Cbpeia, 1997 (1), pp. 227-232 Peruana, Iquitos, Peru and MUSM for Museu de Historia Natural de la Universidad Nacional Generic Reassignment of the Loricariid Mayor de San Marcos Lima, Peru. Monistiancistrus carachama Fowler 1940, Plecostomus lacerta Nichols 1919, and Pseudorinelepis carachama (Fowler) levis Pearson 1924 Monistiancistrus carachama Fowler, 1940. Proc. (Teleostei: Siluriformes) Acad. Nat. Sci. Phila. 91:236-238. figs. 26-27.

JONATHAN W. ARMBRUSTER AND Pseudorinelepis carachama. Nijssen and Isbriicker, LAWRENCE M. PAGE 1986. Rev. fro Aquariol. 13:93. carachama. Isbriicker, 1992. Harnis­ The contains approxi­ chwelse. September 1992:71. mately 600 species of from South and Central America (lsbriicker, 1980). The family Monistiancistrus was described as a mono typic is diagnosed by several characteristics including from the Rio Ucayali of Peru by Fowler the presence of an armor of bony plates in four (1940). Recently, the validity of Monistiancistrus or more rows, contact of the metapterygoid and the phylogenetic position of the genus have with the lateral ethmoid, an expanded meseth­ been debated. The genus has been placed in moid disk, and an expanded anterior margin of Ancistrinae (Isbriicker, 1980) and Hypostomi­ the anterior ceratohyal (Schaefer, 1987). afthe nae (Isbriicker and Nijssen, 1982; Nijssen and six subfamilies possibly ascribed to Loricariidae, Isbriicker 1986; Schaefer, 1986, 1987). Isbriicker all except are well diagnosed and Nijssen (1982) placed Monistiancistrus in (Schaefer, 1986, 1987). Schaefer hypothesized what they termed the .. Hypostomus group" that Ancistrinae evolved from within Hypostom­ which also included Hypostomus, , inae making the latter subfamily paraphyletic; Cochliodon, , and Pterygoplichthys, however, he continued to recognize both sub­ and Schaefer (1986) stated that the dif­ families until a thorough study of Hypostomi­ fered from Hypostomus only by the loss of an ad­ nae could be completed. Despite the apparent ipose fin. Nijssen and Isbriicker (1986) placed paraphyly of Hypostominae and the fact that Monistiancistrus into synonymy with the hypos­ few of the genera of the subfamily are diag­ tomine genus Pseudorinelepis Bleeker 1862 with nosed, it is still possible to identify most genera little comment. Finally, Isbriicker (1992) re­ by referring to Gosline (1947) and examining moved Monistiancistrus from Pseudorinelepis and type-specimens of type-species or using pub­ placed it into the synonymy of Hypostomus with lished diagnoses. no comment. While examining type-specimens of loricariid Formerly, Monistiancistrus carachama was catfishes, we have discovered three taxa that known only from the holotype (ANSP 68654) were placed by the original authors or recent which is a juvenile, 72.2 mm standard length. workers into the wrong genera. In this paper, However, Isbriicker and Nijssen (1982) noted we present evidence to place Monistiancistrus the existence of a second specimen from the Fowler 1940 into the synonymy of Pseudorinelepis upper Amazon of Peru, and we recently ac­ Bleeker 1862, Plecostomus lacerta Nichols 1919 quired new material assignable to Monistiancis­ into Kronichthys Miranda Ribeiro 1908, and Rhi­ trus from the Amazon near Iquitos, Peru. Upon nelepis levis Pearson 1924 into Cochliodon Heckel examination of new material, the holotype of 1854. M. carachama, and comparative specimens of Pseudorinelepis, we have determined that Monis­ Materials and methods.-Type-specimens were ex­ tiancistrus should be returned to the synonymy amined for various morphological characteris­ of Pseudorinelepis. tics associated with the genera of Hypostomi­ Species of Pseudorinelepis are large-river nae. Placement of species into genera was ac­ found in backwaters and floodplain lakes of the complished by comparison to the type-speci­ Rio Amazonas and its major tributaries (Fig. 1). mens of the type-species of the genus when Most specimens are dark brown, occasionally possible and published descriptions or topotyp­ with slightly darker spots on the dorsal-fin mem­ ic material when direct examination of the type brane (Fig. 2A). In breeding males, the body is specimens of the type species of the genus was tan with dark spots located at the junction of not possible. Institutional abbreviations are as in the lateral plates and on all fins, and the edges Leviton et al. (1985) with the addition of IIAP of the cheek, dorsal, caudal, and pectoral fins for Instituto de Investigaciones de la Amazonia are orange (see photograph in Burgess, 1989: 228 COPEIA, 1997, NO.1

The type of M. carachama is a heavily armored fish that putatively differs from Pseudorinelepis by lacking plates on the ventral surface of the head and by the absence of elongate odontodes on the cheek. However, both of these characteris­ tics are associated with the size and age of the type of M. carachama which is a juvenile. Plates on the ventral surface of the head and abdo­ men develop rather late ontogenetically in hy­ postomine species (Regan, 1904) and are only just beginning to form on the type of M. cara­ chama. A series of small (85.7-113.1 mm) Pseu­ dorinelepis (INHS 36938) from the Rio Amazo­ nas near Iquitos, Peru, demonstrates that plates on the throat first develop in three patches, one on each side and one in the middle just above the cleithrum. As the fish grows, plates are add­ Fig. 1. Type localities of Cochliodon levis (square) ed mesially and anteriorly from these three and Kronichthys lacerta (diamond) and range of Pseu­ patches (Fig. 3) such that, in adults, the entire durinelepis (circles). Open circle is the type locality of throat region is covered with small plates except P. carachama. Some symbols represent more than one for a small bare patch under the lower lip. The locality. type of M. carachama does possess small patches of plates on the throat, but the patches are not as well developed as they are on the smaller 748). In one specimen (IIAP 114), the anterior specimen in Figure 3. part of the body is uniformly dark, and the pos­ The only characteristic other than plates on terior half is spotted whereas all other speci­ the ventral surface of the head used to distin­ mens in the same lot are uniformly dark brown. guish Monistiancistrus from Pseudorinelepis is the One specimen (INHS 36938) was olive-green latter's possession of elongate odontodes on the when alive. cheeks (Burgess, 1989); however, this also is a Pseudorinelepis is distinguishable from most characteristic that is added late ontogenetically, Hypostominae by the lack of an adipose fin. and the holotype of Monistiancistrus is too small The only other Hypostominae that lack an adi­ to have developed odontodes. In the larger pose fin are Rhinelepis, Pogonopomoides, Pareior­ specimens of INHS 36938, development of the hina, Corymbophanes andersoni, and Cochliodon lev­ cheek odontodes has just begun. Both males is. Pseudorinelepis can be distinguished from and females have elongate cheek odontodes, these other hypostomines that lack adipose fins but the patch of odontodes in males is larger by a combination of the following characteris­ and the odontodes are longer. tics: adipose fin completely absent and not re­ Because the characteristics used to diagnose placed by a low ridge as in C. andersoni, well­ Monistiancistrus are found only in juveniles and, keeled plates (vs flat in all others), odontodes in fact, are identical to those of juvenile Pseu­ forming tall ridges on the pterotic-supracleith­ dorinelepis, we conclude that Monistiancistrus be­ rum (vs nearly flat in C. levis, Rhinelepis, and Po­ longs in the synonymy of Pseudorinelepis. With gonopomoides), head short but deep with depth the addition of P. carachama, Pseudorinelepis con­ to length ratio of 0.66-0.76 (x = 0.71, n = 11) tains four nominal species, P. agassizi, P. caracha­ whereas the head is long and depressed in Rhi­ ma, P. genibarbis, and P. pellegrini, all of which nelepis with a depth to length ratio of 0.47-0.59 come from the Rio Amazonas. (x = 0.52, n = 6), abdomen of adults complete­ ly encased in plates (abdomen either without Kronichthys lacerta (Nichols) plates or not completely encased in C. levis, Par­ eiorhina, and Pogonopomoides), posterior margin Plecostomus lacerta Nichols, 1919. Revista Mus. of the pterotic-supracleithrum not bordered by Paul. 11:424-425. a patch of numerous small plates as in Pogono­ pomoides and Rhinelepis, teeth thin and bicuspid Hypostomus lacerta. Isbriicker, 1980. Versl. Techn. (vs spoon-shaped and usually unicuspid in C. Gegevens, Univ. van Amsterdam. No. 22:25. levis), and an anal fin with Qne unbranched and five branched rays (vs four branched rays in C. Plecostomus lacerta Nichols (1919) was de­ levis) . scribed from the rio Juquia at Po~o Grande, Sao SHORTER CONTRIBUTIONS 229 A

B

c

Fig. 2. (A) PseudorinelefJis pellegrini (lNHS 36938, 95.0 mm), dorsal view; (8 ) Kronichthys la certa (CAS 5671 8, 39.1 mm), dorsal view, and (C) Cochliodon levis (CAS 77350, paralype, 150.6 mm), lateral view.

Paulo, Brazil (Fig. I). The o nly type numbered transferred to Hypostomus when Plecostomus was by Nichols ( 1919) in h is text is the holotype synonymized with H),postomus by Boeseman (AMNH 7 15 1); however, he examined three ( 1968). more specime ns, and MZUSP 964 (n = 1) is Bo th specime ns are cylindrical in bod y shape labeled as a pal'atype fro m Esta~ao de Rai z da and possess a naked abdomen; mo rc than three Serra, Sao Paulo, Brazil. Plecoslomus lacerta was plates (four and six) along the do rsal midline 230 COPEIA , 1997, NO. I

B

Fig. 3. Fo rmation of plates on the throat of Pseudorillele/Jis (INHS 36938) . (A) 85.7 mm SL. (B) 99.4 mm 51.. White line points to scuteless patch.

between the supraoccipital tip and the dorsal­ amined have a single, medium to large plate fin spine; a bare patch just postnior to the pte­ posterior to the pterolic-suprac1eilhrum and a rOlic-supracleithrum; five branched anal-fin V-shaped spinclct. Given the characters, it is rays; a spine1el in the first that has necessary to remove the species from flyposto­ become reduced to a small, rectangular, plate­ ?nus. Kmnichthys is the only genus that agrees like struclllI"e; shon pelvic-fll1 spines that are with all of the characters of P. lacerta, and wide and c1ublike; and the plate that bears the throug h the usc of the key of Gosline (1947), it anterior segment of the preopcrcular canal is is apparent that P. lacerta can only be ascribed large and is refl ected mesially. Both specimens to Kronichthys. lack elongate odontodes o n the cheek. Species The color of both specimens of K. lacerta are o f I-Iypostomus usually have [our branched anal­ considt:rably faded. Nicho ls (1919) gave litLl e in fin rays (Schaefer, 1986), and all Hypostomus ex- the way of color characteristics other than La say SHORTER CONTRIBUTIONS 231 that the backs of the specimens were dusky. The From examination of the types, it appears paratype appears to have had several saddles that C. levis is light brown with a lighter ventral along its back. Specimens of Kronichthys from surface. Small spots are present on the anterior near the type locality (CAS 56718) have 4-5 part of the body and the head but are absent dark saddles (Fig. 2B). The sides of the body from the abdomen. The dorsal, anal, and below the saddles are uniformly dark brown, paired fins all have large, diffuse spots. The cau­ and the abdomen is white. The fins have large dal fin is dusky and no spots are visible; how­ spots with the spots on the caudal fin often co­ ever, Pearson (1924) reported that the speci­ alescing to form a dark wash. Upper lobe of the mens had spots on the caudal fin and that there caudal fin lighter than the lower lobe. Head were fewer than on the other fins. The abdo­ mottled with spots along the margin and the men is covered with small plates except for upper lip. broad circular regions around the bases of the Three species of Kronichthys have been de­ pelvic fins, small bare areas at the bases of the scribed, K. heylandii Boulenger 1900, K. subteres pectoral fins, and a small bare patch at the pos­ Ribeiro 1908, and K. lacerta Nichols 1919, all terior margin of the lower lip. As in other spe­ from the state of Sao Paulo. It is unknown how cies of Cochliodon, the dentaries are angled to K. lacerta compares with the two other nominal form a V-shape, and the body is deepest at the species or how any of the species can be distin­ origin of the dorsal fin and tapers dramatically guished from one another because of insuffi­ to the caudal peduncle making the body appear cient original descriptions. Both K. subteres and humped. There are 12-18 teeth per dentary K. lacerta are described from the rio Ribeira do and 10-17 teeth per premaxilla. Iguape system.

Specimens examined.-Cochliodon levis: CAS 77349 Cochliodon levis (Pearson) (IV 17014) (Holotype), CAS 77350 (IV 17014) Rhinelepis levis Pearson, 1924. Revista Mus. Paul. (2 Paratypes), and UMMZ 66496 (Paratype), 11 :424-425. Bolivia, Depto. La Paz, Rio Beni Drainage, San Miguel Huachi at junction of Rio Boopi and Rio Hypostomus levis Isbriicker, 1980. Versl. Techn. Cochabamba. Kronichthys lacerta: AMNH 7151 Gegevens, Univ. van Amsterdam. No. 22:25- (Holotype), .Brazil, Sao Paulo, rio Juquii at 26. Pot;:o Grande; Brazil, Sao Paulo, MZUSP 964 (Paratype), Estat;:ao de Raiz da Serra; CAS 56718 Rhinelepis levis was described by Pearson (3), Brazil, Sao Paulo, rio Ribeira basin, trib. of (1924) from the Rio Beni drainage near San rio Ribeira do Iguape at bridge on Sao Paulo­ Miguel de Huachi at the junction of the Rio Curitiba highway. Pseudorinelepis agassizii: MCZ Boopi and Rio Cochabamba in Bolivia (Fig. 1). 8007 (Syntype?), Brazil, Amazonas, lago Mana­ The species was originally placed by Pearson in capuru, Manacapuru (lago Grande de Manaca­ Rhinelepis because of the lack of an adipose fin puru), 3°6'S, 61°30'W. Pseudorinelepis carachama: (Fig. 2C) and later placed in Hypostomus by Is­ ANSP 68654 (Holotype), Peru, Depto. Loreto, briicker (1980). However, the teeth of the four Rio Ucayali Basin, at Contamana. Pseudorinelepis type specimens (holotype: CAS 77349, para­ genibarbis: FMNH 95569 (1) and MZUSP 6339 types: CAS 77350, UMMZ 66496) are large and (17), Brazil, Amazonas, lago Castro do rio Pu­ spoon-shaped. Among loricariids, spoon-shaped rus. Pseudorinelepis pellegrini: CAS 42325 (1), teeth are found only in Cochliodon and . Peru, Depto. Loreto, Quebrada Yaguas Yacu Because the specimens lack the evertable plates near Pebas; CAS 58801 (1), Peru, Depto. Lore­ and elongate odontodes on the cheek charac­ teristic of Panaque, the specimens are assignable to, Iquitos; HAP 114 (6), Peru, Depto. Loreto, to Cochliodon. No other described species of Rio Samiria (Cauo Ungurahui); INHS 36938 Cochliodon lacks an adipose fin. Although indi­ (6), Peru, Depto. Loreto, Rio Amazonas, at viduals of loricariid species that normally have Pueblo Gallito; INHS 36941 (1), Peru, Depto. adipose fins, such as Hypostomus and Aphanoto­ Loreto, Felipe Cocha (Rio Itaya), 12 km S Iqui­ rulus, are occasionally found without them tos on road to Quistococha near the community (}WA, pers. obs.), all of the specimens in the of 29 Enero 1995; MUSM 1847 (1), Peru, Dep­ type series of C. levis lack an adipose fin, and it to. Ucayali, Ivita, Pucallpa; MUSM 1869 (1), is assumed that the loss of the adipose fin is not Peru, Depto. Ucayali, Santa Carmela de Ma­ a developmental abnormality but a characteris­ changay (laguna), Pucallpa; MUSM 6064 (1), tic that diagnoses C. levis from other species of Peru, Depto. Ucayali, Yarinacocha, Coronel Po­ Cochliodon. tillo. 232 COPEIA, 1997, NO.1

Acknowledgments.-We would like to thank M. E. Vers!' Techn. Gegevens, Univ. van Amsterdam. 22: Retzer, M. H. Sabaj, S. A. Schaefer, and an un­ 1-181. known reviewer for comments and suggestions --. 1992. Uberblick tiber die gtiltigen (Unter-) on improving the manuscript; K. S. Cummings Gattungsnamen der Harnischwelse (Loricariidae) und ihre Synonyme. Harnischwelse, DATZ-Sonder­ for the photographs; and C. W. Ronto for heft, September 1992:71-72. mounting the photographs. Also, thanks to B. --, AND H. NIjSSEN. 1982. Aphanotorulus jrankei, M. Burr, A. Choto, and C. Gonazales for help une espece et un genre nouveaux de poissons-chats in obtaining specimens of Pseudorinelepis in cuirasses du bassin du Ro Ucayali au Perou (Pisces, Peru. Special thanks to the following people for Siluriformes, Loricariidae). Rev. fro Aquario!. 9: 105- invaluable help while visiting institutions and 1l0. for the loans of specimens: R. Arrindell, N. LEVITON, A. E., R. H. GIBBS JR., E. HEAL, AND C. E. Feinberg, G. Nelson, M. Stiassny, W. Saul, S. DAWSON. 1985. Standards in herpetology and ich­ Schaefer, D. Catania, W. Eschmeyer, T. Iwamoto, thyology: Part I. Standard symbolic codes for insti­ tutional resource collections in herpetology and E. Bermingham, M. Rogers, K. Swagel, B. Cher­ ichthyology. Copeia 1985:802-832. noff, N. Flores, E. Isern, H. Sanchez, K. Hartel, NICHOLS, J. T. 1919. Cascudos brasileiros do genero P. Buckup, H. Britski, O. Oyakawa, G. Burgess, Plecostomus do Museu Paulista. Revista Mus. Pau!' H. Ortega, W. Fink, D. Nelson, G. Smith, R. 1l:409-417. (English version follows in 1l:419- Vari, and J. Williams. This research was partially 426). funded by the following grants and awards to NIjSSEN, H., AND IJ.H. ISBRUCKER. 1986. spectracanthi­ JWA: Francis M. and Harlie M. Clark Research cus murinus, nouveaux genre et espece de Poisson­ Support Grants (University of Illinois), Ernst Chat cuirasse du Rio Tapaj6s, Est. Para, Bresil, avec Mayr Grant (Harvard University), Edward C. des remarques sur d'autres genres de Loricariids Raney Memorial Fund Award (American Society (Pisces, Siluriformes, Loricariidae). Rev. fro Aquar­ io!. 13:93-98. of Ichthyologists and Herpetologists), Philip W. PEARSON, N. E. 1924. The of the eastern slope Smith Memorial Fund Award (Illinois Natural of the Andes. I. The fishes of the Rio Beni basin, History Survey), and the University of Illinois, Bolivia, collected by the Mulford Expedition. Indi­ Department of Ecology, Ethology, and Evolu­ ana Univ. Stud. 1l:1-83. tion Graduate Student Research Award. REGAN, C. T. 1904. A monograph of the fishes of the family Loricariidae. Trans. Zoo!. Soc. London 17: 191-350. LITERATURE CITED SCHAEFER, S. A. 1986. Historical biology of the lori­ cariid catfishes: phylogenetics and functional mor­ BOESEMAN, M. 1968. The genus Hypostomus Lace­ phology. Unpub!' Ph.D. diss. Univ. of Chicago, Chi­ pede, 1803, and its Surinam representatives (Silur­ cago. iformes, Loricariidae). Zoologische Verhandelin­ ---. 1987. Osteology of gen 99:1-89. (Linnaeus) with a phylogenetic analysis of the lor­ BURGESS, W. E. 1989. An atlas of freshwater and ma­ icariid subfamilies (Pisces: Siluroidei). Contrib. Sci., rine catfishes, a preliminary survey of the Silurifor­ Nat. Hist. Mus. Los Aangeles Co. 394:1-31. meso T.F.H. Pub!., Neptune City, N.J. FOWLER, H. W. 1940. A collection of fishes obtained Illinois Natural History Survey, 607 East Peabody by Mr. William C. Morrow in the Ucayali River ba­ sin, Peru. Proc. Acad. Nat. Sci. Phila. 91:219-289. Drive, Champaign, Illinois 61820. Send reprint GoSLINE, W. A. 1947. Contributions to the classifica­ requests to JWA. Submitted: 27 Feb. 1996. Ac­ tion of loricariid catfishes. Arq. do Mus. Nac., Rio cepted: 24June 1996. Section editor: R. Win­ de Janeiro 41:79-134. terbottom. ISBRUCKER, I. J. H. 1980. Classification and catalogue of the mailed Loricariidae (Pisces,Siluriformes ) . © 1997 by the American Society of Ichthyologists and Herpetologists