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Caribicus warreni (SCHWARTZ, 1970)

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Higher TaxaDiploglossidae, Diploglossa, Anguimorpha, Sauria, Squamata (lizards)
Subspecies 
Common NamesE: Hispaniolan Giant Galliwasp, Haitian giant Galliwasp 
SynonymDiploglossus warreni SCHWARTZ 1970:780
Diploglossus carraui INCHAÚSTEGUI, SCHWARTZ & HENDERSON 1985: 196
Diploglossus carraui — SCHWARTZ & HENDERSON 1991: 403
Diploglossus warreni — SCHWARTZ & HENDERSON 1991: 406
Celestus warreni — POWELL et al. 1996: 66
Celestus carraui — POWELL et al. 1996: 65
Celestus warreni carraui — HALLERMANN & BÖHME 2002
Celestus warreni warreni — HALLERMANN & BÖHME 2002
Celestus warreni — LANGNER 2019
Caribicus warreni — SCHOOLS & HEDGES 2021 
DistributionHispaniola (N Haiti), Dominican Republic, Ile de la Tortue

Type locality: Palmiste, Île de la Tortue, Département du Nord-Ouest, Haiti.

carraui: Hispaniola (N Dominican Republic); Type locality: Comedero, La Isabela, Puerto Plata Province, República Dominicana.  
Reproductionovovivparous 
TypesHolotype: AMNH 103214.
Holotype: USNM 197369 [carraui] 
DiagnosisDiagnosis (carraui): A species of Diploglossus characterized by: 1) large size (snout-vent length to 283 mm in males; females unknown); 2) throat and venter pale creamy with scattered dark brown markings, either toward the centers of ventral scales or along their edges, the markings randomly placed in each individual; 3) head scales not outlined in black; 4) nuchal scales posterior to enlarged head scales without dark pigment along median sutures so that neck region is not obviously lineate; 5) supralabial intercalated scales absent [from INCHAÚSTEGUI et al. 1985].

Definition: A species of Diploglossus of the Hispaniolan costatus complex which differs from other members of that complex (costatus, stenurus, curtissi) in much larger size (male warreni to 230 mm snout-vent length; female warreni to 227 mm snout-vent length; largest male and female of three other species those of stenurus with maximally sized male 172 mm, female 143 mm), low number of ventral scales between mental and vent ( 84 to 92 in warreni, 11 to 106 in other members of the complex combined), dorsal trunk scales striate and with a strong median keel, dorsal caudal scales likewise striate and strongly keeled, ventral scales finely striate; dorsal pattern consisting of a series of medium dark grayish chevrons, more or less confluent medially (especially anteriorly) and extending the full length of the unregenerated portion of tlie tail, on a pale tan to yellow-tan ground, sides very pale gray, flecked with dark brown and with remnants of vertical bars corresponding to lateral ends of dorsal chevrons, postocular dark mask obsolete and only very faintly indicated, iris brown, and venter immaculate cream to very pale orange tan. (Schwartz 1970: 781)

Comparisons: D. warreni requires comparison only with D. stenurus, apparently its closest relative on Hispaniola, and with D. occiduus in Jamaica, the only Antillean species which is as large as or larger than D. warreni. Comparison with occiduus^ is the more easily made (but I doubt that these two species are at all closely related); ventral scales in occiduus vary between 107 and 121, midbody scales are 49 and 50, the angular subocular lies between supralabials 8 and 9 (unilaterally in UMMZ 53251 and MCZ 74090), fourth toe lamellae vary between 16 and 23, the dorsal body and caudal scales are striate but not keeled, and the ventrals are smooth. UMMZ 53249, an adult male, has a snout-vent length of 305, head length 66.8, head width 55.5, and arm length of 69. Similar measurements on MCZ 74090, a female, are 256, 52.9, 40.5 and 57; both are very much larger and bulkier lizards than D. warreni. All specimens of D. occiduus are presently bleached, so that no data are available from them on color or pattern. Boulenger (1885:290) stated that D. occiduus was "Brownish above, with dark brown spots or cross bands." Structurally, occiduus differs most strik ingly from warreni in that the fonner has striate dorsal body and caudal scales and smooth ventrals, whereas warreni has strongly keeled dorsal body and caudal scales and finely striate ventrals. The higher number of ventral scales and midbody scales in occiduus likewise sep arates the two species; the position of the angular subocular between supralabials 8 and 9 in occiduus differs from its position between 6 and 7 in warreni. D. warreni differs from D. stenurus in several features. First, the strongly keeled dorsal body scales and the very strongly carinate caudal scales differ from the keeled condition in stenurus. Although these scales are keeled in stenurus, even in large adult individuals of the latter species the keels are much lower. This is most especially shown by the caudals in warreni; these scales have such high keels that the tail appears angulate on gross inspection. The ventral scales of warreni are finely striate, whereas these scales are smooth in stenurus. Secondly, the dorsal pattern, although reminiscent of that of stenurus, is distinctive. D. sten urus, D. costatus and D. warreni have a community of dorsal pattern elements; the pattern is composed basically of chevrons or (when these are widely opened) crossbars which extend from the nape to at least the base of the tail. In costatus the chevrons are fine and narrow, giving a herringbone pattern, whereas in stenurus the chevrons are frag mented and are made up of more or less isolated dark squares or rectangles arranged in a chevronate pattern (see Schwartz, 1964:15, figs. 1-4), which, however, lacks the clarity and diagrammatic distinct ness of the costatus pattern. Both these species likewise have, in most subspecies, a pair of anterior paramedian nuchal lines, much better delineated in D. stenurus (especially in D. s. alloeides Schwartz) than in D. costatus. The dorsal pattern of D. warreni likewise is basically chevronate, but the chevrons, since they are widely opened, are almost crossbars. Additionally, these bars are broad and entire (not fragmented as in stenurus) and may be joined medially at their apices in an irregular fashion. There is no indication of paramedian nuchal Unas, although the holotype has dark paramedian blotches on the neck. Thus, although the species warreni, stenurus and costatus are all basically chevronate, the degree and quality of tlie dorsal markings varies with the species. Thirdly, D. warreni is by far the largest member of the costatus complex in Hispaniola. Maximally sized stenurus (males first, females second in each case) are 172 and 143, maximally sized costatus measure 127 and 116, and maximally sized curtissi 86 and 82. The male warreni has a snout-vent length of 230, the larger female 227; in these measurements warreni ranks second only to occiduus in the Antilles.
Scale count differences between warreni and stenurus are difficult to assess, since there are counts available on only tliree warreni in contrast to those from several hundred stenurus. However, tlie warreni counts of ventral and midbody scales fall toward the lower extremes of these counts in D. stenurus, and in the case of midbody scales, tlie counts of warreni ( 33-37 ) lie just below the counts for stenurus ( 37-45 ) . Overlap of counts is of little significance in this group of galliwasps, since even such strikingly different species of stenurus and curtissi have almost identical extremes in ventral and midbody counts. It would be extremely pleasant if warreni differed quantitatively from stenurus in some meristic character, but such is not the case, at least in those counts which I have heretofore employed in differentiating members of this Hispaniolan complex.
One other characteristic separates warreni from stenurus. In the former, the subcaudal scales are very large and almost fanlike in aspect; this resemblance is further enhanced by the dark brown lines of pigment which may radiate from the base of each scale. Such radiations do not occur in stenurus and the ventral caudal scales are relatively (as well as actually) much smaller. Thus the subcaudals in warreni are much larger than are those in stenurus. Finally, the distinctly angulate appear ance of the tail, due to the high median keels on tlie dorsal caudal scales, does not occur in stenurus; in the latter species, although the dorsal caudals are keeled, the keels are not so high and do not impart an angulate or longitudinally keeled aspect to the upper side of the tail. (Schwartz 1970: 782) 
CommentFor illustrations see Henderson and Schwartz, 1984. HALLERMANN & BÖHME 2002 suggested to relegate several species of Celestus to subspecies status of C. warreni, in particular C. carraui and C. anelpistus. For illustrations of C. carraui see Henderson, 1988.

Synonymy: POWELL & HENDERSON (2003) synonymized C. carraui with C. warreni. The relationship of the two is still unclear as C. carraui is only known from 3 specimens (Hallermann & Böhme 2002). 
EtymologyNamed after Mr. C. Rhea Warren, who apparently helped to collect the types or helped to organize the collection trip.

C. carraui was named after Dr. José Antonio Carrau who collected the types. 
References
  • Augstenová, Barbora, Eleonora Pensabene, Lukáš Kratochvíl, and Michail Rovatsos. 2021. Cytogenetic Evidence for Sex Chromosomes and Karyotype Evolution in Anguimorphan Lizards. Cells 10, no. 7: 1612 - get paper here
  • Beolens, Bo; Michael Watkins, and Michael Grayson 2011. The Eponym Dictionary of Reptiles. Johns Hopkins University Press, Baltimore, USA - get paper here
  • Hallermann, J. & W. Böhme 2002. On giant anguids from the West Indies with special reference to Celestus warreni ( Schwartz, 1970) and its relatives from Hispaniola (Reptilia: Squamata: Anguidae). Mitt. Hamb. Zool. Mus. Inst. 99: 169-178
  • Henderson, R W. 1988. Diploglossus warreni Schwartz. Catalogue of American Amphibians and Reptiles (426 - get paper here
  • Henderson, R. W. 1988. Diploglossus carraui Inchaustegui, Schwartz, and Henderson. Catalogue of American Amphibians and Reptiles (425. - get paper here
  • Henderson, R.W., and Schwartz, A. 1984. A guide to the identification of the amphibians and reptiles of Hispaniola. Spec. PubI. Milwaukee Public Mus. Biol. and Geol. (4):1-70.
  • Inchaústegui, S. J., A. Schwartz, and R. W. Henderson. 1985. Hispaniolan giant Diploglossus (Sauria: Anguidae): Description of a new species and notes on the ecology of D. warreni. Amphibia-Reptilia 6: 195-201. - get paper here
  • Langner, Ch. 2019. Ich komme Dir auf die Schliche! Verkanntes Schleichenvolk – die interessanten Echsen der Familie Anguidae. Reptilia (Münster) 24 (136): 16-27 - get paper here
  • Powell, R. & Henderson, R.W. 2003. The taxonomic and conservation status of Giant Hispaniolan Celestus (Anguidae). Carib. J. Sci. 39: 237-240
  • Powell, R. & Incháustegui, S.J. 2009. Conservation of the herpetofauna of the Dominican Republic. Applied Herpetology 6: 103–122 - get paper here
  • Powell, R., R. W. Henderson, K. Adler, And H. A. Dundee. 1996. An annotated checklist of West Indian amphibians and reptiles. In R. Powell and R. W. Henderson (eds.), Contributions to West Indian Herpetology: A Tribute to Albert Schwartz, p.51-93. Society for the Study of Amphibians and Reptiles, Ithaca (New York). Contributions to Herpetology, volume 12. [book review in Salamandra 36 (2): 136]
  • Powell, Robert, José A. Ottenwalder, Sixto J. Incháustegui, Robert W. Henderson and Richard E. Glor. 2000. Amphibians and reptiles of the Dominican Republic: Species of special concern. Oryx 34 (2): 118-128 - get paper here
  • SCHOOLS, MOLLY & S. BLAIR HEDGES 2021. Phylogenetics, classification, and biogeography of the Neotropical forest lizards (Squamata, Diploglossidae). Zootaxa 4974 (2): 201–257 - get paper here
  • Schwartz, A. 1970. A new species of large Diploglossus (Sauria: Anguidae) from Hispaniola. Proc. Biol. Soc. Washington 82: 777-787. - get paper here
  • Schwartz, A. & Henderson, R.W. 1991. Amphibians and Reptiles of the West Indies. University of Florida Press, Gainesville, 720 pp.
 
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