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).

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.

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.

Tables

TABLE 3. Classification and distribution of lizards of the family Diploglossidae (55 species). For common names we use modifying words from literature sources, in some cases modified for consistency following guidelines (Hedges et al. 2019). The IUCN Redlist threat status is indicated in parentheses (NA = not assessed).

Species	Distribution
Celestinae (24 sp.)
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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