Atelopus ardila, Coloma, Luis A., Duellman, William E., C, Ana Almendáriz, Ron, Santiago R., Terán-Valdez, Andrea & Guayasamin, Juan M., 2010

Coloma, Luis A., Duellman, William E., C, Ana Almendáriz, Ron, Santiago R., Terán-Valdez, Andrea & Guayasamin, Juan M., 2010, Five new (extinct?) species of Atelopus (Anura: Bufonidae) from Andean Colombia, Ecuador, and Peru, Zootaxa 2574, pp. 1-54 : 3-12

Atelopus ardila sp. nov.

Holotype. KU 200214, adult gravid female, from 7.5 km E of Pasto, on road to Laguna La Cocha (= Lago Guamuez) (ca. 1˚ 12' N, 77˚ 13' W; 2800 m), Vereda San Fernando, Departamento Nariño, Colombia, obtained on 25 February 1984 by Patricia A. Burrowes and Benjamín del Castillo.

Paratypes. All (except ICN 3406 and specimens from Hacienda San Gerardo at Volcán Galeras) are from the road from Pasto to Laguna de la Cocha, Vereda San Fernando, Departamento Nariño, Colombia; KU 200207 –12, 200215 –18 (3 females, 7 males), with same data as holotype; KU 200213, male, from 7.5 km E of Pasto (2800 m) obtained on 25 February 1984 by Patricia A. Burrowes and B. del Castillo; KU 169267, female, from 8 km NE Pasto (3020 m), obtained on 24 September 1974 by William E. Duellman; KU 169268, female, from 8 km NE of Pasto (3050 m), obtained on 21 May 1975 by William E. Duellman; KU 169269– 335, 169341, QCAZ 24467–71 (10 females, 61 males, 1 juvenile), from 12 km E of Pasto (3050 m), obtained on 24 September 1974 by William E. Duellman, Linda Trueb, Dana T. Duellman, and John E. Simmons, and on 2 October 1974 by William E. Duellman and John E. Simmons; KU 200239, female, from 12.8 km NE of Pasto (3020 m), obtained on 25 February 1984 by Benjamín del Castillo; KU 154583, adult gravid female, from crest between Pasto and Laguna La Cocha, on road to Laguna La Cocha (3150 m), obtained on 2 August 1965 by R. E. Smalley; KU 200219 –38 (19 females, 1 male), from El Tabano, 16 km E of Pasto (3280 m), obtained on 25 February 1984 by Patricia A. Burrowes and Benjamín del Castillo; KU 169262–66, 5 females, from north shore of Laguna La Cocha, (2790 m), obtained on 2 October 1974 by John E. Simmons and Linda Trueb; ICN 3406, female, from Volcán Galeras (Hacienda San Gerardo), obtained on August 1977 by Santiago Díaz; ICN 3633, 3636–38, 4 females, from El Encano (2800 m), obtained on December 1977 by L. Gómez; ICN 450, 459, 469, 471–73, 481 (3 females, 4 males), from El Encano (2800 m), obtained on 19 January 1971 by Nancy Bastidas.

Diagnosis. (1) A species with mean SVL in adult females 46.7 mm (40.3–52.2, SD = 2.852, n = 48) and in adult males 38.6 mm (34.7–41.8, SD = 1.428, n = 76); (2) hind limbs short, tibia length/SVL 0.302–0.406 (n = 121); (3) phalangeal formula of hand 2-2-3-3, webbing absent; (4) foot webbing formula I (0+-1-)—(½-2-) II (½-1+)—(1- -2½-) III (½-2-)—(1- -3) IV (1- -3)—(1- -2-) V; (5) snout acuminate, barely protruding beyond lower jaw; (6) tympanic membrane and tympanic annulus absent; (7) dorsal surfaces of body smooth, bearing spiculae and coni mostly on sacral region, (8) black spiculae and coni (gray in preservative) present on flanks; (9) vertebral neural processes inconspicuous; (10) dorsum orange-red to black (pale creamy brown to black in preservative); (11) minute gray stippling present on dorsum of body; (12) venter orange to orange-red and dull yellow (cream in preservative); (13) gular region with a patch of black spiculae on gular-chest region in females (but see sexual dimorphism variation).

By displaying in life either orange–red, or black or a combination of both colors, Atelopus ardila sp. nov. is most similar to A. guanujo Coloma (some individuals orange-red), A. bomolochos Peters (some individuals orange-red), A. sorianoi (orange-red), A. sp. 6 (some individuals orange, Rueda-Almonacid and Acosta 2005), A. ebenoides Rivero (black), A. nanay Coloma (black), A. marinkellei Cochran and Goin (black), A. guitarraensis Osorno-Muñoz, Ardila-Robayo , and Ruiz-Carranza (some individuals black), A. pastuso sp. nov. (some individuals black), A. patazensis Venegas, Catenazzi, Siu-Ting , and Carrillo (orange and black), A. podocarpus sp. nov. (black), A. carrikeri Ruthven (black to orange and yellow), and A. ignescens (black). Nonetheless, by having a patch of black spiculae on females’ gular-chest region, A. ardila is distinct from all of them, except A. ignescens , A. carrikeri , and potentially A. sp. 6 (presence and characteristics of gular patch unknown). It differs from the latter by lacking a dark brown dorsal pattern. It differs from A. ignescens in SVL, foot and tibia length, and sacrum width (see below), and by nearly lacking posterolateral process on hyoid plate (well developed in A. ignescens ). It differs from A. carrikeri in size and by having minute gray stippling on dorsum of body (absent in A. carrikeri ); A. ardila (mean SVL of females 46.7; SD = 2.852, n = 48) is significantly smaller than A. carrikeri (mean SVL of females 57.0, SD = 4.153, n = 4; data from Coloma 1997) (SVL of females t -test, t = 6.7175, df = 50, P <0.0001).

Herein, we compare SVL and 8 morphological variables of Atelopus ardila vs A. ignescens (see materials and methods). Three axes of the PCA accounted for 58 % of the total variation ( Table 1 ). PC I mainly included tibia, foot, and radio ulna lengths, which are the highest loadings along this axis; in which most A. ardila individuals tended to have larger tibia, feet and forearms ( Fig. 2 ). PC II mainly included sacrum width and internarial distance, whereas in PC III the highest loading was SVL. Although there is some overlap of the morphometric space of A. ardila vs A. ignescens ( Fig. 3 ), significant separation occurred along two axes [overall: F(3) = 43.854, P <0.001; PC 1: F(1) = 59.183, P <0.001; PC 2: F(1) = 31.380, P = 0.009; PC 3: F(1) = 0.990, P = 0.321].

Description of holotype. ( Figs. 1 A–C, virtual animation). Head about as long as wide, HLSQ and HDWD less than one third SVL (HLSQ/SVL = 0.260, HDWD/SVL = 0.256); snout acuminate, its margin rounded in dorsal view; profile of tip of snout in lateral view curved and barely protruding to the anterior margin of jaw; no swollen gland on tip of snout; nostrils slightly protuberant, directed laterally, situated posterior to level of apex of lower jaw; canthus rostralis distinct, nearly straight from eye to nostril; loreal region concave; lips slightly flared; interorbital and occipital regions flat, smooth; eyelid flared without tubercles; postorbital crest flared, not glandular; postorbital, pretympanic and post-tympanic areas bearing coni; tympanic membrane and tympanic annulus absent; choanae small, rounded, widely separated (30.3 % of HW); tongue about twice as long as wide, its posterior half not attached to floor of mouth.

Forearm relatively short (RDUL/SVL = 0.271); palmar tubercle round; thenar, supernumerary palmar, and subarticular tubercles distinct; digital tips with round pads; thumb relatively long (THBL/HAND = 0.611), apparently having two phalanges; webbing on hands absent, fingers lacking lateral fringes; relative length of fingers II<III<V<IV. Tibia relatively short (TIBL/SVL = 0.323); fold on distal half of inner edge of tarsus absent; inner and outer metatarsal tubercles round-oval, distinct, about the same size; supernumerary plantar and subarticular tubercles conspicuous, low raised; digital pads distinct; webbing formula of foot I(0+)—(½) II (½)—(1-) III (½)—(1-) IV (0+–½)—(1) V; relative length of toes I<II<III=V<IV.

Dorsal surfaces mostly smooth, except dorsum of arms, thighs, and sacral region, which are covered by coni; flanks with abundant coni; ventral regions weakly aerolate, except venter of forearm, foot and tarsi; an inverted triangle-shaped patch of spiculae and coni on gular-chest region; cloacal opening a tube at nearly midlevel of thighs, directed posteriorly; rugose skin lateral to cloacal opening, low warts, spiculae and coni at posterior thighs.

In preservative (~70 % ethanol), dorsum of body and limbs brown with gray coni; thin brown line along middorsum from tip of snout to cloacal region; loreal region brown; flanks cream with gray coni; ventral surfaces cream, except slightly darker cream distally on fingers and toes; minute gray stippling on dorsum of body (viewed under a dissecting microscope); spiculae and coni on gular-chest region brown; proximal end of tongue lacking black pigmentation.

Measurements of holotype (mm). SVL 47.3, TIBL 15.3, FOOT 20.4, HLSQ 12.3, HDWD 12.1, ITNR 4.6, EYDM 3.6, EYNO 2.7, RDUL 12.8, HAND 12.6, THBL 7.7, SW 13.5.

Variation. Meristic variation is given in Table 2 . A juvenile female (KU 169341, SVL = 21.3 mm) possesses abundant spiculae and coni on the dorsum and flanks, but lacks spiculae on gular-chest region. It differs from adults by having cream flanks in contrast to gray dorsum.

The paratypes resemble the holotype with the following noteworthy exceptions. Females are larger than males ( Table 2 ). Males have vocal slits and keratinized nuptial pads variably extensive on the dorsal and inner surfaces of the thumb (= Finger II) and Finger III. The forelimbs are relatively long and slender only in females, but they are short with a stout muscular area in males ( Table 2 ); there are significant differences in RDUL between males and females (RDUL Student’s t -test, t = 11.4579, df = 106, P <0.0001). Males lack coni, whereas females have a greater density of spiculae and coni. Females possess an inverted triangular patch of spiculae and coni on the gular-chest region; however, three topotypic females lack spiculae there, whereas in 10 males (out of 61) from 12 km east of Pasto a patch of scattered spiculae is present on gularchest region, or spiculae are nearly absent.

Color variation in preservative (~70 % ethanol): Topotypic specimens and one from a nearby site (KU 169268) (4 females, 7 males) are similar to the holotype. Dorsal color generally varies from pale creamy brown (KU 200217) to dark brown (KU 200212); spiculae and coni vary from dark brown to black. A male (KU 200216) has dark brown marbling on the dorsum, and stripes at joints on the tarsi, feet and hands. The venter is uniform cream in all individuals. A thin brown line along middorsum varies from conspicuous (KU 200207) to barely visible (KU 200217).

Atelopus ardila

Pasto-Cocha Coloration in a series of specimens (KU 169269–335, 169341, 200239, QCAZ 24467–71) from 12.0– 12.8 km east of Pasto (11 females, 61 males, 1 juvenile) varies from plain pale creamy brown (e.g., KU 169270) to plain black (e.g., KU 169322), or to a mosaic pattern of bold black marks (e.g., KU 169303). A narrow middorsal line varies from conspicuous (e.g., KU 169318) to absent (e.g., KU 169317) in males, whereas in females it is absent. The venter is uniform cream; however, in individuals that are darker dorsally the black color partially covers plantar and toe surfaces.

In a series of specimens (KU 200219 –38) from 16 km E of Pasto (19 females, 1 male) the dorsum varies from deep black (12 specimens, e.g., KU 200235 ) to dark brown (e.g., KU 200232 ) to a mosaic pattern of black mottling (e.g., KU 200230 ). In 12 specimens the flanks are paler than the dorsum. In black individuals, the black invades the pelvic patch and the palmar and plantar surfaces, nearly entirely covering the latter, except for some of the tubercles (e.g., KU 200221).

In five females (KU 169262–66) from Laguna La Cocha the dorsum is mostly entirely black with diffuse dark brown mottling.

Color in life. ( Figs. 3 , 4 A–B). The following is based on color slides taken by WED (CT 5932–34, 5735– 42, 7211–12 of KU 169269–73, 169285–88, 200207 –08, 200217), from WED field notes of 24 September and 2 October 1974, 21 May 1975, and from Patricia A. Burrowes’ field notes of 25 February 1984. The dorsal ground color of the body and limbs varies from nearly entirely yellow and red-orange to black. Single individuals or series were described as follows: KU 200207 –18 (from 7.5 km E Pasto, 2800 m): red with or without black suffusion on snout and limbs; belly orange-red; flanks yellow with black lateral tubercles. KU 200239 (from 8 km NE Pasto, 3020 m): body yellow with black markings. KU 169267 (from 8 km NE Pasto, 3020 m): dorsum orange-red with black-tipped tubercles, webbing pale orange, venter deep orange, iris dull brown with minute gold flecks. KU 169268 (from 8 km NE Pasto, 3020 m): dorsum dull orange-red with black-tipped tubercles and brownish black head, venter orange. KU 169269–341 (from 12 km E Pasto, 3050 m): dorsal ground color varying from bright yellow-ochre to medium cadmium-orange and bright venetian red; dorsum plain with slight brownish suffusion and thin vertebral stripe and/or darker colored tubercles to spotted, mottled, or striped with sepia-umber (warm); venters plain with ground color; iris dull brown with minute bronze flecks. KU 200219 –38 (from 16 km E Pasto, 3280 m): black dorsally; venter orange to dull yellow. KU 169262–66 (from North shore Lago de la Cocha, 2790 m): dorsum black; venter red in KU 169262.

Tadpoles. Gómez Castillo (1982) described tadpoles from the Río Pasto and tributaries (2600–3000m). Tadpoles in Gosner Stages 18–29 had total lengths of 12–21 mm. Gómez Castillo (1993) described and illustrated black tadpoles from several creeks around Lago La Cocha (= Lago Guamez). He remarked on their similarity to tadpoles of A. varius and A. ignescens described by Starret (1967) and Duellman and Lynch (1969) respectively.

Distribution, ecology, natural history, and conservation status. Atelopus ardila is known only from the type locality and surrounding areas between Volcán Galeras eastward to Laguna de la Cocha in the Pasto Massif of the Andean Cordillera in Departamento Nariño in southern Colombia. These localities are in subparamo and paramo 2800–3280 m above sea level. The area of its extent of known occurrence is about 51.3 km 2 ( Fig. 5 ). At the type locality, annual mean rainfall is 1388 mm and the annual mean temperature is 12 ºC ( Hijmans et al. 2005). At the time of collection, individuals (KU 200207 –18) were crawling along a grassy irrigation ditch; others (KU 200219 –38) were crawling on black rocks in a subparamo area with a steep incline, orange soils, black loose rocks, bunch grasses, Baccharis , and lots of mosses; an amplectant pair (yellow male, red female) also was found (P.A. Burrowes field notes, 25 February 1984). KU 169268 was under a yucca stump by day in subparamo with dense woody bushes and bamboo (WED field notes, 21 May 1975). KU 169285– 341 were crawling on the ground (open, mossy, grassy) after a rain; some were under rocks by day. Others (KU 169262–66) were crawling across a road by day (WED field notes, 2 October 1974). A female (KU 169267) was under a rock by day (temperature: 7.5–13.0˚C; no rainfall) in a subparamo area with dense woody bushes and bamboo. Other individuals (KU 169269–84) were in a rocky stream, where most were under rocks at the edge of the streambed; a few individuals were crawling amidst rocks on the ground by day (WED field notes, 24 September 1974).

Del Castillo Ch. (1982) and Gómez Castillo (1982; 1993) provided data on the reproductive biology of individuals from several localities and at ex situ conditions. Del Castillo Ch. (1982) induced oviposition using hormones, described behavior, physiology and early development of individuals and eggs from El Encano (locality stated in Gómez Castillo 1993:169). Gómez Castillo (1982) also described courtship, mating and reproduction of individuals from El Encano, San Fernando, and 11 km N of Pasto. Gómez Castillo (1993) provided data on the reproductive biology of individuals from Laguna La Cocha and vicinity and reported additional descriptions of induced oviposition. Three types of calls of individuals from several populations from Departamento Nariño (presumably a mixture of A. ardila and A. pastuso ) were briefly described by Gómez Castillo (1982:61), two of them as peeps and another as a trill. Before axillary amplexus, it was observed that the male walks or jumps over the female from behind or from the side. Two males embracing a female were observed after exposing several males to a female. Amplectant pairs were found throughout the year at El Encano, San Fernando, Veredas Motilón, and Santa Rosa ( Gómez Castillo 1982; 1993). At El Encano a pair remained in amplexus for at least 22 days. Amplexus among pairs maintained in captivity lasted more than 2– 3 months; in one of these cases a pair was in amplexus for more than 74 days, and the male remained in the amplectant position for two days after the female died. Amplectant pairs were observed on two occasions under water (about 25 cm below the surface)—one in October 1978 at El Encano, and another in May 1979 at San Fernando. Another amplectant pair was observed under water at Quebrada Funduyacu ( Gómez Castillo 1993). Tadpoles have been found throughout the year, but they are less abundant in the summer from June to August. Tadpoles were found in clear, non-contaminated waters in mountain creeks, Laguna La Cocha, and the Río Pasto and tributaries, but they were not observed near human habitations in urban areas ( Gómez Castillo 1982; 1993).

Gómez C. and Ramos O. (1982) and Gómez and Ramos O. (1982) analyzed the stomach content of 30 individuals of five populations (San Fernando, 2600 m; El Encano, 2850 m; Daza, 2800 m, Volcán Galeras, 3150 m; Colón-Mocoa, 2200 m) from Municipio de Pasto and Intendencia de Putumayo, presumably belonging to Atelopus ardila (under the name A. ignescens ). No voucher specimens were indicated, but we assume that the specimens were A. ardila based on the localities, measurements and color description of the animals provided. They report the following diet: Diptera 38 %, Coleoptera 33.2 %, Hymenoptera 25.4 %, Lepidoptera and Homoptera in smaller proportions. Chrysomelidae (Coleoptera) were the most abundant prey (26.5 %), followed by Drosophilidae (Diptera) , Pteromalidae (Hymenoptera) , Braconidae (Hymenoptera) , Anthomyiidae (Diptera) , Muscidae (Diptera) , and Tephritidae (Diptera) .

Atelopus ardila is considered to be Critically Endangered (Possibly Extinct) (A2ace, IUCN Red List categories and criteria). The species is tagged as Possibly Extinct until further surveys confirm otherwise. The population has declined dramatically (more than 80 %) in the last two decades probably because of climate change and the impact of pathogens, which have affected many other montane species of Atelopus . No chytrid fungus reports or climate data for its area of distribution are available. The closest report of occurrence of the chytrid fungus is from Provincia Carchi near the Ecuador-Colombia border ( Ron and Merino 2000). The risk factor of potential threat caused by the chytrid for anuran amphibian species calculated by Rödder et al. (2009: Fig. 2 C) is high at the area of its distribution.

Although this species previously was abundant at several localities ( Gómez Castillo, 1982:56), no individuals have been recorded (with voucher specimens) since June 1989 when one individual (IAvH 4767) was collected in Municipio Consacá at the surroundings of Volcán Galeras ( Ardila-Robayo and Maldonado-Silva 2004). Intensive search efforts since the mid-1990s ( Cepeda-Quilindo and Rueda-Almonacid 2005) revealed no individuals. Visual records indicate the presence of a few individuals in 1992 at Colón-La Rejoya in the Sibundoy Valley (Mueses-Cisneros, pers. com. 2009), and unconfirmed visual records by local people were made in 1994 and 2002 ( Mueses-Cisneros, 2005). Intensive searches between December 2000 and February 2004 at 11 historical sites by Mueses-Cisneros (2005) resulted in no individuals.

Etymology. The specific name is a noun in apposition meant as patronym for Cristina Ardila-Robayo , a Colombian herpetologist of Instituto de Ciencias Naturales of the Universidad Nacional de Colombia. We recognize her dedication and valuable contributions to the Herpetology of Colombia. Part of her research efforts resulted in the description of 28 new species of amphibians from Colombia, including 10 Atelopus .

Remarks. Populations herein assigned to Atelopus ardila were excluded from the nominal taxon Atelopus ignescens by Gray (1983), Coloma et al. (2000), and Ron et al. (2003). Nonetheless, these populations were included temporarily under Atelopus ignescens (e.g., cited as Atelopus ignescens complex fide Gray 1983) by Lötters (1996); they were cited as Atelopus “complejo ignescens ” by Ardila-Robayo and Maldonado-Silva (2004) and Cepeda Quilindo and Rueda Almonacid (2005). We provide morphological evidence that justifies the recognition of A. ardila as a different species. Nonetheless, as stated by Coloma et al. (2000), presumably A. ardila is the sister taxon of A. ignescens . Both species share a gular region having the unique presence of a patch of black spiculae in females. However, a phylogenetic analysis of the genus based on genetic markers is needed to enhance our current taxonomic understanding of these species.

Figures

FIGURE 1. — Holotype of Atelopus ardila (KU 200214): (A) adult female, SVL = 47.3 mm; (B) ventral view of right hand, enlarged 3.5 times of A; (C) ventral view of right foot; enlarged 3.2 times of A.

FIGURE 2. — Axes PC I and PC II from Principal Components Analysis based on eight size-corrected morphological variables for populations of Atelopus ardila and A. ignescens from Colombia and Ecuador. Circles = A. ardila; X = A. ignescens.

FIGURE 3. — Intrapopulation color variation of Atelopus ardila from 12 km E of Pasto, Departamento Nariño, Colombia: (A) KU 169270; (B) KU 169272; (C) KU 169271; (D) KU 169273 (Photos by WED).

FIGURE 4. — (A, B) Atelopus ardila from 12 km E of Pasto, Departamento Nariño, Colombia (KU 169287 – 88, photos: WED); (C – D) Atelopus pastuso from Tulcán and 26, 6 km W Tulcán, Provincia Carchi, Ecuador (MNHG 2258.68, QCAZ 1911, photos: LAC); (E – F) Atelopus pastuso from Tulcán and Cerro Cotacachi (MNHG 2258.65, QCAZ 15021 adult male, photos: LAC, Eugenia M. del Pino, respectively); (G) Atelopus pastuso from Hacienda La Esperanza (EPN 3244 adult female, photo: AA); (H) Atelopus angelito from NE Maldonado (KU 3244 adult female, photo: John D. Lynch).

FIGURE 5. — Map of southern Colombia and northern Ecuador showing localities of Atelopus angelito, A. ardila, A. gigas, A. orcesi, A. pastuso and a population of hypothetical hybrids between A. pastuso and A. ignescens. Symbols for A. ardila indicate one or several close localities.

Tables

TABLE 1. Eigenvalues, character loading and percentage of explained variance for principal components (PC) I – III for eight morphometric variables of Atelopus ardila vs A. ignescens. Abbreviations are: SVL = snout – vent length; TIBL = tibia length; FOOT = foot length; HLSQ = head length; HDWD = head width; EYNO = eye – nostril distance; ITNA = Internarial distance; RDUL = radio-ulna length; SW = sacrum width.

Variable	PC I	PC II	PC III
Log SVL	0.00000	-0.00000	1.00000
Residuals TIBL	0.50815	0.09575	-0.00000
Residuals HLSQ	0.43768	-0.26860	0.00000
Residuals HDWD	0.31595	-0.05531	0.00000
Residuals ITNA	0.04993	0.55033	-0.00000
Residuals THBL	0.09394	0.45684	0.00000
Residuals RDUL	0.45037	-0.00874	-0.00000
Residuals FOOT	0.47875	-0.06910	-0.00000
Residuals SW	0.08388	0.63183	-0.00000
Percent %	29.933	16.519	11.111

TABLE 2. Meristic variation of Atelopus ardila. Mean ± one SD, and range are given. Abbreviations are: SVL = snout – vent length; TIBL = tibia length; FOOT = foot length; HLSQ = head length; HDWD = head width; EYDM = eye diameter; EYNO = eye – nostril distance; ITNA = Internarial distance; RDUL = radio-ulna length; THBL = thumb length; SW = sacrum width. All measurements are in mm.

	Males (n = 76)	Females (n = 48)
SVL	38.6 ± 1.43 (34.7–41.8)	46.52 ± 2.85 (40.3–52.2)
TIBL	14.41 ± 0.58 (13.2–16) (n = 73)	15.91 ± 0.81 (13.6–17.8)
FOOT	16.68 ± 0.91 (14–18.7) (n = 71)	19.9 ± 1.34 (16–22) (n = 40)
HLSQ	11.79 ± 0.52 (10.8–13.5) (n = 73)	12.68 ± 0.62 (11.1–14.1)
HDWD	11.44 ± 0.47 (10.4–12.6) (n = 73)	12.34 ± 0.63 (10.6–13.7)
ITNA	4.1 ± 0.27 (3.5–4.8) (n = 72)	4.43 ± 0.3 (3.6–5.1) (n = 40)
RDUL	11.82 ± 0.56 (10.2–13.1) (n = 70)	13.31 ± 0.78 (12–16.4) (n = 38)
THBL	6.72 ± 0.42 (5.9–8.1) (n = 66)	8.48 ± 0.42 (9.7–7.3) (n = 40)
SW	10.48 ± 0.62 (9.2–12.1) (n = 70)	12.66 ± 1.19 (9–15.5) (n = 44)
RDUL/SVL	(0.28–0.34)(n = 70)	(0.26–0.34) (n = 38)
TIBL/SVL	(0.33–0.41) (n = 73)	(0.30–0.38)
HLSQ/SVL	(0.28–0.34) (n = 73)	(0.24–0.32)
HDWD/SVL	(0.27–0.33) (n = 73)	(0.23–0.29)