Podocnemis congolensis Dollo, 1912

GAFFNEY, EUGENE S, TONG, HAIYAN & MEYLAN, PETER A, 2006, EVOLUTION OF THE SIDE-NECKED TURTLES: THE FAMILIES BOTHREMYDIDAE, EURAXEMYDIDAE, AND ARARIPEMYDIDAE, Bulletin of the American Museum of Natural History 300 (300), pp. 1-698 : 99-106

Podocnemis congolensis Dollo, 1912 . Podocnemis congolensis Dollo, 1913 . Bantuchelys congolensis Dollo, 1924 .

TYPE SPECIMEN: Uncataloged specimen in the MRAC, consisting of the first right costal, peripherals 2–4, and fragments of peripheral 1 and the nuchal, figured in Wood (1975: fig. 1) and Dollo (1913: pl. 7, figs. 1, 2). As discussed by Wood (1975), Dollo (1924) designated a new type for Bantuchelys congolensis , but this is preceded by the Dollo (1913) specimen.

TYPE LOCALITY: Presumably ( Dollo, 1913; Wood, 1975) cliff exposures near Landana, Cabinda, a formerly Portuguese colony on the north side of the Congo River (fig. 14).

HORIZON: Presumed to be the sequence consisting of mostly Paleocene sediments ( Dollo, 1913; Wood, 1975). Darteville and Casier (1943, 1959) described 32 beds at Landana, the lower 29 as Paleocene with turtles commonly found in most of them. Most of the turtles come from beds identified as Montian in age (Darteville and Casier, 1959; Wood, 1973, 1975).

DEPOSITIONAL ENVIRONMENT: The Taphrosphys congolensis material is thought to have come mostly from near-shore marine beds (Cahen, 1954) and is associated with rarer cheloniid specimens ( Wood, 1973) and crocodiles ( Dollo, 1914). The fish from these units have been described by Darteville and Casier (1943, 1949, 1959).

DIAGNOSIS: A bothremydid pleurodire of the genus Taphrosphys with these unique characters: premaxilla labial ridge deep not shallow; prefrontal thin and short; distinguished from the other two species as follows: from T. sulcatus : rostrum basisphenoidale a long and prominent, laterally compressed rod; sella turcica deep and narrow; dorsum sellae high; processus clinoideus high and large; processus inferior parietalis closer to midline than in T. sulcatus ; from T. ippolitoi : snout not expanded; quadrate shelf not expanded laterally; sulcus eustachii opens ventrally rather than posteroventrally; fossa precolumellaris absent; sulcus eustachii without lateral process; premaxilla narrow; fenestra postotica not subdivided; tuberculum basioccipitale smaller; lateral surface of maxilla convex rather than concave; cavum tympani shallow rather than deep.

ETYMOLOGY: Presumably in allusion to the Congo River, adjacent to the type locality.

REFERRED MATERIAL: Uncataloged skull (figs. 187–189) in the MRAC collections, ‘‘Landana’’ presumably Paleocene. Although there is a report (Wood, personal commun.) that shell material was found with this skull, we have not seen it. The identification of this skull with the type shell material is based only on their common occurrence at the same locality.

The following specimens described in Wood (1975), all from the Montian Paleocene, Landana cliffs, Cabinda: Uncataloged MRAC: an eighth cervical vertebra, parts of the carapace including the nuchal, first and second neurals, the pygal, the first and fifth pleurals, and various peripherals (first through third, eighth, ninth, and eleventh) and a pelvis ( Wood, 1975: pl. 6; all listed, without numbers, in Dollo, 1924); MRAC 3086A, two adjacent peripherals; MRAC 4794 ( Wood, 1975: pls. 3, 4), a nearly complete plastron lacking the right xiphi- and hypoplastron, the lateral portion of the right hyoplastron, and medial parts of the left hypo- and hyoplastron; MRAC 4795 ( Wood, 1975: pls. 1, 2), the posterior third of a carapace; MRAC 6316, fragment of a peripheral; MRAC 6320 ( Wood, 1975: pl. 6), posterior portion of a pair of xiphiplastra with associated left pelvis; MRAC 6322, nearly complete left xiphiplastron; MRAC 6323, fragment of a right xiphiplastron; MRAC 6328, fragment of the proximal end of a pleural; MRAC 6329, fragment of a peripheral; MRAC 6337 ( Wood, 1975: pl. 6), complete pair of xiphiplastra; and MRAC 6340, fragment of a peripheral. Wood (1975) also referred the following undeterminable shell fragments to this species on the basis of their size and surface texture: MRAC 6295, 6313–6315, 6317, 6319, 6321, 6324–6326, 6331–6335, 6338, 6339, 6341– 6344, 16024, and 16025.

MRAC 3090, lower jaw (fig. 250; Dollo, 1924: fig. 1; Wood, 1973: pl. 3), bed 12, Landana, Cabinda ( Wood, 1973), originally identified as Bantuchelys congolensis by Dollo (1924), then redescribed as a lophochelyine toxochelyid by Wood (1973). However, in light of the narrow triturating surfaces of the lower jaw in the only other Taphrosphyini known with a jaw, Rhothonemys ; we think that there is a good basis for identifying MRAC 3090 as Taphrosphys congolensis (see below).

PREVIOUS WORK: First named Podocnemis congolensis by Dollo in 1912, without a specimen or diagnosis. The actual description dates from Dollo (1913), the first time the name was used legally ( Wood, 1975). Dollo (1913) provided figures of shell pieces and a rudimentary description. Dollo was correct in relating the form to Podocnemis - like Pelomedusoides, but he apparently did it on the basis of an axillary musk duct, a character also found in cryptodires. Dollo (1924) identified a lower jaw as belonging to his new genus Bantuchelys , which he created for congolensis . Wood (1973) identified the lower jaw as a toxochelyid (which we do not agree). Wood (1975) considerably increased knowledge of congolensis by describing the plastron, much of the carapace, and reassigning the species to Taphrosphys on the basis of plastral morphology (with which we agree).

DISCUSSION: The skull identified here as Taphrosphys congolensis (MRAC uncataloged) was originally associated with a shell fragment (Wood, personal commun.) having the typical Taphrosphys surface texture. The skull is clearly very similar to skulls of Taphrosphys sulcatus , and the most common shells in the Landana sequence are very similar to Taphrosphys sulcatus , which is the basis for the identification.

The lower jaw described by Dollo (1924) as the pleurodire ‘‘ Bantuchelys congolensis ’’ and by Wood (1973) as a possible toxochelyid is probably the lower jaw of Taphrosphys congolensis . We agree with Dollo’s identification and corroborate it with new evidence. The association of the jaw in the same beds as the skull and shell of Taphrosphys congolensis supports this idea as argued by Dollo (1924), but as Wood (1973) demonstrated, there are also sea turtles in these units. The discovery of a lower jaw of a closely related member of the tribe Taphrosphyini , the Moroccan Rhothonemys brinkmani , adds new support to Dollo’s contention. The very narrow lower jaw, MRAC 3090, inconsistent with African pleurodires then known to Wood (1973), was a reason for Wood to reject Dollo’s assertion. The discovery of the Rhothonemys lower jaw, however, shows that at least one Taphrosphyini also has very narrow lower jaws. Furthermore, MRAC 3090 has a large processus retroarticularis, absent in chelonioids. We have not directly examined MRAC 3090 and cannot present a detailed description of it, but based on figures in Dollo (1924) and Wood (1973) (reproduced here as fig. 250) we can list the following similarities between these specimens: very narrow triturating surface; narrow jaw rami; symphyseal hook; processus retroarticularis present; and V-shaped sulcus cartilaginis meckelii. These similarities are consistent with MRAC 3090 being identified as a Taphrosphyini , but they are not conclusive. We think that Dollo’s original identification is well enough supported to include this jaw in the dataset for the few characters that it provides. Exclusion of it does not alter the MPC.

Another lower jaw that may belong to the tribe Taphrosphyini was described by Bardet et al. (2000: 281, fig.7d, e) as a ‘‘Chelonioidea gen. and sp. indet.’’ Associated with this jaw are shell elements also described and figured ( Bardet et al., 2000: 281, fig.7a–c, g), one of which (fig. 7a) has the iliac scar small, round, and at the shell margin, features probably diagnostic for the tribe Taphrosphyini . The lower jaw is very similar to those of Rhothonemys and Taphrosphys congolensis (see figs. 248–250).

See table 17 for a comparison of the species in Taphrosphys .

Labrostochelys , new genus

TYPE AND ONLY INCLUDED SPECIES: Labrostochelys galkini , n. gen. et sp.

DISTRIBUTION: Paleocene of Morocco.

ETYMOLOGY: Labrostos, Greek for rushing furiously, in allusion to its sharply pointed head.

DIAGNOSIS: A bothremydid pleurodire of the tribe Taphrosphyini with these unique characters among Taphrosphyini : skull very long and narrow with an elongate, tapering preorbital region; extremely long prefrontal, maxilla, and premaxilla; premaxilla with anterior projection extending anterior to labial ridge unique among turtles; apertura narium externa partially or completely divided by prefrontal and premaxilla; long squamosal projection extending posteriorly to an extent unique among turtles (except for some trionychoids); large, triangular basisphenoid nearly separating pterygoids; interorbital distance relatively shorter than in any other Taphrosphyini ; fenestra postotica more horizontal than in other Taphrosphyini ; processus trochlearis pterygoidei parasagittal and very small; other differentiating characters are: narrow jugal, in contrast to Azabbaremys and Phosphatochelys ; squamosal with vertical flange, as in Taphrosphys and Phosphatochelys and in contrast to Azabbaremys ; triturating surface a broadly curved trough, as in Taphrosphys and in contrast to all other Taphrosphyini ; labial ridge on maxilla thin, as in Taphrosphys and in contrast to all other Taphrosphyini ; sulcus eustachii with dorsal process; foramen posterius canalis carotici interni formed almost completely by quadrate with small contribution of pterygoid; posteroventrally opening pocket on posterior surface of quadrate, as in Taphrosphys ; postorbital with medial process; sulcus palatinopterygoideus wide; ventrally opening channel at back of skull, as in Taphrosphys ; condylus mandibularis well anterior to condylus occipitalis, as in Taphrosphys ; fossa pterygoidea absent.

DISCUSSION: See table 16 for a comparison of the genera in the tribe Taphrosphyini .

Labrostochelys galkini , new species

TYPE SPECIMEN: AMNH 30043, a nearly complete skull (figs. 192, 193, 195, 287) purchased from M. Hammer, 1998.

TYPE LOCALITY: Phosphates near Oued Zem, Ouled Abdoun Basin, Morocco (figs. 14–16).

HORIZON: Presumed to be Tertiary based on matrix.

DEPOSITIONAL ENVIRONMENT: Near-shore marine; discussion of Moroccan phosphate deposits is under Araiochelys hirayamai .

DIAGNOSIS: As for genus.

ETYMOLOGY: For Judy Galkin, in appreciation of her years of efforts in the Department of Vertebrate Paleontology, AMNH, on behalf of this project.

REFERRED MATERIAL: AMNH 29984, a nearly complete skull (fig. 194), Danian based on shark teeth in matrix (Cappetta, personal commun.), near Khouribga, Ouled Abdoun Basin, Morocco, donated by H. Galiano, 1995.

PREVIOUS WORK: None.

DISCUSSION: This very narrow-jawed turtle differs considerably from the other bothremydids, resembling trionychids instead. It illustrates the remarkable degree of diversity present in the pleurodires, particularly in the near-shore seas of the African margin. This unusual species is known from two skulls, collapsed dorsoventrally but relatively well preserved. Labrostochelys is the sister taxon to Taphrosphys , Rhothonemys , Ummulisani , and Phosphatochelys (fig. 288).

Phosphatochelys Gaffney and Tong, 2003

TYPE AND ONLY INCLUDED SPECIES: Phosphatochelys tedfordi Gaffney and Tong, 2003 .

DISTRIBUTION: Eocene of Morocco.

ETYMOLOGY: In allusion to its discovery in the phosphate beds of Morocco.

REVISED DIAGNOSIS: A bothremydid pleurodire of the tribe Taphrosphyini with these unique characters among Taphrosphyini : preorbital region very short with extremely narrow dorsal process of maxilla; wide figure 8-shaped apertura narium externa lying above a premaxilla that slopes posterodorsally to anteroventrally; frontal small and widely separated from orbital margin; broad prefrontal-parietal contact; large quadrate extending anteriorly to cover half of cheek; labial ridge of maxilla deeper than in other Taphrosphyini except Rhothonemys ; quadratojugal widely separated from jugal, as in Rhothonemys ; anterolaterally facing trough developed on pterygoid, parietal, and quadrate extending anterodorsally from foramen nervi trigemini ventrolaterally to condylus mandibularis. Other differentiating characters are: squamosal with vertical flange in contrast to Azabbaremys ; palate dorsally arched, in contrast to Taphrosphys and Labrostochelys ; sulcus eustachii with dorsal process, in contrast to Azabbaremys and Labrostochelys ; foramen posterius canalis carotici interni formed by pterygoid and quadrate; postorbital lacks medial process and postorbital wall open, in contrast to Arenila and Nigeremys ; condylus mandibularis anterior to condylus occipitalis; fossa pterygoidea absent.

DISCUSSION: See table 16 for a comparison of the genera in the tribe Taphrosphyini .

Phosphatochelys tedfordi Gaffney and Tong, 2003

TYPE SPECIMEN: AMNH 30008, complete skull without lower jaws (figs. 198, 199, 203, 279A, 286), gift from François Escuillie´.

TYPE LOCALITY: Oued Zem, Ouled Abdoun Basin, Morocco ( Gaffney and Tong, 2003) (figs. 14–16).

HORIZON: Ypresian phosphates, Eocene, based on shark teeth in matrix (Cappetta, personal commun.) (fig. 17; see Araiochelys

TABLE 18

Comparisons of Two Specimens of Phosphatochelys tedfordi hirayamai for discussion of Moroccan phosphates; see also table 11). The original description of AMNH 30008 stated that it was Paleocene based on shark teeth analysis. Further study of the shark teeth has shown this to be in error and that the age, again based on shark teeth (Cappetta, personal commun.), is Eocene.

DEPOSITIONAL ENVIRONMENT: Near-shore marine; discussion of Moroccan phosphate deposits is under Araiochelys hirayamai .

DIAGNOSIS: As for genus.

ETYMOLOGY: For Dr. Richard H. Tedford, in recognition of his lifelong contributions to vertebrate paleontology in general and to the American Museum of Natural History in particular.

REFERRED MATERIAL: MDEt 26, a nearly complete skull (figs. 200, 201), Ypresian phosphates, based on shark teeth (Cappetta, personal commun.), Ouled Abdoun Basin, Morocco.

PREVIOUS WORK: The species was named and described by Gaffney and Tong (2003). All the figures from that paper, plus new ones, are repeated here. The description and diagnosis have also been revised and updated with the addition of new taxa.

DISCUSSION: It is possible that the two skulls now known for Phosphatochelys represent two species. A list of differentiating characters is in table 18 and features are discussed in the descriptive section. However, in the absence of more material and the probability that at least some of these features are individual variation, we have shown restraint and not created a second species.

Ummulisani , new genus

TYPE AND ONLY INCLUDED SPECIES: Ummulisani rutgersensis , n. gen. et sp.

DISTRIBUTION: Eocene of Morocco.

ETYMOLOGY: Ummu-‘lIhsan, Arabic, ‘‘mother of integrity’’. The senior author is very much indebted to Mark Stephen Caponigro of Columbia University for suggesting this name.

DIAGNOSIS: A member of the tribe Taphrosphyini with the unique feature of a hornlike, anterodorsal process on each prefrontal. Other distinguishing features are septum orbitotemporale open and reduced to low ridge on postorbital and parietal, as in Phosphatochelys , Taphrosphys , and Azabbaremys , but in contrast to Nigeremys and Arenila ; apertura narium externa smaller than in Rhothonemys , but similar in size to Phosphatochelys ; preorbital part of skull short, in contrast to Taphrosphys and Labrostochelys ; triturating surface unique in having very deep labial ridge beneath orbit with very low to absent labial ridge beneath apertura narium externa; labial ridge and maxilla very thin, as in Labrostochelys and in contrast to Phosphatochelys and Rhothonemys ; wide quadrate-basisphenoid contact, as in Taphrosphys and in contrast to all other Taphrosphyini ; foramen posterius canalis carotici interni formed entirely by quadrate, as in Labrostochelys , but in contrast to all other pleurodires.

DISCUSSION: This genus, now known from three skulls, one with a plastron, is one of the more unusual pleurodires. Ummulisani has small hornlike processes on the prefrontals, and these may have borne a scale that would make the process larger in life, as in the squamosal horns of meiolaniids. This is clear evidence for intense mating battles and burrowing. The phylogenetic analysis resolves Ummulisani as the sister taxon to Phosphatochelys .

See table 16 for a comparison of the genera in the tribe Taphrosphyini .

Ummulisani rutgersensis , new species

TYPE SPECIMEN: AMNH 30563, skull, lacking palate (figs. 206, 207), purchased from Adam Aaronson.

TYPE LOCALITY: ‘‘Mrah Iaresh, 20 km south east of Ouled Boali’’ (from Adam Aaronson), Morocco (figs. 14–16).

HORIZON: ‘‘Eocene Phosphates, Upper Ypresian, Couche O’’ (from Adam Aaronson); see figure 17 and Araiochelys hirayamai for discussion of Moroccan phosphates (see table 11).

DEPOSITIONAL ENVIRONMENT: Near-shore marine (see Araiochelys hirayamai for discussion of Moroccan phosphates).

DIAGNOSIS: As for genus.

ETYMOLOGY: For Rutgers, the State University of New Jersey, in gratitude to the faculty of the Department of Geology, Rutgers College, New Brunswick, who from 1961 to 1965 provided the senior author with inspiration, encouragement, and friendship, as well as with an education.

REFERRED MATERIAL: AMNH 30562, skull and plastron (figs. 268, 269), Paleogene phosphates, Mrah Iahresh, Morocco; AMNH 30569, skull, Ypresian (based on shark teeth; Cappetta, personal commun.), phosphates, Oued Zem, Ouled Abdoun Basin, Morocco.

PREVIOUS WORK: None.

DISCUSSION: See above.

Rhothonemys , new genus

TYPE AND ONLY INCLUDED SPECIES: Rhothonemys brinkmani , n. gen. et sp.

DISTRIBUTION: Paleogene of Morocco.

ETYMOLOGY: Rhothon, Greek for nose, beak, in allusion to the gigantic apertura narium externa.

DIAGNOSIS: A bothremydid pleurodire of the tribe Taphrosphyini with these unique characters among Taphrosphyini : apertura narium externa larger than in any other bothremydid; maxilla deeper and longer than in any other bothremydid; anterior half of skull deeper with respect to rest of skull than in any other bothremydid; labial ridge thick in cross section with broadly curved outer surface and slightly concave inner surface. Other differentiating features are: parietal enters orbital margin, in contrast to all bothremydids except Phosphatochelys and Ummulisani ; squamosal with vertical flange, in contrast to Azabbaremys ; frontal shorter than prefrontal (also in Phosphatochelys ), enters orbit for its full length; interorbital width narrower than in other Taphrosphyini except in Labrostochelys ; parietal forms major part of postorbital ridge and pocket, as in Taphrosphys and Phosphatochelys .

DISCUSSION: See table 16 for a comparison of the genera in the tribe Taphrosphyini .

Rhothonemys brinkmani , new species

TYPE SPECIMEN: AMNH 30521 (figs. 209–211), partial skull, lacking palate and basicranium, and lower jaw.

TYPE LOCALITY: Ouled Abdoun Basin, Morocco, based on matrix composition and included fossils.

HORIZON: Within the Paleogene phosphate sequence, based on matrix composition and included fossils (see table 11).

DEPOSITIONAL ENVIRONMENT: Near-shore marine: discussion of Moroccan phosphate deposits is under Araiochelys hirayamai .

DIAGNOSIS: As for genus.

ETYMOLOGY: In recognition of the contributions of Dr. Donald Brinkman, Royal Tyrrell Museum of Palaeontology, to the field of chelonian paleontology and evolution.

REFERRED MATERIAL: None.

PREVIOUS WORK: None.

DISCUSSION: Although the only known specimen of this taxon is an incomplete skull, lacking most of the palate and basicranium, the preserved areas are so different from other pleurodires that it is easily diagnosed. There are also sufficient characters to resolve it in the cladogram (fig. 288) as the sister taxon to Phosphatochelys + Ummulisani .

Azabbaremys Gaffney, Moody, and Walker, 2001

TYPE AND ONLY INCLUDED SPECIES: Azabbaremys moragjonesi Gaffney, Moody, and Walker, 2001 .

DISTRIBUTION: Paleocene of eastern Mali. ETYMOLOGY: Azabbar, a monster in popular Mali folk stories in the Tamasheq language. Thanks to Mr. Ibrahim Litny for suggesting this reference.

REVISED DIAGNOSIS: A bothremydid pleurodire of the tribe Taphrosphyini with these unique characters among Taphrosphyini : triturating surfaces covered with prominent toothlike crenellations forming a corrugated surface; prefrontal extending anteriorly to anterior edge of premaxilla; deep premaxilla with anterior surface sloping anterodorsal to posteroventral; skull roof broadly convex to a greater degree than in any other Taphrosphyini . Other differentiating characters are: short, wedge-shaped skull higher than in Arenila and Nigeremys , differing from the short skull of Phosphatochelys by the presence of a prominent skull roof convexity; broad jugal exposure in orbit, as in Phosphatochelys and Arenila but in contrast to Taphrosphys and Labrostochelys ; squamosal without vertical flange in contrast to Taphrosphys , Labrostochelys , and Phosphatochelys ; labial ridge thicker than in Taphrosphys and Labrostochelys but thinner than in Arenila and Nigeremys ; no maxilla-vomer contact; dorsally arched palate in contrast to Taphrosphys and Labrostochelys ; foramen posterius canalis carotici interni formed by pterygoid and quadrate; postorbital lacking medial process; sulcus palatinopterygoideus wide; vomer narrow; condylus mandibularis anterior to condylus occipitalis; basisphenoid solid triangular not excavated posteriorly; postorbital wall open; fossa pterygoidea absent.

PREVIOUS WORK: See species for Previous Work.

DISCUSSION: See table 16 for a comparison of the genera in the tribe Taphrosphyini .

Azabbaremys moragjonesi Gaffney, Moody, and Walker, 2001

TYPE SPECIMEN: BMNH R 16370, a complete skull lacking lower jaws (figs. 214–218, 280, 281C, 286A).

TYPE LOCALITY: North of In Fargas near Samit, eastern Mali (see Moody and Sutcliffe, 1990, 1991, 1993).

HORIZON: Teberemt Formation, Paleocene (see Moody and Sutcliffe, 1990, 1991, 1993, 1995).

DEPOSITIONAL ENVIRONMENT: Shallow marine ( Moody and Sutcliffe, 1993).

DIAGNOSIS: As for genus.

ETYMOLOGY: For Ms. Morag Jones, a student who participated in the discovery of this specimen; she died tragically on the first Mali expedition ( Gaffney, Moody, and Walker, 2001).

REFERRED MATERIAL: None.

PREVIOUS WORK: This taxon was named and described by Gaffney, Moody, and Walker (2001); the diagnosis and description is updated and revised here.

DISCUSSION: Azabbaremys is the sister taxon to another Mali form, the undescribed CNRST-SUNY 199. Together they are weakly supported as the sister group to the remaining Taphrosphyina .

UNDESCRIBED TAXON: CNRST SUNY 199

SPECIMEN: CNRST SUNY 199, a nearly complete skull, figured in Gaffney, Roberts, Sissoko, Boure´, Tapanila, and O’Leary (in press).

LOCALITY: South of the Adrar des Iforas Mountains, between Saguirilidad and In Fargas, Mali.

HORIZON: Middle to upper portion of the Paleocene Teberemt Formation.

DEPOSITIONAL ENVIRONMENT: Near shore marine.

REVISED DIAGNOSIS: A bothremydid pleurodire of the subtribe Taphrosphydina with these unique characters among the tribe Taphrosphyini : small pit formed by jugal, maxilla, and palatine on triturating surface; jugal exposed on triturating surface; accessory ridge present on anterior triturating surface; wide palatine-basisphenoid contact separating pterygoids on midline; supraoccipital-quadrate contact present; basioccipital narrowly enters condylus occipitalis; palatine-jugal contact in small septum orbitotemporale; other distinguishing characters: skull relatively long and narrow in contrast to all Taphrosphyini except Labrostochelys ; fossa pterygoideus deep and narrow as in Nigeremydina but in contrast to all other Taphrosphydina ; foramen posterius canalis carotici interni formed by pterygoid, basisphenoid, and quadrate in contrast to all Taphrosphyini except Taphrosphys ; small remnant of septum orbitotemporale present consisting of ventral parietal process as in Phosphatochelys but in contrast to other Taphrosphyini .

TABLE 19

Comparison of Arenila and Nigeremys

REFERRED MATERIAL: None.

PREVIOUS WORK: Possibly Gaffney, Roberts, Sissoko, Boure´, Tapanila, and O’Leary (in press), if it is published before the present work.

DISCUSSION: This skull is named and described in a paper that will have a publication date close to the publication date of the present paper, that is, it may be published before or after the present paper. The actual dates of publication are not under the control of the authors. Therefore, we are not mentioning the name of this new taxon here, it is referred to only by its catalogue number, CNRST SUNY 199. Although the description and figures of this skull appear in Gaffney, Roberts, Sissoko, Boure´, Tapanila, and O’Leary (in press), the taxon has been entered into the data set analysed here. The taxon appears as CNRST SUNY 199 in the cladograms in figures 288– 292, 294, 296–314, 317.

SUBTRIBE NIGEREMYDINA , NEW

TYPE GENUS: Nigeremys Broin, 1977 .

INCLUDED GENERA: Nigeremys Broin, 1977 ; Arenila Lapparent de Broin and Werner, 1998 .

DIAGNOSIS: Member of tribe Taphrosphyini with the following characters contrasting with subtribe Taphrosphyina : labial ridge thicker; septum orbitotemporale completely closed; antrum postoticum completely closed (also in Azabbaremys and Ummulisani ); condylus mandibularis posterior to condylus occipitalis; fossa pterygoidea deep and narrow (also in CNRST SUNY 199); basisphenoid ventral outline V-shaped.

DISCUSSION: See table 19 for a comaparison of the genera in the Nigeremydina .