Odorrana fengkaiensis, Wang, Ying-Yong, Lau, Michael Wai-Neng, Yang, Jian-Huan, Chen, Guo-Ling, Liu, Zu-Yao, Pang, Hong & Liu, Yang, 2015
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publication ID |
https://doi.org/ 10.11646/zootaxa.3999.2.4 |
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publication LSID |
lsid:zoobank.org:pub:F9AAACFA-4F59-4D95-8456-220289151E43 |
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DOI |
https://doi.org/10.5281/zenodo.5622858 |
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persistent identifier |
https://treatment.plazi.org/id/272687BD-FFCB-6C02-BEBA-6344FA9FFD0D |
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treatment provided by |
Plazi |
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scientific name |
Odorrana fengkaiensis |
| status |
sp. nov. |
Odorrana fengkaiensis View in CoL sp. nov.
Figs. 3 View FIGURE 3 , 4 View FIGURE 4
Holotype. SYS a002265 ( Fig. 3 View FIGURE 3 ), adult male, from Heishiding Nature Reserve (23°27'40.16" N, 111°54'32.80" E; 253 m a.s.l.), Fengkai County, Guangdong Province, China, collected by Ying-Yong Wang (YYW hereafter).
Paratypes. A total of 41 specimens (10 adult males, 31 adult females): 32 specimens from the same locality as the holotype at elevations between 190–510 m a.s.l. by YYW, Jian-Huan Yang (JHY hereafter), Yu-Long Li (YLL hereafter), Jian Zhao (JZ hereafter) and Run-Lin Li (RLL hereafter): 10 adult males, SYS a000174, SYS a000175 and SYS a000176 ( Fig. 4 View FIGURE 4 : B) collected on 2nd May 2008, SYS a000179 and SYS a000181 on 30th May 2008, SYS a000185 on 31st May 2008, SYS a000413 on 15th July 2008, SYS a000792 on 13th May 2010; SYS a001880 on 28th June 2012, SYS a002263 on 12th August 2013; 22 adult females, SYS a000105, 0 106 ( Fig. 4 View FIGURE 4 : Fe), 0 108, 0 110, 0 111, 0 112, 0 127 and 0 128, collected on 10th–20th July 2007, SYS a000182, 0 183 ( Fig. 4 View FIGURE 4 : B) on 2nd May 2008, SYS a000184 on 29th May 2008, SYS a000143 on 23rd March 2008, SYS a000790 and 0 791 on 13th May 2010, SYS a000793 on 26th May 2010, SYS a001879 and 1881 on 28th June 2012, SYS a002160 and 2161 on 2nd June 2013, SYS a002262 ( Fig. 4 View FIGURE 4 : D) on 13th August 2013; SYS a002273 on 19th August 2013, SYS a003345 on 3rd October 2014. Two adult females from Shiwandashan Forest Park (21° 54' 16.36'' N, 107° 54'11.15'' E; at elevations between 292–310 m a.s.l.), Shangsi County, Guangxi, SYS a001025 ( Fig. 4 View FIGURE 4 : F) on 3rd August 2010, SYS a001352 on 16th August 2011. Seven adult females from Tongling Canyon (23°0'11.07"N, 106°40'2.22"E; 450–550 m a.s.l.), Jingxi County, Guangxi Zhuang Autonomous Region, China by JHY, YLL, JZ and RLL: SYS a001043, 1044 and 1045 on 4–6th August 2010, SYS a001366, 1367, 1368 on 20th–21st August 2011, SYS a002253 on 22nd July 2013.
Diagnosis. Odorrana fengkaiensis sp. nov. can be distinguished from other congeners by the following characters: (1) the body length of adult females approximately twice as long as males (SVL 77.8–111.9 mm in females, 37.4–51.8 mm in males); (2) eye large in males, longer than snout length, ED:SNT ratio 1.01–1.16; (3) tympanum of males large and distinct, extremely close to the eye, 0.7–1.4 mm in tympanum–eye distance (TED); (4) supratympanic fold slightly distinct; (5) dorsolateral folds absent; (6) dorsal skin shagreened with dense granules, only males with several large tubercles on dorsum; (7) flanks with tubercles and scattered larger pustules, 8–10 of which usually arranged in a dorsolateral row; (8) venter smooth; ventral spines present in adult males during the breeding season; (9) the heels significant overlapping, tibio-tarsal articulation stretched forward beyond the tip of snout; (10) disks on digits cordiform, pointed and moderately enlarged, FPW:DPW ratio 1.16–1.52 in males, 1.14–1.57 in females; (11) disks of all digits with circum-marginal grooves; (12) relative finger lengths: II <I <IV <III; (13) feet fully webbed; (14) dorsum brown with irregularly reticulated green markings in males and young females, uniformly brown in some old adult females; dorsal limbs and digits brown with black transverse bands; (15) males with velvety nuptial pad on thumb, paired gular pouches; (16) mature oocytes purely black in life, animal pole dark grey and vegetative pole olive in preservative.
Description of holotype. Size moderate (SVL 51.2 mm); head longer than wide (HDL:HDW = 1.16); snout slightly long (SNT:HDL = 0.41), obtusely rounded in dorsal view, rounded in profile, projecting beyond lower jaw; eye large and convex, eye diameter (ED) 1.03 times of snout length (SNT), upper eyelid wider than interorbital distance; top of head flat; canthus rostralis rounded; lip flared; loreal region concave; internasal distance ( IND 5.2 mm) larger than interorbital distance (IOD 4.0 mm); tympanum circular, large, distinctly visible, 0.54 times of eye diameter; tympanum–eye distance small (TED 1.1 mm, TED: TYD = 0.25); tympanic rim elevated relative to skin of temporal region; choanae large, semicircular or inverted triangular, partly concealed by the maxillary shelves; two vomerine ridges markedly elevated, oblique, posteromedial to choanae, separated by a distance barely equal to length of one, bearing prominent teeth; tongue deeply notched posteriorly; paired gular pouches at corners of throat.
Forelimbs moderately robust, hand moderately long, hand length (HDL) 0.3 times of SVL; fingers slender, relative finger lengths II <I <IV <III; fingers free of webbing, tips of fingers moderately dilated, becoming pointed, cordiform disks; width of disk slightly larger than width of distal phalanx of finger III, FPW:DPW = 1.18; terminal phalanges T-shaped; lateroventral circummarginal grooves not meeting at the tip of dicks; subarticular tubercles ovoid, markedly elevated and prominent; supernumerary tubercle below the base of each finger, distinct, significantly smaller than subarticular tubercles; metacarpal tubercle three, inner one elongated and oblong, other two small and indistinct. Hindlimbs robust, relatively long; tibio-tarsal articulation beyond the tip of snout when hindlimb adpressed along the side of the body; heels overlapped when the flexed hindlimbs held at right angles to the body axis; tibia length (TIB) 0.55 times of SVL; foot length (FTL) 0.78 times of SVL; toes long, slender, relative toe lengths I <II <III <V <IV; tips of toes slightly dilated, becoming narrow and long triangular disks; circummarginal grooves present; feet fully webbed to disks; free edges of toes I and V with wide lateral fringes; subarticular tubercles prominent, ovoid; inner metatarsal tubercle elongated, ellipsoid; outer metatarsal tubercle absent; inner tarsal fold absent.
Skin on dorsum shagreened with dense granules; a number of pustules scattered on flanks and dorsum of trunk, eight large pustules forming a longitudinal row; dorsolateral folds absent; supratympanic folds slightly prominent, from posterior corner of eye, curving posteroventrally, extending to a level above insertion of arm; two rictal glands, the anterior one continuous with upper lip and the posterior one above insertion of arm and continuous with supratympanic fold; venter smooth with ventral spinules; distinct granules present on posterior thighs and around cloaca; cloacal opening unmodified, directed posteriorly, at upper level of thighs.
Measurements of holotype (in mm). SVL 51.2, HDL 19.5, HDW 16.8, SNT 7.9, IND 5.2, IOD 4.0, ED 8.1, TYD 4.4, TED 1.1, HND 15.5, FTL 40.1, TIB 28.0, FPW 1.00, DPW 0.85.
Coloration in life. Dorsum of head and body brown with large dark-brown spots, and reticulated green markings anterior to sacral vertebra; upper part of flank brown, lower part of flank light yellow with several large black spots; upper and lower lips brownish yellow with vertical black bars, the one under the eye largest and distinct; rictal glands brownish yellow; dorsal surface of limbs brown with black transverse bands; posterior surface of thigh brown with black and yellowish irregular markings; webbing on feet dark brown with yellowish marking; ventral surface white to yellowish, chest and chin mottled with light-brown blotches; pupil black edged with a striking bright yellow border; iris brown-yellow with black blotches and dense tiny freckles.
Coloration in preservative. Dorsum, flanks and limbs dark-brown, dorsum with large blackish spots; ventral surface of body and limbs creamy white with faint brown spots on chest and chin.
Variation. Measurements and body proportions of type series of Odorrana fengkaiensis sp. nov. are given in Table 3 View TABLE 3 .
The new species shows obvious differences among females and males. SVL of adult females approximately twice as long as of males, the body size of female individuals from Shiwandashan Nature Reserve and Tongling Canyon, Guangxi, significantly greater than from Heishiding Nature Reserve, Fengkai County, Guangdong (SVL of adult females 89.0–111.9 (103.1 ± 7.8) mm from Shangsi and Jingxi, Guangxi; 77.3–109.9 (93.2 ± 8.9) mm from Heishiding Nature Reserve, Guangdong). Dorsum of young adult females garnished with reticulated green to lightgreen markings ( Fig. 4 View FIGURE 4 : B and D), but in some old adult female individuals ( Fig. 4 View FIGURE 4 : E and F), the dorsum uniform brown without green pattern. In froglet ( Fig. 4 View FIGURE 4 : C), the whole body olive, dorsum with brown irregular spots. Disks of digits of female individuals from Guangxi are larger than from Heishiding Nature Reserve (FPW/DPW ratio 1.79–1.94 vs. 1.35–1.80). Moreover, males with velvety nuptial pad on finger I, paired gular pouches; mature oocytes in life almost purely black (SYS a000111, SYS a00184, SYS a002161, SYS a002262), in preservative showed dark grey animal pole and olive vegetative pole.
Comparisons. Comparative data of O. fengkaiensis sp. nov. with 56 recognized species of the genus Odorrana were obtained from the literature ( Ao et al. 2003; Bain et al. 2003, 2009; Bain and Stuart 2006; Fei et al. 2001, 2009, 2012; Li et al. 2008; Matsui 1994; Matsui & Jaafar 2006; Mo et al. 2015; Orlov et al. 2003, 2006; Stuart et al. 2005, 2006, Stuart & Bain 2005; Tung et al. 2008; Yang & Li 1990). Examined specimens of O. versabilis , O. nasuta , O. exiliversabilis , O. graminea , O. tormota , O. zhaoi , O. hainanensis , O. margaretae , O. jingdongensis , O. huangshanensis , O. schmackeri , O. tianmuii , O. lungshengensis , O. wuchuanensis and O. yizhangensis are listed in the Appendix 1.
Odorrana fengkaiensis sp. nov. is most similar to O. bacboensis , O. hainanensis and O. tiannanensis . It differs from O. hainanensis by having relatively large eye in males, ED longer than SNT (ED: SNT ratio 1.01–1.16, median 1.09) vs. usually ED smaller than SNT, (ED:SNT 0.90–1.03, median 0.97) in O. hainanensis ; disks on fingers of males moderately enlarged, FPW:DPW ratio 1.16–1.52 (median 1.34) vs. disks on fingers significantly enlarged, FPW:DPW ratio 1.79–1.94 (median 1.87) in O. hainanensis (see Fig. 7 View FIGURE 7 ); relative finger lengths II <I <IV <III vs. II <IV <I <III for O. hainanensis ; tibio-tarsal articulation forward beyond the tip of snout vs. reaches between anterior corner of the eye and tip of snout in O. hainanensis ; mature oocytes purely black in life vs.
yellowish white with blackish brown animal pole in O. hainanensis . It differs from O. bacboensis by its eye relatively large in males, ED longer than SNT vs. ED shorter than SNT in O. bacboensis (ED 6.6 mm, SNT 8.8 mm in FMNH 255611 ( Bain et al. 2003); ED 5.2 mm, SNT 5.8 mm in SYS a001048); disks on fingers of males moderately enlarged FPW 0.83–1.57 mm, FPW:DPW ratio 1.16–1.52 vs. disks on fingers significantly enlarged (FPW 1.8 mm in FMNH 255611 ( Bain et al. 2003); FPW:DPW ratio 1.87 in SYS a001048) in O. bacboensis ; ventral spines present in adult mature males vs. absent in O. bacboensis . It differs from O. tiannanensis by its disks of fingers moderate enlarged, (FPW 0.83–1.57 mm, median 1.20 mm) in males vs. significantly enlarged (FPW 2.0 mm), mature oocytes purely black in life vs. yellow in O. tiannanensis .
The absence of dorsolateral folds differentiates O. fengkaiensis sp. nov. from the following 24 congeners: O. absita , O. amamiensis , O. banaorum , O. bolavensis , O. exiliversabilis , O. gigatympana , O. graminea , O. hosii , O. indeprensa , O. khalam , O. leporipes , O. livida , O. monjerai , O. narina , O. nasica , O. nasuta , O. orba , O. supranarina , O. tormota , O. trankieni , O. utsunomiyaorum , O. versabilis , O. yentuensis and O. zhaoi .
O. fengkaiensis sp. nov. can be distinguished from the remaining 29 recognized congeners by the combination of the following characters: dorsum brown with irregularly reticulated green markings in males and young females, becoming uniformly brown in old adult females vs. dorsal ground color always green in O. chloronota , O. sinica , O. aureola , O. anlungensis , O. grahami , O. kuangwuensis , O. margaretae , O. ishikawae , O. splendida , O. huanggangensis , O. hejiangensis , O. nanjiangensis , O. schmackeri , O. tianmuii , O. lipuensis , O. lungshengensis , O. wuchuanensis and O. yizhangensis ; body small-sized in mature males, SVL 37.4–51.8 mm vs. considerably larger in O. cangyuanensis (62–69 mm), O. chapaensis (73–83 mm), O. geminate (71–79 mm), O. grahami (70–84 mm), O. jingdongensis (62–82 mm), O. mawphlangensis (80 mm), O. andersonii (68–76 mm), O. junlianensis (73–80 mm), O. kuangwuensis (57 mm), O. lungshengensis (60–67 mm), O. margaretae (78–88 mm), O. wuchuanensis (62–77 mm); having white pectoral spinules in mature males vs. absent in O. anlungensis , O. geminate , O. chapaensis , O. hejiangensis , O. kuangwuensis , O. lungshengensis , O. nanjiangensis , O. chloronota , O. sinica , O. aureola ; present paired gular pouches in males vs. absent in O. lipuensis , O. margaretae and O. wuchuanensis ; the absence of spinules along the dorsum of body and hindlimbs, flanks of trunk, temporal region and upper lip in mature males vs. the presence of spinules along lateral surface of head, extending to posterior dorsum of body in O. andersonii , on dorsal surface of hindlimbs in O. anlungensis , from jaw angle to level of loreal region in O. chapaensis ; on upper lip from snout tip to level of mid-portion of eye in O. geminata , on lateral parts of body, temporal region, and anterior and posterior edge of tympanum in O. lipuensis , on eyelid and around tympanum in O. hejiangensis , on posterior dorsum of body and dorsal surface of hindlimbs in O. lungshengensis , on dorsum in O. mawphlangensis ; tibio-tarsal articulation forward beyond the tip of snout vs. reaches the nostril in O. anlungensis , O. grahami , O. huanggangensis , O. wuchuanensis and O. kuangwuensis , reaches between eye and nostril in O. schmackeri , O. tianmuii , O. macrotympana , reaches tip of snout or between nostril and tip of snout in O. margaretae , O. swinhoana , O. nanjiangensis , O. margaretae , O. lungshengensis ; outer metatarsal tubercle absent vs. present in O. swinhoana , O. rotodora . The new species possesses purely black mature oocytes in life, differentiating it from O. aureola , O. ishikawae , O. anlungensis , O. hejiangensis , O. jingdongensis , O. kuangwuensis , O. lipuensis , O. lungshengensis , O. margaretae , O. swinhoana , O. nanjiangensis , O. wuchuanensis , O. yizhangensis (all with purely whitish yellow to yellow oocytes), O. macrotympana (grey oocytes), O. andersonii , O. grahami , O. huanggangensis , O. junlianensis , O. schmackeri , O. tianmuii (their mature oocytes with dark colored animal pole and light colored vegetative pole in life). Further, the new species differs from O. rotodora by its snout obtusely rounded vs. obtusely pointed; upper lips brownish yellow with vertical black bars vs. light colored without bars in the latter.
Distribution and habitats. Odorrana fengkaiensis sp. nov. is a lowland dweller. At present, it was only found in three localities: Heishiding Nature Reserve (23°25'– 23°30' N, 111°48'– 111°55' E), located in the northern region of the Yunkai Mountains, Fengkai County, Guangdong Province; Shiwandashan Nature Reserve (21°40'– 22°04' N, 107°29'– 108°13' E) and Tongling Canyon (23°0'11.07"N, 106°40'2.22"E), located in the China-Vietnam border area, namely in Shangsi and Jingxi Counties, Guangxi Zhuang Autonomous Region, China. The population from Heishiding Nature Reserve inhabits broad slow-flowing mountainous streams, rivers, pools and near the riparian areas, surrounded by moist subtropical evergreen broadleaved forests at elevations between 190–510 m a.s.l. ( Fig. 6 View FIGURE 6 ); the populations from Shiwandashan Nature Reserve and Tongling Canyon inhabit around broad streams, surrounded by moist subtropical evergreen broadleaved forests or secondary limestone forests at elevations between 290–550 m a.s.l..
Remarks. O. fengkaiensis sp. nov. was found on forest floor, rocks or low shrubs at night, and diving into water by disturbance. They take the shelter under submerged rocks or perhaps as entering holes in the banks during the day. All females were found on roadside and bare forest floor; all males were found on top of the bushes and scrubby branches at height of 10–30 cm, closed to adult females. The adult male individuals frequently emit a short whistling call from end of May to mid-July, and rarely emit voices for the rest of the year.
The diet and behavior of O. fengkaiensis sp. nov. remains poorly known. We dissected SYS a000174 and found seven termites in its stomach. Currently, the tadpoles and eggs of O. fengkaiensis have not been found in the field, showed that the reproduction behavior of this species is rather cryptic. The female individuals were commonly found in May to September, occasionally in March, April, October and November. The male juvenile was found occasionally in September. The female paratypes, including SYS a000111, SYS a00184, SYS a002161, SYS a002262 possessed mature black oocytes in oviduct. While the females, collected before May and after August, possessed under-developed oocytes in ovary. Therefore, the breeding season of this species may be during May to August.
Etymology. The specific epithet “ fengkaiensis ” refers to the locality of the holotype, Fengkai County, Guangdong Province, China. We propose the common English name “Feng-kai Odor Frog” for this species.
TABLE 3. Measurements (in mm) (minimum-maximum [mean ± standard deviation]) of the Odorrana fengkaiensis sp. nov. from Heishiding Nature Reserve, Fengkai Co., Guangdong, Shiwandashan Nature Reserve, Shangsi Co. and Tongling Canyon, Jingxi Co., Guangxi; O. hainanensis from Hainan Island, O. bacboensis from Bainan Village, Napo Co., Guangxi. See materials and methods for abbreviations.
| Odorrana fengkaiensis sp. nov. | Odorrana hainanensis | Odorrana bacboensis | ||
|---|---|---|---|---|
| SVL | Males (n=11) from Fengkai, Females (n=22) from Guangdong Fengkai, Guangdong 37.4–51.8 Ä 48.4 ±4.1Å 77.3–109.9 (93.2 ± 8.9) | Females (n=9) from Shangsi and Jingxi, Guangxi 89.0–111.9 (103.1 ± 7.8) | Males (n=6) from Females (n=5) from Hainan Hainan 49.1–60.7 (52.6 ± 4.3) 94.2–120.4 (109.3 ± 10.9) | Male (n=1) Females (n=2) from from Napo, Napo, Guangxi Guangxi 35.6 78.0–95.0 |
| HDL | 14.8–19.7 Ä 18.9 ± 1.4Å 30.4–39.1 (35.4 ± 2.4) | 31.4–40.5 (36.5 ± 3.5) | 19.2–23.6 (21.1 ± 1.6) 37.3–43.4 (40.5 ± 2.6) | 14.1 26.9–35.4 |
| HDW | 12.4–17.3 Ä 16.3 ± 1.4Å 36.8–36.0 (32.1 ± 2.5) | 29.8–38.4 (34.4 ± 3.2) | 16.5–20.0 (18.0 ± 1.4) 33.9–40.6 (37.7 ± 2.8) | 11.7 23.5–32.8 |
| SNT | 5.8–7.9 Ä 7.2 ± 0.5Å 12.2–16.4 (14.2 ± 1.1) | 14.1–16.9 (15.4 ± 1.1) | 7.0–9.3 (7.9 ± 0.9) 15.3–16.6 (15.9 ± 0.5) | 5.8 12.8–14.4 |
| IND | 4.4–6.0 Ä 5.2 ± 0.5Å 8.2–11.6 (9.8 ± 0.9) | 9.0–11.5 (10.2 ± 0.8) | 5.0–6.6 (5.7 ± 0.6) 9.0–11.7 (10.6 ± 1.1) | 4.2 8.1–9.2 |
| IOD | 3.3–4.3 Ä 4.1 ± 0.3Å 6.6–11.1 (8.6 ± 1.0) | 7.6–10.4 (9.6 ± 1.1) | 3.6–4.6 (4.0 ± 0.4) 8.5–9.8 (9.2 ± 0.6) | 2.5 7.1–7.5 |
| ED | 5.9–8.1 Ä 7.7 ± 0.7Å 10.0–13.5 (11.8 ± 0.9) | 10.5–12.9 (11.7 ± 0.9) | 6.4–9.6 (7.6 ± 1.1) 10.7–12.8 (11.8 ± 1.0) | 5.2 10.1–10.4 |
| TYD | 2.9–4.6 Ä 4.2 ± 0.5Å 5.2–6.9 (6.0 ± 0.5) | 4.6–6.1 (5.5 ± 0.5) | 3.6–5.8 (4.5 ± 0.9) 5.5–6.7 (6.2 ± 0.5) | 3.2 4.7–5.4 |
| TED | 0.7–1.4 Ä 1.2 ± 0.2Å 2.5–5.0 (3.7 ± 0.6) | 2.9–4.5 (3.8 ± 0.4) | 1.7–2.0 (1.9 ± 0.1) 4.7–6.0 (5.2 ± 0.5) | 0.8 2.9–3.4 |
| HND | 11.9–16.3 Ä 15.4 ± 1.2Å 24.7–28.7 (26.8 ± 1.2) | 26.5–30.9 (28.7 ± 1.7) | 14.5–17.9 (16.1 ± 1.2) 27.3–33.1 (29.8 ± 2.3) | 11.1 22.2–26.4 |
| TIB | 22.9–29.7 Ä 27.7 ± 1.9Å 46.0–58.9 (54.8 ± 3.1) | 50.0–64.8 (58.0 ± 4.4) | 28.2–35.3 (31.4 ± 2.9) 58.9–68.6 (63.2 ± 3.8) | 22.0 45.8–52.9 |
| FTL | 31.2–41.8 Ä 39.2 ± 2.8Å 64.7–79.7 (74.1 ± 3.9) | 70.0–85.1 (78.0 ± 4.9) | 38.5–47.9 (42.6 ± 3.3) 77.9–90.9 (84.5 ± 6.1) | 30.4 60.8–71.3 |
| FPW | 0.83–1.57 Ä 1.06 ± 0.19Å 1.87–2.86 (2.32 ± 0.24) | 2.64–3.29 (3.07 ± 0.24) | 1.8–2.1 (1.9 ± 0.1) 3.4–3.9 (3.7 ± 0.2) | 1.25 2.5–2.9 |
| DPW | 0.56–1.03 Ä 0.81 ± 0.12Å 1.39–2.18 (1.72 ± 0.19) | 1.64–2.26 (2.00 ± 0.21) | 0.9–1.1 (1.0 ± 0.1) 2.0–2.4 (2.3 ± 0.2) | 0.67 1.3–1.8 |
| HDL/SVL | 0.37–0.41 Ä 0.39 ± 0.01Å 0.35–0.42 (0.38 ± 0.02) | 0.33–0.37 (0.35 ± 0.01) | 0.38–0.43 (0.40 ± 0.02) 0.36–0.40 (0.37 ± 0.01) | 0.40 0.34–0.37 |
| HDW/SVL | 0.32–0.35 Ä 0.34 ± 0.01Å 0.32–0.37 (0.35 ± 0.01) | 0.32–0.35 (0.33 ± 0.01) | 0.32–0.38 (0.34 ± 0.02) 0.33–0.36 (0.35 ± 0.01) | 0.33 0.30–0.35 |
| HDL/HDW | 1.13–1.20 Ä 1.16 ± 0.03Å 1.04–1.16 (1.10 ± 0.03) | 1.03–1.10 (1.06 ± 0.02) | 1.13–1.20 (1.17 ± 0.02) 1.05–1.10 (1.08 ± 0.03) | 1.21 1.08–1.14 |
| SNT/HDL | 0.36–0.42 Ä 0.38 ± 0.01Å 0.38–0.42 (0.40 ± 0.01) | 0.39–0.47 (0.42 ± 0.02) | 0.36–0.40 (0.38 ± 0.02) 0.37–0.42 (0.39 ± 0.02) | 0.41 0.41–0.48 |
| IND/HDW | 0.29–0.35 Ä 0.32 ± 0.F02Å 0.28–0.33 (0.31 ± 0.02) | 0.27–0.31 (0.30 ± 0.01) | 0.28–0.35 (0.32 ± 0.03) 0.26–0.30 (0.28 ± 0.01) | 0.36 0. 34–0.38 |
| IOD/HDW | 0.24–0.27 Ä 0.25 ± 0.01Å 0.25–0.31 (0.27 ± 0.02) | 0.26–0.30 (0.28 ± 0.02) | 0.21–0.23 (0.22 ± 0.01) 0.22–0.26 (0.24 ± 0.02) | 0.21 0.23–0.30 |
| ED/SNT | 1.01–1.16 (1.07 ± 0.05) 0.74–0.92 (0.84 ± 0.05) | 0.69–0.82 (0.76 ± 0.04) | 0.90–1.03 (0.96 ± 0.05) 0.69–0.79 (0.74 ± 0.04) | 0.90 0.72–0.79 |
| ED/HDL | 0.38–0.43 Ä 0.41 ± 0.01Å 0.30–0.38 (0.33 ± 0.02) | 0.29–0.34 (0.32 ± 0.02) | 0.33–0.41 (0.36 ± 0.03) 0.27–0.31 (0.29 ± 0.01) | 0.37 0.29–0.38 |
| TED/TYD | 0.24–0.33 Ä 0.28 ± 0.03Å 0.39–0.87 (0.63 ± 0.11) | 0.58–0.88 (0.70 ± 0.10) | 0.35–0.53 (0.43 ± 0.07) 0.74–0.95 (0.84 ± 0.10) | 0.26 0.62–0.63 |
| TYD/ED | 0.50–0.61 Ä 0.55 ± 0.03Å 0.43–0.58 (0.51 ± 0.04) | 0.44–0.51 (0.47 ± 0.03) | 0.55–0.68 (0.59 ± 0.05) 0.51–0.55 (0.53 ± 0.02) | 0.62 0.47–0.52 |
| HND/SVL | 0.30–0.35 Ä 0.32 ± 0.01Å 0.25–0.33 (0.29 ± 0.02) | 0.26–0.30 (0.28 ± 0.02) | 0.29–0.34 (0.31 ± 0.02) 0.26–0.29 (0.27 ± 0.01) | 0.31 0.28–0.29 |
| TIB/SVL | 0.54–0.61 Ä 0.57 ± 0.03Å 0.53–0.67 (0.59 ± 0.04) | 0.54–0.59 (0.56 ± 0.02) | 0.56–0.68 (0.60 ± 0.04) 0.53–0.65 (0.58 ± 0.04) | 0.62 0.56–0.59 |
| FTL/SVL FPW/DPW | 0.76–0.85 Ä 0.81 ± 0.03Å 0.72–0.88 (0.80 ± 0.05) 1.16–1.52 Ä 1.30 ± 0.13Å 1.14–1.57 (1.35 ± 0.11) | 0.74–0.79 (0.76 ± 0.02) 1.35–1.80 (1.54 ± 0.15) | 0.77–0.87 (0.81 ± 0.04) 0.76–0.83 (0.75 ± 0.03) 1.79–1.94 (1.87 ± 0.06) 1.54–1.67 (1.61 ± 0.05) | 0.85 0.75–0.78 1.87 1.61–1.92 |
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