Lebetus patzneri, Schliewen & Kovačić & Cerwenka & Svensen & Ordines, 2019

Schliewen, Ulrich K., Kovačić, Marcelo, Cerwenka, Alexander F., Svensen, Rudolf & Ordines, Francesc, 2019, Lebetus patzneri (Teleostei: Gobiidae), a new goby species from the Balearic Islands, western Mediterranean, with first records of Lebetus guilleti (Le Danois, 1913) from this area and Norway, and with notes on its biology, Zootaxa 4706 (2), pp. 231-254 : 236-243

Lebetus patzneri sp. nov. Schliewen, Kovačić & Ordines

( Figs. 2–9 )

Holotype. ZSM 47486, female, 16.6 mm SL (20.8 mm TL, Spain, Balearic Islands, Menorca Channel (39.859°N, 3.5413°E), rodolith beds, 72 m depth, coll. F. Ordines during INTEMARES-CANAL0419 survey on April 27th 2019 with haul #16 ( Figs. 1–3 ; 6 ). Preserved in 4% formol solution after capture, later transferred to 75% EtOH, right pectoral fin removed as DNA-tissue voucher ZSM-PIS-GO 2124.

Paratype. ZSM 46865, female, 16.8 mm SL (caudal fin damaged), Spain, Balearic Islands, off the southern coast of Mallorca (39.2372°N, 2.9777°E), rodolith beds, 60 m depth, coll. F. Ordines during DRAGONSAL0914 survey on September 8 th 2014 with haul #25 ( Figs. 4 , 5 , 7 , 8a,b ). Preserved in 4% formol solution after capture, later transferred to 75% EtOH, and now stored as a critical-point-dried and gold-sputtered SEM specimen mounted on SEM mount (details of preparation for SEM see Material and Methods) .

Non-type material. Photographs of two specimens ( Figs. 8c,d ) of unknown size and sex, Spain, Balearic Islands, Menorca channel (39.8628°N, 4.0747°E), red algae beds, 67m depth, coll. F. Ordines during the INDE- MARES-CANAL0411 survey on September 7 th 2011 with beam trawl #26. Not preserved.

Diagnosis. L. patzneri sp. nov. is distinguished from the one or both other Lebetus species by each of following characters: (1) P 15–16 vs. 17–21 in L. scorpioides ; (2) D2 segmented fin rays 7–8 vs. 9–10 in L. scorpioides ; (3) snout slightly larger than eye diameter vs. snout clearly shorter than eye diameter in L. guilleti ; (4) eyes small, 23.3–25.5% of head length vs. eyes large, 29.4–35.7% of head length in L. scorpioides and 28.8–35.5% of head length in L. guilleti ; (5) body height low, body depth at pelvic fin 14.9–16.4% of SL vs. 16.6–21.1% of SL in L. guilleti ; (6) lips thick, giving mouth a broad, almost rectangular shape in dorsal and lateral view vs. lips of normal size, mouth of oval shape in dorsal and lateral view in both L. scorpioides and L. guilleti ; (7) minute black “pepper” spots on predorsal area, dorsum, flanks and caudal peduncle, visible in fresh and preserved specimens vs. no similar spots in L. guilleti and L. scorpioides ; (8) the most of the caudal peduncle similarly colored as the lateral area below D2, reddish or brown vs. sharp dark-pale boundary between caudal peduncle and the rest of the body in L. guilleti and L. scorpioides .

Description (all morphometric values and meristics in the text are presented as holotype first and paratype in parentheses). General Morphology ( Figs. 2–4 ; Table 2 ). Body proportions and meristic counts of holotype and paratype are given in Table 2 . Body elongate, subcylindrical and slightly laterally compressed towards caudal fin. Head fairly long 32.2 (32.5) % SL, slightly depressed, with a nearly horizontal predorsal profile. Snout moderately long and pointed, about as long as eye 105.2 (106.3) % in eye length, dorsal profile of snout gently sloping. Eyes dorsolaterally extending in fresh specimens above dorsal profile, large 25.5 (23.3) % of head length, interorbital space narrow, about one-fourth of eye length. Anterior and posterior nostrils both short, not reaching upper lip, tubular, erect, without process from rim; anterior nostrils slightly longer than posterior ones. Mouth oblique from lateral view, posterior angle of jaws below anterior third of eye. Lips ending about equally anteriorly, thick, giving mouth a broad, almost rectangular shape in dorsal and lateral view. Minimum teeth counts of inner rows were made for paratype only, because for the holotype no high-resolution MicroCT image was available; however roughly estimated numbers for holotype are similar to the paratype: lower jaw with three rows of unicuspid teeth at symphysis, the outer row with more than twenty (left: 22 (25), right; 26 (21)) upright teeth, the second (intermediate) row with very small and irregularly set teeth along the anterior jaw part, and the third row with teeth as large as in the first row, but more slender, the first reaching back to the jaw angle, the third row reaching back almost as far as the first row. Upper jaw with two rows, the outer row with slender and backwards curved teeth (more than 20 on both sides in holotype, at least 17 on each side in paratype), and an interior row of numerous small to minute and irregularly set teeth, reaching farther back to the jaw angle than the anterior row. Chin without mental flap (fold) or barbels. Cranial roof covered with dorsal axial musculature. Pectoral girdle without flaps on anterior edge. Genital papilla short, broad and with terminal villose processes on the posterior and lateral parts, but without on the anterior part of the rim.

Fins ( Fig. 5 ). D 1 VI (VII); D 2 I /7 (I/8) (last bifid); A I/6; C 9 branched rays, 13 segmented rays (caudal fin of paratype damaged, therefore identification as branched and segmented tentative); P 16 (16 left and 15 right), V I/5 + 5/I. Fin-bases and lengths in proportion to standard length are given in Table 2. D 1 II and III the longest spine of D1, their tips just reaching D 2 I anterior origin. Fin membrane of D 1 VI not connected with base of D 2 I. D2 commences at vertical of urogenital papilla, with its last ray over vertical of last A ray. A commences below third (holotype) or second (paratype) segmented ray of D2. C rounded, shorter than head length. Uppermost P rays within membrane, all P rays unbranched, ending at vertical of D2 spine (holotype), or just before it (paratype, but potential damage has to be taken into account). V without an anterior membrane; V fin membranes strongly damaged, i.e. it is not possible to infer shape and completeness of pectoral disc. V longest ray either not reaching genital papilla (holotype; longest ray damaged?) or reaching slightly beyond (paratype).

Squamation ( Figs. 3 , 4a ): Body scaled. Since both specimens lost scales, scale counts are based on scale pouches visible on the stained holotype and on the REM image of the paratype and scale types could not be identified. Head including opercle and cheek naked. Predorsal area, including nape, and the area below D1 downwards to the horizontal level of upper edge of pectoral fin base naked. Breast and belly naked. Scales in lateral series 25 on both sides plus one on caudal fin origin (23 plus one on caudal fin count on the right side). Scales in transverse series 7.

Osteology. Postcranial skeleton and pterygiophore insertion pattern ( Fig. 5 ). 11 (12) precaudal and 15 caudal vertebrae including urostyle; total count 26 (27). In lateral view, first three vertebral centra short, becoming progressively longer in vertebrae 4 and 5, and approximately of equal lengths further posteriorly. First three vertebrae substantially, fourth vertebrae slightly shorter than posterior ones. Neural spines and haemal spines of last four vertebrae before urostyl increasingly enlarged posteriorly, especially last two haemal spine. D1 pterygiophore formula 3- 221100 (3-2220100); one prehaemal A pterygiophore. Four branched caudal fin rays inserting at hypurals 1 + 2 (fused), five at hypurals 3 + 4 (fused with each other and with urostyl); 1 branched and 1 unbranched ray in hypural 5; one branched ray in parhypural; each one unbranched caudal ray beyond hypural 5 and parhypural, respectively; total number of fin rays inserting in hypuralia and parhypural 12. Hypural 5 and parhypural small splint-like bones. One epural. Five dorsal and six ventral procurrent rays.

Lateral line system ( Fig. 6 ). No head canals present. Suborbital sensory papillae of the head lateral line system were identified on the holotype and confirmed in part on the paratype where many rows are lost due to the heavily damaged surface. The sensory papillae of the head lateral line system are large and reduced in number in all rows, many represented by a single papilla. Rows with count of sensory papillae in parentheses: (1) preorbital: snout with two rows in median preorbital series visible, row r not visible, superior row s 1 (1) close to posterior nostril, inferior row s 3 (1) above upper lip visible only in holotype. Lateral series c: superior c 2 as single papilla visible only in holotype, superior c 1 (2) behind and below anterior nostril, upper inferior c 2 not visible, lower inferior c 1 (1) above upper lip. (2) suborbital: rows a and c, including cp, without transverse proliferation; row a longitudinal (3) below rear part of eye; row c (4) longitudinal below frontal and middle part of eye anteriorly with two papillae superimposed each other between anterior part of eye and posterior part of upper lip, the row c visible only in holotype; two cp larger papillae bellow a row anterior part, lower cp’ anterior to upper cp. Longitudinal row b as two papillae below rear border of eye, only one papilla visible in paratype. Longitudinal row d with single papilla above and behind mouth corner. (3) preoperculo-mandibular: internal row i divided into anterior (i: 6 in holotype, 5 in paratype), and posterior sections, posterior section visible only in holotype (i: 2 + 2); external row e divided into anterior and posterior section (2 + 1), anterior visible only in holotype; mental row f (1) present; two larger papillae marked pc present on the place of the absent PC. (4) oculoscapular: longitudinal row marked aoc (2) starting behind from posterior edge of orbit on the place of the lost AOC; anterior row x 1 as single visible papilla above preopercle visible also in paratype, all other oculoscapular rows recorded only on the holotype: posterior row x 2 (1) above posterior opercle, row u (2) on the place of the lost POC, axillary rows as 1 (2) and as 2 (1) present, no other oculoscapular papillae are visible; (5) opercular: two larger papillae marked pc present on the place of the absent PC are visible, transverse row ot (3) present with lower part; superior longitudinal row os and inferior longitudinal row oi both as single papilla in holotype while the surface of the opercular area was destroyed on both sides in the paratype, so no papillae are visible; (6) anterior dorsal: rows n (1) at posterior edge of orbit, another single visible papilla is assigned to row g since row m is usually lower and first lost in species with reduced lateral line system and row o more anteriorly placed, so no rows o and m, row h (1) in front of the first dorsal fin visible only in holotype. (7) interorbital: interorbital papillae not visible.

Otoliths ( Fig. 7 ). The Micro-CT of the paratype ZSM 46865 shows a right otolith in medial/inner view ( Fig. 7a ) and a left otolith seen from the anterior margin ( Fig. 7b ). A small utricular otolith is visible in the ventral sector of each saccular otolith. The otolith shown in Fig. 7a displays smooth margins and an almost quadratic outline. The dorsal margin ascends slightly toward the posterior margin, at the highest tip it declines and forms a slightly bulged posterodorsal projection. The posterior margin is slightly concave in the middle, its transition to the ventral margin is skewed. The ventral margin is straight, only the passage to the anterior margin is slightly curved. The anterior margin is straight and displays a small predorsal angle. A clear ventral line extends with some distance from the ventral rim and extends up to the tip and end of the sulcus, respectively. The sulcus is relatively small, its length is 30% of otolith length, its height equals 10% of the otolith length. The orientation of the sulcus is horizontal. It appears that the cauda is wider than the ostium, but this would have to be checked with an SEM image. The otolith shown in Fig. 7b reveals a bent, thickened lateral face and a relatively plane medial face; the total thickness of the otolith is approx. 20% of otolith length. The general outline of this otolith conforms well to that of the otoliths of Oxudercidae , and in particular to that of the otoliths of species of the Pomatoschistus -lineage (see Gierl et al. 2018: Figs. 2 , 3 ). The quadratic otolith shape of L. patzneri sp. nov. resembles that of Pomatoschistus marmoratus (imaged in Gierl et al. (2018), Figs. 3 d–f). However, a horizontal position of the sulcus as seen in the present otolith does not occur in otoliths of the species of Pomatoschistus .

Coloration. Color of the freshly collected females, holotype ( Figs. 2 .a–d) and paratype ( Figs. 8a,b ) and photographs of two additional specimens ( Figs. 8c,d ). Holotype and paratype: ground color of body, head, snout and lips orange-red except for white areas of lower head, lower postopercular region belly and ventral caudal peduncle; back with four whitish-beige saddle marks, one before D1 origin, one below D1, one at D2 origin and one forming a broad vertical band on caudal peduncle, the latter vermiculated in the holotype; additional very small whitish blotches in the predorsal area and along posterior lower flanks. Chin, suborbital region and lower parts of preopercle porcelain white with two well defined red bands almost meeting at a moderately acute angle below eye diverging radially downwards; ventral part of preopercle fringed with a narrow red margin in the paratype, but reduced to a red narrow spot in the holotype. Iris orange-red, partially blackish or with a golden hue; interior iris margin golden. Dorsal area and flanks with many minute blackish spots, fewer in the paratype on predorsal region than in the holotype, and generally more densely aggregated on caudal peduncle and anal fin base, and forming in the paratype two vertical narrow bands at vertical of D1 anterior and posterior origins. Head, preopercle, opercle, pectoral and anterior two thirds of ventral area almost without minute small black spots. D1 black (folded down on photos), in paratype with two narrow white bands indicated at distal margin; D2 (folded down on photos) red with white bands indicated. P base white with scattered orange-red spots, P transparent with few thin reddish bands indicated in paratype. C with a porcelain white basal blotch, sharply delineated anteriorly, otherwise transparent orange and with a whitish band at about middle part (visible only in holotype and one of the additional only photographed specimens). The coloration of the two additional specimens is similar, matching the holotype coloration, D1 of the one unpreserved only photographed specimen ( Fig. 8c ) is partially unfolded and exhibits an overall black coloration and small pinkish-red elements along the base of two spines as well as a white blotch at the upper posterior area and around the membrane between the last D1 spine and the dorsum; furthermore, whitish-transparent V are visible in the holotype ( Fig. 2d ) and second unpreserved and only photographed specimen ( Fig. 8d ). In the second photographed only specimen the C base has an inverted brownish triangular bar, darker at the caudal and ventral margins, more reddish on the dorsal side, and the posterior base of the same vertical boundary as in the other specimens.

Color of preserved specimens in alcohol. Body of holotype yellowish white, with orange-red coloration completely lost. The minute small black spots on the dorsal area and flanks with still clearly visible. D1 black and other fins transparent. The paratype was completely discolored (whitish) before preparation of the specimen for REM, except for the distinct and scattered and minute black spots on predorsal dorsal and posterior flanks. D1 black.

Etymology. The species is named patzneri , to honor Prof. Robert (“Bobby”) Patzner, Professor Emeritus at the University of Salzburg ( Austria), in recognition for his dedication to the study of gobies in the Mediterranean including the Balearic Islands, see e.g. Patzner et al. (2011).

Genetics and phylogenetic position ( Fig. 9 ). Partial cytochrome oxidase subunit I (COI) sequences (520 base pairs) of Lebetus patzneri sp. nov. compared with five L. guilleti and one L. scorpioides sequences, as well as with those of members of all other sand goby and DLBS genera provide good support for the phylogenetic position as the sisterclade of the L. scorpioides from Norway (BS 87), as well as for its species-level distinctiveness. K2P-distance between the single L. patzneri and the single L. scorpioides COI-haplotype is 0.150 (±0.018), and between L. patzneri and each of the five L. guilleti haplotypes it ranges from 0.195 (± 0.019 S.E.) to 0.196 (± 0.019 S.E.). Thus, pairwise K2P-distances between the three congeneric Lebetus taxa range well above distances between other congeneric sand goby and DLBS-taxa, e.g. Pomatoschistus species ( Knebelsberger & Thiel 2014; Öztürk & Engin 2019) or Buenia species ( Kovačić et al. 2017, 2018).

Ecological and geographical distribution. Lebetus patzneri sp. nov. has only been caught so far from sampling stations on red algae beds in south Menorca, the Menorca Channel and off south Mallorca, at depths ranging between 60 and 72 m. The first two specimens caught, not preserved, were collected from a Peyssonnelia bed off the southern coast of Menorca ( Fig. 1 ). In that sample 87 kg of algae were collected from a swept area of 555 m 2, with dominance of red algae species belonging to the Peyssonneliaceae family (79 kg), mainly Peyssonnelia inamoena but also with some presence of rhodolith forming species such as the coralline algae Lithothamnion valens and Spongites fruticulosus (3 and 1.2 kg, respectively). The paratype was collected from a rhodolith bed off the south coast of Mallorca ( Fig. 1 ). In that sample 166 kg of algae were collected from a swept area of 273 m 2, with dominance of coralline red algae species, mainly Lithothamnion corallioides , Phymatolithon calcareum and Lithophyllum racemus (39, 10.6 and 4.1 kg, respectively), but also with brown algae, mainly Dictyota dichotoma and Dictyopteris lucida (21.9 and 3.1 kg, respectively). Finally, the holotype was collected from a rhodolith bed in the Menorca Channel ( Fig. 1 ). In that sample 56 kg of algae were collected from a swept area of 400 m 2, with dominance of coralline red algae species, mainly L. valens , unidentified crustose coralline algae and S. fruticulosus (22.6, 19.9 and 12.4 kg, respectively). In all three sampling stations, the specimens were also accompanied by a large diversity of benthic animals (both sessile and mobile) with most abundant species including the decapods crustaceans Inachus dorsettensis , Inachus thoracicus and Pagurus prideaux , the teleost fish Buenia massutii , Odondebuenia balearica , Diplecogaster bimaculata and Serranus hepatus , the echinoderms Sphaerechinus granularis , Ophiura albida and Echinaster sepositus , the annelids Hyalinoecia tubicola and Laetmonice hystrix , the ascidians Aplidium nordmanni , Ciona spp. and Ascidia mentula , and several unidentified sponge species.

Figures

FIGURE 1. Map showing the three sampling stations where all the individuals of Lebetus patzneri sp. nov. (stars) and L. guilleti newly recorded individuals (circles) from the Balearic Islands were collected. The holotype and the paratype of L. patzneri sp. nov. were collected from the sampling station in the Menorca Channel and off southeast Mallorca, respectively. The holotype, paratype and the two unpreserved specimens off south Menorca were collected at 72, 60 and 67 m, respectively. The 50, 100, 500 and 800 m isobaths are shown. Map drawn by F. Ordinas.

FIGURE 2. Lebetus patzneri sp. nov., aspects of the freshly collected holotype, ZSM 47486, female, 16.6 mm SL, from off SW Mallorca Island, 72 m depth, preserved. a: left lateral view; b: right lateral view; c: ventral view; c: dorsal view; d: ventral view left lateral view. Photographs by F. Ordinas.

FIGURE 3. Lebetus patzneri sp. nov., aspects of the preserved holotype, reversibly stained with Cyanine Blue, ZSM 47486, female, 16.6 mm SL, from off SW Mallorca Island, 72 m depth, preserved. a: left lateral view b: dorsal view; c: ventral view. Photographs by M. Kovačić.

FIGURE 4. Lebetus patzneri sp. nov., SEM aspects of the paratype ZSM 46865, female, 16.8 mm SL; from off SW Mallorca Island, 60m depth. a: right lateral view; b: ventral view of pelvic fins and urogenital papilla. SEM photo by A. Cerwenka.

FIGURE 5. Lebetus patzneri sp. nov., MicroCT reconstruction of the paratype ZSM 46865, female, 16.8 mm SL; from off SW Mallorca Island, 60m depth. a: left lateral view; b: dorsal view; c: ventral view. MicroCT reconstruction by A. Cerwenka.

FIGURE 6. Lebetus patzneri sp. nov., head lateral line system of the holotype ZSM 47486, female, 16.6 mm SL, from off SW Mallorca Island, 72 m depth. Terminology in text. Drawing by M. Kovačić.

FIGURE 7. Lebetus patzneri sp. nov., right saccular otolith of the paratype ZSM 46865, female, 16.8 mm SL; from off SW Mallorca, 60m depth. a: in medial/inner view; b: left otolith in anterior view. Orientation of otolith with respect to fish symmetry is indicated as follows: a, anterior; d, dorsal; l, lateral; m, medial; p, posterior; v, ventral. MicroCT reconstruction by A. Cerwenka.

FIGURE 8. Lebetus patzneri sp. nov., aspects of the paratype and two unpreserved specimens in life coloration shortly after collection. a: left lateral view of paratype ZSM 46865, female, 16.8 mm SL; from off SW Mallorca Island, 60m depth (based on montage of two separate photographs (slight artifacts at montage suture at about D1 origin due to slightly different position of the specimen between the two photographs); b: left dorsolateral view of head and anterior flank view of paratype ZSM 46865; c: left lateral view of the second known specimen (unpreserved), unknown size, Menorca channel, 67m depth (based on montage of two separate photographs; stitching artifacts visible before D1 origin); d: left slightly dorsolateral view of third known specimen (unpreserved) of unknown size, Menorca channel, 67m depth. Photographs by F. Ordines.

FIGURE 9. Maximum likelihood phylogenetic tree topology depicting the phylogenetic position and small intraspecific genetic distance of COI-barcode haplotypes (520 bp) of the holotype of Lebetus patzneri sp. nov. and five Lebetus guilleti from the northeastern Atlantic (Norway, two specimens), the Western Mediterranean (Baleares, one specimen) and from the northern Adriatic (Croatia, two specimens). All sand goby genera are included, except for Hyrcanogobius and Pseudaphya. Numbers on branches are bootstrap support values (%) for the Maximum Likelihood analysis (values below 50% not shown). Tip labels are composed of the genus and species name followed by ZSM collection number (if newly sequenced in this study) or of the GenBank accession number. Inserted photos depict the second known specimen of L. patzneri sp. nov. (as Fig. 8c; upper photograph) and the first L. guilleti specimen photographed and collected in 15m depth in Norway (ZSM-PIS-GO 1227; lower photograph). Photographs above and below by F. Ordines and R. Svensen, respectively.

Tables

TABLE 2. Morphometric characters of Lebetus patzneri sp. nov. holotype (ZSM 47486; female) and paratype (ZSM 46865; female); absolute values and % values in brackets. An asterik (*) indicates a damaged character.

		Holotype	Paratype
	Standard length (Sl) in mm	16.56	16.79
% Standard length	Anal fin base	2.48 (14.9)	2.70 (16.1)
	Anal fin spine length	1.40 (8.4)	1.36 (8.1)
	Body depth at anal fin origin	2.68 (16.1)	2.39 (14.2)
	Body width at anal fin origin	1.65 (9.9)	1.30 (7.7)
	Body depth at pelvic fin origin	2.73 (16.4)	2.50 (14.9)
	Body width at pelvic fin origin	2.91 (17.6)	2.59 (15.4)
	Caudal fin length (longest ray)	4.22 (25.4)	*
	Caudal peduncle depth	1.61 (9.7)	1.38 (8.2)
	Caudal peduncle length	3.79 (22.8)	3.91 (23.3)
	Eye diameter (horizontal)	1.36 (8.2)	1.27 (7.6)
	Eye diameter (vertical)	1.06 (6.4)	0.88 (5.2)
	First dorsal fin base	1.72 (10.4)	1.90 (11.3)
	Head length	5.34 (32.2)	5.45 (32.5)
	Head width	3.22 (19.4)	3.30 (19.7)
	Pectoral fin length	4.87 (29.3)	4.50 (26.8)
	Pelvic fin length	3.50 (21.1)	3.91 (23.3)
	Pelvic to anus	4.45 (26.8)	4.80 (28.6)
	Second dorsal fin base	3.38 (20.4)	3.28 (19.5)
	Second dorsal fin spine length	1.32 (8.0)	*
	Snout to origin of first dorsal fin	6.53 (39.4)	6.23 (37.1)
	Snout to origin of second dorsal fin	9.59 (57.9)	9.69 (57.7)
	Snout to vertical of anal fin origin	10.4 (62.8)	10.32 (61.5)
	Snout to vertical of pelvic fin origin	5.75 (34.7)	5.80 (34.5)
	1st spine length of 1st dorsal fin	2.41 (14.5)	*
	2nd spine length of 1st dorsal fin	2.93 (17.7)	*
	3rd spine length of 1st dorsal fin	2.64 15.9)	*
	4th spine length of first dorsal fin	2.17 (13.1)	*
	5th spine length of first dorsal fin	2.08 12.5)	*
	6th spine length of first dorsal fin	0.92 (5.5)	*
	1st ray length of second dorsal fin	1.88 (11.3)	1.83 (10.9)
% Caudal peduncle	Caudal Peduncle depth	1.61 (42.5)	1.38 (35.3)
length % Head length	Cheek depth	0.97 (18.2)	1.06 (19.5)
	Eye diameter	1.36 (25.5)	1.27 (23.3)
	Head depth	2.55 (47.8)	2.10 (38.5)
	Head width	(expanded head)	3.62 (66.4)
	Postorbital length	2.39 (44.8)	2.50 (45.9)
	Snout length	1.43 (26.8)	1.35 (24.8)
% Eye length	Interorbital width	0.24 (17.7)	0.33 (26.0)
	Snout length	1.43 (105.2)	1.35 (106.3)
% Pelvic to anus	Pelvic fin length	3.5 (78.7)	5.91 (122.6)

46865; female); absolute values and % values in brackets. An asterik (*) indicates a damaged character.