Gymnura lessae Carvalho 2017, sp. nov.
Gymnura lessae , sp. nov. (Figs. 11–20, 34b, 36b – 38b, 42b – 44b, 46b – 48b; Tabs. 3–4) Pteroplatea maclura : Jordan & Gilbert, 1883: 46 (description, North America, distributional range including G. micrura ); Jordan & Evermann, 1896: 86 (description, North and Middle America, distribution...
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Online Access: | https://dx.doi.org/10.5281/zenodo.6039807 https://zenodo.org/record/6039807 |
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Biodiversity Taxonomy Animalia Chordata Elasmobranchii Myliobatiformes Gymnuridae Gymnura Gymnura lessae |
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Biodiversity Taxonomy Animalia Chordata Elasmobranchii Myliobatiformes Gymnuridae Gymnura Gymnura lessae Carvalho, Marcelo Rodrigues De Gymnura lessae Carvalho 2017, sp. nov. |
topic_facet |
Biodiversity Taxonomy Animalia Chordata Elasmobranchii Myliobatiformes Gymnuridae Gymnura Gymnura lessae |
description |
Gymnura lessae , sp. nov. (Figs. 11–20, 34b, 36b – 38b, 42b – 44b, 46b – 48b; Tabs. 3–4) Pteroplatea maclura : Jordan & Gilbert, 1883: 46 (description, North America, distributional range including G. micrura ); Jordan & Evermann, 1896: 86 (description, North and Middle America, distributional range including G. micrura ); Linton, 1905: 348 (stomach contents, Beaufort, N. Carolina); Smith, 1907: 45 (description, North Carolina, maximum size probably wrong); Wilson, 1908: 626 (parasites, North Carolina); Weymouth, 1910: 135 (Louisiana); Coles, 1915: 94 (C. Lookout, North Carolina); Nichols & Breder, 1927: 27 (Sandy Hook Bay and Woods Hole). Pteroplatea maclura (in part): Müller & Henle, 1841: 169 (description, only specimen(s) from New York); Duméril, 1865: 614 (description, only specimens from USA, size probably erroneous as the New York specimen recorded with a width of 2 meters is the same analyzed by Müller & Henle 1841, who had earlier termed it as small); Günther, 1870: 487 (description, only specimen from Texas). Pteroplatea micrura : Garman, 1913: 414, pl. 33, figs. 3–4 (description, color, illustration); Fowler, 1920: 146 (listed, C. May County, New Jersey); Hildebrand & Schroeder, 1928: 67, fig. 37 (description, Chesapeake Bay); Breder, 1931: 39 (listed, Sandy Hook Bay, New Jersey); Fowler, 1931: 46 (listed, Port Aransas, Texas); Beltran, 1934: 10 (listed, Mexico, not seen); Gunter, 1935: 39 (listed, Louisiana & Gulf of Mexico); Fowler, 1936: 132 (West Africa references for G. sereti , sp. nov., description of mid-Atlantic U.S. specimen); Gunter, 1941: 196 (listed, Louisiana). Pteroplatea micrura (in part): Meek & Hildebrand, 1923: 87 (no Panama record, color description matching G. lessae , nov. sp., probable area of occurrence of G. micrura ). Gymnura micrura : Bigelow & Schroeder, 1953: 408 –416, figs. 97, 98 (description, illustrations, U.S. [Gulf of Mexico and North Coast] and Mexico); Joseph & Yerger, 1956: 119 (biological notes, Alligator Harbor, Florida); Daiber & Booth, 1960: 138 –139 (biological notes, Delaware and Chincoteague Bays); Schwartz, 1961: 390 (size, embryos, listed, Chincoteague & Sinepuxent Bays); Hoese, 1962: 170 –171 (occurrence, Virginia’s Seaside); Snelson & Williams, 1981: 6, 8 (biological notes, Florida); Schmid et al . 1988: 123 –124 (biological notes, Florida); Wourms, 1993: 271, 283 (embryonic development, South Carolina); Maruska, 2001: 52, 55–57, 59, 62, figs. 6, 9b (lateral line canal system); Rosenberger, 2001b (locomotion, pectoral fin); González-Isáis, 2003: 260, 262, 264–266, 269–271, figs. 2a, 5 (cephalic musculature; Veracruz, Mexico); Wigley et al . 2003: 7, 17 (length-weight relationships, Northeast Coast of the U.S.); González-Isáis & Domínguez, 2004: 518, 521–524, 527, 528, 530, 532, tab. 1, figs. 3a, 4b, 5b, 6b, 7a, 9a (skeletal morphology, cephalic musculature; Veracruz, Mexico); Kobelkowsky, 2004: 185 –192, figs. 1–3 (morphology of urogenital system; Veracruz, Mexico); McEachran & Aschliman, 2004: 87, fig. 3.8 (clasper; Florida, Gulf of Mexico); Shepherd & Myers, 2005: 1097 – 1100, 1102 (population decline, Gulf of Mexico); Smith et al. , 2009: 761 –763, 765, 766, 768, 772–778 (morphometry, molecular data); Schwartz, 2011: 35 –36 (distributional pattern, North Carolina); Kobelkowsky, 2012: 36, 38–41 (heart anatomy, Mexico); Cu-Salazar et al. 2014: 109 –117 (reproductive biology; Tabasco and Campeche, Mexico). Gymnura micrura (in part): McEachran et al . 1996: 67 –68, 70–75, 83 (lateral line, skeletal morphology, cephalic and branchial musculature; only specimen TCWC 642.08 [Florida, Gulf of Mexico],? TCWC [uncataloged specimen]); Rosenberger, 2001a: 615, 616, 618, 620, 621, 623–627 (in systematics analysis; only specimen FMNH 46250 [Florida, Gulf of Mexico] and probably FMNH 40363 [Dominican Republic],? FMNH 89992); McEachran & Carvalho, 2002: 577 (identification guide, illustration, Western Central Atlantic, G. micrura also included). Gymnura altavela : Fowler, 1945: 162 (embryos, South Carolina, Bigelow & Schroeder analyzed the specimens and found them to be G. micrura [= G. lessae , sp. nov.]). Holotype. USNM 51940 (420 mm DW, adult male), North Carolina, Beaufort, Collector: G. Bean, Accession num. 0 43551, USNM 440342 recatalogued from USNM 51940 (Fig. 11). Paratypes. USNM 440342 (4 specimens: adult male, 370 mm DW; adult male, 342 mm DW; juvenile female, 280 mm DW; skeleton of a dissected specimen), North Carolina, Beaufort, collector: G. Bean, accession num. 0 43551, date cat: 0 9 Feb 2017, recatalogued from USNM 51940 (Figs. 12–14) Diagnosis. A medium size butterfly ray occurring in the central and northwestern Atlantic Ocean. Gymnura lessae , sp. nov. may be distinguished from other species of butterfly rays (except G. micrura and G. sereti , sp. nov. ) by the combination of the following characters: absence of spiracular tentacle, caudal stings, and dorsal fin; tail relatively short (mean post-cloacal length 19% DW) and prominently banded, presenting 3 to 5 black bands that may be less distinct in large adults. Gymnura lessae , sp. nov. may be distinguished from G. micrura and G. sereti , sp. nov. by the dorsal contour of its hyomandibula with a conspicuous proximal protuberance followed by a inconspicuous distal protuberance ( vs . dorsal contour of hyomandibula with two conspicuous protuberances); projection of the anteroventral fenestra on the floor of the coracoid bar taking the form of a funnel with relatively narrower opening and end (in G. micrura this projection takes the shape of a funnel with large opening and closing; in G. sereti , sp. nov. the funnel has a large opening, but presents an abrupt tapering at its half length, with the end of the funnel relatively narrow), and higher number of pectoral radials (ranges 118–127 vs . 112–119 in G. micrura and 109–115 in G. sereti , sp. nov. ). Gymnura lessae , sp. nov. may be further distinguished from G. sereti , sp. nov. by the following characters: clasper of mature males more slender and longer, Lclasper 9.4–14.3% DW ( vs . clasper stouter and shorter, Lclasper 6.8–9.2% DW); cranial fontanelle U-shaped ( vs . keyhole-shaped); mesopterygium divided into one anterior solid element and 5–7 smaller fragments ( vs . mesopterygium divided into two solid elements); distance between anteroventral and posteroventral fenestrae of scapulocoracoid representing 35–40% of scapulocoracoid length ( vs . distance between AVF and PVF 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 at its half length); lower number of diplospondylous vertebrae (means 97 vs . 110), and higher number of radials in the mesopterygial fragments compared to the second mesopterygial element of G. sereti , sp. nov. (ranges 14–19 vs . 10–15). Gymnura lessae , sp. nov. is most similar to G. micrura , and may be further differentiated from it by its brownish dorsal side with a vermiculate background ( vs . dorsal side usually uniformly brown or gray without any vermiculate pattern). Description. Measurements are presented in Table 3; meristic data in Table 4. The following description is based on all specimens examined. Throughout the text the proportions are presented as: minimum value–maximum value (mean). See Figs. 11–17, 18a, and 19 and 20 for external morphology and color, Fig. 34b for ventral lateralline pattern and Figs. 18b, 36b – 38b, 42b – 44b and 46b – 48b for skeletal morphology. External morphology. Disc lozenge shaped, 1.54–1.94 (1.75) times broader than long [1.54–1.70 (1.61) for adult males; 1.71–1.89 (1.80) 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 their disc (Fig. 15). Preoral snout length 7.6–16 (12)% DW [8–11 (10)% DW in adult females; 13–16 (14)% DW in adult males], preorbital snout length 6.8–14 (9.7)% DW [7–9 (8)% DW in adult females; 10–14 (12)% DW in adult males], preorbital snout width 31–41 (36)% DW, postspiracle snout width 40–54 (48)% 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. 15); 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 disc half-length. Pelvic fin single lobed, rectangular, their corners rounded (Fig. 16). Skin entirely smooth, without denticles on the dorsal and ventral surface of the disc. Interorbital space broad, interorbital width 7.4–10 (8.6)% DW. Eyes dorsolateral, small, oval and protruding slightly (Fig. 17a); eye diameter 1.1–2.2 (1.7)% DW, representing 11.5–26.3 (19.5)% of interorbital width; orbit diameter 2–3.9 (2.8)% DW, representing 61–175 (96)% of spiracle length. Eyes more protruded and relatively larger in embryos. Spiracles immediately posterior to eyes, relatively large, lozenge-shaped, contour of the inner spiracular margin concave to rectilinear (Figs. 17b, c); spiracle length 2.1–3 (1.8)% DW, 1.1–3 (1.8) times eye diameter; its inner posterior margin without tentacles. Ventral head length 20–28 (23)% DW. Nostril narrowly oval, diagonally directed, only the circular distal margin not covered by nasal curtain, posterior lateral margin with lobe; nostril length 2.0–4.1 (3.2)% DW [2.7–3.4 (3.1)% DW in adult females; 3.1–4.1 (3.5)% DW in adult males], 0.9–2.7 (1.7) times internasal width; internasal width 1.4–3.1 (1.9)% DW. Anterior nasal flaps medially expanded and fused into a broad, skirt-shaped, posteriorly expanded nasal curtain that covers internasal space and reaches mouth (Fig. 17d). Nasal curtain skirt shaped, weakly to moderately bilobed, wide, its width 2.4–4.3 (3.1) times its length; lateral margin concave; posterolateral apices rounded; posterior margin straight to weakly concave, a small medial gap may be present. Mouth relatively wide, its width 7.9–11 (9.1)% DW, 30–48 (39)% head length, 0.9–1.7 (1.2) times nasal curtain width; without papillae on floor or labial folds, although some striations may originate radially from mouth corners (Fig. 17d); lower lip arched rearward toward corners, uniformly convex or weakly to moderately concave along medial region, without any lump or knob (Fig. 17d); a strip of corrugate skin forms a half circle below mouth, ending at mouth corners. Small, numerous and closely crowded teeth in bands; teeth similar to teeth in G. micrura , with one medial, pointed cusp directed towards inside of mouth and the base somewhat expanded, entirely twisted in opposite direction; in labial face, tooth resembling a three-armed boomerang; 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. 17d). Distance between gill slits decreasing posteriorly; distance between inner ends of fifth pair 62–79 (73)% distance between inner ends of first pair; distance between inner ends of first pair 15–19 (16) % DW, 57–84 (70)% ventral head length, 4.8–13.0 (8.5) times internasal width; distance between inner ends of fifth pair 10–13 (12)% DW, 40–60 (51)% ventral head length, 4.0–9.2 (6.2) times internasal width. Tail slender and short, whip-like, cross-banded (see color description), tapering toward tip, without caudal stings and dorsal fin (Figs. 20d, e); postcloacal tail 15–24 (19)% DW, 28–42 (34)% DL, 32–48 (39)% precloacal length; low longitudinal skin folds present on dorsal and ventral surfaces, skin folds not reaching tail tip. Clasper of mature males cylindrical, somewhat depressed, relatively slender and long (compared to G. sereti , sp. nov. ) (Fig. 18); distance from posterior margin of cloaca to clasper tip 9.4–14.3 (11.9)% DW; clasper tapering slightly distally; tip conical, bluntly pointed to pointed, not calcified; non-calcified portion relatively long compared to G. micrura , but shorter than in G. sereti , sp. nov. apopyle (on the anterior dorsal surface) connected to hypopyle by a long, dorsomedial, posteriorly curved clasper groove; spermatic duct ending before reaching clasper tip; rhipidion and pseudorhipidion absent; a long ventral pseudosiphon laterodistally situated to hypopyle; welldeveloped dorsal pseudosiphon 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 brownish, grayish or purplish, vermiculated. The vermiculate pattern varying from less conspicuous with lighter and darker shades to a conspicuous pattern where the lighter areas become larger and more remarkable (Figs. 19a–h), with light bands regularly distributed on outer anterior margins of disc (Figs. 20a, b). Living specimens may vary their color according to the environment, but maintain the color pattern described above when preserved. Ventral surface whitish to brownish, copper; generally darkening toward edges. Tail prominently cross-banded, with 3–5 black bands (Fig. 20d); bands diffuse or less distinct in large specimens (Fig. 20e). Size, reproduction, diet and habitat. Small juveniles with 168, 171 and 174 mm DW from Gulf of Mexico and Bahamas were analyzed by us, at the same time we have observed term embryos from Virginia and North Carolina ranging from 235 to 258 mm DW, so the birth size may be estimated as ranging from 170 to 260 mm DW. The largest juvenile male was 342 mm DW (FLMNH 45943, Georgia), while the smallest adult male measured 274 mm DW (BMNH 1948.8.6.20, Texas). Three individuals with 306, 333 and 333 mm DW were near maturity; all males larger than 342 mm DW were adults. The maturation size of males is estimated between 270–340 mm DW. The largest male analyzed by us measured 420 mm DW. Schwartz (2011) mentioned males up to 609 mm DW, which should be close to the maximum size for males of this species. Bigelow & Schroeder (1953) state that females produce young when reaching 625–650 mm DW, and maybe even smaller. Daiber & Booth (1960) reported that a female with 669 mm DW was becoming sexually mature and observed pregnant females only with 800 mm WD. Cu-Salazar et al . (2014), working in the southern Gulf of Mexico, estimated the size at 50% maturity at 568 mm DW. Therefore, the size at maturity should be between 550–600 mm DW. The largest female examined by us was 732 mm DW (FLMNH 29943, Florida, Atlantic coast), and the largest female reported that could be referred with confidence to the present species rather than to G. altavela measured 899 mm DW (Hoese 1962). Females of G. lessae , sp. nov. should attain a maximum breadth of 1000–1100 mm. Schwartz (2011) comments on a female of 1660 mm DW caught in North Carolina, a size that is well above the size that has been undoubtedly reported for this species (as G. micrura ). Individuals from North Carolina, Virginia and regions farther north reach larger sizes (birth, maturity and maximum) than individuals from Gulf of Mexico. The species is matrotrophic viviparous (lipid histotrophy). Bigelow & Schroeder (1953) suggest that the young are produced from late May through July and into August, and perhaps even later in the season. Daiber & Booth (1960) indicate that pregnancy occurs during the summer months for this species. In the few analyzed stomachs were found fishes, crustaceans, mollusks and other invertebrates (Bigelow & Schroeder 1953, Daiber & Booth 1960, Schmid et al . 1988). Also a demersal species, living along the coast on sandy and muddy bottoms. Geographical distribution. Gymnura lessae , sp. nov. is distributed in the north and central western Atlantic from Woods Hole in southern Massachusetts and Buzzards Bay to the Gulf of Mexico (including the Gulf Coast of the United States and Mexico) (Fig. 21). Both previously unidentified specimens from Bahamas and Jamaica (SMF 2398 and ZMB 34702, respectively) were classified by the discriminant model as G. lessae , sp. nov. (99.9 and 99.8% probability, respectively), so the distributional range of the species may be extended to the northern Caribbean Sea. The occurrence of G. lessae , sp. nov. in North Carolina and regions farther north seems to be seasonal, related to summer months (Bigelow & Schroeder 1953). The northernmost specimen analyzed in this study was MNHN 7938 from New York, however there are records of the species (as G. micrura ) from Chincoteague, Sinepuxent and Delaware Bays, Cape May and Sandy Hook Bay (New Jersey), New York, Narragansett and Buzzards Bays, and Woods Hole (Massachusetts) (Daiber & Booth 1960, Schwartz 1961, Fowler 1920, Nichols & Breder 1927, Breder 1931, Müller & Henle 1841, Bigelow & Schroeder 1953). Although we have not analyzed any specimens from Mexico, the species is commonly caught in Mexican waters of the Gulf of Mexico (Bigelow & Schroeder 1953, González-Isáis 2003, González-Isáis & Domínguez 2004, Kobelkowsky 2004, Cu-Salazar et al . 2014). Etymology. The name lessae is a tribute to Dr. Rosângela Lessa, a highly esteemed Brazilian researcher who has been struggling valiantly for the conservation of sharks and rays in Brazil, and has contributed significantly to the knowledge of this group. Prof. Rosângela Lessa has been a valued mentor and friend of the first author. Remarks. Müller & Henle (1841) included the specific name maclura in G. micrura , their material of which comprised both G. micrura and G. lessae (these authors also included G. poecilura in G. micrura see above). This error was perpetuated by subsequent studies until the end of the 1930s (Duméril 1865, Günther 1870, Jordan & Gilbert 1883, Jordan & Evermann 1896, Linton 1905, Smith 1907, Wilson 1908, Weymouth 1910, Fowler 1913, Gudger 1913, Coles 1915, Breder 1926, Nichols & Breder 1927, Breder & Nigrelli 1934, Puyo 1936, Breder 1938). Raja maclura (= Gymnura maclura ) was described by Lesueur (1817) from the Northeastern coast of North America and i : 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 20-34, DOI: 10.11646/zootaxa.4332.1.1, http://zenodo.org/record/1052041 : {"references": ["Jordan, D. S. & Gilbert, C. H. (1883) Synopsis of the fishes of North America. Bulletin of the United States National Museum, 16, 1 - 1018. https: // doi. org / 10.5479 / si. 03629236.16. i", "Jordan, D. S. & Evermann, B. W. (1896) The fishes of North and Middle America: a descriptive catalogue of the species of fishlike vertebrates found in the waters of North America, north of the Isthmus of Panama. Bulletin of the United States National Museum, 47 (1), 1 - 1240.", "Linton, E. 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(2004) El sistema excretor de la raya mariposa Gymnura micrura (Pisces: Rajiformes). Anales del Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Serie Zoologia, 75 (1), 185 - 192.", "McEachran, J. D. & Aschliman, N. (2004) Phylogeny of Batoidea. In: Carrier, J. C., Musick, J. A. & Heithaus, M. R. (Eds.), Biology of Sharks and their Relatives. Chapter 3. CRC Press, Boca Raton, pp. 79 - 113. https: // doi. org / 10.1201 / 9780203491317. ch 3", "Shepherd, T. D. & Myers, R. A. (2005) Direct and indirect fishery effects on small coastal elasmobranchs in the northern Gulf of Mexico. Ecology Letters, 8, 1095 - 1104. https: // doi. org / 10.1111 / j. 1461 - 0248.2005.00807. x", "Smith, W. D., Bizarro, J. J., Richards, V. P, Nielsen, J., Marquez-Farias, F. & Shivji, M. S. (2009) Morphometric convergence and molecular divergence: the taxonomic status and evolutionary history of Gymnura crebripunctata and Gymnura marmorata in the eastern Pacific Ocean. 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Academic Press, San Diego / London, pp. 63 - 84. https: // doi. org / 10.1016 / B 978 - 012670950 - 6 / 50005 - 9", "Rosenberger, L. J. (2001 a) Phylogenetic relationships within the stingray genus Dasyatis (Chondrichthyes: Dasyatidae). Copeia, 3, 615 - 627.", "McEachran, J. D. & Carvalho, M. R. de (2002) Gymnuridae: Butterfly Rays. In: Carpenter, K. E. (Ed.), The Living Marine Resources of the Western Central Atlantic. Volume 1: Introduction, Mollusks, Crustaceans, Hagfishes, Sharks, Batoid fishes and Chimaeras. FAO, Rome, pp. 575 - 577.", "Fowler, H. W. (1945) A study of the fishes of the southern Piedmont and coastal plain. Monographs of the Academy of Natural Sciences of Philadelphia, 7, 1 - 408.", "Puyo, J. (1936) Contribution a l'etude ichthyologique de la Guyane francaise. Peches et pecheries. Bulletin de la Societe d'Histoire Naturelle de Toulouse, 70, 5 - 258.", "Lesueur, C. A. (1817) Description of three new species of the genus Raja. Journal of the Academy of Natural Sciences, 1 (3), 41 - 45.", "Krefft, G. & Stehmann, M. (1973) Pristidae, Rhinobatidae, Torpedinidae, Dasyatidae, Myliobatidae, Rhinopteridae, Mobulidae. In: Hureau, J. C. & Monod, T. (Eds.), Check - list of the fishes of the north - eastern Atlantic and of the Mediterranean. Vol. 1. CLOFNAM, Paris, pp. 51 - 77"]} |
format |
Text |
author |
Carvalho, Marcelo Rodrigues De |
author_facet |
Carvalho, Marcelo Rodrigues De |
author_sort |
Carvalho, Marcelo Rodrigues De |
title |
Gymnura lessae Carvalho 2017, sp. nov. |
title_short |
Gymnura lessae Carvalho 2017, sp. nov. |
title_full |
Gymnura lessae Carvalho 2017, sp. nov. |
title_fullStr |
Gymnura lessae Carvalho 2017, sp. nov. |
title_full_unstemmed |
Gymnura lessae Carvalho 2017, sp. nov. |
title_sort |
gymnura lessae carvalho 2017, sp. nov. |
publisher |
Zenodo |
publishDate |
2017 |
url |
https://dx.doi.org/10.5281/zenodo.6039807 https://zenodo.org/record/6039807 |
long_lat |
ENVELOPE(170.033,170.033,-72.117,-72.117) ENVELOPE(-58.250,-58.250,-63.917,-63.917) ENVELOPE(77.955,77.955,-68.605,-68.605) ENVELOPE(-69.117,-69.117,-68.517,-68.517) ENVELOPE(8.224,8.224,63.072,63.072) ENVELOPE(156.450,156.450,-83.050,-83.050) ENVELOPE(-64.246,-64.246,-65.246,-65.246) ENVELOPE(-66.511,-66.511,-68.993,-68.993) ENVELOPE(-67.338,-67.338,-68.785,-68.785) ENVELOPE(130.500,130.500,-66.217,-66.217) ENVELOPE(-55.798,-55.798,52.533,52.533) ENVELOPE(-57.233,-57.233,-63.900,-63.900) |
geographic |
Pacific Indian Myers Gonzalez Lookout Perez Midland Cape Lookout Meek Gunter Keyhole Cape May Sandy Hook Dominguez |
geographic_facet |
Pacific Indian Myers Gonzalez Lookout Perez Midland Cape Lookout Meek Gunter Keyhole Cape May Sandy Hook Dominguez |
genre |
North Atlantic Copepods |
genre_facet |
North Atlantic Copepods |
op_relation |
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op_rights |
Open Access Creative Commons Zero v1.0 Universal https://creativecommons.org/publicdomain/zero/1.0/legalcode cc0-1.0 info:eu-repo/semantics/openAccess |
op_rightsnorm |
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op_doi |
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1766137984628817920 |
spelling |
ftdatacite:10.5281/zenodo.6039807 2023-05-15T17:37:48+02:00 Gymnura lessae Carvalho 2017, sp. nov. Carvalho, Marcelo Rodrigues De 2017 https://dx.doi.org/10.5281/zenodo.6039807 https://zenodo.org/record/6039807 unknown Zenodo http://zenodo.org/record/1052041 http://publication.plazi.org/id/FF9EFFCAFF9DFFC7A229883A3533FFC0 http://zoobank.org/C7D6FF64-5813-4AAB-AE40-C24ABB74433F https://zenodo.org/communities/biosyslit https://dx.doi.org/10.11646/zootaxa.4332.1.1 http://zenodo.org/record/1052041 http://publication.plazi.org/id/FF9EFFCAFF9DFFC7A229883A3533FFC0 https://dx.doi.org/10.5281/zenodo.1052063 https://dx.doi.org/10.5281/zenodo.1052065 https://dx.doi.org/10.5281/zenodo.1052067 https://dx.doi.org/10.5281/zenodo.1052069 https://dx.doi.org/10.5281/zenodo.1052071 https://dx.doi.org/10.5281/zenodo.1052073 https://dx.doi.org/10.5281/zenodo.1052075 https://dx.doi.org/10.5281/zenodo.1052077 https://dx.doi.org/10.5281/zenodo.1052079 https://dx.doi.org/10.5281/zenodo.1052081 https://dx.doi.org/10.5281/zenodo.1052109 https://dx.doi.org/10.5281/zenodo.1052113 https://dx.doi.org/10.5281/zenodo.1052117 https://dx.doi.org/10.5281/zenodo.1052125 https://dx.doi.org/10.5281/zenodo.1052129 https://dx.doi.org/10.5281/zenodo.1052133 https://dx.doi.org/10.5281/zenodo.1052137 https://dx.doi.org/10.5281/zenodo.1052083 http://zoobank.org/C7D6FF64-5813-4AAB-AE40-C24ABB74433F https://dx.doi.org/10.5281/zenodo.6039808 https://zenodo.org/communities/biosyslit Open Access Creative Commons Zero v1.0 Universal https://creativecommons.org/publicdomain/zero/1.0/legalcode cc0-1.0 info:eu-repo/semantics/openAccess CC0 Biodiversity Taxonomy Animalia Chordata Elasmobranchii Myliobatiformes Gymnuridae Gymnura Gymnura lessae article-journal ScholarlyArticle Taxonomic treatment Text 2017 ftdatacite https://doi.org/10.5281/zenodo.6039807 https://doi.org/10.11646/zootaxa.4332.1.1 https://doi.org/10.5281/zenodo.1052063 https://doi.org/10.5281/zenodo.1052065 https://doi.org/10.5281/zenodo.1052067 https://doi.org/10.5281/zenodo.1052069 https: 2022-04-01T09:51:46Z Gymnura lessae , sp. nov. (Figs. 11–20, 34b, 36b – 38b, 42b – 44b, 46b – 48b; Tabs. 3–4) Pteroplatea maclura : Jordan & Gilbert, 1883: 46 (description, North America, distributional range including G. micrura ); Jordan & Evermann, 1896: 86 (description, North and Middle America, distributional range including G. micrura ); Linton, 1905: 348 (stomach contents, Beaufort, N. Carolina); Smith, 1907: 45 (description, North Carolina, maximum size probably wrong); Wilson, 1908: 626 (parasites, North Carolina); Weymouth, 1910: 135 (Louisiana); Coles, 1915: 94 (C. Lookout, North Carolina); Nichols & Breder, 1927: 27 (Sandy Hook Bay and Woods Hole). Pteroplatea maclura (in part): Müller & Henle, 1841: 169 (description, only specimen(s) from New York); Duméril, 1865: 614 (description, only specimens from USA, size probably erroneous as the New York specimen recorded with a width of 2 meters is the same analyzed by Müller & Henle 1841, who had earlier termed it as small); Günther, 1870: 487 (description, only specimen from Texas). Pteroplatea micrura : Garman, 1913: 414, pl. 33, figs. 3–4 (description, color, illustration); Fowler, 1920: 146 (listed, C. May County, New Jersey); Hildebrand & Schroeder, 1928: 67, fig. 37 (description, Chesapeake Bay); Breder, 1931: 39 (listed, Sandy Hook Bay, New Jersey); Fowler, 1931: 46 (listed, Port Aransas, Texas); Beltran, 1934: 10 (listed, Mexico, not seen); Gunter, 1935: 39 (listed, Louisiana & Gulf of Mexico); Fowler, 1936: 132 (West Africa references for G. sereti , sp. nov., description of mid-Atlantic U.S. specimen); Gunter, 1941: 196 (listed, Louisiana). Pteroplatea micrura (in part): Meek & Hildebrand, 1923: 87 (no Panama record, color description matching G. lessae , nov. sp., probable area of occurrence of G. micrura ). Gymnura micrura : Bigelow & Schroeder, 1953: 408 –416, figs. 97, 98 (description, illustrations, U.S. [Gulf of Mexico and North Coast] and Mexico); Joseph & Yerger, 1956: 119 (biological notes, Alligator Harbor, Florida); Daiber & Booth, 1960: 138 –139 (biological notes, Delaware and Chincoteague Bays); Schwartz, 1961: 390 (size, embryos, listed, Chincoteague & Sinepuxent Bays); Hoese, 1962: 170 –171 (occurrence, Virginia’s Seaside); Snelson & Williams, 1981: 6, 8 (biological notes, Florida); Schmid et al . 1988: 123 –124 (biological notes, Florida); Wourms, 1993: 271, 283 (embryonic development, South Carolina); Maruska, 2001: 52, 55–57, 59, 62, figs. 6, 9b (lateral line canal system); Rosenberger, 2001b (locomotion, pectoral fin); González-Isáis, 2003: 260, 262, 264–266, 269–271, figs. 2a, 5 (cephalic musculature; Veracruz, Mexico); Wigley et al . 2003: 7, 17 (length-weight relationships, Northeast Coast of the U.S.); González-Isáis & Domínguez, 2004: 518, 521–524, 527, 528, 530, 532, tab. 1, figs. 3a, 4b, 5b, 6b, 7a, 9a (skeletal morphology, cephalic musculature; Veracruz, Mexico); Kobelkowsky, 2004: 185 –192, figs. 1–3 (morphology of urogenital system; Veracruz, Mexico); McEachran & Aschliman, 2004: 87, fig. 3.8 (clasper; Florida, Gulf of Mexico); Shepherd & Myers, 2005: 1097 – 1100, 1102 (population decline, Gulf of Mexico); Smith et al. , 2009: 761 –763, 765, 766, 768, 772–778 (morphometry, molecular data); Schwartz, 2011: 35 –36 (distributional pattern, North Carolina); Kobelkowsky, 2012: 36, 38–41 (heart anatomy, Mexico); Cu-Salazar et al. 2014: 109 –117 (reproductive biology; Tabasco and Campeche, Mexico). Gymnura micrura (in part): McEachran et al . 1996: 67 –68, 70–75, 83 (lateral line, skeletal morphology, cephalic and branchial musculature; only specimen TCWC 642.08 [Florida, Gulf of Mexico],? TCWC [uncataloged specimen]); Rosenberger, 2001a: 615, 616, 618, 620, 621, 623–627 (in systematics analysis; only specimen FMNH 46250 [Florida, Gulf of Mexico] and probably FMNH 40363 [Dominican Republic],? FMNH 89992); McEachran & Carvalho, 2002: 577 (identification guide, illustration, Western Central Atlantic, G. micrura also included). Gymnura altavela : Fowler, 1945: 162 (embryos, South Carolina, Bigelow & Schroeder analyzed the specimens and found them to be G. micrura [= G. lessae , sp. nov.]). Holotype. USNM 51940 (420 mm DW, adult male), North Carolina, Beaufort, Collector: G. Bean, Accession num. 0 43551, USNM 440342 recatalogued from USNM 51940 (Fig. 11). Paratypes. USNM 440342 (4 specimens: adult male, 370 mm DW; adult male, 342 mm DW; juvenile female, 280 mm DW; skeleton of a dissected specimen), North Carolina, Beaufort, collector: G. Bean, accession num. 0 43551, date cat: 0 9 Feb 2017, recatalogued from USNM 51940 (Figs. 12–14) Diagnosis. A medium size butterfly ray occurring in the central and northwestern Atlantic Ocean. Gymnura lessae , sp. nov. may be distinguished from other species of butterfly rays (except G. micrura and G. sereti , sp. nov. ) by the combination of the following characters: absence of spiracular tentacle, caudal stings, and dorsal fin; tail relatively short (mean post-cloacal length 19% DW) and prominently banded, presenting 3 to 5 black bands that may be less distinct in large adults. Gymnura lessae , sp. nov. may be distinguished from G. micrura and G. sereti , sp. nov. by the dorsal contour of its hyomandibula with a conspicuous proximal protuberance followed by a inconspicuous distal protuberance ( vs . dorsal contour of hyomandibula with two conspicuous protuberances); projection of the anteroventral fenestra on the floor of the coracoid bar taking the form of a funnel with relatively narrower opening and end (in G. micrura this projection takes the shape of a funnel with large opening and closing; in G. sereti , sp. nov. the funnel has a large opening, but presents an abrupt tapering at its half length, with the end of the funnel relatively narrow), and higher number of pectoral radials (ranges 118–127 vs . 112–119 in G. micrura and 109–115 in G. sereti , sp. nov. ). Gymnura lessae , sp. nov. may be further distinguished from G. sereti , sp. nov. by the following characters: clasper of mature males more slender and longer, Lclasper 9.4–14.3% DW ( vs . clasper stouter and shorter, Lclasper 6.8–9.2% DW); cranial fontanelle U-shaped ( vs . keyhole-shaped); mesopterygium divided into one anterior solid element and 5–7 smaller fragments ( vs . mesopterygium divided into two solid elements); distance between anteroventral and posteroventral fenestrae of scapulocoracoid representing 35–40% of scapulocoracoid length ( vs . distance between AVF and PVF 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 at its half length); lower number of diplospondylous vertebrae (means 97 vs . 110), and higher number of radials in the mesopterygial fragments compared to the second mesopterygial element of G. sereti , sp. nov. (ranges 14–19 vs . 10–15). Gymnura lessae , sp. nov. is most similar to G. micrura , and may be further differentiated from it by its brownish dorsal side with a vermiculate background ( vs . dorsal side usually uniformly brown or gray without any vermiculate pattern). Description. Measurements are presented in Table 3; meristic data in Table 4. The following description is based on all specimens examined. Throughout the text the proportions are presented as: minimum value–maximum value (mean). See Figs. 11–17, 18a, and 19 and 20 for external morphology and color, Fig. 34b for ventral lateralline pattern and Figs. 18b, 36b – 38b, 42b – 44b and 46b – 48b for skeletal morphology. External morphology. Disc lozenge shaped, 1.54–1.94 (1.75) times broader than long [1.54–1.70 (1.61) for adult males; 1.71–1.89 (1.80) 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 their disc (Fig. 15). Preoral snout length 7.6–16 (12)% DW [8–11 (10)% DW in adult females; 13–16 (14)% DW in adult males], preorbital snout length 6.8–14 (9.7)% DW [7–9 (8)% DW in adult females; 10–14 (12)% DW in adult males], preorbital snout width 31–41 (36)% DW, postspiracle snout width 40–54 (48)% 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. 15); 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 disc half-length. Pelvic fin single lobed, rectangular, their corners rounded (Fig. 16). Skin entirely smooth, without denticles on the dorsal and ventral surface of the disc. Interorbital space broad, interorbital width 7.4–10 (8.6)% DW. Eyes dorsolateral, small, oval and protruding slightly (Fig. 17a); eye diameter 1.1–2.2 (1.7)% DW, representing 11.5–26.3 (19.5)% of interorbital width; orbit diameter 2–3.9 (2.8)% DW, representing 61–175 (96)% of spiracle length. Eyes more protruded and relatively larger in embryos. Spiracles immediately posterior to eyes, relatively large, lozenge-shaped, contour of the inner spiracular margin concave to rectilinear (Figs. 17b, c); spiracle length 2.1–3 (1.8)% DW, 1.1–3 (1.8) times eye diameter; its inner posterior margin without tentacles. Ventral head length 20–28 (23)% DW. Nostril narrowly oval, diagonally directed, only the circular distal margin not covered by nasal curtain, posterior lateral margin with lobe; nostril length 2.0–4.1 (3.2)% DW [2.7–3.4 (3.1)% DW in adult females; 3.1–4.1 (3.5)% DW in adult males], 0.9–2.7 (1.7) times internasal width; internasal width 1.4–3.1 (1.9)% DW. Anterior nasal flaps medially expanded and fused into a broad, skirt-shaped, posteriorly expanded nasal curtain that covers internasal space and reaches mouth (Fig. 17d). Nasal curtain skirt shaped, weakly to moderately bilobed, wide, its width 2.4–4.3 (3.1) times its length; lateral margin concave; posterolateral apices rounded; posterior margin straight to weakly concave, a small medial gap may be present. Mouth relatively wide, its width 7.9–11 (9.1)% DW, 30–48 (39)% head length, 0.9–1.7 (1.2) times nasal curtain width; without papillae on floor or labial folds, although some striations may originate radially from mouth corners (Fig. 17d); lower lip arched rearward toward corners, uniformly convex or weakly to moderately concave along medial region, without any lump or knob (Fig. 17d); a strip of corrugate skin forms a half circle below mouth, ending at mouth corners. Small, numerous and closely crowded teeth in bands; teeth similar to teeth in G. micrura , with one medial, pointed cusp directed towards inside of mouth and the base somewhat expanded, entirely twisted in opposite direction; in labial face, tooth resembling a three-armed boomerang; 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. 17d). Distance between gill slits decreasing posteriorly; distance between inner ends of fifth pair 62–79 (73)% distance between inner ends of first pair; distance between inner ends of first pair 15–19 (16) % DW, 57–84 (70)% ventral head length, 4.8–13.0 (8.5) times internasal width; distance between inner ends of fifth pair 10–13 (12)% DW, 40–60 (51)% ventral head length, 4.0–9.2 (6.2) times internasal width. Tail slender and short, whip-like, cross-banded (see color description), tapering toward tip, without caudal stings and dorsal fin (Figs. 20d, e); postcloacal tail 15–24 (19)% DW, 28–42 (34)% DL, 32–48 (39)% precloacal length; low longitudinal skin folds present on dorsal and ventral surfaces, skin folds not reaching tail tip. Clasper of mature males cylindrical, somewhat depressed, relatively slender and long (compared to G. sereti , sp. nov. ) (Fig. 18); distance from posterior margin of cloaca to clasper tip 9.4–14.3 (11.9)% DW; clasper tapering slightly distally; tip conical, bluntly pointed to pointed, not calcified; non-calcified portion relatively long compared to G. micrura , but shorter than in G. sereti , sp. nov. apopyle (on the anterior dorsal surface) connected to hypopyle by a long, dorsomedial, posteriorly curved clasper groove; spermatic duct ending before reaching clasper tip; rhipidion and pseudorhipidion absent; a long ventral pseudosiphon laterodistally situated to hypopyle; welldeveloped dorsal pseudosiphon 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 brownish, grayish or purplish, vermiculated. The vermiculate pattern varying from less conspicuous with lighter and darker shades to a conspicuous pattern where the lighter areas become larger and more remarkable (Figs. 19a–h), with light bands regularly distributed on outer anterior margins of disc (Figs. 20a, b). Living specimens may vary their color according to the environment, but maintain the color pattern described above when preserved. Ventral surface whitish to brownish, copper; generally darkening toward edges. Tail prominently cross-banded, with 3–5 black bands (Fig. 20d); bands diffuse or less distinct in large specimens (Fig. 20e). Size, reproduction, diet and habitat. Small juveniles with 168, 171 and 174 mm DW from Gulf of Mexico and Bahamas were analyzed by us, at the same time we have observed term embryos from Virginia and North Carolina ranging from 235 to 258 mm DW, so the birth size may be estimated as ranging from 170 to 260 mm DW. The largest juvenile male was 342 mm DW (FLMNH 45943, Georgia), while the smallest adult male measured 274 mm DW (BMNH 1948.8.6.20, Texas). Three individuals with 306, 333 and 333 mm DW were near maturity; all males larger than 342 mm DW were adults. The maturation size of males is estimated between 270–340 mm DW. The largest male analyzed by us measured 420 mm DW. Schwartz (2011) mentioned males up to 609 mm DW, which should be close to the maximum size for males of this species. Bigelow & Schroeder (1953) state that females produce young when reaching 625–650 mm DW, and maybe even smaller. Daiber & Booth (1960) reported that a female with 669 mm DW was becoming sexually mature and observed pregnant females only with 800 mm WD. Cu-Salazar et al . (2014), working in the southern Gulf of Mexico, estimated the size at 50% maturity at 568 mm DW. Therefore, the size at maturity should be between 550–600 mm DW. The largest female examined by us was 732 mm DW (FLMNH 29943, Florida, Atlantic coast), and the largest female reported that could be referred with confidence to the present species rather than to G. altavela measured 899 mm DW (Hoese 1962). Females of G. lessae , sp. nov. should attain a maximum breadth of 1000–1100 mm. Schwartz (2011) comments on a female of 1660 mm DW caught in North Carolina, a size that is well above the size that has been undoubtedly reported for this species (as G. micrura ). Individuals from North Carolina, Virginia and regions farther north reach larger sizes (birth, maturity and maximum) than individuals from Gulf of Mexico. The species is matrotrophic viviparous (lipid histotrophy). Bigelow & Schroeder (1953) suggest that the young are produced from late May through July and into August, and perhaps even later in the season. Daiber & Booth (1960) indicate that pregnancy occurs during the summer months for this species. In the few analyzed stomachs were found fishes, crustaceans, mollusks and other invertebrates (Bigelow & Schroeder 1953, Daiber & Booth 1960, Schmid et al . 1988). Also a demersal species, living along the coast on sandy and muddy bottoms. Geographical distribution. Gymnura lessae , sp. nov. is distributed in the north and central western Atlantic from Woods Hole in southern Massachusetts and Buzzards Bay to the Gulf of Mexico (including the Gulf Coast of the United States and Mexico) (Fig. 21). Both previously unidentified specimens from Bahamas and Jamaica (SMF 2398 and ZMB 34702, respectively) were classified by the discriminant model as G. lessae , sp. nov. (99.9 and 99.8% probability, respectively), so the distributional range of the species may be extended to the northern Caribbean Sea. The occurrence of G. lessae , sp. nov. in North Carolina and regions farther north seems to be seasonal, related to summer months (Bigelow & Schroeder 1953). The northernmost specimen analyzed in this study was MNHN 7938 from New York, however there are records of the species (as G. micrura ) from Chincoteague, Sinepuxent and Delaware Bays, Cape May and Sandy Hook Bay (New Jersey), New York, Narragansett and Buzzards Bays, and Woods Hole (Massachusetts) (Daiber & Booth 1960, Schwartz 1961, Fowler 1920, Nichols & Breder 1927, Breder 1931, Müller & Henle 1841, Bigelow & Schroeder 1953). Although we have not analyzed any specimens from Mexico, the species is commonly caught in Mexican waters of the Gulf of Mexico (Bigelow & Schroeder 1953, González-Isáis 2003, González-Isáis & Domínguez 2004, Kobelkowsky 2004, Cu-Salazar et al . 2014). Etymology. The name lessae is a tribute to Dr. Rosângela Lessa, a highly esteemed Brazilian researcher who has been struggling valiantly for the conservation of sharks and rays in Brazil, and has contributed significantly to the knowledge of this group. Prof. Rosângela Lessa has been a valued mentor and friend of the first author. Remarks. Müller & Henle (1841) included the specific name maclura in G. micrura , their material of which comprised both G. micrura and G. lessae (these authors also included G. poecilura in G. micrura see above). This error was perpetuated by subsequent studies until the end of the 1930s (Duméril 1865, Günther 1870, Jordan & Gilbert 1883, Jordan & Evermann 1896, Linton 1905, Smith 1907, Wilson 1908, Weymouth 1910, Fowler 1913, Gudger 1913, Coles 1915, Breder 1926, Nichols & Breder 1927, Breder & Nigrelli 1934, Puyo 1936, Breder 1938). Raja maclura (= Gymnura maclura ) was described by Lesueur (1817) from the Northeastern coast of North America and i : 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 20-34, DOI: 10.11646/zootaxa.4332.1.1, http://zenodo.org/record/1052041 : {"references": ["Jordan, D. S. & Gilbert, C. H. (1883) Synopsis of the fishes of North America. Bulletin of the United States National Museum, 16, 1 - 1018. https: // doi. org / 10.5479 / si. 03629236.16. i", "Jordan, D. S. & Evermann, B. W. (1896) The fishes of North and Middle America: a descriptive catalogue of the species of fishlike vertebrates found in the waters of North America, north of the Isthmus of Panama. Bulletin of the United States National Museum, 47 (1), 1 - 1240.", "Linton, E. 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CLOFNAM, Paris, pp. 51 - 77"]} Text North Atlantic Copepods DataCite Metadata Store (German National Library of Science and Technology) Pacific Indian Myers ENVELOPE(170.033,170.033,-72.117,-72.117) Gonzalez ENVELOPE(-58.250,-58.250,-63.917,-63.917) Lookout ENVELOPE(77.955,77.955,-68.605,-68.605) Perez ENVELOPE(-69.117,-69.117,-68.517,-68.517) Midland ENVELOPE(8.224,8.224,63.072,63.072) Cape Lookout ENVELOPE(156.450,156.450,-83.050,-83.050) Meek ENVELOPE(-64.246,-64.246,-65.246,-65.246) Gunter ENVELOPE(-66.511,-66.511,-68.993,-68.993) Keyhole ENVELOPE(-67.338,-67.338,-68.785,-68.785) Cape May ENVELOPE(130.500,130.500,-66.217,-66.217) Sandy Hook ENVELOPE(-55.798,-55.798,52.533,52.533) Dominguez ENVELOPE(-57.233,-57.233,-63.900,-63.900) |