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Pimelodella Spelaea: a New Cave Catfish from Central Brazil, with Data on Ecology and Evolutionary Considerations (Siluriformes: Heptapteridae)

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Pimelodella Spelaea: a New Cave Catfish from Central Brazil, with Data on Ecology and Evolutionary Considerations (Siluriformes: Heptapteridae) Copeia, 2004(2), pp. 315±325 Pimelodella spelaea: A New Cave Cat®sh from Central Brazil, with Data on Ecology and Evolutionary Considerations (Siluriformes: Heptapteridae) ELEONORA TRAJANO,ROBERTO E. REIS, AND MARIA ELINA BICHUETTE Pimelodella spelaea, new species, is described from a subterranean stream tributary to the SaÄo Bernardo River inside the SaÄo Bernardo Cave, in the SaÄo Domingos karst area, upper Tocantins River basin, central Brazil. The new species can be distin- guished from its epigean (surface) congeners by derived troglomorphic features such as smaller eyes and fainter coloration, which are nevertheless less reduced than in the other known troglobitic (exclusively subterranean) species, Pimelodella kronei, from southeast Brazil. Ecological data such as habitat characteristics, population data based on mark-recapture techniques, and distributions of frequencies of stan- dard length, weight, and condition factor, were obtained during the dry season of 2000 (May to September). A relatively high population density (about one individual per m22) was recorded for the accessible habitat, which is probably much lower in the nonaccessible, phreatic area of distribution. The low condition factor recorded for P. spelaea, which tended to decrease along the study period, indicates a regime of severe food limitation, intensifying along the dry season. An allopatric model of differentiation is hypothesized, either because of topographic isolation or because of local extinction of epigean relatives, or both factors combined. The conservation status of the new species is also discussed. Pimelodella spelaea, espeÂcie nova, e descrita de um riacho subterraÃneo tributaÂrio do rio SaÄo Bernardo dentro da caverna SaÄo Bernardo, na aÂrea caÂrstica de SaÄo Domingos, alto rio Tocantins, Brasil. A nova espeÂcie pode ser distinguida dos seus congeÃneres epõÂgeos (de superfõÂcie) por caracterõÂsticas troglomoÂr®cas derivadas tais como olhos e pigmentacËaÄo reduzidos. Estes saÄo, entretanto, menos reduzidos que na outra espeÂcie trogloÂbia (exclusivamente subterraÃnea) conhecida, Pimelodella kro- nei, do sudeste do Brasil. Dados ecoloÂgicos tais como caracterõÂsticas do haÂbitat, dados de populacËaÄo baseados em teÂcnicas de marcacËaÄo e recaptura, e distribuicËaÄo de frequÈeÃncias de comprimento-padraÄo, peso e fator de condicËaÄo, foram obtidos durante a estacËaÄo seca de 2000 (maio a setembro). Uma densidade populaciona relativamente alta (cerca de 1 ind. m22) foi registrada na porcËaÄo acessõÂvel do haÂbitat da espeÂcie, que e possivelmente muito menor na aÂrea freaÂtica, naÄo acessõÂvel da sua distribuicËaÄo. Os baixos fatores de condicËaÄo registrados para P. spelaea, que tende- rem a diminuir ao longo do perõÂodo de estudo, indicam um regime de severa li- mitacËaÄo alimentar, intensi®cando-se ao longo da estacËaÄo seca. Um modelo alopaÂtrico de diferenciacËaÄo e sugerido, seja pelo isolamento topogra®co ou pela extincËaÄo local de parente epõÂgeos, ou ainda por ambos os fatores combinados. O estado de con- servacËaÄo da nova espeÂcie tambeÂm e discutido. HE Neotropical region is remarkable for its prevalence of these groups in the subterranean T rich subterranean ichthyofauna, only com- biotope is an evidence of high potential for col- parable to that from China and southeastern onization of the subterranean habitats, includ- Asia. More than 35 among approximately 100 ing caves, probably facilitated by their predom- troglobitic (exclusively subterranean) ®sh spe- inantly nocturnal activity and generalized car- cies occurring around the world are known nivorous (heptapterids, trichomycterids) or de- from Mexico, Caribbean, Venezuela, Ecuador, tritivorous (Ancistrus) diet (Trajano, 2001). Peru, Bolivia and Brazil, including several un- Troglobitic species are generally characterized described taxa (Weber et al., 1998; Romero and by autapomorphies related to the isolation in Paulson, 2001; ET and MEB, unpubl. data). The the perpetually dark subterranean habitat (tro- great majority are siluriforms belonging to the glomorphisms), the most widespread being the Heptapteridae (Rhamdia and Pimelodella, among reduction or even loss of eyes and pigmenta- others), Trichomycteridae (Trichomycterus and tion. Ituglanis), and Loricariidae (Ancistrus). The The SaÄo Domingos karst area, a limestone re- q 2004 by the American Society of Ichthyologists and Herpetologists 316 COPEIA, 2004, NO. 2 gion in Central Brazil crossed by tributaries of marked by subcutaneous injection of ¯uores- the upper Tocantins River, Amazon basin, pre- cent pigments (``Photonics marking,'' from sents the most diversi®ed cave ichthyofauna in New West Co.), and released. Association of the country. In the course of our studies in this mark color and position, and ®sh size, pro- area, we discovered a new cave species of hep- duced individual marks. Some specimens were tapterid cat®sh, genus Pimelodella, living in slow- sacri®ced by overanesthesia and preserved in moving pools in an upper conduit of the SaÄo formalin (later transferred to alcohol 70GL) for Bernardo Cave. Herein we describe this new description. Only eight specimens were pre- species and present data on population ecology served to avoid unnecessary mortality in the gathered during the dry season of 2002. This is population, whose total size was unknown. the second troglobitic species known for the ge- Mark-recapture data from May to July and nus, the ®rst one being Pimelodella kronei (Ribei- from July to September were used separately to ro, 1907), from southeastern Brazil. estimate the number of individuals in the study The genus Pimelodella, with 71 valid species section using the Lincoln-Petersen Index (Be- and a cis- and trans-Andean distribution from gon, 1979; Blower et al., 1981). We did not use Panama to Paraguay and southern Brazil, is the the whole set of data for a single estimation us- largest among the Heptapteridae (Bockmann ing a multiple-recapture method (such as the and Guazelli, 2003). Pimelodella is currently in triple-catch) because there is evidence of some need of a taxonomic revision being last re- loss of marks by pigment reabsorption during viewed by Eigenmann (1917) who diagnosed our four-month observation period. the genus based on several not unique features Standard length and weight data were used but including a few that provide a diagnosis to calculate the allometric condition factors (K when in combination. These features are ``. 5 100.W.SL2b; Le Cren, 1951). For these cal- a long, adnate, adipose ®n; caudal ®n deeply culations, units of millimeters and grams were forked; a frontal and a parietal fontanel, the lat- used. The power coef®cient (b) was estimated ter reaching the base of the occipital process, from the slope of the regression of log weight which is narrow and reaches, or nearly reaches on log standard length using the pooled data. the plate in front of the dorsal; and humeral Dice-Leraas diagrams (Dice and Leraas, 1936) process spine-like ...''(Eigenmann, 1917:229). were used to show tendencies along the study More recently, however, Bockmann (1998) pre- period. Graphs with the distributions of fre- sented two characters that may represent syna- quencies of size (SL), weight and condition fac- pomorphies for Pimelodella or to a subgroup of tor (K) in July and in September were built; May species within Pimelodella. These are (1) the two was not included here because of the low sam- innermost caudal-®n rays not directly articulat- ple size. July and September samples were com- ing with hypural bones (character 220), and (2) pared by two-sample t-test, with a55%. those same rays united by an incomplete caudal- For comparison, we estimated the average ®n membrane (character 223). The new species condition factor for P. kronei from the Areias described below shares the second character cave, upper Ribeira karst area, in the Atlantic but not the ®rst, because the two innermost cau- rain forest in southeastern Brazil, based on data dal-®n rays directly articulate with the hypural gathered between May and September of 1984 bones. and 1985 (ET, unpubl. data), a period of low rainfall locally corresponding to the dry season. MATERIALS AND METHODS Measurements of preserved ®sh, taken with electronic calipers (0.1 mm precision), followed The ®eldwork was conducted in May, July, basically Lundberg et al. (1991) and Trajano and September 2000 and was restricted to the and Britski (1992). In the description, meristic dry season because of the danger of being data are followed by number of specimens dis- caught by ¯oods in the rainy period (October playing each count in parentheses. Density of to March). Collections were made with hand chromatophores was estimated by direct counts nets in May and with funnel-type minnow traps under stereomicroscope on a 1-mm2 area locat- baited with chicken liver in July and September. ed immediately posterior of the dorsal-®n ori- Baits were kept in small nylon bags to avoid be- gin, midway between the lateral line and the ing swallowed by the ®sh and interfering with dorsal ®n. A cleared-and-stained specimen was weight measurements. Fish were anesthetized in prepared according to Taylor and Van Dyke a solution of MS-222 (tricaine-methanesulpho- (1985). nate), measured (standard length, SL) to the No Pimelodella species has been described nearest 0.1 mm, and weighed with a Pesola dy- from the upper Tocantins River to date. How- namometer, 0.1 g precision. Most cat®shes were ever, watersheds around the upper Tocantins in- TRAJANO
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