Redescription of Pseudorinelepis Genibarbis (Loricariidae: Hypostominae) with Comments on Behavior As It Relates to Air-Holding
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53 Ichthyol. Explor. Freshwaters, Vol. 10, No.1, pp. 53-61, 4 figs., 1 tab., January 1999 © 1999 by Verlag Dr. Friedrich Pfeil, Miinchen, FRG - ISSN 0936-9902 Redescription of Pseudorinelepis genibarbis (Loricariidae: Hypostominae) with comments on behavior as it relates to air-holding Jonathan W. Armbruster* and Michael Hardman** The loricariid catfish genus Pseudorinelepis is revised. Analysis revealed no difference between the nominal species of Pseudorinelepis and only P. genibarbis is recognized as valid. The Pseudorinelepis from the Rio Branco of Brazil may represent an undescribed species. Pseudorinelepis was examined for its behavior as it relates to the ability of a large esophageal diverticulum to hold air. Pseudorine/epis was shown to breathe air, and, when disturbed, would swim through the water column remaining neutrally buoyant. Introduction logical, and physiological evolution of the lung/ swim bladder in sarcopterygians and basal actin Pseudorinelepis Bleeker is a small genus (4 nomi opterygians. Unfortunately, very little informa nal species) of suckermouth armored catfish from tion is available on the life history characteristics the Amazon basin characterized by a peculiar of members of the Rhinelepis group, and only modification of the digestive tract. At the distal Rhinelepis is known to breathe air (Santos et aI., end of the esophagus is a large, U-shaped diver 1994). It has been suggested that Pseudorinelepis ticulum that was hypothesized by Armbruster and Rhinelepis use their diverticula mainly for (1998b) to be an accessory respiratory organ. This respiration, but may also use it for hydrostatic diverticulum is also found in Pogonopoma Regan, control. The diverticulum in Pogonopoma and Pogonopomoides Gosline, and Rhinelepis Spix Pogonopomoides is similar to a swim bladder and which, together with Pseudorinelepis, represent is suggested to function only as a hydrostatic the Rhinelepis group of Loricariidae (Armbruster, organ (Armbruster, 1998a-b). 1998a-b). The evolution of the diverticulum ap The four species currently recognized in Pseu pears to mirror the evolution of the lung/swim dorinelepis are: P. genibarbis (Valenciennes, 1840), bladder in other fishes, and may represent a use P. agassizii (Steindachner, 1877), P. pellegrini (Re ful model for studying the behavioral, morpho- gan, 1904), and P. carachama (Fowler, 1940) (Arm- * lllinois Natural History Survey, 607 E. Peabody, Champaign, IL 61820, USA. Present address: Department of Zoology and Wildlife Science, 331 Funchess Hall, Auburn University, AL 36849-5414, USA. ** Department of Zoology, Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom. Present address: lllinois Natural History Survey, 607 E. Peabody, Champaign, IL 61820, USA. Ichthvol. Explor. Freshwaters, Vol. 10, No.1 54 bruster & Page, 1997). Despite this small number Pseudorinelepis specimens and provided the im of species, the taxonomic history of Pseudorinel petus to examine the species of Pseudorinelepis, epis is complex. Pseudorinelepis genibarbis was orig examine behavior in aquaria, and to examine the inally described by Valenciennes (in Cuvier & ability to breathe air. In this study, 41 specimens Valenciennes, 1840) in Rhinelepis. Bleeker (1862) of Pseudorinelepis, including all types except that described Pseudorinelepis to include P. genibarbis of P. genibarbis (which is lost) were examined to which is the type species by monotypy; however, determine if P. agassizii, P. carachama, and P. pel Pseudorinelepis was not recognized by subsequent legrini are valid. A neotype is designated for authors until Isbriicker (1980). Steindachner (1877) Rhinelepis genibarbis. The behavior of Pseudorinel described P. agassizii in Rhinelepis. Regan (1904) epis as it relates to buoyancy and air-breathing is placed P. agassizii into the synonymy of P. geni also described. barbis and described P. pellegrini based on minor differences with the types of P. agassizii and P. genibarbis. Regan (1904) placed P. genibarbis and Methods P. pellegrini in the subgenus Pogonopoma of Ple costomus Gronovius (= Hypostomus Lacepede). Ei Morphometric features were measured with dig genmann (1910) described Canthopomus to include ital calipers to the nearest 0.1 mm. Measurements P. genibarbis as the type species, P. agassizii as a are as defined below or as defined in Boeseman synonym of P. genibarbis, and P. pellegrini as val (1968) or Armbruster & Page (1996) and include: id. Fowler (1940) described Monistiancistrus cara standard length, head length (from the snout tip chama and suggested that it was closely related to to the posterior tip of the supraoccipital), clei Ancistrus. Eigenmann & Allen (1942) recognized thral width (width of the body just posterior to all four species currently in Pseudorinelepis; how the posterior margin of the cleithral process that ever, they placed P. agassizii in Canthopomus with is just dorsal to the pectoral fin), snout length, it now erroneously cited as the type species, re orbit diameter, interorbital width, dorsal spine turned P. genibarbis to Rhinelepis and retained length, folded dorsal fin length, dorsal base length P. pellegrini in the subgenus Plecostomus (Pogono (from the point between the dorsal-fin spinelet poma) and P. carachama in Monistiancistrus. Eigen and spine to the posterior end of the membrane mann & Allen (1942) incorrectly suggested that connecting the last dorsal-fin ray to the body), Canthopomus Eigenmann (1910) was a nomen dorsal-caudal length (from the posterior end of nudum. In Eigenmann (1910), the genus name the membrane that connects the last dorsal-fin Canthopomus is clearly stated as a new genus and ray to the body to the posterior edge of the pe described species are indicated as belonging to nultimate dorsal procurrent caudal-fin spine), Canthopomus, thereby satisfying Article 12 of the thorax length (from the insertion of the pectoral International Code of Zoological Nomenclature fin spine to the insertion of the pelvic-fin spine for genus-group names described prior to 1931 [pelvic insertion defined as a bony, subcutane (ICZN, 1985). Gosline (1947) returned P. genibar ous process just anterior to the point where the bis, and P. pellegrini to Canthopomus in which he pelvic-fin spine passes through the skin]), pecto also recognized P. agassizii as valid. Isbriicker ral spine length, abdominal length (from the in (1980) recognized Canthopomus as a synonym of sertion of the pelvic-fin spine to the insertion of Pseudorinelepis, retained P. carachama in Monis the anal-fin spine), pelvic spine length (from the tiancistrus, and concluded that Monistiancistrus insertion of the pelvic-fin spine to the tip of the belonged in the Ancistrinae. Isbriicker & Nijssen spine), postanallength (from the posterior inser (1982) moved Monistiancistrus to the Hypostom tion of the anal fin to the posterior margin of the inae and later placed Monistiancistrus into the penultimate ventral procurrent caudal-fin spine), synonymy of Pseudorinelepis (Nijssen & Isbriick caudal peduncle depth (measured vertically from er, 1986). Isbriicker (1992) removed Monistiancis the posterior margin of the adipose fin mem trus from the synonymy of Pseudorinelepis and brane), anal width (body width at the anal-fin placed it in the synonymy of Hypostomus, and, spine), snout-opercle length, and head width. finally, Armbruster & Page (1997) returned Mon Counts are as in Armbruster & Page (1996) istiancistrus to the synonymy of Pseudorinelepis. except that the unbranched ray of the anal fin Recent collecting efforts in the Iquitos region was counted as a spine and dorsal-caudal plates of Peru has resulted in the collection of several are the number of plates in a dorsal series start- Armbruster & Hardman: Redescription of Pseudorineiepis genibarbis 55 ing from the plate just posterior to the last plate 70 contacted by the dorsal-fin membrane up to and 0 including the elongate plate covering the base of 0 60 0 0 the dorsal-most caudal-fin ray. Values for the 0 neotype are given in parentheses when different. ..c oeg 0 Q) 0 Maximum number of teeth is the number of teeth -Q) l- 50 000 00 in the jaw ramus with the most teeth. Institution 0 0 E 0 abbreviations are as in Leviton et al. (1985) with ~ 00 the addition of HAP, Instituto de Investigaciones E 0000 'x 40 de la Amazonia Peruana, Iquitos, Peru. Numbers ctI o 0 following catalog numbers are the number of ::E. 0 0 specimens examined. Specimens were grouped 30 into five popUlations: the Rio Branco (upper Rio 0 Negro drainage), the Brazilian Amazon (in and 20 near the mainstem Amazon), the Rio Madeira, 0 100 200 300 400 the Peruvian Amazon (in and near the mainstem Amazon from the Brazilian border to the conflu SL (mm) ence of the Rios Marafton and Ucayali), and the Ucayali/Maranon (all collections examined were Fig. 1. Maximum number of teeth vs. standard length from the Rio Ucayali proper or the swampy area in PseudorineZepis genibarbis. between the two rivers near their confluence). The locality of the type of P. pellegrini is given only as Upper Amazon (Regan, 1904); P. pellegrini dorinelepis. Color is variable, but observations on was coded as an unknown population. the live specimens suggests that they may change All measurements were natural log-trans their color to match the substrate. Color patterns formed and a principal components analysis was in live specimens range from tan to almost black, performed using a covariance matrix in Systat and specimens may have spots