Mazarunia charadrica, López-Fernández & Taphorn & Liverpool, 2012

López-Fernández, Hernán, Taphorn, Donald C. & Liverpool, Elford A., 2012, Phylogenetic diagnosis and expanded description of the genus Mazarunia Kullander, 1990 (Teleostei: Cichlidae) from the upper Mazaruni River, Guyana, with description of two new species, Neotropical Ichthyology 10 (3), pp. 465-486 : 475-479

Mazarunia charadrica new species

Figs. 2b , 3b , 6a , 8-10

Mazarunia sp. 1 López-Fernández et al., 2010: 1072, 1075, 1077, 1079-1081 [relationships within Geophagini].

Holotype. UG/ CSBD 1667 , (ex-ROM 83808), 65.2 mm SL, Guyana, Cuyuni-Mazaruni (Region 7), upper Mazaruni River, Waruma Creek , shallow pool at HLF08-19, between 2-10m wide by about 20m long, 5°28’31.8”N, 60°46’46.812”W, 21 Apr 2008, H. López-Fernández, D. C. Taphorn, E. Liverpool, C. Thierens & K. Kramer.

Paratypes. Guyana, upper Mazaruni River , Cuyuni-Mazaruni (Region 7) : ROM 83687 , 2 , 22.0- 24.7 mm SL, Kukui River, upstream of Philipai , 5°22’5.304”N, 60°21’59.1114”W, 16 Apr 2008, H. López-Fernández, D. C. Taphorn, E. Liverpool & C. Thierens . ROM 83693 , 11 , 52.0-66.0 mm SL, Kukui River, in and around Philipai , 5°21’37.116”N, 60°22’18.408”W, 16 Apr 2008, D. C. Taphorn, E. Liverpool, C. Thierens & local fisherman . ROM 83705 , 8 , 20.4-78.5 mm SL (1, 77.6 mm SL), Kukui River, day sample around Philipai , 5°21’37.116”N, 60°22’18.408”W, 16 Apr 2008, H. López-Fernández, D. C. Taphorn, E. Liverpool & C. Thierens . ROM 83709 , 16 , 21.9 -67.0 mm SL (1, 57.8 mm SL), Kukui River, upstream around the camp on Philipai , 5°21’37.116”N, 60°22’18.408”W, 17 Apr 2008, H. López-Fernández, D. C. Taphorn, E. Liverpool, K. Kramer & C. Thierens . ROM 83741 , 1 , 16.97 mm SL, Mazaruni River, sandy beach at the confluence of the Mazaruni River and the Kukui River , 5°40’21.2154”N, 60°28’58.5834”W, 17 Apr 2008, H. López-Fernández, D. C. Taphorn, E. Liverpool, C. Thierens & K. Kramer . ROM 83781 , 6 , 49.7-83.1 mm SL (all measured), Waruma Creek at campsite, 5°28’40.2954”N, 60°46’45.2994”W, 20 Apr 2008, H. López-Fernández, D.C. Taphorn, E. Liverpool, C. Thierens & K. Kramer . ROM 83796 , 1 , 42.2 mm SL, Waruma Creek , down stream from camp at the complex of riffles and shallow, semi-isolated pools, 5°29’9.816”N, 60°47’22.416”W, 21 Apr 2008, H. López-Fernández, D.C. Taphorn, E. Liverpool, C. Thierens & K. Kramer . ROM 83808 , 17 , 20.9-65.1 mm SL, same as holotype ; ROM 83825 , 3 , 19.5 - 25.5 , Paikwa River rapids, 5°28’59.916”N, 60°43’55.704”W, 22 Apr 2008, H. López-Fernández, D. C. Taphorn & E. Liverpool . ROM 83840 , 1 , 53.4 mm SL (1,54.0 mm SL), Paikwa River rapids, 5°29’10.392”N, 60°43’57.1074”W, 22 Apr 2008, H. López-Fernández, D. C. Taphorn & E. Liverpool . ROM 83863 , 2 , 76.7-76.8 mm SL (all measured), Membaru creek , at an artificial sandy beach, 5°55’33.996”N, 60°35’26.808”W, 23 Apr 2008, H. López-Fernández, D. C. Taphorn & E. Liverpool . ROM 89535 , 15 , 32.5-68.5 mm SL (3, 48.8-53.8), Kamarang River, beach on left bank, just downstream from Kelly Kramer’s house, 5°51’59.5002”N, 60°37’14.4984”W, 14 Mar 2011, H. López-Fernández, S. Refvik, D. C. Taphorn, E. Liverpool, K. Kramer & G. Kramer . ROM 89553 , 78 , 20.9-70.2 mm SL (5, 45.0- 68.1 mm SL), Kamarang River, near its mouth at Kelly Kramer’s house ( Kelly Landing ), 5°51’59.5002”N, 60°37’14.4984”W, 12 Mar 2011, E. Liverpool, H. López-Fernández, D. C. Taphorn & S. Refvik . ROM 89559 , 1 , 29.7 mm SL, Mazaruni River, main channel on a mine tailings beach just downstream from Abbou Creek on left bank, 5°42’30.4554”N, 60°21’39.564”W, 4 Mar 2011, D. C. Taphorn, E. Liverpool, H. López-Fernández & S. Refvik . ROM 89585 , 2 , 68.6 - 71.0 mm SL (1, 66.9 mm SL), Kamarang River, outside Kelly Kramer’s house, 5°51’59.5002”N, 60°37’14.4984”W, various dates between 2008-2011, K. Kramer . ROM 89622 , 2 , 38.8-42.8 mm SL, Waruma Creek , riffles and shallow rapids upstream from camp, 5°28’31.8”N, 60°46’46.812”W, 11 Mar 2011, H. López-Fernández, D. C. Taphorn, E. Liverpool & S. Refvik . ROM 89644 , 1 , 41.8 mm SL, Sandaa Creek , run between the lowermost and second set of rapids, 5°34’52.7514”N, 60°48’31.1034”W, 9 Mar 2011, D. C. Taphorn, E. Liverpool, H. López-Fernández & S. Refvik . ROM 89647 , 3 , 35.3-53.1 mm SL (1, 52.0 mm SL), Mazaruni River at Kamarang , 15 Mar 2011, D. C. Taphorn, E. Liverpool, H. López-Fernández & S. Refvik . ROM 89659 , 6 , 32.7-58.6 mm SL (1, 50.0 mm SL), Mazaruni River, Warwata village, near mouth of Kamarang River , 5°55’1.776”N, 60°36’13.4994”W, 16 Mar 2011, D. C. Taphorn, E. Liverpool, H. López-Fernández & S. Refvik . ROM 89673 , 6 , 42.9-55.3 mm SL (1, 55.0 mm SL), Mazaruni River, backwater channel on right bank in front of Warwata village, near mouth of Kamarang River , 24 Mar 2011, H. López-Fernández, D. C. Taphorn & E. Liverpool . ROM 89697 , 3 , 51.5-65.2 mm SL (3, 50.0- 63.4 mm SL), Abbou Creek , approx. 1.2 km walking E from camp, 5°44’10.4274”N, 60°21’34.416”W, 5 Mar 2011, H. López-Fernández, D. C. Taphorn & E. Liverpool . ROM 89701 , 2 , 59.3- 70.7 mm SL (2, 59.6-68.3 mm SL), Sanda Creek tributary about 500 m downstream from HLF11-17, 9 Mar 2011, H. López-Fernández, D. C. Taphorn, E. Liverpool & S. Refvik . ROM 89703 , 13 , 48.7- 70.9 mm SL (1, 69.6 mm SL), Kamarang River, right bank, across the channel from Kelly Kramer’s house, 5°51’59.5002”N, 60°37’14.4984”W, 12 Mar 2011, E. Liverpool & D.C. Taphorn . ROM 89735 , 2 , 69.7-74.2 mm SL (2, 69.5-73.9 mm SL), Mazaruni River , river channel on right bank, across from HLF11-29, 5°55’1.776”N, 60°36’13.4994”W, 13 Mar 2011, D. C. Taphorn, H. López-Fernández, E. Liverpool & S. Refvik . ROM 89749 , 2 , 40.3-64.3 mm SL (all measured), Mazaruni River, channel on left bank, downstream from Kamarang , 1 km upstream from Membaru, 5°55’1.776”N, 60°36’13.4994”W, 13 Mar 2011, H. López-Fernández, D. C. Taphorn, E. Liverpool, S. Refvik & K. Kramer . ANSP 192031 (ex. ROM 89553 ), 5, 46.1-64.7 mm SL, Kamarang River, near its mouth at Kelly Kramer’s house ( Kelly Landing ), 5°51’59.5002”N, 60°37’14.4984”W, 12 Mar 2011, E. Liverpool, H. López-Fernández, D. C. Taphorn & S. Refvik .

Diagnosis. Mazarunia charadrica can be distinguished from all other cichlids by the generic diagnostic characters. Phylogenetic analysis indicates it can be distinguished from the other species of Mazarunia by the apomorphic structure of its first epibranchial bone (100-1: uncinate process and anterior arm of approximately equal width and 101-2: reduced anteroventral expansion vs. uncinate process narrower and complete anteroventral expansion in M. mazarunii and M. pala ); a dorso-ventrally flattened maxillary process of the palatine is a plesiomorphic condition restricted to M. mazarunii (vs. palatine maxillary process cylindrical in the other two species). Externally, M. charadrica is distinguished by having cycloid scales in the opercular, postorbital, lateral chest and anal-genital regions (vs. ctenoid in the other species). M. charadrica is further diagnosed by the lack of infraorbital and supraorbital bars present in M. mazarunii and the mid-lateral spot present in both M. mazarunii and M. pala . In M. charadrica the supraorbital stripe is diffuse and expanded into a dark area that covers the dorsal portion of the head in an ascending line from the lower lip to the ventral edge of the orbit and from there to a point about mid-way between the posterior edge of the orbit to the insertion of the dorsal fin, giving the impression that the fish has a “black cap” ( Figs. 8-10 ). M. charadrica has a unique pattern of seven vertical bars which are most obvious in juveniles but are not completely lost in adults. Bars 3-6 in antero-caudal direction are most visible in juveniles and medium-sized specimens but become fainter and almost disappear in adults. Many specimens show only bar number 3 (midlateral bar).

Description. Measurements are summarized in Table 1. Males with relatively small nuchal hump anterior to first dorsal-fin spine, possibly with longer dorsal and anal-fin filaments. Head slightly broader ventrally. Mouth subterminal. Dorsal head profile gently convex, from tip of snout to dorsal-fin origin; dorsal base slightly convex, descending gently to last dorsal-fin spine, then steeply to dorsal-fin caudal insertion; dorsal caudal peduncle straight or slightly convex to caudal-fin base. Ventral head profile straight or lightly convex, descending to vertical line from pectoral-fin base; flat or slightly convex to anal-fin origin; anal-fin base straight, ascending; ventral caudal peduncle profile slightly concave, ascending caudal-fin base. Tip of snout slightly anterior to upper lip in some specimens; upper lip about half as wide as lower; lower lip widest caudally, sometimes covering caudal-most edge of upper lip when mouth closed. Lower lip fold broad, discontinuous at dentary symphysis. Maxilla reaching from one third to less than one half distance between nostril and orbit. Opercle, preopercle, cleithrum, supracleithrum and post-temporal smooth.

E1 24(3), 25(27)*, 26(4); scales between upper lateral line and dorsal fin 3(11)*, 3½(23) anteriorly and 1(4), 1½(30)* posteriorly. Scales on upper lateral line 15(2), 16(12), 17(15)*, 18(5), and lower lateral line 7(6), 8(20)*, 9(8). Circumpeduncular scale rows 7 above, 7 below lower lateral line, ctenoid. Scale rows between lateral lines 2. Opercle, subopercle and cheek fully scaled, cycloid or slightly ctenoid; interopercle caudally scaled, cycloid. Occipital and flank scales ctenoid. One column of postorbital scales, cycloid. Lateral chest scales cycloid, smaller than flank scales. Dorsal, anal, pectoral and pelvic fin naked; caudal fin with row of ctenoid scales between rays to one third of fin length. Accessory caudal-fin extensions of lateral line restricted to one tubed scale at base of dorsal-fin between D3 and D4 and ventrally between V4 and V5.

Dorsal fin XV,8(1), XVI,8(28)*, XVII,8(5); anal fin III,7(6)*, III,8(28). Dorsal spines increasing in length from first to about 7 th- 8 th, then decreasing slightly; last spine about two thirds of first ray length, about half length of longest soft ray; membranous lappets behind all spines. Soft portion of dorsal fin pointed, 4 th ray longest, sometimes slightly produced as filament reaching one third to half of caudal fin length, especially in males with nuchal humps. Caudal fin symmetrical, distal margin almost straight, indented slightly at middle caudal-fin rays. Anal fin pointed, 3 rd- 4 th rays longest, sometimes extended as filaments, reaching up to half caudal-fin length. Pectoral fin asymmetrical, rounded dorsally, more or less straight ventrally from 4 th ray, which is longest; longest rays not reaching pelvic-fin or anus. Pelvic-fin spine shorter than first ray which is longest, barely reaching anus unless produced into filament, which may reach 1 st anal spine; each subsequent pelvic-fin ray shorter till last, which is about half length of longest.

Outer premaxilla hemiseries with 7-8 teeth, progressively smaller distally from symphysis. Three irregular rows of much smaller, unicuspid teeth, separated from outermost row by gap. Dentary hemiseries with 11-16 thin, sharp, slightly recurved, unicuspid teeth in outer row, two irregular inner rows with much smaller teeth separated from outermost row by gap, widest anteriorly and progressively narrow caudally. Dentary hemiseries separated by small symphyseal gap, particularly in outermost row.

External rakers on first gill arch 8(1), 9(6), 10(23)*, 11(4). Lower pharyngeal tooth plate wider than long, length of bone 79% of width; dentigerous area 66% of width, 62% of length; 9-12 teeth in posterior row; 5-7 in median row. Lateral outer teeth unicuspid, cylindrical, slightly recurved, progressively smaller caudally, larger medially; posteromedial teeth much larger, molariform, unicuspid, with medial cusps slightly curved anteriorly ( Fig. 6a ). Two tooth plates on fourth ceratobranchial. Vertebrae. 12+14=26(9). One supraneural bone.

Color in alcohol. Figs. 8-9 . Base color pinkish brown to beige; snout, region between posterior margin of maxilla to eye and top of head near supraoccipital dark brown or gray. Lips light gray. Side of head, throat, interopercle, lower region of preopercle and chest light beige; gill cover with brown-black blotch, most pronounced dorsally. Infraorbital stripe absent; supraorbital stripe present, directed caudad, distinct in small juveniles ( Fig. 9c ), progressively expanding with growth until forming dark “cap” extending diagonally from dorsal midline caudad to orbit to caudo-ventral margin of orbit and then to corner of mouth ( Fig. 9 a-b); groove from posterior margin of nare to anterior margin of eye light beige. Branchiostegal membrane brown, greatly contrasting with light chest. Body counter shaded, darkest at dorsal midline, most scales outlined with gray to black, outlines darker and thicker dorsally, lightest below pectoral fin; centers of scales light, forming rows of light spots, most notable on caudal peduncle. Flanks with seven vertical bars, variably expressed, most obvious in juveniles but not completely lost in many adults (including holotype).Anteriormost bar coincides with dorsal fin insertion, continuous with gill cover blotch in larger individuals ( Fig. 9a ), but separated by lighter area in smaller specimens ( Fig. 9 b-c); second bar extending anteroventrally from about 4 th and 5 th dorsal spines to E1, forming broad “U-shape” with first bar just caudad of postcleithrum; third bar approximately between 7 th and 9 th dorsal spines, 2 scales wide, extending vertically under pectoral fin to H2, or reaching H 4 in smaller juveniles ( Fig. 9c ); fourth bar approximately below 12 th to 14 th dorsal-fin spines, two scales wide, extending ventrally, slightly caudad to posterior edge of pectoral-fin to E0 in small adults, to H 2 in small juveniles; fifth bar below last two dorsal spines and first soft ray, two scales wide dorsally, extending slightly caudally to E1 then slightly rostrally to H1; sixth bar below caudal insertion of dorsal fin, about two scales wide, extending antero-ventrally to E0, then caudo-ventrally to H1, forming “H-figure” with fifth bar; seventh bar most noticeable on E0, forming faint blotch in middle of caudal peduncle, about 1-1½ scales wide and deep. Bars 3-6 most visible in juveniles and medium-sized specimens ( Fig. 9 b-c), tend to almost disappear in larger adults ( Fig. 9a ). Many specimens show only bar 3 (midlateral bar). Spinous portion of dorsal fin and base of soft portion black, rest of fin plain, dusky gray. Caudal fin gray, rays lighter than membranes. Anal fin gray, darker pigment along rays. Pelvic fin gray, first rays darkest.

Color in life. From photographs of live and freshly preserved specimens ( Fig. 10 ). Melanic coloration as for preserved specimens. Background color pinkish-tan to purple, most scales with lighter centers, often forming horizontal rows of light spots, and reticulate pattern on sides, more apparent posteriorly. Head from tip of snout above eye and posterior to tip of supraoccipital dark brown to gray, anteriorly tapering light stripe from dorsal anterior margin of eye to posterior naris. Iris greenish with golden highlights. Upper lip brown at tip, white laterally, lower lip white. Head below eye bright pink, yellow or orange, extending onto opercle and sides of body posterior to opercle and anterior to pectoral-fin base. Pink or orange coloration extending more or less continuously to bright orange blotch just dorsal to insertion of pectoral fin. Branchiostegal membrane pink to orange or maroon. White on ventral portion of head and ventrum between pectoral-fin base and posteriorly onto abdomen. Dorsal fin gray to maroon brown on spinous portion and base of soft rays, rest of soft portion hyaline or slightly dusky. Anal-fin spines blackish gray, soft rays pink to maroon, membranes lighter. Pectoral fin transparent, rays thinly outlined with black. Pelvic whitish, anterior rays outlined in black. Caudal fin dusky gray, membranes weakly pink.

Geographic distribution. Mazarunia charadrica has the broadest distribution of all species in the genus, known from the upper Mazaruni River main channel, its direct tributaries, the Kamarang and Kukui Rivers and Abbou creek, and from several tributaries of the Kako River, including Waruma, Sandaa and Paikwa creeks. It is presumably present in the Kako River itself, but conditions in the field prevented us from performing collections. Areas without sites on the map have not been sampled by ichthyologists, but we expect the species to be present in all upper Mazaruni tributaries ( Fig. 7 ).

Etymology. From the Greek charadra (χαραδρα), a mountain stream or a torrent. In reference to its apparent preference for reophilous habitats; the species is often found in the steep tributaries of the Mazaruni River that flow from the slopes of the Roraima massif and other mountainous sources in the area. To be regarded as an adjective in feminine form.

Habitat. Mazarunia charadrica is the most reophilic of all three species. Although it can be collected with the other species in main channel sites with sandy bottoms and sluggish currents, this species is abundant in the entire upper Mazaruni River basin, and especially in tributaries with rocky substrates and rich in rapids. Captured in black water sites with fairly high transparency (0.55-1.0 m Secchi depth), none to relatively fast currents (0-0.6 m /s), temperature (21.7- 24.5ºC), pH (4.4- 4.8), dissolved oxygen (5.2- 9.3 mg /L) and very low conductivity (<10 μS).

Figures

Fig.2. Infraorbital series in left, lateral view, illustrating morphological character 116; a) Guianacara dacrya,character state 116-1, paratype, 58.0 mm SL,ANSP 179495; b) Mazarunia charadrica, character state 116-3, 56.0 mm SL, ROM 83808; c) Mazarunia mazarunii, character state 116-3, 50.1 mm SL, ROM 83869 d) Mazarunia pala, character state 116-3, 50.8 mm SL, ROM 83753.

Fig. 3. First epibranchial and pharyngobranchial bones in right, approximately antero-dorsal view illustrating morphological phylogenetic characters 100 and 101; a) Guianacara dacrya, character states 100-0, 101-0, 58.0 mm SL, ANSP 179495; b) Mazarunia charadrica, character states 100-1, 101-2,56.0 mm SL, ROM 83808; c) Mazarunia mazarunii, character states 100- 2,101-4, 50.1 mmSL,ROM 83869d)Mazarunia pala,character states 102-0, 101-4, 50.8 mm SL, ROM 83753; UP, uncinate process of the first epibranchial; AA, anterior arm of the first epibranchial; PhB, first pharyngobranchial.

Fig. 6. Lower pharyngeal jaw of species of Mazarunia in occlusal view; a) Mazarunia charadrica, paratype, ROM 89553,67.1mm SL, b) Mazaruniamazarunii, ROM 83737,58.5 mmSL, c) Mazarunia pala,paratype, ROM 83753,43.8 mmSL.

Fig. 8. Mazarunia charadrica, holotype, UG/CSBD 1667, 65.2 mm SL, Guyana, Cuyuni-Mazaruni (Region 7), Waruma Creek, tributary of Kako River, upper Mazaruni River drainage.

Fig. 9. Mazarunia charadrica, ontogenetic changes in overall body shape and melanic coloration; a) adult, 67.2 mm SL, b) young adult, 45.3 mm SL, c) juvenile, 21.9 mm SL.All specimens from ROM 83709.

Fig. 10. Mazarunia charadrica, 65.0 mm SL, ROM 89553, live coloration in a specimen from the Kamarang River, photographed shortly after capture.

Fig. 7. Map of the upper Mazaruni River basin showing collection localities and known distribution of the three species of Mazarunia. Blue squares: Mazarunia charadrica, Black circles: Mazarunia mazarunii; Purple triangles: Mazarunia pala. One symbol may represent more than one collection locality.