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1 Zootaxa 4136 (1): Copyright 2016 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) Oxyropsis ephippia, a New Hypoptopomatine Catfish (Siluriformes: Loricariidae) from Guyana ADRIANA E. AQUINO 1,3 & MARK H. SABAJ PÉREZ 2 1 Division of Vertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, New York, aaquino@amnh.org 2 The Academy of Natural Sciences of Drexel University, 1900 Benjamin Franklin Parkway, Philadelphia, PA sabaj@ansp.org 3 Corresponding author Abstract Oxyropsis ephippia, a new species of loricariid catfish in the subfamily Hypoptopomatinae, is described from the Essequibo and Branco basins, Guyana. Based on the presence of a keel-like row of relatively enlarged odontodes on trunk median plates immediately above the lateral line canal, the new species is assigned to the genus Oxyropsis Eigenmann & Eigenmann It can be distinguished from its congeners by having the keel-like row on each median plate dominated by a single odontode (one closest to the posterior margin of the plate) conspicuously larger than preceding ones, and by having the sides of the trunk below the median series (approximately between median plates 7 to 12) shielded only by plates of the midventral series, with the exclusion of plates from the ventral series. The report of O. ephippia from the Essequibo is the first record of the genus in a river system that drains the north-central portion of the Guiana Shield into the Atlantic Ocean. Key words: Essequibo, armored catfish, taxonomy, new species, key, distribution Introduction Aquino & Schaefer (2002) recognized three species of Oxyropsis Eigenmann and Eigenmann 1889 in their taxonomic revision of the genus: two species in the middle and upper Amazon system (O. carinata (Steindachner 1879) and O. wrightiana Eigenmann and Eigenmann 1889), and one in the upper Orinoco and rio Negro basins (O. acutirostra Ribeiro 1951). The latter species occurs in systems draining the Western Guiana Shield and its outliers further west (i.e., Colombian Guiana Shield; Funk & Kelloff 2009). Hereby we describe a new species of Oxyropsis based, in part, on the first records of the genus from the Essequibo basin, Guyana, which drains the north-central portion of the Guiana Shield into the Atlantic Ocean. The new species is assigned to Oxyropsis based on a shared feature that is autapomorphic for the genus in Loricariidae (Aquino & Schaefer 2002): single row of enlarged odontodes along the trunk midline, lying adjacent and immediately dorsal to the lateral-line canal (Figs. 1B D). Material and methods Counts follow Schaefer (1997) and Aquino & Schaefer (2002) (Fig. 2A). Only the right side series of plate counts are included in the table. The series of median dermal plates bears the lateral-line canal. Total lateral plate counts include all plates in the median series. The first in the ventral series is a triangular plate between the 4th and 5th plates of the midventral series, above the pelvic fin, and that, unlike adjacent midventral plates, does not articulate 1 to 1 with a median plate. Measurements were taken following Buckup (1981) and Aquino & Schaefer (2002) using digital calipers to the nearest 0.1 mm. Throughout the text, BD denotes body depth, HL, head length, CP, Accepted by M.R. de Carvalho: 23 Dec. 2015; published: 5 Jul

2 FIGURE 1. Left lateral view of the posterior caudal peduncle of (A) Oxyropsis ephippia, (B) O. wrightiana, (C) O. carinata, and (D) O. acutirostra. Scale is 1 mm. 130 Zootaxa 4136 (1) 2016 Magnolia Press AQUINO & PÉREZ

3 FIGURE 2. Left lateral view of the trunk plates in (A) O. ephippia, (B) Oxyropsis wrightiana, (C), O. carinata, and (D) O. acutirostra. Vertical lines identify first plate in corresponding series of trunk plates. Scale is 5 mm. caudal peduncle, SL, standard length, alc, alcohol preserved specimen(s) and cs, cleared and stained specimen. The term marginal odontodes refers to the column of enlarged odontodes on the posterior margin of the trunk dermal plates; the expression ventral canal-bearing plate refers to the large triangular dermal plate of the head located ventrally between cleithrum and infraorbital bones that bears the terminus of the preoperculo-mandibular canal; compound pterotic is the term applied to the complex of bones located in the temporal region of the head composed of posttemporal, supracleithrum, pterotic, and ossified Baudelot s ligament; nuchal plate is applied to the plate anterior to the dorsal-fin spine; thoracic plates refers to the dermal plates anterior to the paired cleithra, abdominal plates to the ventral shield of dermal plates between the pectoral and pelvic fins, and anal plate to the plate between the anus, abdominal plates, and pelvic fins. Throughout the text, fully developed individuals (mature specimens) are recognized by having complete development of abdominal plates, which are the last plates to form during ontogeny. Institutional abbreviations are as listed as in Sabaj Pérez (2010). NEW OXYROPSIS EPHIPPIA FROM GUYANA Zootaxa 4136 (1) 2016 Magnolia Press 131

4 Oxyropsis ephippia Aquino & Sabaj Pérez sp. nov. (Figs. 3 5) Holotype: ANSP , female 42.0 mm SL, Guyana, Burro Burro River (Essequibo Drainage), Deer Falls between Deer Creek and Water Dog Falls, 4 39'53"N, 58 50'28"W, G. Watkins et al., 21 November Paratypes (49, mm SL). Guyana, Essequibo Atlantic Ocean Drainage: AMNH , 3 (1 cs) mm SL, Rupununi River 4.6 km NW of village of Massara, 3 55'34"N 59 16'49"W, M. Sabaj et al., 26 October 2002; ANSP , 1, 30.8 mm SL, small creeks crossing Kurupukari Surama River road ca. 3.0 miles from Kurupukari field station, N W, W.G. Saul et al., 4 February 1997; ANSP , 1, 29.0 mm SL, Essequibo River approximately three hours upstream from Kurupukari field station, N W, W.G. Saul et al., 3 January 1997; ANSP , 1, 27.0 mm SL, clearwater creek at campsite 3.1 miles from Kurupukari field station on Kurupukari Surama River road, 4 38'0"N 58 42'59"W, W.G. Saul et al., 5 February 1997; ANSP , 1, 29.5 mm SL, Essequibo River approximately two hours downstream from Kurupukari field station, 4 47'44"N 58 48'52"W, W.G. Saul et al., 20 January 1997; ANSP , 4, mm SL, Essequibo River, sandbar ca. 800 m downstream from Essequibo campsite (Maipuri), N W, W.G. Saul, et al., 29 January 1997; ANSP , 3, mm SL, Essequibo River (east bank) at Kurukupari, 4 39'41"N 58 40'31"W, J.W. Armbruster et al., 24 October 2002; ANSP , 3, mm SL, Rupununi River, 3.7 km SSE village of Massara, 3 51'44"N 59 17'4"W, M. Sabaj et al., 27 October 2002; ANSP , 2, mm SL, AUM 35652, 12, mm SL, INPA 50886, 3, mm SL, and ROM 91534, 3, mm SL, same data as AMNH ; ANSP , 5, mm SL, Simoni River (trib Rupununi), four stations along river from 6.6 km SE to 3.2 km W of Karanambo Ranch, ca. 3 43'9"N 59 15'40"W, M. Sabaj et al., 29 October 2002; ANSP , 1, 33.0 mm SL, Araquai Creek (trib Rupununi), 77.3 km SSE of Lethem, 2 45'45"N 59 27'60"W, M. Sabaj et al. 15 November 2003; MCP 49016, 2 alc, mm SL, same data as ANSP ; MCP 49017, 2, same data as ANSP Branco Negro Amazonas Drainage: ANSP , 2, mm SL, Yuora River (trib Ireng, Takutu drainage), 6.7 km NE of village of Karasabai on road to Tiger Creek village, 4 3'14"N 59 29'7"W, M. Sabaj et al., 31 October Diagnosis. Oxyropsis ephippia is distinguished from its congeners by having a keel-like row of odontodes above the lateral-line canal on each median plate that is dominated by a single odontode (one closest to posterior margin of plate) conspicuously enlarged, typically one-third to one-half larger (along base-tip axis) than preceding one in same row (Figs. 1A and 4A) (vs.keel-like row immediately above lateral-line with odontodes of roughly equal size; Figs. 1B D). It is further distinguished by having the sides of the trunk below the median series, approximately between plates 7 to 12, shielded by plates of the midventral series, with the exclusion of plates from the ventral series, which are visible only in ventral view of the trunk (Fig. 2A) (vs. sides of the trunk shielded by plates of the midventral series and ventral series, the latter visible in lateral and ventral views of the trunk; Figs. 2B D). Oxyropsis ephippia is further distinguished by having 23 plates along the trunk median series (vs in O. acutirostra and in O. carinata and O. wrightiana) and the caudal peduncle in cross section slightly compressed at the level of the median plates (vs. posterior caudal peduncle depressed); from O. acutirostra by having median series of plates complete (vs. series incomplete; Fig. 2D); from both O. acutirostra and O. wrightiana by having a deeper caudal peduncle (mean depth 4.8% SL vs. 2.9% and 2.0%, respectively), and shorter caudal peduncle (mean length 36.0% SL vs. 42.4% and 44.4%, respectively); and from O. carinata by having a larger eye (mean orbital diameter 18.2% HL vs. 15.7% HL). Description. General size small; largest specimen examined 48.7 mm SL. Head and trunk slender, moderately depressed (cleithral width % HL); greatest body depth at dorsal-fin origin ( % SL), only slightly deeper than head ( % SL). Dorsal profile straight to slightly convex from tip of snout to dorsal-fin origin (Figs. 3-4). Dorsal profile of trunk straight from dorsal-fin origin to anteriormost procurrent ray of caudal fin. Ventral profile of head and abdomen approximately straight to slightly convex from snout tip to procurrent caudalfin rays. Dorsal and ventral profiles gently diverging from procurrent towards marginal rays of caudal fin. Snout smoothly rounded, parabolic in dorsal view; dorsal rostrum smooth, with paired smooth concavity in region anterior to naris. Dorsal surface between eyes and origin of dorsal fin convex in cross section, flattening caudally. Cross section of body between pectoral and pelvic-fin origins ovoid, becoming progressively fusiform-shaped (tapering posteriorly) between verticals lines through approximately tip of pectoral fin and median plates 17 18; gradually 132 Zootaxa 4136 (1) 2016 Magnolia Press AQUINO & PÉREZ

5 compressed at level of 6 7 most posterior median plates; CP mean width 75.5% (range %) of CP depth at plates 21 23). FIGURE 3. Oxyropsis ephippia, ANSP , holotype, 42.0 mm SL, Guyana: Burro Burro River (Essequibo Basin). Photos by K. Luckenbill and T. Tran. Eyes moderately large, orbital diameter % HL, center of eye positioned closer to posterior tip of pterotic than to tip of snout. Eye dorsolateral on head; fourth infraorbital lateroventral. Tip of adpressed dorsal fin reaching vertical through midlength of adpressed anal-fin spine. Pelvic fin not reaching plate anterior to base of anal-fin spine. Pectoral fin reaching vertical through posterior half of pelvic-fin spine length. Serrae (teeth) along posterior aspect of pectoral-fin spine present, less developed toward basal fifth and distal fifth of spine; best developed in specimens shorter than about 27.0 mm SL, wherein serrae are conical, moderately acute, each with tip oriented toward base of spine (retrorse); serrae become less conspicuous, more blunt with increasing standard length, and tips variably slanted toward base of spine. Dermal plates on trunk arranged in five longitudinal rows (Fig. 2A): dorsal series 20; middorsal series 3; median series 23; midventral series 13 (including 4 plates between cleithrum lateral process and first plate of ventral series); ventral series typically Ventral plates positioned either lateroventrally (1 to 4 and 11 to 20) or ventrally (5 to 10). Midventral plates deep above ventrally positioned plates in ventral series, shielding side of trunk below median series (Fig. 2A). Lateral line complete, continuous along median series. Midventral plates NEW OXYROPSIS EPHIPPIA FROM GUYANA Zootaxa 4136 (1) 2016 Magnolia Press 133

6 articulating with median plates via interlocking digit-like projections; ventral and median plates in contact via overlapping smooth edges. Thoracic plates absent. Abdominal plates fully developed at 29 mm SL (development of plates incipient at about mm SL); abdominal plates arranged in paired series of 5 9 elongate lateral plates and medial series of 4 9 polygonal plates; three or more polyhedral plates posterior to pectoral girdle; patch of small polyhedral plates between abdominal series and anal plate. Single anal plate. Canal-bearing plate between cleithrum and infraorbital bones well developed, mostly ventral except for posterior ventro-lateral portion in contact with opercle, and with lateral pore of canal close to pore of infraorbital canal between infraorbital plates 4 and 5. Odontodes uniformly small and evenly distributed dorsally on head, progressively arranged in longitudinal rows toward sides and dorsum of trunk. Subtle keel-like longitudinal row of odontodes immediately above lateralline canal on each median plate is dominated by single odontode (close to posterior plate margin above pore of lateral-line canal) that is conspicuously enlarged, typically one-third to one-half longer (along base-tip axis) than preceding one in same row (Fig. 1A); keel-like row most noticeable in dorsal view of trunk between median plates 8 and 18. Enlarged odontode usually paired with similarly enlarged odontode below lateral-line canal. Midventral plates 8 to 13 with area of slightly enlarged odontodes, particularly along margin of plates; odontodes less noticeable with increasing SL. On ventral side of body, odontodes tightly and evenly arranged over ventral canal-bearing plates, abdominal plates, anal plate, and ventral series of plates posterior to anal plate. Odontodes on ventral margin of pelvic-fin spine enlarged; odontodes on ventral margin of pectoral-fin spine moderate in size. Anterior tip of rostral plate with narrow pad of soft tissue; odontodes dorsal and ventral to tip irregularly arranged. Anterior and lateral rostral snout plates not noticeably reflected ventrally. Nuchal plate width about 2.5 times width of dorsal-fin spine at origin. Teeth small, slender; premaxillary teeth 19 27, mandibular teeth Oral disk round, papillose. Maxillary barbels present, short. Dorsal fin i,7; pectoral fin i,6; pelvic fin i,5, anal fin i,5. Adipose fin absent. Sexual dimorphism. Male genital papilla present. In males, odontodes small and tightly packed on plates 2 4 of trunk ventral series, lateral to anal plate and anus region; odontode arrangement more spaced on anal plate and on plates between anal fin and anus. In females, distribution of odontodes on same ventral plates not noticeably different from patterns on adjacent plates. Coloration. Ground color in ethanol tan (Figs. 3 4). Dark brown punctiform epidermal melanophores in small clusters irregularly distributed on dorsal and lateral portions of head (Fig. 4), sometimes leaving pale middorsal region on snout (Fig. 3). Dark brown deep-lying punctiform melanophores roughly arranged in six middorsal blotches (typically, one anterior to dorsal fin, one at base of dorsal fin, and four posterior to dorsal fin). Brown midlateral stripe through cleithrum lateral process, along trunk median and midventral series of plates to base of caudal fin; stripe more diffuse posteriorly, variably connected to darkened lanceolate plates at base of caudal fin. Dorsal portion of cleithrum lateral process, in contact with first midventral plate, dark brown. Ventral surface of head and trunk, between posterior lip and anus, pale, scarcely pigmented. Punctiform melanophores concentrated on anterior lip, extended laterally to barbels; scattered at bases of anal, pectoral and pelvic fins. Uneven sprinkling of deep-lying dark melanophores over ventral canal-bearing plates and along ventral series of plates. Caudal-fin with 6 7 vertical bands of brown melanophores restricted to fin rays; proximal bands variably coalesced into darker, slightly asymmetrical blotch on basal lower half of caudal fin; basal blotch variably connected with dark lanceolate plates (Fig. 5A). Other fins with faint dark bands along anterior edge of spine, continuous with diffuse bands on branched rays (roughly, 8 9 bands on dorsal fin, 3 4 on anal fin, five on pectoral fin, and three on pelvic fin). Interradial membranes of all fins clear, unpigmented. Distribution. Oxyropsis ephippia is recorded from Guyana (Fig. 6) on both sides of the shallow divide separating the Essequibo watershed (Atlantic Drainage) from the Branco (Negro Amazonas Drainage). In the Essequibo Basin, it is known from the mainstem Essequibo, and two of its left bank tributaries (Burro Burro, Rupununi, and their tributaries). In the Branco Basin it is know from the Yuora River, a small tributary of the Ireng draining the southeastern limits of the Pakaraima Mountains. The cichlid Guianacara dacrya described by Arbour & López Fernández (2011) occurs in many of the same streams (e.g., Burro Burro, Rupununi and Yuora), and shares a similar distribution pattern on both sides of the Essequibo Branco divide. The two watersheds are intermittently connected in wet years (between May and September) when the Rupununi savannas between the Pakaraima and Kanuku mountains becomes a vast flooded plain known as Lake Amaku (Lowe McConnell 1964). 134 Zootaxa 4136 (1) 2016 Magnolia Press AQUINO & PÉREZ

7 TABLE 1. Holotype N Range Mean sd Standard length PERCENT OF STANDARD LENGTH Predorsal length Head length Body depth Dorsal-fin spine length Trunk length Pectoral-fin spine length Abdominal length Caudal peduncle length Caudal peduncle depth Head depth Snout length Horizontal eye diameter Least orbit-nare distance Dorsal least interobital distance Ventral least interobital distance Cleithral width Head width Interpelvic distance PERCENT OF HEAD LENGTH Body depth Head depth Snout length Horizontal eye diameter Least orbit-nare distance Dorsal least interobital distance Ventral least interobital distance Cleithral width Head width Interpelvic distance LATERAL SERIES OF PLATES Dorsal series Middorsal series Medial series Midventral series First midventral series Ventral series ABDOMINAL SERIES OF PLATES Right lateral series Left lateral series Medial series TEETH Premaxillary teeth Dentary teeth NEW OXYROPSIS EPHIPPIA FROM GUYANA Zootaxa 4136 (1) 2016 Magnolia Press 135

8 FIGURE 4. Oxyropsis ephippia, ANSP , paratype, XX mm SL, Guyana: Yuora River (trib Ireng, Takutu Branco Negro Dr., Amazon Basin). Habitat. Oxyropsis ephippia generally inhabits small to medium-sized streams with moderate current and riparian forest; water ranges from moderately turbid (e.g., Essequibo mainstem), to clear (Yuora), to black (Burro Burro). A variety of substrates are found at most collecting localities, sand being particularly common. Etymology. The specific epithet ephippia (L. ephippium, Gr. ephippion, meaning saddle) refers to the saddlelike middorsal blotches. Discussion Species of Oxyropsis are sympatric with similar-looking species of Hypoptopoma, and both genera are members of the loricariid subfamily Hypoptopomatinae (sensu Roxo et al. 2014). For example, O. ephippia was collected with H. guianensis in the Rupununi River and its tributary the Simoni in Guyana. Oxyropsis wrightiana and H. psilogaster occur in proximity in the lower río Nanay basin, a moderate blackwater tributary to the río Amazonas in Peru. Oxyropsis wrightiana, however, appears restricted to small tributary creeks whereas H. psilogaster occurs over sandy beaches in the main channel of the Nanay. Diagnostic features of the Oxyropsis aside, abdominal plating and caudal fin pigmentation can be useful to distinguish sympatric species of Oxyropsis and Hypoptopoma. In both genera, the abdomen becomes fully plated in adults (Aquino & Schaefer 2002; 2010); however, Oxyropsis matures at a smaller size. Accordingly, O. ephippia has abdomen fully plated, medial abdominal plates wide, contacting lateral ventral plates in specimens >29 mm SL 136 Zootaxa 4136 (1) 2016 Magnolia Press AQUINO & PÉREZ

9 (Figs. 3 4) vs. medial abdominal plates reduced, leaving unplated gaps on either side in specimens of H. guianense <41.5 mm SL. Similarly, specimens of O. wrightiana >40 mm SL have wide medial abdominal plates contacting lateral ventral plates vs. medial abdominal plates highly reduced or lacking entirely, leaving abdomen naked in specimens of H. psilogaster <47 mm SL. Subtle differences in caudal-fin pigmentation also exist between the aforementioned species (Fig. 5). In O. ephippia, the caudal fin has 6 7 faint vertical bands of brown melanophores restricted to fin rays with proximal bands variably coalesced into darker, slightly asymmetrical blotch on basal lower half of caudal fin (i.e., lower half of 2 3 most proximal bands more intensely pigmented; Fig. 5A). In H. guianense, the caudal fin typically has fewer ( 5), wider brown bands that coalesce along the middle rays and membranes, forming an elongate blotch; the lower half of each band appears darker with melanophores occurring on both the rays and membranes, and the distal tips of the upper and lower caudal lobes are somewhat darkened (Fig. 5C). In O. wrightiana, the caudal fin has 3 4 wide vertical bands of brown melanophores with lower half of two most proximal bands coalesced into a darker, slightly asymmetrical blotch; melanophores occur on both the rays and membranes in the proximal bands, but are restricted to the rays of in the distal bands (Fig. 5B). In H. psilogaster, the vertical bands are scarcely evident in adults, and the pigmentation pattern is dominated by a dark elongate blotch on the middle rays and membranes, a large dark blotch on the distal lower lobe, and a smaller dark blotch on the distal upper lobe (Fig. 5D). There are noteworthy convergences in coloration, morphology and habitat between Oxyropsis ephippia and a species of hillstream lizard loach in the Asian genus Balitoropsis (Cypriniformes: Balitoridae). For example, O. ephippia and Balitoropsis ophiolepis (Bleeker 1853) share six dark brown middorsal saddles and dark bands in the medial and paired fins (see Fig. 11 in Randall & Riggs 2015:220). Shared morphological features include small size, depressed head with elongate, parabolic snout, ventral mouth, ventrally flattened body, and a long and slender caudal peduncle. Balitoropsis also has scales highly textured with longitudinal keels, up to eight keels per scale (Randall & Riggs 2015); the central keel of each scale is enlarged and aligned along the dorsal, lateral and posterior ventral sides. In Oxyropsis, odontodes on the plates are aligned to form longitudinal keels, effecting a similar surface texture. In Oxyropsis, the vent is located near the center of its standard length, midway between the origins of the pelvic and anal fins. In Balitoropsis, the vent is closer to midbody than in other balitorins (i.e., nearer to pelvic-fin base vs. anal-fin origin; Z. Randall, pers. comm. 2015). Balitoropsis and Oxyropsis both occur in relatively small tropical streams in southeastern Asia and cis-andean South America, respectively, and are often associated with woody debris in swift flowing water. The parent taxon of balitorids, order Cypriniformes, is entirely absent from South America whereas the parent taxon of Oxyropsis, family Loricariidae, is endemic to Central and South America. Accordingly, characteristics shared by Oxyropsis and Balitoropsis may represent a globally distributed, strong evolutionary convergence as discussed by Winemiller et al. (2015). In light of the niche classification scheme advanced by Winemiller et al. (2015), one would predict that O. ephippia and B. ophiolepis share a discrete niche defined by a particular combination of functional traits associated with life history, trophic position, habitat, defense and metabolism. Key to the species of Oxyropsis 1 Median plate series truncated, plates along median series (Fig. 2D); caudal peduncle extremely shallow, depth % SL O. acutirostra Ribeiro (Upper Orinoco basin; upper and lower Negro, Amazon basin) - Median plate series complete, plates along median series; caudal peduncle depth greater than 2.5% Plates along median series 23 (Fig. 2A) O. ephippia Aquino & Sabaj Pérez sp. nov. (Essequibo basin; trib Ireng, Takutu Branco Negro drainage, Amazon basin) - Plates along median series (Figs. 2B C) Caudal peduncle depth % SL ( % BD); premaxillary teeth, mandibular teeth; last four median series plates with 1 2 rows of odontodes above and below mid-lateral row of enlarged odontodes (Fig. 1B) O. wrightiana Eigenmann and Eigenmann (Middle and upper Amazon basin) - Caudal peduncle depth % SL ( % BD); premaxillary teeth, mandibular teeth; last four median series plates with 5 6 rows of odontodes above and below mid-lateral row of enlarged odontodes (Fig. 1C) O. carinata (Steindachner) (Middle and upper Amazon basin; lower Negro, Amazon basin) NEW OXYROPSIS EPHIPPIA FROM GUYANA Zootaxa 4136 (1) 2016 Magnolia Press 137

10 FIGURE 5. Pigmentation pattern in caudal fin of (A) Oxyropsis ephippia, 29.1 mm SL (ANSP ) (B) Oxyropsis wrightiana, 50.1 mm SL (ANSP ), (C) Hypoptopoma guianense, 41.4 mm SL (AUM 35652), (D) Hypoptopoma psilogaster, 34.9 mm SL (ANSP ). Vertical line points to enlarged odontode on posterior margin of median plate, immediately dorsal to pore of lateral-line canal. Scale is 5 mm. Photos by K. Luckenbill and T. Tran. 138 Zootaxa 4136 (1) 2016 Magnolia Press AQUINO & PÉREZ

11 FIGURE 6. Drainage map of northern South America showing distribution of species of Oxyropsis (modified from Aquino & Schaefer, 2002). Symbols may indicate more than one collection locality; open symbol represents type locality of O. ephippia. Comparative Material (Additional examined material listed in Aquino & Schaefer, 2002). Hypoptopoma guianense: (all specimens from Guyana) AUM 35652, 3, Rupununi River 4.6 km northwest of Massara, N W; AUM 35658, 26, Simoni River, N, Hypoptopoma psilogaster: (all specimens from Peru, Loreto) ANSP , 1, stream ca. 70 km south of Iquitos, 5 0 S W; ANSP , 2, río Itaya at Iquito Nauta hwy bridge; ANSP , 26, río Nanay near Santa Clara, S, W; ANSP , 1, río Nanay at Pampa Chica, S W; ANSP , 1, río Nanay 7 km west of Iquito, S W. Oxyropsis acutirostra: ANSP , 22, Colombia, Guainía, río Inirida, caño Caranacoa, upstream of Puerto Inirida, N W; ANSP , 1, ANSP , 1, Venezuela, Amazonas, caño Marujeta, trib Ventuari, N W; ANSP , 9, Venezuela, río Guapuchi, trib Ventuari, N W; AUM 43090, 1, Venezuela, Amazonas, caño Grulla, trib Orinoco, N W. Oxyropsis wrightiana: (all specimens from Perú, Loreto) ANSP , 1, creek trib to río Momon, Nanay Dr., 3 42 S W; ANSP , 3, río Yanayacu, 25 miles south of Iquitos; ANSP , 1, caño Shirui, trib río Nanay, S W; ANSP , 1, Moena caño and mouth of Ullpa caño near Iquitos, S W. NEW OXYROPSIS EPHIPPIA FROM GUYANA Zootaxa 4136 (1) 2016 Magnolia Press 139

12 Acknowledgments We thank the following individuals and institutions for relevant information and field assistance: C. Allison, J. Armbruster, N. Lujan, and D. Werneke (AUM), W. Eschmeyer (CAS), R. Ota, M. Rocha (INPA), E. Holm (ROM), M. Thomas (SIUC), Z. Randall (UF), and M. Hardman. Thanks to Kyle Luckenbill (ANSP) and Trung Tran (Drexel University) for photographic assistance. AEA also thanks B. Brown and S. Schaefer (AMNH) for their support in her research. Project supported in part by grants to MHSP from U.S. National Science Foundation (DEB , DEB ). References Aquino, A.E. & Schaefer, S.A. (2002) Revision of Oxyropsis Eigenmann & Eigenmann, 1889 (Siluriformes, Loricariidae). Copeia, 2002, Aquino, A.E. & Schaefer, S.A. (2002) Systematics of the genus Hypoptopoma Günther, 1868 (Siluriformes, Loricariidae). Bulletin of the American Museum of Natural History, 336, Buckup, P.A. (1981) Microlepidogaster taimensis sp. n., novo Hypoptopomatinae da Estação Ecológica do Taim, Rio Grande do Sul, Brasil (Ostariophysi, Loricariidae). Iheringia, Série Zoology, 60, Arbour, J.H. & López-Fernández, H. (2011) Guianacara dacrya, a new species from the rio Branco and Essequibo River drainages of the Guiana Shield (Perciformes: Cichlidae). Neotropical Ichthyology, 9, Funk, V.A. & Kelloff, C.L. (2009) Introduction. In: Vari, R.P., Ferraris, C.J. Jr., Radosavljevic, A. & V. & Funk, A. (Eds.), Checklist of the freshwater fishes of the Guiana Shield. Bulletin of the Biological Society of Washington, No. 17, pp Lowe-McConnell, R.H. (1964) The fishes of the Rupununi savanna district of British Guiana, South America. Part 1. Ecological groupings of fish species and effects of the seasonal cycle on the fish. Journal of the Linnean Society (Zoology), 45, Randall, Z.S. & Riggs, P.A. (2015) Revision of the hillstream lizard loaches, genus Balitoropsis (Cypriniformes: Balitoridae). Zootaxa, 3962 (1), Roxo, F.F., Albert, J.S., Silva, G.S., Zawadzki, C.H., Foresti, F. & Oliveira, C. (2014) Molecular phylogeny and biogeographic history of the armored neotropical catfish subfamilies hypoptopomatinae, neoplecostominae and otothyrinae (siluriformes: loricariidae). PLoS One, 9 (8), e Sabaj Pérez, M.H. (Ed.) (2014) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 5.0 (22 September 2014). American Society of Ichthyologists and Herpetologists, Washington, DC. Electronically accessible. Available from: (accessed 26 January 2016) Schaefer, S.A. (1997) The Neotropical cascudinhos: systematics and biogeography of the Otocinclus catfishes (Siluriformes: Loricariidae). Proceedings of the Academy of Natural Sciences of Philadelphia, 148, Vari, R.P & Ferraris, C.J. (2009) Fishes of the Guiana Shield. In: Vari, R.P., Ferraris, C.J. Jr., Radosavljevic, A. & Funk, V.A. (Eds.), Checklist of the freshwater fishes of the Guiana Shield. Bulletin of the Biological Society of Washington, No. 17, pp Winemiller, K.O., Fitzgerald, D.B., Bower, L.M. & Pianka, E.R. (2015) Functional traits, convergent evolution, and periodic tables of niches. Ecological Letters 18, Zootaxa 4136 (1) 2016 Magnolia Press AQUINO & PÉREZ