Creagrutus anary Fowler 1913

Creagrutus anary Fowler, 1913

Creagrutus anary Fowler. 1913:552, fig. 16 [type locality: Brazil, Madeira River (=Rio Madeira), approximately 200 miles (=320 km) E of longitude 62°20′W; see also under “Remarks,” below, concerning reported locality]; 1948:82, fig. 87 [literature compilation]; 1975:25 [literature compilation].—Eigenmann, 1927:423 [Madeira River, based on Fowler, 1913].—Géry, 1964:62 [in key]; 1977:407 [Rio Madeira, C. hildebrandi Schultz cited as possible junior synonym].—Böhlke, 1984:42 [holotype reported missing, paratype cited as cleared and stained].—Cala, 1990:92 [as possible synonym of C. hildebrandi following Géry, 1977].—Chang and Ortega, 1995:2 [Peru, Department of Madre de Dios].—Chang, 1998:22 [southeasternPeru].

DIAGNOSIS.—The combination of the possession of premaxillary dentition arranged in the three ecomponents generalized for most species of Creagrutus without a pronounced gap between the first and second teeth of the primary tooth series of the premaxilla, 6 teeth in the primary series of each premaxilla, 2 to 4 maxillary teeth, 4 or 5 teeth on each dentary, 8 or 9 median predorsal scales, 40 to 43 lateral line scales without a lamellar process over each pore, 4 scale rows between the dorsal-fin origin and the lateral line, 10 to 12 branched anal-fin rays, 7 or 8 gill rakers on the upper limb of the first gill arch, the distance from the snout to the pectoral-fin insertion (21.0%–23.8% of SL), the snout length (24.9%–26.4% of HL), the bony orbital diameter (33.9%–37.4% of HL), the interorbital width (30.7%–33.0% of HL), the lack of contact between the ventral margin of the third infraorbital and the horizontal limb of the preopercle, the possession of a distinct spot of dark pigmentation at the base of the middle caudal-fin rays, the moderately vertically elongate humeral mark without a secondary, dorsally situated patch of pigmentation, the absence of a distinct patch of pigmentation on the dorsal fin, and the lack of a series of dark spots along the midlateral surface of the body distinguishes Creagrutus anary within the clade formed by Creagrutus and Piabina.

Characters A B

Morphometrics

Standard length ∼25.0 25.0–44.5

1. Snout to anal-fin origin – 60.7–64.5

2. Snout to pelvic-fin insertion – 44.3–47.5

3. Snout to pectoral-fin insertion – 21.0–23.8

4. Snout to dorsal-fin origin – 44.0–48.3

5. Dorsal-fin origin to hypural joint – 54.0–58.3

6. Dorsal-fin origin to anal-fin origin – 28.5–32.3

7. Dorsal-fin origin to pelvic-fin insertion – 24.1–27.5

8. Dorsal-fin origin to pectoral-fin insertion – 30.5–32.6

9. Caudal peduncle depth – 9.7–11.4

10. Pectoral-fin length – 18.9–19.4

11. Pelvic-fin length – 14.8–16.2

12. Dorsal-fin length – 21.8–23.5

13. Anal-fin length – 18.4–19.5

14. Head length – 23.5–26.1

15. Postorbital head length – 39.4–42.5

16. Snout length – 24.9–26.4

17. Bony orbital diameter – 33.9–374

18. Interorbital width – 30.7–33.0

Meristics

Lateral line scales –1 40–43

Scale rows between dorsal-fin origin and lateral line –2 4

Scale rows between anal-fin origin and lateral line –3 3

Predorsal median scales –4 8–9

Branched dorsal-fin rays 85 8

Branched anal-fin rays 126 10–12

Branched pelvic-fin rays 77 6–7

Pectoral-fin rays 138 12–14

Vertebrae 39 38–40

1Reported by Fowler (1913) as 40 in paratype and 42 in holotype.

2Reported by Fowler (1913) as 5 in both paratype and holotype

3Reported by Fowler (1913) as 3 in paratype and 4 in holotype.

4Reported by Fowler (1913) as 9 in paratype and 8 in holotype.

5Same value reported by Fowler (1913) for holotype.

6Reported by Fowler (1913) as 11 in both paratype and holotype.

7Same value reported by Fowler (1913) for holotype.

8Same value reported by Fowler (1913) for holotype.

DESCRIPTION.—Morphometric and meristic data for Creagrutus anary in Table 5. Head and body moderately robust, body more so in larger females. Greatest body depth at vertical through dorsal-fin origin in most individuals, shifted distinctly anteriorly in specimens with distended abdomens. Dorsal profile of head convex from margin of upper lip to vertical through anterior margin of orbit, nearly straight from that point to tip of supraoccipital spine. Interorbital region distinctly convex transversely. Predorsal profile slightly convex, without obvious change in alignment relative to that of head. Predorsal region of body with obtuse median ridge proximate to dorsal-fin origin. Ventral profile of head slightly convex anteriorly, with barely obvious obtuse angle at anteroventral corner of dentary (angle in ventral profile of dentary not apparent in fig. 16 of Fowler, 1913); head profile nearly straight from that point to isthmus. Profile of prepelvic region of body variably convex, more so in larger specimens. Prepelvic region of body distinctly flattened transversely.

Head obtusely pointed in lateral view, moderately compressed in dorsal view. Upper jaw slightly longer than, and overhanging, lower jaw. Anteromedial portion of snout fleshy with scattered papillae. Scattered papillae distributed over lateral surface of jaw with papillae more concentrated along ventral margin of jaw anteriorly and on plicae and folds extending between outer and medial premaxillary teeth. Lower lip fleshy, with papillae concentrated along dorsal margin and scattered papillae anteromedially.

Infraorbital series relatively well developed. Ventral margin of third infraorbital contacting or nearly contacting horizontal limb of preopercle. Posterior margins of third and fourth infraorbitals falling slightly short of vertical limb of preopercle; posterior margin of fifth infraorbital in, or nearly in, contact with vertical limb of preopercle.

Premaxillary dentition in three series: primary row slightly sigmoid, with 6 teeth, without pronounced gap between first and second tooth of series and with medial teeth of contralateral series distinctly separated; triangular cluster of 3 larger teeth with medial teeth of contralateral clusters nearly in contact; and single tooth of form similar to that of primary series occurring lateral to or slightly anterolateral to fourth tooth of premaxillary row. Maxilla with 2 to 4 tricuspidate teeth. Dentary with 4 or 5 teeth; 3 anterior teeth distinctly larger and tricuspidate with middle cusp much larger; second tooth about twice as high and wide as first tooth and three times as high and four times as wide as third tooth; fourth and fifth teeth (when 5 teeth present) graded in size, fourth always tricuspidate, fifth tricuspidate or with small posterior cusp missing.

Dorsal-fin rays ii,8 in all examined specimens. Dorsal-fin origin at, or approximately at, vertical through pelvic-fin insertion. Profile of distal margin of dorsal fin slightly concave. Anal-fin rays ii or iii,10–12. Anal-fin hooks present on first branched anal-fin ray in mature males. Pectoral-fin rays i,11–13. Tip of pectoral fin extending approximately two-thirds to three-fourths of distance to pelvic-fin insertion. Pelvic-fin rays i,6,i or i,7. Tip of pelvic fin extending to anus or to anal-fin origin.

Gill rakers 7–8+10–11.

COLORATION IN ALCOHOL.—Ground coloration of specimens yellow-tan. Dorsal surface of the head with few surface dark chromatophores scattered over anterodorsal region of head and over snout, otherwise without surface pigmentation. Series of very dark, deep-lying stellate chromatophores overlying dorsal portion of brain. Anterior portion of brain overlain by contralateral, horizontally elongate patches of dark chromatophores. Region anterior to nostrils with crescent-shaped patch of dark chromatophores. Anteroventral region of orbit outlined by variably obvious narrow series of dark chromatophores. This series not continuous with chromatophore field anterior to nostrils, nor continuing around ventral and dorsal margins of orbits as in many congeners. Sixth infraorbital and proximate portion of opercle with few scattered dark chromatophores.

Scales of anterodorsal portion of body outlined by single series of dark chromatophores. Middorsal region with field of scattered chromatophores. Obscure midlateral stripe formed of scattered dark chromatophores extending from approximately vertical through dorsal-fin origin to caudal peduncle. Humeral mark moderately obvious and vertically elongate, with variable anterior and posterior margins. Ventral margin of humeral mark extending approximately to lateral line; dorsal margin less discrete, with variably developed series of scattered dark chromatophores extending dorsally from main body of mark.

Dorsal fin with anterior margin of second unbranched ray outlined by dark chromatophores and with membranes between distal one-half to two-thirds of anterior branched rays with scattered chromatophores. Anal-fin rays with chromatophores at base of anterior branched rays. Middle caudal-fin rays with obscure horizontal stripe; stripe most concentrated anteriorly, giving appearance of distinct spot. Pectoral and pelvic fins hyaline.

ECOLOGY.—Stomach contents of two specimens prepared for clearing and staining consisted mostly of parts of small seeds, with limited amounts of insect parts and insect larvae.

DISTRIBUTION.—Creagrutus anary is known from the middle portions of Rio Madeira basin in Brazil (Figure 22, squares). The species also has been reported from further upstream in the Department of Madre de Dios, Peru (Chang and Ortega, 1995:2).

COMPARISONS.—In addition to Creagrutus anary, only one other Creagrutus species, C. petilus, is known to occur in the central portions of the Rio Madeira basin. Creagrutus anary and C. petilus are readily distinguishable in overall appearance, number of lateral line scales, and head length, snout length, and interorbital width (compare Tables 5 and 46).

MATERIAL EXAMINED.—19 specimens (10, 25.0–44.5).

BRAZIL. Amazonas: Madeira River (=Rio Madeira), approximately 200 miles (=320 km) E of longitude 62°20′W (see under “Remarks,” above, concerning the reported locality), ANSP 39291, 1 (approximately 25 mm, paratype of Creagrutusanary; specimen stained for bone and perhaps cleared, in poor condition). Mouth of Igarapé Puruzinho (7°24′S, 63°00′W), MZUSP 35590, 1. Rio Madeira, Ilha do Puruzinho (7°24′S, 63°00′W), MZUSP 35615, 5 (1, 30.5). Rondônia: Rio Madeira, Calama (8°04′S, 62°52′W), MZUSP 30575, 4 (1, 29.0). Rio Madeira, Paraná do Caraparu, Calama (8°04′S, 62°52′W), MZUSP 31875, 2 (1, 25.0). Rio Madeira, Cachoeira de Santo Antônio (8°43′S, 63°55′W), MZUSP 35604, 6 (34.1–44.5; 2 specimens cleared and counterstained for cartilage and bone).

Creagrutus anary

DIAGNOSIS.—The presence of four median scales between the posterior margin of the anus and the anal-fin origin distinguishes Creagrutus ungulus from all congeners. The only other members of the genus with more than 2 scales in that series (with three scales as a occasional variant) are C. cracentis and C. maxillaris, which have 5 post-anal median scales to the anus. Those species are readily distinguished from C. ungulus in having only two rows of premaxillary teeth contrary to the typical Creagrutus dentition with three major components found in C. ungulus, higher numbers of maxillary and dentary teeth, and in having the third infraorbital well developed contrary to the distinctly reduced third infraorbital of C. ungulus.

In addition to this post-anal scale character, the combination of the possession of premaxillary dentition arranged in the three components generalized for most of the species of Creagrutus and Piabina without a distinctly larger gap between the first and second teeth of the primary series, 4 or 5 teeth on the maxilla, 6 teeth in the primary tooth row of the premaxilla, 5 or 6 dentary teeth, 37 to 41 lateral line scales without a lamellar process over each pore, 9 to 12 predorsal median scales, 4 or 5 scale rows between the dorsal-fin origin and the lateral line, 3 scale rows between the anal-fin origin and the lateral line, 37 to 40 vertebrae, 9 or 10 branched anal-fin rays, 4 post-anal median scales to the anal-fin origin, the distance from the snout to the anal-fin origin (58.4%–63.4% of SL), the distance from the dorsal-fin origin to the pelvic-fin insertion (25.7%–30.7% of SL), the caudal peduncle depth (11.6%–13.0% of SL), the head length (26.8%–29.1% of SL), the postorbital head length (45.2%–51.0% of HL), the snout length (24.2%–27.9% of HL), the bony orbital diameter (27.7%–32.2% of HL), the highly reduced third infraorbital with its posteroventral margin concentric with the orbital rim, the lack of a series of dark midlateral spots on the body, the vertically elongate, ventrally attenuating humeral mark, and the absence of a discrete patch of dark pigmentation on the middle portion of the anterior dorsal-fin rays distinguishes Creagrutus ungulus within the clade composed of Creagrutus and Piabina.

DESCRIPTION.—Morphometric and meristic data for Creagrutus ungulus in Table 55. Head relatively large, more so in larger individuals; body robust anterior to dorsal-fin origin, less so posteriorly at all sizes. Greatest body depth at dorsal-fin origin in smaller specimens, distinctly shifted anterior of that point in larger individuals. Dorsal profile of head smoothly rounded from margin of upper lip to vertical through posterior nostril, slightly convex from that point to tip of supraoccipital spine in small specimens, nearly straight in that region in larger examined individuals. Dorsal profile of body convex from tip of supraoccipital spine to dorsal-fin origin at all body sizes, but with overall convexity of profile of posterior portion of head and predorsal region of body more pronounced in larger specimens. Dorsal profile of body posteroventrally angled along dorsal-fin base and nearly straight from rear of dorsal-fin base to caudal peduncle. Ventral profile of head with slight change in angle at anteroventral corner of dentary. Head and body profile gently convex from that point to anal-fin origin, convexity more pronounced in larger specimens.

Head obtusely pointed in both lateral and dorsal views. Upper jaw somewhat longer than, and overhanging, lower jaw. Snout slightly fleshy anteromedially, with scattered papillae anteriorly; papillae continue onto fleshy upper lip and onto folds and plicae extending between outer and medial premaxillary teeth. Lower lip fleshy, particularly anteriorly, with few scattered papillae.

Infraorbital series poorly developed. Ventral margin of third infraorbital falling distinctly short of horizontal limb of preopercle, more so posteriorly. Posterior margins of third through fifth infraorbitals distinctly separated from vertical limb of preopercle. Distal margins of third and fourth infraorbitals roughly concentric with margin of orbit.

Premaxillary dentition in three series: primary row consisting of 6 teeth arranged in slightly curved series without pronounced gap between first and second tooth of series but with anterior tooth distinctly separated from first tooth of contralateral row; cluster of three teeth with posterolateral tooth distinctly larger; and single tooth of form comparable to those of primary row, lateral to fourth tooth of that series. Maxilla with 4 or 5 tricuspidate teeth. Dentary with 5 or 6 teeth, all teeth tricuspidate when 5 teeth present, sixth tooth, when present, conical or sometimes with only slight indication of side cusps. First through third teeth largest, with first and second teeth subequal or second tooth somewhat larger than first tooth. Both first and second dentary teeth much larger than third tooth. Fourth through last tooth of series progressively decreasing in size.

Dorsal-fin rays typically ii,8, rarely ii,9. Dorsal-fin origin at vertical through pelvic-fin insertion. Profile of distal margin of dorsal fin straight to slightly convex. Anal-fin rays ii,9–10. Distal margin of anal fin distinctly convex, with anterior rays slightly longer but not forming distinct lobe. Anal-fin hooks present on only few examined mature males, located along posterior margin of first through fourth branched rays. Pectoral-fin rays i,12–14. Tip of pectoral fin extending posteriorly from slightly before, to slightly beyond, pelvic-fin insertion. Pelvic-fin rays typically i,6,i, with i,5,i in only 2 examined specimens. Tip of pelvic fin extending posteriorly slightly beyond anal-fin origin. Pelvic-fin hooks present in only few mature males, limited to medial margins of all branched pelvic-fin rays.

Gill rakers 5–7 + 9–11.

COLORATION IN LIFE.—No information available on coloration of live or recently collected specimens. Specimens fixed in formalin and preserved in alcohol examined approximately one month following their capture (MUSM 10362) with yellowish tint on dorsal fin and with yellow pigmentation most intense on basal one-half of branched and anterior unbranched dorsal-fin rays and membranes. Caudal fin with hemicircular patterns of yellow pigmentation on dorsalmost and ventralmost rays and intervening membranes. Caudal fin mostly yellow, pigmentation field most elongate dorsally and ventrally, and gradually shortening towards middle rays of fin, which lack yellow coloration. Middle portions of unbranched and anterior one or two branched anal-fin rays and membranes yellow. Middle portions of the lateral unbranched and first branched pelvic-fin rays with distinct patch of yellow pigmentation. No yellow coloration apparent on pectoral fins. Dark pigmentation as described under “Coloration in Alcohol,” below.

COLORATION IN ALCOHOL.—Overall ground coloration tan to light brown. Dorsal surface of head with field of scattered, dark chromatophores in juveniles; field coalesced into continuous patch of dark pigmentation in larger individuals. Snout and lateral surface of upper jaw with scattered, dark chromatophores. Region in front of anterior nostril with chromatophores more concentrated, but not forming distinct crescent-shaped patch present in many Creagrutus species. Distinct band of dark chromatophores along ventral and posterior margins of orbit in smaller specimens, these subsumed into broader, more diffuse band in larger specimens. Dorsal portion of opercle and region posterior of orbit with scattered, dark chromatophores; chromatophore field in larger specimens extending further ventrally, particularly on posterior portion of check and on opercle. Variably developed dark spot on midlateral surface of opercle; spot typically less developed in smaller specimens. Lateral and dorsolateral surface of body with scattered, dark chromatophores; more concentrated in crescent-shaped patches over center of scales. Chromatophore field denser midlaterally, forming dark midlateral body stripe. Midlateral body stripe beginning about two scales posterior of humeral mark, somewhat diffuse anteriorly and most intense on posterior portion of body and along caudal peduncle. Humeral mark vertically elongate in all examined specimens, with deep-lying, more intensely pigmented dark central region. Humeral mark with less intensely pigmented areas extending dorsally and ventrally from dark central portion resulting in overall ventrally attenuating form, particularly in larger individuals. Dorsal portion of humeral mark merging into overall dark pigmentation of dorsal portion of body in larger specimens.

Dorsal-fin rays outlined with small dark chromatophores, with additional chromatophores scattered over membranes, giving fin dusky appearance, particularly in larger specimens. Anal fin with dark chromatophores along branched rays and intervening membranes, particularly anteriorly; pigmentation more pronounced in larger individuals. Caudal-fin rays delimited by dark chromatophores in smaller specimens, and with chromatophores scattered over membranes, increasingly so in larger specimens. Pectoral- and pelvic-fin rays slightly outlined with dark chromatophores, more so in case of pectoral fins. Pectoral fin in larger specimens with faint indication of dark transverse band.

ETYMOLOGY.—The specific name, ungulus, from the Latin for ring, refers to the narrow ring of infraorbitals bordering the ventral and posterior margins of the orbit.

ECOLOGY.—The type locality for the species, Quebrada Soga, is a small, rocky, rainforest stream approximately 5 m wide and 0.6 deep with clear water and rapid water flow over a substrate of rocks, sand, and detritus (Ortega, 1996:461). Most other specimens of Creagrutus ungulus captured near the type locality, and for which habitat information is available, came from comparable habitats, although some specimens were captured at the mouth of an stream tributary to the Alto Río Madre de Dios, a habitat with less gradient and slower water flow over a mud and cobble bottom.

DISTRIBUTION.—Creagrutus ungulus is known only from the Río Madre de Dios basin of southeastern Peru (Figure 90, diamonds).

COMPARISONS.—The only other species of Creagrutus known to occur within the range of C. ungulus are C. manu and C. occidaneus. Neither of those species has the narrow infraorbitals of C. ungulus, and both have only 2 median scales between the anus and the anal-fin origin contrary to the 4 post-anal scales in C. ungulus. These species also differ from C. ungulus in various meristic and morphometric features.

MATERIAL EXAMINED.—185 specimens (42, 25.0–74.9).

HOLOTYPE.—PERU. Madre de Dios: Provincia Manu, Quebrada Soga, tributary of Río Alto Madre de Dios, 1 km upstream from Erika (opposite Salvacion; approximately 12°53′S, 71°12′W), collected by M. Rauchenberger et al., 5 Sep 1988, MUSM 8878, 1 (66.8).

PARATYPES.—30 specimens (30, 25.0–73.9).

PERU. Madre de Dios: Provincia Manu, Quebrada Soga, tributary of Río Alto Madre de Dios, 1 km upstream from Erika (opposite Salvacion; approximately 12°53′S, 71°12′W), collected with holotype, MUSM 8879, 7 (36.4–66.0); USNM 303066, 10 (47.5–73.9; 3 specimens cleared and counterstained for cartilage and bone). Provincia Manu, stream tributary to Río Alto Madre de Dios, at Erika (opposite Salvacion; approximately 12°53′S, 71°12′W; locality = Quebrada Soga according to Ortega, 1996:465), collected by M. Rauchenberger et al., 5 Sep 1988, MUSM 8880, 4 (47.8–52.8); USNM 302788, 3 (42.2–56.1). Provincia Manu, stream tributary to Río Alto Madre de Dios, approximately 1.5 km upstream from Erika (latter locality at approximately 12°53′S, 71°12′W), between Salvacion and Atalaya, collected by H. Ortega et al., 6 Sep 1988, MNRJ 14481, 6(25.0–47.2).

NONTYPE SPECIMENS.—154 specimens (11, 41.8–74.9).

PERU. Madre de Dios: Provincia Manu, stream tributary to Río Alto Madre de Dios, approximately 1.5 km upstream from Erika (latter locality at approximately 12°53′S, 71°12′W), between Salvacion and Atalaya, USNM 302792, 36. Provincia Manu, Río Manu basin, Quebrada Pachija, at and above its mouth (approximately 11°57′S, 71°17′W), USNM 302785, 2. Provincia Manu, Quebrada Culli, left bank tributary of Río Alto Madre de Dios, 1.5 km upriver from Erika (latter locality at approximately 12°53′S, 71°12′W), USNM 317781, 6. Provincia Manu, Río Alto Madre de Dios, 15 km upstream from Boca Manu (12°19′30″S, 70°58′W), ANSP 143755, 7. Carabaya, Río Inambari, BMNH 1902.11.28:24, 1 (74.9). Cusco: Provincia Quispicanchis, Valley of Río Marcapata, BMNH 1902.5.29:208–209, 2 (60.5–73.8). Provincia Paucartambo, Río Tono, at Hacienda San Jorge, 4 km W on road from Patria (12°53′S, 71°27′W), ANSP 143762, 2 (44.0–56.8); ANSP 151518, 2. Provincia Paucartambo, Río Pilcopata, 3 km above Pilcopata (12°56′30″S, 71°24′W), ANSP 143752, 1 (65.5). Provincia Paucartambo, Pilcopata, Río Hospital, MUSM 10757, 1. Puno: Provincia Sandia, Río Inambari, Muspaypampa, MUSM 5673, 12 (5, 41.8–72.3). Provincia Sandia, Santa Elena, Huinchusmayo, MUSM 5674, 15; USNM 340954, 5. Provincia Sandia, Zona Reservada Tambopata Candamo, quebrada tributary to Río Candamo (13°25′S, 70°01′W), MUSM 12931, 14. Provincia Sandia, Zona Reservada Tambopata Candamo, quebrada tributary to Río Candamo (13°24′S, 70°01′W), MUSM 10362, 15; USNM 344537, 5. Provincia Sandia, Zona Reservada Tambopata Candamo, Río Beshuajali (approximately 13°25′S, 70°01′W), MUSM 10341, 23; USNM 344536, 5.

Creagrutus anary, MZUSP 35604, 2, 40.4–44.4 mm; Brazil, Amazonas, Rio Madeira, Cachoeira de Santo Antônio

Creagrutus atratus, USNM 353867, 2, 47.5–63.4 mm; paratypes, Colombia, Cundinamarca, mouth of Río Saname, Río Meta basin.