Research Article |
Corresponding author: L. Lee Grismer ( lgrismer@lasierra.edu ) Academic editor: Uwe Fritz
© 2022 L. Lee Grismer, Attapol Rujirawan, Siriporn Yodthong, Bryan L. Stuart, Minh Duc Le, Dzung Trung Le, Yodchaiy Chuaynkern, Perry L. Wood, Jr., Anchalee Aowphol.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Grismer LL, Rujirawan A, Yodthong S, Stuart BL, Le Duc M, Trung Le D, Chuaynkern Y, Wood, Jr.PL, Aowphol A (2022) The taxonomy and phylogeny of the Cyrtodactylus brevipalmatus group (Squamata: Gekkonidae) with emphasis on C. interdigitalis and C. ngati. Vertebrate Zoology 72: 245-269. https://doi.org/10.3897/vz.72.e80615
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Abstract
Convergent morphological specializations for an arboreal lifestyle in most species of the Cyrtodactylus brevipalmatus group have been a confounding factor for establishing a stable taxonomy among its species. Recent references to C. interdigitalis from throughout Thailand and Laos were made without comparisons to the type material from Tham Yai Nam Nao, Nam Nao National Park, Phetchabun Province, Thailand, but instead, were based on general morphological similarity and distribution. The taxonomy of C. interdigitalis is stabilized here by comparing the paratypes to other specimens from Thailand and Laos and recovering their phylogenetic relationships based on newly acquired genetic data, including those from the type locality. The phylogeny recovered all specimens outside the type locality to be either C. ngati from Vietnam or new species closely related to C. ngati. Cyrtodactylus interdigitalis is shown here to be a range-restricted upland endemic on the Phetchabun massif of northern Thailand. The phylogeny also indicates that C. ngati extends hundreds of kilometers farther south into northern Thailand and central Laos. We hypothesize that the significant morphological divergence in body shape of the types of C. ngati, compared to that of the Lao and Thai populations, may be due to local adaptions for utilizing karst (C. ngati) rather than vegetation (Lao and Thai populations). Additionally, phylogenetic and multivariate analyses identified a potentially new species from Phu Hin Rong Kla National Park, Phitsanulok Province, in northern Thailand and another from the Khlong Naka Wildlife Sanctuary, Ranong Province, in southern Thailand. A series of newly examined specimens from Kaeng Krachan National Park, Phetchaburi Province, Thailand represents a possible ~82 km range extension to the southeast of C. rukhadeva. This research continues to underscore the high diversity of range-restricted upland endemics in Thailand and the importance of examining type material (if possible) in the context of a phylogeny so as to construct proper taxonomies that reveal, rather than obscure, diversity.
Convergent evolution, Gekkota, integrative taxonomy, multivariate analysis, phylogenetics, Thailand, Vietnam
The bent-toed gecko genus Cyrtodactylus is an ecologically and morphologically diverse gekkotan lineage (
Distribution of nominal species and unnamed populations and specimens of the Cyrtodactylus brevipalmatus group. A single asterisk denotes specimens in the phylogenetic tree (Fig.
The general lineage concept (GLC:
These boundaries were cross-checked using a Generalized Mixed Yule Coalescent (GMYC) approach (
The morphological data set of
Material examined in this study. Institutional abbreviations follow
Species | Catalog no. | Location | GenBank no. |
C. brevipalmatus | LSUHC 1899 | Thailand, no data | not in tree |
C. brevipalmatus | THNHM 10670 | Thailand, Nakhon Si Thammarat Province, Nopphitam District, Khao Nan National Park, Huay Lak Protected Unit | not in tree |
C. brevipalmatus | THNHM 14112 | Thailand, Nakhon Si Thammarat Province, Lan Saka District, Khao Luang National Park | not in tree |
C. brevipalmatus | AUP-00573 | Thailand, Nakhon Si Thammarat Province, Khao Ram Mt. | OK626313 |
C. brevipalmatus | LSUHC 11788 | Penisular Malaysia, Kedah State, Pulau Langkawi, Gunung Raya | not in tree |
C. cf. brevipalmatus | USMHC 2555 | Penisular Malaysia, Kedah State, Pulau Langkawi, Gunung Raya | OK626314 |
C. elok | ZRC 2.6091 | Penisular Malaysia, Pahang State, Fraser‘s Hill, the Gap | JQ889180 |
C. elok | LSUHC 12180 | Penisular Malaysia, Pahang State, near Cameron Highlands | not in tree |
C. elok | LSUHC 12181 | Penisular Malaysia, Pahang State, near Cameron Highlands | not in tree |
C. elok | ZMMU R-16144 | Malaysian pet trade, no data | not in tree |
C. interdigitalis | THNHM 20226 paratype | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | not in tree |
C. interdigitalis | THNHM 20228 paratype | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | not in tree |
C. interdigitalis | THNHM 20229 paratype | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | not in tree |
C. interdigitalis | THNMH 20227 paratype | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | not in tree |
C. interdigitalis | YC000952 | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | ON055281 |
C. cf. interdigitalis | ZMMU R-16492 | Thailand, Phitsanulok Province, Phu Hin Rong Kla National Park | MW792061 |
C. ngati | FMNH 255454 | Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation Area | JQ889181 |
C. ngati | FMNH 270493 | Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation Area | not in tree |
C. ngati | FMNH 270492 | Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation Area | OK626315 |
C. ngati | FMNH 265806 | Thailand, Loei Province, Nam San Noi River, Phu Luang Wildlife Sanctuary | JX51947 |
C. ngati | HNUE-R00111 holotype | Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, Karst forest near Pa Thom Cave | ON411220 |
C. ngati | IEBR 4829 paratype | Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, Karst forest near Pa Thom Cave | OK626318 |
C. ngati | VNUF R.2020.12 paratype | Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, Karst forest near Pa Thom Cave | OK626319 |
C. ngati | HNUE-R00112 paratype | Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, Karst forest near Pa Thom Cave | not in tree |
C. ngati | VNUF R.2014.50 | Laos, Khammoue Province, Hin Nam No National Protected Area | ON411221 |
C. cf. ngati 1 | NCSM 79472 | Laos, Xaignabouli Province, Ban Pha Liep, Houay Liep Stream | OK626316 |
C. cf. ngati 2 | ZMMU R-14917 | Vientiane Province, Laos | not in tree |
C. cf. ngati 2 | NCSM 80100 | Laos, Vientiane Province, tributary of Nam Pha River, Houay Wan Stream | OK626317 |
C. cf. rukhadeva | THNHM 24622 | Thailand, Phetchaburi Province, Kaeng Krachan National Park | not in tree |
C. cf. rukhadeva | THNHM 24838 | Thailand, Phetchaburi Province, Kaeng Krachan National Park | not in tree |
C. cf. rukhadeva | THNHM 03251 | Thailand, Phetchaburi Province, Kaeng Krachan National Park | not in tree |
C. cf. rukhadeva | THNHM 03252 | Thailand, Phetchaburi Province, Kaeng Krachan National Park | not in tree |
C. cf. rukhadeva | THNHM 03253 | Thailand, Phetchaburi Province, Kaeng Krachan National Park | not in tree |
C. cf. rukhadeva | THNHM 03254 | Thailand, Phetchaburi Province, Kaeng Krachan National Park | not in tree |
C. cf. rukhadeva | THNHM 01807 | Thailand, Phetchaburi Province, Kaeng Krachan National Park | not in tree |
C. rukhadeva | ZMMU R-16851 holotype | Thailand, Ratchaburi Province, Suan Phueng District, Khao Laem Mountain | OK626320 |
C. rukhadeva | ZMMU R-16852 paratype | Thailand, Ratchaburi Province, Suan Phueng District, Hoop Phai Tong | not in tree |
sp. 9 | AUP-01715 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Thong Pha Phum National Park | MT468909 |
sp. 13 | THNHM 00104 | Thailand, Tak Province, Umphang District, Thung Yai Naresuan Wildlife Sanctuary | not in tree |
sp. 13 | THNHM 27821 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Ban Saphan Lao | not in tree |
sp. 10 | AUP-00680 | Thailand, Tak Province, Tha Song Yang District, Mae Moei National Park, Chao Doi Waterfall | MT468902 |
sp. 14 | THNHM 01667 | Thailand, Ranong Province, Khlong Nakha Wildlife Sanctuary | not in tree |
sp. 12 | ZMMU R-16492 | Phu Hin Rong Kla National Park, Phitsanulok, Province, Thailand | ON411222 |
Morphological data included both meristic and morphometric characters. To reduce the degree of researcher bias, data were taken using the protocol of
Morphometric data taken were: snout-vent length (SVL), taken from the tip of the snout to the vent; tail length (TL), taken from the vent to the tip of the tail—original or partially regenerated; tail width (TW), taken at the base of the tail immediately posterior to the postcloacal swelling; humeral length (HumL), taken from the proximal end of the humerus at its insertion point in the glenoid fossa to the distal margin of the elbow while flexed 90°; forearm length (ForL), taken on the ventral surface from the posterior margin of the elbow while flexed 90° to the inflection of the flexed wrist; femur length (FemL), taken from the proximal end of the femur at its insertion point in the acetabulum to the distal margin of the knee while flexed 90°; tibia length (TibL), taken on the ventral surface from the posterior margin of the knee while flexed 90° to the base of the heel; axilla to groin length (AG), taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body; head length (HL), the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HW), measured at the angle of the jaws; head depth (HD), the maximum height of head measured from the occiput to base of the lower jaw posterior to the eyes; eye diameter (ED), the greatest horizontal diameter of the eye-ball; eye to ear distance (EE), measured from the anterior edge of the ear opening to the posterior edge of the bony orbit; eye to snout distance or snout length (ES), measured from anteriormost margin of the bony orbit to the tip of snout; eye to nostril distance (EN), measured from the anterior margin of the bony orbit to the posterior margin of the external nares; interorbital distance (IO), measured between the dorsomedial-most edges of the bony orbits; internarial distance (IN), measured between the external nares across the rostrum; and ear length (EL), greatest oblique length across the auditory meatus.
Meristic characters evaluated were the number of supralabial scales (SL), counted from the largest scale at the corner of the mouth or posterior to the eye, to the rostral scale; infralabial scales (IL), counted from termination of enlarged scales at the corner of the mouth to the mental scale; number of paravertebral tubercles (PVT) between the limb insertions counted in a straight line immediately left of the vertebral column; number of longitudinal rows of body tubercles (LRT) counted transversely across the body midway between the limb insertions from one ventrolateral body fold to the other; number of longitudinal rows of ventral scales (VS) counted transversely across the abdomen midway between limb insertions from one ventrolateral fold to the other; number of transverse rows of ventral scales (VSM) counted along the midline of the body from the postmentals to just anterior to the cloacal opening, stopping where the scales become granular; number of expanded subdigital lamellae on the fourth toe proximal to the digital inflection (TL4E) counted from the base of the first phalanx where it contacts the body of the foot to the largest scale on the digital inflection—the large contiguous scales on the palmar and plantar surfaces were not counted; number of small, generally unmodified subdigital lamellae distal to the digital inflection on the fourth toe (TL4U) counted from the digital inflection to the claw including the claw sheath; total number of subdigital lamellae (TL4T) beneath the fourth toe (i.e. TL4E + TL4U = TL4T); number of expanded subdigital lamellae on the fourth finger proximal to the digital inflection (FL4E) counted the same way as with TL4E; number of small generally unmodified subdigital lamellae distal to the digital inflection on the fourth finger (FL4U) counted the same way as with TL4U; total number of subdigital lamellae (FL4T) beneath the fourth toe (i.e. FL4E + FL4U = FL4T); total number of enlarged femoral scales (FS) from each thigh combined as a single metric; number of enlarged precloacal scales (PCS); number of precloacal pores (PP) in males; the number of femoral pores (FP) in males; and the number of dark body bands (BB) between the dark band on the nape and the hind limb insertions on the body. Categorical characters evaluated were the presence or absence of tubercles on the flanks (FKT; Fig.
Subcaudal scale and ventrolateral caudal fringe morphology. A Cyrtodactylus cf. rukhadeva (THNHM 03251) from Kaeng Krachan National Park, Phetchburi Province, Thailand. B Cyrtodactylus interdigitalis (THNHM 20226) from Tham Yai Nam Nao, Nam Nao National Park, Phetchabun Province, Thailand. C Cyrtodactylus sp. 13 (THNHM 00104) from Thung Yai Naresuan Wildlife Sanctuary, Tak Province, Thailand.
part a. Sex, meristic, categorical, and raw morphometric data used in the analyses from specimens in the Cyrtodactylus brevipalmatus group. Abbreviations for morphometric characters are in the Materials and methods. / = data unavailable; m = male; f = female.
Species | brevipalmatus | brevipalmatus | brevipalmatus | brevipalmatus | brevipalmatus | elok | elok | elok | elok | interdigitalis | interdigitalis | interdigitalis | interdigitalis | cf. interdigitalis |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Institutional catalog number | LSUHC 1899 | LSUHC 15076 | LSUHC 11788 | THNHM 10670 | THNHM 14112 | LSUHC 8238 | LSUHC 12180 | LSUHC 12181 | ZMMU R-16144 | paratype THNHM 20226 | paratype THNHM 20228 | paratype THNHM 20229 | paratype THNHM 20227 | ZMMU R-16492 |
sex | m | f | f | f | f | f | m | m | f | f | f | f | f | m |
Meristic data | ||||||||||||||
supralabials (SL) | 11 | 12 | 10 | 14 | 12 | 11 | 8 | 13 | 9 | 14 | 12 | 11 | 12 | 8 |
infralabials (IL) | 8 | 10 | 9 | 11 | 11 | 11 | 8 | 11 | 9 | 9 | 8 | 8 | 7 | 9 |
paravertebral tubercles (PVT) | 39 | 37 | 38 | 29 | 23 | 0 | 0 | 0 | 0 | 27 | 27 | 24 | 23 | 30 |
longitudinal rows of tubercles (LRT) | 15 | 16 | 17 | 16 | 14 | 6 | 7 | 4 | 4 | 19 | 20 | 19 | 19 | 18 |
ventral scales (VS) | 38 | 38 | 38 | 36 | 39 | 45 | 45 | 47 | 36 | 42 | 40 | 42 | 43 | 34 |
ventral scales along middle of body (VSM) | 176 | 170 | 182 | 154 | 160 | 190 | 225 | 234 | 192 | 187 | 170 | 187 | 178 | 160 |
expanded subdigital lamellae on 4th toe (TL4E) | 7 | 8 | 9 | 8 | 8 | 10 | 9 | 9 | 9 | 12 | 10 | 10 | 11 | 9 |
unmodified subdigital lamellae on 4th toe (TL4U) | 13 | 11 | 11 | 11 | 12 | 11 | 10 | 11 | 9 | 14 | 13 | 12 | 14 | 10 |
total subdigital lamellae 4th toe (TL4T) | 20 | 19 | 20 | 19 | 20 | 21 | 19 | 20 | 18 | 26 | 23 | 22 | 23 | 19 |
expanded subdigital lamellae on 4th finger (FL4E) | 8 | 8 | 8 | 7 | 8 | 8 | 9 | 9 | 9 | 9 | 8 | 9 | 9 | 10 |
unmodified subdigital lamellae on 4th finger (FL4U) | 9 | 11 | 10 | 10 | 10 | 12 | 13 | 9 | 8 | 12 | 11 | 12 | 13 | 9 |
total subdigital lamellae 4th finger (FL4T) | 17 | 19 | 18 | 17 | 18 | 20 | 22 | 18 | 17 | 21 | 21 | 21 | 22 | 19 |
enlarged femoral scales (R/L) | 0 | 0 | 0 | 8R/ 8L | 7R/ 7L | 0 | 0 | 0 | 0 | 11R/8L | 10R/9L | 8R/ 8L | 9R/10L | 9R/8L |
total enlarged femoral scales (FS) | 16 | 10 | 11 | 16 | 14 | 0 | 0 | 0 | 0 | 14 | 19 | 19 | 19 | 17 |
total femoral pores (FP) | 7 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 17 |
enlarged precloacal scales (PCS) | 7 | 7 | 7 | 8 | 7 | 8 | 8 | 8 | 7 | 14 | 15 | 13 | 19 | 13 |
precloacal pores (PP) | 7 | 0 | 0 | 0 | 0 | 0 | 8 | 8 | 0 | 0 | 0 | 0 | 0 | 13 |
postcloacal tubercles (PCT) | 3 | 3 | 2 | 3 | 3 | 3 | 2 | 3 | 3 | 3 | 2 | 3 | 3 | 3 |
body bands (BB) | 4 | 6 | 3 | 5 | 5 | 5 | 5 | 3 | 3 | 5 | 5 | 5 | 5 | 3 |
Categorical data | ||||||||||||||
small tubercles on flank (FK-tub) | present | present | present | present | present | absent | absent | absent | absent | present | present | present | present | present |
dorsolteral caudal tubercles (DCT) | small | small | small | / | small | large | large | large | large | small | / | small | small | large |
ventrolateral caudal fringe (VLF1) | small | small | small | / | small | large | large | large | large | small | / | small | small | large |
ventrolateral caudal fringe scales generally homogenous (VFL2) | no | no | no | / | no | no | no | no | no | yes | yes | yes | yes | yes |
tail crossection (TLcross) | circular | circular | circular | / | circular | square | square | square | square | circular | / | circular | circular | square |
slightly enlarged medial subcaudals (SC1) | present | present | present | / | absent | absent | absent | absent | absent | absent | / | absent | absent | / |
single enlarged medial subcaudal (SC2) | absent | absent | absent | / | absent | absent | absent | absent | absent | absent | / | absent | absent | / |
single enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) | no | no | no | / | no | no | no | no | no | yes | / | yes | yes | / |
Morphomeric data | ||||||||||||||
SVL | 68.8 | 70.8 | 64.1 | 66.0 | 63.8 | 80.2 | 78.2 | 84.8 | 78.6 | 81.2 | 74.8 | 78.6 | 59.7 | 68.1 |
AG | 35.7 | 33.4 | 30.1 | 30.0 | 26.5 | 39.7 | 37.8 | 41.5 | 36.2 | 34.5 | 33.7 | 32.7 | 24.6 | 34.6 |
HumL | 9.7 | 9.3 | 8.0 | 9.6 | 9.5 | 10.2 | 9.1 | 10.1 | 1.7 | 9.8 | 10.2 | 11.2 | 7.4 | 10.3 |
ForL | 9.9 | 9.8 | 8.9 | 8.2 | 8.7 | 11.5 | 11.7 | 11.8 | 10.2 | 10.6 | 10.5 | 11.1 | 8.4 | 8.5 |
FemL | 12.0 | 12.6 | 11.5 | 11.7 | 9.8 | 12.9 | 14.2 | 14.6 | 13.1 | 14.7 | 13.2 | 12.7 | 10.2 | 12.6 |
TibL | 11.6 | 12.2 | 10.5 | 9.7 | 8.2 | 13.5 | 14.0 | 13.8 | 12.3 | 13.1 | 11.9 | 12.9 | 10.2 | 11.4 |
HL | 19.3 | 19.3 | 19.0 | 17.9 | 18.2 | 21.8 | 21.6 | 21.9 | 21.7 | 20.8 | 19.9 | 21.7 | 16.7 | 18.4 |
HW | 13.2 | 13.8 | 12.3 | 12.3 | 12.0 | 15.6 | 16.1 | 15.9 | 15.1 | 14.0 | 13.4 | 14.2 | 11.4 | 13.1 |
HD | 8.0 | 7.6 | 7.6 | 7.3 | 7.0 | 9.6 | 9.8 | 10.4 | 9.8 | 3.4 | 8.6 | 8.7 | 6.6 | 8.3 |
ED | 5.2 | 4.5 | 4.3 | 5.3 | 4.4 | 4.8 | 5.0 | 5.7 | 5.0 | 5.3 | 5.5 | 5.9 | 4.4 | 4.4 |
EE | 5.7 | 5.9 | 4.9 | 5.7 | 5.7 | 6.4 | 7.1 | 7.0 | 6.8 | 5.8 | 6.2 | 6.4 | 4.8 | 6.2 |
ES | 7.4 | 7.6 | 7.0 | 7.0 | 7.2 | 8.6 | 8.7 | 9.5 | 8.6 | 8.3 | 7.8 | 9.1 | 6.8 | 7.7 |
EN | 5.7 | 5.4 | 4.9 | 5.3 | 5.4 | 6.0 | 6.2 | 6.5 | 6.2 | 6.0 | 5.5 | 6.8 | 5.1 | 5.5 |
IO | 5.4 | 4.7 | 4.7 | 4.2 | 5.2 | 5.7 | 5.4 | 5.4 | 3.9 | 4.8 | 4.7 | 5.5 | 4.3 | 2.9 |
EL | 1.0 | 1.4 | 1.1 | 1.3 | 1.0 | 1.9 | 1.4 | 1.5 | 1.4 | 1.3 | 1.3 | 1.6 | 1.2 | 0.9 |
IN | 1.7 | 2.1 | 2.3 | 2.1 | 2.2 | 2.7 | 2.6 | 2.5 | 3.1 | 2.1 | 2.2 | 2.5 | 1.8 | 2.3 |
part b. Sex, meristic, categorical, and raw morphometric data used in the analyses from specimens in the Cyrtodactylus brevipalmatus group. Abbreviations for morphometric characters are in the Materials and methods. / = data unavailable; m = male; f = female.
Species | ngati | ngati | ngati | ngati | ngati | ngati | ngati | ngati | cf. ngati 1 | cf. ngati 2 | cf. ngati 2 |
---|---|---|---|---|---|---|---|---|---|---|---|
Institutional catalog number | holotype HNUE-R00111 | paratype IEBR 4829 | paratype VNUF R.2020.12 | paratype HNUE-R00112 | FMNH 255454 | FMNH 270493 | FMNH 270492 | FMNH 265806 | NCSM 79472 | ZMMU R-14917 | NCSM 80100 |
sex | m | f | f | f | f | m | m | m | f | f | f |
Meristic data | |||||||||||
supralabials (SL) | 10 | 10 | 10 | 10 | 13 | 13 | 13 | 10 | 14 | 9 | 12 |
infralabials (IL) | 9 | 9 | 9 | 9 | 10 | 9 | 11 | 8 | 11 | 10 | 12 |
paravertebral tubercles (PVT) | 39 | 40 | 38 | 40 | 28 | 27 | 26 | 27 | 28 | 32 | 29 |
longitudinal rows of tubercles (LRT) | 18 | 18 | 17 | 22 | 19 | 18 | 17 | 19 | 18 | 24 | 19 |
ventral scales (VS) | 38 | 36 | 35 | 32 | 37 | 36 | 36 | 33 | 33 | 36 | 35 |
ventral scales along middle of the body (VSM) | 168 | 164 | 178 | 158 | 159 | 166 | 156 | 158 | 164 | 166 | 165 |
expanded subdigital lamellae on 4th toe (TL4E) | 8 | 10 | 9 | 9 | 10 | 10 | 8 | 10 | 9 | 8 | 10 |
unmodified subdigital lamellae on 4th toe (TL4U) | 11 | 10 | 11 | 10 | 11 | 11 | 11 | 11 | 12 | 10 | 10 |
total subdigital lamellae 4th toe (TL4T) | 13 | 16 | 17 | 16 | 21 | 21 | 19 | 21 | 21 | 18 | 20 |
expanded subdigital lamellae on 4th finger (FL4E) | 16 | 16 | 17 | 16 | 8 | 8 | 8 | 8 | 9 | 7 | 9 |
unmodified subdigital lamellae on 4th finger (FL4U) | 9 | 9 | 9 | 9 | 10 | 10 | 10 | 10 | 8 | 9 | 10 |
total subdigital lamellae 4th finger (FL4T) | 15 | 15 | 18 | 15 | 18 | 18 | 18 | 18 | 17 | 16 | 19 |
enlarged femoral scales (R/L) | 10R/10L | 9R/8L | 10R/9L | 8R/9L | 9R/7L | 8R/9L | 9R/9L | 8R/8L | 9R/8L | 7R/8L | 7R/8L |
total enlarged femoral scales (FS) | 20 | 17 | 19 | 17 | 16 | 17 | 18 | 16 | 17 | 15 | 15 |
total femoral pores (FP) | 14 | 0 | 0 | 0 | 0 | 14 | 15 | 13 | 0 | 0 | 0 |
enlarged precloacal scales (PCS) | 13 | 13 | 13 | 13 | 15 | 13 | 13 | 13 | 12 | 13 | 13 |
precloacal pores (PP) | 0 | 0 | 0 | 0 | 13 | 13 | 13 | 13 | 0 | 0 | 0 |
postcloacal tubercles (PCT) | 3 | 2 | 1 | 2 | 0 | 0 | 0 | 0 | 2 | 3 | 4 |
body bands (BB) | 6 | 6 | 6 | 6 | 3 | 4 | 3 | 3 | 3 | 3 | 3 |
Categorical data | |||||||||||
small tubercles on flank (FK-tub) | present | present | present | present | present | present | present | present | present | present | present |
dorsolteral caudal tubercles (DCT) | small | small | small | small | small | small | small | small | small | small | small |
ventrolateral caudal fringe (VLF1) | small | small | small | small | small | small | small | small | small | small | small |
ventrolateral caudal fringe scales generally homogenous (VFL2) | no | no | no | no | yes | yes | yes | yes | yes | yes | yes |
tail crossection (TLcross) | circular | circular | circular | circular | circular | circular | circular | circular | circular | circular | circular |
slightly enlarged medial subcaudals (SC1) | present | present | present | present | / | present | present | present | present | present | present |
single enlarged medial subcaudal (SC2) | absent | absent | absent | absent | / | absent | absent | absent | absent | absent | absent |
single enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) | no | no | no | no | / | no | no | no | no | no | no |
Morphometric data | |||||||||||
SVL | 66.5 | 68.1 | 69.3 | 46.6 | 83.56 | 70.24 | 74.13 | 73.76 | 77.95 | 87.1 | 77.66 |
AG | 28.8 | 29.8 | 30.2 | 19.7 | 41.3 | 35.4 | 37.0 | 31.3 | 38.2 | 41.9 | 36.8 |
HumL | 7.9 | 8.1 | 8.5 | 5.6 | 8.6 | 8.7 | 8.6 | 6.9 | 8.7 | 11.5 | 9.2 |
ForL | 9.2 | 10.0 | 10.1 | 6.5 | 10.2 | 9.3 | 10.4 | 10.0 | 10.3 | 10.4 | 10.7 |
FemL | 11.5 | 11.5 | 11.5 | 7.6 | 13.7 | 12.7 | 13.0 | 13.1 | 13.1 | 15.2 | 14.2 |
TibL | 10.8 | 11.1 | 11.8 | 7.8 | 12.5 | 11.8 | 11.2 | 11.1 | 12.8 | 12.6 | 12.7 |
HL | 20.1 | 20.4 | 20.7 | 16.1 | 21.7 | 20.6 | 20.3 | 20.7 | 21.2 | 22.1 | 21.4 |
HW | 12.6 | 12.0 | 11.8 | 8.8 | 13.8 | 12.5 | 13.0 | 12.3 | 12.7 | 14.8 | 13.5 |
HD | 7.4 | 7.2 | 6.6 | 5.1 | 9.2 | 8.4 | 9.1 | 7.6 | 8.3 | 8.7 | 9.2 |
ED | 3.8 | 4.1 | 3.4 | 2.6 | 4.9 | 4.9 | 4.9 | 4.8 | 6.5 | 4.6 | 6.0 |
EE | 5.8 | 5.5 | 5.9 | 4.4 | 6.9 | 6.1 | 6.2 | 5.7 | 5.3 | 6.5 | 6.2 |
ES | 7.5 | 7.6 | 6.9 | 5.0 | 9.0 | 8.3 | 8.3 | 8.2 | 8.7 | 8.8 | 8.4 |
EN | 6.7 | 6.3 | 6.2 | 4.5 | 6.5 | 6.2 | 6.1 | 6.2 | 6.2 | 6.6 | 6.0 |
IO | 5.6 | 5.4 | 5.6 | 4.2 | 6.6 | 5.6 | 5.4 | 5.1 | 4.9 | 3.5 | 5.7 |
EL | 0.8 | 0.8 | 0.7 | 0.3 | 1.3 | 1.1 | 1.2 | 1.0 | 1.5 | 1.2 | 0.9 |
IN | 2.8 | 2.6 | 2.6 | 2.0 | 2.8 | 2.5 | 2.5 | 2.3 | 2.7 | 2.2 | 2.5 |
part c. Sex, meristic, categorical, and raw morphometric data used in the analyses from specimens in the Cyrtodactylus brevipalmatus group. Abbreviations for morphometric characters are in the Materials and methods. / = data unavailable; m = male; f = female.
Species | rukhadeva | rukhadeva | cf. rukhadeva | cf. rukhadeva | cf. rukhadeva | cf. rukhadeva | cf. rukhadeva | cf. rukhadeva | cf. rukhadeva | sp. 13 | sp. 13 | sp. 14 |
---|---|---|---|---|---|---|---|---|---|---|---|---|
Institutional catalog number | Holotype ZMMU R-16851 | Paratype ZMMU R-16852 | THNHM 24622 | THNHM 24838 | THNHM 03251 | THNHM 03252 | THNHM 03253 | THNHM 03254 | THNHM 01807 | THNHM 00104 | THNHM 27821 | THNHM 01667 |
sex | m | f | m | f | m | m | f | m | m | f | f | m |
Meristic Data | ||||||||||||
supralabials (SL) | 11 | 9 | 11 | 13 | 13 | 11 | 12 | 13 | 12 | 12 | 15 | 12 |
infralabials (IL) | 10 | 11 | 10 | 10 | 10 | 10 | 10 | 11 | 10 | 10 | 11 | 10 |
paravertebral tubercles (PVT) | 27 | 30 | 26 | 28 | 27 | 27 | 30 | 30 | 26 | 33 | 29 | 29 |
longitudinal rows of tubercles (LRT) | 19 | 20 | 18 | 19 | 18 | 18 | 19 | 19 | 19 | 18 | 20 | 19 |
ventral scales (VS) | 34 | 43 | 38 | 36 | 37 | 37 | 39 | 34 | 35 | 37 | 36 | 34 |
ventral scales along middle of the body (VSM) | 154 | 152 | 162 | 158 | 157 | 159 | 168 | 160 | 161 | 159 | 165 | 159 |
expanded subdigital lamellae on 4th toe (TL4E) | 9 | 9 | 8 | 9 | 9 | 10 | 9 | 10 | 10 | 9 | 7 | 8 |
unmodified subdigital lamellae on 4th toe (TL4U) | 11 | 11 | 11 | 13 | 12 | 12 | 15 | 13 | 13 | 12 | 12 | 13 |
total subdigital lamellae 4th toe (TL4T) | 20 | 18 | 19 | 22 | 21 | 22 | 14 | 23 | 23 | 21 | 19 | 21 |
expanded subdigital lamellae on 4th finger (FL4E) | 9 | 8 | 7 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 |
unmodified subdigital lamellae on 4th finger (FL4U) | 10 | 9 | 10 | 11 | 10 | 10 | 12 | 12 | 12 | 11 | 10 | 12 |
total subdigital lamellae 4th finger (FL4T) | 19 | 17 | 17 | 17 | 18 | 18 | 20 | 20 | 20 | 19 | 18 | 20 |
enlarged femoral scales (R/L) | 9R/8L | 8R/8L | 9R/L | 9R/ 9L | 9R/ 7L | 7R/ 7L | 6R/ 7L | 5R/ 8L | 7R/ 7L | 9R/ 9L | 7R/10L | 7R/ 7L |
total enlarged femoral scales (FS) | 17 | 16 | 18 | 18 | 16 | 14 | 13 | 13 | 14 | 18 | 17 | 14 |
total femoral pores (FP) | 17 | 0 | 14 | 0 | 12 | 13 | 0 | 11 | 13 | 0 | 0 | 7 |
enlarged precloacal scales (PCS) | 17 | 13 | 15 | 15 | 14 | 13 | 15 | 15 | 14 | 14 | 16 | 16 |
precloacal pores (PP) | 17 | 13 | 15 | 0 | 14 | 13 | 0 | 15 | 14 | 0 | 0 | 16 |
postcloacal tubercles (PCT) | 3 | 2 | 3 | 2 | 3 | 2 | 2 | 3 | 2 | 3 | 3 | 3 |
body bands (BB) | 3 | 3 | 3 | 3 | 4 | 4 | / | / | 5 | 3 | / | 5 |
Categorical data | ||||||||||||
small tubercles on flank (FK-tub) | present | present | present | present | present | present | present | present | present | present | present | present |
dorsolteral caudal tubercles (DCT) | small | small | small | small | small | small | small | small | / | small | small | large |
ventrolateral caudal fringe (VLF1) | small | small | small | small | small | small | small | small | / | small | small | large |
ventrolateral caudal fringe scales generally homogenous (VFL2) | yes | yes | yes | yes | yes | yes | yes | yes | / | yes | yes | no |
tail crossection (TLcross) | square | square | square | square | square | square | square | square | / | circular | circular | / |
slightly enlarged medial subcaudals (SC1) | absent | absent | absent | absent | absent | absent | absent | absent | / | present | present | absent |
single enlarged medial subcaudal (SC2) | present | present | present | present | present | present | present | present | / | absent | absent | absent |
single enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) | no | no | no | no | no | no | no | no | no | no | no | no |
Morphomeric data | ||||||||||||
SVL | 74.9 | 71.7 | 68.3 | 71.8 | 73.6 | 75.3 | 74.7 | 73.2 | 61.5 | 63.7 | 72.9 | 70.2 |
AG | 34.6 | 32.6 | 27.3 | 29.9 | 30.9 | 31.3 | 32.2 | 30.3 | 26.2 | 25.8 | 30.6 | 31.5 |
HumL | 10.7 | 10.4 | 9.8 | 8.3 | 12.2 | 11.3 | 11.8 | 11.0 | 10.1 | 7.6 | 10.1 | 10.2 |
ForL | 8.6 | 7.9 | 8.7 | 8.5 | 9.0 | 10.6 | 9.6 | 9.2 | 7.9 | 8.1 | 9.6 | 8.6 |
FemL | 12.6 | 11.8 | 10.8 | 10.9 | 11.5 | 10.2 | 11.9 | 12.1 | 9.5 | 10.7 | 12.8 | 12.1 |
TibL | 10.1 | 9.3 | 9.7 | 10.7 | 10.9 | 11.7 | 11.3 | 11.1 | 9.1 | 10.1 | 10.2 | 11.8 |
HL | 20.2 | 19.2 | 19.7 | 19.9 | 20.8 | 21.3 | 20.8 | 21.5 | 17.9 | 17.6 | 19.9 | 18.3 |
HW | 14.6 | 13.4 | 13.1 | 13.9 | 14.9 | 15.0 | 13.1 | 14.1 | 11.8 | 11.9 | 13.8 | 12.1 |
HD | 9.2 | 8.5 | 7.3 | 8.9 | 8.2 | 8.2 | 8.1 | 8.9 | 7.5 | 7.7 | 8.4 | 7.8 |
ED | 4.6 | 4.3 | 4.9 | 5.1 | 5.8 | 5.4 | 5.0 | 5.5 | 4.7 | 4.1 | 5.3 | 5.2 |
EE | 6.2 | 6.2 | 5.1 | 6.2 | 5.6 | 5.7 | 5.4 | 6.2 | 4.3 | 4.9 | 6.3 | 4.9 |
ES | 8.3 | 7.7 | 7.4 | 8.1 | 8.4 | 8.8 | 8.1 | 8.6 | 7.3 | 7.2 | 8.0 | 7.5 |
EN | 6.3 | 5.7 | 5.4 | 6.0 | 6.2 | 6.4 | 5.8 | 6.2 | 5.3 | 5.6 | 5.9 | 5.5 |
IO | 3.3 | 3.1 | 4.5 | 4.7 | 5.6 | 5.7 | 5.7 | 5.6 | 4.2 | 4.8 | 6.1 | 4.0 |
EL | 1.2 | 1.0 | 1.6 | 1.5 | 1.2 | 1.3 | 1.2 | 1.2 | 0.9 | 1.4 | 1.4 | 1.3 |
IN | 2.2 | 2.1 | 2.0 | 2.2 | 2.4 | 2.5 | 2.4 | 2.3 | 2.0 | 2.1 | 2.3 | 2.2 |
Ingroup samples consisted of 16 individuals of the brevipalmatus group representing five nominal species (Table
All statistical analyses were conducted using
Small sample sizes (n=1 or 2) for some of the species/ populations precluded standard statistical analyses for relevant groups closely related to C. interdigitalis. However, morphospatial clustering and positioning among the species/populations was analyzed using multiple factor analysis (MFA) on a concatenated data set comprised of 15 meristic characters, 16 normalized morphometric characters, and seven categorical characters (Table
In order to further examine the morphometric dissimilarity among C. ngati and closely related specimens from Laos and Thailand that have been referred to as C. interdigitalis (see below), a PCA and discriminant analysis of principal components (DAPC) of the 16 normalized morphometric characters was employed. PCA is a dimension reducing algorithm that decreases the complexity of a data set by finding a subset of input variables that contain the most relevant information (i.e. the greatest variance in the data) while de-emphasizing those characters that do not, thus increasing the overall accuracy of the results by eliminating noise and the potential of overfitting (
A two-sample Student t-test on data meeting the assumptions of normality and homogeneity of variance, and a Welch’s two-sample t-test on data violating those assumptions, were run for each morphometric character to test for the presence or absence of significantly different mean values (p<0.05). The ranges, means, medians, and 50% quartiles of each character were visualized using violin plots with embedded boxplots. All analyses were performed in R [v3.4.3].
Following the MFA and PCA, a non-parametric permutation multivariate analysis of variance (PERMANOVA) from the vegan package 2.5-3 in R (
Mismatch distributions using the PopGenome package version 2.7.5 in R (Pfeifer et al. (2020) based on the infinite-sites model (
A Mantel randomization test from the adegent package 2.1.5 in R (
To complement the Mantel test, a dbRDA analysis was performed on the two matrices using the capescale() function from the vegan package. Statistical significance using 999 permutations was assessed by calculating and R2, R2adj, and F test p-values. The data matrices were regressed in a linear regression analysis using the lm() function and plotted on a heat map using a 2-dimensional kernal density estimation (kde2d function) from the MASS package 7.3-54 (
The ML and BI analyses recovered trees with well-supported (UFB 100/BPP 1.00) identical topologies (Fig.
The GMYC species delimitation independently recovered FMNH 265806, 255454, 270492 and VNUF R.2014.50 and C. ngati (IBER 4829 and VNUF R.2020.12) as conspecific. However, the likelihood ratio test was insignificant (p=0.10000) which is not surprising given the high percentage of singletons in the data set (
A Maximum likelihood consensus tree based on 1399 base pairs of ND2 and flanking tRNAs with UFBS and BBP nodal support. B Histogram of uncorrected pairwise sequence divergence of specimens in the interdigitalis clade from Cyrtodactylus interdigitalis YC00952 from the type locality of Tham Yai Nam Nao, Nam Nao National Park, Phetchabun Province, Thailand.
The MFA of the concatenated data sets corroborate the phylogenetic analyses, in part, in that C. brevipalmatus (combined with C. cf. brevipalmatus from here on out), C. elok, C. interdigitalis, and C. rukhadeva (including the type series and specimens from Kaeng Krachan National Park; see below), and sp. 9. were recovered as distinct well-separated species along the combined axes of Dim-1 and Dim-2 (Fig.
A. MFA of all species/populations of the brevipalmatus group. B MFA of all species/populations of the brevipalmatus group minus Cyrtodactylus elok. C Histograms of the percent contribution of each data type to the variation in dimensions 1 (Dim-1) and dimensions 2 (Dim-2) based on 6B. The dashed red line represents the average of percent contribution.
Summary statistics of the PERMANOVA test. Shaded cells denote significant p values (< 0.05)
species pairs | F Model | R 2 | p value | p-adjusted |
rukhadeva vs ngati | 11.2567 | 0.5058 | 0.0014 | 0.0142 |
rukhadeva vs interdigitalis | 4.6290 | 0.2962 | 0.0016 | 0.0158 |
rukhadeva vs ngati types | 9.1022 | 0.4765 | 0.0046 | 0.0458 |
rukhadeva vs brevipalmatus | 7.9317 | 0.3979 | 0.0006 | 0.0056 |
ngati vs interdigitalis | 3.8323 | 0.3898 | 0.0286 | 0.2857 |
ngati vs ngati types | 15.6712 | 0.7581 | 0.0286 | 0.2857 |
ngati vs brevipalmatus | 18.0303 | 0.7203 | 0.0080 | 0.0804 |
interdigitalis vs ngati types | 7.7098 | 0.6066 | 0.0286 | 0.2857 |
interdigitalis vs brevipalmatus | 5.9252 | 0.4584 | 0.0082 | 0.0820 |
ngati types vs brevipalmatus | 5.9327 | 0.4972 | 0.0179 | 0.1786 |
The MFA recovered the holotype of C. rukhadeva from Khao Laem Mountain, Suang Phueng District, Ratchaburi Province, Thailand and the paratype from Hoop Phai Tong, Suang Phueng District 7.7 km to the east, as overlapping along Dim-2 and in close morphospatial proximity to each other and a series of seven specimens (THNHM 01807, 03251–54, 24622, 24838) of C. cf. rukhadeva (
The MFA recovered the specimen from the Khlong Nakha Wildlife Sanctuary, Ranong Province, Thailand (sp. 12, THNHM 01667) as widely separated from C. brevipalmatus (Fig.
The MFA recovered significant morphological distinction between the type series of C. ngati from Vietnam and the geographically distant Thai and Lao populations of C. ngati plus C. cf. ngati 1 and 2 (Figs
A PCA of specimens in the interdigitalis clade. B DAPC of specimens in the interdigitalis clade. C Violin plots overlain with box plots showing the range, frequency, mean (white dot), and 50% quartile (black rectangle) of the size-adjusted morphometric characters. * denotes characters with significantly different mean values.
PCA summary statistics for Cyrtodactylus ngati and C. cf. ngati 1 and 2. Shaded cells denote heavy loadings. Abbreviations are in the Materials and methods.
PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | PC7 | PC8 | PC9 | PC10 | |
Standard deviation | 3.02514 | 1.43855 | 1.31700 | 1.05543 | 0.94514 | 0.75287 | 0.64766 | 0.22607 | 0.00000 | 0.00000 |
Proportion of variance | 0.57197 | 0.12934 | 0.10841 | 0.06962 | 0.05583 | 0.03543 | 0.02622 | 0.00319 | 0.00000 | 0.00000 |
Cumulative proportion | 0.57197 | 0.70131 | 0.80971 | 0.87933 | 0.93516 | 0.97059 | 0.99681 | 1.00000 | 1.00000 | 1.00000 |
Eigenvalue | 9.15147 | 2.06943 | 1.73449 | 1.11394 | 0.89329 | 0.56681 | 0.41947 | 0.05111 | 0.00000 | 0.00000 |
SVL | –0.24017 | 0.02365 | –0.05714 | –0.04626 | 0.38558 | –0.03327 | 0.88620 | –0.03510 | 0.00000 | 0.00000 |
AG | –0.31222 | –0.03444 | –0.16700 | –0.13169 | –0.04196 | 0.24410 | –0.07073 | 0.08044 | –0.09704 | –0.44157 |
HumL | –0.18395 | 0.30455 | –0.04150 | –0.10087 | –0.70280 | 0.16579 | 0.23479 | –0.28513 | 0.10144 | –0.10135 |
ForL | –0.20015 | –0.17542 | 0.37514 | 0.50929 | 0.06400 | 0.22482 | –0.00663 | 0.29195 | 0.33516 | –0.05149 |
FemL | –0.31397 | 0.15040 | 0.00333 | 0.10602 | 0.09091 | –0.19909 | –0.12166 | 0.22019 | –0.43592 | –0.21044 |
TibL | –0.26011 | –0.21195 | 0.05811 | –0.33668 | –0.28102 | –0.36533 | 0.04617 | 0.41117 | –0.28757 | 0.16460 |
HL | –0.29184 | 0.00874 | –0.02861 | –0.13911 | 0.05482 | –0.57390 | –0.13660 | –0.06990 | 0.65217 | –0.23562 |
HW | –0.29884 | 0.18474 | –0.19528 | 0.18133 | –0.09028 | –0.00066 | –0.04357 | 0.15712 | 0.11422 | 0.76179 |
HD | –0.30883 | –0.11021 | –0.13985 | 0.18125 | –0.07375 | 0.20441 | –0.04838 | –0.19786 | –0.14403 | –0.03920 |
ED | –0.28495 | –0.23283 | 0.25136 | –0.15942 | 0.04516 | –0.02978 | –0.08982 | –0.14963 | –0.07552 | 0.08892 |
EE | –0.13329 | 0.05995 | –0.61810 | 0.38078 | –0.05532 | –0.05066 | –0.02928 | 0.15765 | 0.06903 | –0.15484 |
ES | –0.30780 | –0.06844 | –0.06403 | –0.13606 | 0.24823 | 0.07530 | –0.22076 | –0.57475 | –0.01069 | 0.20829 |
EN | 0.20879 | 0.23130 | –0.41436 | –0.33049 | 0.26567 | 0.06299 | –0.10032 | 0.16319 | 0.09653 | 0.05554 |
IO | 0.07527 | –0.57805 | –0.32974 | 0.20819 | 0.01806 | –0.17406 | 0.00256 | –0.21397 | –0.17053 | 0.00329 |
EL | –0.26849 | –0.09245 | –0.05184 | –0.34304 | 0.19539 | 0.49703 | –0.14735 | 0.26629 | 0.13461 | –0.01479 |
IN | 0.14076 | –0.54969 | –0.17904 | –0.20525 | –0.26817 | 0.16311 | 0.15877 | 0.13640 | 0.26272 | 0.06895 |
The mismatch analyses returned multimodal distributions for the interdigitalis clade and C. ngati, indicating they are not undergoing range expansion (Fig.
Based on eigenvalue decomposition, the loadings of the first two principal components from the PCoA for the geographic data were retained as spatial variables. The very low calculated Mantel r statistic and insignificant p-value (r=-0.1578; p=0.7438) between the genetic and transformed geographic distance (dis)similarity matrices of individuals in the interdigitalis clade fell midway within the range of the observed permutations, indicating there is no correlation between genetic and geographic distances (i.e. potentially no IBD; Fig.
A Observed simulations of the Mantel test of the individuals in the interdigitalis clade based on 10,000 permutations. B Regression analysis of the geographic and genetic distance matrices and heat map of clusters of the individuals in the interdigitalis clade. C Observed simulations of the Mantel test of the individuals of Cyrtodactylus ngati based on 10,000 permutations. D Regression analysis of the geographic and genetic distance matrices and heat map of clusters of individuals of C. ngati. Dashed red line is the regression line. Vertical line with diamond tip refers to the value of the r statistic.
The lack of correlations between the genetic and geographic (dis)similarity matrices recovered in the Mantel and dbRDA tests for both data sets are consistent with a phylogeographic structure that is not a function of IBD (e.g.
The significant morphological distinction between the type series of C. ngati from Vietnam and the geographically distant Thai and Lao populations of C. ngati plus C. cf. ngati 1 and 2 (Figs
t-test summary statistics for Cyrtodctylus ngati and C. cf. ngati 1 and 2. Shaded cells denote significant p values. Abbreviations are in the Materials and methods.
character | Welch’s t | Student t | p value |
SVL | 2.9327 | 0.0189 | |
AG | 10.47 | 3.91E-05 | |
HumL | 1.1501 | 0.2833 | |
ForL | 1.7757 | 0.1137 | |
FemL | 13.081 | 1.235E-05 | |
TibL | 2.7255 | 0.026 | |
HL | 5.1505 | 0.0009 | |
HW | 7.6325 | 0.0001 | |
HD | 7.3555 | 8.20E-05 | |
ED | 3.8713 | 0.0047 | |
EE | 1.3404 | 0.2169 | |
ES | 7.1308 | 0.0107 | |
EN | 1.4927 | 0.1727 | |
IO | -64882 | 0.5346 | |
EL | 4.5441 | 0.0019 | |
IN | -1.5426 | 0.1615 |
An alternative, but less preferred hypothesis, is that mitochondrial DNA introgression (through hybridization) or incomplete lineage sorting (very recent speciation precluding the accumulation of mutations) is obscuring species boundaries in this group (e.g.,
These analyses demonstrate the necessity of examining type material (when possible) in the context of a phylogeny in order to correctly identify similarly appearing specimens (see Fig.
A Cyrtodactylus interdigitalis from Phetchabun Province, Thailand. Photo by Montri Sumonta. B Cyrtodactylus cf. interdigitalis (ZMMU R-16492) from Phu Hin Rong Kla National Park, Phitsanulok Province, Thailand. Photo by Nikolay A. Poyarkov. C Cyrtodactylus cf. ngati 2 (NCSM 80100) from Houay Wan Stream, tributary of Nam Pha River, Vientiane Province, Laos. Photo by Bryan L. Stuart. D Cyrtodactylus ngati (VNUF R.2020.12) from Pa Thom Cave, Pa Xa Lao Village, Pa Thom Commune, Dien Bien District, Dien Bien Province, Vietnam. Photo by Dzung T. Le. E Cyrtodactylus cf. ngati 1 (NCSM 79472) Houay Liep Stream, Paklay: Ban Pha Liep, Xaignabouli Province, Laos. Photo by Bryan L. Stuart. F Adult male C. cf. rukhadeva from Kaeng Krachan National Park, Phetchaburi Province, Thailand. Photo from Creative Commons Attribution Share Alike.
This work (Grant No. RGNS 64-038) was financially supported by Office of the Permanent Secretary, Ministry of Higher Education, Science, Research and Innovation. AR and AA were supported by Kasetsart University Research and Development Institute (KURDI). YC was financial supported by the Research Administration Division (RAD), Khon Kaen University. Wachara Sanguansombat and Sunchai Makchai (Thailand Natural History Museum) made specimens in their care available for study. Sengvilay Seateun, Khampong Thanonkeo, Tanya Chan-ard and David Emmett assisted with collecting NCSM and FMNH specimens in Laos.