25 September 1990 PROC. BIOL. SOC. WASH. 103(3), 1990, pp. 550-557 PANTOSTICTOS, A NEW (PISCES: OSTARIOPHYSI: : ) FROM THE WESTERN PORTIONS OF THE AMAZON BASIN

Richard P. Vari and Ramiro Barriga S.

Abstract. — Cyphocharax pantostictos, a species of curimatid characiform with a distinctive pattern of dark spots arranged in longitudinal series along the sides of the body, is described as new from the Rio Napo, Rio Putumayo, Rio Ucayali, and Rio Nanay in Ecuador and northern Peru. Cyphocharax multi- lineatus (Myers), the only other species in that questionably monophyletic with a similar pigmentation pattern has dark wavy horizontal lines, rather than discrete spots arranged in horizontal patterns. The dark body pigmentation in the two species also differs in its relative position on the scales. The pigmen- tation pattern and overall external appearance of C. pantostictos are nearly identical to that of fasciata (Vari & Gery) a species endemic to the upper Rio Madeira system in Brazil. The two species can be readily distinguished on the basis of a series of meristic and morphometric features, and in differences in the portion of buccopharyngeal complex on the roof of the oral cavity. A series of polarized characters indicate furthermore that the two species are not closely related.

The myriad drainage systems, range of azonian endemic area" based on distribu- stream gradients, and complexity of aquatic tional data from various neotropical genera habitats found in the drainage basins of the of the perciform family Cichlidae. In the western portions of the Amazon basin are course of investigations of the fauna of reflected in the remarkable diversity of the eastern Ecuador and northern Peru we in- fish fauna in that region (Ortega & Vari 1986, dependently collected a distinctive species Stewart et al. 1987). This region is also one of curimatid with an unusual pigmentation of the areas of greatest diversity for the fam- pattern consisting of seven or eight longi- ily Curimatidae, involving both species tudinal series of dark spots along the sides widely distributed within the Amazon basin of the body. This material first appeared to (e.g., aspera Gunther (Vari 1988: represent a major extension in the known fig.8) and C. vittata Kner (Vari 1989b:42)), distribution of Curimata fasciata which Vari or ranging north and south of that system & Gery (1985) described from the Rio Ma- into the Rio Orinoco or Rio de La Plata deira system in Brazil, a considerable dis- systems (e.g., Steindachnerina guentheri tance southeast of the region from which (Eigenmann & Eigenmann) (Vari 1990)). the Ecuadorian and Peruvian specimens Other curimatids in this area of high species originated. More recently Vari (1989a:ta- diversity have much more restricted ranges bles 2, 3), in an analysis of intrafamilial phy- (e.g., Steindachnerina quasimodoi Vari & logenetic relationships, restricted Curimata Vari (see Vari & Vari 1989:477)) and are to a single lineage within the family and known only to occur in the region that Kul- reassignedfasciata to Steindachnerina Fow- lander (1986:40) termed the "Western Am- ler (1906) on the basis of a series of derived

VOLUME 103, NUMBER 3 551 characters. Further examination of our 58-59) noted was not defined on the basis specimens surprisingly showed that they of known derived features. Cyphocharax of neither constitute a major range extension that classification was rather an assemblage for Steindachnerina fasciata, nor do they of species lacking the derived features di- even represent a species of Steindachnerina. agnostic of the three other genera (Stein- Rather they are a species of Cyphocharax dachnerina, Eigenmann & Ei- Fowler (1906) previously unknown to sci- genmann, and Fernandez- ence. This new curimatid is described herein Yepez) which together with Cyphocharax and is yet another fish species with a known form an unresolved terminal polytomy in range limited to the Western Amazonian Vari's hypothesis of intrafamilial relation- endemic area identified by Kullander. ships within the Curimatidae. The absence Materials and methods. —Counts and of identified synapomorphies for Cypho- measurements were made following meth- charax increases the likelihood that the ge- ods outlined in Van (1989b, 1989c, 1990). nus may not be monophyletic. Ongoing Ranges for meristic and morphometric fea- studies by the senior author focus on the tures include values of all examined speci- question of the monophyly of Cyphocharax mens. The values in square brackets are and its subunits. In the interim we assign those of the holotype. Subunits of the head the new species to Cyphocharax given that are presented as proportions of head length C. pantostictos shares the synapomorphies (HL). Head length itself and measurements for the clade formed by Cyphocharax, of body parts are presented as proportions Steindachnerina, Curimatella, and Pseu- of standard length (SL). docurimata, but lacks the derived features The following abbreviations for institu- that diagnose each of Steindachnerina, Cu- tions are used: Academy of Natural Sciences rimatella, and Pseudocurimata. The strik- of Philadelphia (ANSP); British Museum ing pattern of seven or eight horizontal se- (Natural History), London (BMNH); Cali- ries of prominent dark spots aligned along fornia Academy of Sciences, San Francisco the center of the body scales is unique to (CAS); Stanford University, collections now the Cyphocharax pantostictos within the ge- deposited at CAS (CAS-SU); Instituto Na- nus (Fig. 1). Only one other Cyphocharax cional de Pesquisas da Amazonia, Manaus species, C. multilineatus (Myers 1927) of (INPA); Indiana University, collections now the Rio Negro system in Venezuela and Bra- deposited at various repositories (IU); Mu- zil, has a pattern of horizontal dark body seo de Biologia, Universidad Central de pigmentation reminiscent of that in C. pan- Venezuela, Caracas (MBUCV); Museo de tostictos. The pattern of dark body pigmen- Biologia de la Escuela Politecnica Nacional, tation in C. multilineatus (Fig. 2) differs from Quito (MEPN); Museum d'Histoire Natu- that in C. pantostictos in forming solid wavy relle, Geneva (MHNG); Museu Nacional, horizontal lines rather than a series of dis- Rio de Janeiro (MNRJ); Museu de Zoolo- crete rotund spots (compare Figs. 1 and 2). gia, Universidade de Sao Paulo, Sao Paulo Furthermore, the dark stripes in C. multi- (MZUSP); Naturhistoriska Riksmuseet, lineatus are positioned along the area of Stockholm (NRM); and National Museum overlap of horizontal rows of scales along of Natural History, Smithsonian Institu- the body, rather than being aligned along tion, Washington, D.C. (USNM). the center of the scale rows as are the spots in C. pantostictos. Thus the patterns of lon- Cyphocharax pantostictos, new species gitudinal dark pigmentation on the bodies Figs. 1, 4 in the two species are apparently non-ho- Diagnosis. —The new species is assigned mologous. Cyphocharax multilineatus also to Cyphocharax, a genus that Van (1989a: has a discrete dark band across the mid- 552 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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Fig. 1. Cyphocharax pantostictos, new species, holotype, USNM 306594, 72.4 mm SL; Ecuador, Napo, Laguna de Jatuncocha. lateral surface of the head anterior and pos- rina fasciata (Fig. 3), an endemic of the Rio terior to the orbit, a pigmentation pattern Madeira basin, and have lead to misiden- lacking in C. pantostictos. Cyphocharax tifications of the two species. Cyphocharax pantostictos, in turn, is characterized by a pantostictos lacks the derived features that well-developed, mid-lateral, horizontally diagnose Steindachnerina (see Vari 1989a: elongate patch of dark pigmentation on the 58, 1990), and lacks the intrageneric syn- caudal peduncle that is absent in C. multi- apomorphies for the clades that include S. lineatus. It is likely that various meristic and fasciata (see Vari 1990 for details). One of morphometric features further distinguish the most obvious differences between the C. pantostictos within Cyphocharax. Iden- two species involves the form of the buc- tification of those characters must await the copharyngeal complex on the roof of the completion of revisionary studies within that oral cavity. Cyphocharax pantostictos has speciose genus. three simple longitudinal fleshy folds in that The overall pigmentation pattern and region. Steindachnerina fasciata, in con- overall external appearance of C. pantostic- trast, has a mass of lobulate fleshy bodies tos are strikingly similar to that of one other that extend ventrally into the oral cavity, a member of the Curimatidae, Steindachne- hypothesized derived condition unique to a

Fig. 2. Cyphocharax multilineatus, USNM 269987, 111.8 mm SL; Venezuela, Territorio Federal Amazonas, Departamento Rio Negro, Caiio Tremblador where crossed by road from San Carlos de Rio Negro to Solano. VOLUME 103, NUMBER 3 553

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Fig. 3. Steindachnerina fasciata, MNRJ 11208, 89.6 mm SL, holotype of Curimata fasciata Vani and Gery; Brazil, Territorio de Rond6nia, Municipio de Ouro Preto do Oeste, Rio Roman (or Sao Domingo) near Nova Unido. subunit of Steindachnerina (see Van 1989a: larged relative to those on lateral surfaces 31-35, 1990, for a discussion of the buc- of body. Median pre-pelvic scale series copharyngeal complex). somewhat irregular, particularly near origin The two species also differ in various me- of pelvic fin. No distinct median keel pos- ristic and morphometric values including terior to origin of pelvic fin. Barely discern- the number of vertebrae (31 in Cyphocha- able secondary obtuse keel on each side of rax pantostictos versus 32 to 34, typically post-pelvic portion of body about two scales 33, in Steindachnerina fasciata), the num- dorsal of ventral midline ber of scales in a longitudinal series to the Greatest body depth at origin of dorsal hypural joint (29 to 31 versus 32 to 37), the fin, depth 0.35-0.40 [0.37], relatively deep- number of scale rows above the lateral line er in larger specimens; snout tip to origin to the origin of dorsal fin (41/2 versus 51/2 or of dorsal fin 0.47-0.52 [0.50]; snout tip to 61/2), the relative length of the pelvic fin origin of anal fin 0.83-0.85 [0.83]; snout tip (0.20-0.23 of SL versus 0.23-0.25), and the to origin of pelvic fin 0.52-0.57 [0.56]; snout relative gape width (0.24-0.28 of HL versus tip to anus 0.78-0.79 [0.78]; origin of dorsal 0.28-0.32). fin to hypural joint 0.54-0.58 [0.56]. Margin Description. —Body moderately elongate, of dorsal fin rounded posteriorly; anterior- somewhat compressed. Dorsal profile of most rays approximately two to two and head straight overall, slightly convex ante- one-half times length of ultimate ray. Mar- riorly. Dorsal profile of body smoothly con- gin of pectoral fin pointed; length of pectoral vex from posterior portion of head to origin fin 0.18-0.21 [0.20], extending slightly over of dorsal fin; straight or slightly convex, pos- one-half distance to vertical line through teroventrally slanted at base of dorsal fin, origin of pelvic fin. Margin of pelvic fin greatly convex from base of last dorsal-fin pointed, length of pelvic fin 0.20-0.23 [0.22], ray to caudal peduncle. Dorsal surface of tip reaches to anus in holotype, falls some- body with indistinct median keel anterior what short of that point in larger specimens. to dorsal fin, smoothly rounded transversely Caudal fin forked. Adipose fin well devel- posterior to fin. Ventral profile of body gently oped. Anal fin emarginate, anteriormost curved from tip of lower jaw to caudal pe- branched rays about two and one-half times duncle. Pre-pelvic region very obtusely flat- length of ultimate ray. Caudal peduncle tened, scales of that area not notably en- depth 0.12-0.14 [0.14]. 554 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Head profile distinctly pointed anteriorly, rows ventral of lateral line; very poorly de- head length 0.29-0.33 [0.31]; upper jaw veloped in series starting immediately dor- somewhat longer than lower, mouth sub- sal to origin of pectoral fin. Intense dark terminal; snout length 0.27-0.31 [0.30]; spots also progressively less pronounced in nares of each side very close, anterior ro- horizontal series dorsal to lateral line. tund, posterior crescent-shaped with aper- Patches of dark pigmentation located on ture partially closed by thin flap of skin sep- center of scale, with midpoint of spots lying arating nares; orbital diameter 0.27-0.32 medial of margin of preceding scale. Scales [0.31]; adipose eyelid present, moderately dorsal to lateral line with secondary area of developed, with rotund opening over center diffuse dark pigmentation posterior to dis- of eye; length of postorbital portion of head crete central dark spot; secondary dark pig- 0.42-0.46 [0.44]; gape width 0.24-0.28 mentation increasingly pronounced on dor- [0.27]; interorbital width 0.39-0.43 [0.43]. sal portions of body. Dark pigmentation Pored lateral-line scales from supraclei- patches on scales along lateral line merging thrum to hypural joint 29 to 31 [29]; all posteriorly into distinct horizontal stripe on scales of lateral line pored, canals in scales mid-lateral surface of caudal peduncle; stripe straight; 2 or 3 pored scales extend beyond continuing onto base of middle caudal-fin hypural joint onto caudal-fin base; 41/2 [41/2] rays. Deeper lying, dusky band extends along scales in transverse series from origin of mid-lateral surface of body from supra- dorsal fin to lateral line; 31/2 to 41/2 [4] scales cleithrum to caudal peduncle. in transverse series from lateral line to or- Caudal fin with small streak of dark pig- igin of anal fin. mentation on basal portions of middle rays; Dorsal-fin rays ii,9 [ii,9]; anal-fin rays ii,7 basal two-thirds of fin somewhat more dus- or iii,7 [ii,7]; pectoral-fin rays 13 to 15 [15]; ky than remainder of fin. Median and paired pelvic-fin rays i,8 or i,7,i [i,7,i]. fins somewhat dusky. Total vertebrae 31 in 8 specimens. Distribution.—Rio Napo, Rio Putumayo, Color in life. —(Based on photograph of Rio Ucayali, and Rio Nanay systems in Ec- paratype (USNM 280573) from the Rio Na- uador and northern Peru (Fig. 4). nay of Peru taken shortly after capture.) Ecology.—Two of the specimens from Overall coloration silvery with slightly ol- Peru (USNM 280573, NRM SOK/ ive-grey cast on dorsal portions of head and 1986293.5292) were collected in acidic black body. Series of black spots arranged in hor- waters among grass and submerged vege- izontal series along dorsal and lateral sur- tation. The specimens from the holotype faces of body. Distinct black mid-lateral locality were collected in submerged vege- stripe on caudal peduncle. Fins hyaline. tation in blackwater of pH 5.5 at a depth of Color in alcohol.—See Fig. 1 for pre- 1.5 m. The Rio Yasuni specimens came from served color pattern. Available specimens a slow flowing turbid stream with a pH of largely lacking guanine on scales. Overall 6.0 lacking submerged vegetation. ground coloration yellowish-tan, darker on Etymology. — Pantostictos, from the dorsal portions of head and body. Scales on Greek for "spotted all over" refers to the lateral and dorsal surfaces of body with dark prominent dark spots on the lateral and dor- patch of pigmentation on each scale; size of sal surfaces of the body. spots largest mid-laterally; overall intensity Remarks. —As noted above, Cyphocha- of spots not as pronounced in smaller in- rax pantostictos is very similar in body form dividuals. Spots forming 7 or 8 horizontal and pigmentation to Steindachnerina fas- series, dorsal most series not apparent in ciata which is apparently endemic to upper smaller specimens. Series of dark spots on portions of the eastern drainages of the Rio scales less developed posteriorly on scale Madeira basin in Brazil (see Van 1990:fig.

VOLUME 103, NUMBER 3 555

Fig. 4. Map of Amazon basin and adjoining areas showing distribution of Cyphocharax pantostictus (dots; localities: 1 = Rio Napo above Coca; 2 = Rio Yasuni; 3 = Laguna Jatuncocha, type locality; 4 = Rio Putumayo, El Estrecho; 5 = Rio Nanay, Nanay beach), Steindachnerina fasciata (squares), and Cyphocharax multilineatus (stars) (see under "Material examined" for additional locality information).

40). Nonetheless those two species differ in river kilometers. Thus mimicry would not many meristic and morphometric features, appear to be involved in the remarkable and Cyphocharax pantostictos furthermore external resemblance between Cyphocharax lacks the derived characters that both define pantostictos and Steindachnerina fasciata. Steindachnerina and clades within the ge- Material examined. —10 specimens, 34.4- nus that include S. fasciata (see Van (1990) 98.2 mm SL. for details). Holotype. —Ecuador: Napo. Laguna de Such pronounced superficial similarities Jatuncocha (01°00'S, 75°29'W), collected by between two distantly related species would R. Barriga, 29 Sep 1988, USNM 306594, at first consideration apparently represent a 72.4 mm SL. case of intergeneric mimicry. Interestingly, Paratypes. —Ecuador: Napo. Laguna de however, there is no overlap in the distri- Jatuncocha (1°00'S, 75°29'W) collected with butions of the two species, whose known holotype, MEPN 4554, 1 specimen, 74.9 ranges are separated by over two thousand mm SL. Estero Culebrero, tributary of Rio 556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Yasuni (0°54'45"S, 76°13'03"W) collected Ouro Preto do Oeste, MNRJ 11208, 1 (ho- by R. Barriga, 9 May 1988, USNM 305617, lotype of Gurimata fasciata); USNM 1 specimen, 98.2 mm SL; MEPN 4557, 1 270377, 4 (paratypes of Gurimatafasciata); specimen, 81.8 mm SL. Rio Napo, 2.7 km MNRJ 11271, 4 (paratypes of Gurimata along river above the bridge at Coca fasciata). Jiparand, Rio Urupá, tributary of (0°29.0S, 77°04.0'W), collected by D. Stew- Rio Jiparana, USNM 273306, 3. Rio Ma- art, R. Barriga, and M. Ibarra, 2 Oct 1983, chado system, 20 km upstream ofhparand, USNM 305616, 1 specimen, 66.7 mm SL; USNM 295126, 1. Mato Grosso. Rio Ari- MEPN 4558, 1 specimen, 63.0 mm SL. puand, above Cachoeira de Dardanelos (ap- Peru: Loreto. Rio Nanay, Nanay beach prox. 10°19'42"S, 59°12'30"W), USNM along river west of Iquitos (approx. 3°50'S, 270375, 2 (paratypes of Curimata fasciata); 073°11'W), collected by R. P. Van, H. Or- INPA, 3 (paratypes of Curimata fasciata). tega, A. Gerberich, and J. A. Louton, 17 Rio Aripuana, approximately 10 km above Aug 1986, USNM 280573, 1 specimen, 72.0 Cachoeira de Dardanelos, Cidade de Hum- mm SL. Small stream approx. 65 km up- boldt, USNM 270376, 2 (paratypes of Curi- stream from mouth of Rio Nanay, collected mata fasciata); INPA, 2 (paratypes of Curi- by P. Fromm et al., 18 Aug 1989, ANSP mata fasciata); BMNH 1985.2.5:1-2, 2 164981, 1 specimen, 34.4 mm SL. Rio Pu- (paratypes of Curimata fasciata); MZUSP tumayo drainage, El Estrecho, Quebrada de 28724, 2 (paratypes of Curimata fasciata). Las Granjas, collected by S. 0. Kul- Rio Aripuand, above Cachoeira das An- lander et al., 16 July 1986, NRM SOK/ dorinhas, MHNG 2226.24, 6 (paratypes of 1986293.5292, 1 specimen, 94.0 mm SL. Curimata fasciata). Along road from Genero Herrera towards Peruvian-Brazilian border, Rio Ucayali Acknowledgments drainage, collected by P. Fromm et al., 23 Aug 1989, ANSP 164980, 1 specimen, 39.3 Research associated with this study in mm SL. Peru was supported by the Neotropical Other material examined. — Cyphocha- Lowland Research Program of the Inter- rax multilineatus. Brazil: Amazonas. Rio national Environmental Sciences Program Negro below Daraa, USNM 274102, 1. Rio of the Smithsonian Institution. Collecting Negro at Bucuri, CAS 58605, 1 (holotype activities in Peru were made possible by the of Curimatus multilineatus Myers, formerly generous assistance of Prof. Hernan Ortega IU 17672); CAS-SU 58986, 1. Rio Paduari, (Museo de Historia Natural de la Univer- MZUSP 21161, 1. sidad Nacional Mayor de San Marcos) and Venezuela: Territorio Federal Amazonas. the Consejo Nacional de Ciencia y Tecno- Cañ° La Esmeralda, tributary of Rio Ori- logia of Peru. Prof. Ortega and Dr. Andres noco, SE of La Esmeralda, MBUCV V-4479, Urteaga (Universidad Nacional de la Ama- 1. Rio Mawarinuma (0°55'N, 66°10'W), zonia Peruana) facilitated and participated AMNH uncat., 5. Rio Urumi, tributary to in collecting efforts that resulted in the cap- Rio Negro upstream of Santa Lucia (1°! TN, ture of the new species. Mr. William G. Saul 66°51'W), USNM 270241, 1. Canso Trem- directed my attention towards material of blador where crossed by road from San Car- the new species in the ANSP holdings. Dr. los de Rio Negro to Solano, USNM 269987, Jerry A. Louton (USNM) and Mr. Andrew 7. Rio Barria (0°50'N, 66°10'W), MBUCV Gerberich (USNM) contributed to the suc- V-14898, 1. cess of the expedition. Dr. Sven 0. Kullan- Steindachnerina fasciata. Brazil: Terri der and Mr. E. Ahlander (NRM) arranged tÓnio de Rondonia, Rio Romari (or Sao Do- for the loan of a series of curimatids which mingo) near Nova Unido, MunIcipio of included the specimen of Cyphocharax pan- VOLUME 103, NUMBER 3 557 tostictos from the Rio Putumayo. Figures 1 patterns: proceedings of a workshop. Academia to 3 were prepared by Mr. Theophilus Britt Brasiliera de Ciencias, Rio de Janeiro. Griswold (USNM). This paper benefitted . 1989a. A phylogenetic study of the neotrop- ical Characiform family Curimatidae (Pisces: from the comments and suggestions of Dr. Ostariophysi). —Smithsonian Contributions to Stanley H. Weitzman (USNM) and Dr. Zoology 471:iv +1-25. Thomas A. Munroe (National Marine Fish- . 1989b. Systematics of the neotropical Cha- eries Service, Systematics Laboratory), and raciform genus Curimata Bose (Pisces: Characi- two anonymous reviewers. formes).—Smithsonian Contributions to Zool- ogy 474:iii +1-63. . 1989c. Systematics of the neotropical Cha- Literature Cited raciform genus Eigenmann and Eigenmann (Pisces: Characiformes). — Smith- Fowler, H. W. 1906. Further knowledge on some sonian Contributions to Zoology 481:iii+ 1-42. heterognathus , Part. I. —Proceedings of the . 1990. Systematics of the neotropical Characi- Academy of Natural Sciences of Philadelphia form genus Steindachnerina Fowler (Pisces: Os- 58:293-351. tariophysi). —Smithsonian Contributions to Zo- Kullander, S. 0. 1986. Cichlid fishes of the Amazon ology (in press). River drainage of Peru. Swedish Museum of , & J. Gery. 1985. A new curimatid fish (Cha- Natural History, Stockholm, 431 pp. raciformes: Curimatidae) from the Amazon ba- Myers, G. S. 1927. Descriptions of new South Amer- sin.—Proceedings of the Biological Society of ican fresh-water fishes collected by Dr. Carl Ter- Washington 98:1030-1034. netz. — Bulletin of the Museum of Comparative , &A. W. Van. 1989. Systematics of the Stein- Zoology 68:107-135. dachnerina hypostoma complex (Pisces, Ostar- Ortega, H., & R. P. Van. 1986. Annotated checklist iophysi, Curimatidae), with the description of of the freshwater fishes of Peru. — Smithsonian three new species.—Proceedings of the Biolog- Contributions to Zoology 437:1-25. ical Society of Washington 102:468-482. Stewart, D., R. Barriga S., & M. Ibarra. 1987. Ictio- fauna de la cuenca del rio Napo, Ecuador ori- ental: lista anotada de especies. —Politecnica, Revista de Informacion Tecnico-Cientifica, (RPV) Department of Vertebrate Zoology Quito 12(4):9-63. (Fishes), National Museum of Natural His- Van, R. P. 1988. The Curimatidae, a lowland neo- tory, Smithsonian Institution, Washington, tropical family (Pisces: Characiformes); distri- bution, endemism and phylogenetic biogeog- D.C. 20560; and (RBS) Departamento de raphy. Pp. 313-348 in P. E. Vanzolini, and W. Ciencias Biologicas, Escuela Politecnica Ronald Heyer, eds., Neotropical distribution National, Quito.