Celestinae, Schools & Hedges, 2021

Schools, Molly & Hedges, S. Blair, 2021, Phylogenetics, classification, and biogeography of the Neotropical forest lizards (Squamata, Diploglossidae), Zootaxa 4974 (2), pp. 201-257 : 212-217

publication ID	https://doi.org/10.11646/zootaxa.4974.2.1
publication LSID	lsid:zoobank.org:pub:0CCA430E-5601-42CB-847F-87B22BFD3112
persistent identifier	https://treatment.plazi.org/id/DA66BA10-FFD5-FFE9-0DF1-0CFC03C8D3E4
treatment provided by	Plazi (2021-05-20 11:24:58, last updated 2021-05-20 11:39:49)
scientific name	Celestinae
status	subfam. nov.

Treatment

Subfamily Celestinae subfam. nov.

Caribbean Forest Lizards

Figs. 5–6 View FIGURE 5 View FIGURE 6

Type genus. Celestus Gray, 1839 .

Diagnosis. Members of this subfamily have (1) claw sheath, present or absent, (2) contact between the nasal and rostral scales, absent, (3) scales in contact with the nasal scale, four, (4) postnasal scales, one, (5) position of the nostril in the nasal scale, central, (6) keels on dorsal body scales, present or absent, (7) digits per limb, four or five, (8) longest toe lamellae, 8–26, (9) dorsal scale rows, 80–233, (10) relative head width, 8.52–20.0, (11) relative rostral height, 37.6–66.5, (12) relative frontonasal length, 1.47–3.65, (13) relative interparietal distance, 0–1.42, (14) relative axilla-groin distance, 49.7–71.4.

The subfamily Celestinae subfam. nov. is distinguished from the other two subfamilies by scales in contact with the nasal scale (four versus 5–6 in Diploglossinae and Siderolamprinae ), the postnasal scales (one versus two in Siderolamprinae ), and the position of the nostril in the nasal scale (central versus posterior in Diploglossinae and Siderolamprinae ).

Content. Twenty-four currently recognized species in seven genera ( Table 3): Advenus gen. nov., Caribicus gen. nov., Comptus gen. nov., Celestus Gray, 1839 , Panolopus Cope, 1862 , Sauresia Gray, 1852 , and Wetmorena Cochran, 1927 .

Distribution. One species in this subfamily occurs in eastern Panama, near the border with Colombia. All others occur on Caribbean islands, including Cayman Brac, Little Cayman, Jamaica, Navassa, and Hispaniola.

Etymology. As for the type genus.

Remarks. The subfamily Celestinae subfam. nov. is a monophyletic clade that has a support value of 100% in Bayesian and ML analyses ( Fig. 2 View FIGURE 2 ). We have defined seven genera that fall into this subfamily based on molecular and morphological evidence ( Advenus gen. nov., Caribicus gen. nov., Comptus gen. nov., Celestus , Panolopus , Sauresia , and Wetmorena ). Our phylogeny includes six of the seven genera, with Advenus gen. nov. not being present. Here, we resurrect one of the seven genera, Panolopus (Cope, 1862) . Previous authors have accepted or disregarded Sauresia and Wetmorena based on usage of the presence or absence of a claw sheath as a diagnostic characteristic trait (see above). For most of the time since being named, Sauresia and Wetmorena have been recognized as a valid genera, distinct from Celestus . The genera were synonomized with Celestus by Hass et al. (2001) because their study using immunological data found relationships that differed from previous studies ( Savage & Lips 1993). Advenus montisilvestris ( Myers 1973) ; Pirre Mountain Forest Lizard (DD) Panama

Caribicus anelpistus ( Schwartz et al. 1979) ; Altagracia Giant Forest Lizard (CR) Hispaniola

Caribicus darlingtoni ( Cochran 1939) ; Hispaniolan Striped Forest Lizard (EN) Hispaniola

Caribicus warreni ( Schwartz 1970) ; Hispaniolan Giant Forest Lizard (VU) Hispaniola, Ile de la Tortue

Celestus barbouri ( Grant 1940a) ; Limestone Forest Lizard (EN) Jamaica

Celestus crusculus ( Garman 1887) ; Jamaican Forest Lizard (LC) Jamaica

Celestus duquesneyi ( Grant 1940b) ; Blue-tailed Forest Lizard (CR) Jamaica

Celestus fowleri ( Schwartz 1971) ; Bromeliad Forest Lizard (VU) Jamaica

Celestus hewardii ( Gray 1845) ; Red-spotted Forest Lizard (EN) Jamaica

Celestus macrolepis ( Gray 1845) ; Large-scaled Forest Lizard (NA) Jamaica

Celestus macrotus ( Thomas & Hedges 1989) ; La Selle Forest Lizard (EN) Hispaniola

Celestus microblepharis ( Underwood 1959) ; Small-eyed Forest Lizard (CR) Jamaica

...Continued onthe next page

Species Distribution

Celestus molesworthi ( Grant 1940b) ; Portland Coast Forest Lizard (EN) Jamaica

Celestus occiduus ( Shaw 1802) ; Jamaican Giant Forest Lizard (CR) Jamaica

Celestus striatus ( Gray 1839) ; Golden Forest Lizard (NA) Jamaica

Comptus badius ( Cope 1868) ; Navassa Forest Lizard (LC) Navassa Island

Comptus maculatus ( Garman 1887) ; Lesser Cayman Forest Lizard (EN) Cayman Islands

Comptus stenurus (Cope 1862) ; Hispaniolan Keeled Forest Lizard (LC) Hispaniola, Ile-a-Vache, Ile a Cabrit,

and Ile Grande Cayemite

Panolopus costatus (Cope 1862) ; Hispaniolan Smooth Forest Lizard (LC) Hispaniola

Panolopus curtissi ( Grant 1951) ; Hispaniolan Khaki Forest Lizard (LC) Hispaniola

Panolopus marcanoi ( Schwartz & Incháustegui 1976) ; Pico Duarte Forest Lizard Hispaniola

(LC)

Sauresia sepsoides ( Gray 1852) ; Common Four-toed Forest Lizard (LC) Hispaniola

Wetmorena agasepsoides ( Thomas 1971) ; Serpentine Four-toed Forest Lizard Hispaniola

(EN)

Wetmorena haetiana ( Cochran 1927) ; Earless Four-toed Forest Lizard (EN) Hispaniola

Diploglossinae (16 sp.)

Diploglossus delasagra (Cocteau & Bibron 1838) ; Cuban Pale-necked Forest Cuba

Lizard (LC)

Diploglossus fasciatus ( Gray 1831) ; Banded Forest Lizard (LC) Brazil

Diploglossus garridoi ( Thomas & Hedges 1998) ; Cuban Small-eared Forest Cuba

Lizard (NT)

Diploglossus lessonae ( Peracca 1890) ; Brazilian Forest Lizard (LC) Brazil

Diploglossus microlepis ( Gray 1831) ; Small-scaled Forest Lizard (NA) Unknown

Diploglossus millepunctatus ( O’Shaughnessy 1874) ; Malpelo Island Forest Malpelo Island

Lizard (LC)

Diploglossus monotropis ( Kuhl 1820) ; Yellow-headed Forest Lizard (LC) Colombia, Costa Rica, Ecuador, Nicaragua, Panama

Diploglossus montisserrati ( Underwood 1964) ; Montserrat Forest Lizard (CR) Montserrat

Diploglossus nigropunctuatus ( Barbour & Shreve 1937) ; Cuban Spotted Forest Cuba

Lizard (LC)

Diploglossus pleii ( Duméril & Bibron 1839) ; Puerto Rican Forest Lizard (LC) Puerto Rico

Ophiodes enso ( Entiauspe-Neto et al. 2017) ; Pelotas Glass Lizard (NA) Brazil

Ophiodes fragilis ( Raddi 1826) ; Yacupoi Glass Lizard (NA) Argentina, Bolivia, Brazil, Paraguay

Ophiodes intermedius ( Boulenger 1894) ; Asuncion Glass Lizard (LC) Argentina, Bolivia, Paraguay, Uruguay

Ophiodes luciae ( Cacciali & Scott 2015) ; Pale-striped Glass Lizard (NA) Paraguay

Ophiodes striatus ( Spix 1824) ; Dark-striped Glass Lizard (LC) Brazil, Paraguay, Uruguay

Ophiodes vertebralis ( Bocourt 1881) ; Uruguayan Glass Lizard (NA) Argentina, Brazil, Uruguay

Siderolamprinae (15 sp.)

Mesoamericus bilobatus ( O’Shaughnessy 1874) ; Talamancan Forest Lizard (LC) Costa Rica, Nicaragua, Panama

Siderolamprus adercus ( Savage et al. 2008) ; Panamanian Forest Lizard (DD) Panama

Siderolamprus atitlanensis ( Smith 1950) ; Atitlan Forest Lizard (DD) El Salvador, Guatemala, Mexico

Siderolamprus bivittatus ( Boulenger 1895) ; Two-striped Lesser Forest Lizard El Salvador, Guatemala, Honduras,

(EN) Nicaragua

Siderolamprus cyanochloris ( Cope 1894) ; Irazu Forest Lizard (LC) Costa Rica

...Continued onthe next page Species Distribution

Siderolamprus enneagrammus ( Cope 1861) ; Huasteca Lesser Forest Lizard (LC) Mexico

Siderolamprus hylaius ( Savage & Lips 1993) ; Costa Rican Forest Lizard (NT) Costa Rica

Siderolamprus ingridae ( Werler & Campbell 2004) ; Veracruz Forest Lizard (DD) Mexico

Siderolamprus laf ( Lotzkat et al. 2016) ; Chiriqui Forest Lizard (NA) Panama

Siderolamprus legnotus ( Campbell & Camarillo 1994) ; Campbell’s Forest Lizard Mexico

(LC)

Siderolamprus montanus ( Schmidt 1933) ; Montane Lesser Forest Lizard (EN) Honduras

Siderolamprus orobius ( Savage & Lips 1993) ; Hortensia Forest Lizard (DD) Costa Rica

Siderolamprus owenii ( Duméril et al. 1839) ; Owen’s Forest Lizard (NA) Guatemala, Mexico

Siderolamprus rozellae ( Smith 1942) ; Mayan Forest Lizard (LC) Belize, Guatemala, Mexico Siderolamprus scansorius ( Mccranie & Wilson 1996) ; Yoro Forest Lizard (EN) Honduras

Myers (1973) used a single specimen of Advenus montisilvestris to define this species that he collected on the southeastern slope of Cerro Pirre, Serranía de Pirre, Province of Darién, Republic of Panama. Morphologically, it groups with Celestinae subfam. nov. because of contact between the nasal and rostral scale, the scales in contact with the nasal scale, the number of postnasal scales, and the position of the nostril in the nasal scale.

We have chosen to use generic names to classify species in this subfamily for several reasons. First, we do so for a more manageable classification. While the current number of species (24) is not excessive, we know that the actual number is more than twice that amount. Four species in this subfamily are not monophyletic and there are deep divergences among populations within most species ( Fig. 3 View FIGURE 3 ). Our greater sampling of populations has confirmed this, indicating that more than 30 additional species of celestines are undescribed (Schools & Hedges, unpubl.). Secondly, the times of divergence of the celestine genera (10–15 Mya) are typical of squamate reptile genera (mode of 485 genera, 13.3 Mya; Hedges et al. 2015). Thirdly, the use of genera for clades in this subfamily has been the status quo. Four of the six clades ( Celestus , Panolopus , Sauresia , and Wetmorena ) already had generic names, and three of them ( Celestus , Sauresia , and Wetmo rena) have been in use, on and off, for much of the last two centuries.

References

Barbour, T. & Shreve, B. (1937) Novitates Cubanae. Bulletin of the Museum of Comparative Zoology, 80, 377 - 387.

Bocourt, M. F. (1881) s. n. In: Dumeril, A., Bocourt, M. F. & Moquard, F. (Eds.), Etudes sur les reptiles. In: Recherches Zoologiques pour sevir a l'Histoire de la Faune de l'Amerique Centrale et du Mexique. Mission Scientifique au Mexique et dans l'Amerique Centrale. Troisieme partie, Primiere Section. Livrason 7. Imprimerie Nationale, Paris, pp. 1 - 459.

Boulenger, G. A. (1894) List of reptiles and batrachians collected by Dr. J. Bohls near Asuncion, Paraguay. Journal of Natural History, 13, 342 - 348. https: // doi. org / 10.1080 / 00222939408677709

Boulenger, G. A. (1895) Second report on additions to the lizard collection in the Natural History Museum. Proceedings of the Zoological Society of London, 1894, 722 - 736.

Cacciali, P. & Scott, N. J. (2015) Key to the Ophiodes (Squamata: Sauria: Diploglossidae) of Paraguay with the description of a new species. Zootaxa, 3980 (1), 42 - 50. https: // doi. org / 10.11646 / zootaxa. 3980.1.2

Campbell, J. A. & Camarillo, J. L. (1994) A new lizard of the genus Diploglossus (Anguidae: Diploglossinae) from Mexico, with a review of the Mexican and northern Central American species. Herpetologica, 50, 193 - 209.

Cochran, D. M. (1927) A new genus of anguid lizards from Haiti. Proceedings of the Biological Society of Washington, 40, 91 - 92.

Cochran, D. M. (1939) Diagnoses of three new lizards and a frog from the Dominican Republic. Proceedings of the New England Zoological Club, 18, 1 - 3.

Cope, E. D. (1861) Descriptions of reptiles from tropical America and Asia. Proceedings of the Academy of Natural Sciences of Philadelphia, 12, 368 - 374.

Cope, E. D. (1868) An examination of the Reptilia and Batrachia obtained by the Orton Expedition to Ecuador and the Upper Amazon, with notes on other species. Proceedings of the Academy of Natural Sciences of Philadelphia, 20, 96 - 140.

Cope, E. D. (1894) Third addition to a knowledge of the batrachia and reptilia of Costa Rica. Proceedings of the Academy of Natural Sciences of Philadelphia, 46, 194 - 206.

Dumeril, A. M. C. & Bibron, G. (1839) Erpetologie Generale on Histoire Naturelle Complete des Reptiles. Vol. 5. Roret / Fain et Thunot, Paris, 871 pp.

Entiauspe-Neto, O. M., Quintela, F. M., Regnet, R. A., Teixeira, V. H., Silveira, F. & Loebmann, D. (2017) A new and microendemic species of Ophiodes Wagler, 1828 (Sauria: Diploglossinae) from the Lagoa dos Patos Estuary, Southern Brazil. Journal of Herpetology, 51, 515 - 522. https: // doi. org / 10.1670 / 17 - 007

Garman, S. (1887) On West Indian Geckonidae and Anguide. Bulletin of the Essex Institute, 19, 17 - 24.

Grant, C. (1940 a) II. The reptiles. In: Lynn, W. G. & Grant, C. (Eds.), The Herpetology of Jamaica. The Institute of Jamaica, Kingston, pp. 61 - 148.

Grant, C. (1940 b). Notes on the reptiles and amphibians of Jamaica, with diagnoses of new species and subspecies. In: Jamaica Today. Hazell, Watson, and Viney, London and Aylesbury, pp. 151 - 157.

Grant, C. (1951) The specific characters of the Celesti, with description of a new species of Celestus (Sauria: Anguidae). Copeia, 1, 67 - 69. https: // doi. org / 10.2307 / 1438056

Gray, J. E. (1831) A synopsis of the species of Class Reptilia. In: Griffith, E. & Pidgeon, E. (Eds.), The animal kingdom arranged in conformity with its organisation by the Baron Cuvier with additional descriptions of all the species hither named, and of many before noticed, V Whittaker, Treacher and Co., London, pp. 1 - 481 + 1 - 110.

Gray, J. E. (1839) Catalogue of the slender-tongued saurians, with descriptions of many new genera and species. Annals & Magazine of Natural History, Series 1, 2 (11), 331 - 337. https: // doi. org / 10.1080 / 00222933909512395

Gray, J. E. (1845) Catalogue of the specimens of lizards in the collections of the British Museum. British Museum, London, xxvii + 289 pp.

Gray, J. E. (1852) Description of Sauresia, a new genus of Scincidae from St. Domingo. Annals and Magazine of Natural History, 10, 281 - 282. https: // doi. org / 10.1080 / 03745485609495697

Hass, C. A., Maxson, L. R. & Hedges, S. B. (2001) Relationships and divergence times of West Indian amphibians and reptiles: Insights from albumin immunology. In: Woods, C. A. & Sergile, F. E. (Eds.), Biogeography of the West Indies: Patterns and Perspectives. 2 nd Edition. CRC Press, Boca Raton, Florida, pp. 15 - 33. https: // doi. org / 10.1201 / 9781420039481 - 11

Hedges, S. B., Marin, J., Suleski, M., Paymer, M. & Kumar, S. (2015) Tree of life reveals clock-like speciation and diversification. Molecular Biology and Evolution, 32, 835 - 845. https: // doi. org / 10.1093 / molbev / msv 037

Hedges, S. B., Powell, R., Henderson, R. W., Hanson, S. & Murphy, J. C. (2019) Definition of the Caribbean Islands biogeographic region, with checklist and recommendations for standardized common names of amphibians and reptiles. Caribbean Herpetology, 67, 1 - 53. https: // doi. org / 10.31611 / ch. 67

Kuhl, H. (1820) Beitrage zur Zoologie und vergleichenden Anatomie. Hermannsche Buchhandlung, Frankfurt, 152 pp.

Lotzkat, S., Hertz, A. & Kohler, G. (2016) A new species of Celestus (Squamata: Anguidae) from western Panama. Mesoamerican Herpetology, 3, 962 - 975. https: // doi. org / 10.1163 / 157075407780681356

McCranie, J. R. & Wilson L. D. (1996) A new arboreal lizard of the genus Celestus (Squamata: Anguidae) from northern Honduras. Revista de Biologia Tropical, 44, 259 - 264.

Myers, C. W. (1973) Anguid lizards of the genus Diploglossus in Panama, with the description of a new species. American Museum Novitates, 2523, 1 - 20.

O'Shaughnessy, A. W. E. (1874) A description of a new species of Scincidae in the collection of the British Museum. Annals & Magazine of Natural History, Series 4, 13 (76), 298 - 301. https: // doi. org / 10.1080 / 00222937408680864

Peracca, M. G. (1890) Descrizione di una nuova specie del gen. Diploglossus Wiegm. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino, 5, 1 - 5.

Raddi, G. (1826) Di alcune specie nuovi di rettili e piante brasiliana. Atti ella Societa Italiana di Scienze Moderna, 18, 1 - 39.

Savage, J. M. & Lips, K. R. (1993) A review of the status and biogeography of the lizard genera Celestus and Diploglossus (Squamata: Anguidae), with description of two new species from Costa Rica. Revista de Biologia Tropical, 41, 817 - 842.

Savage, J. M., Lips, K. R. & Ibanez, D. (2008) A new species of Celestus from west-central Panama, with consideration of the status of the genera of the Anguidae: Diploglossinae (Squamata). Revista de Biologia Tropical, 56, 845 - 859.

Schmidt, K. P. (1933) New reptiles and amphibians from Honduras. Field Museum of Natural History, 20, 15 - 22.

Schwartz, A. (1970) A new species of large Diploglossus (Sauria: Anguidae) from Hispaniola. Proceedings of the Biological Society of Washington, 82, 777 - 788.

Schwartz, A. (1971) A new species of bromeliad-inhabiting galliwasp (Sauria: Anguidae) from Jamaica. Breviora, 371, 1 - 10.

Schwartz, A. & Inchaustegui, S. I. (1976) A new species of Diploglossus (Reptilia, Lacertilia, Anguidae) from Hispaniola. Journal of Herpetology, 10, 241 - 246. https: // doi. org / 10.2307 / 1562985

Schwartz, A., Graham, E. D. Jr. & Duval, J. J. (1979) A new species of Diploglossus (Sauria: Anguidae) from Hispaniola. Proceedings of the Biological Society of Washington, 92, 1 - 9.

Shaw, G. (1802) General zoology or systematic natural history. Vol. 3. Thomas Davison, London, viii + 312 pp.

Smith, H. (1942) Mexican herpetological miscellany. Proceedings of the United States National Museum, 92, 349 - 395. https: // doi. org / 10.5479 / si. 00963801.92 - 3153.349

Smith, H. M. & Taylor, E. H. (1950) An annotated checklist and key to the reptiles of Mexico exclusive of the snakes. Bulletin of the United States National Museum, 199, 1 - 253. https: // doi. org / 10.5479 / si. 03629236.199

Spix, J. B. (1824) Animalia nova sive species nova lacertarum quas in itinere per Brasiliam annis MDCCCXVII - MDCCCXX jussu et auspiciis Maximiliani Josephi I. Bavariae Regis, suscepto, collegit et descripsit Dr. J. B. de Spix. Typis Francis Seraphici Hubschmanni, Munich, 75 pp. https: // doi. org / 10.5962 / bhl. title. 5117

Thomas, R. (1971) A new species of Diploglossus (Sauria: Anguidae) from Hispaniola. Occasional papers of the Museum of Zoology Louisiana State University, 40, 1 - 9.

Thomas, R. & Hedges, S. B. (1989) A new Celestus (Sauria: Anguidae) from the Chaine de Ia Selle of Haiti. Copeia, 4, 886 - 891. https: // doi. org / 10.2307 / 1445973

Thomas, R. & Hedges, S. B. (1998) A new anguid lizard (Diploglossus) from Cuba. Copeia, 1, 97 - 103. https: // doi. org / 10.2307 / 1447704

Underwood, G. (1959) A new Jamaican galliwasp (Sauria, Anguidae). Brevoria, 102, 1 - 13.

Underwood, G. (1964) An anguid lizard from the Leeward Islands. Breviora, 200, 1 - 10.

Werler, J. E. & Campbell, J. A. (2004) New lizard of the genus Diploglossus (Anguidae: Diploglossinae) from the Tuxtlan faunal region, Veracruz, Mexico. The Southwestern Naturalist, 49, 327 - 333. https: // doi. org / 10.1894 / 0038 - 4909 (2004) 049 % 3 C 0327: NLOTGD % 3 E 2.0. CO; 2

Figures

FIGURE 2. Phylogenetic tree of diploglossid lizards based on sequences of nine genes: four mitochondrial genes (CytB, ND2, 12S rRNA, and 16S rRNA) and five nuclear genes (AMEL, BDNF, PLPR, RAG1, and ZFP36). Maximum likelihood tree obtained from the nine-gene dataset (59 individuals; 6,949 sites).A scale bar indicates 5% sequence divergence. The numbers at nodes are ML bootstrap values, followed by Bayesian posterior probabilities; asterisks indicate significant (≥ 95%) support, and a dash or zero value indicates weak (<50%) support. The tree is rooted with Pseudopus apodus (Anguidae).

FIGURE 3. Timetree of Diploglossidae based on sequences of nine genes: four mitochondrial genes (CytB, ND2, 12S rRNA, and 16S rRNA) and five nuclear genes (AMEL, BDNF, PLPR, RAG1, and ZFP36). Nodes show divergence times in millions of years. A scale bar indicates time (6 million years ago, Mya). We show Bayesian credibility ranges as gray bars at nodes.

FIGURE 5. Head scalation of celestine genera from top (left) and side (right) views. (A–B) Caribicus warreni (ANSP 38502), HW 31.8 mm, HL 44.2 mm. (C–D) Celestus barbouri (MCZ R-45169). (E–F) Comptus stenurus alloeides (MCZ R-77152), HW 14.1 mm, HL 19.3 mm. (G–H) Panolopus curtissi hylonomus (MCZ R-77160), HW 10.1 mm, HL 13.2 mm.

FIGURE 6. Head scalation of celestine genera from top (left) and side (right) views. (A–B) Sauresia sepsoides (ANSP 38667), HW 8.15 mm, HL 9.37 mm. (C–D) Wetmorena haetiana mylica (MCZ R-77049), HW 12.0 mm.