A New Extinct Species of Solenodon (Mammalia: Insectivora: Solenodontidae) from the Late Quaternary of Cuba
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; ANNALS OF CARNEGIE MUSEUM VOL 62, NUMDER 2, PP. 151-164 28 MAY 1993 A NEW EXTINCT SPECIES OF SOLENODON (MAMMALIA: INSECTIVORA: SOLENODONTIDAE) FROM THE LATE QUATERNARY OF CUBA GARY S. MORGAN1 J osE A. OrrENWALDER 1 Visiting Museum Specialist, Section of Mammals ABSTRACT An ex1inct species of giant Solenodon. S. arredondoi. ts described on the basis of a partial skull and a small sample of postcranial elements from three Late Quaternary fossil deposits in western Cuba. S. arredondoi can be distinguished from all other species of Solenodon by its considerably larger size. Cramal features in which S. arredondoi differs from the two Hispaniolan species of Solenodon, the living S. paradoxus and the extinct S. marcanoi, include the absence of an os proboscis, relatively broader frontals at the anterior edge of the orbits, more pronounced interorbital constriction, con stricted internal narial opening and anterior portion of pterygoid fossa, and much larger posterov entrally expanded pterygoid processes. Dental characters separating S. arredondoi from S. paradoxus are the greatly enlarged and in Hated C', lack of accessory cusps on C', and presence of a noticeable diastema between P and C'. S. arredondoi is closely related to the living Cuban Solenodon, S. cubanus, but differs from that species in its larger size, as well as the somewhat more constricted interorbital region, narrower internal narial opening, more prominent pterygoid processes, comparatively more inflated C', and broader upper molars. The diverse fauna of extinct birds and mammals collected in association with S. arredondoi indicates a Late Quaternary age (late Pleistocene or early Holocene) for this species. The giant Cuban Solenodon is one of the largest known members of the order lnsectivora, living or extinct. It was probably carnivorous, preying on larger invertebrates and a variety of small to medium-sized terrestrial vertebrates. INTRODUCTION The existence of a very large extinct species of Solenodon from Cuba was first mentioned by Arredondo ( 1970) based on a femur he collected 16 years earlier from a fossil deposit at Abra de Andres, in the Sierra de Anafe, La Habana Province, western Cuba. This femur was later figured and described in detail by Morgan et al. (1980). These authors also mentioned the existence of two other large fossil femora of Solenodon from western Cuba, one each from Cavema de Pio Domingo, Pinar del Rio Province and Cueva Paredones, La Habana Province. Both of these specimens were intermediate in size between the extremely large fossil femur from Abra de Andres and the living Cuban species, Solenodon cubanus Peters. Morgan et al. (1980) did not name a new species because they felt that a single incomplete femur was an inadequate element upon which to describe a new taxon. In early 1991 , while examining specimens in the Museo Nacional de Historia Natural in La Habana, Cuba, Oscar Arredondo discovered a partial skull 1 Florida Museum of Natural History, University of Florida, Gainesville, Florida 326 I I -2035. Submitted 6 August 1992. 151 V> -N Table I.- Cranial measurements offossi l and Recent Solenodon from Cuba. Statistics for the samples offo ssil and Recent S. cuban us are (in order) mean, standard deviation (± 1 SD). observed range, coefficient of variation, and sample size (N). Alveolar length of Anto rbital z~;goma tic Interorbital Squamosal Palatal Palatal maxillary > Species and locality constrict ion readth constric tion breadth length breadth toothrow z ;;.z Solenodon arredondoi (type) [;; Cueva Paredones 0.... MNHNC 421/123 19.0 39.0 16.3 35.4 40.7 25.2 28.0 () Solenodon cuban us (fossil) 17. 1 ± 0.5 34.3 ± 0.6 15.7 ± 0.7 31.2 35.7 ± 1.6 23 .7 ± 0.8 25.2 ± 0.6 z~ 16.8-17.8 34.0- 35.0 15.3-16.2 - 33.9-37.2 22.9- 24.3 24.5-26.0 ~ 3.3 1.7 4. 1 - 4.4 3.2 2.5 iii N = 3 N = 3 N = 2 N= l N = 4 N= 3 N= 4 a!: c Solenodon cubanus (Recent) 15.0 ± 0.7 32.4 ± 1.3 15.2 ± 0.6 30.9 ± 0.8 34.3 ± 1. 1 21.5 ± 1.2 23.7 ± 0.9 "'M 14.3-16.4 30.5- 35.2 14.5- 16.4 29.6-31.9 3 1.8-35.7 20.3- 23.9 21.8-24.7 ~ 4.8 4.0 4. 1 2.6 3.1 5.4 3.7 N= 14 N=9 N = 13 N = 12 N = 14 N = 14 N= 14 1993 MORGAN AND 0TTENWALDER- NEW EXTINCT S OLENODON FROM CUBA 153 of a very large Solenodon from a fossil deposit in Cueva Paredones. The discovery of this skull permits a thorough description of the previously unnamed giant Solenodon. We have also re-examined and measured the three large Solenodon femora mentioned in the literature, as well as several additional cranial and postcranial elements that pertain to this new species. Comparisons and measurements of the Solenodon skull from Cueva Paredones reveal that this specimen, like the femur from Abrade Andres reported by Ar redondo (1970) and Morgan et al. (1980), is considerably larger than either of the two extant species of Solenodon, S. cubanus Peters from Cuba and S. paradoxus Brandt from Hispaniola. We present a morphological description and measure ments (Tables 1-4) of this new large Solenodon, as well as comparisons with the two living and one other extinct species in the genus, S. marcanoi Patterson from Hispaniola. The Solenodon fossils from western Cuba described here belong to a new species that is not only much larger than any previously described member of the genus, but is also one of the largest known species in the order Insectivora. METHODS AND ABBREVIATIONS Ex. planations of the cranial characters and measurements of Solenodon used herein are presented by Ottenwalder ( 1991). Excellent descriptions and illustrations of the cranial anatomy and dentition of Solenodon have been published by McDowell (1958). We follow the standardized dental nomen clature for mammals (e.g., Szalay, 1969). However, the cusp homologies of the upper molars and the molariform p• of So!enodon are in question. In attempting to demonstrate a close phylogenetic re lationship between Nesophontes and Sole nodon, McDowell ( 1958) interpreted the large central cusp on the P•-M' of Solr!nodon to be the protocone. He also proposed that the metacone on these teeth was lost and that the paracone was represented by a tiny cusp located on the paracrista anterolabial to the protocone. However, most other authors (Butler, 1937, 1939; Van Valen, 1966, 1967; Hersh kovitz, 197 1) considered the primary cusp on the upper molariform teeth of S olenodon to be the paracone. According to these workers, a metacone is absent and the two cusps on the lingual cingulum are the slightly larger anterolingual protocone and the smaller posterolingual hypocone. We follow the latter dental terminology for Solenodon molariform teeth, as it appears to be the most commonlv accepted and the least controversial. It should also be noted that mammalian paleon to lo~ i s ts have Ion~ disagreed on the origin and homology of the central cusp in zalambdodont molars, such as those of Solenodon. There is also some disagreement among previous authors on the dental formula of Solendon. which lacks one tooth in each jaw quadrant compared to the primitive eutherian dentition. The missing 2 3 tooth is obviously a premolar, but it is unclear whether this is the P / , or P / 3• We follow McDowell (1958) who tentatively regarded the missing premolars as the P' /3 • Poduschka and Poduschka (1983) examined the external, cranial, and dental characters used by previous workers to distinguish Solenodon cubanus from S. paradoxus. In particular, they evaluated the validity of the genus Atopogale, established by Cabrera (1925) for S. cuba nus. It is not our intent to comment extensively on the relationship between S. cubanus and S. paradoxus. as a thorough systematic review of the genus Solenodon has already been undertaken (Ottenwalder, 1991 ). Our discussion and comparisons are primarily limited to characters that can be observed in the fossil material of the new species of Solenodon from Cuba. The following museums, institutions, and collections possess specimens of Recent and fossil So !enodon examined during this study. The abbreviations used in this paper for each of these collections are as follows: Institute de Ecologia y Sistematica, Academia de Ciencias de Cuba, Havana. Cuba (IES/ ACC);Jose A. Ottenwalder, private field collections, Santo Domingo, Dominican Republic (JAO): Museum of Comparative Zoology, Harvard University (MCZ); Museo Nacional de Historia Natural de Cuba, Havana, Cuba (MNHNC); Personal collection of Oscar Arredondo, Havana. Cuba (OA): Florida Museum of Natural History, University of Florida (UF); United States National Museum of Natural History, Smithsonian Institution (USNM). 154 ANNAlS OF CARNEGIE M USEUM VOL. 62 Table 2. -Measurements ofupper teeth offossil and Recent Solenodonfrom Cuba. Statistics for samples offossil and Recent S. cubanus are (in order) mean, standard deviation (± I SDj, observed range, coefficient of variation, and sample size (N). Species and Length Width Length Width Length locality if.~~~ c, c• P' P' P' So!enodon arrendondoi (type) Cueva Paredoncs MNHNC 421 / 123 9.7 6.2 4.0 4.0 2.9 5.2 Solenodon cubanus 4.6 ± 0.2 3.1 ± 0.1 3.4 ± 0.2 2.3 ± 0.1 3.8 ± 0.6 (fossil) 4.4-4.8 3.0-3.2 3.2-3.7 2.2-2.4 3.4-4.6 4.7 4.2 6. 1 5.1 16.2 lV = 3 N =3 N= 4 N=4 N = 3 Solenodon cubanus 8.6 ± 0.4 4.5 ± 0.3 3.0 ± 0.2 3.1 ± 0.