Gymnura micrura (Bloch & Schneider 1801) Smooth (or Lesser) Butterfly Ray (Figs. 1–10, 34a, 35, 36a – 38a, 39–41, 42a – 44a, 45, 46a – 48a; Tabs. 1–2) Raja micrura Bloch & Schneider, 1801: 360 (original description, type locality: Suriname) ? Trygon micrura: Müller, 1837: 40 (not seen: Rio de Janeiro, according to Bigelow & Schroeder [1953] identification probable because said to agree with description of Raja micrura of Bloch & Schneider 1801). Pteroplatea maclura: Puyo, 1936: 79, 250 (French Guiana; not seen). Pteroplatea maclura (in part): Müller & Henle, 1841: 169 (description, specimens from Suriname and Brazil only); Duméril, 1865: 614 (description, specimens from Brazil only, size probably erroneous as the New York specimen [= G. lessae, sp. nov.] recorded with a width of 2 meters is the same specimen analyzed by Müller & Henle, 1841, who had earlier characterized it as small); Günther, 1870: 487 (description, specimen from Brazil only). Pteroplatea micrura: Metzelaar, 1919: 8, 199 (listed, Trinidad, perhaps Curaçao). Pteroplatea micrura (in part): Meek & Hildebrand, 1923: 87 (not present in Panamá, color description matching G. lessae, sp. nov.). Gymnura micrura: Nishida, 1990: 65, 66, 83, 84, fig. 59B (skeletal morphology, clasper;?Suriname); Lovejoy, 1996: 214, 215, 223, 224, 227, 228, 230, 236, 242, 256, fig 7E (lateral line canals, skeletal morphology; Guianas and Suriname); Mazzoleni & Schwingel, 1999: 144, 116 (listed, Southern Brazil); Menni & Stehmann, 2000: 92 (distribution, South America); Furtado Jr. et al. 2003: 8, 9 (fisheries, North Brazil); Carvalho et al., 2004: 10, 33, 38, 39, 74, 79, 82, 86, 87, figs. 15, 31, 35B, 36C-B, 38C, 39A-B (skeletal morphology, Suriname); Meneses et al. 2005: 80, 82 (listed; Sergipe, Brazil); Nunes et al. 2005: 51, 52, 53 (inventory; Maranhão, Brazil); Yokota & Lessa, 2006: 349 –360 (nursery area, Northeastern Brazil); Yokota & Lessa, 2007: 249 –257 (reproductive biology, Northeastern Brazil); Basílio et al. 2008: 67 – 69 (checklist; Curu River estuary, Ceará, Brazil); Lessa et al. 2008 (fisheries, Northeastern Brazil); Basilio et al. 2009: 41 (listed; Ceará, Brazil); Bornatowski et al. 2009: 3 (listed; Southern Brazil); Nunes & Piorski, 2009: 479 –482 (presence dorsal fin; Maranhão, Brazil); Reis et al. 2012: 217 –219 (albinism; Alagoas, Brazil); Yokota et al., 2012: 1315 –1326 (reproductive biology, Northeastern Brazil); Ragno, 2013: 57, 58, 130, 177, fig. 46 (lateral line; Rio Grande do Norte, Brazil); Yokota et al., 2013: 1325 –1329 (diet, Northeastern Brazil); Fontenelle & Carvalho, 2015: 7 –9, figs. 8, 9E, 10E (brain morphology; Rio Grande do Norte, Brazil); Garcia Jr. et al., 2015: 4 (checklist; Rio Grande do Norte, Brazil); Willems et al., 2016: 36, 38, 40, 41 (fisheries, Suriname). Gymnura micrura (in part): Rosenberger, 2001: 615, 616, 618, 620, 621, 623–627 (systematics, specimen FMNH 89993 only [Suriname],?FMNH 89992); McEachran & Carvalho, 2002: 577 (identification guide, Western Central Atlantic; illustration is of G. lessae, sp. nov.). Not Raja maclura: Lesueur, 1817: 41, pl. not numbered (original description, Newport, Rhode Island, junior synonym of G. altavela). Not Pteroplatea micrura: Müller & Henle, 1841: 169 (Java, India, Red Sea; = Gymnura poecilura [Shaw, 1804]); Cantor, 1849: 1409 (= G. poecilura [Shaw, 1804]); Blyth, 1860: 29 (= G. poecilura [Shaw, 1804]); Day, 1865: 278 (= G. poecilura [Shaw, 1804]); Duméril, 1865: 613 (description, mouth of Ganges, = G. poecilura, [Shaw, 1804]); Günther, 1870: 487 (= G. poecilura [Shaw, 1804]); Day, 1878: pl. CXCIV, fig. 2 (= G. poecilura [Shaw, 1804]); Day, 1889: 56, fig. 23 (= G. poecilura [Shaw, 1804]); Wood-Mason & Alcock, 1891: 359 –367, pl. 7–8 (embryonic development, = G. poecilura [Shaw, 1804]); Alcock, 1892: 1 –8, pl. 4 (embryonic development, = G. poecilura [Shaw, 1804]); Annandale, 1909: 39 (= G. poecilura [Shaw, 1804]); Pillay, 1929: 353 (= G. poecilura [Shaw, 1804]). ?Not Gymnura sp. cf. micrura: Compagno & Last, 1999: 1507, 1509 (identification guide, Western Central Pacific); Randall & Lim, 2000: 583 (checklist, South China Sea); Jacobsen & Bennett, 2009: 1, 21, 22, 24, 25 (taxonomic revision, Indo-West Pacific). Not Gymnura micrura: Fowler, 1941: 455 (description based on a G. australis [Ramsay & Ogilby] specimen [USNM 39978]); Talwar & Kacker, 1984: 181 (= Gymnura poecilura [Shaw, 1804]); Raje, 2003: 89, 91–95 (probably equals Gymnura poecilura [Shaw, 1804]); Raje et al. 2007 (= Gymnura poecilura [Shaw, 1804]). Neotype. USNM 156714 (289 mm DW, adult male), Suriname, 6o20’45”N, 54o56’30”W, 0 to 26m, Vessel Coquette, station 145, trawl, 30.v.1957, accession num. 215120, USNM 440341 recatalogued from USNM 156714 (Fig. 1). Designated herein. Diagnosis. A small to medium size butterfly ray occurring in the Western South Atlantic distinguished from congeners (except the two new species described in this study) by the combination of the following characters: absence of spiracular tentacle and caudal stings; dorsal fin usually absent (a small dorsal fin may be present in some males); tail relatively short (mean post-cloacal length 22% DW) and prominently banded, presenting 3 to 5 black bands that may be less distinct in large adults. Gymnura micrura is distinguished from G. lessae, sp. nov. and G. sereti, sp. nov. by its usually uniformly brown or gray dorsal disc, without any vermiculate pattern (vs. dorsal side brownish, usually with a vermiculate pattern in both new species). Gymnura micrura may be further distinguished from G. sereti, sp. nov. by the following characters: clasper of mature males more slender and longer, Lclasper 9.3–11% DW (vs. clasper stouter and shorter, Lclasper 6.8–9.2% DW); cranial fontanelle U-shaped (vs. keyhole-shaped fontanelle); mesopterygium divided into one anterior solid element and 5–7 smaller fragments (vs. mesopterygium divided into two solid elements); distance between anteroventral (AVF) and posteroventral (PVF) fenestrae of scapulocoracoid representing 30–35% of scapulocoracoid length (vs. distance between AVF and PVF representing about 20% of scapulocoracoid length); lateral projection of the base of synarcual starting at its anterior third (vs. lateral projection of the base of synarcual starting synarcual half-length); thicker proximal portion of Meckel’s cartilage representing 28–40% of Meckel’s cartilage width (vs. 10–24%); lower number of diplospondylous vertebrae (mean 98 vs. 110, respectively), and higher number of radials associated to the mesopterygial fragments compared to the second mesopterygial element of G. sereti (range 15–17 vs. 10–15, respectively). Gymnura micrura is most similar to G. lessae, sp. nov., but may be further differentiated from it by the following characters: dorsal contour of hyomandibula with two conspicuous humps (vs. dorsal contour of the hyomandibula with a conspicuous proximal protuberance followed by an inconspicuous distal protuberance); projection of the anteroventral fenestra of coracoid bar in the form of a funnel with a large opening and closing (vs. projection of the anteroventral fenestra with a relatively narrower opening and end) and lower number of pectoral radials (range 112–119 vs. 118–127, respectively). Other diagnostic differences are described below. Description. Measurements are presented in Table 1; meristic data in Table 2. The following description is based on all specimens examined. Throughout the text the proportions are presented as: minimum value–maximum value (mean). See Figs. 1–4, 5a–h, 6a and 7–9 for external morphology and color, Fig. 34a for ventral lateral-line pattern, Figs. 5i, 6b, 35, 36a – 38a, 39–41, 42a – 44a, 45 and 46a – 48a for skeletal morphology and Fig. 10 for geographical distribution. External morphology. Disc lozenge shaped, 1.54–1.92 (1.75) times broader than long [1.54–1.72 (1.64) for adult males; 1.75–1.92 (1.81) for adult females]. Trunk strongly flattened, slightly raised above scapular region and posterior head. Snout relatively short and obtuse with a subtle lobe at snout tip. Sexual dimorphism evident in disc shape, mainly between adults, with adult males presenting proportionally longer snouts with smaller angles, which confers a triangular shape to the anterior half of the disc (Fig. 2). Preoral snout length 8.6–14 (11)% DW [8–11 (9)% DW in adult females; 12–14 (13)% DW in adult males], preorbital snout length 6.1–13 (9.4)% DW [7–10 (8)% DW in adult females; 10–13 (11)% DW in adult males], preorbital snout width 31–40 (37)% DW, postspiracle snout width 42–54 (49)% DW. Anterior margins of disc with a slight medial concavity and becoming weakly convex towards extremities (in adult males these curves may be less noticeable, with the anterior margins almost straight in some males) (Fig. 2); pectoral-fin apices acutely angular (sometimes moderately angular in adult males); posterior margins weakly convex; free rear tip broadly rounded; axis of greatest width positioned posteriorly to one-half disc length. Pelvic fin single lobed, rectangular, their corners rounded (Fig. 3). Skin entirely smooth, without denticles on dorsal and ventral surface of the disc. Interorbital space broad, interorbital width 7.9–9.8 (8.7)% DW. Eyes dorsolateral, small, oval and protruding slightly; eye diameter 1.2–2.2 (1.7)% DW, representing 13–25 (19)% of interorbital width; orbit diameter 1.9–3.5 (2.5)% DW, representing 59–108 (86)% of spiracle length. Eyes more protruded and relatively larger in embryos. Spiracles immediately following the eyes, relatively large, lozenge-shaped, contour of the inner spiracular margin concave (Fig. 4a); spiracle length 2.1–4 (2.9)% DW, 1.1–2.7 (1.8) times eye diameter; its inner posterior margin without tentacles. Ventral head length 20–27 (23)% DW. Nostrils narrowly oval, diagonally directed, only the circular distal margin not covered by nasal curtain, posterior lateral margin with lobe (Fig. 4c); nostril length 1.8–4.3 (3.2)% DW [2.5–3.4 (2.9)% DW in adult females; 3.1–4.3 (3.7)% DW in adult males], 1.0–3.0 (1.8) times internasal width; internasal width 1.1–2.7 (1.8)% DW. Anterior nasal flaps medially expanded and fused into a broad, skirt-shaped, posteriorly expanded nasal curtain that covers the internasal space and reaches mouth (Fig. 4b). Nasal curtain skirtshaped, weakly to moderately bilobed, wide, its width 1.9–4.3 (3.0) times length; lateral margin concave; posterolateral apices rounded; posterior margin straight to weakly concave, a small gap may be present medially. Mouth relatively wide, its width 7.5–10 (8.5)% DW, 32–45 (38)% head length, 0.9–1.8 (1.2) times nasal curtain width; without papillae on floor or labial folds, although some striations may originate radially from mouth corners; lower lip arched rearward toward corners, uniformly convex or weakly to moderately concave along medial region, without any lump or knob (Fig. 4b); a strip of corrugated skin forms a half-circle below mouth; strips ends at the mouth corners. Small, numerous and closely crowded teeth in bands; teeth with one medial, pointed cusp directed towards inside of mouth; tooth base somewhat swollen, entirely twisted to the opposite direction; in labial face, tooth resembling a three-armed boomerang (Fig. 4d); teeth similar between jaws and sexes; their number increasing with growth. Gill slits moderately S-shaped, first four gill slits markedly larger than fifth (Fig. 4b). Distance between gill slits decreasing posteriorly; distance between inner ends of fifth pair 60–74 (67)% distance between inner ends of first pair; distance between inner ends of first pair 15–18 (16)% DW, 58–82 (72)% ventral head length, 6.3–14.0 (9.3) times internasal width; distance between inner ends of fifth pair 10–12 (11)% DW, 40–56 (48)% ventral head length, 4.1–9.3 (6.3) times internasal width. Tail slender and short, whip-like, cross-banded (see section on color), tapering toward tip, without caudal stings at any age (Fig. 9); postcloacal tail 17–30 (22)% DW, 30–47 (38)% DL, 30–56 (45)% precloacal length; tail usually lacking dorsal fin, however some males (about 25% of males) may present a small plesodic dorsal fin at tail base with varying degrees of development (Figs. 5a–h); when present the dorsal fin is situated at the level of free rear tips of pelvic fin, distance from snout tip to anterior base of the dorsal fin representing 76–80 (78)% of total length; low longitudinal skin folds present on dorsal and ventral surfaces of tail, skin folds not reaching the tail tip. Clasper of mature males cylindrical, somewhat depressed, relatively slender and longer (compared to G. sereti, sp. nov.) (Fig. 6); distance from posterior margin of cloaca to clasper tip 9.3–11.0 (9.9)% DW; clasper tapering slightly distally; clasper tip conical, bluntly pointed to pointed, not calcified; portion not calcified relatively shorter than in G. lessae, sp. nov. and G. sereti, sp. nov.; apopyle (on the anterior dorsal surface) connected to hypopyle by a long, dorsomedial, posteriorly curved clasper groove; spermatic duct almost reaching clasper tip; rhipidion and pseudorhipidion (“small flap” of Nishida 1990) absent; a long ventral pseudosiphon (“SAC 1” of Nishida 1990) laterodistally situated to hypopyle; a well-developed dorsal pseudosiphon (“SAC 2” of Nishida 1990) on inner margin of clasper; dorsal pseudosiphon more posteriorly located when compared to G. sereti, sp. nov.; ventral surface of clasper entirely smooth. Coloration. Dorsal side usually uniformly light to dark brown, grayish, sometimes slightly roseate, not vermiculated (Figs. 7a, 8); irregular dark patches or irregular dark round spots may be present, especially in freshly caught specimens (Figs. 7b, c). Living specimens may be variable in color according to the environment (Figs. 7d, e), but they tend to become uniformly brown or gray when preserved. Ventral surface whitish to brownish, creamy, slightly roseate, yellowish or copper, generally darkening toward edges. Tail prominently cross-banded, with 3–5 black bands (Fig. 9a); bands diffuse or less distinct in large specimens (Fig. 9b). Size, reproduction, diet and habitat. Size at birth ranging from 135 to 185 mm DW; size at 50% maturity was estimated at 269 (265–273) mm and 405 (401–409) mm for males and females, respectively (Yokota et al. 2012). The largest male analyzed was 360 mm DW (MZUSP 9926, from Espírito Santo, Brazil), and males of this species shall reach a maximum of 450 mm DW. The largest female analyzed was 629 mm DW (sampled in Rio Grande do Norte, Brazil, by the first author) and the females reach around 800 mm DW. This species is matrotrophic viviparous (lipid histotrophy) and appears to be reproductively active throughout the year, producing 1–6 young per litter (Yokota et al. 2012). Diet consists mainly of bony fishes with evidence of feeding specialization, ingesting intermittently proportionally larger fishes (Yokota et al. 2013). This is a demersal species living along the coast on sandy and muddy bottoms, but also occurs in estuaries. Geographical distribution. According to material examined, G. micrura is distributed in the Atlantic coast of South America from Venezuela (including Trinidad & Tobago) to Rio de Janeiro State, Brazil (Fig. 10). The species is also registered in São Paulo and Paraná States, Brazil (Gonzalez 1995, P. Charvet, pers. comm.). The species is relatively common in the north of South America and northern and northeastern regions of Brazil, with its abundance declining towards the south, where its occurrence may be related to seasonal warmer water. There are no confirmed records of this species in Colombia and Panama (Meek & Hildebrand 1923, Mejía-Falla et al. 2007), and it is not known if the species occurs in the Caribbean Sea, beyond the Venezuelan coast. Remarks. The original description of Gymnura micrura (as Raja micrura Bloch & Schneider 1801) is brief, nondiagnostic, presenting characters that are commonly shared by all species in the genus, and is not illustrated; however, the type locality is clearly stated as Suriname (“Habitat in Surinamo”, p. 360). This fixed the name micrura to the butterfly ray without caudal stings and spiracular tentacles commonly found in the region, in contrast to G. altavela (Linnaeus 1758), which possesses spiracular tentacles and caudal stings and does not occur in the area. Bloch & Schneider mention that type material would probably be housed in the “Museo Vaillanti Parisiis”. After an exhaustive search in the Muséum national d'Histoire naturelle in Paris and other collections, we are not able to locate any type material for G. micrura. At the same time, this type material was never mentioned in any study subsequent to the original description (e.g. Séret & McEachran 1986) and is considered to be unknown in Eschmeyer et al. (2017); the type material was probably lost or destroyed. As this species is central to a complex taxonomic problem, in which three distinct species were being identified under the name G. micrura (two of these are described as new here), the designation of a neotype for this species is expressly required to define the nominal taxon objectively and clarify its taxonomic status. The designed neotype matches the available information from the original type locality. Fowler (1941) characterized the type locality of Bloch & Schneider as “likely erroneous” but without further comment. This author probably made this comment because the species he named G. micrura is not that of Bloch & Schneider, but a misidentification of Gymnura from the Indian Ocean and Indo-Pacific region. His description of G. micrura, for example, is based on a specimen of G. australis (Ramsey & Ogilby 1886) (USNM 39978). Hence, a type locality in South America for a species from the Indo-Pacific region is erroneous. Müller & Henle (1841) mistakenly applied the specific name micrura to G. poecilura of Shaw (1804), and were followed by many others authors (e.g. Cantor 1849, Blyth 1860, Day 1865, Duméril 1864, Günther 1870, Day 1878, 1889, Wood-Mason & Alcoc, Published as part of Carvalho, Marcelo Rodrigues De, 2017, Taxonomic and morphological revision of butterfly rays of the Gymnura micrura (Bloch & Schneider 1801) species complex, with the description of two new species (Myliobatiformes: Gymnuridae), pp. 1-74 in Zootaxa 4332 (1) on pages 6-19, DOI: 10.11646/zootaxa.4332.1.1, http://zenodo.org/record/1052041, {"references":["Bloch, M. E. & Schneider, J. G. (1801) M. E. Blochii, Systema Ichthyologiae Iconibus cx Ilustratum. Post obitum auctoris opus inchoatum absolvit, correxit, interpolavit Jo. Gottlob Schneider, Saxo. Sumtibus Auctoris Impressum et Bibliopolio Sanderiano Commissum, Berolini, 584 pp. [in Latin]","Bigelow, H. B. & Schroeder, W. C. 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