Research Article |
Corresponding author: Shu-Ping Wu ( shupingwu@ntu.edu.tw ) Academic editor: Franco Andreone
© 2016 Shu-Ping Wu, Chuan-Chin Huang, Chi-Li Tsai, Te-En Lin, Jhih-Jia Jhang, Sheng-Hai Wu.
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:
Wu S-P, Huang C-C, Tsai C-L, Lin T-E, Jhang J-J, Wu S-H (2016) Systematic revision of the Taiwanese genus Kurixalus members with a description of two new endemic species (Anura, Rhacophoridae). ZooKeys 557: 121-153. https://doi.org/10.3897/zookeys.557.6131
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Two new species of rhacophorid tree frog were identified in Taiwan. In both new taxa, derived reproductive characteristics of laying eggs in tree holes and oophagous tadpoles are shared with Kurixalus eiffingeri, but they are divergent from each other in molecular genetics, mating calls, and tadpole and adult morphology. The morphological characteristics and the molecular phylogenetic evidence support the hypothesis that the two new species, Kurixalus berylliniris sp. n. and Kurixalus wangi sp. n., are both monophyletic lineages.
Kurixalus berylliniris sp. n., Kurixalus wangi sp. n., oophagous tadpoles
There are four genera (Buergeria, Tschudi, 1838, Kurixalus, Ye, Fei, and Dubois In Fei, 1999, Polypedates, Tschudi, 1838, and Rhacophorus, Kuhl and Van Hasselt, 1822) and eleven species of rhacophorid tree frogs on the island of Taiwan (
Kurixalus eiffingeri, a native species in the island of Taiwan, is the only rhacophorid within the genus Kurixalus that has a tree-hole breeding reproductive mode and oophagous tadpoles (
In our study, Kurixalus treefrog specimens were collected from the island of Taiwan. Additionally, the specimens of Kurixalus eiffingeri were collected from the type localities, Iriomote and Ishigaki isles. In the field, we noticed that some of the Kurixalus populations in eastern and southern Taiwan resembled K. eiffingeri in external morphology but differed in their reproductive season (November to February) from K. eiffingeri (from February to August). One group has an extraordinarily small body size, and the other group has green irises. Further examination of the samples from the two populations and K. eiffingeri revealed the differences in external morphology, tadpole morphology, comparative anatomy, mating call analysis, and molecular genetic evidence. From these results, the two populations of rhacophorid frogs are describe as new species.
The type specimens of frogs and tadpoles of the two new Kurixalus species were collected by hand, euthanized using a dilute chloretone solution, and fixed in 10% buffered formalin. Frogs were later transferred to 70% ethanol, and tadpoles were stored in 10% buffered formalin. In addition to the type specimens described in this study, 343 samples that consisted of Kurixalus eiffingeri and related taxa were collected from 22 locations throughout the island of Taiwan. Furthermore, three specimen of Kurixalus eiffingeri were collected from the type locality. One was from Iriomote isle and the other two were from Ishigaki isle (Fig.
Sampling localities of this study. Localities 1 through 22 are around Taiwan island, locality 23 from Iriomote isle, locality 24 from Ishigaki isle. The two isles belong to the southern end of Ryukyu archipelago. Color refers to the geographical distribution of the three Kurixalus species. Red: K. eiffingeri; Green: K. berylliniris sp. n. (Taxon 1); B: K. wangi sp. n. (Taxon 2). Loc. 20: Ligia, type locality of K. berylliniris sp. n.; Loc. 21: Shouka, type locality of K. wangi sp. n.
Morphometric characteristics of adult specimens: snout-vent length (SVL), head width (HW), head length (HL), internarial distance (IN), eye-narial distance (EN), horizontal eye diameter (ED), distance between the anterior margins of eyes (DFE), distance between the posterior margins of eyes (DBE), upper eyelid width (UEW), interorbital distance (IO), tympanic annulus diameter (TAD), distance between the axillae, between posterior margins of the upper arm (AXI), axilla-groin distance (AGD), forearm length (UAW), manus length (PAL), length of first finger from base of palmar tubercle to tip of third finger disc (F1L), width of third finger disc (D3L), femur length (FEL), tibia length (TBL), tarsus length (TSL), foot length from proximal margin of inner metatarsal tubercle to tip of fourth toe (FOL), first toe length (TL), inner metatarsal tubercle length (IML), and disc width of fourth toe (T4D) (Table
In addition, the eggs and tadpole morphometric characteristics were measured comprising total length (TL), body length (BL), tail length (CL), tail height (TH), tail muscle height (TM), internarial distance (NA), distance between eyes (IN), and tail muscle width (MW) (
All measurements of morphometric characteristics were taken using a dial caliper under a dissecting microscope, and measurements were rounded to 0.1 mm. Digital webbing of the adults was recorded using Savage and Heyer’s formula (1997).
T-tests were used to examine whether body size varied by gender within each taxon. An analysis of covariance (ANCOVA) method was used to compare the size-adjusted means of morphometric characteristics. Morphometric characteristics that satisfied the normality assumption were included in a multivariate principal component analysis (PCA) based on the correlation matrix of size-standardized measurements (all measurements divided by SVL). Scatter plots of the scores of the first three factors of PCA were used to examine the differentiation among specimens. All of these tests and analyses were applied separately to male and female specimens. The statistical analyses were performed using SigmaPlot 12 (Systat Software, Inc.).
Frog mating calls were recorded using a digital recorder (Fostex FR-2LE) and a microphone (Sennheiser ME 67/k6). Calls were recorded in the native habitats of these tree frogs, and environmental parameters including temperature and humidity were also recorded. Avisoft SASLab Pro 5.2.08 (Avisoft Bioacoustics) was used to extract the maximum and minimum frequencies, as well as the width of frequency, the single note duration, and the time interval between notes of the mating calls. A rapid call and a slow call were identified. Slow mating calls were compared among the subtypes in a pair-wise manner using Wilcoxon-Mann-Whitney odds (WMWodds) calculations (
Whole genomic DNA was extracted from muscle tissue of fresh or ethanol-preserved specimens using the procedure originally described by
A general time reversible model with a proportion of invariable sites and a gamma shaped distribution of rates across sites (GTR + I + G, I = 0.4402, G = 0.4519) was determined as the best-fitting model for the aligned sequences of the combined dataset using a hierarchical likelihood ratio test performed with the program MrModeltest 2.2 (
The Bayesian tree and the posterior probability distribution were determined using the program MrBayes 3.1 (
Partial sequences of mtDNA CO1 gene were used as haplotypes to examine the genetic structures of the three subtypes. We calculated the Fst, Nm (number of immigrants per generation,
Holotype. ASIZAM 0053, an adult male (Figs
Measurements (in mm) of type series and other specimens of Kurixalus berylliniris sp. n. and K. wangi sp. n. Abbreviations as in Materials and methods.
K. berylliniris sp. n. | K. wangi sp. n. | |||||||||
---|---|---|---|---|---|---|---|---|---|---|
male | male | female | male | male | female | |||||
no. | ASIZAM 53 | Mean±SD |
range | Mean±SD | range | ASIZAM 55 | Mean±SD |
range | Mean±SD | range |
holotype | (n=13) | (n=7) | holotype | (n=17) | (n=8) | |||||
SVL | 40.0 | 34.4±4.1 | 29.0–42.3 | 37.8±7.1 | 27.6–46.3 | 29.3 | 30.0±0.9 | 28.6–31.6 | 34.3±1.8 | 30.8–37.1 |
HW | 13.1 | 11.7±1.3 | 7.4–13.7 | 12.9±1.8 | 10.4–14.6 | 11.3 | 11.2±0.6 | 10.1–12.2 | 12.5±0.6 | 11.4–13.2 |
HL | 9.9 | 9.0±1.1 | 7.8–12.0 | 9.6±0.9 | 8.3–10.4 | 8.6 | 7.9±0.5 | 6.7–8.6 | 9.3±1.0 | 8.3–11.2 |
IN | 4.5 | 3.6±0.5 | 3.0–4.5 | 4.0±0.7 | 3.1–4.7 | 3.5 | 3.3±0.2 | 2.9–3.7 | 3.7±0.3 | 3.2–4.2 |
EN | 4.5 | 3.5±0.5 | 2.9–4.5 | 3.7±0.6 | 3.0–4.4 | 3.7 | 3.2±0.3 | 2.6–3.7 | 3.4±0.2 | 3.2–3.8 |
ED | 5.2 | 4.3±0.5 | 3.5–5.2 | 4.5±0.8 | 3.6–5.9 | 3.5 | 4.1±0.3 | 3.5–4.8 | 4.5±0.4 | 4.0–5.0 |
UEW | 2.9 | 3.1±0.4 | 2.8–4.1 | 3.5±0.6 | 2.7–4.0 | 3.6 | 3.0±0.4 | 2.2–3.6 | 3.3±0.1 | 3.0–3.4 |
DFE | 7.9 | 6.7±0.6 | 6.0–7.9 | 7.4±1.1 | 5.7–8.4 | 6.7 | 6.4±0.5 | 5.5–7.2 | 6.9±0.5 | 6.2–7.7 |
DBE | 11.6 | 10.4±1.0 | 8.7–12.8 | 11.5±1.3 | 9.2–13.0 | 11.0 | 9.9±0.5 | 9.1–11.0 | 11.1±0.8 | 9.6–12.1 |
IO | 5.2 | 4.1±0.6 | 3.4–5.2 | 4.4±0.7 | 3.6–5.4 | 3.6 | 4.0±0.2 | 3.6–4.3 | 4.2±0.3 | 3.8–4.8 |
TAD | 2.5 | 2.1±0.2 | 1.7–2.5 | 2.4±0.5 | 1.6–2.9 | 1.9 | 1.9±0.2 | 1.5–2.1 | 2.0±0.1 | 1.9–2.2 |
AXI | 13.8 | 10.8±1.4 | 8.9–13.8 | 12.2±2.6 | 8.4–15.4 | 10.3 | 9.9±0.9 | 7.4–11.0 | 11.0±1.1 | 8.9–12.4 |
AGD | 19.4 | 17.4±1.6 | 15.6–20.6 | 18.2±4.5 | 11.2–24.3 | 14.1 | 13.3±1.6 | 10.4–16.4 | 15.0±2.0 | 12.6–17.8 |
UAW | 7.5 | 6.8±0.9 | 5.6–8.2 | 7.1±1.2 | 5.3–8.4 | 4.5 | 5.6±0.5 | 4.5–6.6 | 6.2±0.4 | 5.7–7.0 |
PAL | 13.9 | 10.7±1.5 | 9.2–13.9 | 12.3±2.2 | 8.9–14.9 | 9.2 | 9.0±0.6 | 8.1–10.3 | 9.5±0.5 | 8.8–10.3 |
F1L | 6.8 | 5.5±0.8 | 4.6–6.8 | 5.9±1.1 | 4.2–7.1 | 4.6 | 4.6±0.4 | 3.7–5.2 | 4.8±0.4 | 4.5–5.7 |
D3L | 2.4 | 1.6±0.4 | 1.1–2.4 | 1.7±0.4 | 1.2–2.6 | 1.8 | 1.5±0.2 | 1.1–1.9 | 1.7±0.1 | 1.5–1.8 |
FEL | 18.8 | 16.1±2.1 | 13.0–18.8 | 17.5±2.7 | 13.8–20.3 | 14.6 | 14.6±0.8 | 13.0–16.0 | 16.3±0.4 | 15.4–16.8 |
TBL | 19.9 | 17.0±2.1 | 14.6–19.9 | 18.1±2.7 | 14.2–21.8 | 14.4 | 14.6±0.8 | 13.3–16.2 | 16.4±0.7 | 15.1–17.1 |
TSL | 9.6 | 8.1±0.8 | 6.9–9.6 | 8.4±1.2 | 6.2–9.4 | 5.9 | 6.9±0.6 | 5.9–7.9 | 7.7±0.5 | 7.2–8.8 |
FOL | 17.9 | 15.4±2.0 | 13.0–18.5 | 17.0±3.0 | 12.6–19.9 | 12.9 | 12.4±0.7 | 11.4–13.9 | 13.8±0.7 | 12.6–14.9 |
TL | 6.7 | 5.7±1.0 | 4.5–7.2 | 6.3±1.5 | 4.3–8.0 | 4.2 | 4.3±0.2 | 3.8–4.6 | 4.8±0.4 | 4.3–5.3 |
T4D | 1.7 | 1.2±0.3 | 0.8–1.7 | 1.3±0.4 | 0.8–2.0 | 1.1 | 1.3±0.2 | 1.0–1.6 | 1.4±0.3 | 1.0–1.7 |
IML | 1.9 | 1.4±0.3 | 1.0–1.9 | 1.6±0.4 | 1.0–2.0 | 1.3 | 1.2±0.2 | 0.9–1.8 | 1.4±0.1 | 1.2–1.5 |
Paratypes. NCHUZOOL 11311-13 collected on 2 August 2005 by Hui-Ming Huang at the type locality; NCHUZOOL 11431, ASIZAM 0054 collected on 15 September 2005 by Shu-Ping Wu at the type locality; NCHUZOOL 11442 (eggs and tadpoles), collected on 7 February 2006 by Shu-Ping Wu at the type locality; NCHUZOOL 11448, collected on 16 February 2006 by Shu-Ping Wu at 425 meters above sea level, at Antong, Hualien County (Fig.
Ligia timber trail, 1250 meters above sea level, Taitung County, Taiwan, Republic of China (Fig.
A moderate-sized Kurixalus. Females average about 41 mm snout-vent length (range: 27.6–46.3 mm); males average about 35 mm (range: 29.0–42.3 mm). Iris emerald to light green. Two dark brown spots on eyelids, separated from each other and from X-shaped blotch on dorsum. Subarticular tubercles on foot rounded and flat. Belly and throat white or faintly-speckled. Prepollex in males squarish, compressed and expanded. About half-webbed between two outer toes. Anterior margin of tadpole dorsal fin extending to body. Tadpole heavily dark brown to black pigmented in gular region and on tail muscle. Upper lip of tadpole with deep transverse furrow, and prominent ridge extending from upper lip to anterior margin of nostril (key of tadpole, 3).
The epithet berylliniris is a compound word formed from beryllin (L.), green-colored, and from iris (L.), iris of the eye, and is treated as a noun in nominative singular in opposition to the generic name.
Habitus moderately slender and somewhat flattened, size moderate (SVL 40.1 mm); head wider than long; tip of snout pointed; snout obtuse in lateral view; nostril barely visible from above; canthus rostralis curved, prominent; loreal region concave, oblique; interorbital distance 1.5 times wider than upper eyelid width; nostril oval, oblique, closer to tip of snout than to eye; internarial distance slightly longer than nostril-eye distance; eye diameter larger than nostril-eye distance; pupil horizontal; tympanic region oblique; diameter of tympanum approximately half of eye diameter; tympanum distinct, round; tympanum to eye distance smaller than half tympanum diameter; supratympanic fold from posterior tip of eye to base of arm; jaw angle almost to posterior rim of tympanum; premaxillary and maxillary teeth present; choana exposed; vomerine teeth present only on left side; tooth patch oval, about half of choana diameter. Vocal slits near commissure of jaw, slit-like.
Limbs slender; tips of all four fingers expanded into discs with ventro-marginal and transverse grooves; disc of finger III about 67% of tympanum diameter; relative finger lengths: I<II<IV<III; relative disc widths I<II<III<IV; disc on finger I small, slightly wider than phalanx width. Webbing more extensive on right hand; only trace of webbing on left hand between fingers III and IV; webbing formula on right hand: I(1.5)–(1.5)II(2)–(2)III(1)–(1.5)IV; subarticular tubercles rounded, elevated, larger under phalanges than at base of fingers; supranumerary tubercles present, smaller than subarticular tubercles; two palmar tubercles, outer longer but narrower than inner. Nuptial pad greatly expanded, proximal edge more flattened than at base; epidermal glands discontinuous, on lateral margin of nuptial pad, and on internal margin of finger I; outer margin of hand with series of longitudinal tubercles somewhat connected to weak skin folds.
Heels overlapping when adpressed; tips of toes expanded into discs with ventro-marginal and transverse grooves; relative length of toes: I<II<V<III<IV; relative width of toe discs: I<II<III<IV<V; disc on toe I small, truncated; disc widest on toe V, less than twice of width of phalanx; webbing formula: I(0.5)–(1)II(0.5)–(1.5)III(1)–(2)IV(1)–(0.5)V; subarticular tubercles rounded, elevated, those at base of toes III, IV, and V smaller than supernumerary tubercles; inner metatarsal tubercle flat, oval, median margin free; outer metatarsal tubercle absent; a series of tubercles on outer surface of tarsus to outer margin of toe V.
Dorsum granular with small tubercles; palpebral tubercles absent; flank and venter smooth or slightly shagreened.
Color. In preservative, two dark brown spots on eyelids; dorsum at shoulder region with a large irregular X-shaped blotch; anterior horn of blotch not continuous with spots on eyelids; two brown blotches on lower back in groin region; flank white with large irregular blotches; dark blotches at cloacal opening, surrounded ventrally by white tubercles; loreal region with dark brown irregular spot; dark spots also present under eye, on posterior part of upper lip near jaw joint, and on supratympanic fold; arm with one thick cross bar on upper arm, two on forearm, one on outer palm; three transverse bars on thigh and on tibia; medial palm and foot white on dorsal surface; venter white; few irregular brown spots on chest, faintly maculated on gular region (Figs
Color in life. iris emerald to light green; dorsum dark green to deep tan with a black X-shaped and irregular blotches; tympanum light yellowish-brown with small dark spots; medial surface of hand and foot creamy white; venter cream sprinkled with minute black spots in gular region (Fig.
Variation. Sexual dimorphism was evident in the possession of nuptial pads and the hypertrophied upper and lower arms in males. Females were 10% larger than males (t-test, p > 0.05). Females possess a supra-cloacal flap (absent in males). The species has dark and white morphs. The dark morph is similar to the holotype (Fig.
Average diameter of the eggs was 4.55 (± 0.25) mm (n = 5) with capsule and 1.79 (± 0.09) mm (n = 8) without capsule. The eggs were creamy yellow with developing embryos. The range of total length of five preserved tadpoles between stages 26–33 was 17.64–30.00 mm (Fig.
Dorsal surface of tadpoles dark brown; ventral surface white; tail fins almost transparent with many faint black flecks; region of tail muscle heavily pigmented, especially anteriorly; body ovoid in lateral view, compressed above, more rounded below; eyes dorsal, not visible from below; eyes on anterior 1/3 of body; nostril lateral, about half way between upper lip and eye; internarial distance 105% of interorbital distance; eye-nostril distance smaller than interorbital distance; a very prominent and elevated ridge extending from nostril to upper lip; a deep transverse groove present in posterior to upper lip; a longitudinal groove on either side of head from lateral rim of upper lip to level between nostril and eye (Fig.
Vent dextral, opening at proximal edge of ventral fin; tail moderately strong, deeper than body; dorsal and ventral fin depth equal, almost symmetrical (or slightly deeper on dorsal fin); origin of dorsal fin anterior to that of ventral fin, on posterior 1/5 of body (Figs
Kurixalus berylliniris sp. n. occurs in eastern Taiwan (at 225 to 1250 meters above sea level). The highest recorded elevation was on the eastern slope of the Central Mountain Range (Taitung County, 1250 meters above sea level), and the lowest recorded elevation was on the western slope of the Coastal Range (Hwalien County, 225 meters above sea level). Specimens were collected near the canopy level in moist broad-leaf forests in Taitung and on forest edges in Hwalien. The northern border of the specimen’s distribution was near the Guangfu township of the central Hualien County (Fig.
Mating calls were heard during the winter months from November through February. Both a slow call and a rapid call consisted of a single beeping sound. Slow calls recorded in the field had an average duration of 158 (± 56) ms (n = 30, equivalent thereinafter); rapid calls had an average duration of 103 (± 42) ms. Intervals between notes were 3195 (± 1060) ms (slow calls) and 1562 (± 1442) ms (rapid calls). For the slow and rapid calls, the maximum frequencies of calls were 2901 (± 89) Hz (slow calls) and 2961 (± 71) Hz (rapid calls); the minimum frequencies of calls were 2517 (± 106) Hz (slow calls) and 2518 (± 124) Hz (rapid calls). (Fig.
Species | MAX (Hz) | MIN (Hz) | WID (Hz) | DUR (msec) | INT (msec) | DF (Hz) |
---|---|---|---|---|---|---|
K. berylliniris sp. n. (slow) | 2901 (89) | 2517(106) | 384 (80) | 158 (56) | 3195(1060) | 2704(35) |
K. berylliniris sp. n. (rapid) | 2961 (71) | 2518 (124) | 443 (97) | 103 (42) | 1562(1442) | 2772(360) |
K. wangi sp. n. (slow) | 3185(194) | 2399 (122) | 786 (192) | 99 (19) | 1122 (230) | 2841(145) |
K. wangi sp. n. (rapid) | 3072 (47) | 2565 (62) | 507 (62) | 57 (15) | 115 (22) | 2848(59) |
K. eiffingeri | 3034 (59) | 2550 (54) | 484 (90) | 154 (27) | 2063 (121) | 2772(260) |
K. idiootocus | 2889 (46) | 2412 (64) | 477 (80) | 48 (16) | 1900 (40) | 2647(62) |
Eggs and tadpoles were found in the pooled water in decaying trunks of tree ferns, Cyathea spinulosa. The eggs were adhered together in a single layer by colloidal gel and attached to the inner roof and wall above the water. A total of 62 eggs were counted in one tree hole (Fig.
Holotype. ASIZAM 0055 (Figs
Paratypes. NCHUZOOL 11161–62, collected on 13 September 2005 by Sheng-Hai Wu at Shuan-Liu, Pingtung County (22°13'15.58"N, 120°49'21.92"E); NCHUZOOL 11314, 11318, 11321-32, collected on 20 October 2005 by Shu-Ping Wu, on Shouka timber trail, Pingtung County, NCHUZOOL 11315, collected on 8 December 2005 by Shu-Ping Wu at Nanjenshan, Pingtung County (22°05'08.32"N, 120°51'24.04"E); NCHUZOOL 11316-17, 11319, collected on 20 December 2005 on Shouka timber trail, Pingtung County; NCHUZOOL 11334-35, collected on 7 December 2005 by Shu-Ping Wu, on Shouka timber trail, Pingtung County; NCHUZOOL 11441 (tadpoles and eggs), ASIZAM 0056 and NCHUZOOL 11445-47, collected on 9–12 February 2006 by Shu-Ping Wu, on Shouka timber trail, Pingtung County.
Shouka timber trail, 400 meters above sea level, Pingtung County, Taiwan, Republic of China (Fig.
A small to moderate-sized Kurixalus. Females snout-vent length averaging about 34 mm (range: 30.8–37.1 mm); males averaging 30 mm (range: 28.6–31.6 mm). Iris golden-yellow. Two anterior horns of the X-shaped marking on back extending to eyelid. Webbing extensive on toes, extending to the toe disc on the inner margin of toe V. Belly and throat whitish. Anterior margin of tadpole dorsal fin extending to posterior body. Tadpole with almost no pigment on region of tail muscle. Upper lip of tadpole with shallow transverse furrow.
The epithet is named and dedicated to Mr. Ching-Shong Wang for his pioneering work and contributions to the herpetology of Taiwan (
Habitus slender, body flat, small (SVL 29.3 mm), head wider than long, snout shape in dorsal view subovoid with pointed tip; profile acuminate, slightly protruding; canthus rostralis distinct, rounded; loreal region oblique, slightly concave; nostril oval and oblique; nostril closer to tip of snout than to eye; internarial distance equals nostril to eye distance; nostril to eye distance smaller than eye diameter; interorbital distance subequal to internarial distance and eyelid width; pupil horizontally oval; tympanum distinct, round, upper margin covered by curved supratympanic fold, which runs from posterior angle of eye to arm; angle of jaw at level of middle of tympanic ring; tympanum less than half of eye diameter; tympanum to eye distance greater than half tympanum diameter; premaxillary and maxillary teeth present; choana exposed; vomerine odontoid in oval patch, equal in diameter to choana; vomerine teeth present; tongue large, forked and shallowly emarginate; no lingual papilla; vocal slits long, near commissure of jaw on floor of mouth.
Limbs moderately robust; forearm shorter than hand; tips of fingers expanded into discs with ventro-marginal and transverse grooves; disc of finger III about 2/3 of tympanum diameter; finger length I<II<IV<III; disc even, truncate, with indistinct transverse groove; size of disc I<II<III<IV; disc of finger I small, same width as phalanx width; phalanges emarginate with trace of webbing; subarticular tubercles prominent, rounded, globular; prepollex expanded, rounded; glandular skin associated with nuptial pad from base of disc I on medial and dorsal side of pollex; palmar tubercle double, oval, subequal in size; supernumerary tubercles small; outer margin of fourth finger with longitudinal flat tubercles connected into a weak flap.
Heels overlapping when adpressed; hind limb moderate in length; shank shorter than thigh and longer than foot; tips of toes expanded into discs with ventro-marginal and transverse grooves; relative length of toes I<II<III<V<IV; relative size of discs I<II<III<IV<V; disc on toe I same width as phalanx width; discs truncate and with indistinct transverse grooves. Webbing moderate on all toes; webbing formula I(1)–(trace)II(0.5)–(1.5)III(1)–(2)IV(1)–(0.2)V; weak dermal fringe on outer side of toe V, from posterior tarsus to base of disc V, formed by continuous elongated papilla; subarticular tubercles rounded, slightly conical; subarticular tubercle on proximal joint on toe IV smaller than the others; inner metatarsal tubercle oval, small; outer metatarsal tubercle absent; supernumerary tubercle absent; small white-tipped tubercle on heel.
Skin shagreen, tubercles not present on back; ventral surface slightly granular, white tipped dermal tubercles on posterior thigh. Series of tubercles near lateral margin of upper eyelids; skin smooth on flank; white tipped tubercles on lateral lower arm in ventral view.
Color. In preservative, dorsum grayish with black irregular spots; patches of dark brown markings on median eyelid, forming triangular X-shaped blotch; two posterior branches of the X marking terminated at middle of dorsum; two dark blotches on posterior back; flank with dark oblique irregular band demarcating grayish dorsum and whitish venter; dark irregular blotches on loreal region, antero-ventral corner of eye, and tympanum; black band from anterior eye angle, through nostril to tip of snout; gular region sprinkled with black spots; upper arm with three wide bands, thigh and shank with three bands; ventral surface orange, speckled with brown spots on gular region; vent with large dark brown blotch over cloacal opening, surrounded ventrally and dorsally by white tubercles (Figs
Color in life, iris golden-yellow; dorsum dark brownish-green scattered with deep brown and black spots, with dark X marking on anterior half of dorsal surface; tympanic membrane light brown to milk-white; white and rounded tubercle located on outer fringe of heel (Fig.
Variation. Females were 14% larger than males (Table
Average diameter of eggs from 4 clutches was 3.37 (± 0.27) mm with capsule (n = 38) and 1.74 (± 0.09) mm without capsules; eggs were creamy yellow with developing embryos. The range of total length of ten tadpoles between stages 27–32 was 13.19–22.64 mm (Fig.
Dorsal surface of tadpoles dark brown; ventral surface white; pigment on tail confined mostly to upper half of tail muscle; tail fins transparent; body ovoid in lateral view, flat and sloping above, rounded below; eye dorsal, not visible from below, located on anterior 1/3 of body; nostril lateral; distance from nostril to upper lip much shorter than to eye; internarial distance subequal to interorbital distance; eye-nostril distance less than internarial distance. Face with slightly elevated ridge, from rostrum to upper lip (Fig.
Kurixalus wangi sp. n. is distributed in the southern part of Pingtung County in southern Taiwan below 500 meters above sea level (Fig.
Mating calls were heard in bushes or on tree branches up to 3 m above the ground between September and March, peaking in December. A slow call and a rapid call were identified. Both types of call consisted of a single beeping sound. Slow calls recorded in the field had an average duration of 99 (± 19) ms (n = 30, equivalent hereafter) and rapid calls had an average duration of 57 (± 15) ms. Intervals between notes were 1122 (± 230) ms (slow calls) and 115 (± 22) ms (rapid calls). For the slow and rapid calls, the maximum frequencies of calls were 3185 (± 194) Hz (slow calls) and 3072 (± 47) Hz (rapid calls); the minimum frequencies of calls were 2399 (±122) Hz (slow calls) and 2565 (± 62) Hz (rapid calls). (Fig.
Eggs were discovered in tree hollows, plastic pipes embedded in retaining walls on slopes (Fig.
Within-species comparisons showed that the body size was differentiated by sex in Kurixalus eiffingeri and K. wangi sp. n., but not in K. berylliniris sp. n. (Table
In the PCA, after eliminating the effect of size by using a normalizing ratio (measurements divided by SVL) and omitting the five non-normal morphometric characteristics (HL, EN, TAD, D3L, TL), 23.98% of the variation was associated with body size (Table
PCA morphometric comparisons of four Kurixalus species from Taiwan. Scatterplots of (A) principal components 1 and 2, and (B) principal components 2 and 3 of size-adjusted morphometric data for male frogs of the four Kurixalus species. The 95% confidence ellipses for each population (ELM) are shown.
The calls of the two new species and of K. eiffingeri were found to be different in maximum frequency, single note duration, and time interval between notes of mating calls. The minimum frequency among the three species was not different (Table
Comparisons of the characteristics of slow mating calls among K. eiffingeri, K. berylliniris sp. n. and K. wangi sp. n.
Characteristics | K. eiffingeri vs. K. berylliniris* | K. berylliniris* vs. K. wangi* | K. eiffingeri vs. K. wangi* |
---|---|---|---|
WMWODDS (95% CI) | WMWODDS (95% CI) | WMWODDS (95% CI) | |
Maximum frequency | 3.4 (1.59–10) | Inf (Inf-Inf) | 2.67 (1.391–5.88) |
Minimum frequency | 1.2 (0.571–2.67) | 0.18 (0–0.54) | 0.53 (0.25–1.04) |
Single note duration | 0.63 (0.29–1.32) | 0 (0–0) | 0.37 (0.17–0.72) |
Time interval between notes of mating | 0.07 (0–0.22) | 0 (0–0) | 0 (0–0) |
Width of frequency | 13.67 (4.5-Inf) | Inf (Inf-Inf) | 12.75 (5.11-Inf) |
Dominant frequency | 2.38 (1.2–6.33) | 3 (1–9) | 1.5 (0.77–3.23) |
As demonstrated by the high bootstrap support, the robustness of the phylogenetic relationship of the three rhacophorid genera is strong. Based on this robust phylogenetic tree, we found that the among-genera genetic distances were greater than the within-genus genetic distance (Fig.
Phylogenetic relationship of all Kurixalus species from Taiwan. A phylogram showing the phylogenetic relationships of the four Kurixalus species, obtained by a maximum likelihood search based on 1207 nucleotides from mtDNA CO1 and 16S rRNA genes. Feihyla palpebralis and Rhacophorus moltrechti were used as outgroups. The three values on each branch are maximum likelihood (ML), maximum parsimony (MP), and neighbor-joining (NJ) analyses with bootstrapping support based on 2000 replicates. Bootstrapping values below 50% are not shown. (JP: Ryukyu Islands of Japan; N. TW: northern Taiwan; C. TW: central Taiwan).
Geographic distribution and genetic structures of K. eiffingeri and the two newly discovered cryptic species from Taiwan and its two adjacent islands. Red: K. eiffingeri; Green: K. berylliniris sp. n.; B: K. wangi sp. n. Bold lines mark the boundaries of each species’ distribution, dotted lines discriminate different genetic groups intra species. Below: a consensus ML tree to show the variation between haplotypes. GenBank number accession numbers KT259055–KT259131.
The phylogenetic consensus tree of K. wangi sp. n. presented a star-like haplotype minimum spanning network with a core ancestral haplotype (W01) and ten derivative haplotypes (W02-W11) (Fig.
The indicators of genetic diversity – haplotype diversity (Hd), nucleotide diversity (Pi), and number of haplotypes of each population – are shown in Table
1 | Dorsum with a brown “) (“ saddle-shaped marking, two arms of the marking not touching each other at mid-dorsum; venter with two large brownish rounded blotches in axilla region; males with very weak nuptial pad; iris golden speckled with brown; cloacal opening of female without supracloacal flap | K. idiootocus |
– | Dorsum with a X- or Y-shaped marking, two arms of the marking touching each other at mid-dorsum; venter without blotches; males with greatly expanded nuptial pad; cloacal opening in females with supracloacal flap | 2 |
2 | Belly smooth; two spots present on upper eyelids, separated from each other, not in contact with marking on back; medial palmar tubercle larger than lateral one; iris emerald to light green | K. berylliniris sp. n. |
– | Belly granular or shagreened; spots on upper eyelids in contact each other, forming a dark bar or connecting with the X-marking on back; two palmar tubercles equal in width; iris golden | 3 |
3 | Tubercles on lateral margin of finger IV connected with dermal fringe; venter whitish with very little pigmentation; loreal region oblique; canthus rostralis curved | K. wangi sp. n. |
– | Tubercles on lateral margin of finger IV separated from each other; venter with numerous fine brownish dots, especially in the gular region; loreal region vertical; canthus rostralis straight | K. eiffingeri |
1 | Lentic tadpole, mouth antero-ventral, tooth formula 5(3-5)/3 or 5(2-5)/3 | K. idiootocus |
– | Oophagous tadpole, mouth terminal or antero-dorsal, tooth row three or less on upper lip, two or less in lower lip | 2 |
2 | Dorsal fin originates at base of tail muscle | K. eiffingeri |
– | Dorsal fin originates on posterior body | 3 |
3 | Dorsum flat in profile; nostril equidistant between upper lip and eye; deep transverse groove on upper lip; a ridge present from lateral margin of upper lip to nostril; gular region and tail muscle heavily pigmented | K. berylliniris sp. n. |
– | Dorsum sloping in profile; nostril closer to upper lip than to eye; inconspicuous transverse groove on upper lip; no ridge from lateral margin of upper lip to nostril; gular region and tail muscle without pigmentation, or with only small scattered spots | K. wangi sp. n. |
Based on the 1) different mating call characteristics, 2) different timing of mating calls, 3) diversified morphological characteristics and genetic composition, 4) no interspecies gene flow indicated by extremely high Fst and low Nm, and 5) sufficient genetic divergences among species (
Unlike previous researchers who did not note the within-species variation of mating calls (
The guts of tadpoles of the two new species contained a yellow ‘yolky’ substance. When the same characteristic was observed in K. eiffingeri it was confirmed as tadpole oophagy (
Previous reports estimated the distribution of K. eiffingeri to be up to 2000 m in mountain forests all over the island of Taiwan. These records were problematic in that they primarily relied on mating call surveys. Our study not only demonstrated the usefulness of advanced voice recording systems in identifying the new species but also highlighted the importance of collecting voucher specimens. In addition to the two newly described species and K. eiffingeri, there is one further species in this genus, K. idiootocus. Until 1987, K. idiootocus was treated as a subgroup within K. eiffingeri (
SP Wu envisioned the original idea, executed this study and wrote the manuscript; CC Huang helped with the statistical methods, data analysis, and paper writing; CL Tsai and TE Lin performed the data analyses; JJ Jhang measured and analyzed the mating call data; SH Wu described adult and tadpole morphology, performed anatomical studies, morphometric analyses and proofread the paper. All the authors contributed to this paper sufficiently.
We would like to show our gratitude to HM Huang for collecting the first specimen of K. berylliniris sp. n.; to JH Chu, CC Hwang, HJ Su and SM Lin for advice on molecular analysis; to H Masaki for providing specimens of K. eiffingeri from the Ryukyu islands, to CY Hsiao for photographs; to YT Chung for his dedicated fieldwork; to CC Wu for performing bench work; and to CF Lin for providing K. eiffingeri tadpoles. Thanks are extended to CJ Huang, CH Chang, CW Lin and YC Liu for assistance with the fieldwork. We appreciate the help of SF Chan with vocal analysis and YH Chen for partial sample contribution. The map of Taiwan was kindly prepared by PF Lee. This study was partially supported by the Council of Agriculture Grant (94–admin–4.1–conserv–03(2)), by Shei-Pa National Park, Ministry of Interior Grant (094-301020500G1-005), and a support from Mr. CC Chen. RE Brown greatly improved an early draft of the manuscript.
Buergeria japonica: NCHUZOOL: 4021, 4825, 4826 (Taiwan). B. robusta: NCHUZOOL 4025, 4027, 4831 (Taiwan). Kurixalus eiffingeri: NTU 927–28, 931-32, 939, 1052, 1054–56, 1649 (Shitou, Nantou); 1058-67, 1645–46, 1769 (Wulai, Taipei); NCHUZOOL: 2502–03, 2508, 2509(C/S), 2514–15, 2517–19, 2523–24, 2525(C/S), 2526–27, 2535, 2536(C/S), 2537, 2538(C/S), 2539–40, 2545–47, 2550–51, 2829, 2831, 4018(C/S), 11320, 11436–40 (Shindian, Taipei); 11333 (Shitou, Nantou). K. idiootocus: NTU 929, 1005–09, 1033, 1035–37, 1038–42 (Shiding, Taipei); 1045 (Suao, Yilan); 1010 (holotype of Chirixalus idiootocus), 1011, 1013–29, 1657, 1695–96, (Yanminshan, Taipei); 1658 (Shindien, Taipei); 1708–09 (Juchi, Chiayi); 1770 (Wulai, Taipei); NCHUZOOL 1010, 2780, 2782, 2784, 2785(C/S), 2787–88, 2792, 2795, 2796(C/S), 2805, 2845, 2847, 2954, 2956(C/S), 2959–63, 3709, 3711(C/S), 4304(C/S), 4990, 4993, 4995(C/S), 7402. K. wangi sp. n.: NTU 1043–1044, 1046, 1647-48 (Manjo, Pingtung County).
Abbreviation | Morphometric characteristic |
---|---|
SVL | snout-vent length |
HW | head width |
HL | head length |
IN | internarial distance |
EN | eye-narial distance |
ED | horizontal eye diameter |
DFE | distance between the anterior margins of eyes |
DBE | distance between the posterior margins of eyes |
UEW | upper eyelid width |
IO | interorbital distance |
TAD | tympanic annulus diameter |
AXI | between posterior margins of the upper arm |
AGD | axilla-groin distance |
UAW | forearm length |
PAL | manus length |
F1L | ength of first finger from base of palmar tubercle to tip of third finger disc |
D3L | width of third finger disc |
FEL | femur length |
TBL | tibia length |
TSL | tarsus length |
FOL | foot length from proximal margin of inner metatarsal tubercle to tip of fourth toe |
TL | first toe length |
IML | inner metatarsal tubercle length |
T4D | disc width of fourth toe |
Species | Sampled locality | Gene sequences | No. of GenBank | Voucher specimen |
---|---|---|---|---|
K. eiffingeri | Ryukyu Islands, Japan | mtDNA CO 1 | DQ468681 | NTUMA 2427 |
mtDNA 16S | DQ468673 | |||
K. eiffingeri | Wulai, Taipei, northen Taiwan | mtDNA CO 1 | DQ468680 | NCHUZOOL 11320 |
mtDNA 16S | DQ468672 | |||
K. eiffingeri | Xitou, Nantou, central Taiwan | mtDNA CO 1 | DQ468678 | NCHUZOOL 11333 |
mtDNA 16S | DQ468670 | |||
K. berylliniris sp. n. | Beinan, Taitung, eastern Taiwan | mtDNA CO 1 | DQ468677 | ASIZAM 00053 |
mtDNA 16S | DQ468669 | |||
K. wangi sp. n. | Shouka, Pintung, southern Taiwan | mtDNA CO 1 | DQ468679 | ASIZAM 00055 |
mtDNA 16S | DQ468671 | |||
K. idiootocus | Wulai, Taipei, northern Taiwan | mtDNA CO 1 | DQ468682 | NA |
mtDNA 16S | DQ468674 | |||
Feihyla palpebralis | Yunnan, China | mtDNA CO 1 | DQ468683 | NA |
mtDNA 16S | DQ468675 | |||
Rhacophorus moltrechti | Antong, Hualien, eastern Taiwan | mtDNA CO 1 | DQ468684 | NA |
mtDNA 16S | DQ468676 |
Genetic distances among Kurixalus species and two outgroup taxa in CO 1 gene (above diagonal) and 16S rRNA gene (below diagonal). Numbers on top row refer to species shown on the left column. Genetic distances are shown as percentage.
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 |
---|---|---|---|---|---|---|---|---|
1. K. eiffingeri (JP) | – | 6.30 | 9.13 | 11.23 | 10.85 | 15.02 | 21.36 | 24.99 |
2. K. eiffingeri (NTW) | 0.92 | – | 9.48 | 9.92 | 10.47 | 16.21 | 20.51 | 25.94 |
3. K. eiffingeri (CTW) | 3.41 | 3.21 | – | 10.84 | 13.12 | 14.20 | 20.51 | 24.33 |
4. K. berylliniris sp. n. | 3.19 | 3.00 | 4.77 | – | 10.65 | 14.36 | 19.21 | 23.89 |
5. K. wangi sp. n. | 3.80 | 3.60 | 4.19 | 3.98 | – | 14.16 | 21.17 | 25.06 |
6. K. idiootocus | 5.97 | 5.97 | 5.76 | 6.15 | 6.59 | – | 18.18 | 23.06 |
7. F. palpebralis | 10.68 | 10.90 | 11.56 | 11.34 | 11.13 | 11.61 | – | 23.79 |
8. R. moltrechti | 14.04 | 13.12 | 14.74 | 14.48 | 14.49 | 14.94 | 15.40 | – |
Comparisons of measurements and ratios of tadpoles of three oophagus Kurixalus species from Taiwan (ANOVA significant level, *: 0.01 < p < 0.05; **: p < 0.01).
species | berylliniris | wangi | eiffingeri | ANOVA |
---|---|---|---|---|
measurements | n=5 | n=10 | n=8 | |
TL | 24.89 ± 5.21 | 18.60 ± 2.44 | 16.75 ± 2.85 | ** |
BL | 7.92 ± 1.72 | 6.68 ± 1.01 | 5.54 ± 1.01 | ** |
CL | 16.97 ± 3.51 | 11.92 ± 1.79 | 11.22 ± 1.87 | * |
TH | 4.84 ± 1.10 | 4.04 ± 0.84 | 2.94 ± 0.66 | ** |
TM | 1.99 ± 0.39 | 1.86 ± 0.38 | 1.63 ± 0.31 | |
NA | 1.99 ± 0.21 | 1.59 ± 0.23 | 1.30 ± 0.20 | ** |
IN | 1.89 ± 0.51 | 1.59 ± 0.27 | 1.38 ± 0.29 | * |
MW | 1.94 ± 0.71 | 1.57 ± 0.38 | 1.37 ± 0.28 | |
BL / TL | 0.32 ± 0.01 | 0.36 ± 0.04 | 0.33 ± 0.01 | * |
CL / TL | 0.68 ± 0.01 | 0.64 ± 0.04 | 0.67 ± 0.01 | * |
TH / TL | 0.19 ± 0.01 | 0.22 ± 0.04 | 0.17 ± 0.01 | * |
TM / TL | 0.08 ± 0.00 | 0.10 ± 0.02 | 0.10 ± 0.00 | * |
NA / TL | 0.08 ± 0.01 | 0.09 ± 0.04 | 0.08 ± 0.00 | |
IN / TL | 0.08 ± 0.01 | 0.09 ± 0.01 | 0.08 ± 0.00 | |
MW / TL | 0.08 ± 0.01 | 0.08 ± 0.01 | 0.08 ± 0.01 | |
TH / CL | 0.28 ± 0.02 | 0.34 ± 0.08 | 0.26 ± 0.02 | * |
TM / MW | 1.08 ± 0.21 | 1.20 ± 0.11 | 1.19 ± 0.11 | |
IN / NA | 0.94 ± 0.16 | 0.99 ± 0.05 | 1.06 ± 0.08 | |
CL / BL | 2.15 ± 0.10 | 1.81 ± 0.27 | 2.03 ± 0.11 | ** |
TH / TM | 2.42 ± 0.19 | 2.18 ± 0.18 | 1.80 ± 0.09 | ** |
Body size variation (SVL) in female and male Kurixalus species from Taiwan (mean ± standard deviation). (**: significant level < 0.01).
Species | female | male | t-test |
---|---|---|---|
K. berylliniris sp. n. | 37.8±7.1 | 34.4±4.1 | 0.178 |
n = 7 | n = 13 | ||
K. eiffingeri | 33.7±2.9 | 31.1±2.3 | ** |
n = 15 | n = 20 | ||
K. wangi sp. n. | 34.3±1.8 | 30.0±0.9 | ** |
n = 8 | n = 17 | ||
K. idiootocus | 36.3±2.5 | 26.9±1.4 | ** |
n = 3 | n = 21 | ||
ANOVA | F3,29 = 1.964 | F3,67 = 30.227** |
Analysis of covariance of morphometric characteristics of males and females among four Kurixalus species.
measurement | berylliniris sp. n. | eiffingeri | idiootocus | wangi sp. n. | equal slope | equal mean |
---|---|---|---|---|---|---|
male | ||||||
HW | 10.59 | 10.83 | 10.56 | 11.21 | ** | 0.001 |
HL | 8.16 | 8.04 | 8.21 | 7.96 | ** | |
IN | 3.29 | 3.41 | 3.20 | 3.31 | ** | 0.028 |
EN | 3.08 | 2.91 | 2.98 | 3.21 | ** | 0.004 |
ED | 3.95 | 4.10 | 4.27 | 4.11 | ** | |
UEW | 2.82 | 3.12 | 3.27 | 3.03 | ** | 0.019 |
DFE | 6.28 | 6.23 | 6.25 | 6.38 | ** | |
DBE | 9.55 | 9.76 | 9.67 | 9.99 | ** | 0.046 |
IO | 3.69 | 3.81 | 3.95 | 4.00 | ** | 0.005 |
TAD | 1.93 | 1.33 | 1.76 | 1.87 | ** | |
AXI | 9.25 | 9.72 | 9.82 | 9.93 | ** | |
AGD | 15.47 | 14.48 | 14.91 | 13.42 | ** | 0.000 |
UAW | 6.12 | 5.40 | 5.70 | 5.60 | ** | 0.000 |
PAL | 9.56 | 10.10 | 9.21 | 9.06 | ** | 0.000 |
F1L | 4.95 | 5.06 | 4.34 | 4.60 | ** | 0.000 |
D3L | 1.30 | 1.86 | 1.78 | 1.56 | ** | 0.000 |
FEL | 14.50 | 14.00 | 13.74 | 14.64 | ** | 0.010 |
TBL | 15.51 | 14.59 | 13.99 | 14.62 | ** | 0.002 |
TSL | 7.37 | 6.94 | 6.78 | 6.93 | ** | 0.028 |
FOL | 13.76 | 13.66 | 12.94 | 12.48 | ** | 0.000 |
TL | 5.00 | 5.00 | 4.43 | 4.29 | ** | 0.000 |
T4D | 0.99 | 1.51 | 1.32 | 1.33 | ** | 0.000 |
IML | 1.27 | 1.36 | 1.39 | 1.23 | ** | |
female | ||||||
HW | 12.20 | 12.22 | 12.16 | 12.67 | ** | 0.030 |
HL | 9.17 | 9.08 | 8.88 | 9.37 | ** | |
IN | 3.77 | 3.75 | 3.73 | 3.74 | ** | |
EN | 3.53 | 3.31 | 3.25 | 3.48 | ** | |
ED | 4.25 | 4.56 | 4.05 | 4.52 | ** | |
UEW | 3.28 | 3.46 | 3.48 | 3.33 | ** | |
DFE | 7.02 | 7.02 | 6.93 | 7.02 | ** | |
DBE | 10.99 | 10.97 | 10.61 | 11.17 | ** | |
IO | 4.19 | 4.27 | 4.57 | 4.30 | ** | |
TAD | 2.20 | 2.23 | 2.21 | 2.09 | ** | |
AXI | 11.27 | 11.43 | 11.57 | 11.22 | ** | |
AGD | 16.49 | 16.94 | 17.55 | 15.36 | ** | |
UAW | 6.65 | 6.36 | 6.89 | 6.31 | ** | |
PAL | 11.47 | 11.35 | 10.22 | 9.71 | ** | 0.000 |
F1L | 5.52 | 5.52 | 5.08 | 4.92 | ** | 0.002 |
D3L | 1.62 | 2.04 | 1.94 | 1.72 | ** | 0.005 |
FEL | 16.51 | 15.60 | 15.93 | 16.53 | ** | 0.025 |
TBL | 17.21 | 16.66 | 16.27 | 16.56 | ** | |
TSL | 8.01 | 7.74 | 8.06 | 7.83 | ** | |
FOL | 15.83 | 15.35 | 14.93 | 14.08 | ** | 0.000 |
TL | 5.85 | 5.79 | 5.25 | 4.94 | ** | 0.001 |
T4D | 1.17 | 1.65 | 1.81 | 1.46 | ** | 0.001 |
IML | 1.48 | 1.49 | 1.60 | 1.43 | ** |
Factor loadings of the first three principal components of 18 size-adjusted morphometric characteristics of males of four Kurixalus species. Absolute values of loadings greater than 0.50 in boldface.
Character | PC 1 | PC 2 | PC 3 |
---|---|---|---|
Eigenvalue | 4.316 | 3.061 | 2.308 |
% variation | 23.98 | 17 | 12.82 |
HW | 0.609 | -0.299 | 0.081 |
IN | 0.615 | 0.147 | -0.237 |
ED | 0.590 | -0.512 | 0.041 |
UEW | 0.581 | -0.471 | -0.144 |
DFE | 0.742 | -0.292 | 0.198 |
DBE | 0.739 | -0.424 | 0.054 |
UEW | 0.449 | -0.273 | 0.245 |
AXI | 0.107 | 0.014 | -0.354 |
AGD | -0.156 | 0.162 | -0.123 |
UAW | 0.112 | 0.089 | 0.682 |
PAL | 0.418 | 0.604 | -0.520 |
F1L | 0.293 | 0.698 | -0.323 |
FEL | 0.431 | 0.413 | 0.479 |
TBL | 0.453 | 0.584 | 0.527 |
TSL | 0.300 | 0.507 | 0.502 |
FOL | 0.413 | 0.686 | -0.150 |
T4D | 0.533 | -0.018 | -0.564 |
IML | 0.582 | 0.015 | -0.206 |
Genetic variation of three Kurixalus species from Taiwan according to mtDNA CO1 gene partial sequence.
N | Number of haplotypes | Haplotype diversity (Hd) | Nucleotide diversity (Pi) | |
---|---|---|---|---|
K. berylliniris sp. n. | 55 | 9 | 0.82088 | 0.02271 |
K. wangi sp. n. | 69 | 11 | 0.85209 | 0.00443 |
K. eiffingeri | 219 | 53 | 0.94600 | 0.05524 |