slightly elevated, forming New World, and is much like Pseudaelurus transitorius in some individuals a medially situated bulge from the Burdigalian ofWintershof-West (Dehm, 1950), that may be a vestige ofthe process. Possibly the only Old World species of the genus in which P2 is the ancestral nimravid stock initially pos- still double-rooted. Dehm regarded this species as tran- sessed a developed VPP as in other aeluroids, sitional between Proailurus and Pseudaelurus, hence such but in time it became reduced in size. an animal might be considered a proailurine. The oldest The auditory bulla of stenoplesictines and known proailurines with unreduced double-rooted P2/ proailurines stands in contrast to the nim- p2 and Pl/pl present are not found outside Europe. Proailurines have not yet been reported in the recently ravid bulla. In stenoplesictines and proailu- excavated, stratigraphically controlled deposits in the rines, the bulla includes an anterior chamber Quercy district (Bonis et al., 1973; Crochet et al., 1981), largely formed by ectotympanic (with prob- although they are known from the older Quercy collec- able rostral entotympanic contribution), and tions (Teilhard, 1915). a posterior chamber formed by a caudal en- 1 989 HUNT: AELUROID CARNIVORA I1I

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.-

Fig. 6. Basicranium of the hyaenid Crocuta crocuta: A, Left auditory region of Pleistocene Crocuta (AMNH 18730), Wanhsien, China, ventral view. The ventral bony floor of the bulla has been removed (including the hyaenid internal septum) to reveal the ventral petrosal process (V) applied against lateral edge of basioccipital. Note caudal entotympanic fitted against the posterior slope of the promontorium, and rostral entotympanic resting on the anterior slope; B, Left auditory region of living Crocuta crocuta (UNSM-ZM 5012), Africa, ventral view. Ventral bony floor of bulla removed, showing ventral petrosal process applied to basioccipital. An opening has been cut in the hyaenid septum to reveal caudal entotympanic chamber dorsal to septum. Stereopairs. totympanic. Ectotympanic in these carnivo- 1952). Behind the anterior chamber, a pos- rans rests on the ventral promontorial surface terior chamber of very small size, enclosed of the petrosal where, in many specimens, a by a caudal entotympanic element, was un- facet is developed at the contact between pro- doubtedly present in stenoplesictines and montorium and ectotympanic rim (Lavocat, primitive proailurines. The caudal entotym- 12 AMERICAN MUSEUM NOVITATES NO. 2930 panic has never been found in place in a skull, rans. Teilhard's stenoplesictoid concept but its presence is documented by indenta- united a group of small Quercy Carnivora on tions registered upon the petrosal and sur- the basis of their similarly specialized shear- rounding bones ofthe auditory region. Earlier ing dentitions developed through parallel workers have suggested that the posterior evolution: "I unite, under the name Stenople- chamber was formed by a cartilaginous [cau- sictoides, a very homogeneous, yet very con- dal] entotympanic, and this indeed may have fused grouping of forms, generally small, been the case, or the entotympanic could as characterized by their common tendency to easily have been a loosely attached bony ele- develop a dentition, at least in the lowerjaw, ment. The double-chambered stenoplesic- approaching that of the Felidae: p3-4 trian- tine-proailurine bulla is similar in configu- gular and denticulate [with accessory cusps]; ration to the double-chambered bullae of ml high and trenchant, with metaconid and living aeluroids, and is a principal reason why talonid very reduced; m2 very small or ab- these groups have been considered aeluroids sent; canine trenchant and grooved." Teil- by earlier workers. hard explained his concept further: "Because If, as bulla morphology indicates, steno- the Stenoplesictoides tend to develop a very plesictines and proailurines are aeluroids, then simplified type of dentition, where conver- other aspects ofauditory anatomy should also gences are inevitable, one must make a spe- reflect this ancestry. In fact, the petrosal in cial attempt to understand, from certain the two groups is characterized by a promi- [morphological] details, often very small, the nent ventral process of the promontorium [evolutionary] pathways they have followed. (VPP) as in the living aeluroid families dis- . . . Such research has a better chance ofguid- cussed earlier. There can be little doubt that ing us toward a natural classification offorms these carnivorans are aeluroids as earlier than simply grouping the fossils on the basis studies have suggested. However, a close ex- ofthe phases ofreduction or ofsimplification amination of the petrosal promontorium in of the teeth." stenoplesictines and proailurines demon- Teilhard never intended to designate strates not only their near identity of form, stenoplesictoids as a formal taxonomic cat- but also their similarly developed VPP, which egory. It was a convenient morphologic is more posteriorly situated in these Oligo- grouping of lower dentitions that reflected cene skulls than in the living aeluroids. VPP similar feeding specializations: the high in these animals is a robust ventral contin- shearing carnassial, reduction ofthe rear mo- uation of the promontorium, not yet altered lars, and the development of multicuspate to the thinner bony lamina found in most tall, thin, catlike premolars. This morpho- living aeluroids, nor suppressed by bulla logic trend, often seen among carnivorans, of growth as in living felids. Furthermore, VPP convergence toward a shearing dentition is is more centrally located along the lateral edge typical of dental patterns that Crusafont and ofthe basioccipital bone, and is not displaced Truyols (1956) defined under their concept into the anterointernal corner ofthe auditory ofhypercarnivory. The original list ofgenera region as in many living aeluroids. placed in the stenoplesictoid division by Teil- Because both stenoplesictines and proai- hard included what today are understood to lurines are characterized by this type of pe- be primitive aeluroids (Stenoplesictis, Pa- trosal with ventral promontorial process, it laeoprionodon, Stenogale), primitive true fel- must be of considerable antiquity within the ids (Proailurus), and a viverravid (Palaeo- Aeluroidea. gale). Hence Teilhard's stenoplesictoid division constitutes not only an adaptive THE STENOPLESICTINE CONCEPT dental grade, but one so broad that it would be meaningless in a formal taxonomic sense. Although Stenoplesictinae was first used as Shortly thereafter, Schlosser (1923) unfor- a subfamily by Schlosser (1923), the concept tunately created the subfamily Stenoplesic- of an allied group ofgenera including Steno- tinae to encompass Teilhard's stenoplesic- plesictis first appeared in Teilhard's classic toid genera, a step at odds with Teilhard's monograph (1915) on the Quercy carnivo- original intent. Moreover, Schlosser failed to 1989 HUNT: AELUROID CARNIVORA 13 clarify the phylogenetic relations of these auditory bulla of Stenoplesictis was detailed small carnivorans. In fact, as Pilgrim (1931) for its time, including the best photographic later made clear, Schlosser gave no apparent illustrations of its auditory region yet pub- reason for this grouping. Despite the fact that lished. He (1943: 69) stressed the strong sim- well-preserved basicrania of stenoplesictines ilarity, in fact to him near morphologic iden- were known, and also that basicranial anat- tity (except in teeth), between the skulls of omy had been emphasized as an important Stenoplesictis and Palaeoprionodon. Later in tool in decoding carnivoran phylogeny by En- 1961, he employed the subfamily Stenople- glish zoologists (Turner, 1848; Flower, 1869; sictinae Schlosser 1923 for only Stenoples- Mivart, 1882a, 1882b), Schlosser placed his ictis and Palaeoprionodon, thus limiting Stenoplesictinae as a subfamily of the Mus- Schlosser's subfamily to just these two gen- telidae, even though the majority of the gen- era, and explicitly stated that these two gen- era considered by Teilhard and by Schlosser era "ont atteint le stade structural viverrien were, from the standpoint ofbasicranial anat- et doivent constituer une sous-famille de omy, aeluroid carnivorans. This preoccupa- Viverrides, les Stenoplesictines." Hence the tion with mustelid affinity was perpetuated studies by Teilhard, Pilgrim, and Piveteau by Helbing (1928) who called Proailurus a are the basis for the current placement of felinoid mustelid. stenoplesictines as Viverridae. Pilgrim (1931) attempted to remedy the In grouping Palaeogale with early aelu- situation by removing some of the "steno- roids such as Stenoplesictis and Palaeoprion- plesictoid" genera to living carnivoran fam- odon, Teilhard necessarily relied on the den- ilies. Pilgrim (1931, 1932) recognized that tal evidence at his disposal. These carnivorans Schlosser gave no valid phylogenetic criteria are simply convergent in their dentitions. for uniting the various stenoplesictine genera Teilhard was aware that two distinctive basi- into a subfamily, and argued that the similar cranial types were included in the stenople- features of these taxa, chiefly the teeth, "are sictoid group: the type "viverrienne" of Ste- no more than parallel adaptations or a mod- noplesictis and Palaeoprionodon, and the ified retention ofprimitive features, in which "weasel-like" basicranium ofPalaeogale fig- case the various stenoplesictine genera should ured earlier by Filhol [Teilhard had only low- properly find a place among the primitive er jaws of Palaeogale from Quercy but was members ofthat modern family to which they aware of the basicranial structure from a most appropriately belong." He placed specimen of this genus figured by Filhol- at Proailurus and Stenogale4 in the Proailuri- that time Teilhard (1915: 70) had knowledge nae, a subfamily of primitive felids, and re- ofbut had not seen the basicranium ofSteno- garded Stenoplesictis and Palaeoprionodon as gale]. primitive viverrids. Today it is clear that, as Teilhard had sup- In 1943, following a thorough study of a posed, stenoplesictoids comprise several di- complete skull of Stenoplesictis, Piveteau verse branches. Basicranial evidence con- concluded, as had Pilgrim, that the steno- firms the wide separation of the viverravid plesictoid group should be subdivided. Pive- Palaeogale from the early aeluroids Steno- teau (1943: 71) thought the similarities be- plesictis and Palaeoprionodon; the affinities tween the auditory bulla of Prionodon and of Stenogale remain to be demonstrated on Stenoplesictis were sufficiently strong to place basicranial evidence, but if the holotype of the latter as a viverrid. His description ofthe "Proailurus" julieni is a Stenogale (Teilhard, 1915), then its basicranium indicates that it 4The phylogenetic position ofStenogale is not clearly is an early aeluroid (see Lavocat, 1952). The resolved even in the more recent literature (compare creation of a subfamily by Schlosser from Bonis, 1973; Sige et al., 1979 with Beaumont, 1964; Ginsburg, 1979). In this report my concept ofStenogale Teilhard's dentally based stenoplesictoid is based upon the holotype skull and jaws of S. julieni, group was premature, since at that time the figured by Viret (1929, pl. 12, figs. 5, 8, 9), which pre- true affinities ofthe member genera were not serves an aeluroid auditory region having a petrosal with clear. However, ifPalaeogaleis removed from the ventral promontorial process. Hence I regard this Teilhard's initial grouping, then only primi- species as an early aeluroid. tive aeluroids sharing a close relationship re- 14 AMERICAN MUSEUM NOVITATES NO. 2930 main: Stenoplesictis, Palaeoprionodon, are aeluroid, not felid, specializations. If we Proailurus, Stenogale. The basicrania ofthese turn to the stenoplesictine basicranium, it four genera are morphologically very similar. seems probable that the similarities shared Beaumont (1964) placed all four genera in with felids are primitive features, such as the a felid subfamily Proailurinae, comprising the shape of the petrosal. At present, only Pa- tribes Stenoplesictini (Stenoplesictis, Palaeo- laeoprionodon among stenoplesictines ap- prionodon) and Proailurini (Proailurus, pears to have a phyletic connection with liv- Stenogale). Ginsburg (1979) adopted a sim- ing viverrids. However, both European ilar arrangement but used a slightly different stenoplesictine genera could eventually prove taxonomy in which a felid subfamily Felinae to be most closely related to Viverridae. Based includes a tribe Proailurini made up of two on the present uncertain state of our knowl- subtribes, one with Stenoplesictis and Pa- edge, I conclude only that stenoplesictines are laeoprionodon, the other with Proailurus, primitive aeluroids, not felids, and that while Stenogale, and Haplogale.5 However, wheth- a relationship of certain stenoplesictines to er one employs the proailurine taxon at the viverrids is probable, it remains to be con- level of subfamily or tribe, the category re- firmed. mains an aggregation of genera primarily based on plesiomorphic features, hence a A NEW STENOPLESICTINE paraphyletic group. Since it is not the intent BASICRANIUM FROM QUERCY ofthis study to establish the detailed cladistic relationships among these forms, I follow Among the most plesiomorphic ofthe early Beaumont and Ginsburg in recognizing a re- aeluroids in terms ofbasicranial structure are lationship between Stenoplesictis and Pa- the stenoplesictines Stenoplesictis and Pa- laeoprionodon on the one hand, and among laeoprionodon. Basicranial structure of Proailurus, Haplogale, and Stenogale on the stenoplesictines is well preserved only in other. I shall call the first two genera "steno- skulls from Quercy, all maintained in Euro- plesictines," and the last three "proailu- pean museums (table 1). The skull, including rines," without formal taxonomic implica- the basicranium, of Stenoplesictis was de- tions, emphasizing that placement of all five scribed and figured by Piveteau (1943: 6-8), genera in the Felidae is inappropriate. It is and the skull ofPalaeoprionodon by Teilhard sufficient to note that only Proailurus (in- (1915: 77-78). These are small carnivorans cluding Haplogale) by current consensus with skull lengths ofabout 7 to 9 cm. Teilhard in (1915: pl. IX, figs. 10, 14, text-fig. 13) illus- seems to be involved the ancestry of the trated two skulls ofPalaeoprionodon laman- true felids (Ginsburg, 1983), whereas the re- dini, both in the Paris Museum, one showing maining genera (Stenoplesictis, Palaeoprion- the basicranium and auditory bulla. Later, odon, Stenogale) appear to include species pl. I, closely related to viverrids, as well as other Piveteau (1943: fig. 3, 3A, text-figs. 2, in 3) illustrated a well-preserved skull, also with early experiments aeluroid evolution that basicranium and bulla, of Stenoplesictis cay- left no known descendants. Therefore, reten- luxi, in the collection of the Faculty of Sci- tion ofa subfamily (Stenoplesictinae) or tribe ences, Marseille. (Stenoplesictini) within Felidae for the I am able to confirm Piveteau's description primitive aeluroids Stenoplesictis and Pa- ofthe auditory region ofStenoplesictis through laeoprionodon is not supported by reliable study ofa partial skull (BMNH M 138 1) from synapomorphies: the hypercarnivorous den- Quercy in the British Museum, discovered titions ofStenoplesictis and Palaeoprionodon during a research visit in 1987. Despite the are only convergent on felids as Teilhard and absence of the teeth and palate, M138 1 rep- others were well aware. The dental special- resents the skull of a small stenoplesictine, izations that are present in these two genera almost certainly Stenoplesictis cayluxi: it is 5 Haplogale is considered a separate proailurine genus nearly identical to the Marseille skull of S. by Ginsburg (1983), and by Beaumont (1961) a synonym cayluxi figured by Piveteau (1943). Although of Proailurus. The oldest proailurines are probably rep- the auditory bullae are missing, the auditory resented by the Quercy sample of Haplogale media. region of Ml 38 1 is unquestionably that of a 1 989 HUNT: AELUROID CARNIVORA 15

TABLE 1 compared in figures 7-10 and in the subse- Basicrania of Early Aeluroid Carnivorans quent discussion. Exclusive of Nimravids A second stenoplesictine skull (Ml 723) in STENOPLESICTIS the paleontological collection of the British 1. Skull of Stenoplesictis cayluxi, Collection de Museum, labeled as Stenoplesictis cayluxi, is Geologie, Faculte des Sciences de Marseille also from Quercy, and was originally figured (Piveteau, 1943). by Lydekker (1885: 97, fig. 9); the specimen 2. Skull of Stenoplesictis cayluxi, collection of A. includes an upper dentition in association Muller, Leipzig, P2004 (Fischer, 1983). with the anterior part ofthe basicranium, in- 3. Skull of Stenoplesictis, not illustrated (Filhol, 1882: 66-67, listed in Fischer, 1983). cluding the petrosal. Lydekker's skull, despite 4. Skull of Stenoplesictis, collection of the British its attribution to Stenoplesictis, is much clos- Museum (Ml 381), not previously described. er in size, skull form, and dentition to the PALAEOPRIONODON stenoplesictine Palaeoprionodon figured by 5-6. Two skulls of Palaeoprionodon lamandini, col- Teilhard (191 5, figs. 10, 14), and it is referred lection of the Paris Museum (Teilhard, 1915: here to this latter genus (table 1). Compara- pl. 9, figs. 10, 14). tive measurements of skull and basicranium 7. Skull of Palaeoprionodon sp., collection of the British Museum (Ml 723) (Lydekker, 1885: 96- in representative stenoplesictines and proai- 98, fig. 9). lurines are presented in table 2. 8-9. Two skulls of Palaeoprionodon, Munich Mu- seum (Schlosser, 1889: 146). PETROSAL FORM IN THE LIVING Additional skull material ofstenoplesictines is AELUROID NANDINIA BINOTA TA conserved at Montauban, Louvain, Lyon, and Basel (G. de Beaumont, personal commun., In a previous discussion of the phyloge- 1985). netic position of Nandinia (Hunt, 1987), I PROAILURUS suggested that numerous plesiomorphic traits 10. Skull ofholotype ofProailurus lemanensis, Paris found in this small African arboreal civet in- Museum (Filhol, 1879: pl. 26). dicate its retention of a primitive aeluroid STENOGALE morphology for much ofCenozoic time. The 11. Skull of holotype of "Proailurus" julieni, Paris animal's auditory bulla, in fact its entire au- Museum, Julien collection (Filhol, 1879: 192, pl. ditory region, is morphologically what one 27, figs. 5, 6, 8, 13; Viret, 1929: pl. 12, fig. 5; would predict in a stem aeluroid, a species Lavocat, 1952: fig. 2). that most probably existed in the early Ce- nozoic. An important morphologic feature men- stenoplesictine camivoran. To demonstrate tioned briefly in my earlier presentation the close correspondence ofpetrosal structure (Hunt, 1987: 44) was the configuration ofthe between the British Museum stenoplesictine petrosal in Nandinia. Here I wish to amplify (Ml 381) and the skull of Nandinia, they are that point. Unlike the majority of living ae-

TABLE 2 Comparative Dimensions (mm) of Stenoplesictine and Proailurine Skulls Width between Width between Greatest width condyloid alisphenoid Taxon Basilar length of braincase foramina canals Stenoplesictis (Marseille) 83.4 28.3 11.3 10.5 Stenoplesictis (M1381) (80-85) 30.6 12.0 12.8 Palaeoprionodon (Paris Mus.) 73.2 28.6 10.3 10.2 (77.6) 28.6 - - Palaeoprionodon (M1723) (70) 27.0 (10) (9.5) Stenogalejulieni (72-73) 25 10 (8.1) Proailurus lemanensis (130) 46 15.7 20.6 () = estimated. 16 AMERICAN MUSEUM NOVITATES NO. 2930

British Museum (Ml 381), and with the pe- trosal retained in the British Museum skull (M1723) of Palaeoprionodon figured by Ly- dekker (1885). Comparisons also were made with descriptions and illustrations of steno- plesictines and proailurines in the literature (table 1). Nandinia's petrosal promontorium retains the robust centrally situated VPP of these early aeluroids, buttressing the edge of the basioccipital. As in stenoplesictines, VPP is applied to the lateral edge ofthe basioccipital about midway along its length. Stereopho- tographs ofthe basicranium ofNandinia (figs. 9A, 1OA) demonstrate that VPP extends ven- trad beyond the plane of the basioccipital; VPP is similarly extended in the stenople- sictine skull (figs. 9B, 10B) but the extension is less obvious due to ventral downturning of the edge of the basioccipital, a condition of the basicranial axis found in stenoplesictines but not pronounced in Nandinia. Anterior to VPP, the surface of Nandinia's promonto- rium forms a smooth, slightly concave slope. On the medial surface of this slope rests the rostral entotympanic (fig. 9A). This slope is Fig. 7. Plesiomorphic features of the auditory similarly configured in the stenoplesictine; region of the living aeluroid Nandinia binotata however in the extinct aeluroid, only frag- (shown in lateral view, anterior to left, bulla re- ments of bone are present at the location of moved) are remarkably similar to equivalent fea- rostral entotympanic. tures of the auditory region of Oligocene steno- Of particular interest is a facet on the plesictine aeluroids (see fig. 8), suggesting the promontorium just anterior to the round antiquity of auditory pattern in Nandinia. Nan- window in the stenoplesictine which has no dinia's petrosal displays both the same general form counterpart in Nandinia. As indicated by and primitively configured ventral petrosal pro- Lavocat (1952), this receives the cess as seen in stenoplesictines. The small size and facet flanged rudimentary development ofcaudal entotympanic edge ofectotympanic where it is seated upon occur in both groups (attachment of caudal en- the promontorium. This is a derived trait; totympanic to petrosal and rostral entotympanic the unfaceted state seen in Nandinia is re- indicated by stippled pattern). Transpromontorial garded as more primitive. In Nandinia the path of the internal carotid artery in Nandinia, posterior ectotympanic rim is very close to running in proximity to the medial wall of the the promontorium but does not rest upon it bulla, is believed to be the primitive course of the or create a facet. artery for Aeluroidea. In living aeluroids, the ectotympanic usu- ally contacts the promontorium adjacent to luroids, Nandinia retains a petrosal shaped the round window where a similar facet often like that of Oligocene stenoplesictines. Pe- is developed (e.g., viverrid Civettictis, her- trosal form appears to have been arrested in pestid Paracynictis, felid Lynx). Because of Nandinia at a phylogenetic stage comparable the apposition ofpetrosal promontorium and to that found in primitive stenoplesictine ectotympanic rim in Aeluroidea, the prom- basicrania from Quercy. ontorial facet is commonplace. Nandinia's petrosal (fig. 7) was compared The development ofa strong promontorial with the petrosal of the recently identified facet correlates with the slightly inflated con- stenoplesictine basicranium (fig. 8) in the dition ofectotympanic documented by Pive- 1 989 HUNT: AELUROID CARNIVORA 17 teau (1943) in Stenoplesictis. In stenople- sictines the ectotympanic is somewhat advanced in its slight to moderate degree of inflation or chambering, and has progressed beyond the primitive nearly planar form pre- dicted in the hypothetical aeluroid ancestor, and closely approached by Nandinia. Nan- dinia remains the most primitive known ae- luroid in this respect in that its ectotympanic is only slightly, if at all, modified from the plesiomorphic aeluroid state, and does not contact the promontorium (fig. I OA). How- ever, Nandinia's ectotympanic most closely approaches the petrosal promontorium at the point on that bone where the facet appears in stenoplesictines, and it is evident that in- flation ofthe ectotympanic could lead to con- tact with petrosal and development of such a facet. Hence the morphology of ectotym- panic rim and petrosal promontorium in Nandinia is arrested in a structural stage that immediately foreshadows the faceted con- dition seen in many living and extinct aelu- roids. In both Nandinia and the stenoplesictine (fig. 1OA, B), the ventrally extended petrosal promontorium divides the auditory region Fig. 8. Auditory region of the extinct Oligo- into two parts, resulting in the segregation of cene stenoplesictine aeluroid Stenoplesictis cay- bulla chambers common to the aeluroid luxi (BMNH M138 1, lateral view, anterior to left, group. In the stenoplesictines, this segrega- bulla removed), from the Quercy district, France, tion is more complete, primarily due to the for comparison with the auditory region of the large dorsoventrally deep petrosals relative living aeluroid Nandinia in figure 7. Stenoplesictis to the small skulls of these animals. The rel- differs in having slightly greater inflation of ecto- ative volumes ofthe bulla chambers also cor- tympanic, resulting in deeper registration marks respond in Stenoplesictis and Nandinia: the for ectotympanic on squamosal and alisphenoid, in and in a prominent facet on the petrosal (F) where posterior chamber is small volume, the the ectotympanic rim makes contact with the pro- plesiomorphic aeluroid condition. montorium. Stipple pattern shows that line of at- Futthermore, the attachment ofcaudal en- tachment for rudimentary caudal entotympanic is totympanic's posterior margin, whether car- developed as in Nandinia. Path ofinternal carotid tilage or bone, produces in Nandinia and in artery restored in conformity to the path known stenoplesictines a diagnostic registration on in Nandinia. the posterior margin ofthe petrosal, best de- scribed as a shallow trough or depression (this identical, bordered by a flanged edge which line of attachment is indicated by stipple in marks the anterior limit of caudal entotym- figs. 7, 8; see also fig. 9A, B, black triangles). panic attachment (figs. 7-10). Lack of caudal In living Nandinia, we know from direct ob- entotympanic inflation in Nandinia and the servation that this trough receives the only stenoplesictine is demonstrated by the very slightly inflected edge of the cartilaginous small space available for the posterior cham- caudal entotympanic (Hunt, 1987, fig. 6). ber of the bulla, and is also indicated by the There is no doubt that this is its function in primitive posteroventrally directed paroccip- stenoplesictines as well, because the config- ital processes present in both animals (fig. 9A, uration of this trough in both Nandinia and B) which fail to contact the bullae. In living the British Museum Stenoplesictis is nearly aeluroids with an enlarged caudal entotym- 18 AMERICAN MUSEUM NOVITATES NO. 2930

Fig. 9. A, Basicranium of the living aeluroid Nandinia binotata (AMNH-M 51492), young adult female, Niapu, Zaire, oblique ventral view, bulla removed from left auditory region, except for rostral entotympanic, and only cartilaginous caudal entotympanic removed from right auditory region. B, Basicranium ofextinct stenoplesictine aeluroid Stenoplesictis (BMNH M138 1), Quercy, France, oblique ventral view, showing bulla removed from auditory region, except for rostral entotympanic fragments. Note in both A and B the correspondence in form ofpetrosal, ventral petrosal process (V), and the small area posterior to petrosal for caudal entotympanic. A similar lin-e of attachment (indicated by six black triangles in each photograph) ofcaudal entotympanic to the petrosal is found in both ofthese carnivorans. Asterisks mark registration ofectotympanic on squamosal and alisphenoid. Stereopairs (negatives printed in reverse for comparison with fig. 12). panic (e.g., figs. 1, 4), the paroccipital process the tensor tympani fossa, epitympanic recess, is contacted by andjoined to the inflated bul- bony canal for the facial nerve, mastoid ge- la, and can become quite thin and expanded ometry, and the configuration of the sur- over the posterior surface of the caudal en- rounding basicranial bones, are very similar totympanic. in Nandinia and Stenoplesictis. Other details of petrosal form, including The path of the internal carotid artery is 1 989 HUNT: AELUROID CARNIVORA 19

Fig. 10. A, Basicranium of Nandinia binotata (AMNH-M 51492), same individual as figure 9A, ventral view, bulla removed from right auditory region, except for rostral entotympanic, and only cartilaginous caudal entotympanic removed from left auditory region. B, Basicranium of Stenoplesictis (BMNH Ml 381), same individual as figure 9B, ventral view, bulla removed from right auditory region except for rostral entotympanic fragments. Note that the ventrally expanded petrosal tends to divide the auditory region into anterior and posterior parts. Asterisks mark registration of ectotympanic on squamosal and alisphenoid. Stereopairs. confirmed in Nandinia by earlier work of inferred for the artery in stenoplesictines (fig. Chapuis (1966) and Wible (personal com- 8). In Nandinia, the artery traverses the mid- mun., 1983). This path is restored in figure dle ear cavity in proximity to (lateral to) the 7 wherein the artery travels lateral to VPP on medial wall ofthe bulla; the path ofthe artery the promontorium, then runs along the lat- is not within the bulla wall, and in fact passes eral face of rostral entotympanic, and turns through it at only one point, just posterior to mediad at the anterior end of rostral ento- VPP, where it enters the middle ear cavity. tympanic to enter the cranial cavity at the This route probably represents the primi- middle lacerate foramen. A similar course is tive course of the internal carotid in the Ae- 20 AMERICAN MUSEUM NOVITATES NO. 2930

Fig. 1 1. Cast of basicranium of adult proailurine felid Proailurus lemanensis (Paris Museum 1903- 20), St.-Gerand-le-Puy (Allier), France, Aquitanian, holotype, ventral view, bulla removed from auditory region. Petrosal form, including the ventral petrosal process (V), is similar to that of Stenoplesictis (compare fig. 9B) but in the proailurine the basicranial bones are emarginated by development of an enlarged caudal entotympanic (white arrows indicate extent of caudal entotympanic encroachment). Strong registration of ectotympanic on auditory region results in a petrosal facet (F) on promontorium and in cavities impressed in squamosal and alisphenoid (asterisks); these same features occur in Steno- plesictis (compare fig. 1OB). Early felids (Proailurus) retain the ventral petrosal process ofthe Aeluroidea but it is lost in living felids due to apparent suppression by the enlarging bulla. Stereopair. luroidea. All viverrids maintain this primi- to the ventral edge of rostral entotympanic tive course, modifying it in a few species in therefore results in complete enclosure ofthe which the artery is enclosed for a short dis- artery within the medial wall of the bulla. tance in a bony tube formed by rostral en- Significantly, the observed courses of the ar- totympanic. This tube forms by lateral ex- tery found in the living aeluroid families, de- tension of the rostral entotympanic around spite what at first appear to be rather different the artery as it passes the lateral face of the arterial paths through the auditory region, are element, so that in fact the arterial path re- in fact basically similar when studied closely, mains the same in all viverrids but enclosure and can be derived from a common mor- of the artery by rostral entotympanic is vari- photypic pattern like that of Nandinia. able (both among and within species). Hyae- In aeluroids, the primitive course of the nids and felids apparently both maintain the internal carotid artery was transpromonto- primitive course, but because the artery is rial, but the path of the vessel lies close to reduced or vestigial in most living members the medial wall ofthe bulla within the middle ofthese families, this is not entirely resolved. ear, and was not diverted far laterad on the In herpestids, on the other hand, the course promontorium. The association ofthe artery of the artery is modified (perbullar carotid, with the bulla wall is so close that in herpes- Hunt, 1987: 45): the vessel runs in a bony tids it becomes incorporated in the bony wall. tube between ectotympanic and rostral en- The artery's proximity to the bulla wall is in totympanic, never entering the middle ear fact an expression of the close association of cavity. In herpestids the artery is situated on the internal carotid with the rostral entotym- the ventral edge ofrostral entotympanic, and panic in all Carnivora (Hunt, 1974a). does not seem to be diverted as far laterad Nandinia is thus confirmed as a primitive as in viverrids. Application of ectotympanic aeluroid based on the morphology of its au- 1 989 HUNT: AELUROID CARNIVORA 21

I I'd BS p BO

Fig. 12. Cast of left auditory region of Proailurus lemanensis, same individual as figure 1 1, lateral view, bulla removed. Note ventral petrosal process (V), deep emargination ofbasi- and exoccipital bones by the bulla, and elliptical promontorial facet (F) produced by the ectotympanic. Asterisks indicate registration of ectotympanic on squamosal and alisphenoid. Proailurus represents a transitional mor- phologic stage in the evolution of the felid auditory region between stenoplesictines and modern felids. Stereopair. ditory region, both in terms ofpetrosal struc- longs to Proailurus lemanensis, a cat with ture and relationships, auditory bulla pattern, basilar length of skull estimated at 13 cm, and internal carotid relations. The degree of about the skull size of the living lynx; the plesiomorphy of Nandinia's auditory region second is the holotype skull of Stenogaleju- is greater than that found in the oldest known lieni, a smaller cat with basilar length esti- confirmed aeluroids, the Oligocene steno- mated at a little more than 7 cm, slightly plesictines, suggesting that Nandinia is a liv- smaller than a domestic cat skull. I have ex- ing representative ofthe basal aeluroid stock amined a cast (figs. 1 1, 12) ofthe basicranium whose origin must predate the Oligocene di- of Proailurus lemanensis from the Aquita- versity of stenoplesictine and proailurine ae- nian of St.-Gerand, France, originally de- luroid carnivorans documented by the Eu- scribed by Filhol (1879: pl. 26, figs. 2-4, 6- ropean fossil record. 8); and have studied photographs ofthe skull of Stenogalejulieni Filhol in Viret (1929: pl. 12), reported to come from the Aquitanian THE PROAILURINE of Chaveroche-sur-Besbre. Neither of these BASICRANIUM AND PETROSAL skulls retains the auditory bulla. Proailurine aeluroids share near identity of As Lavocat (1952) pointed out, these basi- basicranial structure with the stenoplesic- crania are exceedingly similar, not only to tines, indicating that the two lineages are each other but to the basicranium of Steno- closely allied. However, early in their history plesictis (compare figs. 11 and 12 with figs. the proailurines evolved a highly derived au- 9B and 1OB). In fact, the basicrania ofSteno- ditory bulla of felid type, in which an en- plesictis and Stenogalejulieni are more alike larging caudal entotympanic encroached upon than are those of Proailurus lemanensis and and indented the basioccipital medial to VPP S. julieni because the auditory bullae in the (fig. 1 1). By the Aquitanian (early Miocene), first species pair have not developed the strong proailurines have markedly diverged from the inflation and encroachment on the surround- primitive aeluroid auditory structure dis- ing basicranial bones (basioccipital, exoccip- played by Nandinia. ital, mastoid) that occurs in the more ad- I know of only two proailurine skulls in vanced proailurine Proailurus lemanensis which the basicranium, including the petro- from St.-Gerand. Proailurus lemanensis dif- sal, is well preserved (table 1): the first be- fers primarily in the additional registration 22 AMERICAN MUSEUM NOVITATES NO. 2930

marks impressed upon the basicranial bones concolor, F. tigrina, and Lynx rufus, the ec- surrounding the auditory region by the en- totympanic rim or flange is directly applied larged bulla (fig. 11, white triangles). to the promontorium, just anteroventral to If a series of Miocene felid skulls is com- the round window. In lions (Panthera leo) I pared, beginning with the Aquitanian basi- have examined, the ectotympanic rim is crania of the proailurines Stenogale julieni raised a little distance above the petrosal, and and Proailurus lemanensis, and continuing there is no contact. As a result, in living felids, through Hemingfordian and Barstovian the expression of the promontorial facet Pseudaelurus in North America, the gradual ranges from moderately developed to absent. enlargement of the bulla and its encroach- However, it is apparent that most living ment on the bones of the basicranial axis, felids maintain a tight registration between especially basioccipital, are clearly evident. petrosal and ectotympanic rim, resulting in The stenoplesictine basicranium could be a faceted promontorium similar to the type placed as the first and most plesiomorphic found in proailurines and stenoplesictines. stage in this series, since the basicranium lacks During evolution oflater felids from a proai- any significant encroachment of the bulla on lurine, this close ectotympanic-petrosal con- surrounding basicranial bones. Stenogaleju- tact has been retained in many of the de- lieni is very little evolved beyond the steno- scendant species, despite modifications in plesictine condition, and shows no strong petrosal shape. emargination of surrounding basicranial The existence of the flanged and inflated bones by the bulla (Lavocat, 1952); P. le- ectotympanic in early aeluroids is of more manensis (figs. 11, 12), however, is clearly than passing interest. The structural prereq- evolved beyond the more plesiomorphic au- uisite in aeluroids for the formation of the ditory region of S. julieni, and heralds the diagnostic septum bullae is an ectotympanic more advanced bulla and basicranium of resting on the promontorium (Hunt, 1987). Pseudaelurus and the later felids. A mono- An important shared derived character of phyletic felid clade can be identified using as living viverrids, herpestids, felids, and a synapomorphy the medial encroachment of hyaenids is the application of the rim of ec- the bulla on the basioccipital (fig. 11); this totympanic to the promontorium, which clade would include Proailurus lemanensis, closes off an anterior bulla chamber; the ab- Pseudaelurus, and their extinct descendants, sence ofthis contact in lions and some hyaen- as well as the living felids. as appears to be a secondary condition in The proailurine petrosal, including the which the ectotympanic has pulled away a configuration of the robust ventral promon- short distance from the promontorial surface. torial process, is identical to that of steno- Against the intrabullar partition formed by plesictines (compare figs. 9B and 12). Just as the posteromedial wall of this often inflated in stenoplesictines, VPP buttresses the basi- ectotympanic, an anterior surface ofthe cau- occipital in primitive proailurines like S. ju- dal entotympanic is applied. The resulting lieni, but in the more advanced P. lemanen- double-layered internal septum is exclusive sis, the emargination ofthe basioccipital bone to aeluroids.6 by the bulla creates a space between VPP and 6 A true septum bullae occurs only in aeluroids, but the basicranial axis. In both proailurines and not all aeluroids (as recently defined, Hunt, 1987) possess stenoplesictines, the promontorium of the one. Primitive aeluroids such as nimravids, Nandinia, petrosal is faceted to receive the ectotym- and possibly stenoplesictines, lacked the septum. The panic (Lavocat, 1952). aeluroid septum results from a unique juxtaposition of Both stenoplesictines and proailurines pos- the surfaces of the caudal entotympanic and ectotym- sess a flanged ectotympanic. The flange is de- panic. In arctoid and cynoid camivorans, the edges (not veloped on the posteromedial rim of ecto- surfaces) of these two elements join to create an edge- to-edge contact so that the resulting bulla surface is a tympanic and makes contact with the petrosal unilaminar subhemisphere. In aeluroids, the inturned promontorium. The fit between flange and posteromedial face ofectotympanic creates a surface that petrosal is quite close, as demonstrated by meets a second surface produced by the inflected anterior the distinctive promontorial facet (F in figs. margin ofcaudal entotympanic; no edge-to-edge contact 8, lOB, 1 1, 12). In living felids such as the ever persists, but rather two surfaces are brought into domestic cat, Panthera tigris, P. pardus, Felis contact to create a bilaminar septum of varying size. 1 989 HUNT: AELUROID CARNIVORA 23

Did proailurines and stenoplesictines pos- only a modestly inflected margin of caudal sess the septum bullae? Piveteau (1943) and entotympanic, suggesting that the edges of Beaumont (1964) believed that Stenoplesictis caudal entotympanic have not begun to as- did not, interpreting the internal partition as sume the strongly inflected configuration of unilaminar, formed by the ectotympanic only. the living aeluroid families. At best, in these In fact, we are ignorant of not only the type two stenoplesictine genera, the caudal ento- of connective tissue forming the caudal en- tympanic contact with ectotympanic would totympanic of Stenoplesictis, but also do not range from the type found in Nandinia (ru- know ifit was applied as a lamina against the dimentary) to the incipient septum bullae seen ectotympanic. Such application ofcaudal en- in neonatal domestic cats (slightly inflected, totympanic might be very difficult to detect as in Hunt, 1987: fig. 11). in the fossil, especially if cartilaginous. Here The similarity of the proailurine basicra- Nandinia can serve as a guide. In the African nium, especially the petrosal, to that ofsteno- palm civet, the caudal entotympanic does not plesictines indicates that primitive proai- form a bilaminar septum with ectotympanic; lurines such as Stenogale julieni very likely rather its contact with the posteromedial face possessed a bulla in which ectotympanic-cau- of ectotympanic is a linear feature, a line of dal entotympanic contact was rudimentary, attachment (X in figs. 9A, 1OA) that poten- so that a well-developed septum bullae prob- tially could evolve into a septum bullae upon ably did not evolve in felids until at least the the inflation of ectotympanic and inflection stage represented by Aquitanian P. lemanen- ofcaudal entotympanic. Nandinia is remark- sis. A virtually modern felid auditory bulla able for its retention of this plesiomorphic including a true septum bullae is present in pattern, which can be identified as a logical North American mid-Miocene Pseudaelurus. structural predecessor for all other aeluroids Consequently, we can conclude that prim- in terms of the ectotympanic-caudal ento- itive proailurines (S. julieni) and stenople- tympanic contact. Stenoplesictis, on the other sictines possess certain plesiomorphic attri- hand, possesses a more derived (inflated) ec- butes ofthe auditory region that do not occur totympanic; yet the caudal entotympanic in any living aeluroid (except Nandinia), such chamber of the bulla is about the same size as the small size of the caudal entotympanic as in Nandinia. There is no evidence of cau- chamber and the probable lack of, or only dal entotympanic inflation in either animal; incipient development of, a septum bullae. therefore the contact ofcaudal entotympanic However, in their development of a moder- and ectotympanic may not have been con- ately inflated ectotympanic, deeply registered verted into a bilaminar septum bullae in on alisphenoid and squamosal bones, that Stenoplesictis. firmly contacts the petrosal by means of a I am in agreement with Piveteau and Beau- prominent facet, stenoplesictines and prim- mont that the weight of evidence argues itive proailurines are advanced beyond the against the presence in stenoplesictines of a condition seen in Nandinia, having created a typical septum bullae as seen in living felids, somewhat specialized enclosure for the an- herpestids, or viverrids. This is supported by terior chamber of the bulla. the illustrations ofPalaeoprionodon supplied Basicranial evidence derived from these by Teilhard (1915: pl. 9, fig. 10) in which early aeluroids limits the range of possible only a linear attachment of caudal entotym- hypotheses on the origin ofthe aeluroid fam- panic on ectotympanic is suggested, and in ilies which we will next consider. Stenoplesictis illustrated by Piveteau (1943: pl. 1, fig. 3A) where there is very little room adjacent to the flange for application of cau- ORIGIN OF THE LIVING dal entotympanic. This conclusion is also in- AELUROID FAMILIES dicated by the distinctive trough on the pos- The modern felid radiation seems directly terior margin of the petrosal for registration descended from a proailurine ancestry. There of the caudal entotympanic in stenoplesic- is an identifiable continuum of fossils begin- tines (fig. 8, stipple pattern). This trough is ning with Proailurus, leading to Pseudaelu- developed as in Nandinia, where it receives rus, and eventually to the later Cenozoic and 24 AMERICAN MUSEUM NOVITATES NO. 2930 living felids (Ginsburg, 1983). Most impor- genus (Galerella) ofHerpestes, suggesting that tantly, the modification of the auditory re- the basicranium is essentially ofmodem type. gion and bulla can be directly traced from the Recently, two nearly complete herpestid skulls primitive proailurine state to living felids with referable to Herpestes and Helogale have been their enormously inflated bullae. reported in the Pliocene fauna from the upper Because ofthe scarcity offossils, a similarly unit of the Laetolil Beds (Tanzania), dated well-documented history of viverrids and from 3.5 to 3.8 Ma (Petter, 1987). These two herpestids is yet to be worked out. The later skulls are significant in possessing basicrania Cenozoic evolution ofhyaenids is somewhat and auditory bullae of modem herpestid better known, whereas the early history of grade. Thus, in Africa, the evidence suggests that group is only recently emerging. that a herpestid fauna of modem aspect was True herpestids probably first appear as a in existence by the early Pliocene, and that discrete lineage in the earlier Miocene of Eu- the basicranial structure and auditory bulla rope in the form of small mongooses (basilar pattem of the living Herpestidae probably skull lengths of about 6 cm) referred to Lep- evolved in the Miocene. toplesictis. These fossils are similar in size to Herpestids are not recorded in Asia prior the living mongoose Herpestes auropuncta- to the late Miocene. Recent work in the Si- tus. The earliest record of Leptoplesictis in waliks ofnorthem Pakistan by Barry and co- Europe is mammal zone MN4 (Schmidt-Kit- workers (1980) produced herpestids at four tler, 1987) but to my knowledge no well-pre- localities in the Khaur region. These sites fall served basicrania are known. Because the between about 7.1 and 9.5 Ma, hence late basicranium ofLeptoplesictis cannot be com- Miocene (MNl 1-12, Turolian). The fossils pared with stenoplesictine skulls, the rela- have not yet been described. tionship of these earliest herpestids to the Pre-Pliocene herpestid basicrania with pre- stenoplesictines is not apparent. served auditory bullae are unknown to me. Recent discoveries in east Africa (Schmidt- We are left primarily with a Miocene record Kittler, 1987) show that the oldest known of dentitions, referred either to Herpestes or African herpestids found in early Miocene to Leptoplesictis. These teeth are sufficiently rocks of Rusinga Island are nearly the same similar to those ofliving herpestids to suggest age as the oldest European herpestids. The that when good basicrania are found, they Rusinga faunas are radiometrically dated at could have auditory regions similar to the about 17.8 Ma. The African herpestid fossils modem species. This is significant because are dental remains that also can be referred the auditory region ofliving herpestids man- to Leptoplesictis and, as in Europe, no basi- ifests a clearly derivative morphologic pat- crania are known. tem (Hunt, 1987); when this pattem first de- Pleistocene herpestid basicrania from Af- veloped is important in judging the timing of rica are no different from those ofliving her- the herpestid radiation. The herpestid basi- pestids, and, based on the faunas from 01- cranial pattem is distinguished by: anterior- duvai, Omo, and from south Africa, as many posterior alignment of the two bulla cham- as seven of the living herpestid genera (Her- bers, resulting in transverse placement ofthe pestes, Mungos, Ichneumia, Atilax, Helogale, septum bullae; lack of pronounced over- Cynictis, Crossarchus) were in existence by growth of either bulla chamber by the other; the early Pleistocene. considerable distance between the anterior The modern herpestid radiation then must end of the bulla and the postglenoid process begin in the Pliocene or earlier, and may be of the squamosal (except in genera with in- ofconsiderable antiquity, since the E Quarry flated ectotympanic); enclosure of the inter- fauna from Langebaanweg in south Africa nal carotid artery in a bony tube between dates from the early Pliocene (4-5 Ma) and rostral entotympanic and ectotympanic (the preserves a diverse herpestid assemblage that artery never enters the middle ear cavity); may comprise up to five species (Hendey, and the anterior placement and complete en- 1976). A single Pliocene herpestid skull from closure within the basisphenoid ofthe middle Garussi in Tanzania preserves part ofthe au- lacerate foramen (including arterial anasto- ditory region (Dietrich, 1942); Petter (1963) mosis Y of Bugge, 1978). was able to assign the skull to an extant sub- The basicranial pattem of herpestids is so 1989 HUNT: AELUROID CARNIVORA 25 similar among the extant genera that a late chamber by the other, the auditory region Cenozoic divergence would not be surprising: does not possess the derived carotid path the herpestid radiation may be primarily a within the bulla wall typical of living her- Plio-Pleistocene event (Hendey, 1974: 89, pestids. In addition, the isolation ofthe mid- 337f.), geographically largely confined to Af- dle lacerate foramen within the basisphenoid rica. Because of the uniformity of the basi- that characterizes living herpestids does not crania ofliving herpestids, the radiation must seem to be developed in Herpestides. Her- stem from an ancestral herpestid already in pestids also tend to fill in the ectotympanic possession ofthe modern basicranial pattern. crescent with bone, and may laterally extend Hyaenids currently are thought to be de- the ectotympanic to form a short meatal tube, rived from the European early Miocene ae- but this type of closure has not occurred in luroid Herpestides (Beaumont and Mein, Herpestides. 1972). Herpestides already coexists as a sep- Herpestides appears to be an early aeluroid arate lineage in the Aquitanian with primi- that has not yet developed a basicranial pat- tive Proailurus, and must have diverged prior tern typical of one ofthe living families. The to this time in the Oligocene. It appears for strong anteroventral overgrowth of ectotym- the first time in European Neogene mammal panic by caudal entotympanic seen in vi- zone MN2 (Schmidt-Kittler, 1987). The verrids and most felids is not present. Beau- modern hyaenid bulla pattern had evolved mont and Mein's (1972) suggestion that at least by MNl 1-12, about 7 to 9 Ma (Tu- Herpestides is an early hyaenid is not contra- rolian, late Miocene; see Hunt 1 974a: fig. 42). dicted by its auditory region. Its bulla struc- Unpublished observations indicate that the ture could represent an early stage preceding hyaenid bulla type is recognizable somewhat the modern highly derived hyaenid condition earlier in east Asia in deposits of mid-Mio- (fig. 6), particularly since the carotid path in cene age: (a) my examination of the auditory hyaenids, although vestigial, is apprently region of the holotype skull of Tungurictis transpromontorial, just as in Herpestides. (AMNH 26600) from Mongolia (Tung Gur Posterior growth and enlargement of the ec- Formation, Wolf Camp Quarry, zone MN8) totympanic in Herpestides could result in the convinces me that the bulla is of the hyaenid hyaenid bulla configuration. Indeed, the type; (b) hyaenids referable to Percrocuta from comparison ofHerpestides with herpestids is the Tongxin basin of north China, at least as not unreasonable when one realizes that the old as zone MN8, are reported to have rec- bulla elements of a neonatal herpestid (fig. 5) ognizable hyaenid auditory regions (Qiu are structured and positioned to evolve by Zhanxiang, personal commun., 1988). The allometric growth into the hyaenid state (fig. existence in the later mid-Miocene in zone 6). Accordingly, it is possible that Herpestides MN8 of two kinds of hyaenids as morpho- is close to the ancestry of both hyaenids and logically diverse as Tungurictis and Percro- herpestids. cuta that display hyaenid auditory regions is The family Viverridae in the past has been evidence for an even earlier evolution ofthis indiscriminately employed as a taxon encom- bulla type, hence development during or be- passing a variety of primitive and advanced fore the earlier mid-Miocene. aeluroids. With an awareness that true vi- The auditory region of early Miocene verrids can be identified by a derived and Aquitanian Herpestides is preserved in sev- distinctive basicranial anatomy, in particu- eral skulls, and has been described by Beau- lar, the pattern of ontogenetic growth of the mont (1967) and Petter (1974). Beaumont bulla elements (Hunt, 1987), it is possible to initially thought that this carnivoran was separate groups such as herpestids and Nan- closely related to living herpestids. However, dinia from Viverridae. Despite the present the auditory region differs in several respects diversity of true viverrids in the Old World, from those of known herpestids. Both au- there are no described pre-Pliocene viverrid thors restore the internal carotid artery in the skulls with intact basicrania (excluding primitive transpromontorial position. Hence, stenoplesictines) that can be certainly attrib- despite the herpestid-like fore-aft alignment uted to the family on the basis of auditory ofthe anterior and posterior bulla chambers, structure. The viverrid bulla pattern proba- and the apparent lack ofovergrowth ofeither bly first appears in the Miocene, but reliable 26 AMERICAN MUSEUM NOVITATES NO. 2930 paleontological evidence bearing on this issue lassictis. Therefore, both auditory bulla struc- has been difficult to identify, and is only now ture and the dentition of Tungurictis better coming into focus. agree with placement in Hyaenidae, an alli- The principal Old World Miocene viverrid ance initially suggested by Beaumont (1967) genera from Europe (Semigenetta, Jourda- and supported by Qiu et al. (1988). nictis, Viverrictis, Lophocyon) are not repre- By the beginning of the Pliocene, true vi- sented by any well-preserved basicrania verrids with typical basicranial and bullar showing bulla structure. Fossils of these car- structure are known in Africa, and probably nivorans are primarily teeth injaw fragments also were present in south Asia. The three and rare postcranial remains. Dentitions known Pliocene basicrania with preserved strongly indicate these are aeluroid carnivo- auditory bullae belong to two very large true rans, and some may eventually prove to be viverrids, the size of large coyotes or small viverrids on basicranial structure. At least wolves: Vishnuictis durandi (BMNH M1 338, two badly damaged basicrania ofthe dentally M37131) from the Pinjor Siwaliks (Pilgrim, specialized Lophocyon are known (Fejfar and 1932), and "Viverra" leakeyi (SAM L13097) Schmidt-Kittler, 1987); sketches of the au- from Langebaanweg in South Africa (Hen- ditory region suggest it is an aeluroid, but the dey, 1974). The intact well-preserved audi- quality ofpreservation ofthe bulla is not suf- tory bulla of Vishnuictis durandi is essentially ficient to classify it as a true viverrid. that of a modern viverrid, very much like In Asia, the only described Miocene ae- Civettictis, whereas the auditory region ofthe luroid skulls with basicrania that have been Langebaanweg viverrid has yet to be de- attributed to the Viverridae are the holotype scribed in detail. Howell and Petter (1976) of Vishnuictis salmontanus from the Dhok suggested that V. durandi and leakeyi are in Pathan Siwaliks (Pilgrim, 1932), and the ho- fact the same species. lotype of Tungurictis spocki from the mid- Recent discovery of a group of 8.2 Ma late Miocene of Mongolia (Colbert, 1939). These Miocene Siwalik carnivoran skulls by Barry are aeluroids with skull lengths of about 11 and others (personal commun., 1987) may cm, the size of the living Viverra. The basi- include the first definite viverrid skulls of cranium of Vishnuictis is too badly damaged Miocene age with intact auditory regions. The to determine (Pilgrim, 1932), but that ofTun- structure of the auditory bullae of these Si- gurictis, including the auditory bulla, is much walik fossils will provide important con- better preserved. straints on the pattern of viverrid evolution Unfortunately, photographs of the only when described, and may clarify the affinities known skull and basicranium of Tungurictis ofthe Dhok Pathan Vishnuictis salmontanus. have not been published, but I have been able Recently I was able to examine, through to study the skull in the American Museum. the courtesy of Dr. Alan Gentry, a complete The auditory region was originally described fully articulated viverrid skeleton from the and figured by Colbert (1939) who reported freshwater limestone ofOeningen (zone MN8) that a large inflated entotympanic extended preserved in the British Museum, originally forward to cover the ectotympanic; this on- described as a canid. There were no canids togenetic growth pattern ofthe bulla elements in Eurasia at this time, and the teeth which is typical of living members of the Viverri- can be seen in the lower jaw show viverrid dae, and suggested that Colbert was correct dental traits: p4 with anterior accessory cusp; in classifying Tungurictis as a viverrid. How- closed trigonid ofm 1; no m3. This is the most ever, upon firsthand examination ofthe skull, complete European fossil viverrid yet dis- the anterior chamber ofthe bulla is not over- covered, and includes skull and jaws in ar- grown by caudal entotympanic; in fact, the ticulation. The basicranium and bulla are un- anterior chamber is formed entirely by an doubtedly present, but the skull is crushed inflated ectotympanic, as in hyaenids. In ad- flat so that nothing of the basicranial struc- dition, the posterior chamber ofthe bulla pre- ture can be seen without additional prepa- serves an internal floor that represents the ration. This is a large animal, with head-body nearly horizontal hyaenid septum. The form length of about 65 cm (similar in body size and proportions of the teeth are very much to the living Viverra). like, but smaller than, the early hyaenid Tha- As a result of this situation, we cannot be 1 989 HUNT: AELUROID CARNIVORA 27 certain when in the mid-Cenozoic the mod- stenoplesictine presence in the Old World, em viverrid bulla configuration materialized. there must have been somewhere in Eurasia If Palaeoprionodon from Quercy is in fact a or Africa a more basicranially primitive ae- true viverrid, then we can state that in the luroid stock that represents the ancestry of Oligocene its auditory structure was not yet Nandinia. This ancient aeluroid lineage has of modem grade. However, it would require persisted unchanged in basicranial structure, only slight structural changes, primarily a and probably in skull form and postcranial greater degree of development of the caudal anatomy, for at least 35 to 40 million years. entotympanic chamber, to evolve the bulla The only probable modification of this an- ofliving viverrids like Prionodon and Poiana cestral aeluroid necessary to arrive at modem from Palaeoprionodon. The living linsangs Nandinia is the reduction of the dentition (Prionodon, Poiana), the former distributed with development ofa hypocamivorous cusp in Asia from Nepal southward to Java, the pattem. It is remarkable that in the east Af- latter in west Africa, are much like Palaeo- rican Songhor level, dated at about 19.6 Ma, prionodon in bulla structure, dentition, and there has recently been reported a rich di- general skull form (basilar skull lengths in the versity of small aeluroids including Mio- two genera are about 6 cm), and are plausibly prionodon and Stenoplesictis, both described directly descended from the Quercy genus as stenoplesictines (Schmidt-Kittler, 1987). (Gregory and Hellman, 1939). Dissection of Mioprionodon is known only from jaws and a skull of Poiana richardsoni (AMNH-M teeth but its dentition is rather similar to that 236487) from Cameroon shows that the pe- of Nandinia. I think it probable that Nan- trosal possesses a VPP that is more vertical dinia has existed in Africa since at least the and robust than in other viverrids, hence time of Mioprionodon in the early Miocene. closer to the plesiomorphic state found in What then are stenoplesictines? Most cur- Palaeoprionodon. rent workers limit the group to Stenoplesictis Conceivably, the ancestry of many of the and Palaeoprionodon, and ally them with the living viverrid genera may be found in small proailurine felids. Indeed, basicranial struc- aeluroids closely related to Palaeoprionodon ture supports the inference that proailurines that differentiated as distinct phyletic lin- (Proailurus, Stenogale) and stenoplesictines eages during the Oligocene. For the moment, (Stenoplesictis, Palaeoprionodon) are sister- Palaeoprionodon seems to be the most reli- groups, and that they are early derivative lin- able evidence of the first true viverrids doc- eages of a more primitive aeluroid stock- umented by basicranial remains. The best es- whose existence we infer from basicranial timate at present as to the time of origin of structure-that survives in the living Nan- the modem viverrid auditory region is that, dinia (fig. 13). A phyletic connection ofsteno- based on the skull of Palaeoprionodon, the plesictines and proailurines with living ae- modem grade of bulla structure was ap- luroids is documented by the identification proached but not yet achieved in the mid- to of Palaeoprionodon as the oldest probable late Oligocene and early Miocene, so that by viverrid lineage, and Proailurus as an early mid-Miocene we might expect to encounter felid. The anatomical similarity ofbasicrania fossil viverrids with the modem bulla pat- of the stenoplesictine Palaeoprionodon and tem. The Oeningen skeleton and the denti- the proailurine Proailurus indicates the close tions of Viverrictis and Semigenetta may well relationship between these lineages in the represent such animals. Future discoveries of Oligocene, and this relationship is further re- cranial remains will be necessary to confirm flected in the similar ontogenetic pattem of or modify this prediction. bulla growth in living viverrids and felids Nandinia remains as an isolated taxon (Hunt, 1987). Thus a close tie between vi- among living Aeluroidea. It is the only living verrids and felids is supported not only by aeluroid that not only approximates the the similar ontogeny of the auditory bulla of stenoplesictine grade ofbasicranial structure, living forms but also by the paleontological but documents a stage of basicranial orga- evidence allying ancestral species from Quer- nization more plesiomorphic than the sten- cy. Consequently, there is good evidence that oplesictines. The existence of Nandinia im- at least some felids and some viverrids are plies that prior to and during the Oligocene products ofan early aeluroid dichotomy, and 28 AMERICAN MUSEUM NOVITATES NO. 2930

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proailurine~~. stenoplesictine.aeluroids. cer.in. im.fth 1 989 HUNT: AELUROID CARNIVORA 29 this conclusion can be reasonably extended Aquitanian time, probably during the Oli- to unite the two families as sister taxa. gocene, with the origin ofthe aeluroids within In both Europe and Africa, other aeluroids the Carnivora plausibly dating to the early developed into what can tentatively be de- Cenozoic. scribed as protohyaenids (Herpestides) and protoherpestids (Leptoplesictis) by the early Miocene. However, neither ofthese two gen- CONCLUSIONS era can be clearly confirmed as ancestors of A ventral promontorial process of the pe- their respective families based on basicranial trosal characterizes the aeluroid Carnivora, evidence. Nor are they demonstrably derived living and fossil. This process is applied from a common hyaenid-herpestid stem against the lateral edge of the basioccipital, group, although no evidence prevents such and often protrudes below (ventral to) the an interpretation. We must await new fossil basicranial axis. Living aeluroids have a for- discoveries to better determine hyaenid and wardly inclined petrosal process situated near herpestid origins. In the interim, we may con- the basioccipital-basisphenoid suture, but the sider these two genera as hypothetical ances- oldest known generally acknowledged aelu- tral stocks for hyaenids and herpestids. How- roids, the stenoplesictines and proailurines, ever, when additional fossil skulls of early have a more robust and vertically aligned aeluroids are eventually discovered, an im- petrosal process which is applied to basioc- portant criterion needs to be satisfied before cipital about midway along its lateral margin. they can be reliably attributed to the proto- The living African aeluroid Nandinia still re- hyaenid or protoherpestid groups: we should tains a petrosal process configured as in these be able to show that the bulla and basicra- ancient Oligocene aeluroids. In fact, its au- nium ofthe fossil taxon have developed along ditory region in its entirety is more plesio- an ontogenetic trajectory in evidence in the morphic than the auditory regions of these living members of these two families. oldest fossil aeluroids. Nandinia is a "living The coexistence of protofelids, -viverrids, fossil," representative of the earliest stage of -hyaenids, and somewhat later, protoherpes- aeluroid evolution in terms of its basicranial tids in Europe in the late Oligocene and ear- and auditory anatomy. lier Miocene indicates that separation ofthese The time of origin of the living aeluroid lineages had already taken place by Aquita- families, based on auditory anatomy, is dif- nian time in the Old World (fig. 13). Never- ficult to determine because well-preserved theless, the modem grades ofbasicranial and basicrania are scarce in the fossil record, but bulla structure that identify members of the the known paleontological evidence suggests living families were only incipiently devel- that the modem basicranial and bullar pat- oped in these four ancestral lineages, and this terns of most lineages developed during the nascent condition of the auditory region has Miocene. In the Oligocene and early Mio- contributed to the problem of their recogni- cene, ancestral stocks ofthe modem families tion in the fossil record. More importantly, can be cautiously recognized on the basis of the more plesiomorphic structure of the au- basicranial structure likely to precede the dis- ditory region of Nandinia relative to these tinctive basicranial patterns identifying the Aquitanian species implies that it also must living families. In the mid- to late Miocene, have been in existence at that time as an even basicranial patterns typical ofthe living felids more primitively configured lineage, at least and hyaenids were fully developed. Because in terms ofits basicranium. Thus the aeluroid of the paucity of fossil basicrania, modern differentiation must have occurred in pre- herpestid and viverrid basicranial patterns

herpestids (Leptoplesictis) prevents determination of their relationship to proposed oldest hyaenids (Herpestides). The living palm civet Nandinia maintains a basicranial pattern plesiomorphic for all these fossil and living aeluroid groups. A, Stenogalejulieni; B, Stenoplesictis cayluxi; C, Herpestides antiquus; D, Panthera onca; E, Civettictis civetta; F, Galidia elegans; G, Crocuta crocuta; H, Nandinia binotata. 30 AMERICAN MUSEUM NOVITATES NO. 2930 are not certainly recognized until the Plio- Colbert, E. cene, but almost surely were in existence by 1939. Carnivora of the Tung Gur Formation the late Miocene. ofMongolia. Bull. Am. Mus. Nat. Hist. 76: 47-81. Crochet, J.-Y., J.-L. Hartenberger, J.-C. Rage, J.-A. Remy, B. Sige, J. Sudre, and M. Vianey- REFERENCES Liaud Barry, J. C., A. K. Behrensmeyer, and M. Mon- 1981. Les nouvelles faunes de vertebres an- aghan terieures a la "Grande Coupure" de- 1980. A geologic and biostratigraphic frame- couvertes dan les phosphorites du Quer- work for Miocene sediments near Khaur cy. Bull. Mus. Natl. Hist. Nat., Paris, village, northern Pakistan. Peabody ser. 4, 3(3): 245-266. Mus. Postilla 183(1-2): 1-19. Crusafont-Pairo, M., and J. Truyols-Santonja Beaumont, G. de 1956. A biometrical study of the evolution of 1961. Recherches sur Felis attica Wagner du fissiped carnivores. Evolution 10: 314- Pontien eurasiatique avec quelques ob- 332. servations sur les genres Pseudaelurus Dehm, R. Gervais et Proailurus Filhol. Nouv. 1950. Die Raubtiere aus dem Mittel-Miociin Arch. Mus. Hist. Nat. Lyon 6: 17-45. (Burdigalium) von Wintershof-west bei 1964. Remarques sur la classification des Fe- Eichstiitt in Bayern. Abhandl. Baye- lidae. Eclogae. Geol. Helv. 57(2): 837- risch. Akad. d. Wissensch. (n.f.) 58: 1- 845. 141. 1967. Observations sur les Herpestinae (Vi- Dietrich, W. verridae, Carnivora) de l'Oligocene su- 1942. Altestquartire Siiugetiere aus der sild- perieur avec quelques remarques sur des lichen Serengeti, Deutsch-Ostafrica. Pa- Hyaenidae du Neogene. Arch. Sci. Ge- laeontographica 94: 43-133. neve 20(1): 79-107. Fejfar, O., and N. Schmidt-Kittler Beaumont, G. de, and P. Mein 1987. Lophocyon carpathicus n. gen. n. sp. aus 1972. Recherches sur le genre Plioviverrops dem Jungtertiiir der Ostslowakei und Kretzoi (Carnivora, ?Hyaenidae). C. R. eine neue unterfamilie der Schleich- des Seances, SPHN Geneve 25(3): 383- katzen(Viverridae). Palaeontographica 394. 199(1): 1-22. Bonis, L. de Filhol, H. 1973. Contribution a l'etude des mammiferes 1879. Etude des mammiferes fossiles de St.- de 1'Aquitanien de l'Agenais. Mem. Mus. Gerand le Puy (Allier). Bibliotheque de Natl. Hist. Nat., Paris (n.s.), ser. C 28: l'Ecole des Hautes Etudes (Sect. Sci. 1-192. Nat.) 19(1): 1-252. 1974. Premieres donnees sur les carnivores 1882. Memoire sur quelques mammiferes fos- fissipedes provenant des fouilles re- siles des Phosphorites du Quercy. Ann. centes dan le Quercy. Palaeovertebrata Soc. Sci. Phys. Nat., Toulouse: 66-67. 6: 27-32. Fischer, K. Bonis, L. de, J.-Y. Crochet, J.-C. Rage, B. Sige, J. 1983. Stenoplesictis (Viverridae, Carnivora, Sudre, and M. Vianey-Liaud Mammalia) aus dem marinen Mittelo- 1973. Nouvelles faunes de Vertebres oligo- ligoziin der Braunkohlentagebaue des cenes des phosphorites du Quercy. Bull. Weisselsterbeckens (Bezirk Leipzig Mus. Natl. Hist. Nat., Paris, ser. 3, Sci. DDR). Schriftenr. geol. Wiss., Berlin, Terre 28(174): 105-113. 19/20: 209-215. Bugge, J. Flower, W. H. 1978. The cephalic arterial system in carni- 1869. On the value of the characters of the vores, with special reference to the sys- base ofthe cranium in the classification tematic classification. Acta Anat. 101: ofthe Order Carnivora. Proc. Zool. Soc. 45-61. London: 4-37. Chapuis, G. Flynn, J., and H. Galiano 1966. Contribution a l'etude de l'artere caro- 1982. Phylogeny of early Tertiary Carnivora, tide interne des carnivores. Mammalia with a description of a new species of 30: 82-96. Protictis from the middle Eocene of 1989 HUNT: AELUROID CARNIVORA 31

northwestern Wyoming. Am. Mus. nimravid cat Dinictis. Am. Mus. Novi- Novitates, 2725: 64 pp. tates 2886: 74 pp. Ginsburg, L. Lavocat, R. 1979. Revision taxonomique des Nimravini 1952. Sur les affinites de quelques carnassiers (Carnivora, Felidae) de l'Oligocene des de l'Oligocene d'Europe. Mammalia Phosphorites du Quercy. Bull. Mus. 16(2): 62-72. Natl. Hist. Nat., Paris, ser. 4, 1(1): 35- Lydekker, R. 49. 1885. Catalogue of Fossil Mammalia in the 1983. Sur les modalites d'evolution du genre British Museum (Natural History), Neogene Pseudaelurus Gervais (Feli- London. 268 pp. dae, Carnivora, Mammalia). In J. Matthew, W. D. Chaline, ed., Modalites, rythmes, me- 1909. The Carnivora and Insectivora of the canismes de l'evolution biologique. Bridger Basin, middle Eocene. Mem. Gradualisme phyletique ou equilibres Am. Mus. Nat. Hist. 9: 289-567. ponctues? CNRS Colloq. 330: 131-136. Mellett, J. Gregory, W. K., and M. Hellman 1968. The Oligocene Hsanda Gol Formation, 1939. On the evolution and major classifica- Mongolia: a revised faunal list. Am. tion ofthe civets (Viverridae) and allied Mus. Novitates 2318: 16 pp. fossil and recent Carnivora: a phyloge- 1977. Paleobiology of North American netic study of the skull and dentition. Hyaenodon (Mammalia, Creodonta). Proc. Am. Phil. Soc. 81(3): 309-392. Contrib. Vert. Evol. 1: 1-134. Helbing, H. Mivart, St.-G. 1928. Carnivoren des oberen Stampien. Ab- 1882a. On the classification and distribution of handl. schweiz. palaeontol. Ges. 47: 1- the Aeluroidea. Proc. Zool. Soc. Lon- 83. don: 135-208. 1882b. Notes on some points in the anatomy Hendey, Q. B. ofthe Aeluroidea. Proc. Zool. Soc. Lon- 1974. The Late Cenozoic Carnivora of the don: 459-520. Southwestern Cape Province. Ann. South African Mus. 63: 1-369. Petter, G. 1976. The Pliocene fossil occurrences in E 1963. Etude de quelques Viverrides (Mam- Quarry, Langebaanweg, South Africa. miferes, Carnivores) du Pleistocene in- Ann. South African Mus. 69: 215-247. ferieur du Tanganyika (Afrique orien- tale). Bull. Soc. Geol., France, ser. 7, Hough, J. 5(3): 265-274. 1948. The auditory region in some members 1974. Rapports phyletiques des viverrides of the Procyonidae, Canidae, and Ur- (carnivores fissipedes). Les formes de sidae: its significance in the phylogeny Madagascar. Mammalia 38(4): 605-636. of the Carnivora. Bull. Am. Mus. Nat. 1987. Small carnivores from Laetoli. In M. Hist. 92: 67-118. Leakey and J. Harris (eds.), Laetoli: a Howell, F. C., and G. Petter Pliocene site in northern Tanzania, pp. 1976. Carnivora from Omo Group forma- 194-234. Oxford: Oxford Univ. Press. tions, southern Ethiopia. In Y. Cop- Pilgrim, G. E. pens, F. Howell, G. Isaac, and R. 1931. Catalogue of the Pontian Carnivora of Leakey (eds.), Earliest man and envi- Europe in the Department of Geology, ronments in the Lake RudolfBasin, pp. British Museum (Natural History), 314-331. Chicago: Univ. Chicago Press. London. 174 pp. Hunt, R. M., Jr. 1932. The Fossil Carnivora of India. Mem. 1974a. The auditory bulla in Carnivora: an an- Geol. Surv. India (Palaeontol. India) (n. atomical basis for reappraisal of carni- ser.) 18: 1-232. vore evolution. J. Morphol. 143: 21-76. Piveteau, J. 1974b. Daphoenictis, a cat-like carnivore 1943. Etudes sur quelques mammiferes des (Mammalia, Amphicyonidae) from the Phosphorites du Quercy. Ann. Paleon- Oligocene ofNorth America. J. Paleon- tol. 30: 63-72. tol. 48(5): 1030-1047. 1961. Carnivora. In Mammiferes, Origine 1987. Evolution of the aeluroid Carnivora: Reptilienne, Evolution, Traite de Pa- significance of auditory structure in the l6ontol. 6(1): 641-820. 32 AMERICAN MUSEUM NOVITATES NO. 2930

Qiu, Zhanxiang, Yan D., Chen G., and Qiu Zhu- tel-Eoziin von Messel. Palaeontol. Zeit. ding 54: 171-198. 1988. Preliminary report on the field work in 1982. Neue raubtiere (Mammalia: Creodonta 1986 at Tung-Gur, Nei-Mongol. Kexue et Carnivora) aus dem Lutetium der Tongbao 33(5): 399-404. Grube Messel (Deutschland). Palaeon- Schlosser, M. tographica 179: 105-141. 1889. Die Affen, Lemuren, Chiropteren, In- 1985. Zwei neue Skelette von Miacis ?kessleri sectivoren, Marsupialier, Creodonten (Mammalia, Carnivora) aus den lute- und Carnivoren des europiiischen Ter- tischen Olschiefern der "Grube Mes- tiairs, part II, pp. 1-162. Vienna: Alfred sel." Senckenberg. Leth. 66: 121-141. Holder. Tedford, R. H. 1923. Grundzuge der Paliiontologie (Paliio- 1976. Relationships ofpinnipeds to other car- zoologie) von K. A. von Zittel, part II nivores (Mammalia). Syst. Zool. 25(4): (Vertebrata). Neuarbeitet von F. Broili 363-374. und M. Schlosser. Munich and Berlin: Teilhard de Chardin, P. R. Oldenbourg, 706 pp. 1915. Les carnassiers des Phosphorites du Quercy. Ann. Paleontologie 9(3,4): 103- Schmidt-Kittler, N. 192. 1987. The Carnivora (Fissipedia) from the Turner, H. N. Lower Miocene of East Africa. Palae- 1848. Observations relating to some ofthe fo- ontographica 197(3): 85-126. ramina at the base ofthe skull in Mam- Sige, B., J.-Y. Crochet, J.-L. Hartenberger, J.-A. malia, and on the classification of the Remy, J. Sudre, and M. Vianey-Liaud Order Carnivora. Proc. Zool. Soc. Lon- 1979. Mammiferes du Quercy. Fossilium Ca- don: 63-88. talogus, I. Animalia (F. Westphal, ed.) Viret, J. 126: 1-99. 1929. Les faunes des mammiferes de l'Oli- Springhorn, R. gocene superieur de la Limagne Bour- 1980. Paroodectes feisti, der erste Miacide bonnaise. Ann. l'Univ. Lyon (n. ser.) 47: (Carnivora, Mammalia) aus dem Mit- 1-328.

Note Added in Proof: A primitive mongoose with a head-body length of 50 cm has recently been reported from complete skeletons found at Fort Ternan, Kenya, by R. J. G. Savage (R. J. G. Savage and M. R. Long, 1986, Mammal Evolution, British Museum (Nat. Hist.), London, p. 80). This sample apparently includes at least one skull of a juvenile with a complete or nearly complete auditory bulla (J. Barry, written commun., 1988). The mid-Miocene age ofFort Ternan (about 14 Ma) falls in a critical interval in the course of herpestid basicranial evolution, thus when these specimens are described, they will shed considerable light on the pattern of aeluroid basicranial change.

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