Mazarunia mazarunii Kullander, 1990

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 : 471-475

Mazarunia mazarunii Kullander, 1990 Figs. 2c , 3c , 4 , 5 , 6b

Mazarunia mazarunii Kullander 1990: 3-14 [original description; type species by original designation; relationships to Dicrossus and Crenicara ]. - Kullander, 2003: 638 [summary of type information, body size and distribution]. - López-Fernández et al., 2010: 1072, 1075, 1077, 1079-1081 [relationships within Geophagini].

Diagnosis. Mazarunia mazarunii can be distinguished from all other cichlids by the generic diagnostic characters; additionally, phylogenetic analysis revealed that the species can be distinguished from all other species of Mazarunia by the autapomorphic presence of two foramina (vs. one) on the lateral face of the ascending process of the premaxilla (89-2); a number of plesiomorphic characters in combination are also useful in the species diagnosis: lachrymal bone that is longer than deep (122-0, vs. deeper than long), and by an infraorbital 3 that is contiguous but not overlapping with the lachrymal (125-0, vs. overlapping, Fig. 2c ). Externally, M. mazarunii can be distinguished from its congeners by the presence of ctenoid scales (vs. cycloid) on the subopercle, interopercle and chest, and by unique characters of its coloration, including a complete suborbital stripe extending from the lower edge of the orbit to the interopercle, the presence of a supraorbital stripe directed caudad and for being the only Mazarunia species with a discernible lateral band formed by the mid-line blotching pattern associated with lateral bars. Finally, M. mazarunii has a black or dark bar from the occipital region, along the caudal margin of the opercle to the insertion of the pectoral fin, which in large adult individuals gives the impression of a collar ( Figs. 4a and 5a ).

Description. Measurements are summarized in Table 1. No apparent sexual dimorphism. Head equal or slightly broader ventrally than dorsally. Dorsal head profile ascending, convex from tip of snout to insertion of dorsal fin; dorsal-fin base convex, horizontal to 10-11 spine, descending gently to last spine, then abruptly to last soft ray; dorsal caudal peduncle straight, descending to caudal-fin base. Ventral head profile straight or slightly convex, descending to pelvic-fin insetion; straight or slightly concave, horizontal to anal-fin insertion; anal-fin base convex, ascending; ventral caudal peduncle straight, ascending to caudal-fin base. Upper lip about half as wide as lower; lower lip uniformly wide, covering distal portion of upper lip when mouth closed. Lower lip fold broad, discontinuous at dentary symphysis. Maxilla reaching about one-half of distance between nostril and orbit. Opercle, preopercle, cleithrum, supracleithrum and post-temporal smooth.

E1 scales 24(1), 25(9), 26(3); upper lateral line scales 16(4), 17(4), 18(3), 19(2); lower lateral line scales 8(2), 9(11). Scales between dorsal-fin insertion and upper lateral line 3(5), 3½(9); scales between caudal-most upper lateral line tubed scale and dorsal fin 1(2), 1½(5), 2(6). Circumpeduncular scales ctenoid, 7 rows above and 7 below lower lateral line. Scale rows between lateral lines 2. Opercle fully scaled, ctenoid; subopercle fully scaled, ctenoid; cheek fully scaled, cycloid; interopercle caudally scaled, ctenoid. Occipital and flank scales ctenoid. One column of postorbital scales, ctenoid. Lateral chest scales ctenoid, smaller than flank scales. Dorsal, anal, pectoral, and pelvic fin naked; caudal fin with single row of ctenoid scales between rays, covering basal one quarter 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 (and see Kullander, 1990).

Dorsal-fin rays XVI,6(1), XVI,7(1), XVI,8(1), XVII,7(4), XVII,8(5); anal III,7(3), III,8(7), IV,7(2), IV,8(1). Dorsal spines increasing in length from first to 6 th- 7 th, gradually decreasing to last; short membranous lappets behind all spines. Soft portion of dorsal fin rounded in smaller specimens (<48.0 mm SL), increasingly pointed in larger individuals, 2 nd to 4 th soft ray longest, not produced into filaments, not generally reaching caudal-fin base. Caudal fin symmetrical, distal margin slightly emarginated. Anal fin rounded, 4 th ray longest, not produced even in largest specimen (76.7 mm SL), not reaching caudal-fin. Pectoral fin asymmetrical, slightly rounded dorsocaudally, 2 nd and 3 rd rays longest; longest rays not reaching anus. Pelvic fin triangular, spine about half length of first ray, which is longest, often produced into reaching up to 4 th anal-fin spine.

Premaxillary hemiseries in outer row with 6 to 12 thin, sharp, slightly recurved, unicuspid teeth, progressively smaller distally from symphysis. Three irregular rows of smaller, very thin, straight or very slightly recurved unicuspid teeth, separated from outermost row by wide gap. Dentary hemiseries with 18 to 20 teeth in outer row, size diminishing caudally such that anterior 6 are clearly larger than caudad ones. Two inner rows with smaller teeth, cylindrical, straight, unicuspid, separated from outer row by gap that disappears laterally, teeth eventually line up with smaller teeth of anterior row. Dentary and premaxillary hemiseries approximately continuous with each other, with no readily visible gap between them.

External rakers on first gill arch 11(6), 12(6), 13(1). Lower pharyngeal tooth plate ( Fig. 6b ) slightly wider than long, bone length about 90% of width; dentigerous area 87% of width, 76% of length; 13-15 teeth in posterior row; 8-9 in median row. Lateral outer teeth unicuspid, cylindrical with recurved tips, progressively smaller caudally, larger medially; posteromedial larger, cylindrical, unicuspid, with medial cusps. Three tooth

Mazarunia pala Mazarunia charadrica Mazarunia mazarunii Holotype N Min Max Mean SD Holotype N Min Max Mean SD N Min Max Mean SD

SL 62.7 18 32.5 74.2 47.5 11.7 65.2 33 39.3 84.1 61.2 10.6 13 32.4 76.7 52.2 12.5 Percent of Standard length

Head length 34.9 18 33.2 39.2 36.3 1.7 35.2 33 33.4 38.6 36.1 1.2 13 33.3 36.8 34.9 1.2

Body depth 44.6 18 39.5 46.8 43.0 2.0 36.9 33 35.3 41.6 38.4 1.5 13 37.4 43.0 40.0 1.6

Caudal peduncle depth 19.4 18 16.8 19.7 17.9 0.7 16.8 33 15.7 18.2 16.9 0.6 13 14.5 17.8 16.3 1.0

Caudal peduncle length 12.1 18 10.2 13.7 11.9 1.0 14.8 33 11.4 16.1 13.7 1.1 13 11.1 16.0 13.4 1.3

Pectoral fin length 28.6 18 23.3 31.5 28.4 2.3 27.3 33 18.9 29.7 25.6 1.9 13 24.2 31.7 26.9 2.0

Pelvic fin length 27.4 18 23.7 32.4 27.2 2.3 27.4 33 24.1 35.1 28.5 2.5 13 27.4 37.8 32.4 2.9

Last D spine length 10.0 18 9.8 15.3 11.8 1.6 11.5 33 8.5 15.1 11.7 1.7 13 9.6 14.0 11.5 1.4 Percent of Head length

Snout length 41.5 18 31.6 46.2 39.0 4.3 38.4 33 38.4 46.0 41.2 2.6 13 27.5 37.0 33.4 3.0

Orbital diameter 26.4 18 23.2 31.7 27.7 2.4 24.6 33 24.6 30.9 26.1 2.3 13 27.3 35.8 30.0 2.6

Head width 53.2 18 42.8 58.9 49.1 4.2 45.9 33 45.9 51.4 47.6 1.9 13 32.0 51.8 46.4 5.1

Interorbital width 30.1 18 26.6 36.3 30.7 3.2 27.9 33 27.9 33.0 28.4 2.3 13 28.4 35.5 31.3 2.1

Preorbital depth 26.4 18 19.4 30.3 23.9 2.8 25.4 33 25.4 31.5 25.2 2.7 13 14.5 21.7 18.0 1.8

plates on fourth ceratobranchial. Vertebrae 12+14=26(1), 13+13=26(1), 13+14=27(5).

Color in life and in alcohol. See Kullander (1990) for full description of pigmentation pattern. Previous description herein expanded by incorporating larger adults up to 76.7 mm

SL, largest known individual (ROM 89531). We comment on ontogenetic color variation ( Figs. 4 and 5 ). Life and alcohol coloration treated together as most coloration is melanic and maintained after preservation. Exceptions are gill cover, which has coppery sheen in large adults ( Fig. 4a ), and base color tending to be lighter tan or pinkish color in live juveniles ( Fig. 4b ). Base color grayish-beige, countershaded. Most flank scales in large adults with light centers and grey posterior edge, alignment giving impression of alternating rows of light spots with gray horizontal lines. Head with base color anterior to eye and immediately posterior of orbit; much darker in supraorbital area, making supraorbital bar less distinct. Infraorbital stripe less continuous than in smaller individuals, stronger at corner of preoperculum and at interoperculum, faded along cheek. Whitish stripe from anterior margin of orbit to nostril seen in juveniles changed into darker brown stripe over greyish-beige color of snout. Bar running from dorsal midline just anterior to dorsal-fin origin (Bar 7 in Kullander, 1990) descending vertical to upper lateral line, continuing on caudal edge of opercle, slightly lighter through interopercle and branchiostegal membrane giving impression of collar. Dark vertical bars of juveniles greatly reduced. Following Kullander’s (1990) numbering system, bars 1 and 2 present as diffuse spots or blotches, bars 3 to 6 reduced to wide, faint spots at base of dorsal-fin; in some cases corresponding faint midlateral blotch variably present in bar 4; bar 5 extends ventrally, uniting with large, black midlateral spot, which is about 4 scales wide and 4 scales high; dorsal edge of midlateral spot includes scales of upper lateral line.

Geographic distribution. According to Kullander (1990), the two type specimens were collected from the blackwater Kamarang River , slightly up-stream of Kamarang (the town is located at the mouth of the Kamarang River where it joins the Mazaruni River ).Additional localities from the Mazaruni River and Membaru Creek are shown in Fig. 7 . Mazarunia mazarunii was most frequently found on sandy beaches, many deposited by gold mining dredges, along the main channel of the Mazaruni River and its tributaries .

Habitat. Mazarunia mazarunii was collected mostly in or near the main upper Mazaruni River channel, generally in localities with sandy or slightly muddy substrates, some times with some structure formed by submerged or emerging grass and other riparian vegetation and some drift wood. At night all species of Mazarunia can move into shallow, flooded banks which can have substantial amounts of flooded terrestrial vegetation.All sites had black water with parameters typically ranging from 21.8-24.5ºC, high levels of dissolved oxygen between 5.2 and 9.3 mg /L, acid pH of 4.0-4.8 and none to slow currents usually around 0-0.2 m /s, except for one locality where it reached 0.4 m /s. Conductivity was exceedingly low and ranging between 0-10 μS. The species was frequently caught in artificial gold-mining “tailings” beaches along the Mazaruni main channel; these sites tend to artificially have lower transparency (0.2-0.3 m Secchi depth) than unaltered sites where the species is present (e.g. 0.6- 0.85 m Secchi depth).

Material examined. Guyana, Cuyuni-Mazaruni (Region 7), upper Mazaruni River: ROM 83737, 1, 59.6 mm SL Mazaruni River, sandy beach at the confluence of the Mazaruni River and the Kukui River. ROM 83752, 2, 42.1 and 46.1 mm SL, Mazaruni River, sandy beach and associated embayment both created by gold-mining dredging. ROM 83864, 1, 47.3 mm SL, Membaru creek at an artificial sandy beach. ROM 83869, 4, 40.5-48.5 mm SL, Mazaruni River, beach downstream from Kamarang. ROM 89522, 5, 35.2- 42.7 mm SL (1, 35.6 mm SL), Kamarang River, beach on left bank, just downstream from Kelly Kramer’s house. ROM 89531, 2, 32.4 and 76.7 mm SL, Mazaruni River, backwater channel behind beach sampled at HLF11-06, 2 km downstream from site 5. ROM 89551, 2, 46.8 and 29.6 mm SL, Mazaruni River, main channel on a mine tailings beach just downstream from Abbou Creek on left bank. ROM 89586, 5, 48.1-63.1 mm SL, Kamarang River, outside Kelly Kramer’s house. ROM 89609, 2, 26.4-38.1 mm SL, Mazaruni River, channel at backwaters about 50 m behind HLF11-04. ROM 89674, 1, 42.0 mm SL, Mazaruni River, backwater channel on right bank in front of Warwata village, near mouth of Kamarang River. ROM 89745, 10, 29.8-47.4 mm SL, Mazaruni River, channel on left bank, downstream from Kamarang, 1 km upstream from Membaru.

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. 4. Live colors of Mazarunia mazarunii through ontogeny; a) adult individual, probably male, 76.7 mm SL, ROM 89531 shortly after capture and anesthetized with clove oil, b) juvenile 38.1 mm SL, ROM 89609 shortly after capture.

Fig. 5. Ontogenetic changes in body shape and coloration pattern in Mazarunia mazarunii; a) same adult individual as shown shortly aftercapture and anesthetized in Fig.6a, 76.7 mm SLROM 89531, b) 61.2mm SLROM 89586, c)38.2 mm SLROM 89522.

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. 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.