taxonID	type	description	language	source
9366E1A675D8472F37550BBE31106656.taxon	discussion	Introduction With 650 species currently considered valid (Eschmeyer 2003), the Loricariidae is the most speciose family of catfishes in the world. Loricariids are typically algivorous or detritivorous, but the Hypostomus cochliodon group (formerly the genus Cochliodon Kner) and Panaque Eigenmann are unique among fishes in that they consume wood (Schaefer & Stewart 1993; Nelson et al. 1999). The H. cochliodon group and Panaque share the derived presence of large, spoon-shaped teeth; however, they are unrelated and are placed in two different tribes, the Hypostomini and the Ancistrini, respectively (Armbruster 1997; in press). The original description of Cochliodon was by Heckel (in Kner 1853), but the genus was described in the synonymy of Hypostomus Lacepede. Eigenmann (1922) described Cheiridodus and separated the genus from Cochliodon based on the presence of a small medial tooth cusp (vs. medial cusp absent). Most loricariids have bicuspid teeth (Muller & Weber 1992), and the presence of a mesial cusp represents a plesiomorphic characteristic within the Loricariidae. Cochliodon do actually have a small mesial cusp, but this cusp is occasionally fused into the lateral cusp and visible as a darker, thicker ridge on the tooth (pers. obs.). Isbruecker (1980) recognized Cheiridodus as a synonym of Cochliodon, but did so without comment. Armbruster (1997; in press) provided a phylogeny for the species of the Hypostominae based on morphology and determined that Cochliodon is derived from Hypostomus. In addition, Montoya-Burgos et al. (1998) found Cochliodon to be related to Hypostomus based on sequences of the 12 s and 16 s rRNA genes, Montoya-Burgos et al. (2002) found Cochliodon to be nested within Hypostomus based on sequence data from the mitochondrial D-loop, and Zawadzki (pers comm.) has found Cochliodon to be derived from Hypostomus based on allozymes. Armbruster (1997, in press) recognized Cochliodon as a synonym of Hypostomus and refers to the species formerly in Cochliodon as the H. cochliodon group. Weber and Montoya-Burgos (2002) and Montoya-Burgos et al. (2002) also placed Cochliodon in the synonymy of Hypostomus. The Hypostomus cochliodon group has received little attention from authors except for original species descriptions. The seven currently accepted species of the H. cochliodon group are distributed in the Orinoco, Amazon, Essequibo, Magdalena, Paraguay, and Atrato river basins and in the Lake Maracaibo basin (Lilyestrom 1984; Armbruster & Page 1997). There has only been one modern attempt to examine the species of the H. cochliodon group. Lilyestrom (1984) provides descriptions of the species of the H. cochliodon group in Venezuela, a key to all of the species of the H. cochliodon group, and places Cochliodon pospisili Schultz into the synonymy of H. hondae. The characteristics used in Lilyestrom’s key are mostly proportions and tooth counts and do not adequately separate the species of the H. cochliodon group (pers. obs.). Armbruster and Page (1997) redescribe Rhinelepis levis Pearson, and place the species in Cochliodon. Hypostomus levis is unique among the H. cochliodon group in the absence of an adipose fin. Weber and Montoya-Burgos (2002) describe H. fonchii and suggest that it is related to the H. cochliodon group; however, they present no credible evidence for this assertion and H. fonchii is not considered to be part of the H. cochliodon group in this study. Placing Cochliodon into the synonymy of Hypostomus is further supported by two species described herein. Hypostomus hemicochliodon and H. sculpodon predominantly eat wood, but do not have spoon-shaped teeth. These species have teeth that appear to be intermediate between other Hypostomus and other species of the H. cochliodon group (Fig. 1) and also appear to eat less wood than the other species of the H. cochliodon group (pers. obs. based on gut contents). Although many other Hypostomus will occasionally consume small amounts of wood, wood only amounts to a very small fraction of the diet (pers. obs.). In this manuscript all species of the H. cochliodon group are redescribed, four new species of the H. cochliodon group are described, and distribution maps, a key, and a phylogeny for the species of the H. cochliodon group is provided.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
041EA1EDF99BE4510A1F08CCE85576A4.taxon	diagnosis	Diagnosis: The Hypostomus cochliodon group is diagnosed by the following characteristics, none of which is unique among loricariids: dentary angle averaging less than 80 ° and the preoperculo-hyomandibular ridge deflected posterior to the main body of the hyomandibula (Armbruster, in press). Wood-specializing members of the H. cochliodon group are additionally diagnosed by the following characteristics, none of which are unique among loricariids: the presence of large spoon-shaped teeth (Fig. 1 C), the hyomandibula and metapterygoid sutured to their dorsal margins vs. not sutured dorsally with a notch between the two bones (Fig. 3 B), the presence of a highly curved maxilla (Fig. 3 B), and loss of the buccal papilla (Fig. 4 B). The Hypostomus cochliodon group can be distinguished from most other loricariids by the combination of highly angled jaws and usually less than 20 teeth per jaw ramus (occasional specimens of H. cochliodon, H. hondae, and H. hemicochliodon have been observed with counts of up to 27 teeth). Among the Hypostominae, the H. cochliodon group differs from Pterygoplichthys by having seven dorsal-fin rays (vs. nine or more); from Pogonopoma, Pseudorinelepis, and Rhinelepis by having the dorsal flap of the iris present (vs. flap absent; Armbruster 1998) and by having four anal fin rays (vs. five); from Corymbophanes, by having 0 - 2 median, unpaired plates posterior to the dorsal fin (vs. 3 or more); from most of the Ancistrini by not being able to evert the cheek plates any greater than 30 ° (vs. 75 ° - 90 °); from Spectracanthicus (Ancistrini) by having the dorsal fin separate from the adipose fin (vs. contacting adipose-fin spine) and by having plates on the abdomen in adults; and from Pseudancistrus (Ancistrini) by lacking hypertrophied odontodes on the cheek and snout margin and by having plates on the abdomen in adults. Hypostomus microstomus Weber shares with the H. cochliodon group highly angled jaws and a low number of large teeth (Weber 1987); however, the teeth of H. microstomus are not spoon-shaped or tending towards being spoon-shaped. In addition, H. microstomus is dark gray to black with light spots versus brown with dark spots as in the H. cochliodon group. Some Hypostomus such as H. plecostomus are variable in the angle between the dentaries. Some specimens of H. plecostomus, for example, have a dentary angle less than 80 °. They differ from the H. cochliodon group by having greater than 20 teeth per ramus. Species similar to H. plecostomus also tend to have the head as wide or wider than tall while species of the H. cochliodon group tend to have the head taller than wider.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
041EA1EDF99BE4510A1F08CCE85576A4.taxon	description	Description: Fairly large loricariids, reaching 300 mm SL. Color typically dark brown with spots generally developed over most surfaces. Most species observed have a well developed ability to alter color according to substrate. Body shape in all except Hypostomus sculpodon deep at origin of the dorsal fin (appearing deeper than in other Hypostomus) then narrowing posteriorly more quickly than in Hypostomus causing body to appear humped. Body of H. sculpodon not as deep. Body depth increases from snout to tip of supraoccipital at steep angle, angle of body depth increase decreases from tip of supraoccipital to dorsal-fin spinelet. Rounded ridge present from anterodorsal corner of orbit to posterior margin of nares; ridge widest and tallest posteriorly. Rounded ridge present from posterodorsal corner of eye to end of pterotic-supracleithrum (additional, sharp ridge of bone and moderately enlarged odontodes present on pterotic-supracleithrum in H. cochliodon, H. hemicochliodon, and H. sculpodon). Space between orbits concave such that dorsal rim of orbit raised above medial surface of head. Nares separated by flap of skin held erect in life. Dorsal, supramedian, median, and inframedian plate rows complete from head to caudal fin, ventral plate row begins at insertion of pelvic fin and continues to caudal fin. Lateral plates usually with median keels formed from ridge of bone and enlarged odontodes; height of keels vary from absent to tall with odontodes very sharp; keels may be present or absent on any row of lateral plates; keels of first three plates of supramedian plate row angled dorsally, often confluent with keel of dorsal plate row; keels on first three plates of dorsal row forming angle from tip of supraoccipital to posterolateral corner of nuchal plate, not confluent with keel on dorsal plate row beginning on fourth plate. Base of caudal fin covered in elongate, roughly triangular plates. Entire ventral surface of head and body (including space above pectoral- and pelvic-fin rays) of adults covered in small platelets, platelets often extending onto base of pectoral- and pelvic-fin rays ventrally. Small platelets usually present in skin between dorsal fin and lateral plates of adults. Platelets on abdomen and near fins increase in number with standard length. Head covered in small plates. Frontal, nasal, sphenotic, infraorbitals, pterotic-supracleithrum, suprapreopercle, and supraoccipital supporting odontodes. Opercle supports odontodes in some species. Some odontodes present on posterior margin of preopercle. Platelets that cover anteroventral corner of opercle slightly separated from opercle allowing plates to be marginally everted (angle of eversion less than 30 °). Dorsal fin consisting of small, v-shaped spinelet, fairly strong spine, and seven rays. Caudal fin strongly forked, lower lobe longer than upper. Pectoral-fin spine strong, generally reaches posterior to pelvic-fin rays when depressed ventral to pelvic fin; cleithrum with exposed process dorsal to pectoral-fin rays that tapers posteriorly to point; pectoralfin inserted on same plane as pelvic fin such that spine, when depressed parallel with body, lies on top of and in contact with pelvic fin. Pelvic-fin spine thin, flexible, generally reaches beyond base of anal fin. Anal fin with relatively strong, unbranched first ray that supports odontodes. Adipose fin (when present) consisting of single median, unpaired preadipose plate and a stout, strong, pointed spine; adipose-fin membrane generally extending to anterior-most procurrent caudal-fin spine or just anterior to procurrent caudal-fin spines. Dorsal fin II 7; pectoral fin I 6; pelvic fin I 5, anal fin I 4, caudal fin I 14 I. Jaws strongly angled, dentaries typically forming angle of less than 80 °. Teeth few (5 - 27, mode = 10), generally shorter and wider than most Hypostomus (Fig. 1 B-C); teeth of most species spoon-shaped (Fig. 1 C).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
041EA1EDF99BE4510A1F08CCE85576A4.taxon	distribution	Range: From the Amazon, Aroa, Atrato, Essequibo, La Plata, Magdalena, Orinoco, Sinu, Tocuyo, Tuy, and Yaracuy river drainages and the Lake Maracaibo drainage.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
041EA1EDF99BE4510A1F08CCE85576A4.taxon	biology_ecology	Ecology: Individuals of the Hypostomus cochliodon group are most typically collected in slow-flowing, small- to medium-sized rivers, although they may also be collected in piedmont streams. They are typically associated with submerged logs in the flowing portions of the streams.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
A64BAE23EB74483188A9FCA0B8742B5C.taxon	description	(Fig. 7)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
A64BAE23EB74483188A9FCA0B8742B5C.taxon	materials_examined	Material examined: UNKNOWN LOCALITY. AMNH 97884, 2, (80.9 - 155.5). BRAZIL. Mato Grosso: Rio Cujaba, NMW 46277, 1 (152.9, lectotype) and NMW 44101 (179.3, paralectotype). Near Cuiaba, AMNH 97880, 9, 1 cs, (68.9 - 124.2). Arroio, Rio do Bugre - Rio Jauru - Rio Paraguai dr. at km 165 on Estrada Porto Esperidiao / Pontes e Lacerda (BR 174), ca 48 km de Esperidiao, USNM 326357, 4, 1 cs, (75.8 - 143.6). Arroio Cruzando, Rio Paraguai dr. on estrada Tangara da serra 1, Barra do Bugres, near nova Olimpia, USNM 326319, 2, 1 cs, (70.5 - 96.6). PARAGUAY. Alto Paraguay: Arroyo Ytyguazu, Rio Paraguay dr. Primavera, Caacupe, ANSP 124105, 1, (111.4). Rio Apa, Rio Paraguay dr. ca 0.5 km upstream (= east) of bridge between Brazil and Paraguay in Bella Vista, 22 ° 06 ’ 30 ” S, 56 ° 30 ’ W, UMMZ 206797, 6, (122.6 - 211.38). Amambay: Rio Aquidaban, Rio Paraguay dr. at Paso Horqueta, ca 24 km NNW of Loreto, 23 ° 03 ’ 48 ” S, 57 ° 23 ’ W, UMMZ 207821, 1, (151.4). Rio Aquidaban, Rio Paraguay dr. in Parque Nacional Cerro Cora ca 32 km WSW of Pedro Juan Caballero, 22 ° 38 ’ 12 ” S, 56 ° 03 ’ W, UMMZ 206767, 1, (130.0). Rio Aquidaban, Rio Paraguay dr. Parque Nacional Cerro Cora, USNM 232310, 1, (131.1), USNM 232312, 2, (119.4 - 164.4), and USNM 232311, 1, (146.7). Canendiyu: Rio Jejui, Rio JejuiGuazu - Rio Paraguay dr. ca. 41 km N Curuquaty and 2 km S Ygantini, 24 ° 08 ’ 12 ” S, 55 ° 37 ’ W, UMMZ 206338, 3, 3 cs, (142.2 - 168). Concepcion: Rio Ypane, Rio Paraguay dr. at ferry crossing, ca 2.0 km S by dirt road to Belen (which is ca 18 km ESE Concepcion), 23 ° 29 ’ 30 ” S, 57 ° 15 ’ W, UMMZ 207988, 5, 2 cs, (80.3 - 113.1) and UMMZ 207989, 3, (66.1 - 76.3). Cordillera: Rio La Plata dr. 1.6 km by road S of Tobati, UMMZ 203865, 1, (140.8). Arroyo Tobati, Rio Paraguay dr. small river ca. 0.3 km E of dirt road, 1.6 km S of Tobati, 25 ° 18 ’ 30 ” S, 57 ° 04 ’ W, UMMZ 205771, 1, (151.6) and UMMZ 205770, 1, (148.0). Arroyo Tobati, Rio Piribebuy - Rio Paraguay dr. ca 0.1 km east of Tobati, 25 ° 16 ’ 14 ” S, 57 ° 03 ’ W, UMMZ 205809, 1, (106.5). Arroyo Yaguary-Guazu, Rio Aquidaban dr. at bridge on dirt highway (Route # 5) ca 26.1 km NE of junction with Route #, ca 68 km WSW of Pedro Juan Caballero, 27 ° 47 ’ 24 ” S, 56 ° 21 ’ W, UMMZ 206691, 1, 147.0). Paraguari: Rio Tebicuary dr. Salto de Pirareta, ca. 400 m below falls, 5.8 km S. of Piribebuy and 10 km W on dirt road, UMMZ 205673, 3, (112.4 - 170.5). San Pedro: Rio Ypane, Rio Paraguay dr. on S bank at bridge on dirt highway, 52.7 km S of jct. with highway # 5 at Yby-Yau, 23 ° 27 ’ 18 ” S, 56 ° 31 ’ W, UMMZ 208023, 1, (164.0) and UMMZ 208024, 2, (94.9 - 120.1). Villa Hays: no exact locality, FMNH 59735, 1, (151.7).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
A64BAE23EB74483188A9FCA0B8742B5C.taxon	diagnosis	Diagnosis: Hypostomus cochliodon can be distinguished from all other members of the H. cochliodon group by its almost entirely brown coloration with ventral half of dorsal plate row and dorsal half of supramedian plate row slightly lighter than lower rows forming a tan stripe, and spots, when present, small and widely placed (vs. spots usually welldeveloped and closely placed). Unlike most other members of the H. cochliodon group, H. cochliodon can also be entirely dark brown with no spots anywhere on the body (other members of the H. cochliodon group may be very dark, but will retain spots on fins or the abdomen).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
A64BAE23EB74483188A9FCA0B8742B5C.taxon	description	Description: See Hypostomus cochliodon group description for more details. Morphometric data given in Table 1. Color normally almost entirely dark brown with wide, tan stripes dorsally and ventrally. Tan dorsal stripe on ventral half of dorsal plate row and dorsal half of supramedian plate row; stripe sometimes crossed by dark saddles or with dark spots anteriorly. Tan ventral stripe on ventral half of inframedian plate row and entire ventral plate row. Dorsal stripe forks on nape with one branch continuing straight to eye and other branch ending at posterior tip of supraoccipital. Tan wedge occasionally present at base of supraoccipital crest. Dorsal surface of snout light or mottled. Spots on body often absent, but sometimes present on any part of the body, well separated from one another. Abdomen colored as sides, but usually slightly lighter. Color entirely dark brown without spots in some individuals. Dorsal fin usually short, not reaching preadipose plate when depressed in most specimens. Depressed pectoral-fin spine ventral to pelvic fin reaches beyond bases of pelvic-fin rays. Tip of pectoral-fin spine of nuptial males with stout, recurved, hypertrophied odontodes. Keels moderate to well-developed; keel odontodes occasionally sharp. Orbits forming ridge distinctly raised above medial surface of head; ridges of dorsal and lateral aspect of head well-developed. Longitudinal ridge on pterotic-supracleithrum beginning at posterodorsal corner of eye formed from raised bone and slightly larger odontodes present. Opercle usually supporting much more than 10 odontodes; in some specimens the opercle does not support odontodes (see Comments). Nuptial body odontodes present (Fig. 2 B). Plates in skin anterior to dorsal-fin spines few or absent (Fig. 6 A; Table 7). Each jaw with 6 - 27 teeth (mode = 12). Size of teeth variable (see Comments). Average angle between dentaries 68 ° (SD = 14 °; range: 42 ° - 99 °; N = 30). Lateral line plates 26 - 30; dorsal plates 7 - 8; interdorsal plates 5 - 7; adipose caudal plates 9 - 11.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
A64BAE23EB74483188A9FCA0B8742B5C.taxon	distribution	Range: The upper 2 / 3 of the Rio Paraguay basin of Brazil and Paraguay (Fig. 8). Hypostomus cochliodon is allopatric to all other species of the H. cochliodon group.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
A64BAE23EB74483188A9FCA0B8742B5C.taxon	discussion	Comments: The smallest juveniles often have teeth very similar to algivorous-detritivorous species of Hypostomus. As Hypostomus cochliodon grow, the teeth apparently are replaced by successively larger, more spoon-shaped teeth, and tooth number decreases. Hypostomus cochliodon is variable in color pattern, size of keels, and number of odontodes on the opercle (0 to 30). The variation in number of odontodes may indicate that H. cochliodon as presently delimited may represent more than one species; however, the variation in opercular odontode number does not appear to be correlated with variation in other characteristics.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
43B23634594A5946DD4890010756F37F.taxon	description	(Fig. 9)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
43B23634594A5946DD4890010756F37F.taxon	diagnosis	Diagnosis: Hypostomus ericius can be separated from all other members of the H. cochliodon group by coloration: light tan-gray with well-separated spots and no stripes. In addition, H. ericius differs from H. hemicochliodon by a lack of a buccal papilla and from H. pyrineusi by the presence of strong, sharp keels on the lateral plates. Hypostomus ericius is most similar to H. oculeus from which it differs mainly in coloration and morphometrics (Fig. 10 B).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
43B23634594A5946DD4890010756F37F.taxon	description	Description: See Hypostomus cochliodon group description for more details. Morphometric data given in Table 1. Coloration generally tan-gray with large, widely-spaced spots. Spots present over entire body and becoming larger posteriorly. Caudal-fin membrane and rays often with black wash distally, and lighter area proximally instead of being spotted. Spots on fin spines and rays occasionally large. Spots less pronounced in juveniles, spots may be absent on abdomen. Juveniles with four faint dorsal saddles: first below anterior rays of dorsal fin, second below posterior rays of dorsal fin and slightly posterior to dorsal fin, third slightly anterior to and ventral to adipose fin, and fourth at base of caudal fin. Dorsal fin moderately long, usually just barely reaching preadipose plate when depressed. Depressed pectoral-fin spine ventral to pelvic fin reaches beyond bases of pelvic-fin rays. Pectoral-fin spine supporting numerous stout, recurved, hypertrophied odontodes in nuptial males. Keels sharp, very strongly developed. Orbits forming ridge distinctly raised above medial surface of head; ridges of dorsal and lateral aspect of head well-developed. Longitudinal ridge formed of raised bone and slightly larger odontodes absent on pterotic-supracleithrum beginning at postdorsal corner of orbit. Opercle usually not supporting odontodes, but sometimes up to 10 odontodes may be present on opercle (Fig. 2 B). Nuptial body odontodes present (Fig. 2 B). Plates in skin anterior to dorsal-fin spine usually absent or not numerous (Fig. 6 A; Table 7). Cheek plates generally support several stout odontodes slightly larger than surrounding odontodes. Head appears wider and taller than in other species of Hypostomus. Each jaw with 5 - 9 teeth (mode = 7), teeth large and spoon-shaped. Average angle between dentaries 53 ° (SD = 9 °; range: 40 ° - 69 °; N = 7). Lateral line plates 27 - 28; dorsal plates 7 - 9; interdorsal plates 5 - 7; adipose caudal plates 9 - 10.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
43B23634594A5946DD4890010756F37F.taxon	distribution	Range: Currently known from the Rios Maranon, Itaya, and Nanay of the Upper Rio Amazonas drainage of Peru (Fig. 11). Hypostomus ericius is sympatric with H. hemicochliodon, H. oculeus, and H. pyrineusi.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
43B23634594A5946DD4890010756F37F.taxon	etymology	Etymology: From the Latin, ericius, for hedgehog in reference to the many sharp odontodes on the keels.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
BFFAEFD58F8374B5423CC80E8485EC3F.taxon	description	(Fig. 12)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
BFFAEFD58F8374B5423CC80E8485EC3F.taxon	materials_examined	Nontype material: BRAZIL. Mato Grosso, Rio Juruena, Rio Tapajos dr. upper Rio Juruena, USNM 199225, 1, (119.3). Rio Xingu, Rio Amazonas dr. Sao Felix, USNM 207509, 1, (82.7). ECUADOR: Napo, Rio Jivino, Rio Napo dr. lower 4 km (mostly) to 0.6 km upstream from mouth (most between the two ports of Limoncocha), 00 ° 24 ’ 36 ” S, 76 ° 39 ’ 00 ” W, FMNH 106021, 1, (105.3). Rio Shushufindi, Rio Aguarico dr. lower reaches, about 2 km from mouth in Rio Aquarico, 00 ° 17 ’ 30 ” S, 76 ° 25 ’ 36 ” W, FMNH 106025, 2, (141.5 - 157.9). Tributary, Rio Payamino dr. few km upstream from San Jose de Payamino, 00 ° 20 ’ 12 ” S, 77 ° 18 ’ W, FMNH 106024, 1, (158.0). Tributary, Rio San Miguel dr. km 50, 4.5 km S of Tipischa, FMNH 106022, 1, (208.7). Tributary, Rio Churuyacu - Rio Payamino dr. Tiuyacu, first tributary of Rio Churuyacu upstream from mouth in Rio Payamino and near mouth of Rio Churuyacu, 00 ° 29 ’ 30 ” S, 77 ° 18 ’ W, FMNH 106023, 3, (112.6 - 197.2). Rio Corrientes, Rio Napo dr. eastern Ecuador, FMNH 92633, 3, 1 cs, (78.6 - 89.8). VENEZUELA. Amazonas, Cano Jenita, Rio Ocamo dr. Oramo department, in an old meander 1 km river km above the Rio Ocamo, 02 ° 46 ’ N, 64 ° 54 ’ W, MCNG 24332, 1, (142.3). Laguna Pacon, Rio Orinoco dr. in pools behind beach along Rio Ventuari about 0.5 hr above mouth, MBUCV V- 23092, 1, (165.2). Rio Cataniapo, Rio Orinoco dr. in Las Palvas, MBUCV V- 24501 (none measured). Rio Mavaca, Rio Orinoco dr. river above camp base, MBUCV V- 19239, 1, (186.1). Rio Siapa, Rio Casiquiare - Rio Negro dr. Departmento Rio Negro, at campsite near Laguna Cumicapi between Rio Emoni and Rio Manipitare, MCNG 37041, 1, (100.3). Rio Siapa, Rio Casiquiare dr. near and downstream of confluence with Rio Manipitare, 01 ° 54 ’ 06 ” N, 65 ° 57 ’ 05 ” W, MCNG 37032, 1, (199.5).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
BFFAEFD58F8374B5423CC80E8485EC3F.taxon	diagnosis	Diagnosis: Hypostomus hemicochliodon differs from all other members of the H. cochliodon group except H. sculpodon by the presence of a medium-sized buccal papilla (condition intermediate between those shown in figures 4 A and 4 B), intermediately developed teeth (Fig. 1 B; juvenile H. cochliodon also occasionally have intermediate teeth), and a large patch of odontodes on the opercle. Hypostomus hemicochliodon differs from H. sculpodon by coloration (dark brown with closely spaced spots vs. brownish-red with widely spaced spots), by having a greater dorsal-anal length to anal-fin length ratio (average = 111.1 ± 8.4 %, 92.5 - 132.6 % vs. average = 79.2 ± 7.6 %, 69.6 - 89.7 %), and almost completely by having a larger interorbital width / HL ratio (average = 49.9 ± 2.7 %, 44.1 - 55.6 % vs. average = 51.4 ± 3.1 %, 38.5 - 45.4 %).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
BFFAEFD58F8374B5423CC80E8485EC3F.taxon	description	Description: See description of the Hypostomus cochliodon group for more details. Morphometric data given in Table 2. Color brown with medium to large spots. Spots become larger posteriorly, often coalescing on caudal peduncle to form large, longitudinally oval dashes or stripes. Some adults with spots on body absent. Spots on abdomen variable; fairly large specimens often have few or no spots on the abdomen; abdomen often significantly lighter than sides, almost white. Four faint dorsal saddles often present: first below anterior rays of dorsal fin, second below posterior rays of and slightly posterior to dorsal fin, third slightly anterior to and ventral to adipose fin, and fourth at base of caudal fin. One captive Hypostomus hemicochliodon demonstrated well-developed ability to change color, particularly ability to lighten coloration of abdomen and intensify lateral spots or dorsal saddles (either spots or the saddles dominant, but never both dark at same time). Dorsal fin fairly short, usually falls short of preadipose plate when depressed. Depressed pectoral-fin spine ventral to pelvic fin reaches 2 - 3 plates beyond bases of pelvic-fin rays. Tip of pectoral-fin spine of nuptial males with stout, recurved, hypertrophied odontodes. Keels strong, sharp. Orbit forming ridge distinctly raised above medial surface of head; ridges of dorsal and lateral aspect of head well-developed. Longitudinal ridge formed of raised bone and slightly larger odontodes present on pterotic-supracleithrum beginning at postdorsal corner of orbit. Opercle broadly exposed, always supporting much more than ten odontodes. Nuptial body odontodes absent (Fig. 2 A). Plates in skin anterior to dorsal-fin spine absent or few (Fig. 6 A; Table 7). Each jaw with 7 - 23 teeth (mode = 15), teeth of intermediate size (Fig. 1 B). Average angle between dentaries 63 ° (SD = 7 °; range: 45 ° - 75 °; N = 23). Lateral line plates 28 - 30; dorsal plates 8 - 9; interdorsal plates 6 - 8; adipose caudal plates 8 - 10.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
BFFAEFD58F8374B5423CC80E8485EC3F.taxon	distribution	Range: Has a wide range across central South America in the Rios Maranon, Itaya, Nanay, Napo, and Orosa of the upper Rio Amazonas of Peru, the Rio Napo drainage of the Upper Rio Amazonas drainage of Ecuador, the Rios Tapajos and Xingu of the lower Rio Amazonas drainage of Brazil, and the upper Rio Negro drainage (middle Rio Amazonas drainage) and upper Rio Orinoco drainage of Venezuela. (Fig. 8). Hypostomus hemicochliodon is sympatric with H. ericius, H. oculeus, H. pyrineusi, and H. sculpodon.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
BFFAEFD58F8374B5423CC80E8485EC3F.taxon	etymology	Etymology: From the Latin, hemi, for half, the Latin, coclea (coch-), for spoon, and the Greek, odon, for tooth in reference to the fact that the teeth are about half as spoonshaped as those of wood-specializing members of the Hypostomus cochliodon group.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
4BA3665C2F9A19A50461F6627B07F4EC.taxon	description	(Figs. 13, 14 A)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
4BA3665C2F9A19A50461F6627B07F4EC.taxon	materials_examined	Material examined: COLOMBIA. Rio Sinu dr. Lorica, CAS 149472, 1, 1 cs, (81.7). Probably Rio Sinu, Lorica, CAS 6417, 1, (177.0). Atrato: Quibdo market, STRI 1530, 1, (207.9) and STRI 1531, 1, (205.9). Caldas: Rio Samana La Miel, Rio Magdalena dr. at or near junction of Rio Samana La Miel near La Dorada, CAS 150373, 1, (104.7). Santander: cano Tigre, Rio Sogamosa - Rio Magdalena dr. SIUC 34909, 1, (119.0). Sucre: Pozo del Chorro, Sinclejo, CAS 149472, 4, (81.7 - 107.4) and USNM 175305, 2, (79.5 - 102.1). Tolima: Honda, Colombia, BMNH 1909.7.23.43 (60.8, holotype) and BMNH 1909.7.23.44 (58.4, paratype). VENEZUELA. Merida: Rio Escalante, Lago Maracaibo dr. at bridge of highway 1, 08 ° 31 ’ N, 71 ° 47 ’ W, MCNG 24843, 3, (75.0 - 81.4) and INHS 59864. Tributary, Rio Gavilan - Lago Maracaibo dr. 3 km E Capazon, 08 ° 49 ’ 75 ” N, 71 ° 25 ’ 61 ” W, INHS 59881. Zulia: Maracaibo Expedition, MBUCV V- 23883, 3, (92.5 - 110.5). Cano La Playa, Lago Maracaibo dr. Hacienda San Jose, MCNG 33504, 1, (85.5). Lagunas de Tule, Rio Cachiri dr. MCNG 7487, 1, (226.6). Rio Cachiri, Lago Maracaibo dr. MCNG 7419, 1, (92.6). Rio Cachiri, Lago Maracaibo dr. at Santa Marta bridge, MCNG 33526, 1, (119.8). Rio Catatumbo, Lago Maracaibo dr. at bridge on road to Machiques, MBUCV V- 18491, 1, (81.0). Rio Machango, Lago Maracaibo dr. 20 km above bridge, S Of Lagunillas, UMMZ 142488, 1, (57.4, paratype H. pospisili). Rio Negro, Rio Santa Ana - Lago Maracaibo dr. 12 km S Machiques on road to Tucuco, INHS 59945. Rio Negro, Rio Santa Ana - Lago Maracaibo dr. Bridge ca. 8 km SW Alturnitas, 09 ° 41 ’ 30 ” N, 72 ° 25 ’ 47 ” W, INHS 35436, 1, (130.7). Rio Palmar, Lago Maracaibo dr. in Hacienda el Milagro NW of the village of Rosario, MBUCV V- 18206, 2, (153.2 - 188.0). Rio Palmar, Lago Maracaibo dr. Sierra de Perija, NW Lago Maracaibo, Hacienda el Milagro, MBUCV V- 18411, 1, (168.6). Rio Santa Rosa, Rio Santa Ana - Lago Maracaibo dr. Highway 6 bridge, 09 ° 39 ’ 06 ” N, 72 ° 35 ’ 00 ” W, INHS 35466, 1, (75.9). Rio Yasa, Lago Maracaibo dr. 5 km S Machiques, INHS 60463, 1, 1 cs, (102.2) and MCNG 25011, 1, (82.0). Rio Zulia, Lago Maracaibo dr. MCNG 7486, 1, (237.9). Zulia, Tule Reservoir, Lago Maracaibo dr. lagoon, 10 ° 53 ’ N, 71 ° 12 ’ W, MCNG 746.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
4BA3665C2F9A19A50461F6627B07F4EC.taxon	diagnosis	Diagnosis: Most Hypostomus hondae can be distinguished from all other members of the H. cochliodon group by the presence of more plates in the skin between the dorsal fin and the lateral plates anterior of the dorsal-fin spine (Fig. 6 B; Table 7; some Colombian specimens do not have a large number of plates around the dorsal fin). The number of plates in this area depends on the size of the fish, some very small specimens lack plates in this region, but most small specimens have at least one or more isolated odontodes present. Hypostomus hondae is most similar to H. pagei and H. plecostomoides. Hypostomus hondae further differs from H. pagei by having spots on the caudal-fin spines (except in very melanistic specimens), by having darker coloration, a different juvenile coloration (Fig. 14 A vs. Fig. 14 B; see description of H. pagei), and by having the pectoral-fin spine reach 2 - 3 plates beyond the pelvic fin when depressed ventral to the pelvic fin (vs. 0 - 1). No additional characters can distinguish H. hondae from H. plecostomoides.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
4BA3665C2F9A19A50461F6627B07F4EC.taxon	description	Description: See Hypostomus cochliodon group description for more details. Morphometric data given in Table 2. Body dark brown with round spots present almost everywhere; most specimens with spots fading posteriorly with none present on caudal peduncle. Size of spots on body from medium to large, size increases posteriorly. Some specimens (particularly juveniles) with four dorsal saddles visible: first below anterior rays of dorsal fin, second below posterior rays of dorsal fin and slightly posterior to dorsal fin, third below adipose fin and slightly anterior to adipose fin, and fourth at base of caudal fin; dark bar also present between the eyes. Caudal fin always with spots except in strongly melanistic specimens in which caudal fin appears almost black. Caudal fin often lighter basally than distally. Abdomen slightly lighter than sides in adults; in juveniles, abdomen much lighter than sides and spots may be faint or absent. Dorsal fin usually short, when depressed in most specimens, not reaching preadipose plate. Depressed pectoral-fin spine ventral to pelvic fin reaches 2 - 3 plates beyond pelvicfin rays. Tip of pectoral-fin spine of nuptial males with stout, recurved, hypertrophied odontodes. Keels weak to moderately developed. Orbit forming ridge distinctly raised above medial surface of head; ridges of dorsal and lateral aspect of head well-developed. Longitudinal ridge on pterotic-supracleithrum beginning at posterodorsal corner of eye formed from raised bone and slightly larger odontodes absent. Opercle distinctly exposed, always supporting much more than ten odontodes. Nuptial body odontodes absent (Fig. 2 A). Plates in skin anterior to dorsal-fin spine almost always present and more numerous than in comparatively-sized specimens of other species of the H. cochliodon group (Fig. 6 B; Table 7). Most specimens less than 70 mm SL with at least one or two free odontodes in skin anterior to dorsal-fin spine; some Colombian specimens lack this characteristic. Each jaw with 8 - 22 teeth (mode = 11), teeth almost always large and spoon-shaped, some individuals with smaller, more numerous teeth. Average angle between dentaries 65 ° (SD = 8 °; range: 48 ° - 86 °; N = 27). Lateral line plates 27 - 29; dorsal plates 8 - 10; interdorsal plates 6 - 8; adipose caudal plates 8 - 10.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
4BA3665C2F9A19A50461F6627B07F4EC.taxon	distribution	Range: Lago Maracaibo drainage of Colombia and Venezuela, and the Rio Magdalena, Rio Sinu, and Rio Atrato drainages of Colombia (Fig. 11). Hypostomus hondae is allopatric to all other species of the H. cochliodon group.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
ACE51A6B7A12BFD304FF00923222C560.taxon	description	(Fig. 15)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
ACE51A6B7A12BFD304FF00923222C560.taxon	materials_examined	Material examined: BOLIVIA. La Paz: Rio Cochabamba, Rio Beni dr. Huachi, at junction of Rio Bopi and Rio Cochabamba, CAS 77349, 1, (152.9), Holotype and CAS 77350, 2, (115.0 - 150.8), Paratypes: Rio Popoi, Rio Beni dr. CAS 56747, 2, 1 cs, (77.1 - 92.2) and UMMZ 66492, 1, (57.6).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
ACE51A6B7A12BFD304FF00923222C560.taxon	diagnosis	Diagnosis: Hypostomus levis can be separated from all other members of the H. cochliodon group by the lack of an adipose fin.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
ACE51A6B7A12BFD304FF00923222C560.taxon	description	Description: See Hypostomus cochliodon group description for more details. Morphometric data given in Table 3. Available specimens probably faded considerably; light brown; pectoral and pelvic fins and occasionally head spotted; spots absent elsewhere except for some faint ones in dorsal fin. Abdomen considerably lighter than sides, without spots. Caudal fin with dark wash distally, base of caudal fin significantly lighter than distal margin. Dorsal fin short. Adipose fin and pre-adipose plate absent. Depressed pectoral-fin spine ventral to pelvic fin reaches beyond bases of pelvic-fin rays. Keels absent; lateral plates almost perfectly smooth. Orbits forming slight ridge slightly raised above medial surface of head; ridges of dorsal and lateral aspect of head fairly smooth. Longitudinal ridge on pterotic-supracleithrum beginning at posterodorsal corner of eye formed from raised bone and slightly larger odontodes absent. Opercle usually supporting no odontodes, but occasionally with one or two odontodes present. Presence of unique nuptial body odontodes (Fig. 2 B) in H. levis unknown (see Discussion below). Plates in skin anterior to dorsal-fin spine absent in specimens examined. Each jaw with 11 - 17 teeth (mode = 14), teeth relatively small, but distinctly spoonshaped (Armbruster & Page 1997). Average angle between dentaries 63 ° (SD = 9 °; range: 53 ° - 68 °; N = 3). Lateral line plates 28 - 30; dorsal plates 8; dorsal-caudal plates 16 - 18.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
ACE51A6B7A12BFD304FF00923222C560.taxon	distribution	Range: Only known from the Rios Cochabamba and Popoi of the upper Rio Beni - Upper Rio Madeira drainage of Bolivia (Fig. 8). Hypostomus levis is allopatric to all other members of the H. cochliodon group, but is potentially sympatric with H. pyrineusi.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2643A647C1ED920D0771A1654580AB39.taxon	description	(Fig. 16)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2643A647C1ED920D0771A1654580AB39.taxon	materials_examined	Material examined: AQUARIUM SPECIMEN: AUM 16090, 1 cs. COLOMBIA. Caqueta: Rio Orteguasa - Rio Amazonas dr. Florencia, Caqueta, ANSP 70518, 1, (140.5), Holotype, ANSP 70519, 2, (91.8 - 126.3), Paratypes, and USNM 100773, 1, (143.9). ECUADOR. State unknown: Quebrada, Rio Jivino - Rio Napo dr. ANSP 120350, 1, (163.6). Rio Catapino, Rio Corrientes - Rio Napo dr. MCNG 9724, 2, (145.8 - 161.3). Rio Catapino, Rio Corrientes - Rio Napo dr. Panayaco, MCNG 9725, 2, (149.1 - 190.6). Napo: Rio Aguarico, Rio Napo dr. at campamento Guarmo CEPE, 00 ° 01 ’ S, 76 ° 37 ’ 30 ” W, FMNH 106015, 1, (129.35). Rio Due, Rio Aguarico dr. at site of bridge under construction 8.8 km SW Lumbaqui by road and 0.4 km N of the road, 00 ° 00 ’ 12 ” S, 77 ° 23 ’ 48 ” W, FMNH 106014, 1, (90.8). Rio Lushanta, Rio Napo dr. 4 km E Archidona, AUM 28229, 3, 3 cs, (3). Rio Lushian, Rio Napo dr. 2 km N Archidona, AUM 28221, 1, 2 cs, (1). Rio Lushian, Rio Napo dr. 3 km N Archidona, AUM 28342, 2, (211.9 - 216.9). Rio Pano, Rio Napo dr. 5.5 km NW Tena, elev. 550 m, AUM 28216, 2, (82.9 - 100.1). Tributary, Rio Payamino dr. few km upstream from San Jose de Payamino, 00 ° 20 ’ 12 ” S, 77 ° 18 ’ W, FMNH 106016, 3, (152.2 - 185.4). PERU. Amazonas: Rio Maranon dr. Caterpiza, LACM 41873 - 5, 1, (175.7). Rio Cenepa, Rio Maranon dr. near Rio Tujushiku, LACM 39647 - 2, 2, (161.4 - 167.6). Rio Santiago, Rio Maranon dr. at La Poza, LACM 41724 - 16, 1, (112.3).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2643A647C1ED920D0771A1654580AB39.taxon	diagnosis	Diagnosis: Hypostomus oculeus can be distinguished from all other members of the H. cochliodon group except H. ericius, H. hemicochliodon, and H. sculpodon by having well developed keels. Hypostomus oculeus differs from H. hemicochliodon and H. sculpodon by a lack of a buccal papilla, by having 0 - 10 odontodes on the opercle (versus more than 10), and the presence of nuptial body odontodes (Fig. 2 B); and from H. ericius by being dark brown with closely-placed spots (vs. tan to gray with widely separated spots) and by the following ratios: anal-fin length / SL (10.8 - 13.9 % vs. 13.4 - 17.0 %) and interorbital width / head length (42.4 - 50.2 % vs. 50.5 - 56.6 %).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2643A647C1ED920D0771A1654580AB39.taxon	description	Description: See Hypostomus cochliodon group description for more details. Morphometric data given in Table 3. Body almost entirely dark brown with medium to large spots present. Spots increasing in size posteriorly and almost coalescing to form stripes on caudal peduncle. Spots on abdomen large, often confluent with one another and forming crescents and rings. Abdomen slightly lighter than sides, particularly in juveniles. Individuals below 100 mm SL often entirely dark brown except for faint spots on dorsal and paired fins and with lighter abdomen. Dorsal fin occasionally large, reaching beyond adipose fin when depressed, but dorsal fin generally reaching preadipose plate or slightly beyond when depressed. Pectoral-fin spine when depressed ventral to pelvic fin reaches beyond bases of pelvic-fin rays. Pectoral-fin spine supporting numerous stout, recurved, hypertrophied odontodes in nuptial males. Keels sharp, strongly developed. Orbits forming ridge distinctly raised above medial surface of head; ridges of dorsal and lateral aspect of head well-developed. Longitudinal ridge on pterotic-supracleithrum beginning at posterodorsal corner of eye formed from raised bone and slightly larger odontodes absent. Opercle usually not supporting odontodes, but one to ten odontodes may be present, and odontodes on opercle more numerous in juveniles. Nuptial body odontodes present (Fig. 2 B). Platelets in skin anterior to dorsalfin spine usually absent or not numerous (Fig. 6 A; Table 7). Each jaw with 7 - 18 teeth (mode = 8), teeth large and spoon-shaped. Average angle between dentaries 65 ° (SD = 10 °; range: 48 ° - 89 °; N = 23). Lateral line plates 27 - 30; dorsal plates 7 - 9; interdorsal plates 5 - 7; adipose caudal plates 9 - 11.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2643A647C1ED920D0771A1654580AB39.taxon	distribution	Range: Found in the Rio Maranon drainage of Peru, the Rio Napo drainage of Ecuador, and the Rio Orteguasa drainage of Colombia, all tributaries of the Upper Rio Amazonas drainage (Fig. 17). Hypostomus oculeus is sympatric with H. ericius, H. hemicochliodon, and H. pyrineusi.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
C32AE43BEA5CE55DC8A834818031FDA0.taxon	description	(Figs. 14 B, 18)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
C32AE43BEA5CE55DC8A834818031FDA0.taxon	materials_examined	Nontype material: VENEZUELA. Lara: Rio Curarigua, Rio Tucuyo dr. Paso San Antonio, pozo de la Iguana near Curarigua, MBUCV V- 18450, 4, (83.4 - 187.7). Rio Curarigua, Rio Tucuyo dr. Puente Torres at bridge, INHS 28867, 1, (172.5). Yaracuy: Rio Yurubi, Rio Yaracuy dr. ca. entrance to Guayabito, 10 ° 29 ’ 08 ” N, 68 ° 39 ’ 40 ” W, MCNG 27617, 1, (88.0). Tributary, Rio Yaracuy dr. E Marin, INHS 28902, 2, (84.9 - 91.8).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
C32AE43BEA5CE55DC8A834818031FDA0.taxon	diagnosis	Diagnosis: Hypostomus pagei differs from most other members of the H. cochliodon group mainly in coloration: a light, gray-brown with the spots considerably faded or absent (vs. a deep, dark brown with the spots well-developed). Juvenile H. pagei additionally have the caudal fin darkest along both spines and at the base, clear medially (Fig. 14 B), a color pattern not found in any other members of the H. cochliodon group. Hypostomus pagei is most similar to H. hondae and H. plecostomoides. Hypostomus pagei differs from H. hondae and H. plecostomoides by the general absence of spots on the caudal fin and / or caudal-fin spines (H. pagei may have some very faint spots on the caudal-fin spines; some H. plecostomoides lack spots on the caudal-fin spines, but the spines are almost black instead of light gray-brown) and by having shorter pectoral-fin spines (reaching 0 - 1 plates beyond pelvic fin when depressed below the pelvic fin vs. 2 - 3 plates with the exception of one specimen of H. pagei that has pectoral-fin spines that reach two plates beyond the pelvic fin and the one specimen of H. plecostomoides examined from the Lago Valencia drainage that has pectoral-fin spines that do not reach beyond the pelvic fin).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
C32AE43BEA5CE55DC8A834818031FDA0.taxon	description	Description: See description of the Hypostomus cochliodon group for more details. Morphometric data given in Table 4. Color gray-brown. Spots, if present, usually very faint and restricted to paired fins, dorsal fin, head, anterior part of body and / or abdomen. Spots particularly faint in adults. Caudal fin darkest distomedially. Spots absent from entire caudal fin in adults. Juveniles with faint spots on caudal-fin spines and dark brown along spines and base (Fig. 14 B). Caudal fin coloration darker than body color in juveniles, lightens ontogenetically until equal to body color. Juveniles typically slightly darker overall than adults. Dorsal fin short, falling short of preadipose plate. Depressed pectoral-fin spine ventral to pelvic fin reaches beyond bases of pelvic-fin rays, but to a lesser extent than in other species of the H. cochliodon group (0 - 1 plate vs. 2 - 3 plates in most other specimens of the H. cochliodon group). Keels weak, almost absent. Orbits forming slight ridge slightly raised above medial surface of head; ridges of dorsal and lateral aspect of head fairly smooth. Longitudinal ridge formed of raised bone and slightly larger odontodes absent on pterotic-supracleithrum beginning at postdorsal corner of orbit. Opercle usually broadly exposed, always supporting much more than ten odontodes. Plates in skin anterior to dorsal-fin spine absent or few (Fig. 6 A; Table 7). Each jaw with 7 - 17 teeth (mode = 11), teeth large and spoon-shaped. Average angle between dentaries 63 ° (SD = 5 °; range: 53 ° - 73 °; N = 14). Lateral line plates 27 - 29; dorsal plates 8 - 9; interdorsal plates 6 - 8; adipose caudal plates 9 - 10.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
C32AE43BEA5CE55DC8A834818031FDA0.taxon	distribution	Range: Found in northwestern Venezuela in the Rio Aroa, Rio Tocuyo, and Rio Yaracuy basins, short rivers that drain directly into Caribbean Sea (Fig. 11). Hypostomus pagei is allopatric to all other species of the H. cochliodon group.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
C32AE43BEA5CE55DC8A834818031FDA0.taxon	etymology	Etymology: Named for Dr. Lawrence M. Page for his help and guidance during my career, and to honor the fact that he helped to collect a majority of the specimens of this species.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
D736AC66F543B2A8FD66FD8E8B03D09A.taxon	description	(Fig. 19)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
D736AC66F543B2A8FD66FD8E8B03D09A.taxon	materials_examined	Material examined: COLOMBIA. Meta: Cano Rico, Rio Meta dr. at Brasilia, ca. 03 ° 59 ’ N, 73 ° 08 ’ W, ANSP 133235, 2, 1 cs, (83.6 - 98.8). Quebrada Venturosa, Rio Meta dr. between La Balsa and Puerto Lopez, ca. 04 ° 05 ’ N, 72 ° 58 ’ W, ANSP 133365, 1, (94.4). Rio Negrito, Rio Meta dr. downstream of bridge at La Balsa, ca. 04 ° 04 ’ N, 73 ° 04 ’ W, ANSP 131658, 1, (101.7). VENEZUELA. Anzoategui Atapirire, Rio Orinoco dr. Atapirire road, Manasma, MBUCV V- 16944, 1, (97.2). Rio Moquete, Rio Orinoco dr. Paso bajito, MBUCV V- 16922, 1, (167.0). Rio Tigre, Cano Manamo dr. near El Tigre, 08 ° 55 ’ 38 ” N, 64 ° 11 ’ 11 ” W, INHS 31412, 2, (69.4 - 105.8). Apure: cano, Rio Apure dr. 07 ° 16 ’ 26 ” N, 71 ° 05 ’ 20 ” W, INHS 28385, 2, (113.1 - 113.2). Cano Las Mercedes, Rio Apure dr. dique via Apurito, cerca San Fernando de Apure-Biruaca, 07 ° 51 ’ 30 ” N, 67 ° 22 ’ 30 ” W, MCNG 13923, 1, (173.4). Cano Maporal, Rio Apure dr. at the metal bridge near the Module (a field station of UNELLEZ), 07 ° 25 ’ 30 ” N, 69 ° 35 ’ 40 ” W, MCNG 9134, 1, (222.7). Rio Apure, Rio Orinoco dr. side channel along dike opposite municipal slaughter house, 07 ° 35 ’ N, 67 ° 29 ’ W, USNM 258264, 2, (136.1 - 193.8). Rio Aruaca, Rio Apure dr. in floodplain, paso Aruaca, 07 ° 26 ’ 55 ” N, 68 ° 26 ’ 00 ” W, MCNG 20655, 1, (142.8). Aragua: Rio Bue, Lago Valencia dr. Maracay, CAS 12694, 1, (218.8). Barinas: Cano Arenosa, Rio La Yuca - Rio Masparro dr. 14 km N Barinas, 08 ° 46 ’ 14 ” N, 70 ° 15 ’ 30 ” W, INHS 31837, 3, 1 cs, (105.6 - 119.4). Cano Capa, Rio Masparro - Rio Apure dr. 2 mi E El Tombor, 08 ° 21 ’ 35 ” N, 69 ° 46 ’ 07 ” W, INHS 35487. Cano Guanaparo, Rio Apure dr. N of Apurito, 08 ° 07 ’ 40 ” N, 68 ° 20 ’ 40 ” W, MCNG 13025, 1, (183.2). Cano Hondo, Rio Apure dr. Agua Negra, INHS 30000, 3, 1 cs, (112.7 - 218.4). Rio Chiri Chiri, Rio Apure dr. 0.5 km above mouth, 07 ° 39 ’ 40 ” N, 71 ° 29 ’ 01 ” W, MCNG 10572, 1, (169.5). Rio Mitado-seco, Rio Apure dr. USNM 194210, 1, (205.4). Rio Santa Barbara, Rio Apure dr. at bridge on Barinas to San Cristobal road, km 504, 07 ° 50 ’ 10 ” N, 71 ° 11 ’ 00 ” W, MCNG 11720, 1, (153.8). Rio Santo Domingo, Rio Apure dr. in Torunos, Hacienda La Isla, 08 ° 30 ’ 32 ” N, 70 ° 05 ’ 03 ” W, INHS 28710, 1, (87.1). Tributary of Rio Suripa - Rio Apure dr. 36 km NE La Pedrera on route 5, INHS 28284, 1, (106.4). Tributary of Rio Caipe, Rio Apure dr. 6 km NE Torunos, INHS 28744, 2 cs. Bolivar: Cano Brande, Rio Orinoco dr. Ciudad Bolivar, 08 ° 14 ’ 50 ” N, 63 ° 15 ’ 55 ” W, ANSP 166889, 1, (211.9). Cano Caiman, Rio Orinoco dr. at crossing of Caicara to Puerto Ayacucho highway 191, 2 km W of Ciudad Bolivar to Caicara highway, ANSP 162345, 1, (211.1) and MBUCV V- 16855, 1, (192.0). Cano Garrapata, Rio Orinoco dr. at bridge between Rio Paguaza and Villacoa, 06 ° 19 ’ 20 ” N, 67 ° 07 ’ 00 ” W, MCNG 11176, 1, (125.4). Cano Tabara, Rio Nichare - Rio Orinoco dr. 3 km above mouth with Rio Nichare, 06 ° 22 ’ N, 64 ° 58 ’ W, MCNG 22931, 1, (188.6). Isolated lagoon, Rio Orinoco dr. 200 yards N of Jubillal, 06 ° 57 ’ N, 64 ° 50 ’ W, ANSP 141559, 1, (131.5). Salto Icutu, Rio Orinoco dr. Cedeno district, in front of camp, 05 ° 53 ’ N, 64 ° 51 ’ W, MCNG 21160, 1, (199.8). Cojedes: Rio Camoruco, Rio Cojedes - Rio Portuguesa dr. about 10 km NW Libertad, INHS 29061, 1, (110.1). Rio Tinaco, Rio Portuguesa dr. El Baul, INHS 59831, 1, (116.4). Guarico: Rio Guarico, Rio Orinoco dr. at Flores Morada’s ranch, ca. 3 - 4 km E of road from Calabozo to San Fernando, USNM 260217, 1, (109.6). Rio Guariquito, Rio Orinoco dr. in front of Medana de Gomez, MCNG 32975, 1, (188.8). Rio San Bartolo, Rio Orinoco dr. Parque Nacional Aguaro-Guariquito, Aguas Muertes, 08 ° 04 ’ 40 ” N, 66 ° 40 ’ 00 ” W, MCNG 15027, 1, (140.5). Rio San Bartolo, Rio Orinoco dr. Parque Nacional Aguaro-Guariquito, Aguas Muertes, 08 ° 06 ’ 30 ” N, 66 ° 40 ’ 19 ” W, INHS 34862, 2, (175.7 - 187.1) and MCNG 31701, 2, (149.0 - 248.5). Rio San Bartolo, Rio Orinoco dr. Parque Nacional Aguaro-Guariquito, Aguas Muertes in the Payares, 08 ° 08 ’ 01 ” N, 66 ° 40 ’ 53 ” W, MCNG 31792, 1, (230.9). Rio San Jose, Rio Guariquito - Rio Orinoco dr. Parque Nacional Aguaro-Guariquito, 08 ° 28 ’ 55 ” N, 66 ° 53 ’ 19 ” W, INHS 34614, 1, (107.1). Miranda: Rio Cuira, Rio Tuy dr. FMNH 84616, 1, (69.7) and MBUCV V- 3756, 1, (140.1). Rio Tuy, Caribbean Sea dr. in pool isolated from main stream near Ocumaro de Tuy, MBUCV V- 3701, 10, (96.4 - 103.2). Monagas: Rio Amana, Rio Guanipa dr. 7 km NNW Santa Barbara, INHS 31263, 1, 2 cs, (139.7) and MCNG 29159, 1, (124.4). Rio Amana, Rio Guanipa dr. Las Canitas near El Tefero, MBUCV V- 6755, 1, (107.2). Rio de Oro, Rio Guarapiche - Rio San Juan dr. 4 km SW Jusepin, INHS 31448, 1 cs. Rio Guanipa, Caribbean Sea dr. Highway 5 bridge, 09 ° 22 ’ 06 ” N, 63 ° 46 ’ 47 ” W, INHS 31365, 3, 2 cs, (79.2 - 96.6). Rio Guarapiche, Rio Orinoco dr. Caicara, USNM 163168, 1, (102.9). Portuguesa: Cano El Mamon, Rio Portuguesa dr. 9 km SE of highway 5 on the road to La Quebrada, 09 ° 07 ’ 20 ” N, 69 ° 31 ’ 20 ” W, MCNG 12648, 2, (171.2 - 185.2). Cano Iguecito, Rio Portuguesa dr. on the road to Moritas, 08 ° 38 ’ 00 ” N, 69 ° 29 ’ 56 ” W, MCNG 30207, 1, (107.2). Cano Igues, Rio Portuguesa dr. 2 km from bridge at Paplon, 08 ° 38 ’ 15 ” N, 69 ° 59 ’ 45 ” W, MCNG 30163, 1, (164.8). Cano Igues, Rio Portuguesa dr. La Capilla, INHS 31966, 1, (97.4). Cano Maraca, Cano Igues - Rio Portuguesa dr. on road from Guanare to Guanarito at 60 km marker, 08 ° 49 ’ 34 ” N, 69 ° 20 ’ 45 ” W, INHS 35667, 2, (103.7 - 182.6), MCNG 15696, 1, (131.0), MCNG 8603, 2, (158.6 - 183.5), and MCNG 27275, 1, (135.7). Rio Guanare, Rio Portuguesa dr. Guanarito, INHS 31991, 1, (82.3). Rio Guanare, Rio Portuguesa dr. on S side in Finca Merecure, approximately 45 min from Guanare on the road to Las Moritas, MCNG 33951, 1, (191.66). Rio Orape, Rio Portuguesa dr. 45 km E of San Rafael de Onoto, 09 ° 43 ’ 40 ” N, 68 ° 30 ’ 10 ” W, MCNG 18716, 1, (108.0). Tachira: Rio Teteo, Rio Apure dr. on road to El Nula, 07 ° 28 ’ 40 ” N, 71 ° 55 ’ 40 ” W, MCNG 8086, 2, (122.3 - 154.0). Tributary to Rio Doradas, Rio Apure dr. La Pedrera, INHS 28153, 1, (87.2).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
D736AC66F543B2A8FD66FD8E8B03D09A.taxon	diagnosis	Diagnosis: Hypostomus plecostomoides is the most difficult species of the H. cochliodon group to identify as it is without any known synapomorphies. The most similar species are H. hondae, H. pagei, and H. taphorni. Hypostomus plecostomoides differs from H. pagei by having spots on the caudal-fin spines (except in very melanistic specimens), by having darker overall coloration, a different juvenile coloration (Fig. 14 A vs. Fig. 14 B; see description of H. pagei), and by generally having the pectoral-fin spines reaching 2 - 3 plates beyond the pelvic fin when depressed ventral to the pelvic fin (vs. 0 - 1; one specimen of H. pagei has the pectoral fin reaching two plates beyond the pelvic fin, and the single specimen of H. plecostomoides examined from the Lago Valencia drainage has the pectoral fin not extended beyond the pelvic fin); from H. hondae by having a lower number of plates in the skin around the dorsal-fin spine (Fig. 6 A; Table 7); and from H. taphorni by lacking a bicolored caudal fin and by having no or medium to large spots on the abdomen (vs. small spots; Fig. 22; see Comments). Hypostomus plecostomoides can be distinguished from H. cochliodon by having a different color pattern (lacking a dorsal stripe and generally being well-spotted), from H. ericius and H. oculeus by lacking sharp keels, and from H. ericius, H. oculeus, and H. pyrineusi by having greater than 10 odontodes on the opercle.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
D736AC66F543B2A8FD66FD8E8B03D09A.taxon	description	Description: See Hypostomus cochliodon group description for more details. Morphometric data given in Table 4. Hypostomus plecostomoides is variable in coloration. Typically, body almost entirely dark brown with medium to large spots usually present. Spots on head fairly large compared to H. hondae, H. pagei, and H. taphorni. Spots increasing in size but fading posteriorly, spots absent on caudal peduncle (Figs. 21). Spots on abdomen typically large, but some specimens with medium-sized spots, spots never as small as in H. taphorni (Fig. 22). Spots occasionally entirely absent and entire fish dark brown (almost black). Caudal fin may be entirely spotted, but caudal-fin membrane and rays typically black or black distally and light proximally. Caudal-fin spines typically spotted although some specimens with entirely black spines; caudal fin not bicolored. Specimens from particularly muddy streams gray-brown, but not to extent seen in H. pagei. Abdomen colored as sides, sometimes slightly lighter; adults almost always with welldeveloped spots on abdomen. Color in juveniles as in adults, but spots comparatively larger and abdomen generally white and unspotted. Dorsal fin short, rarely reaching preadipose plate when depressed. Depressed pectoralfin spine ventral to pelvic fin reaches 2 - 3 plates beyond pelvic-fin rays (only one specimen reached just to posterior edge of pelvic fin). Tip of pectoral-fin spine of nuptial males with stout, recurved, hypertrophied odontodes. Keels weak to moderately developed. Orbits forming moderate ridge slightly raised above medial surface of head; ridges of dorsal and lateral aspect of head fairly well-developed. Longitudinal ridge on pterotic-supracleithrum beginning at posterodorsal corner of eye formed from raised bone and slightly larger odontodes absent. Opercle always supporting much more than ten odontodes. Nuptial body odontodes absent (Fig. 2 A). Plates in skin anterior to dorsal-fin spine absent or few (Fig. 6 A; Table 7). Each jaw with 5 - 16 teeth (mode = 9), teeth large and spoon-shaped. Average angle between dentaries 57 ° (SD = 10 °; range: 41 ° - 82 °; N = 30). Lateral line plates 27 - 30; dorsal plates 7 - 9; interdorsal plates 6 - 8; adipose caudal plates 8 - 11.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
D736AC66F543B2A8FD66FD8E8B03D09A.taxon	distribution	Range: Found in the Rio Orinoco basin of Colombia and Venezuela, and the Rio Tuy and Lago Valencia drainages of Venezuela (Fig. 11). Hypostomus plecostomoides is allopatric to all other species of the H. cochliodon group (see Comments).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
D736AC66F543B2A8FD66FD8E8B03D09A.taxon	discussion	Comments: Specimens most appropriately identified as Hypostomus plecostomoides from the Orinoco drainage in northwestern Bolivar state, Venezuela (ANSP 162345, MBUCV V- 16855, MCNG 11176, and MCNG 22931), may provide evidence for intergradation between H. plecostomoides and H. taphorni. Some specimens from this region have the bicolored caudal fin diagnostic for H. taphorni, and few or no odontodes on the opercle and relatively smaller spots on the head than typical in H. plecostomoides. However, the specimens lack the second synapomorphy for H. taphorni, small spots on the abdomen. Alternatively, it is possible that these specimens represent an undescribed species. Additional specimens will have to be examined before the status of the members of the H. cochliodon group from northern Bolivar state can be assessed. For the purposes of this study, specimens from northern Bolivar were excluded from the description above and from Table 4.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2D65F4A558519FDC5229B409E993BC66.taxon	description	(Fig. 20)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2D65F4A558519FDC5229B409E993BC66.taxon	materials_examined	Material examined: BOLIVIA. Beni:, Rio Curiraba, Rio Madeira dr. Campamento Trapiche, USNM 305281, 1, (119.8). Rio Itenez, Rio Beni - Rio Madeira dr. 5 km S Costa Marques, Brasil, 1 km above mouth of Rio Baures, AMNH 39941, 1, (210.1). Rio Itenez, Rio Beni dr. 9 km SE of Costa Marques, Brasil, UMMZ 204399, 5, (139.9 - 226.7). Rio Itenez, Rio Beni - Rio Madeira dr. opposite Costa Marques, Brasil, AMNH 39758, 2, (92.0 - 213.4). Rio Itenez, Rio Beni - Rio Madeira dr. playa pond, 9 km SE Costa Marques, Brasil, AMNH 40108 (none measured). Rio Maniqui, Rio Marmore - Rio Madeira dr. San Borja, INHS 36995, 2, (162.7 - 176.0). BRAZIL. Acre: Rio Macauhan, Rio Yaco - Rio Purus dr. vicinity of Rio Macauhan, USNM 94653, 1, (122.5). Rio Macaua, Rio Iaco dr. near Sena Madueira, 09 ° 20 ’ S, 68 ° 45 ’ W, AMNH 12599, 1, (113.5). Amazonas: Rio Jamari, MNRJ 863, 1, (204.0), Holotype: ECUADOR. Napo: Rio Catapino, Rio Corrientes - Rio Napo dr. MCNG 9724, 1, (174.8). Rio Pucuno, Rio Suno - Rio Napo dr. 00 ° 47 ’ S, 77 ° 16 ’ W, USNM 177222, 1, (71.3). Rio Lushian, Rio Misahuali dr. at bridge E of Rucullacta, 00 ° 54 ’ 42 ” S, 77 ° 48 ’ 48 ” W, CAS 106013. Shansho Cano, USNM 124875, 1, (100.5). PERU. Amazonas: Rio Maranon dr. 1 km upstream from Caterpiza, LACM 42005 - 13, 2, (84.1 - 89.5) and LACM 42003 - 8, 2, (150.6 - 174.3). Rio Maranon dr. 100 m downstream from Caterpiza, LACM 41996 - 9, 1, (137.1) and LACM 41972 - 3, 1, (140.9). Rio Maranon dr. 300 m upstream from Caterpiza, LACM 41813 - 1, 1, (207.7). Rio Maranon dr. 500 m upstream from Caterpiza, LACM 41992 - 7, 4, 1 cs, (162.4 - 190.1) and LACM 42010 - 8, 1, (108.0). Rio Maranon dr. 800 m upstream from Caterpiza, LACM 41834 - 1, 1, (131.2). Rio Maranon dr. Caterpiza, LACM 41873 - 7, 1, (180.7), LACM 41882 - 4, 1, (180.9), LACM 41884 - 1, 1, (198.9), LACM 41887 - 5, 1, (160.3), LACM 42009 - 10, 1, (133.9 - 164.9), LACM 41898 - 2, 1, (140.0), LACM 41893 - 9, 2, 1 cs, (152.8 - 163.2), LACM 42115 - 8, 1, (104.7), LACM 41895 - 6, 1, (152.8 - 163.2), and LACM 42116 - 4, 1, (93.0). Quebrada Kayamasa, Rio Maranon dr. 40 km upstream from Caterpiza, FMNH 97016, 1, (140.9), LACM 41872 - 4, 2, (150.9 - 187.7), LACM 41846 - 4, 1, (199.7), LACM 41852 - 1, 1, (173.6), and LACM 41853 - 4, 2, (136.7 - 156.4). Quebrada Pastazilla, Rio Santiago - Rio Maranon dr. LACM 39947 - 8, 2, (208.5 - 215.1). Quebrada Yutupiza, Rio Maranon dr. LACM 36337 - 10, 1, (159.4). Rio Cenepa, Rio Maranon dr. near San Antonio, LACM 39638 - 3, 3, 1 cs, (94.2 - 202.8). Rio Huampani, Rio Maranon dr. Maja Quebrada, LACM 36359 - 2, 1, (228.0). Rio Maranon, Rio Amazonas dr. at and across from Sta. Maria de Nieva and confluence of Rio Nieva, FMNH 97017, 2, (110.6 - 131.9). Rio Santiago, Rio Maranon dr. at La Poza, LACM 41724 - 17, 1, (131.5), LACM 41723 - 10, 1, (130.5), LACM 41729 - 39, 1, (147.7), LACM 41730 - 11, 1, (128.8), LACM 39883 - 11, 3, 1 cs, (137.9 - 170.8). Huanaco:, Rio Huallaga, Rio Ucayali dr. at Tingo Maria, CAS 56754, 2, 1 cs, (79.6 - 86.1) and CAS 95117, 2, 1 cs, (69.7 - 160.5). Rio Rondos, Rio Monzon dr. vicinity of Tingo Maria, just above new bridge site, ANSP 38873 (none measured). Loreto: Rio Amazonas, Atlantic Ocean dr. INHS 39358 (none measured). Rio Amazonas, Atlantic Ocean dr. 21.1 mi NE Iquitos bearing 22 °, 03 ° 30 ’ 30 ” S, 73 ° 03 ’ 28 ” W, INHS 39122 (65.2). Rio Amazonas, Atlantic Ocean dr. at Pueblo Gallito, INHS 36939, 1, (80.3). Rio Itaya, Rio Amazonas dr. approx. 10 km S Santa Clara, INHS 36877, 1, (94.8). Rio Nanay, Rio Amazonas dr. well above Morona cocha, ANSP 38874 (none measured). Rio Napo, Rio Amazonas dr. at Mazan, INHS 36565 (none measured). Madre de Dios:, Cano Pachija, Rio Madre de Dios dr. Manu National Park, Pakitza, from mouth to 7 km upstream, USNM 319601, (81.2). Cano Picaflor, Rio Madre de Dios dr. Pakitza, Cana Brava Trail # 19, USNM 319353, 1, (114.1). Tributary, Rio Tambopata - Rio Madre de Dios dr. stream enters Rio Tambopata from S bank approximately 500 m downstream of Explorer’s Inn boat landing, 12 ° 49 ’ 35 ” S, 69 ° 17 ’ 30 ” W, USNM 264049, 1, (178.5). Tributary to Rio Madre de Dios, on S side about 10 km downstream of junction of Rio Tambopata and Rio Madre de Dios, 12 ° 30 ’ S, 69 ° 10 ’ W, USNM 263911, 1, (130.1). Tributary, Rio Tambopata - Rio Madre de Dios dr. second stream entering Rio Tambopata on SW shore upstream of mouth of Rio La Torre, 12 ° 49 ’ 40 ” S, 69 ° 18 ’ W, USNM 264050, 2, (143.0 - 202.6).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2D65F4A558519FDC5229B409E993BC66.taxon	diagnosis	Diagnosis: Hypostomus pyrineusi is distinguished from most members of the H. cochliodon group except some H. cochliodon, H. levis, H. ericius, H. oculeus, and H. taphorni by having 0 - 10 (usually 0; Fig. 2 B) odontodes on the opercle (vs. 11 or more). Hypostomus pyrineusi further differs from H. hemicochliodon and H. sculpodon by lacking a ridge on the pterotic-supracleithrum (Fig. 5 B), by the presence of large, spoonshaped teeth (vs. teeth intermediate; Fig. 1 C vs. B), and a lack of a buccal papilla; from H. ericius, H. hemicochliodon, H. oculeus, and H. sculpodon by lacking strong, sharp keels on the lateral plates; from H. levis by having an adipose fin; from H. cochliodon by having the spots more numerous and close-set on the body (vs. spots absent or widely spaced); and from H. taphorni by having pigmentation on the lobes of the caudal fin equally distributed (vs. lower lobe darker than upper).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2D65F4A558519FDC5229B409E993BC66.taxon	description	Description: See Hypostomus cochliodon group description for more details. Morphometric data given in Table 5. Body typically almost entirely dark brown with medium to large spots present. Spots on head fairly large when compared to those of H. hondae, H. pagei, and H. taphorni. Spots increasing in size and fading posteriorly, absent on caudal peduncle. Spots on abdomen typically large, some specimens with medium-sized spots, spots never as small as in H. taphorni; (Fig. 22). Spots occasionally absent and entire fish dark brown (almost black). Caudal fin may be entirely spotted, but caudal-fin membranes and rays typically black or black distally and light proximally; caudal-fin spines typically spotted although some specimens with entirely black caudal-fin spines; caudal fin not bicolored. Abdomen colored as sides, sometimes slightly lighter. Color in juveniles as in adults, but spots comparatively larger and abdomen generally white and unspotted. Dorsal fin short, rarely reaching preadipose plate when depressed. Depressed pectoralfin spine ventral to pelvic fin reaches 2 - 3 plates beyond pelvic-fin rays. Tip of pectoral-fin spine of nuptial males with stout, recurved, hypertrophied odontodes. Keels weak to moderately developed. Orbits forming moderate ridge slightly raised above medial surface of head; ridges of dorsal and lateral aspect of head fairly well-developed. Longitudinal ridge on pterotic-supracleithrum beginning at posterodorsal corner of eye formed from raised bone and slightly larger odontodes absent. Opercle usually supporting no odontodes, but occasionally one to three odontodes are present. Nuptial body odontodes present (Fig. 2 B). Plates in skin anterior to dorsal-fin spine absent or few (Fig. 6 A; Table 7). Each jaw with 5 - 14 teeth (mode = 8), teeth large and spoon-shaped. Average angle between dentaries 55 ° (SD = 7 °; range: 43 ° - 69 °, N = 30). Lateral line plates 26 - 31; dorsal plates 8 - 9; interdorsal plates 4 - 7; adipose caudal plates 9 - 16.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
2D65F4A558519FDC5229B409E993BC66.taxon	distribution	Range: Found in the Rio Madeira drainage of Bolivia and Brazil, the Rios Napo, Ucayali, Maranon and upper Rio Amazonas drainage of Peru, and the Rio Napo drainage of Ecuador, (Fig. 8). Hypostomus pyrineusi is sympatric with H. ericius, H. hemicochliodon, and H. oculeus and is potentially sympatric with H. levis.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
3CACDEC0C7C4A2615B6E10DE98C72185.taxon	description	(Fig. 21)	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
3CACDEC0C7C4A2615B6E10DE98C72185.taxon	diagnosis	Diagnosis: Hypostomus sculpodon differs from all other members of the H. cochliodon group except H. hemicochliodon in the presence of a buccal papilla and intermediately developed teeth (Fig. 1 B; juvenile H. cochliodon and H. taphorni occasionally also have intermediate teeth). Hypostomus sculpodon differs from H. hemicochliodon in coloration (brownish-red with widely spaced spots vs. dark brown with closely spaced spots), by having a smaller dorsal-anal length to anal-fin length ratio (average = 79.2 ± 7.6 %, 69.6 - 89.7 % vs. average = 111.1 ± 8.4 %, 92.5 - 132.6 %), and almost completely by having a smaller interorbital width / HL ratio (average = 51.4 ± 3.1 %, 38.5 - 45.4 % vs. average = 49.9 ± 2.7 %, 44.1 - 55.6 %).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
3CACDEC0C7C4A2615B6E10DE98C72185.taxon	description	Description: See description of the Hypostomus cochliodon group for more details. Morphometric data given in Table 5. Color red-brown. Body and fins well spotted, spots widely spaced. Spots on head and anterior portion of body (to about anterior third of dorsal fin) medium-sized, then shifting abruptly to large spots on posterior part of body. Spots on posterior part of body not progressively increasing in size, generally longitudinally ovoid. Abdomen significantly lighter than sides with spots large. Juveniles colored similarly to adults, but all spots of about the same size (except small spots at anterior margin of head), and abdomen much lighter with spots faint or absent. Dorsal fin fairly large, usually reaching preadipose plate when depressed. Pectoral-fin spine, when depressed ventral to pelvic fin, reaching 2 - 3 plates beyond pelvic-fin rays. Tip of pectoral-fin spine of nuptial males with stout, recurved, hypertrophied odontodes. Keels strong with sharp odontodes. Orbits forming ridge distinctly raised above medial surface of head; ridges of dorsal and lateral aspect of head well-developed, particularly supraoccipital crest. Longitudinal ridge formed of raised bone and slightly larger odontodes present and very well developed on pterotic-supracleithrum beginning at postdorsal corner of orbit. Opercle broadly exposed, always supporting much more than 10 odontodes. Nuptial body odontodes absent (Fig. 2 A). Plates in skin anterior to dorsal-fin spine absent or few (Fig. 6 A; Table 7). Head and body not particularly deep, body more sleek than other members of the H. cochliodon group. Each jaw with 12 - 19 teeth (mode = 14), teeth intermediate in size (Fig. 1 B). Average angle between dentaries 67 ° (SD = 11 °; range: 56 ° - 83 °; N = 5). Lateral line plates 30; dorsal plates 9 - 10; interdorsal plates 7 - 8; adipose caudal plates 9.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
3CACDEC0C7C4A2615B6E10DE98C72185.taxon	distribution	Range: Found in the upper Rio Negro and upper Rio Orinoco basins of Venezuela (Fig. 11). Hypostomus sculpodon is sympatric with H. hemicochliodon.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
3CACDEC0C7C4A2615B6E10DE98C72185.taxon	etymology	Etymology: From the Latin, sculpo, meaning to carve, cut, grave, or chisel in stone, brass, or wood and from the Greek, odon, for tooth in reference to the ability of the species to chisel wood with the teeth.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
9D54D7B8BEEB89222FFD7D8564D412E7.taxon	description	Fig. 22	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
9D54D7B8BEEB89222FFD7D8564D412E7.taxon	materials_examined	Material examined: VENEZUELA. Bolivar: Rio Botanmo, Rio Cuyuni dr. on the road to Bochinche, 07 ° 25 ’ N, 61 ° 11 ’ W, MCNG 8048, 1, (184.8, holotype), MCNG 8049, 1, (76.2, paratype). Cano Negro, Rio Cuyuni dr. cano a little above Isla de Anacoco, 06 ° 47 ’ N, 61 ° 12 ’ W, MCNG 1452, 1, (181.7, paratype). Cano, Rio Cuyuni dr. Highway 1, 20 river km above km 88 of El Dorado - Santa Elena road, MBUCV V- 16491, 1, (151.6). Cano Negro, Rio Cuyuni dr. primary tributary of the Rio Cuyuni above Isla Anacoco, 06 ° 44 ’ N, 61 ° 09 ’ W, MCNG 928. Cano Quebrada Clara, Rio Cuyuni dr. at the moth of Quebrada Clara at Anacoco, cerca the confluence with the Rio Cuyuni, approx. 1 km up, 06 ° 47 ’ N, 61 ° 10 ’ W, MCNG 16901, 1, (149.1). Cano Quebrada Clara, Rio Cuyuni dr. below bridge cerca isla Anacoco, 06 ° 42 ’ N, 61 ° 09 ’ W, MCNG 16450, 1, (83.5). Rio Corumo, Rio Botanmo - Rio Cuyuni dr. 10 km E Tumeremo on road from Tumeremo to Bochinche (just after Fort Terembay), 07 ° 25 ’ 00 ” N, 61 ° 28 ’ 00 ” W, ANSP 168195, 7, 2 cs, (120.4 - 158.7), MBUCV V- 26824, 9, 2 cs, (115.9 - 197.1), INHS 31768, MCNG 29566, 1, (72.3), and MCNG 16038, 1, (80.8). Rio Cuyuni, Rio Essequibo dr. 10 km W km 88 of El Dorado - Santa Elena road, MBUCV V- 16601, 1, (130.4). Rio Cuyuni, Rio Essequibo dr. cerca de la Chalana de Anacoco, 06 ° 43 ’ N, 61 ° 09 ’ W, MCNG 11066, 1, (228.0). Rio Cuyuni, Rio Essequibo dr. near Isla Anacoco, MCNG 7535, 1, (219.2). Rio Yuruari, Rio Cuyuni dr. just E El Manteco, INHS 31600, 1, (87.2).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
9D54D7B8BEEB89222FFD7D8564D412E7.taxon	diagnosis	Diagnosis: Hypostomus taphorni can be distinguished from all other members of the H. cochliodon group by the combination of having a bicolored caudal fin (light above, dark below) and small spots on the abdomen. Some specimens tentatively identified as H. plecostomoides from northern Bolivar State, Venezuela, have a bicolored caudal fin, but have medium to large spots on the abdomen (See Comments in H. plecostomoides).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
9D54D7B8BEEB89222FFD7D8564D412E7.taxon	description	Description: See Hypostomus cochliodon group description for more details. Morphometric data given in Table 6. Body typically almost entirely dark brown with medium to large spots present. Spots on head fairly small when compared to H. plecostomoides, fading posteriorly and absent on caudal peduncle. Spots on abdomen very small (Fig. 22). Caudal fin bicolored, upper lobe tan and without spots, lower lobe black. Color in juveniles as in adults, but spots comparatively larger with spots on abdomen very light or absent. Dorsal fin short, rarely reaching preadipose plate when depressed. Depressed pectoralfin spine ventral to pelvic fin reaches 2 - 3 plates beyond pelvic-fin rays. Tip of pectoral-fin spine of nuptial males with stout, recurved, hypertrophied odontodes. Keels weak to moderately developed. Orbits forming ridge moderately raised above medial surface of head; ridges of dorsal and lateral aspect of head fairly well-developed. Longitudinal ridge on pterotic-supracleithrum beginning at posterodorsal corner of eye formed from raised bone and slightly larger odontodes absent. Opercle supporting up to 10 odontodes in juveniles; number of odontodes on the opercle decreases with age and many adults have no odontodes on the opercle. Nuptial body odontodes present (Fig. 2 B). Plates in skin anterior to dorsal-fin spine absent or few (Fig. 6 A; Table 7). Each jaw with 6 - 12 teeth (mode = 10), teeth large and spoon-shaped. Average angle between dentaries 56 ° (SD = 7 °; range: 44 ° - 66 °; N = 13). Lateral line plates 27 - 29; dorsal plates 8 - 9; interdorsal plates 6 - 8; adipose caudal plates 8 - 10.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
9D54D7B8BEEB89222FFD7D8564D412E7.taxon	distribution	Range: From the Rio Cuyuni drainage of Venezuela (Fig. 11). Recent collecting in Guyana suggests that H. taphorni is limited to the Cuyuni drainage and replaced by two undescribed species in the rest of the Essequibo river drainage. Hypostomus taphorni is allopatric to all other members of the H. cochliodon group (see Comments on page 43).	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
4CDB33DCE60FBFDF63B6C0B704425711.taxon	discussion	Discussion Delimiting the species of the Hypostomus cochliodon group is made difficult by the fact that the species vary little in morphometrics or meristics, and the species have a well developed ability to alter color pattern. However, the species can be divided into four groups and from those groups it is possible to distinguish the species. The four groups are the intermediate group (H. hemicochliodon and H. sculpodon), H. cochliodon with a unique color pattern, the odontodeless opercle group (H. ericius, H. levis, H. oculeus, H. pyrineusi, and H. taphorni), and an undifferentiated northern group (H. hondae, H. pagei, and H. plecostomoides). The odontodeless opercle group can be further subdivided into a highly keeled group (H. ericius and H. oculeus) and a weakly to non-keeled group (H. levis, H. pyrineusi, and H. taphorni). Once in these groups, it is fairly simple to separate the species. Principal components analysis is provided for each of the groups although H. levis is excluded from the analysis because it lacks many of the measurements because of its absence of an adipose fin. The PCA of the intermediate group showed a 100 % separation of the species on principal component 2 (Fig. 10 A). PC 2 is most strongly and negatively effected by dorsal-anal length and anal-fin width and most strongly and positively effected by anal-fin length and folded dorsal-fin length. A ratio of dorsal-anal length to anal-fin length completely separates the two species from one another: average = 111.1 ± 8.4 %, 92.5 - 132.6 % in Hypostomus hemicochliodon and average = 79.2 ± 7.6 %, 69.6 - 89.7 % in H. sculpodon). The PCA of the odontodeless groups showed two trends, a complete or almost complete separation of H. taphorni from H. ericius, H. oculeus, and H. pyrineusi and a complete separation of H. ericius and H. oculeus on a graph of PC 2 vs. PC 3 (Fig. 10 B). PC 2 is most strongly and negatively effected by adipose-caudal length and mouth width and most strongly and positively effected by interdorsal length, eye-nare length and orbit length. PC 3 is most strongly and negatively effected by base of anal-fin length, anal-fin length, and adipose-spine length and most strongly and positively effected by interdorsal length, adipose-caudal length, and postanal length. Although there is no morphometric evidence suggesting that H. ericius and H. oculeus are distinct from H. pyrineusi there are morphological differences (well-developed keels in H. ericius and H. oculeus vs. no or weak keels in H. pyrineusi). The PCA of the northern group showed some general trends separating H. hondae and H. pagei from H. plecostomoides and a complete separation of H. hondae from H. pagei on PC 2 vs. PC 3 (Fig. 10 C). PC 2 is most strongly and negatively effected by eye-nare length and interorbital width and most strongly and positively effected by anal-fin length and interdorsal length. PC 3 is most strongly and negatively effected by anal-fin length and folded dorsal-fin length and most strongly and positively effected by adipose-caudal length and interdorsal length. For the most part, the species of the Hypostomus cochliodon group are allopatric (Figs. 8, 11, 17). It is only in the Amazon basin that several species coexist. Panaque, the other wood-eating genus of loricariid catfishes, also reaches its peak of diversity in the Amazon basin (Schaefer & Stewart 1993; Schaefer & Stewart 2002). In the upper Rio Maranon drainage of Peru, there are at least nine species of wood-eaters: H. ericius, H. hemicochliodon, H. oculeus, H. pyrineusi, P. albopunctatus, P. dentex, P. gnomus, P. nocturnus, and probably at least one species of the P. nigrolineatus clade (distributions based on this study and Schaefer & Stewart 1993). How these species subdivide the wood-eating niche is unknown, and ecological studies on wood-eating by fishes in the upper Rio Maranon would represent a very interesting study. Weber and Montoya-Burgos (2002) recently described Hypostomus fonchii and suggested that the species was derived from the H. cochliodon group. The teeth described for H. fonchii are elongate and unicuspid and it is suggested that the unicuspid teeth represent a synapomorphy for H. fonchii and the H. cochliodon group. This is problematic in that I have not observed truly unicuspid teeth in any members of the H. cochliodon group. In all specimens of the H. cochliodon group that I have examined, the medial cusp remains present, but is generally fused with the lateral cusp. The mesial cusp remains visible as a slightly darker, thicker ridge on the medial side of the tooth. I have not examined any specimens of H. fonchii, and I cannot determine whether or not this is the case with this species. The only truly unicuspid teeth that have been reported for the Hypostominae are the teeth found in nuptial male Aphanotorulus (Armbruster & Page 1996). I do not consider H. fonchii as a member of the H. cochliodon group as its body shape is that of a generalized, fast-water dwelling Hypostomus and not the derived shape of H. cochliodon. Exact placement of H. fonchii awaits an analysis of its diet, osteology, and external anatomy for the synapomorphies of the H. cochliodon group. Recent expeditions into poorly collected regions where no members of the Hypostomus cochliodon group have been reported from in the past are finding more species of the H. cochliodon group. Two undescribed species are now known from the Essequibo and Takutu River drainages of Guyana (pers. obs.) and one species from the Rio Tocantins in Brazil (Reis, pers. comm.). These species and a revised key will be presented in future publications. Phylogeny Character 1: Teeth - 0: elongate (Fig. 1 A); 1: intermediate throughout life (Fig. 1 B); 2: spoon-shaped at least in adults (Fig. 1 C). Character 2: Maxilla - 0: straight to moderately curved (Fig. 3 A); 1: greatly curved, almost forming right angle (Fig. 3 B). Character 3: Odontodes on opercle - 0: 11 + (Fig. 2 A); 1: 0 - 10 (Fig. 2 B). Character 4: Longitudinal ridge formed from bone and slightly enlarged odontodes on pterotic-supracleithrum - 0: present (Fig. 5 A); 1: absent (Fig. 5 B). Character 5: Nuptial body odontodes - 0: absent (Fig. 2 A); 1: present (Fig. 2 B). Character 6: Notch between the hyomandibula and metapterygoid - 0: present (Fig. 3 A); 1: absent (Fig. 3 B). Character 7: Buccal papilla - 0: present (Fig. 4 A); 1: absent or extremely small (Fig. 4 B). Character 8: Dentary angle - 0: averaging greater than 90 °; 1: averaging less than 80 °. Character 9: Sharp keel odontodes - 0: present; 1: absent. Character 10: Body shape deep and narrow with the head taller than wide - 0: absent; 1: present. Very few morphological characters are useful in elucidating the relationships of the species of the Hypostomus cochliodon group. Based on the ten characters found, a phylogeny is produced for the species of the H. cochliodon group (Fig. 23). The single tree found has 12 steps and CI = 1.00. Character state data is in Table 8. No skeletal material is available for H. sculpodon or H. ericius; however, external evidence does provide information as to the potential relationships of these species with other members of the H. cochliodon group. Considering there are so few characters available for analysis, the phylogeny should be considered tentative. Also providing support for the basal position of Hypostomus hemicochliodon and H. sculpodon is diet. In all of the specimens of the H. cochliodon group examined, almost the entire intestine is filled with small wood chips suggesting that the fishes consume little other than wood. In H. hemicochliodon and H. sculpodon the intestine mostly contains wood, but other organic matter makes up a major portion of the diet (not greater than 50 %). It is apparent that H. hemicochliodon and H. sculpodon are wood eaters but not wood specialists.	en	Jonathan W. Armbruster (2003): The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae). Zootaxa 249: 1-60, URL: http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDFAA9D6-E4FA-4C3C-9749-CF0313D30F3C
