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

Subfamily Celestinae subfam. nov.

Caribbean Forest Lizards

Figs. 5–6

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). 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

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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

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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). 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.