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First Description of Small Juveniles of the Primitive Catfish Diplomystes (Siluriformes: Diplomystidae)

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First Description of Small Juveniles of the Primitive Catfish Diplomystes (Siluriformes: Diplomystidae) 71 Ichthyol. Explor. Freshwaters, Vol. 15, No. 1, pp. 71-82, 8 figs., 2 tabs., March 2004 © 2004 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902 First description of small juveniles of the primitive catfish Diplomystes (Siluriformes: Diplomystidae) John G. Lundberg*, Tim M. Berra** and John P. Friel*** At 13 mm SL young Diplomystes nahuelbutaensis are free living and, judging by their greatly diminished yolk mass, probably foraging. External development of form from these small fish to subadults is smooth and continuous, without distinct juvenile stages or specializations. The major changes of body and fin shape shown by Diplomystes are those common to most siluriforms and other fishes: juveniles have large heads and eyes, and expanded posterior median-fin membranes relative to larger specimens. Pigmentation pattern is similar in juveniles and adults in this species. Definitive pectoral- and dorsal-fin spines, and fin ray counts, except for caudal-fin procurrent rays, are established before firm ossification and segmentation of lepidotrichia. The postcleithral process develops relatively late. Introduction The largest specimen of D. nahuelbutaensis col- lected with the juveniles is 131 mm SL, thus yield- In December, 1992 T. M. Berra and V. H. Ruiz ing a growth series from a local population that collected specimens of the catfish Diplomystes na- ranges across one order of magnitude (Fig. 3). huelbutaensis in the clear, rocky and fast-flowing Because of the wide interest in Diplomystes as a Río Laja (Fig. 1) of central Chile, ca. 37° S (Ruiz & phylogenetically basal, relatively rare and poten- Berra, 1994). Diplomystes nahuelbutaensis is an en- tially threatened group of siluriforms, we here demic and the only diplomystid species known describe and illustrate these specimens empha- from the Río Bío Bío basin (Arratia, 1987; Ruiz & sizing the juveniles. Further, we offer general Berra, 1994). To our knowledge, the specimens in comparisons of the juvenile diplomystids to oth- this collection measuring about 13-14 mm SL are er siluriforms based largely on a synthesis of the smallest Diplomystes known (Figs. 2-3). The scattered published illustrations that summarize smallest diplomystid specimens reported in the the diversity and commonality of juvenile cat- literature are twice this size at about 30 mm SL fishes. (Arratia, 1983; Azpelicueta & Gosztonyi, 1998). * Department of Ichthyology, The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103, USA. E-mail: [email protected] ** Department of Evolution, Ecology and Organismal Biology, The Ohio State University, Mansfield, Ohio 44906, USA. E-mail: [email protected] *** Cornell University Museum of Vertebrates, E151 Corson Hall, Ithaca, NY 14853, USA. E-mail: [email protected] Ichthyol. Explor. Freshwaters, Vol. 15, No. 1 72 Siluriformes Diplomystidae Siluroidei † Hypsidoridae Siluroidea (all other catfishes) In phylogenetic studies of siluriforms Diplomystes and †Hypsidoris have been used as a basal ‘dou- blet’ outgroup for inferring polarities of charac- ter state transformation (Bornbusch, 1991; Friel, 1994). De Pinna (1993, 1998) argued that cetopsid catfishes are also relatively basal among sil- uriforms, and he questioned resolution of the Fig. 1. Habitat of juvenile Diplomystes nahuelbutaensis diplomystid-hypsidorid basal ‘doublet’ below on Río Laja at La Cantera, 7 km below Salta Laja, Río Siluroidea. In any case, the basal position of Diplo- Bío Bío basin, Chile, December 1992. Photo by T. M. mystes is well corroborated, and we expect that Berra. additional features of these fishes will be identi- fied as plesiomorphic among siluriforms. Phylogenetic context. Eigenmann & Eigenmann Diplomystid distribution and natural history. (1890) and Regan (1911) called attention to the Six species of Diplomystidae are currently recog- primitive nature of Diplomystes, emphasizing its nized (Arratia, 1987; Azpelicueta, 1994a-b). Berra dentigerous maxillary bone. The Eigenmanns (2001) depicts the family distribution. Half of the placed the Diplomystidae at the base of their species occur west of the Andes in south central evolutionary tree diagram of Neotropical catfish- Chile. The trans-Andean species are D. chilensis es (1890, p. 7). The recognition of Diplomystes as from rivers near Valparaíso and Santiago, D. na- primitive among all catfishes has been supported huelbutaensis from the Bío Bío basin, and D. cam- and extended by many subsequent workers: Berg posensis from the Valdivia region (Arratia, 1987). (1940), Chardon (1968), Greenwood et al. (1966), Three other species occur east of the Andes in Lundberg & Baskin (1969), Lundberg & Case southern Argentina. Arratia (1987) erected the (1970), Fink & Fink (1981, 1996), Arratia (1987, genus Olivaichthys for the Argentinian specimens. 1992), Grande (1987), Mo (1991), Azpelicueta We follow Azpelicueta (1994a-b) and most subse- (1994a-b), de Pinna (1993, 1998). Diplomystids quent authors who treat Olivaichthys as a syno- retain more plesiomorphic characteristics than nym of Diplomystes. The cis-Andean species are any other siluriforms, Recent or fossil, including D. viedmensis from the Río Negro system, D. cuya- aspects of the maxillary bone, barbels, nares, otic nus from the Río Colorado and the Desaguadero- capsule, anterior pterygoid bones, Weberian com- Salado basin, and D. mesembrinus known only plex centra, caudal skeleton and fin rays, and from relatively few specimens from Chubut and pectoral girdle. Nevertheless, monophyly of Diplo- Senguerr rivers (Azpelicueta, 1994a-b). mystes is well supported by synapomorphies of Relatively little is known of the habits and life the vomerine and palatine shapes, cranial articu- history of diplomystid species. In Chile Diplo- lation of the hyomandibula, and heavily papil- mystes are mostly found to be benthic in fast lose skin (Arratia, 1987; Arratia & Huaquin, 1995). moving streams, and D. camposensis also occurs One of the oldest and best preserved fossil in lakes (Arratia, 1983, 1987). Diplomystes vied- siluriforms, †Hypsidoris, from the Eocene of North mensis has been taken from rivers near sea level America is the only catfish besides Diplomystes to about 1,900 m (Azpelicueta, 1994a). Diplomyst- that has a toothed maxilla (Lundberg & Case, ids are generalized carnivores that consume an- 1970; Grande, 1987). Otherwise †Hypsidoris shares nelids, mollusks, and arthropods (Arratia, 1987; derived characters that diagnose all other recent Azpelicueta & Gosztonyi, 1998). Specimens of catfishes (Grande, 1987; Grande & de Pinna, 1998). D. nahuelbutaensis from fast flowing moderate el- Grande (1987) proposed a classification to reflect evation (370-520 m) tributaries of the Río Bío Bío this phylogenetic sequence: (Ruiz & Berra, 1994) had eaten aquatic insect larvae, especially chironomids, and the relatively large decapod crustacean Aegla (Arratia, 1987; Lundberg et al.: Diplomystes juveniles 73 Ruiz & Berra, 1994). Reproduction occurs at least during the Austral summer based on captures of females with maturing eggs, and the juveniles reported here were collected in December. All diplomystids are considered to be potentially or actually threatened or endangered due to habitat deterioration and predation or competition by introduced trout, Oncorhynchus mykiss and Salmo trutta. Diplomystes chilensis may be extinct (Arra- tia, 1987). Material and methods Material examined: 7 specimens, 12.9-130.6 mm SL, all from Chile: ANSP 177914, 2, 30.7-37.9 mm SL; Río Laja at La Cantera, 7 km below Salta Laja; 9 Dec 1992. – ANSP 177913, 5, 12.9-130.6 mm SL; Río Laja 2 km S of Tucapel, 14 Nov 1992. An Fig. 2. Diplomystes nahuelbutaensis, three freshly cap- additional 27 specimens (not examined in this tured juveniles from series collected in Río Laja, Río Bío study but 3 shown in Fig. 2) collected at these Bío basin, Chile, December 1992. Photo by T. M. Berra. same localities are deposited at Ohio State Uni- versity or Universidad de Concepción, Instituto de Zoología, Chile. hypurals; 9, caudal-fin ventral origin at anterior- In the descriptions below, specimens of the most declination of caudal-fin membrane; 10, size series are ordered by standard length (SL) anal-fin posterior end behind insertion of last and, where necessary, identified by SL rounded lepidotrichium; 11, anal-fin origin at insertion of to nearest integer: 13, 14, 31, 38, 55, 71, 131 mm. first lepidotrichium; 12L, 12R, left and right pel- The 13 and 14-mm specimens are staged as ‘juve- vic-fin origins at insertions of their first (outer) niles’ (Kendall et al., 1984; Fuiman, 1984). Speci- lepidotrichium. mens larger than 31 mm SL in our series lack Bivariate allometric growth relationships are juvenile features and look like small adults, but visualized in scatterplots of ratios between meas- the series does not include large sexually mature urement pairs vs. SL or head length. Multivariate individuals. allometric growth patterns among a subset of 43 Fifty seven measurements (Table 1) were made non-overlapping measurements were inferred of the seven specimens in the 13-131 mm size from a Principal Components Analysis of the series including straight-line dimensions com- covariance matrix of log transformed measure- monly used in catfish taxonomy and dimensions ment data (Table 2). The scaled factor loadings between 14 homologue landmark points arranged (coefficients; Sneath & Sokal, 1973) of the 43 di- in a truss set (Strauss & Bookstein, 1982). The mensions on the first Principal Component
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