Psammobatis extenta (Garman, 1913)

Psammobatis extenta (Garman, 1913) View in CoL

Raja erinacea (non Mitchill). — Miranda Ribeiro, 1907:176, pl. XII, XIII ( Rio de Janeiro, Brazil; catalogue).

Raia extenta Garman, 1913:356 View in CoL (original description; type-locality: off the coast of Ilha Rasa, Rio de Janeiro, Brazil).

Psammobatis extenta View in CoL — Fowler, 1942:131 ( Brazil; listed). —Menni, 1972 (clasper morphology). —Carvalho, Figueiredo, 1994 (systematic study and lectotype selection). — McEachran, Dunn, 1998:286 (systematic study). — Mazzoleni , Schwingel , 1999:114 ( Itajaí , Santa Catarina, Brazil; listed). — Compagno , 1999:489 (listed). — Cousseau et al., 2000:30 ( Argentina and Uruguay; catalogue). — Menni , Stehmann , 2000:87 ( Argentina, Uruguay and Brazil; listed). — Muto et al., 2001 ( Ubatuba , São Paulo, Brazil; diet). — Paragó , 2001 (systematic study). — Gomes , 2002 (systematic study). — Braccini , Chiaramonte , 2002 ( Argentina; reproductive biology). — Braccini , Chiaramonte , 2005 ( Argentina; morphometry). — Gomes , Gadig , 2003:29 ( São Paulo, Brazil; listed). — Mabragaña , 2007 ( Argentina; biological and ecological study). — Cousseau et al., 2007:62 ( Mar del Plata , Argentina; catalogue). —Rocha et al., 2010 (egg capsules). — Weigmann , 2016:100 (listed). — Last et al., 2016:457 (listed). — Nión et al., 2016:23 ( Uruguay; listed). — Viana et al., 2017:3 ( Rio de Janeiro, Brazil; diet). — Cordeiro , Oddone , 2019:47 ( Rio Grande do Sul, Brazil; reproductive biology). — Gomes et al., 2019:299, fig. 291 ( Rio de Janeiro, Brazil; catalogue). — Brum-Neto , Lucena , 2020 ( Brazil; tooth morphology). — Sabadin et al., 2020:1901 ( Argentina, Uruguay and Brazil; listed). — Mabragaña , Cousseau , 2021:64 ( Argentina; listed). — Hoff et al., 2023 ( Brazil; listed) .

Psammobatis sp. —Figueiredo, 1977:34, fig. 75 (Southern Brazil; catalogue).

Psammobatis glansdissimilis —McEachran, 1983:67, fig. 18 (systematic study). —Mazzoleni, Schwingel, 1999 (Itajaí, Santa Catarina, Brazil; listed). —Menni, Stehmann, 2000:87 ( Argentina, Uruguay and Brazil; listed).

Diagnosis. Psammobatis extenta is distinguished from P. lentiginosa and P. rutrum by the dark brown coloration pattern of tiny dark spots ( vs. black in P. lentiginosa and whitish in P. rutrum ); tiny spots distributed around the orbits ( vs. absent in P. lentiginosa and concentrated in the preorbital region in P. rutrum ); presence of thorns on pectoral fins ( vs. reduced or absent in P. lentiginosa ); pectoral thorns with tip oriented towards the caudal region ( vs. tip oriented towards the lateral margin of disc in P. rutrum ); presence of a mid-pectoral thorn ( vs. absent in P. lentiginosa and P. rutrum ); continuous rows of dorsal thorns posterior to the scapular region ( vs. interruption only from the middorsal to the pelvic insertion in P. lentiginosa and interruption of the three rows in P. rutrum ); spermatic groove of claspers positioned dorsally with dermal denticles ( vs. lateral spermatic groove and denticles absent in P. lentiginosa and P. rutrum ); presence of ocelli on the posterior region of pectoral fins in adults and juveniles ( vs. absent in P. lentiginosa and P. rutrum ); anterior fontanelle rectangular ( vs. trapezoidal in P. rutrum and P. lentiginosa ); basal fenestrae rectangular and narrow ( vs. kidney-shaped in P. rutrum and P. lentiginosa ); posterior nasal cartilage perpendicular in relation the anteroposterior axis ( vs. oblique in relation the anteroposterior axis in P. rutrum ); dorsal terminal cartilage 2 and accessory terminal cartilage 2 absent ( vs. present in P. lentiginosa and P. rutrum ); accessory terminal cartilage 1 continuous with the ventral marginal cartilage ( vs. articulated in P. lentiginosa and P. rutrum ).

External description. Disc 1.4–1.5 times wider than long ( Tab. 1). Heart-shaped disc in juvenile and adult specimens; subtle in the former and accentuated in the latter. Adults with wide pectoral fins and convex margins in the posterior region, while the anterior region narrows towards the snout. Adult males with concave anterior disc margin close to the orbital region; disc width at eye line in females is 1.2 times longer than in males.

Preorbital length 3.0–3.6 times the eye horizontal diameter and 2.5–2.6 times the interorbital distance. Preoral length 1.4–1.6 times the mouth width. Upper jaw arched, especially in middle region; lower jaw convex. First pair of gill openings 1.3 times larger than the fifth. Distance between first gill slits 1.8–2.2 times the distance between the fifth.

Pelvic fins convex externally with a notch forming anterior and posterior lobes. Pelvic anterior lobe shorter with a convex anterior margin; posterior margin distinguished by radial tips and continuously connected to the lateral margin of posterior lobe. Posterior lobe with a convex lateral margin and a straight inner margin. Dorsal fins similar in size and shape, with rounded apex. Distance from pelvic posterior margin to origin of first dorsal fin 2.2 times the distance from first dorsal origin to posterior margin of caudal fin. Tail long, 1.7–1.8 times in total length and 1.4 times the snout-vent distance. Caudal fin rounded.

Coloration in alcohol. Dorsal coloration dark brown, its intensity varying according to development stage ( Fig. 3A). In adults, a grouping of tiny dark spots distributed over the dorsal surface produces a reticulated appearance, contributing to the darkening of this region. Spots can still form circular borders, smaller or equal to the eye diameter. Preorbital region with a lighter triangular area contrasting to the predominant background color and tiny spots distributed around the orbits, projecting towards disc margin ( Fig. 4A). An ocellus is observed near the posterior region of each pectoral fin in both adults and juveniles ( Figs. 3A, 5A). In juveniles, a dark spot located in the central region of each pectoral fin and a cluster of dark spots located on the posterior region of each pectoral fin as in P. rutrum , but spots can be less intensely colored ( Figs. 3E, 5A). Some specimens present a black spot on the snout tip as in P. lentiginosa and P. rutrum ( Figs. 3A, C, E). Tail region presenting the same predominant background color, but also with lighter bands formed by the union of spots on dorsal region. In adults, some bands form an hourglass. Ventral region uniformly cream ( Fig. 3B).

Dermal denticles. Rostral and malar thorns forming rows with no differences between sexes; star-shaped base and a curved crown that narrows towards its distal end ( Figs. 6C–D). Rows of alar thorns of adult males situated posterior to the malars and positioned horizontally, close to pectoral lateral margin, with distal end oriented medially; elongated base and a thorn-shaped crown, arranged in two to six alternating rows, five to 20 denticles ( Fig. 6A; Tab. 2).

Head with two to five preorbital thorns, none to three midorbital thorns, none to two postorbital thorns, and two to four spiracular thorns. Thorns similar in structure to the nuchal and scapular but smaller. Nuchal and scapular thorns forming a triangular grouping, organized into one to three nuchal thorns and five to 21 scapular thorns. Scapular thorns more concentrated in the central region, resulting in a smaller base for the triangle; leaf-shaped base with shallow grooves and a curved crown with an extremely sharp distal end ( Figs. 6E–F).

Mid-pectoral thorns located on the medial region of each pectoral fin. Stellate base with well-defined grooves between the dorsal ridges, a slightly curved crown, and a tapered distal end with a straight orientation; occurring individually or grouped into two to four thorns ( Figs. 7A–B). Smaller, cross-shaped dermal denticles with a tapered distal end around the mid-pectoral thorns, giving a rough texture to this region. Posterior pectoral thorns present in both males and females; cross-shaped or star-shaped base, with a curved crown and tapered distal end ( Fig. 6B).

Well-developed dermal denticles located along the margin of clasper spermatic groove ( Fig. 8A). Middorsal row of thorns presenting or not a small interruption, while dorsolateral rows extend to the tail region, although there is considerable individual variation in this pattern ( Fig. 4D). Middorsal and dorsolateral thorns similar in structure to the nuchal and scapular ones. Smaller denticles present posteriorly to the scapula and adjacent to thorn rows, giving a rough texture to this region. Cross-shaped or plateaulike bases and crown curved with a tapered distal end.

One row of mid-caudal thorns and two caudo-lateral thorns, one at each side of the tail, all oriented anteroposteriorly; denticles arranged randomly between rows. Caudo-lateral thorns with base and crown flattened dorsolaterally. Mid-caudal thorns similar in structure to middorsal thorns and forming a continuous row with them, 26 to 39 denticles ( Tab. 2).

Geographical distribution. Examined specimens with a wide geographical distribution, extending along the coasts of Rio de Janeiro, São Paulo, Paraná, and Santa Catarina ( Fig. 9). This species is also recorded in the Gulf of San Jorge, Argentina (Mabragaña, 2007).

Material examined. MNRJ 12317 View Materials , lectotype . MNRJ 12318 View Materials , paralectotype , 4, off the coast of Ilha Rasa , Rio de Janeiro, Brazil, 23°S 43°W. Santa Catarina GoogleMaps : DBAV. UERJ 1545 *, male, 241 mm TL, 110 mm DW, Itajaí . DBAV. UERJ 1624.7 , male, 190 mm TL, 94.5 mm DW, DBAV. UERJ 1624.11 , female, 196 mm TL, 111 mm DW, DBAV. UERJ 1624.12 , male, 180 mm TL, 94 mm DW, DBAV. UERJ 1624.14 , female, 180 mm TL, 107 mm DW, DBAV. UERJ 1624.16 , male, 200 mm TL, 111 mm DW, DBAV. UERJ 1624.17 *, male, 184 mm TL, 107 mm DW, DBAV. UERJ 1624.18 , female, 230 mm TL, 126 mm DW, DBAV. UERJ 1624.19 , female, 220 mm TL, 118 mm DW, DBAV. UERJ 1624.20 , male, 225 mm TL, 127 mm DW, São Francisco do Sul. São Paulo : DBAV. UERJ 883 , female, 256 mm TL, 144 mm DW, between Paranaguá, Paraná and Ilha do Bom Abrigo, São Paulo . DBAV. UERJ 1623.1 , male, 210 mm TL, 120 mm DW, DBAV. UERJ 1623.2 , male, 225 mm TL, 138 mm DW, Santos . DBAV. UERJ 1627.4 , female, 267 mm TL, 135 mm DW, Santos . DBAV. UERJ 1720.1 , female, 242 mm TL, 135 mm DW, DBAV. UERJ 1720.2 , male, 185 mm TL, 106 mm DW, DBAV. UERJ 1720.3 , female, 178 mm TL, 87 mm DW, DBAV. UERJ 1720.5 , female, 154 mm TL, 82 mm DW, DBAV. UERJ 1720.6 , female, 235 mm TL, 133 mm DW, Arquipélago de Alcatrazes . DBAV. UERJ 1868 , male, 220 mm TL, 118 mm DW, Santos . DBAV. UERJ 1871.1 , male, 84 mm TL, 44 mm DW, DBAV. UERJ 1871.2 , female, 272 mm TL, 148 mm DW, between Paranaguá, Paraná and Ilha do Bom Abrigo , São Paulo . DBAV. UERJ 1874.3 , male, 255 mm TL, 135 mm DW, DBAV. UERJ 1874.4 , male, 251 mm TL, 140 mm DW, Ubatuba . DBAV. UERJ 1885.9 *, female, 200 mm TL, 115 mm DW, Barra de Santos . DBAV. UERJ 1886.1 , female, 195 mm TL, 110 mm DW, Barra de Santos . DBAV. UERJ 1886.8 , male, 202 mm TL, 120 mm DW, Barra de Santos. Rio de Janeiro : DBAV. UERJ 759 , female, 176 mm TL, 100 mm DW, Maricá . DBAV. UERJ 763 , female, 176 mm TL, 100 mm DW, Ilha Grande . DBAV. UERJ 2124 , female, 263 mm TL, 145 mm DW, DBAV. UERJ 2125 , female, 261 mm TL, 138 mm DW, DBAV. UERJ 2126 , female, 247 mm TL, 138 mm DW, DBAV. UERJ 2127 , male, 258 mm TL, 140 mm DW, DBAV. UERJ 2128 , female, 231 mm TL, 134 mm DW, DBAV. UERJ 2129 , female, 206 mm TL, 115 mm DW, DBAV. UERJ 2139 , female, 262 mm TL, 135 mm DW, Angra dos Reis . DBAV. UERJ 2144.2 *, female, 275 mm TL, 165 mm DW, 23º27.4’S 44º13.4’W GoogleMaps . DBAV. UERJ 2146.8 , female, 240 mm TL, 149 mm DW, Angra dos Reis . MNRJ 14159 View Materials , male, 254 mm TL, 141 mm DW, Ilha Feia . MNRJ 17763 View Materials , 2 View Materials , 246–253 mm TL, 143–153 mm DW, Macaé . MNRJ 32323 View Materials , 5 View Materials , 125.8 View Materials – 267 mm TL, 77–152 mm DW, Rio de Janeiro. MNRJ 32468 View Materials , 3 View Materials , 233–262 mm TL, 129.85–141.5 mm DW, Rio de Janeiro. MNRJ 32595 View Materials , 2 View Materials , 107.95 View Materials – 141.9 mm TL, 61.85–77.65 mm DW, Rio de Janeiro. MNRJ 33103 View Materials , 3 View Materials , 225–245 mm TL, 129–140 mm DW, between Ilha Rasa and Marambaia. MNRJ 49365 View Materials , male, 235 mm TL, 115 mm DW, Armação dos Búzios . MNRJ 49227 View Materials , male, 245 mm TL, 131.5 mm DW, Armação dos Búzios. Locality undetermined : DBAV. UERJ 1645 , male, 245 mm TL, 137 mm DW. DBAV. UERJ uncataloged*, male, 262 mm TL, 144.7 mm DW.