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Cyrtodactylus hantu DAVIS, DAS, LEACHÉ, KARIN, BRENNAN, JACKMAN, NASHRIQ, CHAN & BAUER, 2021

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Higher TaxaGekkonidae, Gekkota, Sauria, Squamata (lizards: geckos)
Subspecies 
Common NamesMalaysia (Sarawak: Kapit Division, Sarawak) 
SynonymCyrtodactylus hantu DAVIS, DAS, LEACHÉ, KARIN, BRENNAN, JACKMAN, NASHRIQ, CHAN & BAUER 20121 
DistributionMalaysia (Sarawak: Kapit Division, Sarawak)

Type locality: Pelagus Resort, Nanga Merit, Kapit Division, Sarawak, Malaysia  
Reproduction 
TypesHolotype: UNIMAS 9615
Paratypes: UNIMAS 9616; UNIMAS 9617; UNIMAS 9618; UNIMAS 9619; UNIMAS 9631; UNIMAS 9639 
DiagnosisDiagnosis: Cyrtodactylus hantu can be distinguished from all of Cyrtodactylus species by a combination of the following characters: maximum SVL of at least 73 mm; 10–12 supralabials; 9–12 infralabials; weak tuberculation on dorsal surface of body; no tubercles on ventral surface of body; 37–48 paravertebral tubercles; 13–19 longitudinal tubercle rows; 34–46 ventral scales; 19–22 subdigital lamellae on fourth toe; no femoral pores; no enlarged femoral scales; 0–6 precloacal pores; precloacal slit; blotches, indistinct dorsal body bands, and/or longitudinal stripes; no rostral chevron; and no single row of enlarged caudal scales. The species can further be distinguished using fixed genetic differences (Davis et al. 2021).

Comparisons: Cyrtodactylus hantu differs from most of their Bornean congeners by one or more morphological characteristics (Table 6). The new species is distinguished from C. baluensis (Mocquard, 1890) in having a precloacal slit as opposed to a pit, fewer precloacal pores (0–6 versus 9–12), and a lower number of paravertebral tubercles (37–48 versus 47–60); it is distinguished from C. cavernicolus Inger & King, 1961 in having fewer ventral scales (34–46 versus 51–58) and fewer subdigital lamellae on the fourth toe (19–22 versus 22–26); it is distinguished from C. consobrinus (Peters, 1871) in having a smaller maximum SVL (73 mm versus 125 mm), no reticulated pattern on the parietal, and a precloacal slit as opposed to a pit; it is distinguished from C. ingeri Hikida, 1990 in having a precloacal slit as opposed to a pit, fewer subdigital lamellae (19–22 versus 23–27), and ventral scales slightly imbricate as opposed to non-overlapping; it is distinguished from C. limajalur Davis, Bauer, Jackman, Nashriq & Das, 2019 in having a smaller maximum SVL (73 mm versus 94 mm), no enlarged femoral scales as opposed to 5–6, and a precloacal slit as opposed to a pit; it is distinguished from C. malayanus (de Rooij, 1915) in having a lower number of ventral scales (34–46 versus 58–62), a smaller maximum SVL (73 mm versus 83 mm), and no reticulated pattern on the parietal; it is distinguished from C. muluensis Davis, Bauer, Jackman, Nashriq & Das, 2019 in having a varied color pattern with dark blotches as opposed to bands, fewer precloacal pores (0–6 versus 4–5), and a smaller maximum SVL (73 mm versus 88 mm); it is distinguished from C. matsuii Hikida, 1990 in having a smaller SVL (73 mm versus 105 mm) and no single row of enlarged subcaudals; and it is distinguished from C. yoshii Hikida, 1990 in having a smaller maximum SVL (73 mm versus 96 mm), fewer subdigital lamellae on the fourth toe (19–22 versus 25–30), and fewer ventral scales (34–46 versus 50–58). Comparisons to C. miriensis and C. pubisulcus are provided below. No morphological characters consistently distinguish C. hantu from C. pubisulcus or C. miriensis. Noticeable trends separating the three lineages are seen: C. miriensis and C. hantu males can have zero precloacal pores opposed to C. pubisulcus which has at least five pores. All three species have a high degree of variation in dorsal coloration but C. hantu seems to express longitudinal stripes more frequently than C. miriensis sp. nov. or C. pubisulcus. Cyrtodactylus miriensis has a higher maximum number of paravertebral tubercles (47–57) than C. pubisulcus (41–55) and C. hantu (37–48). The combination of these features can often be used to identify each species, but there is definitive genetic data to distinguish them. Using genetic differences in our ND2 alignment, C. pubisulcus can be differentiated from the other two species by an amino acid change at base pair position 105 from a Methionine to an Alanine; C. miriensis sp. nov. can be distinguished by a three nucleotide insertion at position 633, and an amino acid shift from Alanine to Leucine at position 454 (Alignment S1) (Davis et al. 2021).

Color in life: Dorsal color of head, body, limbs, and tail brown; wide dark-brown nuchal loop that extends to the tip of the snout, edged by white line; seven dark-brown bands between nuchal loop and the posterior portion of the hindlimb insertion, each edged anteriorly and posteriorly by thin dark-brown lines; body bands wider than interspaces; limbs with light-brown band/blotch pattern; ventral portion of body bearing uniform light cream color; and tail bearing ten dark bands separated by eleven narrower grey bands dorsally, uniform beige coloration ventrally (Davis et al. 2021). 
CommentSimilar species: C. pubisulcus; C. miriensis 
EtymologyThe specific epithet hantu is in reference to the Malay word for ghost. We chose this specific epithet for two reasons: 1) the species was found around the Pelagus Resort, a now abandoned resort in the middle of the rainforest that is said to be haunted; 2) the cryptic characteristics of this species have enabled it to hide in plain sight. 
References
  • Davis, H. R., Das, I., Leaché, A. D., Karin, B. R., Brennan, I. G., Jackman, T. R., ... & Bauer, A. M. 2021. Genetically diverse yet morphologically conserved: Hidden diversity revealed among Bornean geckos (Gekkonidae: Cyrtodactylus). Journal of Zoological Systematics and Evolutionary Research - get paper here
 
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