Haplochromis argens

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de Zeeuw M, Westbroek I, van Oijen M, Witte F (2013) Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania. ZooKeys 256 : 1–34, doi. Versioned wiki page: 2013-01-02, version 29887, https://species-id.net/w/index.php?title=Haplochromis_argens&oldid=29887 , contributors (alphabetical order): Pensoft Publishers.

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BibTeX:

@article{de2013ZooKeys256,
author = {de Zeeuw, Marnix P. AND Westbroek, Irene AND van Oijen, Martien J.P. AND Witte, Frans},
journal = {ZooKeys},
publisher = {Pensoft Publishers},
title = {Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania},
year = {2013},
volume = {256},
issue = {},
pages = {1--34},
doi = {10.3897/zookeys.256.3871},
url = {http://www.pensoft.net/journals/zookeys/article/3871/abstract},
note = {Versioned wiki page: 2013-01-02, version 29887, https://species-id.net/w/index.php?title=Haplochromis_argens&oldid=29887 , contributors (alphabetical order): Pensoft Publishers.}

}

RIS/ Endnote:

TY - JOUR
T1 - Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania
A1 - de Zeeuw M
A1 - Westbroek I
A1 - van Oijen M
A1 - Witte F
Y1 - 2013
JF - ZooKeys
JA -
VL - 256
IS -
UR - http://dx.doi.org/10.3897/zookeys.256.3871
SP - 1
EP - 34
PB - Pensoft Publishers
M1 - Versioned wiki page: 2013-01-02, version 29887, https://species-id.net/w/index.php?title=Haplochromis_argens&oldid=29887 , contributors (alphabetical order): Pensoft Publishers.

M3 - doi:10.3897/zookeys.256.3871

Wikipedia/ Citizendium:

<ref name="de2013ZooKeys256">{{Citation
| author = de Zeeuw M, Westbroek I, van Oijen M, Witte F
| title = Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania
| journal = ZooKeys
| year = 2013
| volume = 256
| issue =
| pages = 1--34
| pmid =
| publisher = Pensoft Publishers
| doi = 10.3897/zookeys.256.3871
| url = http://www.pensoft.net/journals/zookeys/article/3871/abstract
| pmc =
| accessdate = 2025-04-01

}} Versioned wiki page: 2013-01-02, version 29887, https://species-id.net/w/index.php?title=Haplochromis_argens&oldid=29887 , contributors (alphabetical order): Pensoft Publishers.</ref>

See also the citation download page at the journal.


Taxonavigation

Ordo: Perciformes
Familia: Cichlidae
Genus: Haplochromis

Name

Haplochromis argens de Zeeuw, Westbroek & Witte sp. n. – Wikispecies link – ZooBank link – Pensoft Profile

Cheironyms used

Haplochromis argens: Goldschmidt 1994[1]: 179, 199–200; Haplochromis argens; Verheyen et al. 1989: 93, 94, 96; van der Meer 1995[2]: 26, 30–32; van der Meer and Bowmaker 1995[3]: 232–239; Goldschmidt 1996[4]: 164, 184–185; Hilder and Pankhurst 2003[5]: 194; Collin et al. 2004[6]: 769; Carleton et al. 2008[7].
Haplochromis “argens”; Zihler 1982[8]: 568; Barel 1983[9]: 384, 385, 407, 418; Witte 1984a[10]: 604, 611; Witte 1984b[11]: 159–161, 163; Barel 1985[12]: chapter 1, 384, 385, 407, 418; Gottfried 1986[13]: 1029; Witte 1987[14], chapter 1: 611, chapter 2: 67, 76, chapter 3: 8–13, 19; Goldschmidt 1989a[15]: 24, 27, 29–39, 41, 42, 44, 45, 53, 55, 58, 59, 61–63, 65–69, 71–75, 77, 82, 84, 86–89, 97–101, 103, 118, 119, 148, 158, 160, 162, 166; Goldschmidt 1989b[16]: 122–126, 129–131; Wanink et al. 1989[17]: 25; van der Meer 1989[18]: 52, 53; Goldschmidt and de Visser 1990[19]: 129, 130, 132; Goldschmidt et al. 1990[20]: 344, 346–351, 353; Goldschmidt and Witte 1990[21]: 356–367; Barel et al. 1991[22]: 262; Goldschmidt 1991[23]: 181, 182, 185, 187; van der Meer 1991a[24]: 91–94, 96; van der Meer 1991b[25]: 3, 5, 6, 9–12, 14–22, 24–26, 28, 68, 76, 77, 83, 85–87, 89–93, 99, 101, 104; Goldschmidt and Witte 1992[26]: 104; Kaufman 1992[27]: 848, 849; Witte et al. 1992b[28]: 11, 13, 17, 25, 27, 28; Barel 1993[29]: 366; Smith and Wootton 1994[30]: 99–103; Balshine-Earn 1995[31]: 5; Seehausen 1995[32]: 143, 146; Seehausen and Witte 1995[33]: 101; van der Meer et al. 1995[2]: 116–129; Smith and Wootton 1995[34]: 12; Anker and Dullemeijer 1996[35]: 4–11; de Visser and Barel 1996[36]: 4; Seehausen 1996[37]: 51, 55, 255, 256; Smit and Anker 1997[38]: 9–17; Seehausen et al. 1997b[39]: 899; Witte et al. 1997[40]: 591; Wanink 1998[41]: 159, 232; Rinkes 1999[42]: 46, 47, 54, 56–58, 60, 62, 76, 77, 79, 97; de Visser 2000[43]: 18; Tacon et al. 2000[44]: 63; Witte and Wanink 2000[45]: 78–81, 84; Seehausen et al. 2003[46]: 272; Witte et al. 2005[47]: 320; Kishe-Machumu 2012[48]: 35; van Rijssel and Witte 2012[49].
H. “Argens”; Kaufman and Seehausen 1995[50]: 148.
Haplochromis (?) “argens”; Wanink and Witte 2000[51]: 563; Witte et al. 2000[52]: 234, 235, 237; Witte et al. 2003[53]: 108; Niemantsverdriet 2005[54]: 151, 155; Witte et al. 2007a[55]: 1153.
Haplochromis sp. “argens”; Mizoiri et al. 2008[56]: 228, 241, 254.
Yssichromis argens; van Staaden et al. 1995[57]: 168; Chapman et al. 1995[58]: 1277, 1279–1282, 1285; Huber et al. 1997[59]: 170; Rosenberger and Chapman 2000[60]: 498; Melnychuk and Chapman 2002[61]: 107.
Yssichromis sp. “argens”; Mizoiri et al. 2008[56]: 228.
Type-locality. Tanzania, Lake Victoria, Mwanza Gulf (ca 2°29'–2°36'S; 32°48'–32°54'E) and Emin Pasha Gulf (ca 2°18'–2°41'S; 31°47'–31°59' E).
Holotype. RMNH.PISC.835884, ♂, 67.3 mm SL, Tanzania, Lake Victoria, Mwanza Gulf, 8.iii.1979, HEST.
Paratypes. All type specimens collected by Haplochromis Ecology Survey Team (HEST) in Mwanza Gulf, Tanzania, Lake Victoria, except where noted otherwise. Size of specimens given as standard length. RMNH.PISC.728315, ♀, 71.0 mm, 30.v.1980; RMNH.PISC.728845, 6, ♂, 74.4 mm, 31.v.1980; RMNH.PISC.73097, ♀, 71.3 mm, 27.ix.1977; RMNH.PISC.812025, ♀, 65.6 mm, 11.v.1978; RMNH.PISC.83587, ♂, 77.4 mm, 31.v.1975; RMNH.PISC.835894, ♀, 75.5 mm, 21.iv.1980; RMNH.PISC.835904, ♂, 68.2 mm, 22.iv.1980; RMNH.PISC.836064, ♂, 73.7 mm, 22.iv.1980; RMNH.PISC.836074, ♂, 65.7 mm, 15.iv.1980; RMNH.PISC.836085, ♂, 65.7 mm, 31.v.1975; RMNH.PISC.836094, ♂, 67.4 mm, 22.iv.1980; RMNH.PISC.836104, ♂, 66.2 mm, 30.ix.1977; RMNH.PISC.836114, ♂, 69.5 mm, 30.ix.1977; RMNH.PISC.836124, ♀, 71.1 mm, 30.ix.1977; RMNH.PISC.836134, ♀, 68.1 mm, 30.ix.1977; RMNH.PISC.836144, ♀, 71.9 mm, 8.ix.1977; RMNH.PISC.836154, ♂, 57.7 mm, 8.ix.1977; RMNH.PISC.836164, ♂, 70.2 mm, 19.viii.1977; RMNH.PISC.836174, ♀, 71.5 mm, 19.viii.1977; RMNH.PISC.836184, ♂, 66.1 mm, 7.ix.1977; RMNH.PISC.836194, ♂, 66.5 mm, 21.xii.1977; RMNH.PISC.836204, ♂, 65.8 mm, 10.x.1977; RMNH.PISC.836211, ♂, circa 70 mm, 30.ix.1977; RMNH.PISC.836225, ♂, 59.3 mm, 22.vi.1985, Emin Pasha Gulf; RMNH.PISC.836235, ♂, 57.2 mm, 22.vi.1985, Emin Pasha Gulf; RMNH.PISC.836245, ♂, 59.6 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836254, ♂, 58.3 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836264, ♂, 58.5 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836274, ♂, 54.3 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836284, ♂, 59.6 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836294, ♂, 55.4 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836304, ♂, 53.1 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836314, ♂, 66.8 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836324, ♂, 67.0 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836334, ♂, 60.4 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836344, ♂, 53.9 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836354, ♂, 62.6 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836364, ♂, 66.6 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836374, ♂, 69.2 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836384, ♂, 65.8 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836394, ♂, 61.3 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836404, ♂, 73.4 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836414, ♂, 64.1 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836424, ♂, 63.1 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836434, ♂, 63.7 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836444, ♂, 62.1 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836454, ♂, 64.1 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836464, ♂, 61.6 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836474, ♂, 59.8 mm, 25.viii.1981; RMNH.PISC.836484, ♂, 61.6 mm, 10.x.1977; RMNH.PISC.836494, ♂, 61.5 mm, 21.xii.1977; RMNH.PISC.836504, ♂, 57.7 mm, 6.iii.1979; RMNH.PISC.836514, ♂, 60.6 mm, 13.vii.1979; RMNH.PISC.836524, ♂, 58.0 mm, 1.ii.1979; RMNH.PISC.836534, ♂, 53.5 mm, 18.xi.1981; RMNH.PISC.836554, ♂, 61.8 mm, 14.v.1979; RMNH.PISC.836564, ♂, 59.0 mm, 27.iii.1979; RMNH.PISC.836574, ♂, 61.6 mm, 27.iii.1979; RMNH.PISC.836604, ♂, 56.8 mm, 12.iii.1979; RMNH.PISC.836614, ♂, 57.9 mm, 12.iii.1979; RMNH.PISC.836624, ♂, 59.7 mm, 14.v.1979; RMNH.PISC.836634, ♂, 58.6 mm, 14.v.1979; RMNH.PISC.836734, ♂, 55.5 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836881–3, ♂, 61.4 mm, 6.iii.1979; RMNH.PISC.836891–3, ♂, 62.2 mm, 14.v.1979; RMNH.PISC.836941, 5, ♂, 63.1 mm, 21.vi.1985, Emin Pasha Gulf; RMNH.PISC.836971, 5, ♂, 59.1 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.836981–3, 5, ♂, 61.0 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.837033, 6, ♂, 66.4 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.837041, 3, 6, ♂, 68.4 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.837051–3, 6, ♂, 67.5 mm, 23.vi.1985, Emin Pasha Gulf; RMNH.PISC.837061, 5, ♂, 68.8 mm, 27.vi.1985, Emin Pasha Gulf; RMNH.PISC.840675, 7, ♂, 72.1 mm, 15.viii.1986; AMNH 2550354, ♂, 62.2 mm, 14.v.1979; BMNH 2012.1.5.2 4, ♂, 62.8 mm, 3.v.1979; NSMT-P 1069594, ♂, 62.6 mm, 3.v.1979.
1 dissected to describe oral jaws; 2 dissected to describe pharyngeal jaws; 3 dissected to count gill filaments; 4 proportional measurements taken (Table 1); 5colour picture available; 6colour picture of anal fin available; 7 specimen of which Dr E. Verheijen, Royal Belgian Institute of Natural Sciences, has taken a tissue sample for DNA analysis.

Diagnosis

Small sized (< 8 cm SL), slender (BD < 31% SL), micrognathic (LJL < 45% HL) zooplanktivorous Haplochromis species with slightly curved to straight dorsal head profile. Relatively long and slender, mainly bicuspid, teeth in oral jaws. Premaxillary dentigerous area extending almost to caudal end of dentigerous arm. Both males and females silvery with conspicuously ivory-white lips. Three to five, generally faint vertical stripes on flank; faint traces of a dark mid-lateral band occasionally present. Males with yellow to greenish sheen on flank.

Description

Proportional measurements of type material given in Table 1.
Habitus. See Fig. 2. Body slender. Dorsal head profile straight to slightly curved, occasionally moderately curved. Premaxillary pedicel slightly prominent. Mouth oblique. Lips not thickened. Medial part of premaxilla slightly expanded. Caudal part of maxilla not bullate. Vertical through caudal tip of maxilla running through iris, just rostral to pupil. Lateral snout outline isognathous and obtuse, in larger specimens slightly prognathous. Jaws equal anteriorly or lower jaw slightly protruding. Mental prominence slightly pronounced. Retro-articular processes of right and left mandible touching each other, interrupting ventral body outline. Eye approximately circular and medium to large. Generally an aphakic aperture present in pupil. Cephalic lateral line pores not enlarged. Scales. Cheek, gill cover and rostral part of dorsal head surface covered with cycloid scales. Nape and rostral part of dorsum with mixture of cycloid and weakly ctenoid scales. Chest with ctenoid, weakly ctenoid and some cycloid scales. Scales on remaining part of body mainly ctenoid. Scales on chest smaller than those on ventral and ventro-lateral part of body; size transition gradual. Small elongated scales on basal quarter to half of caudal fin. Three to seven (mode 6) scales between upper lateral line and dorsal-fin origin, four to eight (mode 6) between pectoral- and pelvic-fin bases.
Fins. Pelvic fins just reaching or slightly surpassing rostral-most point of anal-fin origin. Pelvic fins with first soft rays slightly produced in both sexes, in males occasionally filamentous. Caudal tip of anal fin not reaching caudal-fin origin. Caudal-fin outline truncate to slightly emarginate.
Gill apparatus. Description based on lateral gill rakers and lateral hemibranch of first gill arch. Number of gill rakers on lower part of gill arch 11–12. Lower two to three rakers reduced (= very short), next one to two short, followed by two to six slender and longer ones. Remaining rakers hooked, bifid, trifid or quadrifid. Rakers generally closely set, viz. touching each other over major part of length. Number of gill filaments 94 to 106.
Viscera. Ratio between intestine length and SL: 1.0–1.4 (n = 25).
Oral jaws. (Fig. 3 A–C) Premaxillary ascending arm equal to or longer than dentigerous arm (asc./dent. arm ratio 1.0 to 1.1). Angle between the arms 77° to 81°. Symphyseal articulation facet not present. Lower jaw slightly more elongated than generalized type (length/height ratio 2.3 to 2.5). Upper half of dentary with distinct outwardly directed flare. Mental prominence slightly pronounced. Oral teeth shape. (Fig. 3 A–C) Generally teeth in outer row of both premaxilla and lower jaw bicuspid or weakly bicuspid, with some unicuspid or tricuspid teeth interspersed. In specimens over 65 mm SL, weakly bicuspids and unicuspids may dominate. Major cusp of bicuspids isoscelene to subequilateral, protracted and acutely pointed. Flange generally absent, when present very small. Minor cusp weakly developed to distinct, relatively short compared to major cusp. Cusp gap wide. In labial view, neck slender to moderately slender, crown not or slightly expanded. In lateral view, crown compressed. Outer-row teeth in both premaxilla and lower jaw recurved. Inner rows in both jaws with mainly tricuspid or weakly tricuspid teeth.
Oral teeth size. Outer-row teeth relatively long and slender, gradually decreasing in size from rostral to caudal.
Dental arcade and tooth band. (Fig. 3B) Rostrally dental arcade rounded. Outer row generally occupying almost total length of dentigerous arm of premaxilla, in two specimens (RMNH.PISC.83697 and RMNH.PISC.83621; Fig. 3B) edentulous part about 25% of arm. Outer row in lower jaw not, or just, reaching coronoid wing in most dissected specimens. In one case caudal-most tooth relatively high on coronoid wing. One or two inner rows in rostral part of both jaws, decreasing to zero in caudal part.
Teeth counts and setting. Outer row of upper jaw (l+r premaxilla) with 30–52 teeth. In both jaws outer-row teeth regularly set, their placement wider rostrally than laterally.
Tooth implantation. Outer-row teeth of premaxilla rostrally erect. Inner-row teeth recumbent. Outer-row teeth of lower jaw slightly procumbent, inner-row teeth erect.
Lower pharyngeal element. (Fig. 3 D, E) Lower pharyngeal element relatively small and slender (length/width ratio 1.2–1.3). Dentigerous area slightly broader than long (length/width ratio = 0.7–0.9). Suture straight.
Pharyngeal teeth counts. Caudal-most transverse row with about 30–38 teeth, medial longitudinal rows with eight to 11 teeth.
Pharyngeal teeth shape. Teeth in caudal-most transverse row hooked, major cusp only slightly incurved, blunt to slightly acute. Other teeth bevelled or pronounced. All teeth relatively fine and slender, medial teeth not coarser than other teeth.
Vertebrae. Total number of vertebrae in 57 specimens: 30 (12), 31 (39) or 32 (6), comprising 13–14 abdominal and 16–19 caudal vertebrae.
Live colouration males. (Fig. 4 A, B). Sexually active males with ivory to grey snout and cheek. Lips remarkably ivory-white with no or few pigment spots. Eye with grey outer ring and silver to golden inner ring. Lower jaw and interoperculum whitish. Gill cover silver, sometimes with grey to dusky flush. Dorsal head surface, dorsum and flank silvery-grey, dorsum with bluish to purplish sheen, flank with yellow to greenish sheen. Chest, belly and ventral side silvery-white. Pelvic fins black; in specimens of Emin Pasha Gulf, medial side sometimes red. Anal fin rostrally faintly to distinctly red, rest of fin hyaline. One to two dark yellow to orange egg dummies with hyaline ring present on caudal part of anal fin. Caudal fin orange-red to wine-red. Dorsal fin hyaline with red streaks and spots. Lappets hyaline or reddish, rostral lappets sometimes dusky.
Dark grey to blackish markings: Nostril-, interorbital-, and supraorbital stripes, sometimes rather distinct. Lachrymal stripe distinct, but relatively short (i.e. small blotch at caudal end of lachrymal generally not reaching caudal tip of maxilla), sometimes extending over iris. Irregular preopercular vertical bar generally present. Opercular blotch distinct. Three to five, generally faint vertical stripes on flank. Traces of dark-grey mid-lateral band occasionally present.
Live colouration females. Live females basically with same colours as males, lacking bluish-purplish and yellow-greenish sheens and distinct red colouration in fins, but sometimes with faint red flush in caudal fin. In females upper lip usually with more pigment than in males. Of markings on head, only lachrymal stripe and opercular blotch distinct. Mid-lateral band sometimes more distinct than in males, vertical stripes faint.
Preserved colouration of males and females. (Fig. 5) Body light brown, dorsally darker than ventrally. Snout, lips and lower jaw coloured as old ivory. Fins hyaline and light grey-brown in both sexes, except for black pelvic fins in adult males. Same markings, but slightly more distinct, as in live specimens.

Distribution

Haplochromis argens is only known from the Tanzanian part of Lake Victoria. Specimens were caught in the Mwanza Gulf (from entrance of Stuhlmann Sound to entrance of gulf in north), in the south-western part of the Speke Gulf (near its entrance), in the area around Kome Island, and in the Emin Pasha Gulf (Fig. 1).

Habitat

Haplochromis argens is a pelagic species from the littoral and sub-littoral zone. At night the species is virtually restricted to the two upper metres of the water column (Witte 1984b[11], Goldschmidt et al. 1990[20]). By day, the highest densities were found at two to three metres from the surface, but individuals of this species were also caught with bottom trawls over sand and mud bottoms, and in gill nets and traps near rocks (Goldschmidt et al. 1990[20], Witte et al. 1992b[28]).

Abundance

During 1979–1982, Haplochromis argens was present in 70% of the bottom-trawl tows by day and in 100% of the surface trawl tows at night; the mean numbers of individuals per tow ranged from 6.2 to 18.3, respectively (Table 2). In 1987–1988, it occurred in 3% of the bottom trawl tows and the mean number of individuals per tow was 0.03; the species was absent in surface trawls. Thirty-four bottom-trawl tows in 1990–1999 captured no Haplochromis argens. From 2001 to 2011 more than 150 bottom-trawl tows contained about 15 individuals of Haplochromis argens, corresponding to a decline in catch per unit effort of more than 50 times compared to the 1979–1982 captures; no individuals were caught with surface trawls.

Table 2. Frequencies of occurrence (Foo) in trawl tows and mean numbers (± SD) per tow of Haplochromis argens. Tows of 10 minutes duration were made with bottom trawls (day) at stations E to J (depth range 7–14 m) and with surface trawls (night) at station G (14 m deep); in the period 1990-1999 no surface trawls were made; n indicates number of trawl tows.
Station Bottom trawl day 1979–1982† n = 104 1987–1988† n = 29 1990–1999‡ n = 34 2001–2011§ n > 150
E–J Foo 70 % 3 % 0 % < 10 %
Mean nr 6.2 ± 12.2 0.03 ± 0.19 0 < 0.1
Surface trawl night 1981/1982 n = 8 1987-19888| n = 26 - 2001-20111§ n = 15
G Foo 100 % 0 % - 27 %
Mean nr 18.3 ± 12.4 0 - 0.5 ± 0.9
Food. Before the ecological changes in Lake Victoria,the diet of Haplochromis argens comprised mainly zooplankton during the day; Chaoborus larvae were important at night (Goldschmidt et al. 1990[20]). The current diet is unknown, but all the studied resurgent species in the Mwanza Gulf changed their diet (e.g. van Oijen and Witte 1996[62], Katunzi et al. 2003[63], Kishe-Machumu et al. 2008[64]).

Breeding. Haplochromis argens is a female mouth brooder. Spawning sites are located at depths < 9 m (Goldschmidt and Witte 1990[21]).

Etymology

In reference to the silver male colouration, Haplochromis argens was given the nickname “argens” under the false assumption it was Latinized Greek for silver. Since this species is well known under its cheironym, we think it is best to upgrade the nickname to the species’ epithethon.

Comparisons

The zooplanktivorous species Haplochromis (Yssichromis) laparogramma Greenwood & Gee, 1969, Haplochromis (Yssichromis) pyrrhocephalus Witte & Witte-Maas, 1987 and Haplochromis (Yssichromis) heusinkveldi, Witte & Witte-Maas, 1987, have shorter bicuspid teeth in the oral jaws than Haplochromis argens, and generally the premaxillary dentigerous arm is edentulous over the caudal 1/4 -1/3 versus the dentigerous portion extending almost to the caudal end of the dentigerous arm. The dental features of the zooplanktivorous/insectivorous Haplochromis tanaos van Oijen & Witte, 1996 and Haplochromis thereuterion van Oijen & Witte, 1996, are more or less similar to those of Haplochromis argens, but the former two species have more unicuspids. Haplochromis argens is further distinguished from these and other species by its colouration. Sexually active males of Haplochromis tanaos are dark blue, the females silvery with a distinct mid-lateral band and slightly less distinct dorso-lateral band. Sexually active males of Haplochromis thereuterion are black, the females coloured like females of Haplochromis tanaos (van Oijen and Witte 1996[62]). The body of Haplochromis argens is less slender (BD 26.0–30.7% of SL, mean 28.2%, Table 1) than that of Haplochromis tanaos and Haplochromis thereuterion (22.1–27.1% and 24.4–27.6% of SL, respectively; Tables 3 and 7 in van Oijen and Witte 1996[62]). Live Haplochromis argens is similar to (juvenile) Haplochromis cassius in colouration and general habitus. However, Haplochromis cassius has a broad, well defined mid-lateral band (Greenwood and Barel 1978[65]), more and distinctly longer unicuspid teeth, more curved and more widely set teeth, and thicker lips than Haplochromis argens. The maximum size of Haplochromis cassius (99.0 mm SL) is larger than that of Haplochromis argens, but when comparing similar size ranges (see material and methods) Haplochromis argens has: a smaller head (ratio HL/SL: 31.0 – 35.6%, mean 32.9% versus 35.0 – 36.0%, mean 35.5% in Haplochromis cassius); a shorter snout (ratio SnL/HL: 23.6 – 29.7%, mean 26.6% versus 29.8 – 32.6%, mean 31.2% in Haplochromis cassius); larger eyes (ratio EyL/HL: 30.9 – 39.0%, mean 35.7% versus 26.8 – 28.8%, mean 28.3% in Haplochromis cassius). For comparison with Haplochromis goldschmidti sp. n., see below.

Original Description

  • de Zeeuw, M; Westbroek, I; van Oijen, M; Witte, F; 2013: Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania ZooKeys, 256: 1-34. doi

Other References

  1. ↑ Goldschmidt T (1994) Darwins Hofvijver. Uitgeverij Prometheus, Amsterdam, 286 pp.
  2. ↑ 2.0 2.1 van der Meer H (1995) Visual resolution during growth in cichlid fish: A morphological and behavioural case study. Brain, Behaviour and Evolution 45: 25-33. doi: 10.1159/000113383
  3. ↑ van der Meer H, Bowmaker J (1995) Interspecific variation of photoreceptors in four co-existing haplochromine cichlid fishes. Brain, Behaviour and Evolution 45: 232-240. doi: 10.1159/000113552
  4. ↑ Goldschmidt T (1996) Darwin’s Dreampond. MIT Press, Cambridge, Massachusetts, 277 pp.
  5. ↑ Hilder M, Pankhurst N (2003) Evidence that temperature change cues reproductive development in the spiny damselfish, Acanthochromis polyacanthus. Environmental Biology of Fishes 66: 187-196. doi: 10.1023/A:1023601729203
  6. ↑ Collin S, Hart N, Wallace K, Shand J, Potter I (2004) Vision in the southern hemisphere lamprey Mordacia mordax: Spatial distribution, spectral absorption characteristics, and optical sensitivity of a single class of retinal photoreceptor. Visual Neuroscience 21: 765–773. doi: 10.1017/S0952523804215103
  7. ↑ Carleton K, Spady T, Streelman J, Kidd M, McFarland W, Loew E (2008) Visual sensitivities tuned by heterochronic shifts in opsin gene expression. BioMed Central Biology 6: 22. doi: 10.1186/1741-7007-6-22
  8. ↑ Zihler F (1982) Gross morphology and configuration of the digestive tract of Cichlidae (Teleostei, Perciformes): phylogenetic and functional significance. Netherlands Journal of Zoology 32: 544-571. doi: 10.1163/002829682X00210
  9. ↑ Barel C (1983) Towards a constructional morphology of cichlid fishes (Teleostei, Perciformes). Netherlands Journal of Zoology 33: 357-424. doi: 10.1163/002829683X00183
  10. ↑ Witte F (1984a) Consistency and functional significance of morphological differences between wild-caught and domestic Haplochromis squamipinnis (Pisces, Ciclidae). Netherlands Journal of Zoology 34: 596-612. doi: 10.1163/002829684X00308
  11. ↑ 11.0 11.1 Witte F (1984b) Ecological differentiation in Lake Victoria haplochromines: comparison of cichlid species flocks in African Lakes. In: Echelle A Kornfield I (Eds). , Evolution of Fish Species Flocks. Orono Press, University of Maine, Maine: 155-167.
  12. ↑ Barel C (1985) A matter of space, constructional morphology of cichlid fishes. PhD Thesis, Leiden, the Netherlands: Leiden University.
  13. ↑ Gottfried M (1986) Developmental transition in feeding morphology of the Midas cichlid. Copeia 1986: 1028-1030. doi: 10.2307/1445308
  14. ↑ Witte F (1987) From form to fishery. An ecological and taxonomical contribution to morphology and fishery of Lake Victoria cichlids. PhD Thesis, Leiden, the Netherlands: Leiden University.
  15. ↑ Goldschmidt T (1989a) An ecological and morphological field study on the haplochromine cichlid fishes (Pisces, Cichlidae) of Lake Victoria. PhD Thesis, Leiden, the Netherlands: Leiden University.
  16. ↑ Goldschmidt T (1989b) Reproductive strategies, subtrophic niche differentiation and the role of competition for food in haplochromine cichlids (Pisces) from Lake Victoria, Tanzania. Annalenvan het Koninklijk Museum voor Midden-Afrika, Zoologische Wetenschappen 257: 119-132.
  17. ↑ Wanink J, Goldschmidt T, Witte F (1989) Recent changes in the zooplanktivorous-/insectivorous fish community from the Mwanza Gulf of Lake Victoria. In: Anonymous (Eds). WOTRO Report for the year 1988. Netherlands Foundation for the Advancement of Tropical Research, The Hague, the Netherlands: 22-28.
  18. ↑ van der Meer H (1989) Ecological significance of retinal receptor patterns in 4 sympatric haplochromine cichlids. Annalenvan het Koninklijk Museum voor Midden-Afrika, Zoologische Wetenschappen 257: 51-56.
  19. ↑ Goldschmidt T, de Visser J (1990) On the possible role of egg mimics in speciation. Acta Biotheoretica 38: 125-134. doi: 10.1007/BF00047549
  20. ↑ 20.0 20.1 20.2 20.3 Goldschmidt T, Witte F, de Visser J (1990) Ecological segregation of zooplanktivorous haplochromine species (Pisces, Cichlidae) from Lake Victoria. Oikos 58: 343-355. doi: 10.2307/3545226
  21. ↑ 21.0 21.1 Goldschmidt T, Witte F (1990) Reproductive strategies of zooplanktivorous haplochromine species (Pisces, Cichlidae) from Lake Victoria before the Nile perch boom. Oikos 58: 356-368. doi: 10.2307/3545227
  22. ↑ Barel C, Ligtvoet W, Goldschmidt T, Witte F, Goudswaard P (1991) The haplochromine cichlids in Lake Victoria: an assessment of biological and fisheries interests. In: Keenleyside M (Ed). Cichlid Fishes, Behaviour, Ecology and Evolution. Chapman & Hall, London: 258-279.
  23. ↑ Goldschmidt T (1991) Egg mimics in haplochromine cichlids (Pisces, Perciformes) from Lake Victoria. Ethology 88: 177-190. doi: 10.1111/j.1439-0310.1991.tb00273.x
  24. ↑ van der Meer H (1991a) Determination of photopic thresholds in two sympatric cichlids using optomotor response. Annalenvan het Koninklijk Museum voor Midden-Afrika, Zoologische Wetenschappen 263: 91-96.
  25. ↑ van der Meer H (1991b) Ecomorphology of photoreception in haplochromine cichlid fishes. PhD Thesis, Leiden, the Netherlands: Leiden University.
  26. ↑ Goldschmidt T, Witte F (1992) Explosive speciation and adaptive radiation of haplochromine cichlids from Lake Victoria: an illustration of the scientific value of a lost species flock. Mitteilungen Internationale Vereinigung für theoretische und angewandte Limnologie 23: 101–107.
  27. ↑ Kaufman L (1992) Catastrophic change in species-rich freshwater ecosystems. BioScience 42: 846–858. doi: 10.2307/1312084
  28. ↑ 28.0 28.1 Witte F, Goldschmidt T, Wanink J, van Oijen M, Goudswaard K, Witte-Maas E, Bouton N (1992b) The destruction of an endemic species flock: quantitative data on the decline of the haplochromine cichlids of Lake Victoria. Environmental Biology of Fishes 34: 1-28. doi: 10.1007/BF00004782
  29. ↑ Barel C (1993) Concepts of an architectonic approach to transformation morphology. Acta Biotheoretica 41: 345-381. doi: 10.1007/BF00709371
  30. ↑ Smith C, Wootton R (1994) The cost of parental care in Haplochromis ‘argens’ (Cichlidae). Environmental Biology of Fishes 40: 99-104. doi: 10.1007/BF00002184
  31. ↑ Balshine-Earn S (1995) The costs of parental care in Galilee St Peter’s fish, Sarotherodon galilaeus. Animal Behaviour 50: 1-7. doi: 10.1006/anbe.1995.0214
  32. ↑ Seehausen O (1995) Cichliden von der tansanischen Victoriasee-Küste. In: Stawikowski, R (Ed). Cichliden, Festschrift zum 25jährigen Jubiläum der DCG. Deutsche Cichliden-Gesellschaft, Frankfurt am Main: 143-146.
  33. ↑ Seehausen O, Witte F (1995) Extinction of many and survival of some: the current situation of the endemic cichlids in southern Lake Victoria. Tropical Fish Hobbyist 43 (7): 96-105.
  34. ↑ Smith C, Wootton R (1995) The costs of parental care in teleost fishes. Reviews in Fish Biology and Fisheries 5: 7-22. doi: 10.1007/BF01103363
  35. ↑ Anker G, Dullemeijer P (1996) Transformation morphology on structures in the head of cichlid fishes. In: Datta M Dutta H (Eds). Fish Morphology – Horizon of New Research. Oxford & IBH Publishing, New Delhi: 1-20.
  36. ↑ de Visser J, Barel C (1996) Architectonic constraints on the hyoid’s optimal starting position for suction feeding of fish. Journal of Morphology 228: 1-18. doi: <1::AID-JMOR1>3.0.CO;2-B 10.1002/(SICI)1097-4687(199604)228:1<1::AID-JMOR1>3.0.CO;2-B
  37. ↑ Seehausen O (1996) Lake Victoria Rock Cichlids: Taxonomy, Ecology and Distribution. Verduijn Cichlids, Zevenhuizen, 304 pp.
  38. ↑ Smit S, Anker G (1997) Photopic sensitivity to red and blue light related to retinal differences in two zooplanktivorous haplochromine species (Teleostei, Cichlidae). Netherlands Journal of Zoology 47: 9-20. doi: 10.1163/156854297X00201
  39. ↑ Seehausen O, Witte F, Katunzi E, Smits J, Bouton N (1997b)Patterns of the remnant cichlid fauna in southern Lake Victoria. Conservation Biology 11: 890-904. doi: 10.1046/j.1523-1739.1997.95346.x
  40. ↑ Witte F, Barel C, van Oijen M (1997) Intraspecific variation of haplochromine cichlids from Lake Victoria and its taxonomic implications. South African Journal of Science 93: 585–594.
  41. ↑ Wanink J (1998) The pelagic cyprinid Rastrineobola argentea as a crucial link in the disrupted ecosystem of Lake Victoria. PhD Thesis, Leiden, the Netherlands: Leiden University.
  42. ↑ Rinkes M (1999) Conflicts in cichlid head morphology due to optimization of functional demands on suction feeding. PhD Thesis, Leiden, the Netherlands: Leiden University.
  43. ↑ de Visser J (2000) Tongue in cheek. A study in biological engineering of fish. PhD Thesis, Leiden, the Netherlands: Leiden University.
  44. ↑ Tacon P, Baroiller J, Le Bail P, Prunet P, Jalabert B (2000) Effect of egg deprivation on sex steroids, gonadotropin, prolactin, and growth hormone profiles during the reproductive cycle of the mouthbrooding cichlid fish Oreochromis niloticus. General and Comparative Endocrinology 117: 54-65. doi: 10.1006/gcen.1999.7388
  45. ↑ Witte F, Wanink J (2000) De zoöplanktivore cichliden van het Victoriameer: ontdekking, verdwijning en terugkeer. Cichlidae 26: 77-84.
  46. ↑ Seehausen O, van Alphen J, Witte F (2003) Implications of eutrophication for fish vision, behavioral ecology and species coexistence: A theoretical framework. In: Crisman T Chapman L Chapman C Kaufman L (Eds). Conservation, Ecology, and Management of African Fresh Waters. University Press of Florida, Florida: 268-287.
  47. ↑ Witte F, Wanink J, Rutjes H, van der Meer H, van den Thillart G (2005) Eutrophication and its influences on the fish fauna of Lake Victoria. In: Reddy V (Ed). Restoration and Management of Tropical Eutrophic Lakes. Science publishers, Inc., Enfield (NH): 301-338.
  48. ↑ Kishe-Machumu M (2012) Inter-guild differences and possible causes of the recovery of cichlid species in Lake Victoria, Tanzania. PhD Thesis, Leiden, the Netherlands: Leiden University.
  49. ↑ van Rijssel J, Witte F (2012) Adaptive responses in resurgent Lake Victoria cichlids over the past 30 years. Evolutionary Ecology. doi: 10.1007/s10682-012-9596-9
  50. ↑ Kaufman L, Seehausen O (1995) Cichliden von der ugandischen und der kenianischen Victoriaseeküste. In: Stawikowski R (Ed). Cichliden, Festschrift zum 25jährigen Jubiläum der DCG. Deutsche Cichliden-Gesellschaft, Frankfurt am Main: 147-151.
  51. ↑ Wanink J, Witte F (2000) The use of perturbation as a natural experiment: effects of predator introduction on the community structure of zooplanktivorous fish in Lake Victoria. Advances in Ecological Research 31: 553-570. doi: 10.1016/S0065-2504(00)31030-3
  52. ↑ Witte F, Msuku B, Wanink J, Seehausen O, Katunzi E, Goudswaard P, Goldschmidt T (2000) Recovery of cichlid species in Lake Victoria: an examination of factors leading to differential extinction. Reviews in Fish Biology and Fisheries 10: 233-241. doi: 10.1023/A:1016677515930
  53. ↑ Witte F, van der M, Barel K (2003) Door cichlidenogen bezien: het belang van het gezichtsvermogen bij haplochromine cichliden voor het ontstaan en in stand houden van diversiteit. Cichlidae 29: 104-114.
  54. ↑ Niemantsverdriet P (2005) Victoriacichliden houden en kweken in het laboratorium. Cichlidae 31: 150-156.
  55. ↑ Witte F, Wanink J, Kishe-Machumu M (2007a) Species distinction and the biodiversity crisis in Lake Victoria. Transactions of the American Fisheries Society 136: 1146-1159. doi: 10.1577/T05-179.1
  56. ↑ 56.0 56.1 Mizoiri S, Aibara M, Okada N (2008) Live Cichlids in the Southern Lake Victoria - Ongoing Speciation. (in Japanese) Report of Grant-in-Aid for Scientific Research on Priority Areas 2002–2007 from The Ministry of Education, Culture, Sports, Science and Technology (Japan). Kougakutosho Ltd., Tokyo, 278 pp.
  57. ↑ van Staaden M, Huber R, Kaufman L, Liem K (1995) Brain evolution in cichlids of the African Great Lakes: brain and body size, general patterns, and evolutionary trends. Zoology 98: 165-178.
  58. ↑ Chapman L, Kaufman L, Chapman C, McKenzie F (1995) Hypoxia tolerance in twelve species of East African cichlids: potential for low oxygen refugia in Lake Victoria. Conservation Biology 9: 1274-1287. doi: 10.1046/j.1523-1739.1995.9051262.x-i1
  59. ↑ Huber R, van Staaden M, Kaufman L, Liem K (1997) Microhabitat use, trophic patterns, and the evolution of brain structure in African cichlids. Brain, Behaviour and Evolution 50: 167-182. doi: 10.1159/000113330
  60. ↑ Rosenberger A, Chapman L (2000) Respiratory characters of three species of haplochromine cichlids: Implications for use of wetland refugia. Journal of Fish Biology 57: 483-501. doi: 10.1111/j.1095-8649.2000.tb02187.x
  61. ↑ Melnychuk M, Chapman L (2002) Hypoxia tolerance of two haplochromine cichlids: swamp leakage and potential for interlacustrine dispersal. Environmental Biology of Fishes 65: 99-110. doi: 10.1023/A:1019602403098
  62. ↑ 62.0 62.1 62.2 van Oijen M, Witte F (1996) Taxonomical and ecological description of a species complex of zooplanktivorous and insectivorous cichlids from Lake Victoria. Zoologische Verhandelingen Leiden 302: 1-56.
  63. ↑ Katunzi E, Zoutendijk J, Goldtschmidt T, Wanink J, Witte F (2003) Lost zooplanktivorous cichlid from Lake Victoria reappears with a new trade. Ecology of Freshwater Fish 12: 237-240. doi: 10.1046/j.1600-0633.2003.00023.x
  64. ↑ Kishe-Machumu M, Witte F, Wanink J (2008) Dietary shift in benthivorous cichlids after the ecological changes in Lake Victoria. Animal Biology 58: 401-417. doi: 10.1163/157075608X383700
  65. ↑ Greenwood P, Barel C (1978) A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae), Part VIII. Bulletin British Museum of natural History (Zoology), 33: 141–192.

Images

Figure 1. Tanzanian part of Lake Victoria. A, distribution of Haplochromis argens sp. n.(hatched area) B Mwanza Gulf with catch localities of type specimens of Haplochromis argens sp. n.(hatched area); E to J represent stations along the research transect where data on abundance of Haplochromis argens sp. n. were collected C Emin Pasha Gulf with catch localities of type specimens of Haplochromis argens sp. n. (filled circles) and type specimens of Haplochromis goldschmidti sp. n. (filledtriangles); open triangles indicate other catch localities of Haplochromis goldschmidti sp. n.
Figure 1. Tanzanian part of Lake Victoria. A, distribution of Haplochromis argens sp. n.(hatched area) B Mwanza Gulf with catch localities of type specimens of Haplochromis argens sp. n.(hatched area); E to J represent stations along the research transect where data on abundance of Haplochromis argens sp. n. were collected C Emin Pasha Gulf with catch localities of type specimens of Haplochromis argens sp. n. (filled circles) and type specimens of Haplochromis goldschmidti sp. n. (filledtriangles); open triangles indicate other catch localities of Haplochromis goldschmidti sp. n. 
Figure 2. Habitus of Haplochromis argens sp. n. ♂ (holotype, RMNH.PISC.83588). Scale bar equals 10 mm. Drawing by I. Westbroek, missing (= dotted) scales added by M. van Oijen.
Figure 2. Habitus of Haplochromis argens sp. n. ♂ (holotype, RMNH.PISC.83588). Scale bar equals 10 mm. Drawing by I. Westbroek, missing (= dotted) scales added by M. van Oijen. 
Figure 3. Skeletal elements of Haplochromis argens sp. n. A Right premaxilla, lateral view (RMNH.PISC.83705) B Right premaxilla, lateral (top) and occlusal (bottom) views (RMNH.PISC.83621), illustrating, for this species, a rare case of a posteriorly edentulous premaxilla C Right lower jaw, lateral view (RMNH.PISC.83697) D Lower pharyngeal element, dorsal view (RMNH.PISC. 83706) E Lower pharyngeal element, lateral view (RMNH.PISC.83706). Scale bars equal 1 mm. A, C, D drawn by I. Westbroek, B by M. van Oijen.
Figure 3. Skeletal elements of Haplochromis argens sp. n. A Right premaxilla, lateral view (RMNH.PISC.83705) B Right premaxilla, lateral (top) and occlusal (bottom) views (RMNH.PISC.83621), illustrating, for this species, a rare case of a posteriorly edentulous premaxilla C Right lower jaw, lateral view (RMNH.PISC.83697) D Lower pharyngeal element, dorsal view (RMNH.PISC. 83706) E Lower pharyngeal element, lateral view (RMNH.PISC.83706). Scale bars equal 1 mm. A, C, D drawn by I. Westbroek, B by M. van Oijen. 
Figure 4. Live colours of Haplochromis argens sp. n. A sexually active ♂, 59.3 mm SL (paratype, RMNH.PISC.83622), Emin Pasha Gulf B sexually active ♂, 72.1 mm SL (paratype, RMNH.PISC.84067), Mwanza Gulf.
Figure 4. Live colours of Haplochromis argens sp. n. A sexually active ♂, 59.3 mm SL (paratype, RMNH.PISC.83622), Emin Pasha Gulf B sexually active ♂, 72.1 mm SL (paratype, RMNH.PISC.84067), Mwanza Gulf. 
Figure 5. Preserved colours of Haplochromis argens sp. n. ♂ 67.3 mm SL (holotype, RMNH.PISC.83588). Scale bar equals 10 mm.
Figure 5. Preserved colours of Haplochromis argens sp. n. ♂ 67.3 mm SL (holotype, RMNH.PISC.83588). Scale bar equals 10 mm. 
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