Revision of southern African species of the anglerfish genus Chaunax (Lophiiformes: Chaunacidae), with descriptions of three new species Ho, Hsuan-Ching Ma, Wen-Chun Zootaxa 2016 4144 2 175 194 [151,402,359,385] Actinopterygii Chaunacidae Chaunax Animalia Lophiiformes 1 176 Chordata species atimovatae sp. nov.     Holotype.  SAIAB81724(149), 26°10.2'S, 33°17.6'E, off Mozambique,  505–509 m,  30 Sep. 2007.    Paratypes. 13 specimens, 104–211 mmSL. AMS I.28137-006 (2, 108–111), 22°19' 06"S, 43°06' 06"E, NW Tulear, Madagascar,  330–335 m,  2 Dec. 1988.   MNHN1977-0030(1, 119), 22°17'10"S, 43°4'1"E, Madagascar,  450 m,  29 Nov. 1973.   MNHN1977-0036(1, 105), 12°40'59"S, 48°16'1"E, Madagascar,  314 m,  Apr. 1971.   SAIAB3306(1, 118), KwaZulu-Natal, South Africa,  238 m,  10 Jan. 1973.   SAIAB11484(1, 139), off Natal, KwaZulu- Natal, South Africa,  146 m,  11 Jan. 1973.   SAIAB34164(1, 211), Maputo, southern Mozambique,  Dec. 1987.   SAIAB74601(3, 104–131), 18°2.1'S, 37°37.2'E, Beira, northern Mozambique,  162–200 m,  12 Aug. 2002.   SAIAB81904(1, 106), 25°5.5'S, 35°18.4'E, off Mozambique,  347–353 m,  7 Oct. 2007.   USNM307534(2, 122–162), 22°19'51"S, 43°06'06"E, northern Tulear, Madagascar,  330–335 m, Vityaz, cr. 17, st. 2644,  2 Dec. 1988.     Non-types(*indicates specimens with meristic data taken).  Madagascar:  MNHN1977-0023(2, 69.6–82.1), 25°28'59"S, 46°46'1"E,  350–360 m,  30 May 1973. MNHN 1977-0027 (1, 72.8), 23°36'0"S, 43°31'5"E, 395–410 m, 26 Feb. 1973. MNHN 1977-0029 (3, 43.1–111), 22°18'0"S, 43°4'8"E, 400 m, 27 Nov. 1973. MNHN 1977-0033 (2, 35.9–67.8), 18°55'01"S, 43°55'59"E, 250 m, 24 Nov. 1973.   MNHN1994-0312(1, 107), 20°57' 0"S, 55°13'59"E, RéunionIsland,  450–580 m,  1 Sep. 1982.   MNHN1994-0314(1, 76.4), 20°49' 59"S, 55°34'59"E, RéunionIsland,  605–620 m,  4 Sep. 1982. MNHN 1994-0315 (2, 67.0–131), 20°57'0"S, 55°13'59"E, 300–410 m, 9 Sep. 1982. MNHN 2013-1225 (2, 56.7–83.8), 22°14'6"S, 43°4'5"E, 470–475 m, 2 Dec. 1973. MNHN 2014-1107 (1, 123), 15° 35'8.4"S, 45°41'55.8"E, 197–217 m, 10 Jul. 2009. MNHN 2014-1169 (5, 39.1–71.1), 15°34'45"S, 45°42'30"E, 227–283 m, 10 Jul. 2009. MNHN 2014-1211 (1, 87.6), 15°33'37.2"S, 45°42'5.4"E, 350–580 m, 11 Jul. 2009. MNHN 2014-1264 (2, 31.3–35.2), 14°29'3"S, 47°26'50.4"E, 257– 251 m, 6 Jul. 2009. MNHN 2014-1602 (1, 68.5), MNHN 2014-1628 (1, 37.7), 12°30'16.8"S, 48°23'12"E, 301– 298 m, 27 Jun. 2009. MNHN 2014-1664 (5, 68.8– 101), 15° 34'12.612"S, 45°43'10.812"E, 287–450 m, 10 Jul. 2009. MNHN 2014-1699 (1, 39.5), 14°51'56.4"S, 46°56'34.2"E, 236–297 m, 13 Jul. 2009. MNHN 2014-1719 (1, 95.5), 13°23'31.2"S, 47°58'4.2"E, 3 Jul. 2009. MNHN 2014-1756 (2, 52.6–79.7), 14°29'39.6"S, 47°26'27"E, 379– 332 m, 14 Jul. 2009. MNHN 2014-1764 (1, 87.8), 12°32'7.8"S, 48°12'27"E, 485– 468 m, 27 Jun. 2009. MNHN 2014-1781 (4, 38.7–105), MNHN 2014-1815 (1, 95.4), MNHN 2014-1838 (2, 33.5–60.1), MNHN 2014-1840 (1, 40.9), 14°49'53.4"S, 46°59'9"E, 446– 340 m, 7 Jul. 2009. MNHN 2014-1846 (1, 110), 12°46'18"S, 48°12'48.6"E, 355–380 m, 2 Jul. 2009. MNHN 2014-1855 (3, 31.3–32.7), 14°30'10.8"S, 47°26'35.4"E, 325– 274 m, 6 Jul. 2009. MNHN 2014-1878 (2, 22.0–22.9), 14°48'12"S, 46°59'9"E, 620–637 m, 7 Jul. 2009. MNHN 2014-1900 (1, 69.3), 15°25'4.8"S, 46°0'16.2"E, 313– 274 m, 12 Jul. 2009. MNHN 2014-1962 (2, 38.0–40.0), 12°31'18.6"S, 48°19'33"E, 346–376 m, 27 Jun. 2009. MNHN 2014-2169(1, 79.2), 12°43'15.6"S, 48°13'1.8"E, 442–491 m, 29 Jun. 2009.   SAIAB189524(3, 31.5–45.2), 25°28'S, 44°25'E, Sud-Est Pointe Barrow,  203–210 m,  10 May 2010, coll. M. Lee& S. Ribes.   SAIAB189525(1, 31.5), 25°03'S, 43°59'E, Sud-Ouest Pointe Barrow,  300–309 m,  11 May 2010, coll. M. Lee& S. Ribes. *  USNM307546(2, 60.0– 100), 22°19'30"S, 43°03'12"E, northern Tulear,  450–500 m, Vityaz, Cr.17, st. 2645,  2 Dec. 1988.   South Africa: *  SAIAB4709(1, 76.0), 34°02'S, 25°42'E, Algoa Bay, Eastern Cape,  27 May 1970. *  SAIAB4710(1, 150), 32°09'S, 29°00'E, Xora Rivermouth, Eastern Cape,  Aug. 1947. *  SAIAB16533(1, 77.0), 29°13'S, 31°29'E, Tugela Rivermouth, KwaZulu-Natal, 1920. *  SAIAB49355(1, 84.0), 35°1.3'S, 19°44'E, off west coast,  145 m,  5 Jul. 1986.   Mozambique: *  SAIAB28513(1 of 3, 104), S of Maputo,  Dec. 1987. No data: *SAIAB 4713 (1 of 5, 125).   Diagnosis.A species of the  Chaunax abeispecies group with the following combination of characters: mixture of bifurcate and simple spinules on dorsal surface; when fresh, dorsal surface with green spots surrounded by yellow, mostly inter-connected; when preserved, dorsal surface with brown spots loosely arranged; 9 or 10 gill rakers on second gill arch; and usually 12 or 13 neuromasts in pectoral series and 34–36 inlateral-line proper.   FIGURE 1.  Chaunax atimovatae  sp. nov.A. SAIAB 81724, holotype, 149 mm SL. B. SAIAB 81904, paratype, 106 mm SL. C. Uncatalogued specimen, ca. 21 cm in total, off Maputo, southern Mozambique.   FIGURE 2.Underwater photograph of  Chaunax atimovatae sp. nov., 29°40.616’S, 31°31.223’E, 220 m.   FIGURE 3.Distribution map of  Chaunax atimovatae  sp. nov.(red squares),  Chaunax heemstraorum  sp. nov.(green circles) and  Chaunax hollemani  sp. nov.(black star).   TABLE 1.Morphometric đata of three new  Chaunaxspecies đescribeđ in present stuđy. H=holotype.   Chaunax atimovatae Chaunex heemstraorum Chaunax hollemani   sp. nov. sp. nov. sp. nov. H TypesH All typesH All types (mm) 149 104.0$216.0 (n=13) 163 95.1$163.0 (n=9) 194.5 114.9$178.7 (n=5) SL Mean (Range) SD Mean (Range) SD Mean (Range) SD Description.Morphometric and meristic data are provided in Tables 1–3; data for holotypeis provided below followed by range for typesin parentheses.    Heađ length 39.6 40.6 (38.1$42.7) 1.2 39.3 40.4 (38.8$43.2) 1.8 40.3 39.7 (39.1$40.5) 0.6  Heađ wiđth 17.4 17.2 (16.1$19.5) 0.9 15 16.0 (15.0$16.9) 0.8 17.2 17.2 (16.0$18.5) 0.9  Pre-preopercular length 26.8 27.5 (26.0$29.4) 1 26.1 27.5 (25.6$30.2) 1.8 27.1 27.5 (26.8$28.5) 0.7  Eye điameter 6 7.3 (6.0$8.5) 0.7 8.9 9.2 (8.5$10.9) 1 7.6 7.7 (7.3$8.4) 0.4  Illiical length 4.7 4.8 (4.3$5.7) 0.4 3.7 2.8 (2.5$3.7) 0.5 4.6 4.9 (4.2$5.6) 0.5  Illicial trough length 7.4 7.5 (6.2$9.2) 0.8 6.1 5.5 (4.7$6.7) 0.8 7.4 7.5 (7.1$7.9) 0.4  Pre-illicial trough length 5.4 5.0 (3.8$6.9) 0.9 5.8 5.7 (4.7$6.0) 0.6 3.8 3.4 (2.2$3.9) 0.7  Upper jaw length 19.5 20.3 (19.4$21.6) 0.9 18.4 19.2 (18.4$20.3) 0.8 21.7 21.6 (20.8$23.0) 0.9  Pre-đorsal length 51 50.5 (45.8$53.2) 2.5 46.6 48.4 (46.6$50.0) 1.3 46.9 48.0 (46.0$50.0) 1.6  Pre-gill opening length 66.4 64.1 (61.2$67.4) 2.1 60.1 57.8 (52.9$60.4) 3.3 62.9 62.1 (60.7$63.9) 1.5  Tail length 1 30.9 31.2 (27.3$33.9) 2 34.4 36.3 (34.4$38.8) 2 16.0 17.7 (15.2$19.4) 2.0  Tail length 2 16.1 17.2 (14.6$20.9) 1.7 19.6 19.3 (17.9$21.5) 1.4 28.9 30.3 (28.9$31.8) 1.2  Tail length 3 15.4 14.5 (13.2$16.0) 1 17.8 18.5 (17.0$20.8) 1.5 14.5 14.9 (13.7$15.9) 0.9  Cauđal-peđuncle đepth 8.1 8.1 (7.7$8.6) 0.3 8 7.7 (7.4$8.1) 0.3 8.4 8.7 (8.4$9.0) 0.2  Cauđal-fin length 28.2 30.2 (28.0$32.1) 1.5 30.7 29.5 (28.5$30.7) 0.9 27.8 29.9 (27.8$31.5) 1.4 Dorsal-fin rays III, 12; pectoral-fin rays 13 (13, rarely 12); anal-fin rays 7; caudal-fin rays 9. Head length 2.5 (2.3–2.6) in SL; head width 5.7 (5.6–6.2) in SL and 2.3 (2.3–2.5) in HL; pre-dorsal length 1.9 (1.9–2.2) in SL; pregill opening length 1.5 (1.5–1.6) in SL; pre-preopercular length 3.7 (3.5–3.9) in SL and 1.5 (1.4–1.6) in HL; upper jaw 5.1 (4.6–5.0) in SL and 2.0 (1.9–2.1) in HL; illicial length 8.4 (7.0–9.4) in HL; eye diameter 6.6 (5.2–6.6) in HL; post-dorsal-fin length 6.2 (5.4–6.4) in SL and 2.5 (2.0–2.8) in HL; post-anus length 3.2 (3.0–3.7) in SL and 1.3 (1.1–1.5) in HL; post-anal-fin length 6.5 (6.5–7.6) in SL and 2.6 (2.6–3.1) in HL; caudal-peduncle depth 4.9 (4.8– 5.3) in HL; caudal-fin length 3.5 (3.1–3.6) in SL and 1.4 (1.3–1.5) in HL. Head globular, skull elevated posteriorly; trunk and tail robust, weakly compressed, tapering posteriorly; ventral surface of belly flattened; skin thick, loose and flaccid; caudal peduncle short, slightly depressed. Eyes rounded, directed laterally, covered by dermal membrane, broadly connected to adjacent skin, forming clear window; interspace between eyes broad, convex. Illicium relatively slender, its length less than eye diameter; esca depressed, forming a large central plate bearing many dark brown cirri; second dorsal-fin spine close to illicium, embedded under skin and not detectable externally; third dorsal-fin spine situated at about midpoint of pre-dorsal distance, embedded beneath skin. Illicial trough oval, slightly concave, uniformly narrow, its length about twice its width. Origin of soft dorsal fin slightly behind middle of body; pectoral fin emerging laterally near middle of body, slightly anterior to vertical through gill opening; pelvic fin on breast, well anterior to pectoral fin; anus situated about three-fourths of way along body; anal-fin origin about four-fifths of way along body, tip nearly reaching caudal-fin base when depressed. Both nostrils anterior to eye; anterior nostril surrounded by fleshy membrane, its rim taller posteriorly than anteriorly; posterior nostril a circular depression; mouth wide, superior, its opening nearly vertical; lower jaw robust, protruding slightly in front of upper jaw; maxilla tapering, narrow dorsally, broadly expanded ventrally; blunt spine on symphysis of lower jaw. Broad transparent membrane on first gill arch; first ceratobranchial well-connected to opercular wall, and first epibranchial entirely free of it; gill filaments present on second to fourth gill arches, two rows of gill filaments on second and third gill arches, single row of gill filaments on fourth gill arch; filaments on inner rows of third and fourth gill arches about two-thirds length of filaments on other arches; inner surface of fourth gill arch completely connected to body. Single row of 14 (12–15) rakers on 1st gill arch, 3 (3–4) on upper limb and 11 (8–11) on lower limb, 10 (9 or 10) rakers on outer row of 2nd arch, 9 (9 or 10) rakers on outer row of 3rd arch, and single row of 8 (7 or 8) rakers on 4th arch. Interspaces of lateral-line neuromast complex slightly longer than width of complex; 3 (2–4, usually 3) pairs of short spines bridging each neuromast. Lateral-line neuromast counts: supraorbital (AB) 11 (11 or 12); premaxillary (AC) 8; upper preopercular (BD) 2; infraorbital (CD) 7 (6 or 7); lower preopercular (DG) 3; mandibular (EF) 6 (5 or 6); hyomandibular (FG) 3; pectoral (GH) 14 (11–14); anterior lateral-line proper (BB') 4 (4 or 5); supratemporal (BB) 6; and lateral-line proper (BI) 35 (32–37), including 3 (2 or 3) on caudal-fin base. Dorsal surface covered by simple, stout spinules, except for eye window, lips, distal fifth of dorsal surface and entire ventral surface of pectoral fin, distal half of dorsal surface and entire ventral surface of pelvic fin, entire anal fin and its base, membranes of dorsal fin, anus, and caudal-fin rays. Broad band of 4–9 rows of spinules in front of illicial trough (3 or 4 rows in specimens smaller than 100 mmSL). Ventral surface covered by slightly shorter, firm spinules. Jaws and body margin along lateral line densely covered with simple, stout cirri; cirri absent from head, jaws and ventral surface.  Coloration.When fresh, background of dorsal surface reddish gray dorsally and pale ventrally; small irregular green spots surrounded by yellow, mostly inter-connected on dorsal surface; bright white flap-like cirri along margins of body and jaws. When preserved, dorsal surface covered by small irregular gray spots on uniformly grayish background; pale ventrally. Cirri on esca dark brown. Gill chamber, buccal cavity and gill rakers pale; peritoneum black with small pale patches.  Size.The largest known specimen has a length of 211 mmSL.   Distribution.Known from the Western Indian Ocean off Madagascar, Mozambiqueand South Africa, westernmost record appears to be on the southern coast of South Africa( Fig. 3), collected at depths 145– 637 m.   Etymology.The specific name is from a field trip of the "Atimo Vatae" (Madagascan for "deep south"), a trawler from Fort Dauphin, from which many Madagascan specimens were collected, including those in the typeseries.   TABLE 2.Distribution of gill-rakers counts of  Chaunaxspecies recognized from the study area. Star indicates the value of holotype. Rakers of 1st gill arch n 11 12 1314 15 16 17 18 19 20  C. apus5 1 3 2  C. atimovatae1 4 1 8 4* 1  C. flammeus1 1   C. heemstraorum31 4 20 7*  C. hollemani5 1* 3 1  C. penicillatus5 1 3 1    C. russatus2 2  C. suttkusi7 1 3 3 Rakers of 2nd gill arch n 8 9 1 0 1 1 1 2 1 3 1 4 1 5  C. apus6 5 1  C. atimovatae 14 6 8*   C. flammeus1 1   C. heemstraorum 31 3 22* 6  C. hollemani5 5*   C. penicillatus5 2 3   C. russatus2 2  C. suttkusi7 2 4 1 Rakers of 3rd gill arch n 8 9 1 0 1 1 1 2 1 3 1 4  C. apus6 5 1  C. atimovatae14 10* 4   C. flammeus1 1   C. heemstraorum 31 21 10*   C. hollemani5 1 4*   C. penicillatus5 2 3   C. russatus2 2  C. suttkusi7 2 4 0 1 Rakers of 4th gill arch n 7 8 9 1 0 1 1 1 2????  C. apus6 4 2   C. atimovatae 14 3 11*   C. flammeus1 1   C. heemstraorum 31 10 20* 1   C. hollemani5 1 4*   C. penicillatus5 5   C. russatus2 2  C. suttkusi7 1 5 1   Remarks.  Chaunax atimovatae  sp. nov.is most similar to  Chaunax abeifrom the northwestern Pacific region in having bifurcated spinules mixed with simple ones on its dorsal surface, but these bifurcated spinules are much less numerous and quite scattered. The background of body of  C. atimovatae  sp. nov.is grayish with numerous tiny black melanophores (visible under magnification) whereas that of  C. abeiis pale and lacks this pigmentation. The new species is also similar to  C. breviradiusfrom the South ChinaSea region in having a similar coloration when fresh; it differs from  C. breviradiusin having bifurcated spinules mixed with simple ones on dorsal surface (vs. simple spinules only), and relatively more neuromasts in the lateral-line proper (BI) (usually 34–36 vs. 30–32). 1288101093 [151,1260,502,527] 2007-09-30 SAIAB Mozambique 507 -26.17 Mozambique 124 33.293335 1 176 SAIAB 81724 1 holotype 1288101088 1988-12-02 Madagascar 333 -22.318335 Tulear 21 43.101665 1 176 13 paratype 1288101095 1973-11-29 MNHN Madagascar 450 -22.28611 Madagascar 21 43.066944 1 176 MNHN 1977-0030 1 paratype 1288101107 [365,1338,610,635] 1971-04 MNHN Madagascar 314 -12.683056 Madagascar 21 48.266945 1 176 MNHN 1977-0036 1 paratype 1288101094 1973-01-10 SAIAB South Africa 238 KwaZulu-Natal 1 176 SAIAB 3306 1 KwaZulu-Natal paratype 1288101083 1973-01-11 SAIAB South Africa 146 Natal Natal 1 176 SAIAB 11484 1 paratype 1288101099 [651,1432,682,707] 1987-12 SAIAB Mozambique Maputo 1 176 SAIAB 34164 1 Maputo paratype 1288101090 [151,1342,718,743] 2002-08-12 SAIAB Mozambique 181 -18.035 Beira 127 37.62 1 176 SAIAB 74601 1 paratype 1288101086 2007-10-07 SAIAB Mozambique 350 -25.091667 Mozambique 124 35.306667 1 176 SAIAB 81904 1 paratype 1288101087 1988-12-02 USNM Madagascar 333 -22.330833 northern Tulear 21 43.101665 Vityaz 1 176 USNM 307534 1 paratype 1288101105 1973-05-30 MNHN Madagascar 355 -25.483055 Non-types 21 46.766945 1 176 MNHN 1977-0023 1 1288101098 1982-09-01 MNHN Réunion 515 -20.95 Reunion 21 55.233055 1 176 MNHN 1994-0312 1 1288101096 1982-09-04 MNHN Réunion 613 -20.833055 Reunion 21 55.583054 1 176 MNHN 1994-0314 1 1288101085 2010-05-10 SAIAB M. Lee & S. Ribes Réunion 207 -25.466667 Sud-Est Pointe Barrow 1246 44.416668 1 176 SAIAB 189524 1 1288101092 2010-05-11 SAIAB M. Lee & S. Ribes Réunion 305 -25.05 Sud-Ouest Pointe Barrow 1248 43.983334 1 176 SAIAB 189525 1 1288101102 1988-12-02 USNM Vityaz & Cr. Réunion 475 -22.325 northern Tulear 21 43.053333 1 176 USNM 307546 1 1288101091 1970-05-27 SAIAB South Africa -34.033333 Algoa Bay 1202 25.7 1 176 SAIAB 4709 1 Eastern Cape 1288101101 1947-08 SAIAB South Africa -32.15 Xora River 1212 29.0 1 176 SAIAB 4710 1 Eastern Cape 1288101100 1986-07-05 SAIAB South Africa 145 -35.021667 Tugela River 764 19.733334 1 176 SAIAB 16533, SAIAB 49355 1 KwaZulu-Natal 1288101097 [221,1001,1798,1823] 1987-12 SAIAB Mozambique Maputo 1 176 SAIAB 28513 1 Maputo