Copeia 2013, No. 4, 641–646
A New Species of Listrura from Laguna dos Patos System, Brazil: The Southernmost Record of the Glanapteryginae (Siluriformes: Trichomycteridae) Leandro Villa-Verde1, Juliano Ferrer2, and Luiz R. Malabarba2 Listrura depinnai, new species, is described from a small swampy area belonging to Laguna dos Patos system, Municipality of Viama ˜ o, Rio Grande do Sul state, southern Brazil. It occurs within the wildlife reserve Refu ´ gio de Vida Silvestre Banhado dos Pachecos, and represents the southernmost record of the glanapterygine catfishes. The new species is easily distinguished from remaining Listrura by having the following putative autapomorphies: the region below the midline of flank covered by several spots (vs. unpigmented or with a slender row of small spots); the caudal peduncle at procurrent-ray folds deeper than the preanal portion of body (vs. approximately the same depth); and the presence of a conspicuous intumescence adjacent to the posterodorsal margin of the pectoral-fin base (vs. swelling slight or absent). Putative phylogenetic positions of Listrura depinnai are herein discussed. Listrura depinnai, espe´cie nova, e´ descrita de uma pequena a´rea alagada pertencente ao sistema da Laguna dos Patos, Municı´pio de Viama ˜ o, Estado do Rio Grande do Sul, sul do Brasil. A espe´cie ocorre dentro da unidade de conservac¸a ˜o Refu ´ gio de Vida Silvestre Banhado dos Pachecos, e representa o registro mais ao sul dos bagres glanapterigı´neos. A nova espe´cie e´ facilmente distinguida das demais espe´cies de Listrura pelas possı´veis autapomorfias a seguir: regia ˜ o abaixo da linha mediana do flanco coberta por va´rias pintas (vs. despigmentada ou com uma faixa de pequenas pintas); pedu ´ nculo caudal na regia ˜ o das membranas dos raios procurrentes mais alto do que a porc¸a ˜ o pre´-anal do corpo (vs. aproximadamente a mesma altura); e presenc¸a de uma conspı´cua intumesce ˆ ncia adjacente a` margem po ´ stero-dorsal da base da nadadeira peitoral (vs. levemente inchada ou ausente). Possı´veis posic¸o ˜ es filogene´ticas de Listrura depinnai sa ˜o aqui discutidas.
T
HE Glanapteryginae includes four genera, three of which occur in the Rio Amazonas and Rio Orinoco basins (i.e., Glanapteryx, Pygidianops, and Typhlobelus; Myers, 1927, 1944; Nico and de Pinna, 1996; Schaefer et al., 2005). The fourth genus, Listrura, occurs in the Atlantic Forest of southeastern and southern Brazil (de Pinna and Wosiacki, 2002) and is recognized as the sister group of the remaining genera (de Pinna, 1988, 1998). The knowledge about the biodiversity of the glanapterygines is markedly incomplete (de Pinna and Wosiacki, 2002, 2003), inasmuch as most species of the subfamily (ten of the 17) were only described this century. Listrura includes six species of small size catfishes: L. boticario de Pinna and Wosiacki, L. camposi (Miranda Ribeiro), L. costai Villa-Verde, Lazzarotto, and Lima, L. nematopteryx de Pinna, L. picinguabae Villa-Verde and Costa, and L. tetraradiata Landim and Costa (Villa-Verde et al., 2012). Those species inhabit interstitial environments and are usually found in small and shallow water bodies characterized by leaf litter and soft mud bottom, commonly associated with small mountain streams (Nico and de Pinna, 1996). The recent discoveries of new populations of Listrura are possibly a consequence of sampling directed to these habitats (Villa-Verde et al., 2012). Until now, the southernmost record of Listrura was from a small coastal stream in Floriano´polis, Santa Catarina state, Brazil (Nico and de Pinna, 1996; Villa-Verde et al., 2013). A recent inventory of the fish species from a swamp close to a dense fragment of wetland forest, forming the Rio Gravataı´, Laguna dos Patos system, near Porto Alegre, Rio Grande do
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Sul state, Brazil, revealed the presence of a distinctive new species of Listrura, representing the southernmost record of the genus and subfamily. Repeated collecting efforts made monthly for one year in the type locality yielded only two specimens. The distinctiveness of the species and its biogeographic and conservation relevance, however, lead us to describe the new taxon even with the few available specimens. MATERIALS AND METHODS Measurements were taken point-to-point using a digital caliper, according to de Pinna (1988), except for body width, caudal peduncle width, eye diameter, and mouth width which were taken following Costa (1992). Measurements are presented as percents of standard length (SL), except for subunits of head, which are presented as percents of head length (HL). Counts of premaxillary and dentary teeth, branchiostegal rays, opercular and interopercular odontodes, and pectoral-fin rays were taken from alcoholpreserved specimens. Counts of anal- and caudal-fin rays, and respective dorsal and ventral procurrent rays were taken on both radiographs and alcohol-preserved specimens, with the higher values reported. Counts of vertebrae and ribs, as well as osteological observations were taken only from radiographs. Number of vertebrae excludes those incorporated into the Weberian apparatus and considers the caudal complex as one element. Number of principal caudal-fin rays includes all those articulating with the hypurals; counts are given for each lobe (upper first) separated by a plus sign
Setor de Ictiologia, Departamento de Vertebrados, Museu Nacional da Universidade Federal do Rio de Janeiro, 20940-040 Rio de Janeiro, RJ, Brazil; E-mail:
[email protected]. Send reprint requests to this address. 2 Departamento de Zoologia, Instituto de Biocieˆncias, Universidade Federal do Rio Grande do Sul, Av. Bento Gonc¸alves 9500, 91501-970 Porto Alegre, RS, Brazil; E-mail: (JF)
[email protected]; and (LRM)
[email protected]. Submitted: 12 March 2013. Accepted: 27 May 2013. Associate Editor: R. E. Reis. DOI: 10.1643/CI-13-027 F 2013 by the American Society of Ichthyologists and Herpetologists
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Fig. 1. Listrura depinnai, holotype, UFRGS 17135, 30.8 mm SL, Brazil, Rio Grande do Sul state, Municipality of Viama˜o, in lateral (top), dorsal (middle), and ventral (bottom) views.
(+), with Roman numeral representing the unbranched rays and Arabic numeral representing the branched rays. Counts of the holotype were marked with an asterisk (*). Institutional abbreviations follow Sabaj Pe´rez (2013). Map was performed on Geographical Information System (GIS) with Quantum GIS version 1.8.0 software (Sherman et al., 2013). Listrura depinnai, new species Figures 1, 2; Table 1 Holotype.—UFRGS 17135, 30.8 mm SL, Brazil, Rio Grande do ´ guas Claras, Rio Gravataı´ Sul state, Municipality of Viama˜o, A basin, Laguna dos Patos system, swamp on the wildlife reserve Refu´gio de Vida Silvestre Banhado dos Pachecos, 30u059520S, 50u519050W, J. Wingert and J. P. Miranda, 21 October 2011. Paratype.—UFRGS 16383, 1, 27.7 mm SL, collected with the holotype. Diagnosis.—Listrura depinnai is easily distinguished from its congeners by the following putative autapomorphies: the region below the midline of the flank is covered by several spots (Fig. 1, vs. unpigmented in L. boticario, L. camposi, L. costai, and L. tetraradiata or with a slender row of small spots in L. nematopteryx and L. picinguabae); the caudal peduncle, including procurrent-ray folds, is deeper than the preanal portion of body (Fig. 1 top, vs. approximately same depth);
and there is a conspicuous intumescence adjacent to the posterodorsal margin of the pectoral-fin base (Fig. 1 middle and bottom, vs. swelling slight or absent). It shares the absence of a dorsal fin with L. boticario and the glanapterygines Glanapteryx, Pygidianops, and Typhlobelus, but can be readily distinguished from the former species by the features mentioned above and from the remaining genera by having a larger eye (9.0–10.5% HL vs. less than 6.0%, or absent) and a longer pectoral fin (11.1–11.6% SL vs. less than 5.0%, or absent). The new species differs further from all species of Listrura, except L. tetraradiata, by having a low number of vertebrae (45–46 vs. 48–55). It differs further from L. tetraradiata by having more dorsal and ventral procurrent caudal-fin rays (33–39 and 31–34 rays vs. 21–28 and 20–24, respectively), only unbranched rays on the anal and pectoral fins (vs. many branched rays), and the sensory pore of the sphenotic branch absent (vs. present). It differs further from L. costai, L. nematopteryx, and L. picinguabae by having more pectoral-fin rays (two or three vs. one) and a slightly elongated, relatively deep body (body depth 12.2–12.7% SL vs. less than 10.5%). It differs further from L. camposi by having all caudal elements almost entirely fused (Fig. 2, vs. hypurals 1+2 not fused to hypurals 3+4+5). Description.—Morphometric data in Table 1. Body elongate, subcylindrical on anterior portion of trunk, to strongly compressed on caudal peduncle. Head broad, wider than body. Dorsal body profile from head to above anal-fin origin
Villa-Verde et al.—A new trichomycterid catfish
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Fig. 2. Radiographic image of Listrura depinnai, holotype, UFRGS 17135, 30.8 mm SL.
and ventral preanal profile straight. Dorsal and ventral postanal profiles (including procurrent-ray folds) slightly convex, with small concavity where joining to caudal fin. Anterior portion of caudal peduncle excluding procurrent rays with fold as deep as body, gradually tapering toward caudal skeleton. Greatest depth including procurrent-ray folds about 15% deeper than preanal portion of body. Dorsal keel gently increasing posterior to middle of body length onto anteriormost dorsal procurrent ray and continuous with respective fold. Anal opening on posterior two-thirds of standard length, with integument surrounding vent. Small ventral keel from middle of body length to anterior margin of anus. Conspicuous intumescence (axillary organ?) adjacent to posterodorsal margin of pectoral-fin base (see Discussion). Skin covered with minute papillae. Table 1. Morphometric Data of Holotype and Paratype of Listrura depinnai.
Holotype
Paratype
30.8 4.0
27.7 3.8
Subunits of body (%) Total length Body depth Caudal peduncle depth Body width Caudal peduncle width Caudal peduncle length Anal-fin base length Pectoral-fin length Preanal length Head length
119.8 12.7 14.2 4.0 1.5 24.2 5.0 11.6 71.5 13.0
116.3 12.3 14.5 4.2 1.7 25.3 5.7 11.1 69.5 13.6
Subunits of head (%) Head depth Head width Interorbital width Preorbital length Eye diameter Mouth width Anterior internarial width Posterior internarial width Nasal barbel length Maxillary barbel length Rictal barbel length
39.5 85.7 27.0 35.3 10.5 50.0 38.3 24.8 103.7 100.3 95.8
40.9 91.1 27.6 38.7 9.0 50.1 32.4 19.6 90.7 92.3 87.8
Standard length (mm) Head length (mm)
Head depressed and trapezoidal. Snout blunt and slightly elongate. Mouth subterminal, narrow, and straight. Upper lip narrow and continuous with dorsal surface of head. Lower lip fleshy, well differentiated from ventral surface of head. Premaxilla with at least 22 teeth, dentary with at least 18 teeth; all conical, tips pointed and curved, surrounded by fleshy rim of integument, and distributed on two rows in jaw. Eye well formed, anteriorly located on head, nearer snout tip than to opercular patch of odontodes. Nasal, maxillary and rictal barbels well developed. Tip of nasal barbel reaching vertical through pectoral-fin base. Tips of maxillary and rictal barbels reaching to posterior margin of opercular patch of odontodes. Anterior nostril opening anteromesially at base of nasal barbel. Posterior nostril round and opening midway between anterior nostril and eye. Low rim of integument surrounding posterior nostril, longer anteriorly. Interopercle with at least four odontodes, opercle with at least seven odontodes; all odontodes conical, tips pointed and slightly curved, and surrounded by fleshy rim of integument. Dorsal fin, pelvic fin, and pelvic girdle absent (Fig. 2). Anal fin small and round on distal margin, with six segmented unbranched rays plus one or two anterior unsegmented rays embedded in integument. Anal-fin origin adjacent to posterior margin of anus, continuous with integument surrounding vent, and at vertical through 29th vertebra. Caudal fin elongate and round on posterior margin. Principal caudal-fin rays i,5+5,i* or ii,3+4,ii, dorsal procurrent rays 33*–39, ventral procurrent rays 31*–34. Pectoral fin narrow with two segmented unbranched rays (holotype with three rays in right side); first about 50% longer than second. Ribs 2. Total vertebrae 45* or 46. Branchial membranes attached only at anteriormost point of isthmus. Six branchiostegal rays (holotype with seven rays in right side) visible through overlying skin; lateralmost ray behind interopercular patch of odontodes. Four visible pores, all paired, on laterosensory system. Two anteriormost pores corresponding to cephalic system, located above dorsal margin of opercular patch of odontodes, and homologous to preopercular and postotic canal pores, respectively. Two posteriormost pores located immediately posterior of head and above pectoral fin, and homologous to two pores of main lateral line; posterior smaller than anterior. Coloration.—Color in alcohol-preserved specimens very similar to coloration in life. General body color yellowish with vermicular or roundish, partly coalescent, dark brown spots over entire flank and dorsum, more concentrated along lateral midline forming discontinuous row of dark brown
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Fig. 3. Current geographic distribution of Listrura. All localities in Brazil: (1) swamp in Laguna dos Patos system, Municipality of Viama˜o, Rio Grande do Sul state; (2) stream in Ribeira˜o da Ilha, Municipality of Floriano´polis, Santa Catarina state; (3) coastal streams draining to Paranagua´ Bay, Municipalities of Guaraquec¸aba and Antonina, Parana´ state; (4) streams in Rio Ribeira de Iguape basin, Municipalities of Juquia´ and Pedro de Toledo, Sa˜o Paulo state; (5) streams in coastal Rio da Fazenda drainage, Picinguaba, Municipality of Ubatuba, Sa˜o Paulo state; (6) streams in coastal Rio Jurumirim drainage, Municipality of Angra dos Reis, Rio de Janeiro state; (7) coastal streams draining to Guanabara Bay, Municipality of Mage´, Rio de Janeiro state; (8) stream in Rio Sa˜o Joa˜o basin, Municipality of Cachoeiras de Macacu, Rio de Janeiro state; (9) stream in Lagoa de Araruama system, Municipality of Saquarema, Rio de Janeiro state. Drainages are drawn only to Brazil. Delimitation of the biomes was based on IBGE (2004). Listrura sp. marked in locality (2) is an undetermined-status population as discussed in Villa-Verde et al. (2013). Listrura sp. marked in locality (8) is a potential new species and equals the Listrura sp. previously referred in Villa-Verde et al. (2012).
blotches; coalesced dots above lateral midline of trunk and dorsum larger and denser than below lateral midline (Fig. 1 top). Some dark brown coalesced dots above and below lateral midline of caudal peduncle usually arranged in oblique rows, outlining myomeres. Abdomen whitish with few small dark brown marks (Fig. 1 bottom). Dorsal and lateral surfaces of head predominantly brown to black, with yellowish areas on ventrolateral region of eye and on mesial region between eyes and anterior nostrils in dorsal view (Fig. 1 middle). Unpigmented areas on part of cheek posterior of maxillary barbel, on posterior region of interopercular and opercular patches of odontodes, and on branchial membrane. Margins of posterior nostril, sensory pores, and neuromasts finely delineated in white. Nasal barbel whitish hyaline, with elongate scattering of dark chromatophores along posterior margin of dorsal side on basal half; ventral side with sparse dark fields near base. Maxillary and rictal barbels with similar pigmentation pattern, but streak outlining anterior margin of dorsal side and faintest on latter. Ventral surface of head whitish with irregular dark brown blotches over mental region. Lower lip whitish, with dark coalesced dots at symphysis. Pectoral fin hyaline with faint streak of minute dark brown dots outlining inter-ray region. Anal fin hyaline with brown blotches on base. Region of procurrent caudal-fin rays hyaline with sparse yellowish to dark brown fields, denser on base. Caudal fin hyaline with concentration of dark brown chromatophores on base, faint dark streaks outlining inter-ray region on anterior half of fin, and two semicircular vertical bands of dark brown chromatophores on posterior half of fin; posteriormost weakly distinguished.
Distribution.—Listrura depinnai is known from its type locality, a swamp on the wildlife reserve Refu´gio de Vida Silvestre Banhado dos Pachecos, Rio Gravataı´ basin, Laguna dos Patos system, Brazil (Fig. 3). Habitat and ecological notes.—The swamp where the new species was collected had clear black water with mud and litter bottom, covered with a large amount of emerging vegetation adjacent to a dense and isolated fragment of wetland forest. The level of the swamp varies along the year, being flooded from April to September and becoming almost completely dry from October to March. The two specimens of Listrura depinnai were captured at a shallow site (about 10 cm water depth above nearly 50 cm depth of submerged litter deposits) among the emerging vegetation. During the collection, the water temperature was 25.5uC with pH 5.25 at approximately 1100 hr. A species of Cynopoecilus and Gymnotus aff. pantherinus (both potentially new species) were collected in the same swamp. Etymology.—The species epithet is given for Ma´rio C. C. de Pinna, in recognition of his significant contributions to the knowledge of fish systematics, especially on trichomycterid catfishes. DISCUSSION Although only two specimens are currently available, Listrura depinnai is easily diagnosed and identified by its external morphology. Its pigmentation is denser than the remainder species of Listrura, especially over the region below the midline
Villa-Verde et al.—A new trichomycterid catfish
of the flank (Fig. 1 top; for comparisons, see figures in de Pinna, 1988:116; Landim and Costa, 2002:153; de Pinna and Wosiacki, 2002:721; Villa-Verde and Costa, 2006:45; VillaVerde et al., 2012:529). Listrura picinguabae and L. nematopteryx also display brown to dark spots below the midline of flank, but these are sparse and are aligned to form a slender longitudinal row just above the dorsal limit of the abdomen. A conspicuous sac-like intumescence on posterodorsal margin of pectoral-fin base is present in Listrura depinnai (Fig. 1 middle, bottom). Similar structures with different sizes are also observed in other trichomycterids, including many sarcoglanidines (Myers and Weitzman, 1966; de Pinna, 1989a) and the miniature catfishes Trichomycterus hasemani (Eigenmann), T. johnsoni (Fowler), and T. anhanga Dutra, Wosiacki, and de Pinna. Whereas those miniature trichomycterids have been recently hypothesized to be closely related to the derived clade formed by Tridentinae, Stegophilinae, Vandelliinae, Glanapteryginae, and Sarcoglanidinae (Wosiacki, 2002; Dutra et al., 2012), and the two latter subfamilies comprise a monophyletic clade (Baskin, 1973; de Pinna, 1998; Datovo and Bockmann, 2010), this structure can supply an additional character for better understanding of the relationships among these taxa. Nevertheless, it is unclear if it is a hyper-developed axillary glandular organ or another structure. The shape of the caudal peduncle in Listrura depinnai is peculiar, being deeper on anterior half than in other species of the genus. The neural and hemal spines of the first caudal-peduncle vertebrae are at least three times longer than respective vertebral-centrum length in L. depinnai (Fig. 2), contrasting with the condition present in congeners, in which the corresponding spines are two times (or less) longer than respective vertebral-centrum length. The caudal-peduncle shape as well as the pigmentation and intumescence characters described above are apparently autapomorphies for the new species. The phylogenetic position of Listrura depinnai is not as obvious as its distinctiveness. Its inclusion in the Glanapteryginae is confirmed by the following synapomorphies proposed by de Pinna (1988, 1989b), with further modifications by Costa and Bockmann (1994), Landim and Costa (2002), and Schaefer et al. (2005): (1) four or fewer pectoralfin rays; (2) ten or fewer principal branched rays on caudal fin (Pygidianops and Typhlobelus display a further derived condition of all rays unbranched); and (3) all caudal elements almost entirely fused (for specific condition in L. camposi, see Villa-Verde et al., 2013). Synapomorphies for Listrura are scarce since its description by de Pinna (1988). As discussed by de Pinna and Wosiacki (2002), the extremely short cephalic laterosensory system restricted to pterotic branch requires further investigation, since some Pygidianops and Typhlobelus display a similar condition (see Schaefer et al., 2005) and L. tetraradiata displays a somewhat longer system, entering the sphenotic bone (see Landim and Costa, 2002). Two other proposed synapomorphies, the bottle-shaped vomer (de Pinna, 1988) and the dorsally curved posterolateral process of the autopalatine (Villa-Verde et al., 2013), cannot be seen on radiographs of L. depinnai. Its placement into Listrura among the glanapterygine genera is justified by the general morphological aspect, such as the general color pattern, the presence of opercular and interopercular odontodes in adult individuals, and the procurrent-ray folds extending anteriorly with numerous rays (24–39 dorsally and 21–35 ventrally; for Glanapteryx, see Landim and Costa, 2002).
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Although the inclusion of the new species in Listrura is tentative, L. depinnai shares the absence of dorsal fin with L. boticario and with one undescribed species listed by VillaVerde et al. (2012). This character is considered apomorphic by de Pinna and Wosiacki (2002), suggesting a close relationship among the three species. The lower number of vertebrae found in L. depinnai (45–46, Fig. 2) and L. tetraradiata (43–45), however, is interpreted as a plesiomorphic condition in the genus (see Landim and Costa, 2002), while the remaining species of Listrura present 50–55 vertebrae, rarely 48 or 49. It suggests a basal position for L. depinnai and L. tetraradiata. Villa-Verde et al. (2012) further found (L. picinguabae, L. costai (L. nematopteryx, Listrura sp.)) forming a clade supported by molecular evidence, but such information is unavailable for L. depinnai and L. boticario. Those four species, however, share at least one morphological synapomorphy: the presence of a single ray on the pectoral fin. Listrura depinnai and L. boticario have two or three pectoral-fin rays, refuting their inclusion in that clade. All populations of Listrura previously known occur in the Atlantic Forest biome associated with forested areas. In contrast, L. depinnai is restricted to a transition area between the Atlantic Forest and Pampa biomes (IBGE, 2004), as shows the Figure 3. The type locality of L. depinnai is within a dense isolated fragment of wetland forest; an uncommon environment in the predominantly grassland Pampa biome wherein dense forests represent about 5% of whole biome (IBGE, 2004). Thus, that transition area is probably the southernmost distributional limit of the genus and it is unlikely that it occurs widely in the Pampa biome. MATERIAL EXAMINED Trichomycteridae. ‘‘Trichomycterinae’’: Trichomycterus hasemani: LBP 4198, 160 (10 CS). Sarcoglanidinae: Microcambeva barbata: UFRJ 684, 2 CS paratypes; MZUSP 43679, 1 paratype; MNRJ 37572, 3. Microcambeva ribeirae: MNRJ 14304, 4 (1 CS) paratypes; MNRJ 32443, 39 (2 CS); MNRJ 37165, 1. Glanapteryginae: Glanapteryx anguilla: MZUSP 36530, 21 (2 CS). Listrura boticario: MHNCI 11689, 6 (1 CS); MHNCI 11690, 2 (1 CS); MNRJ 32442, 1; MNRJ 32444, 21 (2 CS); MZUSP 69573, holotype. Listrura camposi: MNRJ 37023, 6 (2 CS); MNRJ 33031, 14 (1 CS); MZUSP 3426, holotype; MZUSP 95189, 25 (2 CS); UFRJ 1278, 1; UFRJ 1279, 1 CS. Listrura costai: MNRJ 31917, holotype; MNRJ 31535, 3 (2 CS) paratypes; MNRJ 31918, 5 paratypes; UFRJ 7214, 3 paratypes; UFRJ 7215, 4 paratypes; UFRJ 6577, 3 CS paratypes. Listrura nematopteryx: MNRJ 10970, 8 (1 CS) paratypes, UFRJ 0707, 9; UFRJ 5952, 5 CS. Listrura picinguabae: MCP 38921, 2 paratypes; UFRJ 6111, holotype; UFRJ 5948, paratype; UFRJ 5949, 2 paratypes; UFRJ 5950, 15 paratypes; UFRJ 5951, 4 CS paratypes; UFRJ 5991, 2 CS paratypes; UFRJ 6138, 5 CS paratypes. Listrura sp.: MNRJ 32026, 46 (3 CS); MZUSP 93882, 21 (2 CS). Listrura tetraradiata: MNRJ 31534, 17 (4 CS) paratypes; MNRJ 39068, 6; MZUSP 52572, holotype; UFRJ 4586, 17 paratypes; UFRJ 4588, 3 CS paratypes; UFRJ 4590, 7 paratypes. Pygidianops amphioxus: INPA 35500, 3 paratypes; INPA 35503, 10 paratypes. ACKNOWLEDGMENTS Special thanks to J. Wingert and J. Miranda for their help in field work. We are also grateful to P. Buckup for providing
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the radiographic images of the specimens, and to M. de Pinna for important discussions about the new species. We acknowledge the Secretaria do Meio Ambiente do Estado do Rio Grande do Sul that issued the license to develop collection and research in the state reserve Refu´gio de Vida Silvestre Banhado dos Pachecos. This work was also supported by CNPq (Conselho Nacional de Desenvolvimento Cientı´fico e Tecnolo´gico, Ministe´rio de Cieˆncia e Tecnologia, Process 300705/2010-7 to JF and 457503/2012-2 to LRM), CAPES (Coordenac¸a˜o de Aperfeic¸oamento de Pessoal de Nı´vel Superior), and FAPERJ (Fundac¸a˜ o de Amparo a` Pesquisa do Estado do Rio de Janeiro). LITERATURE CITED Baskin, J. N. 1973. Structure and relationships of the Trichomycteridae. Unpubl. Ph.D. diss., City University of New York, New York. Costa, W. J. E. M. 1992. Description de huit nouvelles espe`ces du genre Trichomycterus (Siluriformes: Trichomycteridae), du Bre´sil oriental. Revue Franc¸aise d’Aquariologie et Herpe´tologie 18:101–110. Costa, W. J. E. M., and F. A. Bockmann. 1994. A new genus and species of Sarcoglanidinae (Siluriformes: Trichomycteridae) from southeastern Brazil, with a re-examination of subfamilial phylogeny. Journal of Natural History 28:715–730. Datovo, A., and F. A. Bockmann. 2010. Dorsolateral head muscles of the catfish families Nematogenyidae and Trichomycteridae (Siluriformes: Loricarioidei): comparative anatomy and phylogenetic analysis. Neotropical Ichthyology 8:193–246. Dutra, G. M., W. B. Wosiacki, and M. C. C. de Pinna. 2012. Trichomycterus anhanga, a new species of miniature catfish related to T. hasemani and T. johnsoni (Siluriformes: Trichomycteridae) from the Amazon basin, Brazil. Neotropical Ichthyology 10:225–231. IBGE (Instituto Brasileiro de Geografia e Estatı´stica). 2004. Mapa dos Biomas e de Vegetac¸a ˜ o (21 March 2004). Rio de Janeiro. IBGE, Diretoria de Geocieˆncias. Electronically accessible at http://www.ibge.gov.br/home/presidencia/ noticias/21052004biomashtml.shtm (accessed 28 February 2013). Landim, M. I., and W. J. E. M. Costa. 2002. Listrura tetraradiata (Siluriformes: Trichomycteridae): a new glanapterygine catfish from the southeastern Brazilian coastal plains. Copeia 2002:152–156. Myers, G. S. 1927. Descriptions of new South American fresh-water fishes collected by Dr. Carl Ternetz. Bulletin of the Museum of Comparative Zoology 68:107–135. Myers, G. S. 1944. Two extraordinary new blind nematognath fishes from the Rio Negro, representing a new subfamily of Pygidiidae, with a rearrangement of the genera of the family, and illustrations of some previously described genera and species from Venezuela and Brazil. Proceedings of the California Academy of Sciences 23:591–602. Myers, G. S., and S. H. Weitzman. 1966. Two remarkable new trichomycterid catfishes from the Amazon basin in Brazil and Colombia. Journal of Zoology (London) 149:277–287. Nico, L. G., and M. C. C. de Pinna. 1996. Confirmation of Glanapteryx anguilla (Siluriformes, Trichomycteridae) in the Orinoco River basin, with notes on the distribution
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