Karstic Landscapes Are Foci of Species Diversity in the World’s Third-Largest Vertebrate Genus Cyrtodactylus Gray, 1827 (Reptilia: Squamata; Gekkonidae)
Abstract
:1. Introduction
2. Materials and Methods
2.1. Habitat Preferences and Ecotypes
- General (Figure 4A). Species that use the majority of the microhabitats in their immediate surroundings in whatever environment they inhabit. The microhabitats may include rocks of all types (when present), logs, tree trunks (with or without holes and crevices), and all vegetative structures of various dimensions, the ground, and human-made structures in many cases. No particular microhabitat is notably preferred over any other although some species may be most often observed in low vegetation.
- Trunk (Figure 4B). These are species generally found on the trunks and large branches of large trees at varying heights and often take refuge in cracks, crevices, or holes in the trunks. They may occasionally occur on large granite rocks but only if the rocks are near the trees. These species are generally the largest and most robust species in the genus [22,23,24]. None have been reported to have prehensile tails although some species may coil the tail horizontally similar to that seen in arboreal species.
- Karst (Figure 4C). These are generally more gracile species that are restricted to habitats where limestone rock (karst) is present. Individuals use this substrate (including cliff faces, small rocks, and boulders) as well as adjacent vegetation. If caves are present, they will enter only into the twilight zone and usually no deeper than 50 m from the entrance [14]. Despite what has been written about many karst-associated species being cave species or cave adapted (e.g., [25]), none truly are and most are more commonly found on the outside of caves (see below). These species do not occur in habitats lacking karstic substrates.
- Cave (Figure 4D). These are species that occur exclusively in the cave-like environments formed by large granite boulders. Open spaces between the boulders can be quite extensive and contain areas where very little light penetrates. These species rarely occur on the out-facing (i.e., the forest-side) surfaces of the boulders and for the most part, are restricted to the spaces between the boulders at varying depths below the surface of the ground in extremely low levels of illumination. These are truly cave-adapted species with notably thin, gracile bodies, long limbs, flat heads, large eyes, and faded color patterns [13,26,27].
- Terrestrial (Figure 4E). These are species that generally occur only on the ground and may take refuge beneath natural and human-made surface objects. They may occasionally be found on the tops of small rocks (when present) or on the bases of small trees and shrubs but never higher than 1 m above the ground. These species are relatively small and notably squat, with short fat tails, thick heads, and short digits [28,29].
- Arboreal (Figure 4F). These are cryptically colored species [30,31] generally restricted to small branches, leaves, trunks of varying sizes, and shrubs. Some may take refuge beneath exfoliating bark often as high or higher than three meters above the ground. These species are rarely observed on the ground or lower than 1.5 m above the ground. In such instances, it is usually during windy and/or rainy nights (perhaps forced down from higher up; [32]; authors pers. obs.) or during egg laying. All species have a prehensile tail used as a climbing aid [31,32,33] that is often carried in a coiled, elevated position.
- Granite (Figure 4H). These are generally more robust, strongly tuberculated species found in forested habitats bearing large granite boulders (not just small, scattered, granite rocks or rocks of other types). Vegetation is often used, especially by hatchlings and juveniles, but individuals occur more commonly on the granite boulders in all planes of orientation. These species do not occur in forested areas lacking granite boulders.
- Intertidal (Figure 4I). This category contains a single species that occurs exclusively in the rocky intertidal zones of small islands in the Seribuat Archipelago off the southeastern coast of Peninsular Malaysia and avoids nearby forested regions even if they lack other species of Cyrtodactylus [19,36].
- Sandstone (Figure 4J). This category was not included in Grismer et al. [6]. It contains a single species endemic to a forested sandstone massif isolated in the lowlands of northwestern Cambodia [11]. This species is known to forage only on the surface or within crevices of sandstone rocks and was not observed on the nearby vegetation [37]. This species is similar in body shape to closely related granite-associated species (Grismer unpublished).
2.2. Mitochondrial DNA
2.3. Phylogenetic Analyses
2.4. Ancestral State Reconstruction
3. Results
4. Discussion
5. Conclusions
Supplementary Materials
Author Contributions
Funding
Institutional Review Board Statement
Informed Consent Statement
Data Availability Statement
Acknowledgments
Conflicts of Interest
References
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Grismer, L.; Wood, P.L.; Poyarkov, N.A.; Le, M.D.; Karunarathna, S.; Chomdej, S.; Suwannapoom, C.; Qi, S.; Liu, S.; Che, J.; et al. Karstic Landscapes Are Foci of Species Diversity in the World’s Third-Largest Vertebrate Genus Cyrtodactylus Gray, 1827 (Reptilia: Squamata; Gekkonidae). Diversity 2021, 13, 183. https://doi.org/10.3390/d13050183
Grismer L, Wood PL, Poyarkov NA, Le MD, Karunarathna S, Chomdej S, Suwannapoom C, Qi S, Liu S, Che J, et al. Karstic Landscapes Are Foci of Species Diversity in the World’s Third-Largest Vertebrate Genus Cyrtodactylus Gray, 1827 (Reptilia: Squamata; Gekkonidae). Diversity. 2021; 13(5):183. https://doi.org/10.3390/d13050183
Chicago/Turabian StyleGrismer, Lee, Perry L. Wood, Nikolay A. Poyarkov, Minh D. Le, Suranjan Karunarathna, Siriwadee Chomdej, Chatmongkon Suwannapoom, Shuo Qi, Shuo Liu, Jing Che, and et al. 2021. "Karstic Landscapes Are Foci of Species Diversity in the World’s Third-Largest Vertebrate Genus Cyrtodactylus Gray, 1827 (Reptilia: Squamata; Gekkonidae)" Diversity 13, no. 5: 183. https://doi.org/10.3390/d13050183