Liolaemus pacha, Heredia, Viviana Juárez, Robles, Cecilia & Halloy, Monique, 2013
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publication ID |
https://doi.org/ 10.11646/zootaxa.3681.5.2 |
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publication LSID |
lsid:zoobank.org:pub:A3546543-6904-4971-9E2A-A6500F9A3FAE |
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DOI |
https://doi.org/10.5281/zenodo.5620043 |
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persistent identifier |
https://treatment.plazi.org/id/8E08B516-7715-FFC7-FF17-FB3FFEE5F854 |
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treatment provided by |
Plazi |
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scientific name |
Liolaemus pacha |
| status |
sp. nov. |
Liolaemus pacha sp. nov.
( Figs. 2 to 4).
1843. Liolaemus darwinii, Bell, T., Smith, Elder & Co., London, pp. 1–55.
1993. Liolaemus quilmes, Etheridge, R., Boll. Mus. Reg. di Sc. Naturali, Torino, 11: 137–199.
1993. Liolaemus darwinii (partim), Cei, J. M., Mus. Reg. di Sc. Naturali, Torino, Monografie XIV, 949 pp.
1996. Liolaemus quilmes, Halloy, M., Bull. of the Maryland Herp. Soc., 32: 43–57.
1998. Liolaemus quilmes, Halloy, M., Etheridge, R. and Burghardt, G., Herpetological Monographs, 12: 1–37.
2002. Liolaemus quilmes, Halloy, M. and Robles, C., Bull. of the Maryland Herp. Soc., 38: 118–129.
2003. Liolaemus quilmes, Halloy, M. and Robles, C., Cuad. Herp., 17: 67–73.
2004. Liolaemus quilmes, Halloy, M., Acta Zool. Lilloana, 48 (1–2): 103–121.
2004. Liolaemus quilmes, Martins E. P., Labra, A., Halloy, M. and Tolman Thompson J., Animal Behaviour, 68 (3): 453–463.
2005. Liolaemus quilmes (partim), Abdala, C. S., Rev. Esp. de Herp., 19: 5–17.
2006. Liolaemus quilmes (partim), Abdala, C. S. and Lobo, F., Cuad. Herp., 19 (2): 3–18.
2006. Liolaemus quilmes (partim), Abdala, C. S. and Díaz Gómez, J., Zootaxa, 1317: 21–33.
2006. Liolaemus quilmes (partim), Avila, L. J., Morando, M. and Sites, J. W. Jr., Biol. Journal of the Linnean Society, 89: 241– 275.
2006. Liolaemus quilmes, Halloy, M., Herp. Review, 37: 88–89.
2006. Liolaemus quilmes, Halloy, M. and Castillo, M., Herp. Nat. History, 9 (2): 127–133.
2006. Liolaemus quilmes, Halloy, M., Robles, C. and Cuezzo, F., Rev. Esp. Herp., 20: 47–56.
2007. Liolaemus quilmes (partim), Abdala, C. S., Zootaxa, 1538: 1–84.
2007. Liolaemus quilmes, Halloy, M., Guerra, C. and Robles, C., Bull. of the Maryland Herp. Soc., 43: 110–118.
2008. Liolaemus quilmes, Robles, C. and Halloy, M., Cuad. de Herpetología, 22: 73–79.
2009. Liolaemus quilmes, Robles, C. and Halloy, M., South Am. Journal of Herp., 4: 253–258.
2009 a. Liolaemus quilmes, Salica, M. J. and Halloy, M., Acta Zool. Lilloana, 53: 89–96.
2009 b. Liolaemus quilmes, Salica, M. J. and Halloy, M., Rev. Esp. Herp., 23: 141–149.
2010. Liolaemus quilmes, Halloy, M., Robles, C. and Salica, M. J. Anales de la Academia Nac. Ciencias, Buenos Aires, Argentina, 44: 3–16.
2010. Liolaemus quilmes, Robles, C. and Halloy, M., Herpetological Journal, 20: 243–248.
2011. Liolaemus quilmes, Salica, M. J. and Halloy, M., Herp. Review, 42 (4): 606.
2012. Liolaemus quilmes, Halloy, M., Herpetological Journal, 22 (4): 267–270.
2012. Liolaemus quilmes, Robles, C. and Halloy, M., Salamandra, 48 (2): 115–121.
2013. Liolaemus quilmes, Halloy, M., Robles, C., Salica, M. J., Semhan, R., Juárez Heredia, V. and Vicente, N., Cuadernos de Herpetología, 27 (1): 15–26.
Holotype. Fundación Miguel Lillo ( FML) 02448/1. Km 98, Ruta Provincial 307 (Los Cardones), Tafí del Valle Department, Tucumán Province, 26°40’1.5” S 65°49’5.1” W, datum: WGS84, 2725 m, collector, O. Pagaburo, 21/ 04/1999.
Paratypes. FML 2448/3–4/6–9 Km 98, Ruta Provincial 307, Amaicha del Valle, Tafí del Valle Department, Tucumán Province. O. Pagaburo, Col. 21/04/1999.
Etymology. Pacha comes from the Aimara and Quechua indigenous languages from northwestern Argentina. It means “earth” and by extension “world” or “cosmos”. In this part of the country, social gatherings called Pachamama are celebrated every year in the autumn to thank and bless the earth and its products.
Diagnosis. Liolaemus pacha sp. nov., belongs to the L. boulengeri group because it has a patch of enlarged scales on the posterior region of the thigh (Etheridge 1995; Abdala et al. 2006). Within this group, it belongs to the L. laurenti group (Abdala 2007) because its posterior teeth have straight edged crowns and sexual dichromatism is evident. Within the L. laurenti group, Liolaemus pacha sp nov. is a member of the L. darwinii group (Abdala 2007) or the darwinii complex (Etheridge 1993) because it has a black line which crosses the eye vertically (except L. chacoensis ) and pre-scapular spots in both males and females (Abdala 2007). Within the L. darwinii group (Abdala 2007), L. pacha differs from the members of the L. ornatus group ( L. albiceps , L. calchaqui , L. crepuscularis , L. irregularis , L. lavillai , and L. ornatus ) because it has an oviparous reproductive mode. It also differs from L. albiceps and L. irregularis because it is smaller (max SVL 67.9 mm vs 82.5 and 86.1 mm, respectively). Liolaemus pacha differs from L. abaucan and L. uspallatensis in having imbricate and keeled dorsal scales, whereas in those two species the dorsal scales are smooth or weakly keeled. Liolaemus chacoensis exhibits keeled temporal scales and mucronate dorsal scales, absent in L. pacha . The presence of a large prescapular spot distinguishes L. pacha from L. abaucan , L. darwinii and L. koslowskyi , which have large and evident postscapular spots. It differs from L.
cinereus in having a black stripe in the lateral field and in lacking melanism on the throat. Liolaemus darwinii , L. grosseorum , L. laurenti , L. montanezi and L. olongasta have a black antehumeral arch, absent in L. pacha .
Liolaemus pacha sp. nov. differs from L. quilmes mainly because of its size (see below). It has a larger prescapular spot ( Fig. 5 View FIGURE 5 , left) than L. quilmes ( Fig. 6 View FIGURE 6 , left) and numerous light blue scales on the sides of the body and tail ( Fig. 2). It has a Y-shaped mark on the snout ( Fig. 7 View FIGURE 7 ), reduced in L. quilmes ( Table 1 View TABLE 1 ). In L. pacha , scales on dorsum are uniformly colored ( Fig. 8) whereas in L. quilmes scales on dorsum are irregularly variegated with diffuse spots ( Fig. 9). Both species present alternating light and dark spots on supra and infralabials, more diffuse on infralabials in L. pacha sp. nov. ( Fig. 10 View FIGURE 10 , top), well marked in L. quilmes ( Fig. 10 View FIGURE 10 , bottom). Anterior limbs with lightly visible bands and posterior limbs without these ( Fig. 2 and 5 View FIGURE 5 , left), whereas in L. quilmes both anterior and posterior limbs with visible bands ( Fig. 6 View FIGURE 6 , left). The new species presents yellow dorsolateral spotted bands ( Fig. 2), different from L. quilmes which are continuous. Liolaemus pacha sp. nov. has ventrally scattered spots in the jaw area, light yellow on the throat and thigh areas ( Fig. 3 and 5 View FIGURE 5 , right), in L. quilmes throat with spots that reach to the neck, ventral area white ( Fig. 6 View FIGURE 6 , right, Table 1 View TABLE 1 ).
Moreover, among its distinctive characteristics, L. pacha sp. nov. differs significantly from 3 populations of L. quilmes in 10 characters ( Table 2 View TABLE 2 , P <0.05, shown with asterisk): greater snout-vent-lengths in both males and females; higher, wider and longer head in males; larger and wider male torso; longer hand; more scales on average around the body; more gular scales. The new species differs in four other characters ( Table 2 View TABLE 2 , P <0.05, shown without asterisk) from one or two of these populations: in two cases, it differed from El Tio y Las Ruinas de Quilmes but not from Animaná (ventral scales, length of female torso); in one case, it differed from Animaná and El Tio but not from Ruinas de Quilmes (length of humerus); and lastly, it differed from Animaná but not from El Tío and Ruinas de Quilmes (head width of females).
Description of the holotype. Adult male. SVL 63.1 mm. Trunk length 33.8 mm. Head longer (15.1 mm) than wide (11.5 mm). Head height 7.8 mm. Eye diameter 3.8 mm. Interorbital distance 8.6 mm. Orbit–auditory meatus distance 5.8 mm. Auditory meatus 2.5 mm high, 1.3 mm wide. Orbit–commissure of mouth distance 3.3 mm. Internares distance 2.6 mm. Subocular scale length 3.1 mm. Femur length 13.5 mm, tibia 10.4 mm, and foot 17.3 mm. Humerus length 9.4 mm. Forearm length 7.6 and hand 9.1 mm. Tail length 95.5 mm. Length of the pigal region 6.4 mm and cloaca width 8.5 mm.
Dorsal surface of the head smooth, with 17 scales. Rostral wider than long, bordered by six scales. Mental larger than rostral, trapezoidal, bordered by four scales. Nasal not in contact with rostral. Nasal surrounded by six scales, separated from canthal by one scale. Five scales between frontal and supercilliaries. Five scales between frontal and rostral. Frontal not divided. Two postrostrals. Interparietal smaller than parietals, in contact with eight scales. Orbital semicircles complete. Preocular separated from lorilabial row by one scale. Three scales in anterior margin of auditory meatus. Fifteen smooth temporals. Four lorilabials in contact with subocular. Seven supralabials, none in contact with subocular. Seven supraoculars. Eight lorilabials. Six infralabials, second ventrally in contact with two scales. Seventy scales around midbody. Seventy-five round, slightly imbricated, slightly keeled dorsals from occiput to hind limbs. Twenty-four rows of scales in dorsum. Scales of flank same size and shape as dorsals. Ninety-three ventrals same size as dorsals, flat, imbricate. Thirty smooth, weakly imbricated gulars. Four precloacal pores. Antehumeral scales flat, larger than or equal in size to dorsals. Postauricular, rectal, and longitudinal folds present. Scales on the longitudinal fold granular and smooth. Fourth finger with 17 subdigital lamellae; fourth toe with 25. Infracarpals flat, imbricate, trifid; infratarsals flat, imbricate, trifid. Without gular fold. With 36 scales on the neck, from the posterior border of the ear to the shoulder, through the longitudinal fold. With femoral parch. Dorsal scales of the tail without mucro and with keel, ventrals smooth.
Color in alcohol. Dorsum and sides of the head brown colored with numerous scales and small spots dark brown. Subocular light brown colored with dark spots. Lorilabials and supralabials alternate between light and dark color, infralabials with alternating diffuse spots ( Fig. 10 View FIGURE 10 top). Snout with diffuse inverted Y shaped mark ( Fig. 7 View FIGURE 7 ). Body dorsally with 8 series of paravertebral spots, subquadrangular shaped, with white-colored, spotted, dorsolateral bands. Paravertebral spots with white posterior elongated spots. Dorsum and sides of the body grey colored to light brown, scattered irregularly with numerous white colored scales ( Fig. 5 View FIGURE 5 left). Scales on dorsum, uniformly colored ( Fig. 8). With white-colored spotted vertebral line. Presence of prescapular spot. With many scattered light blue spots on the sides of the body ( Fig. 2). Anterior and posterior limbs light brown colored, with white spots distributed irregularly. Femur with obvious light blue spots. Tail grey to brown-colored, with elongated spots and numerous light blue spots, mainly distributed at the base. Ventrally the mandibular region is white colored with diffuse spots from infralabials ( Fig. 3 and 5 View FIGURE 5 , right). Ventrum, anterior and posterior limbs, cloaca and tail completely white.
Color in life ( Table 1 View TABLE 1 ). Dorsum and sides of the head light chestnut brown, with dark and/or black chestnut brown scales and spots. Supralabials with alternating dark and light chestnut brown spots, infralabials with light and dark spots diffuse. Chestnut-brown inverted Y-mark, on the tip of the snout ( Fig. 7 View FIGURE 7 ). Body with 8 pairs of paravertebral spots of subquadrangular shape, black colored, with elongated posterior white spot ( Fig. 5 View FIGURE 5 , left). With prescapular black spot. Lateral region of the body beige reddish with numerous white and light blue spots ( Fig. 2). With vertebral line and dorsolateral bands discontinuous yellow colored. With diffuse subquadrandular dorsolateral spots. Neck with yellow band from above the ear to the anterior limb. Scales on the dorsum with a homogeneous colored pattern. Limbs and tail chestnut-brown colored, scattered with white, yellow light blue and dark brown scales. Arm and forearm with two diffuse rings ( Figs. 2 and 5 View FIGURE 5 , left). Posterior members without rings. Tail with numerous light blue scales. Ventrally, the mandibular area, anterior limbs, ventrum, cloaca and tail are white ( Fig. 3 and 5 View FIGURE 5 , right). In the mandibular region, with disseminated spots from the infralabials to the anterior region of the neck ( Fig. 3 and 5 View FIGURE 5 , right). Gular region, sides and femur with diffuse yellow spots ( Fig. 3). Obvious orange-colored precloacal pores.
Variation ( Table 2 View TABLE 2 ). Based on 25 specimens (15 males, 10 females) (Appendix 1). Head longer (10.9–15.1 mm; mean = 13.3 mm) than wide (9.1–13.4 mm; mean = 11.4 mm). Head height 6.4–9.9 mm (mean = 7.9 mm). Neck narrower than head. SVL 53.6–67.9 mm (mean = 61.2 mm), averaging 1.7 times longer than trunk. Tail length 63.9–103.8 mm (mean = 85.1 mm). Humerus length 5.9–9.7 mm (mean = 8.1 mm). Dorsal surface of head smooth, with 12–20 scales. One row of lorilabials, same size as supralabials. Seven to ten lorilabials. Five to eight supralabials. Frontal not divided. Interparietal always smaller than parietals, surrounded by 5–7 scales. Nasal surrounded by 6–9 scales. Four to seven infralabials. Mental in contact with four scales. Ten to 13 smooth round temporals. Longitudinal, postauricular, and antehumeral folds present. Horizontal fold Y-shaped between shoulder and auditory meatus. Scales around midbody 46–64 (mean = 58.5). Gulars 26–44 (mean = 31.7). Dorsal scales between occiput and hind limbs 62–77 (mean = 69.0). Ventrals 79–97 (mean = 90.2). Males with 4–7 (mean = 5.5) precloacal pores. Females with 1–2 (mean = 1.7) precloacal pores. Dorsals flat, slightly imbricate, slightly keeled. Twelve to 20 infradigital lamellae on fourth finger and 21–28 on fourth toe.
CHARACTER / SPECIES L. pacha sp. nov. Animaná El Tio Ruinas P Variation in color pattern. With clear sexual dichromatism. Background color of the head may vary, from grey to light chestnut-brown to dark. Dorsum of the head presents occasionally a greater number of irregular black spots. Laterally the head may present conspicuous yellow spots. On the sides of the head, alternating clear and dark spots, diffuse on the infralabials and well marked on the supralabials ( Fig. 10 View FIGURE 10 , top), stand out. The inverted Yshaped mark on the snout diffuse or absent ( Fig. 7 View FIGURE 7 ). The color of the body may vary between brownish and reddish tones. Without a black antehumeral arc. With discontinuous or absent vertebral line. Paravertebral spots shaped as horseshoe with posterior outline or subquadrangulars, black colored, with posterior white spot elongated or circular shaped. Dorsal body scales with a homogeneous pattern ( Fig. 8). With diffuse dorsolateral subquadrangular spots, smaller than paravertebrals and sometimes absent. White line discontinuous on the sides of the body. Conspicuous black colored prescapular spot present. Limbs vary from grey to brown. Arm and forearm with two diffuse or absent rings ( Fig. 5 View FIGURE 5 , left). The tail is lighter colored than the dorsal region of the body, with numerous light blue scales. With discontinuous spots.
Ventrally white colored occasionally the mandibular, gular and femur regions, light yellow colored. Jaw with diffuse or absent spots. Precloacal pores conspicuous orange color. Head and body of females varying from grey to dark brown. Inverted Y-spot on snout diffuse. Paravertebral lines light ochre which unite on the tail. Paravertebral and subquadrandular dorsolateral spots black with thin posterior white spot, occasionally very diffuse. Lateral white line diffuse. Laterally body with white and light blue rare or absent spots. Anterior limbs with diffuse or absent rings. Posterior limbs without rings. Laterally the neck varying between yellow to intense orange. Ventrally white colored. Precloacal pores rare, barely visible.
Distribution. The new species is known from the type locality Km 98, Ruta Provincial 307 (Los Cardones), Tafí del Valle Department, Tucumán Province (26°40’1.5” S 65°49’5.1” W, datum: WGS84; 2725 m).
Natural history. Liolaemus pacha is found on rocky to sandy subtrates. The type locality is characterized by firm substrate, large rocks, scattered shrubs and tall cacti (Thrichocereus pasacana, Cactaceae ). It corresponds to the Monte and Prepuna ecoregions (Cabrera & Willink 1980). They are mainly insectivorous occasionally eating small plant parts depending on the season and availability (Halloy et al. 2006). It is an oviparous species (Ramirez Pinilla 1992). Males defend territories which generally overlap with two to three females (Halloy 1996; Halloy & Robles 2002; Robles & Halloy 2009; Robles 2010). Both males and females use visual displays, such as headbobs, forelimb waves, and tail movement, during encounters as well as during courtship (Halloy 1996; Martins et al. 2004; Halloy & Castillo 2006; Halloy 2012).
It is a dimorphic species (Etheridge 1993; Halloy 1996; this study), males being slightly larger and more colorful than females. Females present during the reproductive season nuptial coloration on the side of the neck, going from yellowish to intense orange (Halloy et al. 2007; Salica 2008; Salica & Halloy 2009b). Males emerge from hibernation in September whereas females emerge one to two months later (Halloy & Robles 2003). Matings occur end of October, beginning of November. Neonates are observed by the end of December and in January (Ramírez Pinilla 1992; pers. obs.). Males and females remain active through March and April when they return to hibernation.
TABLE 1. Markings and coloration patterns in Liolaemus pacha sp. nov. (Km 98, Ruta Provincial 307, “ Los Cardones ”, Tafí del Valle Department, Tucumán Province) and in three populations of L. quilmes combined (Animaná, Salta [Holotype / Paratypes]; El Tío and Ruinas de Quilmes, Tucumán; Appendix I).
| CHARACTER / SPECIES | L. pacha sp. nov. | L. quilmes |
|---|---|---|
| Y-shaped mark on snout | Present, conspicuous | Present, diffuse |
| Supralabials and infralabials alternately spotted | Diffuse on infralabials | Yes |
| Rings on the anterior limb | Present, diffuse | Present |
| Rings on the posterior limb | Absent | Present |
| Prescapular spots | Present, large | Present |
| Shape of paravertebral spots | Large subquadrangular | Subquadrangular |
| White spots posterior to paravertebrals | Present (circle) | Present, circular and/or surrounding paravertebrals |
| Dorsolateral bands | Dotted | Continuous |
| Dorsolateral spots | Present, diffuse | Present, conspicuous |
| Spots on the sides of the body | Numerous white spots | White spots distributed unevenly |
| Light blue scales on body | Numerous on the sides of the body | Present, distributed irregularly |
| Light blue scales on sides of tail | Present, abundant | Present |
| Ventral pattern of jaw | Inmaculate or with diffuse dark dots dispersed irregularly | Dots that can reach neck |
| Color of ventrum | White or Light yellow | White |
| Sexual dichromatism | Present, more conspicuous | Present |
TABLE 2. Ranges and averages (in parentheses) of various character states in Liolaemus pacha sp. nov. (Km 98 [Los Cardones], Ruta Provincial 307, Tafí del Valle Department, Tucumán; 15 males, 10 females) and in three populations of L. quilmes (Animaná, Salta [Holotype / Paratypes], 12 males, 12 females; El Tío, Tucumán, 8 males, 3 females; and Ruinas de Quilmes, Tucumán, 10 males, 5 females; Appendix I). P-values for Kruskal-Wallis one-way analysis of variance are provided in the last column. An asterisk is shown when L. pacha sp. nov. differed significantly from all three populations of L. quilmes (more details in text).
| scales around the body | 46–70 (58.9) | 42–60 (53.9) | 50–63 (55.8) | 52–59 (55.0) | 0.0001 * |
|---|---|---|---|---|---|
| ventral scales | 79–97 (90.3) | 74–101 (82.3) | 81–98 (87.6) | 78–90 (83.8) | 0.0001 |
| dorsal scales | 62–77 (69.3) | 57–75 (67.7) | 62–74 (68.4) | 60–72 (66.2) | 0.185 |
| SVL in males | 59.0–67.9 (63.3) | 49.9–62.1 (56.0) | 47.6–57.3 (54.4) | 49.0–57.9 (54.2) | 0.0001 * |
| SVL in females | 53.6–62.9 (58.2) | 48.9–57.6 (53.2) | 47.0–52.9 (49.1) | 45.8–54.1 (50.9) | 0.003 * |
| gular scales | 26–44 (31.7) | 23–30 (26.4) | 24–31 (27.7) | 22–30 (26.1) | 0.0001 * |
| N° scales neck | 29–49 (35.7) | 19–38 (32.9) | 28–37 (32.7) | 23–38 (31.7) | 0.09 |
| length of male torso | 30.6–39.8 (35.2) | 26.4–34.0 (31.0) | 26.3–31.9 (30.2) | 25.1–32.6 (29.3) | 0.0001 * |
| length of female torso | 30.3–36.9 (34.3) | 27.7–35.0 (31.8) | 28.0–31.3 (29.3) | 26.6–32.8 (30.6) | 0.019 |
| length/width male head | 1.0–1.3 (1.1) | 1.1–1.2 (1.1) | 1.1–1.3 (1.2) | 1.1–1.2 (1.2) | 0.83 |
| hand length | 5.5–10.6 (8.5) | 6.8–9.4 (7.9) | 7.3–8.1 (8.0) | 7.4–9 (7.3) | 0.0011 * |
| length humerus | 5.9–9.7 (8.1) | 5.7–8.5 (7.4) | 6.5–8.5 (7.4) | 5.8–8.7 (7.6) | 0.017 |
| length of tail | 63.9–103.8 (85.1) | 44.7–80.3 (69.8) | 56.6–85.6 (72.7) | 60.5–90.3 (76.5) | 0.11 |
| head width of males | 10.3–13.4 (12.0) | 9.6–12.3 (10.8) | 9.1–11.4 (10.3) | 9.1–11.2 (10.5) | 0.0001 * |
| head width of females | 10.2–11.7 (10.4) | 8.2–9.8 (9.1) | 8.7–10.2 (9.2) | 9.0–9.7 (9.3) | 0.0004 |
| head height | 6.4–9.9 (7.9) | 5.5–8.1 (6.8) | 5.8–8.0 (6.8) | 6.0–7.9 (6.8) | 0.0002 * |
| torso width of males | 12.7–18.0 (15.3) | 12.0–16.3 (13.9) | 12.3–15.8 (14.3) | 11.7–16.0 (13.9) | 0.0001 * |
| body/tail | 1–1.5 (1.4) | 0.7–1.5 (1.3) | 1.2–1.5 (1.4) | 1.1–1.6 (1.4) | 0.54 |
| Humerus/radius | 0.9–1.4 (1.4) | 1.0–1.4 (1.2) | 1.1–1.2 (1.1) | 1.0–1.3 (1.2) | 0.785 |
| head length | 12.4–15.0 (14.0) | 11.4–14.1 (12.8) | 10.9–13.7 (14.4) | 11.6–13.1 (12.6) | 0.0002 * |
| FML |
Fundacion Miguel Lillo |
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