Hemibates koningsi spec. nov: a new deep-water cichlid (Teleostei: Cichlidae) from Lake Tanganyika Author Schliewen, Ulrich Kurt text Zootaxa 2017 2017-08-28 4312 1 92 112 journal article 32280 10.11646/zootaxa.4312.1.4 3f438ffd-0acb-4805-85cb-096bf46c56f7 1175-5326 892855 Hemibates koningsi sp. nov. Hemibates sp. “ stenosoma zambia” Konings 2015 Holotype . ZSM 45056 ( 1, 192.2 mm SL, male, DRC-2012/3138) Zambia , Northern ProvinCe , Lake Tanganyika , Mpulungu basin, no exaCt loCality data available, purChased in Inguenyo fish market, Mpulungu (-8.73°/31.11°), F. SChedel, 28.VIII.2015 Paratypes . BMNH 2016.9 .1.2. ( 1, 153.5 mm SL, males, DRC-2012/3242), Zambia , Northern ProvinCe , Lake Tanganyika , Mpulungu basin, no exaCt loCality data available, purChased in Inguenyo fish market, Mpulungu (- 8.73°/31.11°), F. SChedel, 05.IX.2015 . MRAC 2017-005 -P-0001 ( 1, 154.2 mm SL, males, DRC-2012/3103), Zambia , Northern ProvinCe , Lake Tanganyika , Mpulungu basin, no exaCt loCality data available, purChased in Inguenyo fish market, Mpulungu (-8.73°/31.11°), F. SChedel, 23.VIII.2015 . ZSM 44570 ( 1, 162.5 mm SL, male, DRC-2012/3186), Zambia , Northern ProvinCe , Lake Tanganyika , Mpulungu basin, no exaCt loCality data available, purChased in Inguenyo fish market, Mpulungu (-8.73°/31.11°), F. SChedel, 01.IX.2015 . ZSM 44565 ( 1, 150.8 mm SL, male, DRC-2012/3243), Zambia , Northern ProvinCe , Lake Tanganyika , Mpulungu basin, no exaCt loCality data available, purChased in Inguenyo fish market, Mpulungu (-8.73°/31.11°), F. SChedel, 05.IX.2015 . ZSM 44566 (1, 141, 9 mm SL, female, DRC-2012/3275) Zambia , Northern ProvinCe , Lake Tanganyika , Mpulungu basin, no exaCt loCality data available, purChased in Inguenyo fish market, Mpulungu (-8.73°/31.11°), F. SChedel, 07.IX.2015 . ZSM 44571 (1, 99.9 mm SL, male, DRC-2012/3179) Zambia , Northern ProvinCe , Lake Tanganyika , Mpulungu basin, no exaCt loCality data available, purChased in Inguenyo fish market, Mpulungu (-8.73°/31.11°), F. SChedel, 31.VIII.2015 . ZSM 44677 ( 1, 106.1 mm SL, male, DRC-2012/3139) purChased with holotype. ZSM 44678 ( 1, 217.4 mm SL, males, DRC-2012/3104), Zambia , Northern ProvinCe , Lake Tanganyika , Mpulungu basin, no exaCt loCality data available, purChased in Inguenyo fish market, Mpulungu (-8.73°/31.11°), F. SChedel, 23.VIII.2015 . Differential diagnosis . Adult males of the Hemibates koningsi Can be distinguished from H. stenosoma , the only Congener, by a flank Color pattern of four to seven blaCk vertiCal bars (two to three flank-sCales wide) on the anterior flank region and five horizontal bands (one to two, rarely three flank-sCales high) on the posterior flank region vs. blaCk blotChes of variable number, size and shape at the anterior part of the flanks and five (rarely four) horizontal bands in H. stenosoma . They Can further be distinguished by the fourth horizontal band (Counted from dorsal to ventral) starting below peCtoral fin base sometimes before and extending to Caudal fin base in H . koningsi vs. normally ending well before Caudal fin base in Hemibates stenosoma . Moreover, both speCies exhibit a blaCk band on dorsal fin membrane along the dorsal-fin base, whiCh is however wider and more pronounCed in H . koningsi males. Females as well as males of the new speCies Can be unambiguously separated from H. stenosoma by their longer lower pharyngeal jaw (37.6–38.2% HL, N=4 vs 27.8–32.5% HL, N=7) with a CharaCteristiCally Curved keel (distinCtive bulge ventrally of the keel), whiCh Contrasts with the more or less straight keel of H. stenosoma , whiCh is only rounded towards the tip ( Fig. 4 ). MeristiCs of H . koningsi overlap with those of Hemibates stenosoma , but the new speCies on average has fewer gill rakers on the first gill arCh (33–37 vs. 35–43). Ranges of morphometriC measurements of the new speCies overlap with those of H. stenosoma , but H . koningsi tend to have longer lower jaws (44.0–47.1% HL vs 39.22–45.9% HL), longer upper lips (32.6–36.7% HL vs 27.6– 33.6% HL) and longer lower lips (34.5–37.8% vs 30.1–36.0% HL). FIGURE 3. Hemibates koningsi sp. nov. A. Holotype, shortly after collection B. Holotype, ZSM 45056, DRC-2012/3138, 192.2 mm SL; Zambia, Northern Province, Lake Tanganyika, Mpulungu basin, no exact locality data available, purchased in Inguenyo fish market, Mpulungu C. radiograph of holotype. Description. MorphometriC measurements and meristiC CharaCters based on type speCimens. Values and Corresponding ranges presented in Table 5 . For general appearanCe see Figs. 3 and 4 . Maximum reCorded total length of wild Caught speCimen (male) 262.2 mm with Corresponding SL of 217.4 mm . Laterally Compressed and relatively deep-bodied speCies with maximum body depth (31.6–34.8% SL) slightly behind dorsal-fin origin, deCreasing towards Caudal pedunCle. Ratio of Caudal pedunCle length to depth: 1.55–1.92. Head length about one third of standard length. Dorsal head profile moderately ConCave with slightly visible premaxillary pediCel prominenCe. No nuChal gibbosity present. Eyes round, eye diameter less than one third of head length (26.2–31.1% HL) and larger than interorbital width. Jaws isognathous to slightly prognathous, lower jaw slightly protruding and Comparatively narrow. Mouth strongly oblique. Posterior tip of maxilla not reaChing level of eye. Lower lip anteriorly wider than upper lip. Lips not notiCeably enlarged or thiCkened. LaChrymal with five sensory-Canal pores. Two separate lateral lines. TABLE 5. Measurements & counts for holotype and paratypes of Hemibates koningsi sp. nov.

Measurements	holotype	holotype +	paratypes
min	Max	SD	n
Total length (mm)	235.5	124.6	262.2	9
Standard length SL (mm)	192.2	99.9	217.4	9
Head length HL (mm)	63.17	33.1	72.3	9
% HL
Interorbital width	21.1	20.9	23.5	0.9	9
Preorbital width	29.7	27.9	31.5	1.2	9
Horizontal eye length	26.5	26.2	31.1	1.4	9
Snout length	32.6	31.5	33.3	0.6	9
Internostril distance	16.5	15.2	18.4	0.9	9
Cheek depth	26.7	22.6	26.7	1.3	9
Upper lip length	36.1	32.6	36.7	1.2	9
Lower lip length	35.8	34.5	37.8	1.1	9
Lower lip width	24.8	21.7	26.6	1.5	9
Lower jaw length	46.0	44..0	47.1	0.8	9
Lower pharyngeal jaw length	-	37.6	38.2	0.3	4
Lower pharyngeal jaw width	-	18.3	21.0	1.2	4
Width of dentigerous area of Lower pharyngeal jaw	-	12.9	14.7	0.8	4
% SL
Predorsal distance	36.3	35.2	37.4	0.8	9
Dorsal fin base length	50.2	47.6	50.5	1.0	9
Last dorsal fin spine length	14.8	11.7	16.2	1.4	9
Anal fin base length	19.0	17.9	19.8	0.7	9
Third anal fin spine length	10.0	9.6	12.3	1.0	9
Pelvic fin length	28.4	24.4	39.1	4.3	9
Pectoral fin length	29.5	27.5	30.7	1.1	9
Caudal peduncle depth	11.2	10.7	12.0	0.4	9
Caudal peduncle length	20.0	18.4	20.9	0.9	9
Body depth (pelvic fin base)	34.2	31.6	34.8	1.1	9
Preanal length	62.5	61.8	67.0	1.7	9
Anus-anal fin base distance	3.3	1.8	3.9	0.6	9
Interpectoral width	12.0	9.7	12.4	0.9	9

......continued on the next page TABLE 5. (Continued)

Counts	holotype	holotype + paratypes
min Max SD	n
Dorsal fin spines	15	14(3); 15(6)	9
Dorsal fin rays	14	13(1); 14(4), 15(4)	9
Anal fin spines	3	3(9)	9
Anal fin rays	13	12(5); 13(2); 14(2)	9
Pelvic fin spines	1	1(9)	9
Pelvic fin rays	5	5(9)	9
Pectoral fin rays	13	13(6); 14 (3)	9
Upper procurrent caudal fin rays	9	8(1); 9(7); 10(1)	9
Lower procurrent caudal fin rays	10	9 (2); 10(7)	9
Caudal fin rays	34	33(1); 34(1); 35(6); 36(1)	9
Scales (horizontal line)	62	60(1); 61(3); 62(2); 63(1); 65(1)	8
Upper lateral line	65	58(1); 62(1); 63(1); 67(2); 69(2)	8
Lower lateral line	39	35(1); 36(4); 39(2); 43(1)	8
Circumpeducular	28	28(3); 30(4); 32(1)	8
Series of Scales on check	3	1(1); 2(5); 3(3)	9
Scales (horizontal line) on operculum	5	3(2); 4(2); 5(3); 6(2)	9
Scales between lateral line and dorsal fin origin	7	7(4); 8(5)	9
Scales between upper lateral line and last dorsal fin spine	5	5 (8)	8
Abdominal vertebrae	16	16(8);17(1)	9
Caudal vertebrae	18	17(2); 18(7)	9
Total number of vertebrae	34	33(1); 34(8)	9
Teeth in upper outer row	69	67(1); 69(1); 70(1); 72(1); 75(1); 77(1); 83(1); 84(2)	9
Teeth in lower outer row	58	41(1); 48(1); 51(1); 52(1); 56(1); 58(1); 59(1); 64(1); 67(1)	9
Gill rakers (ceratobranchial)	28	26(4); 27(2); 28(3)	9
Gill rakers (angle + epibranchial)	8	7(4); 8(3), 9(2)	9

Squamation. Flanks and dorsum Covered with relatively large weakly Ctenoid sCales. Anterior base of some flank sCales Covered with minute, CyCloid or weakly Ctenoid sCales ( Fig. 4 ). CyCloid sCales of ventral belly region smaller than on flanks. CyCloid Chest sCales, espeCially ventrally, smaller than flank sCales; smallest sCales on isthmus. Snout sCaleless. Interorbital, nape and oCCipital region with slightly smaller sCales than on flanks. Cheeks Covered by one to three rows of CyCloid sCales about size of ventral belly sCales. OperCulum Covered with ovoid and CyCloid sCales of variable size (small to about size of flank sCales). Three to six sCales on vertiCal line starting form edge of posterior-dorsal angle of operCulum to anterior edge of operCulum. Caudal fin sCales small and beComing minute more Caudally; sCaled area may extend to more than half of the Caudal fin. Upper lateral line with 58–69 sCales, lower lateral line with 35–43 sCales and horizontal line with 60–65 sCales plus one to three pored sCales on Caudal fin. Upper and lower lateral lines separated by three sCales. Seven to eight sCales between dorsal-fin origin and upper lateral line. 28–32 sCales around Caudal pedunCle. Jaws, dentition and gill rakers. Anterior teeth of outer row of lower and upper jaw uniCuspid and widely and irregularly set. Teeth beComing smaller, less widely spaCed and more regularly set towards mouth angle. Individual uniCuspid teeth Comparatively slender and slightly reCurved with aCutely pointed brownish Crowns. Outer row of upper jaw with 67–84 teeth and outer row of lower jaw with 48–67 teeth in speCimens between 99.9 and 217.4 mm SL. Neither number of teeth in the lower jaw nor of upper jaw is signifiCantly Correlated with SL (Pearson Correlation CoeffiCient: r-sCore for SL against number of teeth in the lower jaw -0,24593, p = 0.174533; r-sCore for SL against number of teeth in upper jaw -0,32063, p= 0.399645). UniCuspid teeth of inner tooth rows smaller than those of outer rows, upper jaw with one or two inner tooth rows and lower jaw with single inner tooth row. Lower pharyngeal bone ( Fig. 5 ) about 1.8 to 2.1 times longer than broad. Width of dentigerous area of lower pharyngeal bone 0.3 to 0.4 times length of lower pharyngeal bone length, with 26–27 teeth along posterior margin of dentigerous area. Individual teeth slender, beveled (i.e. biCuspid), with its labial side featuring several Cusp protuberanCes, and always with a pointed major Cusp. Lower pharyngeal jaw teeth largest along posterior margin of dentigerous area and beComing smaller towards the keel, anterior teeth slender and beveled sometimes uniCuspid. Keel of lower pharyngeal bone CharaCteristiCally Curved with distinCtive bulge on ventral keel. Total gill raker Count 33–37, with six to eight epibranChials, one angle (Cartilaginous plug) and 26–28 CeratobranChial rakers. Individual rakers long and slender. Smallest rakers anterior of CeratobranChial inCreasing in size towards Cartilaginous plug. Gill raker of Cartilaginous plug smaller than neighboring CreatobranChial rakers. EpibranChial rakers deCreasing in size dorsally. FIGURE 4. Hemibates koningsi sp. nov. A. Arrangement and morphology of oral jaw teeth (Holotype, ZSM 45056, DRC- 2012/3138, 192.2 mm SL) B. Flank scales with minute, cycloid to weakly ctenoid scales on their anterior basis (holotype) C. Female Hemibates koningsi sp. nov , shortly after collection, ZSM 44566, DRC-2012/3275 141.9 mm SL; Zambia, Northern Province, Lake Tanganyika, Mpulungu basin, no exact locality data available, purchased in Inguenyo fish market, Mpulungu. Fins. Dorsal fin XIV-XV, 13–15. First dorsal-fin spine shortest. Dorsal-fin base length 47.6–50.5% SL. Dorsalfin rays of mature males extending to around level of Caudal fin base whereas in young males (< 106 mm SL) and in females rays only extending to around the first half of Caudal pedunCle. Anal-fin rays not reaChing Caudal fin base in young males and in females, but reaChing about first half of Caudal pedunCle. Anal fin III, 12–14; third analfin spine longest. Anal-fin base shorter (17.9–19.8% SL) than peCtoral-fin length (27.5–30.7% SL). PeCtoral fin with 13 or 14 rays; longest peCtoral-fin ray (fifth ray, Counted from dorsally to ventrally) more or less slightly exCeeding level of anus. PelviC-fin base posterior to peCtoral-fin base, separated by roughly three to five flank sCale widths. PelviC fin with one spine and five rays; first pelviC-fin ray longest, variably elongated in mature males, either terminating slightly before anal-fin origin or exCeeding it slightly; in females and young males only slightly exCeeding level of anus. Caudal-fin outline furCate and Composed of 34–36 rays (16 prinCipal Caudal fin rays and 18 to 20 proCurrent Caudal fin rays). Vertebrae and caudal fin skeleton ( Fig. 3 ). Total vertebrae 33–34 (exCluding the urostyle), with 16–17 abdominal and 17–18 Caudal vertebrae. Pterygiophore supporting the last dorsal-fin spine loCated between the neural spines of the thirteenth and fourteenth vertebrae or of the fourteenth and fifteenth vertebrae. Pterygiophore supporting the last anal-fin spine loCated between hemal spines of seventeenth and eighteenth vertebrae, rarely between rips of the fourteenth (abdominal) vertebrae and hemal spine of the fifteenth (Caudal) vertebrae. One single predorsal (supraneural) bone present. Hypurals 1 and 2 and hypurals 3 and 4 always fused into a single seamless unit. Coloration in life (based on field photographs of freshly Caught adult speCimens; Fig. 3 and 4 ). PronounCed sexual Color dimorphism present. Males with distinCt Color pattern of blaCk vertiCal bars (between two and three flank-sCales wide) and horizontal bands (between one and two, rarely three flank-sCales high). Body ground Coloration silvery-whitish. Iris of eyes whitish with dusky areas. Cheeks, operCulum and dorsum of freshly Caught speCimens sometimes in light purplish iridesCent. Dorsal head surfaCe, dorsal area of nape and area just below dorsal fin base dusky greyish-brown. Preorbital area whitish to dusky. Cheek with blaCkish horizontal stripe of variable shape and width below eye. SeCond horizontal stripe on ventral part of preoperCulum, intraoperCulum, and lower jaw, fused at the mental area with Corresponding stripe of other side. Area between those stripes whitish inCluding ventral area between lower jaws (area of the musculus genio-hyoideus ) and ventral parts of branChiostegal membrane. OperCulum and suboperCulum with two blaCkish vertiCal elements of variable shape and width. Four to seven blaCkish vertiCal bars at anterior part of flanks with first bar situated just behind gill Cover; if seven vertiCal bars shape of most posterior bar blotChlike. Five horizontal bands present. Dorsalmost one in most speCimens shortest starting on the level of first (anterior) dorsal-fin ray or behind it, extending to Caudal fin base and dorsal Caudal pedunCle area, there fusing with the horizontal band of the other side. SeCond and third horizontal band (Counted from dorsal to ventral) CommenCing anterior to dorsalmost horizontal band; third band sometimes CommenCing before seCond horizontal band and extending to Caudal fin base. Fourth horizontal band CommenCing below peCtoral-fin base, sometimes even before, and extending to Caudal fin base while fusing ventrally of Caudal pedunCle with the Corresponding band of other side. Most ventral horizontal band extending from gill Cover to the anal fin base. VertiCal bars in most Cases ventrally fused with horizontal bands. Dorsal-fin membrane with blaCk band along base, CommenCing at level of third or fourth dorsal-fin spine and extending to soft-rayed area of dorsal fin; above this blaCk band, a smaller whitish or turquoise iridesCent band; margin of dorsal-fin membrane blaCk. Anal-fin membrane with blaCkish or whitish areas of variable extent; soft rayed part with one to three ConspiCuous markings; eggspot-like markings of ovoid shape and of greyish to whitish Color and a blaCkish outline. Caudal-fin membrane white with blaCk horizontal elements, whiCh might be partially fused. PeCtoral fin whitish hyaline. PelviC fin whitish to greyish, posterior part of rays blaCk. Females almost entirely silvery white without any striking Color pattern. Cheeks, operCulum and dorsum of freshly Caught speCimens iridesCent light purple. Chest and belly whitish. Iris of eye whitish. Dorsal head surfaCe, dorsal area of nape and areas just below dorsal-fin base dusky (light greyish-brown to silvery) but less prominent than in males. Anal, pelviC and peCtoral whitish. Caudal fin whitish to dusky. Dorsal fin membrane whitish to dusky, margin of membrane between tips of spines and rays darkish. Coloration of juveniles unknown. Two subadult males (99.9 & 106.1 mm SL) with a less Contrasting flank Coloration than adults males, i.e. with vertiCal bars only slightly visible and horizontal bands almost invisible. Coloration in alCohol. ( Fig. 3 ) Overall Color pattern of vertiCal bars, horizontal bands, and head stripe elements Comparable to fresh male speCimens but dark brown. Body ground Coloration in male and female beige, only ventral region might appear whitish. Dorsal head surfaCe brownish. FIGURE 5. Pharyngeal jaws of the Hemibates stenosoma and H. koningsi sp. nov. A. Lateral view of pharyngeal jaws of H. stenosoma from the top to bottom: BMNH 1906–9–6–77–78 (syntype); ZSM 44567, DRC-2012/3124; ZSM 44564, DRC- 2012/3116 B. Lateral view of pharyngeal jaws of Hemibates koningsi sp. nov. paratypes from the top to bottom: MRAC 2017– 005–P-0001, DRC-2012/3103; ZSM 44570, DRC-2012/3186; ZSM 44565, DRC-2012/3243 C. Dorsal view of the lower pharyngeal jaw of Hemibates koningsi sp. nov. (MRAC 2017–005–P-0001, DRC-2012/3103, paratype), left: radiograph, suture of lower pharyngeal jaw straight, right: same lower pharyngeal jaw photographed through a stereomicroscope. Distribution and biology. Hemibates koningsi is known only from the southern part of Lake Tanganyika from around Mpulungu and the Chituta Bay and appears to be epibenthiC. It is Caught with gill nets at depths between 40 and 150 meters, mainly between 40–60 meters, along with Hemibates stenosoma (pers. Com J. Chanda, fisherman from a village near Kalambo river mouth). No stomaCh investigation was ConduCted, as they appeared to be empty in X-ray piCtures, Hemibates koningsi appears to be rare Compared to H. stenosoma : two and a half weeks of daily inspeCtion of daily artisanal CatChes in August/September ( 23.08.2015 – 09.09.2015 ) yielded only a maximum of zero to two individuals per day of H. koningsi but hundreds of H. stenosoma (identifiCation based on male Color pattern). This might explain why both Hemibates speCies are named Mpande (or “Mhpandi” in Konings (2015) in Bemba language and are not differentiated by the loCal fishermen although both speCies are highly appreCiated. Further, Hemibates CatChes appear to be sex-biased, with either males or females dominating single CatChes; this suggests sex speCifiC sChooling and homing (pers. obs., pers. Comm. Joseph Chanda). Etymology. The speCies is named for the first person who reCognized the new Hemibates speCies as a distinCt phenotype, Ad Konings, in appreCiation of the inspiration to many CiChlidophiles that arose from his Continued popular and sCientifiC Contributions.