Hypostomus khimaera Tencatt, Zawadzki & Froehlich, 2014

Hypostomus khimaera Tencatt, Zawadzki & Froehlich, 2014 View in CoL

( Figs. 1–8, 11; Tab. 1)

Hypostomus cochliodon (non Kner, 1854). ―Armbruster, 2003:19, 21–25 (partim; identification key; redescription).

Hypostomus khimaera . ―Tencatt et al., 2014:597–600 (original description; type-locality: córrego Salobo (= Salobro)). ― Oliveira et al., 2020:493, 496 (diagnosed from H. labyrinthus ; listed as comparative material). ―Carvalho et al., 2019:50, 52 (brief description; photo in life). ―Lopes et al., 2024:594–99 (new record; diagnosis; photos in life; geographic distribution).

Diagnosis. Hypostomus khimaera is distinguished from its congeners, except for the species from the H. cochliodon group sensu Armbruster (2003), by of the following characteristics: (I) absence of a notch between the metapterygoid and the hyomandible ( Fig. 2A) (vs. presence; see Armbruster, 2003:13, fig. 2A), (II) acutely angled dentaries, up to about 90° (vs. obtusely angled dentaries, generally forming a clearly angled greater than 90°), (III) teeth with medial cusp typically spatulate (vs. only villiform teeth); and (IV) exposed portion of opercle relatively small and completely bordered by a thick layer of skin ( Fig. 3) (vs. exposed portion of opercle relatively large and not bordered by a thick layer of skin; see Armbruster, 2003:13, fig. 2A). Hypostomus khimaera is distinguished from species in the H. cochliodon group, except H. basilisko and H. soniae Hollanda Carvalho & Weber, 2005 , by typically presenting a dark stripe along the median portion of the flank; diffuse in some specimens (vs. stripe absent); from H. basilisko and H. soniae , it differs by having dark spots on at least some region of the body (vs. body completely devoid of dark spots). Additionally, H. khimaera differs from H. cochliodon , H. dardanelos Zawadzki & Carvalho, 2014 , H. ericae Hollanda Carvalho & Weber, 2005 , H. ericius Armbruster, 2003 , H. labyrinthus Oliveira, Ribeiro, Canto & Zawadzki, 2020 , H. oculeus (Fowler, 1943) , H. paucipunctatus Hollanda Carvalho & Weber, 2005 , H. pyrineusi (Miranda Ribeiro, 1920), H. taphorni (Lilyestrom, 1984) and H. waiampi Hollanda Carvalho & Weber, 2005 by having more externalized opercle, with exposed portion easily visible (vs. more internalized opercle, with exposed portion, when present, strongly reduced). It can be further distinguished from H. pagei Armbruster, 2003 by having fewer plates between the caudal and adipose fins (6–8 vs. 9–10); from H. kopeyaka Carvalho, Lima & Zawadzki, 2010 and H. labyrinthus by having only rounded dark spots on the body (vs. conspicuously horizontally elongated dark spots in addition to rounded dark spots in H. kopeyaka ; dark vermiculated spots in addition to rounded dark spots in H. labyrinthus ); from H. oculeus , H. plecostomoides (Eigenmann, 1922) and H. simios Hollanda Carvalho & Weber, 2005 by having dark spots on the body in a clearly smaller number and more widely spaced even in the most spotted specimens (vs. body densely covered by closely spaced spots); from H. levis (Pearson, 1924) by having an adipose fin (vs. adipose fin absent); from H. sculpodon by having 26 to 28 plates in the median series of lateral plates (vs. 30).

Description. Morphometric data in Tab. 1. Head large, slightly compressed laterally. Snout and anterior profile generally subtriangular in dorsal view; slightly rounded in some specimens. Eye generally of moderate size, varying slightly in size in some individuals and positioned dorsolaterally. Dorsal margin of orbit elevated. Compound pterotic covered with well-developed odontodes in some individuals. Greater body width at cleithrum, gradually compressing towards base of caudal fin; in some specimens, compression more pronounced posteriorly to first four mid-ventral plates. Opercle partially exposed, completely bordered by thick layer of skin; covered by numerous odontodes; exposed portion small to moderate in size, typically irregular, with ellipsoid, rounded or teardrop shape ( Figs. 2A, 3).

Dorsal profile of head almost straight from tip of snout to anterior margin of parietosupraoccipital, forming angle of approximately 45º with ventral surface of head; convex from this point to origin of dorsal fin; sloping downwards from this point to first dorsal procurrent ray of caudal fin, rising again from this point to insertion of caudal fin. Ventral profile varying from almost straight to slightly convex from tip of snout to insertion of unbranched ray of pelvic-fin; almost straight from this point to first ventral procurrent ray of caudal fin, descending to caudal-fin insertion. Caudal peduncle reduced and laterally compressed, wider in anterior region and very compressed posteriorly to insertion of adipose fin.

Oral disc rounded, sometimes ovoid, relatively small in size; lower lip not reaching transverse line through gill openings; ventral surface covered by numerous small papillae, decreasing in size posteriorly; odontodes generally present on almost entire surface of upper lip in adults; smaller specimens sometimes with wider naked region. Maxillary barbel equal to or slightly longer than eye pupil. Buccal papilla generally absent; papilla present and small in some specimens. Dentaries acutely angled, varying from about 70º to 90º between each ramus (average, 82º) (33); poorly preserved and/ or stored specimens (especially those compressed into the jars) may variably appear to have an angle larger than 90º between dentary rami. Eight to 44 teeth (mode 32; 21*) in premaxillary, 9 to 39 teeth (mode 31; 22*) in dentary. Bicuspid teeth, with medial cusp generally shovel-shaped or spatulate ( Figs. 2B–C); medial cusp varying from rounded to oblong in ventral view; conspicuously more developed than lateral cusp ( Figs. 2B–C, 4); distinct and generally well-developed lateral cusp ( Figs. 2B–C), sometimes fused with medial cusp (similar to condition found in Hypostomus hemicochliodon , see Armbruster (2003, 2004) and H. soniae , see Hollanda Carvalho, Weber (2004); rarely, some larger specimens (119.2 mm to 171.4 mm SL) with villiform medial cusp. Dentition of juvenile specimens (up to approximately 100.0 mm SL) with same morphological variations observed in adults ( Fig. 4).

Body covered by dermal plates bearing odontodes, most concentrated on dorsal surface of head and in middle portion of dermal plates. Dorsal surface of snout covered by dermal plates, except for small naked area on snout tip. Mesethmoid region covered by more developed odontodes, forming median bulge from snout tip to nares. Conspicuous concentration of well-developed odontodes near nostril, passing through dorsal margin of orbit and extending through dorsal portion of compound pterotic, forming keels; less developed odontodes in some juvenile individuals. Predorsal region medially keeled; keels moderately developed, diverging slightly towards dorsal fin; less developed keels in some smaller individuals (up to about 100.0 mm SL). Dorsal surface of trunk covered by dermal plates except region around dorsal-fin. Lateral surface of trunk with five series of dermal plates on anterior portion of flanks and with four series of dermal plates on postdorsal portion. Dorsal and mid-dorsal series of plates generally with moderately developed keels, gradually becoming less developed posteriorly; some larger individuals (around 160.0 mm SL) with keels on dorsal and mid-dorsal series well developed ( Fig. 5B). Median series with lateral line, generally lacking keels; keels, when present, poorly developed. Mid-ventral series with angled plates from first to fifth plate, with keels varying from little developed to moderately developed; keels absent posteriorly. Ventral surface of trunk covered by minute dermal plates in individuals greater than 80.0 mm SL, except on region around pectoral and pelvic fins. Ventral series smoothly angled ventrally towards posterior portion of caudal peduncle. Ventral surface of head covered by small bony plates, with exception of region around mouth.

Pre anal plate present and exposed. Median series with 26 to 28 (mode 27*) plates. Body covered dorsally by longitudinal series of two dermal plates in pre dorsal region, followed posteriorly by nuchal plate; plates between dorsal and adipose fins generally 6–8 (mode 7*); single specimen (CITL 1311, 118.7 mm SL) with 5; plates between adipose and caudal fins 6–8 (mode 7*), and plates at base of dorsal fin generally 6–8 (mode 7*).

Dorsal fin II,7* (53), its origin at vertical through midpoint between pectoral and pelvic fins or slightly posterior to that point, usually passing halfway through second mid-ventral plate; region around dorsal-fin base naked; posterior margin straight or somewhat convex; posterior dorsal-fin rays not reaching adipose-fin spine when adpressed. Adipose-fin spine laterally compressed, ranging from almost straight to slightly curved inwards ( Fig. 6). Pectoral fin I,6* (53), with almost straight posterior margin; pectoral-fin spine slightly curved inwards, covered by odontodes across its entire surface; anterior margin of spine with odontodes gradually becoming more developed towards tip of spine; dorsal surface of spine bearing poorly-developed odontodes with similar size, roughly aligned in longitudinal rows (relative to main axis of spine) on proximal two-thirds of spine, except for its posterior margin, which bears conspicuous line of odontodes gradually becoming more developed towards tip of spine; odontodes on distal portion of spine typically curved towards origin of spine in larger specimens (with more than 100.0 mm SL); ventral surface of spine mostly covered by poorly-developed odontodes with similar size, its distal portion with slightly more developed odontodes; tip of adpressed pectoral-fin spine reaching to anterior third of adpressed pelvic-fin spine. Pelvic fin i,5* (53), its posterior margin almost straight or slightly convex; tip of adpressed spine extending beyond anal-fin origin. Anal fin i,4* (53), its distal margin reaching sixth bony plate posterior to its origin; unbranched rays and fifth branched ray smaller than second, third and fourth branched rays; third branched ray usually largest. Caudal fin i,14,i* (53); posterior margin furcated, with ventral lobe slightly larger than dorsal lobe; ventral lobe conspicuously larger than dorsal lobe in some individuals.

Coloration in alcohol. General color pattern in Figs. 1, 5-8, 11. Ground color of body generally ranging from light to dark yellowish brown; greyish brown in some specimens. Dark brown or black blotches on at least some region of body, typically with numerous closely spaced small rounded dark spots on head region (sometimes fused in some individuals), gradually becoming larger and more spaced posteriorly ( Fig. 5B); in some individuals, dorsal surface of head may present sparse and diffuse black spots on snout, becoming more numerous and conspicuous in region posterior to eyes (compound pterotic and supraoccipital) ( Figs. 1, 6A). Some specimens with darker background color, with few spots on body; dark blotches indistinct in intensely pigmented specimens, possibly due to melanism. Diffuse stripe between keels mid-dorsal and mid-ventral series of plates typically present, except for conspicuously dark specimens (possibly melanic) and specimens from córrego Boa Sentença, which apparently lack midline stripe. Ventral surface of trunk generally with small- to moderate-sized and roughly rounded dark brown or black blotches; some individuals devoid of or with sparse and diffuse blotches ( Fig. 7). Ground color of fins similar to ground color of trunk, sometimes lighter; some specimens with ventral lobe of caudal fin slightly darker than dorsal lobe. Region around base of dorsal fin darker, extending medially along dorsal portion of caudal peduncle, generally forming diffuse dark band in dorsal region; color pattern of anal fin similar to that of dorsal fin, but dorsal fin generally darker. All fins generally covered with small, widely-spaced dark blotches; few, diffuse spots in some individuals; some specimens with conspicuous concentration of dark pigmentation on dorsal-fin membranes; some specimens with closely spaced dark blotches in pectoral and/or caudal fins, transversally aligned, forming dark bands. Color pattern of juvenile similar to that of adults and presenting same variations.

Coloration in life. Similar to color pattern of preserved specimens, with exception of dark stripe along midline of flank, which tends to be more evident. Background color of body yellowish brown in most all individuals; variably reddish brown, dark brown, greyish brown, or bright yellow ( Fig. 8).

Sexual dimorphism. No sexual dimorphism was observed.

Geographical distribution. Hypostomus khimaera is widely distributed in the upper rio Paraguai basin within the Brazilian territory, occurring in the sub-basins of the rivers Aquidauana, Coxipó, Cuiabá, Manso, Jauru, Pari ( LFCT, pers. obs.), Piquiri , Quilombo , Sepotuba , Taquari , Vermelho and Chacororé , in the states of Mato Grosso and Mato Grosso do Sul. Additionally , the species was recently recorded for the upper rio Paraná basin, in the córrego Mimoso , 21°31’48”S 53°26’47”W, a tributary of the rio Anhanduí, southeastern Mato Grosso do Sul (Lopes et al., 2024) ( Fig. 9) GoogleMaps .

Ecological notes. Hypostomus khimaera is a species mainly found in streams and smaller rivers ( Figs. 10A, B, D), but can be occasionally found in the main channel of larger rivers ( Fig. 10C), commonly associated with marginal regions of lentic environments and sandy bottoms, especially with branches/trunks of submerged trees.

In headwater streams, small individuals of this species (up to approximately 100.0 mm SL) can also be found in lotic and shallow environments (up to around 20 cm depth), with a substrate composed by gravel, larger stones and sand. In the rio do Peixe, state of Mato Grosso do Sul, large specimens (about 250.0 mm SL) were found in lotic shallow environments (about 30 cm depth) and rocky substrate, with sections of slab, hiddening under large boulders during the day, an atypical environment for a species from the H. cochliodon group (LFCT, 2024, pers. obs.). Like most loricariids, the species is more easily observed at night. Despite being part of the H. cochliodon group, the morphology of its teeth is clearly different from the typical spoon-shaped pattern of xylophagous species (see Hollanda Carvalho, Weber, 2004: 960, fig. 4b, c), which combined with the common presence of specimens with strongly worn teeth suggests that the species may not be essentially xylophagous. The large morphological variations in body and dentition indicates that H. khimaera is possibly a species with presents high phenotypic plasticity, being opportunistic feeding habits, grazing both wood and periphytic matrix of rocky substrates.

Remarks. Despite being a considerably variable species in both morphology and color pattern, some populations are clearly more distinct from the expected range of variations in Hypostomus khimaera . Specimens from the córrego Fortaleza ( Fig. 11A), a tributary of the rio Taquari in the region of Coxim, Mato Grosso do Sul, have a rounded snout in dorsal view (vs. roughly triangular, pointed), a reduced adipose fin (vs. well developed), and a comparatively shorter and more robust first rib (vs. longer and slender). Despite this, the fact that individuals with this peculiar morphological pattern occur only in this small tributary of the rio Taquari, and that specimens displaying similar morphology to the typical H. khimaera were also captured syntopically (see Fig. 8H), suggests that these individuals only represent another morphotype of H. khimaera . Other point that also raises doubts in this matter is the fact that the c&s individuals apparently have malformations in their ribs (from the second on), as well as other malformations in the cranium of some non c&s specimens. Furthermore, several collections in the rio Taquari main channel and smaller tributaries (see Material Examined section) revealed only individuals clearly compatible with the typical pattern of H. khimaera , which makes this issue even more complex. Despite the morphological differences raised herein, the number of individuals with this morphological pattern is relatively scarce, and broader integrative approaches are still necessary to better understand this population.

The population of the córrego Rio Verde ( Fig. 11B), a tributary of the rio Verde in the municipality of Rio Verde de Mato Grosso, Mato Grosso do Sul, also presents morphological differences when compared to the typical H. khimaera , especially in larger individuals (larger than 120.0 mm SL): shorter and robust caudal peduncle, smoothly narrowing towards caudal-fin base in dorsal and lateral views ( Fig. 8I) (vs. caudal peduncle typically longer, slender, clearly narrowing towards caudal-fin base ( Figs. 8A–H)); rounded snout in dorsal view (vs. roughly triangular, pointed) and the cleithrum apparently wider than other individuals of the same size. Interestingly, smaller individuals (less than 120.0 mm SL) collected at this site have the typical morphology expected for juvenile H. khimaera . Additionally, even with the aforementioned morphological differences, larger specimens (with more than 120.0 mm SL) have the tooth pattern (i.e., number and general morphology of medial cusp) compatible to the typical H. khimaera . In this sense, even though some morphological differences exist, important diagnostic features of H. khimaera are present in individuals from this population (e.g., teeth morphology and number, opercle morphology, dark midline stripe, and dark blotches on body), which led us to consider it as one of the variable forms of H. khimaera .

A third distinct population was found in the córrego Boa Sentença ( Fig. 11C), another tributary of the rio Verde, rio Taquari basin. Individuals from this population have comparatively larger dark blotches (vs. clearly smaller blotches), and apparently lack the dark stripe along midline of flank (vs. midline stripe typically present and evident). Despite the apparent absence of the typical midline stripe in this population, it is important to mention that the available material is scarce, especially regarding adult specimens (largest specimen with about 120.0 mm SL). Therefore, new survey efforts in this drainage remain necessary to safely point the absence of the typical midlateral stripe as well as other putative diagnostic features, allowing us to undoubtedly confirm it as another variable form of H. khimaera . However, considering the overall similarity between the specimens from the córrego Boa Sentença and the typical H. khimaera , we opted to consider them conspecific herein. In any case, the dark longitudinal stripe along midline of flank is still extremely useful to distinguish H. khimaera from its congeners.

In the original description, the number of teeth (including both mandibular rami) was proposed as a diagnostic feature between H. khimaera (12 to 27 teeth) and H. cochliodon (7 to 9) (Tencatt et al., 2014). However, our analysis demonstrates a wider range for H. khimaera (8 to 44 teeth), overlapping with the known range of H. cochliodon . Notwithstanding, specimens of H. khimaera with eight or nine teeth variably presented unusually large gaps between teeth, suggesting tooth loss at these points. In any case, the absolute number of teeth was considered herein to standardize tooth count. The lower counts in tooth range falling within the range of H. cochliodon was found in specimens from different localities (e.g., CPUFMT 2561 and CPUFMT 8344), whereas the higher limit was found in specimens from the córrego Fortaleza (e.g., CITL 1159).

Even though it was possible to recognize some differences in color pattern and/ or external morphology when comparing the populations from the córrego Fortaleza, córrego Rio Verde and córrego Boa Sentença (check previous paragraphs in this section), most of these putative diagnostic features are not exclusive, being variably present in individuals from other populations, although in lower frequency. In this sense, the most reasonable decision right now is to attribute these populations to H. khimaera , something that can eventually change as suitable diagnostic characters are discovered.

Molecular data. The analyses indicate that the sequences examined in this study form a cohesive molecular taxonomic unit (MOTU), identified morphologically as Hypostomus khimaera ( Fig. 12). Furthermore, these analyses reveal that specimens from the córrego Fortaleza, the most morphologically distinct population when compared to typical H. khimaera (see Remarks), exhibit 0% of intraspecific genetic distance when compared to sequences of typical H. khimaera from other localities, such as the Cuiabá and Piquiri river basins ( Tab. 2). Hypostomus khimaera exhibits a genetic distance exceeding 3% when compared to congeners previously deposited in the Genbank ( Tab. 2).

Material examined. In addition to the material examined by Tencatt et al. (2014), a total of 636 specimens were examined, which are listed below. All from Brazil, upper rio Paraguai basin. Mato Grosso. CPUFMT 3028 , 8 , 101.7 – 146.6 mm SL ; CPUFMT 3550 , 5 , 120.0– 153.3 mm SL ; CPUFMT 2729 , 6 , 118.4 – 135.3 mm SL ; CPUFMT 2844 , 3 , 123.3 –157.0 mm SL ; CPUFMT 3417 , 3 , 103.7 – 132.7 mm SL ; CPUFMT 2560 , 2 , 121.9 – 124.9 mm SL ; CPUFMT 3016 , 2 , 90.4–90.9 mm SL ; CPUFMT 3496 , 1 , 96.4 mm SL ; CPUFMT 1454 , 32 , 59.0– 125.3 mm SL ; CPUFMT 4154 , 1 , 97.5 mm SL ; CPFMT 4171 , 1 , 80.6 mm SL ; CPUFMT 183 , 2 , 46.4–47.3 mm SL ; CPUFMT 160 , 2 , 78.5–85.2 mm SL ; CPUFMT 3002 , 2 , 64.3–73.1 mm SL ; CPUFMT 407 , 1 , 76.1 mm SL ; CPUFMT 3164 , 1, 111.6 mm SL ; CPUFMT 1455 , 3 , 32.5–61.6 mm SL ; CPUFMT 1503 , 1, 124.1 mm SL ; CPUFMT 2302 , 1 , 65.8 mm SL ; CPUFMT 2367 , 3 , 109.2 – 132.9 mm SL ; CPUFMT 4624 , 1 , 55.3 mm SL ; CPUFMT 8344 , 2 , 65.6–81.1 mm SL ; CPUFMT 8345 , 1 , 71.4 mm SL ; CPUFMT 8346 , 5 , 47.8–57.2 mm SL ; CPUFMT 8347 , 5 , 47.2–50.9 mm SL ; CPUFMT 8348 , 2 , 72.23–77.1 mm SL. Mato Grosso do Sul . CITL 1137 , 1 , 58.2 mm SL ; CITL1138 , 1 , 59.8 mm SL ; CITL 1139 , 66 , 24.5–98.9 mm SL ; CITL 1140 , 33 , 46.3–113.7 mm SL ; CITL 1141 , 5 , 65.5–69.4 mm SL ; CITL 1142 , 9 , 67.6–99.3 mm SL ; CITL 1143 , 3 , 90.8–91.8 mm SL ; CITL 1144 , 5 , 68.5–75.1 mm SL ; CITL 1145 , 2 , 83.6–88.7 mm SL ; CITL 1146 , 4 , 33.9–64.4 mm SL ; CITL 1147 , 17 , 55.0– 86.1 mm SL ; CITL 1148 , 2 , 60.6–68.2 mm SL ; CITL 1149 , 1 c&s, 134.5 mm SL ; CITL 1150 , 7 , 43.8–102.2 mm SL ; CITL 1151 , 2 , 75.8 – 71.1 mm SL ; CITL 1152 , 1 c&s, 117.9 mm SL ; CITL 1153 , 3 , 49.9–116.5 mm SL ; CITL 1154 , 1 , 77.0 mm SL ; CITL 1155 , 2 , 65.4–91.2 mm SL ; CITL 1156 , 5 , 125.5 – 151.1 mm SL ; CITL 1157 , 1 , 93.4 mm SL ; CITL 1158 , 14 , 55.5–123.8 mm SL ; CITL 1159 , 5 of 7, not measured; 2 c&s of 7, 96.0– 112.2 mm SL ; CITL 1160 , 18 , not measured ; CITL 1161 , 2 , 85.12– 96.4 mm SL ; NUP 21369 , 12 , 28.0–95.0 mm SL ; NUP 21426 , 20 , 28.0–63.0 mm SL ; NUP 21459 , 27 , 22.0–62.0 mm SL ; NUP 21628 , 9 , 40.0–122.0 mm SL ; NUP 25433 , 4 , 68.2–77.7 mm SL . ZUFMS-PIS 1163 , 9 , 55.8–114.6 mm SL ; ZUFMS-PIS 1566 , 6 , 29.8–63.7 mm SL ; ZUFMS-PIS 1807 , 4 , 70.7–91.9 mm SL ; ZUFMS-PIS 1881 , 1 , 82.5 mm SL ; ZUFMS-PIS 1910 , 1 , 101.0 mm SL ; ZUFMS-PIS 1965 , 56 , 26.1–69.2 mm SL ; ZUFMS-PIS 1983 , 12 , 23.5–103.1 mm SL ; ZUFMS-PIS 2034 , 19 , 27.4–69.1 mm SL ; ZUFMS-PIS 3532 , 1 , 56.2 mm SL ; ZUFMS-PIS 3550 , 1 , 60.1 mm SL ; ZUFMS-PIS 3595 , 2 , 20.6–26.6 mm SL ; ZUFMS-PIS 3833 , 2 , 88.0– 101.4 mm SL ; ZUFMS-PIS 3862 , 1, 115.2 mm SL ; ZUFMS-PIS 3874 , 4 , 42.7 –89.0 mm SL ; ZUFMS-PIS 3966 , 7 , 29.0–66.0 mm SL ; ZUFMS-PIS 3967 , 1 , 38.8 mm SL ; ZUFMS-PIS 3976 , 2 , 29.6–56.9 mm SL ; ZUFMS-PIS 3982 , 4 , 32.7–42.4 mm SL ; ZUFMS-PIS 4051 , 3 , 35.9–58.9 mm SL ; ZUFMS-PIS 4219 , 4 , 31.1–44.1 mm SL ; ZUFMS-PIS 4246 , 7 , 53.7–99.7 mm SL ; ZUFMS-PIS 4255 , 29 , 40.4 –89.0 mm SL ; ZUFMS-PIS 4284 , 1 , 39.0 mm SL ; ZUFMS-PIS 4319 , 3 , 64.4–142.2 mm SL ; ZUFMS-PIS 4320 , 2 , 98.0– 146.3 mm SL ; ZUFMS-PIS 4324 , 1 , 91.6 mm SL ; ZUFMS-PIS 4343 , 2 , 40.8–43.4 mm SL ; ZUFMS-PIS 4357 , 8 , 42.6–78.3 mm SL ; ZUFMS-PIS 4368 , 49 , 45.5–107.9 mm SL ; ZUFMS-PIS 4465 , 2 , 48.2–53.3 mm SL ; ZUFMS-PIS 4948 , 1 , 60.0 mm SL ; ZUFMS-PIS 5037 , 2 , 53.3– 71.7 mm SL ; ZUFMS-PIS 5277 , 6, 171.8 mm SL ; ZUFMS-PIS 5949 , 1 , 33.3 mm SL ; ZUFMS-PIS 5962 , 11 , 31.1–59.6 mm SL ; ZUFMS-PIS 5976 , 1 , 46.9 mm SL .