Parapercis maramara, Sparks, John S. & Baldwin, Zachary H., 2012

Sparks, John S. & Baldwin, Zachary H., 2012, A new species of Parapercis (Teleostei: Pinguipedidae) from Madagascar, Zootaxa 3204, pp. 31-39 : 32-38

Parapercis maramara , new species

Figures 1–2 ; Table 1

Holotype. AMNH 243241, 108.2 mm SL, Madagascar, Masoala Peninsula (15° 44’S, 50° 21’E), Masoala National Park, outside Tanjona Marine Reserve, coll. J.S. Sparks, W.L Smith, and K. Tang, 16 November 2003.

Paratypes. AMNH 243242, 4 ex., incl. 1 ex. C&S, 24.5–95.1 mm SL, Madagascar, Masoala Peninsula (15° 44’S, 50° 21’E), Masoala National Park, outside Tanjona Marine Reserve, coll. J.S. Sparks, W.L Smith, and K. Tang, 18 November 2003. AMNH 243243, 1 ex., 53.7 mm SL, Madagascar, Masoala Peninsula (15° 44’S, 50° 21’E), Masoala National Park, outside Tanjona Marine Reserve, coll. J.S. Sparks, W.L Smith, and K. Tang, 14 November 2003.

Diagnosis. The new species is distinguished from all congeners reported from the Western Indian Ocean by a unique pigmentation pattern on the caudal fin comprising a smaller oblong black patch dorsally near the base of the fin and a much larger elongate oval or anvil-shaped black patch ventrally that extends posteriorly over half the length of the fin, the presence of numerous smaller black blotches forming two to three bands distally on the caudal membrane, and a unique spotting pattern on the flank, comprising six to nine black blotches present along the lateral midline over a light background and a second series of eight or nine black blotches (none forming ocelli) on the lower flank at the ventral terminus of the lateral series of vertical brown bands.

The new species is further distinguished from P. hexophtalma , P. maculata , P. punctulata , and P. robinsoni by a unique pattern of large, closely arrayed diffuse brown blotches on the cheek, opercle, lips, and snout (vs. small, densely pigmented, more sparsely arrayed spots in the three former species, and few faint, very large, patches in P. robinsoni ). The new species is distinguished further from P. punctulata and P. robinsoni by the presence of multiple clustered bands of villiform teeth on the vomer (vs. single band in P. punctulata , or four large, stout villiform teeth in P. robinsoni ), and from P. hexophtalma and P. robinsoni by a concave interorbital space (vs. flattened to weakly convex) and pelvic-fin origin located well anterior to pectoral origin, at about 1/3 of the distance from the posterior margin of the interopercle to the base of the pectoral fin (vs. origin of fins at about same vertical). The new species is further distinguished from P. hexophtalma by the absence of strong, dense spotting on the nape and predorsal region (vs. densely spotted), and by the presence of a V-shaped pattern of spots posterior to the orbit in dorsal view (vs. no clear pattern), and from P. robinsoni by a more oblique mouth (vs. horizontal) and longer pelvic fin (24.6– 31.9% vs. 18.4–22.1% SL).

Description. Selected proportional measurements and meristic data presented in Table 1 . Body elongate and moderately dorsoventrally compressed anteriorly. Body becoming rounded and tubular in cross-section medially, and laterally compressed posteriorly. Head dorsoventrally compressed, particularly anteriorly, and elongate. Snout pointed both in lateral profile and to a lesser extent in dorsal view. Predorsal profile weakly curved. Orbit large, eyes strongly oriented dorsolaterally, and fully included (visible) in dorsal view. Interorbital region slightly concave, its width roughly half of orbit diameter. Suborbital distance approximately one and a half times orbit height. Nape broad and somewhat flattened. Mouth large, slightly oblique, with posterior margin of maxilla extending to about mid-orbit. Width of mouth relatively narrow. Upper lip broad throughout. Lower lip broad anteromedially, becoming narrow posteriorly. Lower jaw weakly prognathous, with rostral margin protruding slightly anterior to upper jaw. Symphysis of lower jaw parabolic in shape in ventral view.

Interorbital series comprised of seven elements, including lacrimal, with uppermost element firmly attached to neurocranium. Preopercle with wide sharp serrations along caudal margin. Serrations becoming rounded and less pronounced along ventrocaudal margin of preopercle. Operculum bearing a single well-developed spine, which extends just posterior to pectoral-fin origin. Weak serrations present near ventrocaudal margin of subopercle. Operculum extending just anterior to origin of spinous dorsal fin. Gill membranes free of isthmus, but with broad medial flap of tissue.

Character N Range Holotype Mean SD Standard length (mm) 6 24.5–108.2 108.2 69.7 32.7 Meristics

Vertebrate (precaudal + caudal) 5 10 + 19 = 29 10 + 19 = 29 Dorsal fin 6 V, 2 1 V, 2 1 Anal fin 6 I, 18 I, 18 Pectoral fin 6 16 16 Pelvic fin 6 I, 5 I, 5 Lateral line scales 4 58–60 58 Upper jaw with outer row of large, slightly recurved caniniform teeth, and an anteromedial band comprising several rows (six or seven anteromedially and tapering to a single row posteriorly) of much smaller villiform to small conical teeth. Villiform and small conical upper jaw teeth restricted medially. Outer canine teeth in upper jaw becoming progressively larger posteriorly to the seventh or eighth tooth in series (i.e., the largest) on each side, then becoming progressively smaller posteriorly. Outermost row on lower jaw comprised of three large recurved canine teeth on each side, with posteriormost, third tooth in series largest. A broad medial band of small villiform to conical teeth present with numerous rows medially (four or five), and tapering to a single row posteriorly. Teeth in band somewhat larger and conical anteromedially. A single outer row of smaller canine teeth present posteriorly. Row posterior of outermost anterior series of three enlarged canines in lower jaw comprised of villiform to small conical teeth rostrally, and becoming caniniform posteriorly. Vomer with multiple (two to three) rows of stout slightly recurved conical teeth, with teeth in anterior row much larger than in succeeding rows. Only anterior row of vomerine teeth complete, whereas posterior row(s) only sparsely populated with teeth. Palatine teeth absent in the specimens examined.

First gill arch with eight to nine triangular and highly denticulate outer ceratobranchial rakers, and two smaller hypobranchial rakers of same morphology. Five to six elongate epibranchial rakers present on first arch. Other rakers on arches two to four short and robust, and highly denticulate apically. Fifth ceratobranchial elements separate and boomerang shaped, with three rows of elongate and curved conical teeth; teeth in medial row much larger than those in other rows. Upper pharyngeal toothplates with elongate and curved conical teeth; teeth becoming progressively smaller posteriorly. Two rows of teeth present on second pharyngobranchial toothplate, three to four rows on third pharyngobranchial toothplate (teeth not arranged in well-defined rows), and numerous rows of much smaller teeth on fourth upper toothplate. Dentition on fourth upper toothplate becoming progressively quite small and poorly developed posteriorly. Six branchiostegals present.

Scales on flank and opercle comparatively large and ctenoid. Smaller ctenoid scales present on caudal fin, and extending approximately ¾ length of fin. Scales on nape reduced in size; cycloid anteriorly and becoming weakly ctenoid posteriorly. Scales on cheek numerous and cycloid, much smaller than those on flank, and comprising 15 rows. Ctenoid scales present on opercle. Ventrum fully scaled. Interpelvic chest scales reduced in size, weakly ctenoid posteriorly, and cycloid anteriorly. Lacrimal, snout, and occiput asquamate. Nine or 10 scales in diagonal from dorsal margin of opercle to origin of spinous dorsal fin. Seven scales in diagonal from lateral line to first ray of soft dorsal fin. Eleven or 12 predorsal scales along midline. Pored scales in lateral line number 58 (holotype) to 60.

Fin spine and ray counts as follows: dorsal V, 21; anal I, 18; pectoral 16; pelvic I, 5; principle caudal rays 8 upper + 7 lower, all branched. Vertebral count 10 precaudal + 19 caudal = 29. Pectoral fin originating just anterior to origin of spinous dorsal fin; when adducted extending posteriorly to vertical through fourth or fifth soft dorsalfin ray. Pelvic fin originating just posterior to vertical through caudal margin of interopercle, and extending to second anal-fin ray when adducted. Origin of spinous dorsal fin located at about level of ventral insertion of pectoral fin. Membrane of fifth (occasionally fourth, with fifth spine more or less free within membrane) dorsal spine attached to first ray of soft dorsal at about 80-90% of length of fifth spine (from fin base). Anal-fin origin at vertical through fourth to fifth ray of soft dorsal fin. Soft dorsal and anal fins terminating at more or less similar point posteriorly. Caudal fin relatively short, shallow, and weakly rounded.

Coloration in life. Coloration of a freshly collected specimen (holotype) shown in Figure 1 . Base body coloration light tan with grayish to golden highlights, becoming lighter and whitish ventrally. Pigmentation pattern on flank characterized by two dark brown stripes that extend the length of the flank, and that are connected by seven or eight similarly colored vertical bands that are constricted medially. A horizontal series of six to nine black blotches present along the lateral midline between the vertical bands. A second series of eight or nine larger black blotches present directly below lower brown stripe at point where vertical connecting bands terminate. Iridescent silvery and bluish spangling present on flank, particularly caudally and also dorsal to upper brown stripe. Snout and eyes with a yellowish hue, fading on lacrimal and cheek. Snout, lacrimal, cheek, and opercle with multiple rows of densely arrayed dark brown blotches of varying size. Smaller spotting present on occiput, nape, and extending length of flank dorsal to upper brown stripe. Pectoral fin light brown and hyaline, with dark brown markings on base. Caudal fin light gray with two prominent black blotches proximal to base, a smaller marking dorsally, and a larger and more elongate marking ventrally. Caudal with two or three bands formed by smaller black blotches on membrane between rays.

Pigmentation pattern in alcohol. Coloration in alcohol shown in Figure 2 . Base coloration light tan ventrally to grayish-tan dorsally. Flank pigmentation characterized by a series of seven or eight more or less hourglassshaped brownish bands, and two horizontal series of prominent black blotches, one about mid-flank and the other located ventral to the midline. No spots in ventral series with lighter encircling ring (ocelli). Brown bands on flank do not extend onto ventrum, which is uniform light tan to whitish in coloration. Snout, lips, cheek, opercle, and anterior portion of nape covered with densely arrayed diffuse brownish blotches varying in size and shape. Posterior portion of nape and predorsal region with smaller dark brown to blackish spotting. Spinous dorsal white to grayish-white, with black pigment on membrane near base between first to third spines. Soft dorsal with two or three rows of black spots on membrane between rays. Anal fin with two rows of black spots on membrane between rays; one row near fin base and another distally in smaller specimens (<60 mm SL), whereas one row mid-fin and another distally in larger specimens (~ 90 mm SL and larger). Pectoral and pelvic fins light tan. Pectoral fin somewhat hyaline, and pelvic opaque. Pectoral with brown blotch on membrane near base ventrally. Caudal fin with two prominent black blotches proximal to base; comprising smaller marking dorsally and larger, more elongate marking ventrally. Caudal with numerous smaller black blotches distally, forming two or three bands on membrane.

Distribution. To date known only from the shallow nearshore reefs of northeastern Madagascar. Very little ichthyological survey work has taken place in the region, and it is likely that the new species is more widely distributed in the Western Indian Ocean. Interestingly, we did not collect any other species of Parapercis on the reefs of the Masoala Peninsula during our survey. Parapercis hexophtalma has been collected from reefs off of southwestern (near Tulear) and northwestern Madagascar (pers. obs.; McKenna & Allen, 2005), P. punctulata has recently been collected from northwestern Madagascar ( McKenna & Allen, 2005), and P. maculata was recently collected in the Baie d’Ambodivahibe in far northeastern Madagascar (Randall, 2008). We did not encounter the new species in any of our previous coastal surveys off of southwestern or eastern Madagascar.

Etymology. Named in reference to the abundant and prominent spotting on the head, flank, and caudal fin (from the Malagasy maramara = spotted).

Remarks and comparisons. Although a taxonomically comprehensive comparative anatomical study across Parapercis has not been undertaken since the work of Cantwell (1964), based on the original descriptions, available images of live, fresh, and preserved specimens, and collections available to us, the pigmentation pattern of P. maramara appears to be unique within the genus. Based on pigmentation pattern on the flank comprising two longitudinal series of dark spots, and a dense pattern of spotting and blotching on the head, as well as other anatomical features, including multiple rows of vomerine teeth and pigmentation pattern on the dorsal and anal fins, the new species appears most closely related to the P. hexophtalma species complex, which also comprises P. pacifica , P. queenslandica , and P. xanthogramma ( Imamura & Yoshino, 2007) . Apart from the new species, Parapercis hexophtalma is the only member of the complex that occurs in the Western Indian Ocean, including Madagascar. Parapercis maramara is easily distinguished from members of the P. hexophtalma species complex by the absence of the large oblong black central caudal-fin marking, which is characteristic of all members of that assemblage ( Fig. 3A ). As far as we can determine based on our survey of the genus, the caudal-fin markings of P. maramara , comprising a smaller oblong black patch dorsally near the base of the fin and a much larger elongate oval or anvilshaped black patch ventrally, in addition to prominent black blotches distributed throughout the fin forming a series of two or three bands distally, are unique ( Figs. 1 and 2 ).

Unlike other members of the P. hexophtalma species complex, less males of P. hexophtalma , the new species lacks brownish spots or blotches between the lower row of black spots ( Figs. 1 and 2 ). The lower series of black blotches in the new species is located at the terminus of the vertical brown flank bands and lack a much lighter encircling ring (i.e., do not form ocelli), which is present in members of the P. hexophtalma species complex ( Fig. 3A ). Brownish spots or blotches are present between the ocelli in all four similar species in the P. hexophtalma species complex, except males of P. hexophtalma (contrary to Imamura & Yoshino [2007], spots and blotches are present in females of this species we have to examine, i.e., from Madagascar; Fig. 3A ). Pectoral-fin ray count in P. maramara is also lower (16 vs. 17–19) than in other members of the P. hexophtalma complex (note: a single individual of P. hexophtalma [n = 19] was observed with 16 rays on the left side, as well as a single individual of P. xanthogramma [n = 20] with 15 on the right).

Other species of Parapercis that have been reported from either Madagascar, the Mascarene Islands, South Africa, or East Africa are P. maculata , P. m a r i t z i, P. millepunctata , P. punctata , P. punctulata , P. robinsoni , P. schauinslandii , and P. xanthozona . A number of features have already been presented in the diagnosis to distinguish the new species from P. maculata , P. punctulata , and P. robinsoni . To these, the following additional distinguishing features should be added to aid in identification. Based on caudal pigmentation pattern, the new species is further distinguished from P. hexophtalma by the absence of an expansive black patch that nearly covers the caudal fin ( Fig. 3A ), from P. m a c u l a t a by the presence of two horizontal series of prominent black spots on the flank (vs. absence of longitudinal series of black markings; Fig. 3 B), from P. punctulata by the presence of large oblong black markings on the caudal fin (vs. pair of small spots; Fig. 3 C), and from P. robinsoni by the absence of a large black marking on the dorsal portion of the caudal fin, with a smaller, more diffuse marking ventrally (vs. larger of two markings located on ventral half of caudal; Fig. 3 D). The new species is further distinguished from P. h e x - ophtalma, P. robinsoni , and P. punctulata by a narrower interorbital width (3.8–6.7% [5.1–6.7% less smallest, 24.5 mm, specimen] vs. 7.1–16.3% HL in P. hexophtalma and P. robinsoni ; 5.1–12.6% HL in P. punctulata [7.8–12.6% less smallest, 37.7 mm, specimen]).

Parapercis maramara , new species, can readily be distinguished from the remaining four species reported to occur in the Western Indian Ocean, but that as far as we can determine have not been taken off of Madagascar, by pigmentation pattern. It is distinguished from P. m a r i t z i, a deepwater species known only from South Africa, by pigmentation pattern on the flank. Parapercis maritzi is characterized by five pairs of blotches or short bands (10 in total) on the upper flank, the posteriormost two blotches coalescing to form a single wide band on the upper half of the caudal peduncle ( Anderson, 1992: fig. 1; Randall & Yamakawa, 2006). The lower half of the flank in P. m a r i t z i is without bands or blotches. Parapercis millepunctata , although described from Sri Lanka and not recorded from as far west as Madagascar (i.e., range extending from Mauritius [Mascarenes] and Maldives, eastward to Pitcairn Group, and northward to southern Japan; Eschmeyer & Fricke, 2011), lacks the upper longitudinal series of black blotches present in P. maramara . Additionally, the black markings in the lower series in P. millepunctata are either absent or indistinct from the dark brown lateral bands within which they are located and that extend fully to the ventrum. Finally, P. millepunctata possesses a characteristic large white central patch on the posterior half of the caudal fin, which is lacking in the new species. Parapercis xanthozona was described from Indonesia, but is reported to range across the Indo-West Pacific, from East Africa to Fiji ( Froese & Pauly, 2011). This species is distinguished from P. maramara by the absence of both the upper and lower longitudinal series of black blotches on the flank, and by the presence of a bright white medial stripe across the caudal fin and caudal peduncle that, although fading anteriorly, may extend the length of the flank. Parapercis xanthozona is also characterized by a large bright white patch on the proximal half of the pelvic fin. Parapercis schauinslandii is another congener reported to have an extensive Indo-West Pacific range, from South Africa and the Seychelles to Hawaii, southern Japan, Western Australia, New Caledonia, and Tonga ( Eschmeyer & Fricke, 2011). This species is distinguished from P. maramara by the absence of the upper and lower longitudinal series of black blotches on the flank, and by the presence of broad red to brownish-red bands on the flank, a lunate (strongly concave) caudal fin with highly produced dorsal and ventral trailing margins, a nearly solid black spinous dorsal fin, large black blotches on the membrane of the soft dorsal about halfway from the base, a vivid red terminal band along the margin of the soft (and sometimes also spinous) dorsal fin, and a pair of red to brownish-red spots at the base of the caudal fin.

Lastly, although P. punctata is reported to occur in the Western Indian Ocean (e.g., see Eschmeyer & Fricke, 2011), Cuvier, in Cuvier & Valenciennes, 1829, stated in the original description of this species that they did not know the provenance of the specimen examined and described. Moreover, in his comprehensive review of Parapercis, Cantwell (1964) considered this species to be of doubtful status. It is impossible to determine from the limited information presented regarding pigmentation pattern and external anatomy in the original description, whether the species is valid.

Figures

FIGURE 1. Parapercis maramara, new species, holotype, AMNH 243241, 108.2 mm SL, Masoala Peninsula, northeastern Madagascar. Photographed immediately following capture to illustrate coloration in life.

FIGURE 2. Parapercis maramara, new species, Masoala Peninsula, northeastern Madagascar. A. Holotype, AMNH 243241, 108.2 mm SL. B. Paratype, AMNH 243242, 95.1 mm SL. Preserved in alcohol.

FIGURE 3. A. Parapercis hexophtalma, UMMZ 185962, 150.0 mm SL, Tulear, southwestern Madagascar. B. Parapercis maculata, BPBM 20453, neotype, 120.7 mm SL, Tuticorin, southwestern India. C. Parapercis punctulata, ANSP 126961, 90.4 mm SL, Seychelles Islands. D. Parapercis robinsoni, USNM 266972, 116.7 mm SL, Bahrain.

Tables

TABLE 1. Morphometric and meristic data for Parapercis maramara, new species.

Percentage of SL
Head length	6	29.9–32.1	29.9	31.3	0.8
Caudal peduncle length	6	8.2–11.1	11.1	9.7	1.1
Caudal peduncle depth	6	8.2–9.2	8.9	8.9	0.4
Pre-dorsal length	6	32.1–33.9	32.1	32.9	0.7
Dorsal fin base length	6	57.1–63.5	62.2	61.4	2.4
Pre-anal length	6	41.6–54.2	48.1	49.6	4.7
Anal fin base length	6	40.8–45.8	44.6	43.2	1.9
Pectoral fin length	6	18.9–23.3	18.9	21.1	1.8
Pre-pelvic length	6	23.3–26.6	25.1	25.2	1.2
Pelvic fin length	6	24.6–31.9	24.6	27.6	2.4
Caudal fin length	6	19.0–26.9	19.0	22.1	2.9