Corresponding author: Shu-Ping Wu (
Academic editor: F. Andreone
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
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
There are four genera (
In our study,
The type specimens of frogs and tadpoles of the two new
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
Morphometric characteristics of adult specimens: snout-vent length head width head length internarial distance eye-narial distance horizontal eye diameter distance between the anterior margins of eyes distance between the posterior margins of eyes upper eyelid width interorbital distance tympanic annulus diameter distance between the axillae, between posterior margins of the upper arm axilla-groin distance forearm length manus length length of first finger from base of palmar tubercle to tip of third finger disc width of third finger disc femur length tibia length tarsus length foot length from proximal margin of inner metatarsal tubercle to tip of fourth toe first toe length inner metatarsal tubercle length disc width of fourth toe
In addition, the eggs and tadpole morphometric characteristics were measured comprising total length body length tail length tail height tail muscle height internarial distance distance between eyes tail muscle width
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 principal component analysis
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
Whole genomic DNA was extracted from muscle tissue of fresh or ethanol-preserved specimens using the procedure originally described by cytochrome
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 ( neighbor-joining
The Bayesian tree and the posterior probability distribution were determined using the program MrBayes 3.1 ( Monte Carlo Markov Chain (Maximum Parsimony) tree bisection reconnection
Partial sequences of mtDNA (number of immigrants per generation, nucleotide diversity haplotype diversity
Holotype of
Four
Measurements (in mm) of type series and other specimens of
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) | |||||
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
including the holotype
Ligia timber trail, 1250 meters above sea level, Taitung County, Taiwan, Republic of China (Fig.
A moderate-sized
The epithet
Habitus moderately slender and somewhat flattened, size moderate (
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.
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.
Nesting sites of three tree-hole breeding
Dorsal view of tadpole head region of three oophagus
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
Lateral view of tadpoles of three oophagus
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.
Advertisement calls of four
Measurements of advertisement calls of
Species | ||||||
---|---|---|---|---|---|---|
|
2901 (89) | 2517(106) | 384 (80) | 158 (56) | 3195(1060) | 2704(35) |
|
2961 (71) | 2518 (124) | 443 (97) | 103 (42) | 1562(1442) | 2772(360) |
|
3185(194) | 2399 (122) | 786 (192) | 99 (19) | 1122 (230) | 2841(145) |
|
3072 (47) | 2565 (62) | 507 (62) | 57 (15) | 115 (22) | 2848(59) |
|
3034 (59) | 2550 (54) | 484 (90) | 154 (27) | 2063 (121) | 2772(260) |
|
2889 (46) | 2412 (64) | 477 (80) | 48 (16) | 1900 (40) | 2647(62) |
: maximum frequency : minimum frequency : width of frequency (MAX-MIN) : single note duration : time interval between notes : dominant frequency
Eggs and tadpoles were found in the pooled water in decaying trunks of tree ferns,
Shouka timber trail, 400 meters above sea level, Pingtung County, Taiwan, Republic of China (Fig.
A small to moderate-sized
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 (
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.
Holotype of
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.
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.
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
In the
The calls of the two new species and of
Comparisons of the characteristics of slow mating calls among
Characteristics | |||
---|---|---|---|
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) |
slow call
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 maximum likelihood maximum parsimony neighbor-joining : Ryukyu Islands of Japan : northern Taiwan : central Taiwan
Geographic distribution and genetic structures of
The phylogenetic consensus tree of
The indicators of genetic diversity – haplotype diversity (
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 |
|
– | 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 | 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 |
|
– | 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 | Tubercles on lateral margin of finger IV connected with dermal fringe; venter whitish with very little pigmentation; loreal region oblique; canthus rostralis curved |
|
– | 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 |
|
1 | Lentic tadpole, mouth antero-ventral, tooth formula 5(3-5)/3 or 5(2-5)/3 |
|
– | Oophagous tadpole, mouth terminal or antero-dorsal, tooth row three or less on upper lip, two or less in lower lip |
|
2 | Dorsal fin originates at base of tail muscle |
|
– | Dorsal fin originates on posterior body |
|
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 |
|
– | 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 |
|
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
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
Previous reports estimated the distribution of
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;
We would like to show our gratitude to HM Huang for collecting the first specimen of
Morphometric characteristics and the abbreviations in this study.
Abbreviation | Morphometric characteristic |
---|---|
|
snout-vent length |
|
head width |
|
head length |
|
internarial distance |
|
eye-narial distance |
|
horizontal eye diameter |
|
distance between the anterior margins of eyes |
|
distance between the posterior margins of eyes |
|
upper eyelid width |
|
interorbital distance |
|
tympanic annulus diameter |
|
between posterior margins of the upper arm |
|
axilla-groin distance |
|
forearm length |
|
manus length |
|
ength of first finger from base of palmar tubercle to tip of third finger disc |
|
width of third finger disc |
|
femur length |
|
tibia length |
|
tarsus length |
|
foot length from proximal margin of inner metatarsal tubercle to tip of fourth toe |
|
first toe length |
|
inner metatarsal tubercle length |
|
disc width of fourth toe |
DNA sequences used in this study and their GenBank accession numbers.
Species | Sampled locality | Gene sequences | No. of GenBank | Voucher specimen |
---|---|---|---|---|
|
Ryukyu Islands, Japan | mtDNA CO 1 |
|
NTUMA 2427 |
mtDNA 16S |
|
|||
|
Wulai, Taipei, northen Taiwan | mtDNA CO 1 |
|
NCHUZOOL 11320 |
mtDNA 16S |
|
|||
|
Xitou, Nantou, central Taiwan | mtDNA CO 1 |
|
NCHUZOOL 11333 |
mtDNA 16S |
|
|||
Beinan, Taitung, eastern Taiwan | mtDNA CO 1 |
|
ASIZAM 00053 | |
mtDNA 16S |
|
|||
Shouka, Pintung, southern Taiwan | mtDNA CO 1 |
|
ASIZAM 00055 | |
mtDNA 16S |
|
|||
|
Wulai, Taipei, northern Taiwan | mtDNA CO 1 |
|
|
mtDNA 16S |
|
|||
|
Yunnan, China | mtDNA CO 1 |
|
|
mtDNA 16S |
|
|||
|
Antong, Hualien, eastern Taiwan | mtDNA CO 1 |
|
|
mtDNA 16S |
|
Genetic distances among
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 |
---|---|---|---|---|---|---|---|---|
1. |
– | 6.30 | 9.13 | 11.23 | 10.85 | 15.02 | 21.36 | 24.99 |
2. |
0.92 | – | 9.48 | 9.92 | 10.47 | 16.21 | 20.51 | 25.94 |
3. |
3.41 | 3.21 | – | 10.84 | 13.12 | 14.20 | 20.51 | 24.33 |
4. |
3.19 | 3.00 | 4.77 | – | 10.65 | 14.36 | 19.21 | 23.89 |
5. |
3.80 | 3.60 | 4.19 | 3.98 | – | 14.16 | 21.17 | 25.06 |
6. |
5.97 | 5.97 | 5.76 | 6.15 | 6.59 | – | 18.18 | 23.06 |
7. |
10.68 | 10.90 | 11.56 | 11.34 | 11.13 | 11.61 | – | 23.79 |
8. |
14.04 | 13.12 | 14.74 | 14.48 | 14.49 | 14.94 | 15.40 | – |
Abbreviations: : Ryukyu Islands of Japan : northern Taiwan : central Taiwan
Comparisons of measurements and ratios of tadpoles of three oophagus
species |
|
|
|
ANOVA |
---|---|---|---|---|
measurements | n=5 | n=10 | n=8 | |
|
24.89 ± 5.21 | 18.60 ± 2.44 | 16.75 ± 2.85 | ** |
|
7.92 ± 1.72 | 6.68 ± 1.01 | 5.54 ± 1.01 | ** |
|
16.97 ± 3.51 | 11.92 ± 1.79 | 11.22 ± 1.87 | * |
|
4.84 ± 1.10 | 4.04 ± 0.84 | 2.94 ± 0.66 | ** |
|
1.99 ± 0.39 | 1.86 ± 0.38 | 1.63 ± 0.31 | |
|
1.99 ± 0.21 | 1.59 ± 0.23 | 1.30 ± 0.20 | ** |
|
1.89 ± 0.51 | 1.59 ± 0.27 | 1.38 ± 0.29 | * |
|
1.94 ± 0.71 | 1.57 ± 0.38 | 1.37 ± 0.28 | |
0.32 ± 0.01 | 0.36 ± 0.04 | 0.33 ± 0.01 | * | |
0.68 ± 0.01 | 0.64 ± 0.04 | 0.67 ± 0.01 | * | |
0.19 ± 0.01 | 0.22 ± 0.04 | 0.17 ± 0.01 | * | |
0.08 ± 0.00 | 0.10 ± 0.02 | 0.10 ± 0.00 | * | |
0.08 ± 0.01 | 0.09 ± 0.04 | 0.08 ± 0.00 | ||
0.08 ± 0.01 | 0.09 ± 0.01 | 0.08 ± 0.00 | ||
0.08 ± 0.01 | 0.08 ± 0.01 | 0.08 ± 0.01 | ||
0.28 ± 0.02 | 0.34 ± 0.08 | 0.26 ± 0.02 | * | |
1.08 ± 0.21 | 1.20 ± 0.11 | 1.19 ± 0.11 | ||
0.94 ± 0.16 | 0.99 ± 0.05 | 1.06 ± 0.08 | ||
2.15 ± 0.10 | 1.81 ± 0.27 | 2.03 ± 0.11 | ** | |
2.42 ± 0.19 | 2.18 ± 0.18 | 1.80 ± 0.09 | ** |
Body size variation (
Species | female | male | t-test |
---|---|---|---|
37.8±7.1 | 34.4±4.1 | 0.178 | |
n = 7 | n = 13 | ||
|
33.7±2.9 | 31.1±2.3 | ** |
n = 15 | n = 20 | ||
34.3±1.8 | 30.0±0.9 | ** | |
n = 8 | n = 17 | ||
|
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
measurement |
|
|
equal slope | equal mean | ||
---|---|---|---|---|---|---|
male | ||||||
|
10.59 | 10.83 | 10.56 | 11.21 | ** | 0.001 |
|
8.16 | 8.04 | 8.21 | 7.96 | ** | |
|
3.29 | 3.41 | 3.20 | 3.31 | ** | 0.028 |
|
3.08 | 2.91 | 2.98 | 3.21 | ** | 0.004 |
|
3.95 | 4.10 | 4.27 | 4.11 | ** | |
|
2.82 | 3.12 | 3.27 | 3.03 | ** | 0.019 |
|
6.28 | 6.23 | 6.25 | 6.38 | ** | |
|
9.55 | 9.76 | 9.67 | 9.99 | ** | 0.046 |
|
3.69 | 3.81 | 3.95 | 4.00 | ** | 0.005 |
|
1.93 | 1.33 | 1.76 | 1.87 | ** | |
|
9.25 | 9.72 | 9.82 | 9.93 | ** | |
|
15.47 | 14.48 | 14.91 | 13.42 | ** | 0.000 |
|
6.12 | 5.40 | 5.70 | 5.60 | ** | 0.000 |
|
9.56 | 10.10 | 9.21 | 9.06 | ** | 0.000 |
|
4.95 | 5.06 | 4.34 | 4.60 | ** | 0.000 |
|
1.30 | 1.86 | 1.78 | 1.56 | ** | 0.000 |
|
14.50 | 14.00 | 13.74 | 14.64 | ** | 0.010 |
|
15.51 | 14.59 | 13.99 | 14.62 | ** | 0.002 |
|
7.37 | 6.94 | 6.78 | 6.93 | ** | 0.028 |
|
13.76 | 13.66 | 12.94 | 12.48 | ** | 0.000 |
|
5.00 | 5.00 | 4.43 | 4.29 | ** | 0.000 |
|
0.99 | 1.51 | 1.32 | 1.33 | ** | 0.000 |
|
1.27 | 1.36 | 1.39 | 1.23 | ** | |
female | ||||||
|
12.20 | 12.22 | 12.16 | 12.67 | ** | 0.030 |
|
9.17 | 9.08 | 8.88 | 9.37 | ** | |
|
3.77 | 3.75 | 3.73 | 3.74 | ** | |
|
3.53 | 3.31 | 3.25 | 3.48 | ** | |
|
4.25 | 4.56 | 4.05 | 4.52 | ** | |
|
3.28 | 3.46 | 3.48 | 3.33 | ** | |
|
7.02 | 7.02 | 6.93 | 7.02 | ** | |
|
10.99 | 10.97 | 10.61 | 11.17 | ** | |
|
4.19 | 4.27 | 4.57 | 4.30 | ** | |
|
2.20 | 2.23 | 2.21 | 2.09 | ** | |
|
11.27 | 11.43 | 11.57 | 11.22 | ** | |
|
16.49 | 16.94 | 17.55 | 15.36 | ** | |
|
6.65 | 6.36 | 6.89 | 6.31 | ** | |
|
11.47 | 11.35 | 10.22 | 9.71 | ** | 0.000 |
|
5.52 | 5.52 | 5.08 | 4.92 | ** | 0.002 |
|
1.62 | 2.04 | 1.94 | 1.72 | ** | 0.005 |
|
16.51 | 15.60 | 15.93 | 16.53 | ** | 0.025 |
|
17.21 | 16.66 | 16.27 | 16.56 | ** | |
|
8.01 | 7.74 | 8.06 | 7.83 | ** | |
|
15.83 | 15.35 | 14.93 | 14.08 | ** | 0.000 |
|
5.85 | 5.79 | 5.25 | 4.94 | ** | 0.001 |
|
1.17 | 1.65 | 1.81 | 1.46 | ** | 0.001 |
|
1.48 | 1.49 | 1.60 | 1.43 | ** |
Means adjusted to
Factor loadings of the first three principal components of 18 size-adjusted morphometric characteristics of males of four
Character | PC 1 | PC 2 | PC 3 |
---|---|---|---|
Eigenvalue | 4.316 | 3.061 | 2.308 |
% variation | 23.98 | 17 | 12.82 |
|
|
-0.299 | 0.081 |
|
|
0.147 | -0.237 |
|
|
- |
0.041 |
|
|
-0.471 | -0.144 |
|
|
-0.292 | 0.198 |
|
|
-0.424 | 0.054 |
|
0.449 | -0.273 | 0.245 |
|
0.107 | 0.014 | -0.354 |
|
-0.156 | 0.162 | -0.123 |
|
0.112 | 0.089 |
|
|
0.418 |
|
- |
|
0.293 |
|
-0.323 |
|
0.431 | 0.413 | 0.479 |
|
0.453 |
|
|
|
0.300 |
|
|
|
0.413 |
|
-0.150 |
|
|
-0.018 | - |
|
|
0.015 | -0.206 |
Genetic variation of three
N | Number of haplotypes |
Haplotype diversity |
Nucleotide diversity |
|
---|---|---|---|---|
55 | 9 | 0.82088 | 0.02271 | |
69 | 11 | 0.85209 | 0.00443 | |
|
219 | 53 | 0.94600 | 0.05524 |
|
|||
---|---|---|---|
-- | 0.85922 | 0.60226 | |
0.04 | -- | 0.71956 | |
|
0.17 | 0.10 | -- |