Hydrochaeris gaylordi, R D E Macphee & Ronald Singer & Michael Diamond, 2000

R D E Macphee, Ronald Singer & Michael Diamond, 2000, Late Cenozoic Land Mammals from Grenada, Lesser Antilles Island-Arc, American Museum Novitates 3302, pp. 1-20 : 10-13

Hydrochaeris gaylordi , new species

HOLOTYPE: AMNH-VP 132713 ( fig. 4), partial maxilla retaining M1-M3.

DISCOVERER AND DATE OF DISCOVERY: Ronald Singer and party, 1991.

TYPE LOCALITY AND AGE: Locality 12° North, Lance aux Épines , Grenada (Lesser Antilles); probably Late Pliocene, less likely Early Pleistocene (40 K/ 40 Ar age estimate on hornblende in matrix, 2.7–3.6 Ma) .

ETYMOLOGY: For Mr. Joseph Gaylord, in recognition of his many kindnesses to RS over many years.

DISTRIBUTION: Known only from Grenada in the southern Lesser Antilles.

DIAGNOSIS: A small hydrochaerine that can be distinguished from all other named hydrochaerines, including extant Hydrochaeris hydrochaeris , by its unique occlusal pattern, in which buccal connection of maxillary M2 lamellae is never lost during ontogeny.

DISCUSSION: Although in unerupted cheek teeth of Hydrochaeris hydrochaeris the summits of individual lamellae are typically joined, reflecting the earliest stage of crown ontogeny, these connections are lost with wear. By the young juvenile stage, the notably hyposdont cheek teeth of capybaras appear to be made up of separate enamel-dentine lamellae of complicated shape, united by plaques of cementum. Although M3 may be somewhat variable in that some interlamellar connections may be preserved into later life, observational evidence indicates that M1 and M2 do not vary in this regard. Thus, preservation of a buccal connection between M2 lamellae in the Grenadian specimen appears to be a derived (albeit neotenic) feature within Hydrochaeris , justifying recognition of H. gaylordi . This feature is absent or unrecorded in the other species of Hydrochaeris recognized by Mones (1984): Lujanian H. ballesterensis (Rusconi, 1934) and extant H. isthmius (Goldman, 1912) , sometimes regarded as a valid species (cf. Mares and Ojeda, 1982).

MEGALONYCHID FOSSILS FROM GRENADA

As noted in greater detail in subsequent paragraphs, the three teeth described in this section are referable to Megalonychidae , gen. & sp. indet. The homologs of phyllophagan teeth in other placentals are not certain, and we shall follow the convention of identifying the highly trenchant first tooth as the cani- niform (CF), and the flat-wearing teeth that follow it as the molariforms (MFs).

In this section we will use the phylogenetic framework recently developed by White and MacPhee (in press). Although the grouping ‘‘Antillean members of Megalonychidae ’’ has been used as a taxonomic as well as a biogeographical concept in the past, island sloths actually fall into two subfamilies, each with two contained tribes: Megalocninae , containing Megalocnini (Megalocnus) and Mesocnini (Parocnus) ; and Choloepodinae , containing Acratocnini (Acratocnus) and Cubanocnini (Neocnus) . Extant Choloepus , the two-toed sloth, is the sister group of Acratocnus ; the placement of Paulocnus from Curaçao is still unsettled, although it is definitely a choloepodine (and probably an acratocnin).

MAXILLARY MOLARIFORM (AMNH-VP 132714): The root end of this specimen is incomplete, and there is a small amount of damage evident on its occlusal surface ( fig. 5A, B). On the whole, however, the tooth is well preserved and is immediately recognizable as a right maxillary MF of a megalonychid sloth. Although the specimen was found very close to the larger caniniform ( fig. 3, inset), they are actually isolated finds and for this reason we cannot be certain that they represent the same individual.

The first and the last of the four upper MFs of Antillean megalonychids tend to be subtriangular to ovoid in cross section, while the middle teeth tend to be reniform or meniscoid. In details of shape the Grenadian tooth strongly recalls MF2s/MF3s of the Late Quaternary Cuban megalocnines Megalocnus rodens and Parocnus (= Mesocnus ) browni . The new specimen’s smooth-walled shaft is convex on its mesial surface and concave on its distal; the latter surface is, however, interrupted by a longitudinal groove that parallels the shaft’s mesiobuccal margin ( fig. 5A). The semiserrate wear pattern on the tooth is precisely like that seen in other Antillean sloths. As in the latter, the softer dentine at the center of the tooth is hollowed out by wear, producing a central basin that deepens distolingually.

AMNH-VP 132714 is comparatively large (table 3), and its measurements are close to those cited by Matthew and Paula Couto (1959) for MF2s/MF3s of Parocnus browni . Paulocnus petrifactus from Curaçao is also fairly large for an Antillean sloth (Hooijer, 1962, 1964), but the MF2 and MF3 of this species are not known. The Grenadian specimen is seemingly too large to belong to any recognized species of Acratocnini or Cubanocnini as reorganized by White and MacPhee (in press).

MAXILLARY CANINIFORMS (AMNH-VP 132715 and 132716): There are two maxillary caniniforms (both from the right side) in the sample from Locality 12° North ( fig. 6A– D). These teeth are typically megalonychid in being trigonal, stout, and trenchant, with lengthy wear facets on distal surfaces. However, they differ greatly in size (table 3). The larger caniniform is the size of that of a large Acratocnus (e.g., A. odontrigonus AMNH- VP 17715, M × W = 11.1 mm) while the small tooth is more like that of Neocnus (e.g., N. comes UF 76356, M × W = 7.7 mm). The teeth of extinct megalonychids were nonreplacing and ever-growing, and the evidence is clear that they increased in size during life in conformity with jaw growth (cf. Simpson in Matthew and Paula Couto, 1959: 54). Accordingly, it is often difficult or impossible to decide, when examining isolated megalonychid teeth of different sizes, wheth- er the specimens represent different species or merely different growth stages of the same species. As there is no evidence either way in the present case, we must leave the question undecided.

In discrete features, the CFs from Locality 12° North are rather nondescript, although they are primitive and thus most similar to those of acratocnins and cubanocnins. They were obviously protruding; Naples (1982) has pointed out that CFs extending well beyond the occlusal plane are generally primitive for sloths. The angles at which the three faces meet are slightly rounded, the one bordering the mesial side of the occlusal surface being the most rounded. The tips are sharply pointed rather than chisel-edged, and the large honing facet produced by wear with the lower caniniform faces directly caudad. The Grenadian specimens lack the slight longitudinal groove seen on the external surface of the upper CF of A. odontrigonus (Anthony, 1918) as well as the ‘‘longitudinal ribbing’’ said to be distinctive of the upper CF of A. antillensis (Matthew and Paula Couto, 1959: pl. 27).

A relevant question is whether the large CF represents the same species as the MF from the same locality. Although the large CF compares well with the equivalent tooth of Acratocnus specimens from the Greater Antilles, the MF is much larger than those of any recognized member of that genus. In all Antillean sloths described to date, the maxillary caniniform is the largest tooth in the entire dentition. This is also true in living Choloepus , suggesting that the living twotoed sloth retains the symplesiomorphous condition. However, given the large size of the Grenadian MF, one would expect an even larger CF to be associated with it. As already noted, within-species ontogenetic variation in tooth dimensions is known to be marked in Antillean sloths (cf. Matthew and Paula Couto, 1959; Paula Couto, 1967), making it dif- ficult to set taxonomic boundaries on dental grounds alone (White, 1993). Obviously, the question of how many species are represent- ed in the existing sample must await the recovery of more and better material.

Comparisons to Paulocnus from Curaçao are extremely relevant, because this island is closer to Grenada than are any of the Greater Antilles, and because both of these small islands are situated approximately the same distance from the South American continental shelf. Only one maxillary CF of the Curaçao sloth has been described, from a quarry block containing portions of the skull and mandible of one individual (GIUA X 4781; Hooijer, 1964). Hooijer’s maximum-width measurement of the maxillary CF of this specimen is 11 mm, which compares well with the size of the larger CF from Grenada.

A seeming bar to further comparisons, however, is Hooijer’s (1964) assertion that the CF of Paulocnus is widest on its mesial face (as distinct from its external face, widest in all other Antillean sloths except Megalocnus whose CF is differently derived). Hooijer particularly emphasized this feature in distinguishing Paulocnus from both Acratocnus and Parocnus , which it otherwise resembles. Close study of a photographic illustration of GIUA X 4781 (Hooijer, 1964: pl. X) indicates that Hooijer’s interpretation of caniniform morphology and dimensions cannot be correct. In this plate, the fragmentary upper jaw (essentially corresponding to a palatal fragment with three teeth) rests on the lower jaw, with the first MFs in occlusion. However, the conformation of the occlusal surfaces of the depicted MFs indicates that it is the left lower first molariform that is in occlusion with the right upper, i.e., the palate has been displaced and is not in correct occlusal relationship with the mandible. That being the case, the preserved upper CF is evidently not a left, as Hooijer claimed, but instead a right, and the surface facing the view- er is not the mesial but the external. Although this clarification does not solve the problem of the phylogenetic relationships of the large Grenadian specimen, it does mean that Paulocnus is not barred from being a close relative on the basis of caniniform morphology. Indeed, in view of their metrical similarities, it is possible that they may even represent the same taxon. Once again, further resolution must await new material.

Systematic Paleontology: The three sloth teeth recovered from Locality 12° North are classifiable as follows:

MAGNORDER XENARTHRA COPE, 1889 ORDER PILOSA FLOWER, 1883 SUBORDER PHYLLOPHAGA OWEN, 1842 SUPERFAMILY MEGATHEROIDEA GRAY, 1821 FAMILY MEGALONYCHIDAE GERVAIS, 1855

The teeth are difficult to classify below the family level because they display fairly primitive morphologies. Although the molariform (AMNH-VP 132714) is generally similar to maxillary MF2s/MF3s of megalocnines, this is hardly decisive evidence that its owner was phylogenetically a megalocnine rather than a choloepodine. At the tribal level, megalocnin affiliations can be ruled out for the caniniforms because megalocnin CFs are distinctively oval or crescent-shaped in crosssection. However, a relationship with mesocnins (i.e., Parocnus ) is not so easily dismissed, since CFs are trigonal in the latter tribe as in other Antillean tribes. As already noted, because sloth cheek teeth vary conspicuously in size with ontogenetic age, it is quite possible that all three specimens represent the same species, although the presence of more than one phyllophagan in the sample cannot be ruled out based on this argument alone. In the absence of diagnostic characters, for classificatory purposes it is appropriate to consider all three teeth as Megalonychidae , gen. & sp. indet.

OTHER FOSSIL MATERIAL FROM LOCALITY 12° NORTH

A number of fragments of postcranial bones have been collected at the site, but their condition is such that no positive systematic identifications can be made. A few are clearly rib fragments belonging to large vertebrates (largest fragment, 23.7 mm wide). They may well represent the megalonychid(s) or the capybara already known from the site, or they may relate to some oth- er large vertebrate not yet discovered. (Sirenians are ruled out because the rib fragments are not osteosclerotic.) A few tiny long bone sections, unfortunately lacking articular ends, are provisionally regarded as belonging to small lizards. Their presence indicates that there is also a microfaunal component at Locality 12° North that remains completely unknown.