Styracura schmardae Werner 1904

• Main Authors: De Carvalho, Marcelo R., Loboda, Thiago S., Da Silva, João Paulo C. B.
• Format: Text
• Language: unknown
• Published: Zenodo 2016
• Subjects: Biodiversity; Taxonomy; Animalia; Chordata; Elasmobranchii; Myliobatiformes; Potamotrygonidae; Styracura; Styracura schmardae; Austin; Pacific; De la Costa; Trinidad; Crawford; Lopez; Perez; Fossa; Meek; Aguirre; Garibaldi; North Atlantic
• Online Access: https://dx.doi.org/10.5281/zenodo.5696861; https://zenodo.org/record/5696861

Styracura schmardae (Werner, 1904) (Figs. 1, 3–6; Table 1) Trygon schmardae Werner, 1904: 298 (original description, single specimen, not depicted; type locality: Jamaica). Dasybatus schmardae (Werner, 1904).— Garman, 1913: 386 (verbatim from Werner, 1904; placed in subgenus Pastinachus no specimens examined); Meek & Hildebrand, 1923: 81 (morphological description, Panama Canal, two specimens). Dasybatus torrei Garman, 1913: 386 –388 (original description, denticles, teeth, pelvic girdle, based on single specimen from Tunas de Zaza, Cuba, not depicted; placed in subgenus Pastinachus ). Dasyatis schmardae (Werner, 1904).— Fowler, 1931: 391 (color pattern, Trinidad); Beebe & Tee-Van, 1941: 263 (compared to H. pacifica , described as new); Boeseman, 1948: 31 –33 (morphological description, pelvic girdle, range extension, four specimens). Himantura schmardae (Werner, 1904).— Bigelow & Schroeder, 1953: 390 –394 (morphological redescription, synonymy, distribution); Cervigón, 1992: 200 (Venezuela); Lovejoy, 1996: 220, 221, 229, 230, 246–248 (morphology, relationships, biogeography); McEachran et al. , 1996: 64, 65, 71, 72, 75–81, 83 (morphology, phylogeny); McEachran & Fechhelm, 1998: 182 (Gulf of Mexico); Lovejoy et al ., 1998: 421 (molecular phylogeny); Cervigón & Alcalá, 1999: 190 (Venezuela); Castro-Aguirre et al ., 1999: 72 (Mexico); Camargo & Isaac, 2001: 144 (northern Brazil); McEachran & Carvalho, 2003: 568 (identification, distribution); McEachran & Aschliman, 2004: 80, 86, 88–92, 96, 98, 99, 101 (morphology, phylogeny); Carvalho et al ., 2004: 10, 49, 77, 81, 82, 93, 105, 113 (morphology, relationships); Carvalho & Lovejoy, 2011: 46, 47 (molecular phylogeny); Naylor et al ., 2012a: 43, 46, 48, 49 (mitochondrial DNA-based phylogeny); Naylor et al ., 2012b: 79, 225 (DNA identification); Aschliman et al. , 2012a: 64, 66, 68, 70–72, 75, 77, 79, 86–88, 90, 92–94 (morphology, phylogeny); Aschliman et al. , 2012b: 30, 34–36, 38, 39 (molecular phylogeny); Moral-Flores et al ., 2015: 130 (Mexico); Last et al ., 2016: 352 (molecular phylogeny, classification). Diagnosis . A species of Styracura distinguished from S. pacifica by the following combination of characters: anterior disc margin slightly more straight across (more oblique in S. pacifica ); usually greater enlarged scapular denticles (poorly developed in some specimens of S. pacifica , but always well developed in S. schmardae ); slightly larger eyes (up to one-half spiracle length in S. schmardae vs . one-third in S. pacifica ); slightly smaller preorbital length (range 15.6–20.9% DW and mean 18.3% DW in S. schmardae vs . 21.2% DW and 22.5% DW in holotype and measured specimen of S. pacifica , respectively); smaller prenarial length (range 10.8–13.5% DW and mean 12.2% DW in S. schmardae vs . 14.5% DW and 15% DW in holotype and measured specimen of S. pacifica , respectively); smaller preoral length (range 15.7–18.8% DW and mean 17.5% DW in S. schmardae vs . 20.3% DW in measured specimen of S. pacifica ); shorter distance from cloaca to caudal sting origin (range 51.8–90.9% DW and mean 69.5% DW in S. schmardae [holotype 66.5%] vs . 98.4% DW in measured specimen of S. pacifica ); and brown to grayish- or olivaceous-brown dorsal disc color (frequently purplish-gray or darker brown in S. pacifica ). External morphology . Disc rounded to weakly rhomboidal, slightly broader than long, with average disc length 93.2% DW (range 87.2–98.6% DW). Anterior portion of disc more rounded and broader than posterior portion; posterior portion slightly more oval. Anterior margin of disc straight to weakly convex. Rostral knob small, rounded. Head relatively close to anterior disc margin. Snout relatively short; mean preorbital distance 18.3% DW (range 15.6–20.9% DW); preorbital distance about one and a half times interorbital distance. Mean prenasal distance 12.2% DW (range 10.8–13.5% DW). Mean preoral distance 17.5% DW (range 15.7–18.8% DW). Eyes small, protruding only slightly, their mean diameter 2.6% DW (range 1.8–3.8% DW). Eye diameter from four to five times smaller than interorbital distance. Spiracles very large and rectangular, obliquely positioned in relation to eyes, their mean length 7.7% DW (range 7.2–8.5% DW). Nasal curtain relatively straight, not widening significantly close to mouth. Posterior margin of nasal curtain weakly fringed, straight across. Nostrils anteroposteriorly elongate, slightly more rounded and larger anteriorly; mean distance between anterior margins of nostrils 7.4% DW (range 6.4–8.5% DW). Mouth slightly arched, with a small median concavity on lower jaw; mean mouth width 7.7% DW (range 6.9–8.5% DW). Internarial distance and mouth width about equal. Posterior to mouth integument corrugated. Five papillae on mouth floor. Teeth slightly wider than long, set in quincunx, in up to about 40 rows on upper and lower jaws; a few rows of teeth on upper jaw exposed even with mouth closed. Branchial basket quadrangular; mean distance between first pair of gill slits 20.7% DW (range 19.0–22.4% DW); mean distance between fifth pair of gill slits 15.8% DW (range 13.8–18.0% DW). Branchial basket region relatively long; mean distance between first and fifth pairs of gill slits 15.3% DW (range 14.1–16.3% DW). Pelvic fins protrude only slightly beyond posterior margins of pectoral fins in dorsal view. Pelvic fins broadly triangular, presenting somewhat straight anterior margins and slightly convex posterior margins. Mean length of anterior pelvic margins 17.4% DW (range 15.4–20.1% DW); pelvic fins not particularly broad, mean distance between outer apices of pelvic fins (width of pelvic fins measured together) 44.4% DW (range 36.7–51% DW). Claspers very reduced in juvenile and young specimens, mean external length of clasper in examined specimens 1.7% DW (range 1.5–2.2% DW), and mean internal length 10.9% DW (range 9–12.9% DW); adult males not examined. Tail very long, much longer than disc length; mean distance from posterior margin of cloaca to tail tip 146.7% DW (range 129.2–164.2% DW); tail length as measured from cloaca about twice snout to cloaca distance (mean 76.6% DW, range 64.3–84.9% DW). Tail wide at base, its greatest width slightly smaller than interorbital distance; mean tail width at pectoral fin insertion 10.8% DW (range 9.2–12.2% DW). Tail width tapering mildly from base to distal tip, but reduction slightly more accentuated posterior to caudal stings. Tail weakly rounded anterior to caudal stings, but nearly circular in cross section posterior to stings. Lateral tail ridges low, present from more or less level of posterior pelvic fins to well anterior to caudal sting origin. Ventral tail ridge very low, originating at about level of caudal sting origin to posterior to stings; lacking dorsal tail fold, ridge or keel. Caudal sting positioned far from tail base; mean distance between posterior margin of cloaca and caudal sting origin 69.5% DW (range 51.8–90.9% DW), very close to snout to cloaca distance. Caudal sting origin usually at more or less midlength of tail in specimens with intact tails. Caudal stings elongate, varying from 18.8–26.9% DW, nearly two times interorbital distance (10.8–15.1% DW), and much greater than preorbital snout length. Caudal stings relatively slender, their mean width 1.6% DW (range 1–2% DW). Lateral serrations between 60–70, closely positioned forming an almost continuous edge, especially closer to sting tip; serrations more developed toward sting tip but small serrations also present on sting base. Coloration . Most specimens with a uniform brown, grayish-brown or olivaceous-brown dorsal disc, tail and pelvic fins, sometimes with outer disc contours slightly darker, and dark brown posterior two-thirds of tail. Ventral color creamy white, with lightly darker outer disc margins (sometimes with small, indistinct grayish spots and blotches), and darker posterior pelvic fins; ventral posterior tail darker brown. Dermal denticles . An intense shagreen of dermal denticles covers practically the entire dorsal surface of disc and tail, forming a tough, hardened exterior. Denticles slightly less numerous and less closely packed toward disc margins. Pelvic fins mostly naked. Dermal denticles of at least two main shapes: large stellate denticles with broad basal plates closely packed together on the central area of dorsal disc and all of dorsal tail, and slightly smaller asterisk-shaped denticles present on disc margins and much of tail (Fig. 3). In addition, a pair of larger scapular denticles (or spines) present above shoulders, usually one on each side, but sometimes with two pairs, one more developed than the other. Larger denticles on dorsal disc and tail have broad, subcircular to quadrangular basal plates, these frequently overlapping. Larger denticles with quadriradiate crowns, with primary ridges subdividing distally into smaller ridges, and these frequently also further subdivided. The main dichotomic crown ridges resemble a cross, and bear a central, higher, and smaller pyramidal crown plate. Greatly enlarged scapular denticles above shoulders morphologically similar to larger denticles, but with many more dichotomic ridges radiating from the crown plate, and sometimes with two central, higher, longitudinal ridges forming an inverted V-shape on crown. Smaller denticles asterisk-shaped with several high crown ridges, usually six, branching from a single elevated, acute, cylindrical crown. These denticles present a star-shaped basal plate beneath coronal ridges (in contrast to larger, stellate denticles that have a broad basal plate). On the medial dorsal region of tail, close to caudal stings, these denticles possess a more developed and acute crown with fewer ridges, resembling small thorns. Lateral line canals . The hyomandibular canal ( HMD ) extends close to anterior disc margin, where numerous closely adjacent anterior subpleural tubules branch off in parallel (hyomandibular canal colored pink in Fig. 4); anterior subpleural tubules present only on anterior segment of hyomandibular canal. Hyomandibular canal extends obliquely toward posterior disc; its subpleural component ( spl ) somewhat concave, giving off two small posterior subpleural tubules ( pst ), and ending in an acute triangle; jugular component ( jug ) more or less straight posteriorly, but broadly arched lateral to gill slits, and slightly bulging near first gill slits. Infraorbital ( IOC blue in Fig. 4) canal with small, rounded infraorbital loop posteriorly, and without suborbital loop anteriorly; infraorbital canal highly sinuous, extending anteriorly to ventral midsnout length. Supraorbital canal ( SPO green) positioned medial to infraorbital canal and obliquely positioned, not sinuous; small prenasal loop present; supraorbital forming a small posterior loop just anterior to nasal curtain. Prenasal canal (anterior segment of nasal canal; NAS , yellow in Fig. 4) extends in a vertical line to anterior snout tip, not connecting to its posterior oblique segment lateral to nostril. Mandibular canal (not shown in Fig. 4 due to previous dissection) forming an inverted V-pattern just posterior to mouth (not shown in Fig. 4). Skeletal features . Neurocranium, jaws and hyoid arch . Neurocranium longer than wide, its greatest width at level of preorbital processes; neurocranial greatest width about two-thirds its greatest length (Figs. 5, 6). Nasal capsules with oval and wide nasal apertures ( na ), with broadly rounded anterior wall, but relatively anteroposteriorly short; triangular anterior median indentation present; internasal septum ( is ) moderately broad. Preorbital process ( prp ) projecting laterally, somewhat wide, tapering only slightly. Postorbital process projecting anterolaterally, somewhat straight and rectangular, and poorly calcified, obscured in radiographs by denticles on overlying integument. Supraorbital process ( sp ) broadly triangular. Neurocranium most slender at its posterior fourth, at level of posterior frontoparietal fontanelle. Otic capsule very short, about as wide as orbital region. Precerebral ( pcf ) and frontoparietal ( fpf ) fontenellae about three-fourths length of neurocranium. Precerebral fontanelle relatively large, wider posteriorly, and slightly trapezoidal with rounded margins; precerebral fontanelle wider than long, just greater than half length of frontoparietal fontanelle; frontoparietal fontanelle broadly triangular, longer than wide, not medially constricted and tapering, rounded posteriorly. Antorbital cartilage ( aoc ) very large and laterally compressed, widest at articulation with posterolateral nasal capsule, and extending posteriorly to close to hyomandibula, at level of jaw joint. Prespiracular cartilages ( psc ) weakly calcified, concave and slender. Meckel’s cartilages ( Mc ) greater than palatoquadrates ( pq ), with short and triangular dorsally projecting lateral processes posteriorly; ventral margin of lower jaws lacking distinct projection where lower jaws deflect anteriorly toward midline (Figs. 5, 6); ventrolateral process closer to jaw joint also absent or very reduced. Small triangular gap present between lower jaw symphyses. Jaws about equal in width to widest point of neurocranium. Palatoquadrates markedly slender, stouter at midwidth, and not as wide as lower jaws. Hyomandibulae ( hyo ) laterally compressed, blade-like, slender in dorsoventral view and stouter at midlength. Distal portion of hyomandibulae (where they deflect anteriorly, represented by an asterisk in Figs. 5, 6) stout, very elongate, just under one-third hyomandibular length. Hyomandibulae tightly articulated to neurocranium, extending anterolaterally from it. Well developed and stout ligament between hyomandibulae and lower jaws, usually directed slightly posteriorly toward midline. Small calcified elements present within ligament, including a small, wider than long, obliquely positioned (also directed posteriorly toward midline in Fig. 5), cylindrical angular element ( ac ); rounded calcified element also present adjacent to hyomandibula in some specimens, with smaller rounded cartilages in between (Figs. 5, 6). Pectoral girdle . Coracoid bar dorsoventrally flattened and laterally expanded. Scapulocoracoid in lateral view tall and triangular, with broadly inclined anterodorsal margin bearing anteriorly a triangular projection; scapulae with concave posterodorsal and posteroventral aspects (Fig. 7 a). Pectoral condyles situated on horizontal axis of scapular cartilage. Scapular process ( scp ) bearing a conspicuous fossa perforated by a small foramen (scapular process fenestra, spf ) on its distal portion, broadly triangular and firmly articulated to lateral aspects of synarcual. Procondyle ( pc ) nearly elliptical, vertically oriented and situated at anterolateral surface of scapula. Mesocondyle ( msc ) slightly oval and horizontal. Metacondyle ( mtc ) rounded, just posterior to mesocondyle. Horizontally oriented, expanded, and slightly curved facet ( fct ) present in large gap between pro- and mesocondyle; facet helps support propterygium. Anterior fenestrae much greater than posterior fenestrae. Anterodorsal ( adf ) and anteroventral ( avf ) fenestrae somewhat oval and dorsal and ventral to pectoral condyles, respectively. Posterodorsal fenestra ( pdf ) and posteroventral fenestra ( pvf ) greatly reduced, about equal in size, and much smaller than anterior fenestrae. Pelvic girdle . Puboischiadic bar ( pib ) relatively wide, laterally expanded and dorsoventrally flattened, with relatively oblique but straight anterior margins and posterior margin concave (Fig. 7 b). Five obliquely positioned obturator foramina ( of ) just medial to pelvic condyles ( pvc ) arranged in an arch along the anteroposterior axis of pelvic girdle. Prepelvic process ( ppp ) short and broadly triangular. Iliac processes ( ip ) on posterolateral corners of puboischiadic bar relatively long and wide, laterally flattened, projecting dorsally, and distally tapering. Ischial processes ( isp ) blunt, broadly triangular and posteromedially directed. Lateral prepelvic processes ( lpp ) robust, broadly triangular and anteriorly directed. Distribution . Essentially that given in McEachran & Carvalho (2003) with the noted extension of the northern Brazilian coast (e.g. Camargo & Isaac, 2001). Etymology . Named after zoologist and explorer Ludwig Karl Schmarda (23 August 1819 – 7 April 1908), founder of the zoological museum of the University of Graz, and later professor in the Zoological Institute of the University of Vienna (1861–1883), where he kept his collection of fishes including the specimen that became the type of Trygon schmardae . Franz Werner was on staff in the same institute for over 40 years, eventually retiring as professor in 1939 (Salvini-Plawen & Mizzaro, 1999). Remarks . Werner (1904: 298, 299) used the following characters to diagnose his new species: "front disc not perfectly rounded, but with small snout tip; top completely rough, especially on the pectoral fins, although sparser than the trunk; tail without enlarged thorns; scapular region with two large, round, ribbed tubercles side by side (their distance apart close to distance between nostrils); tail nearly twice as long as disc; color brown uniform, tail darker" [our translation]. Remarkably, Werner (1904: 298) alluded to the similarity between S. schmardae and " Trygon hystrix M. et H." (= Potamotrygon histrix ), and the features listed above were given to distinguish his new species from this species of Potamotrygon (Werner did not report any specimen of Potamotrygon and took his information directly from Müller & Henle, 1841); thus, a putative close relationship between S. schmardae and potamotrygonins was alluded to when S. schmardae was described more than 110 years ago. Werner's (1904) paper on the fishes of the zoological and comparative anatomy collections of the University of Vienna is very thorough, providing morphological details of dozens of species of elasmobranchs, chimaeras, basal actinopterygians, lungfishes, and even agnathans; his paper contains keys to species, and makes constant reference to the works of Cuvier, Günther, Vaillant, and especially Müller & Henle (1841). Concerning dasyatid stingrays, Werner reported on material which had been collected by P. Bleeker and L.K. Schmarda of species, if correctly identified, now classified as Dasyatinae ( Dasyatis pastinaca , Hemitrygon bennetti ), Urogymninae ( Himantura uarnak , Brevitrygon walga , and Urogymnus polylepis ), and Neotrygoninae ( Taeniura lymma ). Garman's (1913) Dasybatus (Pastinachus) torrei clearly equals S. schmardae , a species mentioned by him but not examined. His description highlights typical characters of S. schmardae , such as the denticles (st : Published as part of De Carvalho, Marcelo R., Loboda, Thiago S. & Da Silva, João Paulo C. B., 2016, A new subfamily, Styracurinae, and new genus, Styracura, for Himantura schmardae (Werner, 1904) and Himantura pacifica (Beebe & Tee-Van, 1941) (Chondrichthyes: Myliobatiformes), pp. 201-221 in Zootaxa 4175 (3) on pages 205-214, DOI: 10.11646/zootaxa.4175.3.1, http://zenodo.org/record/267850 : {"references": ["Werner, F. (1904) Die Fische der zoologisch-vergleichend-anatomischen Sammlung der Wiener Universitat. I. Teil. Cyclostomen, Chondropterygier, Ganoiden, Dipnoer. Zoologische Jahrbucher, Abteilung fur Systematik, Geographie und Biologie der Tiere (Jena), 21 (3), 263 - 302.", "Garman, S. (1913) The Plagiostomia (sharks, skates and rays). Memoirs of the Museum of Comparative Zoology at Harvard College, 36, 1 - 528, 77 pl.", "Meek, S. E. & Hildebrand, S. F. (1923) The marine fishes of Panama. Field Museum of Natural History, Publication 215, Zoological Series, Vol. 15, Part 1, 1 - 492.", "Fowler, H. W. (1931) Fishes obtained by the Barber Asphalt Company in Trinidad and Venezuela in 1930. Proceedings of the Academy of Natural Sciences of Philadelphia, 83, 391 - 410.", "Beebe, W. & Tee-Van, J. (1941) Eastern Pacific expeditions of the New York Zoological Society. XXVIII. Fishes from the tropical eastern Pacific. Part 3. Rays, Mantas and Chimaeras. Zoologica, Scientific Contributions of the New York Zoological Society, 26 (3), 245 - 280, pls. 1 - 4.", "Boeseman, M. (1948) Some preliminary notes on Surinam sting rays, including the description of a new species. Zoologische Mededelingen, 30 (2), 31 - 47.", "Bigelow, H. B. & Schroeder, W. C. (1953) Sawfishes, Guitarfishes, Skates and Rays. In: Tee-Van, J., Breder, C. M., Parr, A. E., Schroeder, W. C. & Schultz, L. P. (Eds.), Fishes of the Western North Atlantic. Part 2. Memoirs of the Sears Memorial Foundation for Marine Research, Yale University, New Haven, pp. 1 - 588.", "Cervigon, F. (1992) Tiburones, Peces batoideos y Peces oseos. In: Cervigon, F., Cipriani, R., Fischer, W., Garibaldi, L., Hendrickx, M., Lemus, A. J., Marquez, R., Poutiers, J. M., Robaina, G. & Rodriquez, B. (Eds.), Fichas FAO de identification de especies para los fines de la pesca. Guia de campo de las especies comerciales marinas y de aguas salobres de la costa septentrional de Sur America. Rome, FAO, pp. 163 - 456, Pls. 3 - 40.", "Lovejoy, N. R. (1996) Systematics of myliobatoid elasmobranchs: with emphasis on the phylogeny and historical biogeography of neotropical freshwater stingrays (Potamotrygonidae: Rajiformes). Zoological Journal of the Linnaean Society, 117, 207 - 257. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1996. tb 02189. x", "McEachran, J. D., Dunn, K. & Miyake, T. (1996) Interrelationships of batoid fishes. In: Stiassny, M. L. J., Johnson, G. D. & Parenti, L. (Eds.), Interrelationships of Fishes. Academic Press, San Diego, pp. 63 - 84. http: // dx. doi. org / 10.1016 / B 978 - 012670950 - 6 / 50005 - 9", "McEachran, J. D. & Fechhelm, J. D. (1998) Fishes of the Gulf of Mexico. Volume 1: Myxiniformes to Gasterosteiformes. Univ. of Texas Press, Austin, 1112 pp.", "Lovejoy, N. R., Birminghan, E. & Martin, A. P. (1998) South American rays came in with the sea. Nature, 396, 421 - 422.", "Cervigon, F. & Alcala, A. (1999) Los Peces Marinos de Venezuela. Vol. 5. Tiburones y Rayas. Fundacion Cientifica Los Roques, Caracas, 230 pp.", "Castro-Aguirre, J. L., Espinosa Perez, H. & Schmitter-Soto, J. J. (1999) Ictiofauna estuarino-Lagunar y vicaria de Mexico. Coleccion Textos Politecnicos, Serie Biotechnologias, 711 pp.", "Camargo, M. & Isaac, V. (2001) Os peixes estuarinos da regiao norte do Brasil: lista de especies e consideracoes sobre sua distribuicao geografica. Boletim do Museu Paraense Emilio Goeldi, Nova Serie, Zoologia, 17 (2), 133 - 157.", "McEachran, J. D. & Carvalho, M. R. de (2003) Batoid fishes. In: Carpenter, K. E. (Ed.), The Living Marine Resources of the Western Central Atlantic. Vol. 1. Introduction, Molluscs, Crustaceans, Hagfishes, Sharks, Baroid Fishes, and Chimaeras. FAO Species Identification Guide for Fishery Purposes. American Society of Ichthyologist and Herpetologists Special Publication No. 5. FAO, Rome, pp. 507 - 589.", "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. CRC Press, Boca Raton, pp. 79 - 113. http: // dx. doi. org / 10.1201 / 9780203491317. ch 3", "Carvalho, M. R. de, Maisey, J. G. & Grande, L. (2004) Freshwater stingrays of the Green River Formation of Wyoming (Early Eocene), with the description of a new genus and species and an analysis of its phylogenetic relationships (Chondrichthyes: Myliobatiformes). Bulletin of the American Museum of Natural History, 284, 1 - 136. http: // dx. doi. org / 10.1206 / 0003 - 0090 (2004) 284 % 3 C 0001: FSOTGR % 3 E 2.0. CO; 2", "Carvalho, M. R. de & Lovejoy, N. R. (2011) Morphological and phylogenetic relationships of a remarkable new genus and two new species of Neotropical freshwater stingrays from the Amazon basin (Chondrichthyes: Potamotrygonidae). Zootaxa, 2776, 13 - 48.", "Naylor, G. J. P., Caira, J. N., Jensen, K., Rosana, K. A. M., Straube, N. & Lakner, C. (2012 a) Elasmobranch phylogeny: a mitochondrial estimate based on 595 species. In: Carrier, J. C., Musick, J. A. & Heithaus, M. R. (Eds.), Biology of Sharks and Their Relatives. CRC Press, Boca Raton, Second Edition, pp. 31 - 56. http: // dx. doi. org / 10.1201 / b 11867 - 4", "Aschliman, N. C., Nishida, M., Miya, M., Inoue, J. G., Rosana, K. M. & Naylor, G. J. P. (2012 b) Body plan convergence in the evolution of skates and rays (Chondrichthyes: Batoidea). Molecular Phylogenetics and Evolution, 63 (1), 28 - 42. http: // dx. doi. org / 10.1016 / j. ympev. 2011.12.012", "Salvini-Plawen, L. & Mizzaro, M. (1999) 150 Jahre Zoologie an der Universitat Wien. Verhandlungen der Zoologisch- Botanischen Gesellschaft in Osterreich, 136, 1 - 76.", "Muller, J. & Henle, F. G. J. (1841) Systematische Beschreibung der Plagiostomen. Berlin, Veit, 200 pp. http: // dx. doi. org / 10.5962 / bhl. title. 6906", "Allen, G. R. & Robertson, D. R. (1994) Fishes of the Tropical Eastern Pacific. Crawford House Press, Bathurst, 332 pp. Aschliman, N. C., Claeson, K. M. & McEachran, J. D. (2012 a) Phylogeny of Batoidea. In: Carrier, J. C., Musick, J. A. & Heithaus, M. R. (Eds.), Biology of Sharks and Their Relatives. CRC Press, Boca Raton, Second Edition, pp. 57 - 95. http: // dx. doi. org / 10.1201 / b 11867 - 5", "Bussing, W. A. & Lopez S., M. I. (1994) Demersal and pelagic inshore fishes of the Pacific coast of lower central America. An illustrated guide. Revista de Biologia Tropical, Special Publication, 1 - 164.", "McEachran, J. D. (1995) Dasyatidae. In: Fischer, W., Krupp, F., Schneider, W., Sommer, C., Carpenter, K. E. & Niem, V. H. (Eds.), Guia FAO para la identificacion para los fines de la pesca. Pacifico centro-oriental. Vol. II. Vertebrados. Parte 1. FAO, Rome, pp. 752 - 755."]}