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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.256.3871" ID-GBIF-Dataset="f9361112-c4fb-4c2f-b7f2-90cc1f9fc2f3" ID-PMC="PMC3650828" ID-Pensoft-Pub="1313-2970-256-1" ID-PubMed="23717179" ModsDocAuthor="" ModsDocDate="2013" ModsDocID="1313-2970-256-1" ModsDocOrigin="ZooKeys 256" ModsDocTitle="Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania" checkinTime="1451247797369" checkinUser="pensoft" docAuthor="de Zeeuw, Marnix P., Westbroek, Irene, van Oijen, Martien J. P. &amp; Witte, Frans" docDate="2013" docId="07943F3B81006D47F772A030A0F7AAAB" docLanguage="en" docName="ZooKeys 256: 1-34" docOrigin="ZooKeys 256" docSource="http://dx.doi.org/10.3897/zookeys.256.3871" docTitle="Haplochromis argens de Zeeuw, Westbroek &amp; Witte, sp. n." docType="treatment" docVersion="4" lastPageNumber="12" masterDocId="FFF0FFC2FFE8DF17FFB26F17FFF05401" masterDocTitle="Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania" masterLastPageNumber="34" masterPageNumber="1" pageNumber="5" updateTime="1668155110665" updateUser="ExternalLinkService">
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<mods:title>Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>de Zeeuw, Marnix P.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Westbroek, Irene</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>van Oijen, Martien J. P.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Witte, Frans</mods:namePart>
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<mods:date>2013</mods:date>
<mods:detail type="volume">
<mods:number>256</mods:number>
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<treatment ID-GBIF-Taxon="152039577" LSID="urn:lsid:zoobank.org:act:B0288AFB-9C11-4D81-9B7B-E551951A7B2D" httpUri="http://treatment.plazi.org/id/07943F3B81006D47F772A030A0F7AAAB" lastPageId="12" lastPageNumber="12" pageId="4" pageNumber="5">
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<taxonomicName LSID="urn:lsid:zoobank.org:act:B0288AFB-9C11-4D81-9B7B-E551951A7B2D" authority="de Zeeuw, Westbroek &amp; Witte" class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="argens">Haplochromis argens de Zeeuw, Westbroek &amp; Witte</taxonomicName>
<taxonomicNameLabel pageId="4" pageNumber="5">sp. n.</taxonomicNameLabel>
Figs 2-5, Tables 1, 2
</paragraph>
</subSubSection>
<subSubSection lastPageId="7" lastPageNumber="8" pageId="4" pageNumber="5" type="cheironyms used">
<paragraph pageId="4" pageNumber="5">Cheironyms used.</paragraph>
<paragraph pageId="4" pageNumber="5">
Haplochromis argens:
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Uitgeverij Prometheus, Amsterdam" pageId="23" pageNumber="24" title="Darwins Hofvijver." year="1994">Goldschmidt 1994</bibRefCitation>
: 179, 199-200;
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
; Verheyen et al. 1989: 93, 94, 96;
<bibRefCitation author="van der Meer, HJ" journalOrPublisher="Brain, Behaviour and Evolution" pageId="25" pageNumber="26" pagination="25 - 33" title="Visual resolution during growth in cichlid fish: A morphological and behavioural case study." url="10.1159/000113383" volume="45" year="1995">van der Meer 1995</bibRefCitation>
: 26, 30-32;
<bibRefCitation author="van der Meer, HJ" journalOrPublisher="Brain, Behaviour and Evolution" pageId="25" pageNumber="26" pagination="232 - 240" title="Interspecific variation of photoreceptors in four co-existing haplochromine cichlid fishes." url="10.1159/000113552" volume="45" year="1995">van der Meer and Bowmaker 1995</bibRefCitation>
: 232-239;
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="MIT Press, Cambridge, Massachusetts" pageId="23" pageNumber="24" title="Darwin's Dreampond." year="1996">Goldschmidt 1996</bibRefCitation>
: 164, 184-185;
<bibRefCitation author="Hilder, ML" journalOrPublisher="Environmental Biology of Fishes" pageId="24" pageNumber="25" pagination="187 - 196" title="Evidence that temperature change cues reproductive development in the spiny damselfish, Acanthochromis polyacanthus." url="10.1023/A:1023601729203" volume="66" year="2003">Hilder and Pankhurst 2003</bibRefCitation>
: 194;
<bibRefCitation author="Collin, SP" journalOrPublisher="Conservation Biology" pageId="23" pageNumber="24" title="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." url="10.1017/S0952523804215103" year="2004">Collin et al. 2004</bibRefCitation>
: 769;
<bibRefCitation author="Carleton, KL" journalOrPublisher="Proceedings of the Royal Society of London B" pageId="23" pageNumber="24" title="Visual sensitivities tuned by heterochronic shifts in opsin gene expression. BioMed Central Biology 6: 22." url="10.1186/1741-7007-6-22" year="2008">Carleton et al. 2008</bibRefCitation>
.
</paragraph>
<paragraph lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis" order="Perciformes" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Haplochromis</taxonomicName>
<normalizedToken originalValue="“argens”">&quot;argens&quot;</normalizedToken>
;
<bibRefCitation author="Zihler, F" journalOrPublisher="Netherlands Journal of Zoology" pageId="26" pageNumber="27" pagination="544 - 571" title="Gross morphology and configuration of the digestive tract of Cichlidae (Teleostei, Perciformes): phylogenetic and functional significance." url="10.1163/002829682X00210" volume="32" year="1982">Zihler 1982</bibRefCitation>
: 568;
<bibRefCitation author="Barel, CDN" journalOrPublisher="Netherlands Journal of Zoology" pageId="23" pageNumber="24" pagination="357 - 424" title="Towards a constructional morphology of cichlid fishes (Teleostei, Perciformes)." url="10.1163/002829683X00183" volume="33" year="1983">Barel 1983</bibRefCitation>
: 384, 385, 407, 418;
<bibRefCitation author="Witte, F" journalOrPublisher="Netherlands Journal of Zoology" pageId="26" pageNumber="27" pagination="596 - 612" title="Consistency and functional significance of morphological differences between wild-caught and domestic Haplochromis squamipinnis (Pisces, Ciclidae)." url="10.1163/002829684X00308" volume="34" year="1984 a">Witte 1984a</bibRefCitation>
: 604, 611;
<bibRefCitation pageId="4" pageNumber="5">Witte 1984b</bibRefCitation>
: 159-161, 163;
<bibRefCitation pageId="4" pageNumber="5">Barel 1985</bibRefCitation>
: chapter 1, 384, 385, 407, 418;
<bibRefCitation author="Gottfried, MD" journalOrPublisher="Copeia" pageId="24" pageNumber="25" pagination="1028 - 1030" title="Developmental transition in feeding morphology of the Midas cichlid." url="10.2307/1445308" volume="1986" year="1986">Gottfried 1986</bibRefCitation>
: 1029;
<bibRefCitation pageId="4" pageNumber="5">Witte 1987</bibRefCitation>
, chapter 1: 611, chapter 2: 67, 76, chapter 3: 8-13, 19;
<bibRefCitation pageId="4" pageNumber="5">Goldschmidt 1989a</bibRefCitation>
: 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;
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Annalenvan het Koninklijk Museum voor Midden-Afrika, Zoologische Wetenschappen" pageId="23" pageNumber="24" pagination="119 - 132" title="Reproductive strategies, subtrophic niche differentiation and the role of competition for food in haplochromine cichlids (Pisces) from Lake Victoria, Tanzania." volume="257" year="1989 b">Goldschmidt 1989b</bibRefCitation>
: 122-126, 129-131;
<bibRefCitation pageId="4" pageNumber="5">Wanink et al. 1989</bibRefCitation>
: 25;
<bibRefCitation author="van der Meer, HJ" journalOrPublisher="Annalenvan het Koninklijk Museum voor Midden-Afrika, Zoologische Wetenschappen" pageId="25" pageNumber="26" pagination="51 - 56" title="Ecological significance of retinal receptor patterns in 4 sympatric haplochromine cichlids." volume="257" year="1989">van der Meer 1989</bibRefCitation>
:
<pageBreakToken pageId="5" pageNumber="6" start="start">52</pageBreakToken>
, 53;
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Acta Biotheoretica" pageId="23" pageNumber="24" pagination="125 - 134" title="On the possible role of egg mimics in speciation." url="10.1007/BF00047549" volume="38" year="1990">Goldschmidt and de Visser 1990</bibRefCitation>
: 129, 130, 132;
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Oikos" pageId="24" pageNumber="25" pagination="343 - 355" title="Ecological segregation of zooplanktivorous haplochromine species (Pisces, Cichlidae) from Lake Victoria." url="10.2307/3545226" volume="58" year="1990">Goldschmidt et al. 1990</bibRefCitation>
: 344, 346-351, 353;
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Oikos" pageId="23" pageNumber="24" pagination="356 - 368" title="Reproductive strategies of zooplanktivorous haplochromine species (Pisces, Cichlidae) from Lake Victoria before the Nile perch boom." url="10.2307/3545227" volume="58" year="1990">Goldschmidt and Witte 1990</bibRefCitation>
: 356-367;
<bibRefCitation pageId="5" pageNumber="6">Barel et al. 1991</bibRefCitation>
: 262;
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Ethology" pageId="23" pageNumber="24" pagination="177 - 190" title="Egg mimics in haplochromine cichlids (Pisces, Perciformes) from Lake Victoria." url="10.1111/j.1439-0310.1991.tb00273.x" volume="88" year="1991">Goldschmidt 1991</bibRefCitation>
: 181, 182, 185, 187;
<bibRefCitation author="van der Meer, HJ" journalOrPublisher="Annalenvan het Koninklijk Museum voor Midden-Afrika, Zoologische Wetenschappen" pageId="25" pageNumber="26" pagination="91 - 96" title="Determination of photopic thresholds in two sympatric cichlids using optomotor response." volume="263" year="1991 a">van der Meer 1991a</bibRefCitation>
: 91-94, 96;
<bibRefCitation pageId="5" pageNumber="6">van der Meer 1991b</bibRefCitation>
: 3, 5, 6, 9-12, 14-22, 24-26, 28, 68, 76, 77, 83, 85-87, 89-93, 99, 101, 104;
<bibRefCitation pageId="5" pageNumber="6">Goldschmidt and Witte 1992</bibRefCitation>
: 104;
<bibRefCitation author="Kaufman, L" journalOrPublisher="Brain, Behaviour and Evolution" pageId="24" pageNumber="25" title="Catastrophic change in species-rich freshwater ecosystems. BioScience 42: 846 - 858." url="10.2307/1312084" year="1992">Kaufman 1992</bibRefCitation>
: 848, 849;
<bibRefCitation author="Witte, F" journalOrPublisher="Environmental Biology of Fishes" pageId="26" pageNumber="27" pagination="1 - 28" title="The destruction of an endemic species flock: quantitative data on the decline of the haplochromine cichlids of Lake Victoria." url="10.1007/BF00004782" volume="34" year="1992 b">Witte et al. 1992b</bibRefCitation>
: 11, 13, 17, 25, 27, 28;
<bibRefCitation author="Barel, CDN" journalOrPublisher="Acta Biotheoretica" pageId="23" pageNumber="24" pagination="345 - 381" title="Concepts of an architectonic approach to transformation morphology." url="10.1007/BF00709371" volume="41" year="1993">Barel 1993</bibRefCitation>
: 366;
<bibRefCitation author="Smith, C" journalOrPublisher="Environmental Biology of Fishes" pageId="25" pageNumber="26" pagination="99 - 104" title="The cost of parental care in Haplochromis ' argens' (Cichlidae)." url="10.1007/BF00002184" volume="40" year="1994">Smith and Wootton 1994</bibRefCitation>
: 99-103;
<bibRefCitation author="Balshine-Earn, S" journalOrPublisher="Animal Behaviour" pageId="23" pageNumber="24" pagination="1 - 7" title="The costs of parental care in Galilee St Peter's fish, Sarotherodon galilaeus." url="10.1006/anbe.1995.0214" volume="50" year="1995">Balshine-Earn 1995</bibRefCitation>
: 5;
<bibRefCitation pageId="5" pageNumber="6">Seehausen 1995</bibRefCitation>
: 143, 146;
<bibRefCitation author="Seehausen, O" journalOrPublisher="Tropical Fish Hobbyist" pageId="25" pageNumber="26" pagination="96 - 105" title="Extinction of many and survival of some: the current situation of the endemic cichlids in southern Lake Victoria." volume="43" year="1995">Seehausen and Witte 1995</bibRefCitation>
: 101;
<bibRefCitation author="van der Meer, HJ" journalOrPublisher="Brain, Behaviour and Evolution" pageId="25" pageNumber="26" pagination="25 - 33" title="Visual resolution during growth in cichlid fish: A morphological and behavioural case study." url="10.1159/000113383" volume="45" year="1995">van der Meer et al. 1995</bibRefCitation>
: 116-129;
<bibRefCitation author="Smith, C" journalOrPublisher="Reviews in Fish Biology and Fisheries" pageId="25" pageNumber="26" pagination="7 - 22" title="The costs of parental care in teleost fishes." url="10.1007/BF01103363" volume="5" year="1995">Smith and Wootton 1995</bibRefCitation>
: 12;
<bibRefCitation pageId="5" pageNumber="6">Anker and Dullemeijer 1996</bibRefCitation>
: 4-11;
<bibRefCitation author="de Visser, J" journalOrPublisher="Journal of Morphology" pageId="23" pageNumber="24" pagination="1 - 18" title="Architectonic constraints on the hyoid's optimal starting position for suction feeding of fish." url="10.1002/(SICI)1097-4687(199604)228:1&lt;1::AID-JMOR1&gt;3.0.CO;2-B" volume="228" year="1996">de Visser and Barel 1996</bibRefCitation>
: 4;
<bibRefCitation author="Seehausen, O" journalOrPublisher="Verduijn Cichlids, Zevenhuizen" pageId="25" pageNumber="26" title="Lake Victoria Rock Cichlids: Taxonomy, Ecology and Distribution." year="1996">Seehausen 1996</bibRefCitation>
: 51, 55, 255, 256;
<bibRefCitation author="Smit, SA" journalOrPublisher="Netherlands Journal of Zoology" pageId="25" pageNumber="26" pagination="9 - 20" title="Photopic sensitivity to red and blue light related to retinal differences in two zooplanktivorous haplochromine species (Teleostei, Cichlidae)." url="10.1163/156854297X00201" volume="47" year="1997">Smit and Anker 1997</bibRefCitation>
: 9-17;
<bibRefCitation author="Seehausen, O" journalOrPublisher="Conservation Biology" pageId="25" pageNumber="26" pagination="890 - 904" title="Patterns of the remnant cichlid fauna in southern Lake Victoria." url="10.1046/j.1523-1739.1997.95346.x" volume="11" year="1997 b">Seehausen et al. 1997b</bibRefCitation>
: 899;
<bibRefCitation pageId="5" pageNumber="6">Witte et al. 1997</bibRefCitation>
: 591;
<bibRefCitation pageId="5" pageNumber="6">Wanink 1998</bibRefCitation>
: 159, 232;
<bibRefCitation pageId="5" pageNumber="6">Rinkes 1999</bibRefCitation>
: 46, 47, 54, 56-58, 60, 62, 76, 77, 79, 97;
<bibRefCitation pageId="5" pageNumber="6">de Visser 2000</bibRefCitation>
: 18;
<bibRefCitation author="Tacon, P" journalOrPublisher="General and Comparative Endocrinology" pageId="25" pageNumber="26" pagination="54 - 65" title="Effect of egg deprivation on sex steroids, gonadotropin, prolactin, and growth hormone profiles during the reproductive cycle of the mouthbrooding cichlid fish Oreochromis niloticus." url="10.1006/gcen.1999.7388" volume="117" year="2000">Tacon et al. 2000</bibRefCitation>
: 63;
<bibRefCitation author="Witte, F" journalOrPublisher="Cichlidae" pageId="26" pageNumber="27" pagination="77 - 84" title="De zooeplanktivore cichliden van het Victoriameer: ontdekking, verdwijning en terugkeer." volume="26" year="2000">Witte and Wanink 2000</bibRefCitation>
: 78-81, 84;
<bibRefCitation pageId="5" pageNumber="6">Seehausen et al. 2003</bibRefCitation>
: 272;
<bibRefCitation pageId="5" pageNumber="6">Witte et al. 2005</bibRefCitation>
: 320;
<bibRefCitation pageId="5" pageNumber="6">Kishe-Machumu 2012</bibRefCitation>
: 35;
<bibRefCitation author="van Rijssel, JC" journalOrPublisher="Zoologische Verhandelingen Leiden" pageId="25" pageNumber="26" title="Adaptive responses in resurgent Lake Victoria cichlids over the past 30 years. Evolutionary Ecology." url="10.1007/s10682-012-9596-9" year="2012">van Rijssel and Witte 2012</bibRefCitation>
.
</paragraph>
<paragraph pageId="5" pageNumber="6">
H.
<normalizedToken originalValue="“Argens”">&quot;Argens&quot;</normalizedToken>
;
<bibRefCitation pageId="5" pageNumber="6">Kaufman and Seehausen 1995</bibRefCitation>
: 148.
</paragraph>
<paragraph pageId="5" pageNumber="6">
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis" order="Perciformes" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Haplochromis</taxonomicName>
(?)
<normalizedToken originalValue="“argens”">&quot;argens&quot;</normalizedToken>
;
<bibRefCitation author="Wanink, JH" journalOrPublisher="Advances in Ecological Research" pageId="26" pageNumber="27" pagination="553 - 570" title="The use of perturbation as a natural experiment: effects of predator introduction on the community structure of zooplanktivorous fish in Lake Victoria." url="10.1016/S0065-2504(00)31030-3" volume="31" year="2000">Wanink and Witte 2000</bibRefCitation>
: 563;
<bibRefCitation author="Witte, F" journalOrPublisher="Reviews in Fish Biology and Fisheries" pageId="26" pageNumber="27" pagination="233 - 241" title="Recovery of cichlid species in Lake Victoria: an examination of factors leading to differential extinction." url="10.1023/A:1016677515930" volume="10" year="2000">Witte et al. 2000</bibRefCitation>
: 234, 235, 237;
<bibRefCitation author="Witte, F" journalOrPublisher="Cichlidae" pageId="26" pageNumber="27" pagination="104 - 114" title="Door cichlidenogen bezien: het belang van het gezichtsvermogen bij haplochromine cichliden voor het ontstaan en in stand houden van diversiteit." volume="29" year="2003">Witte et al. 2003</bibRefCitation>
: 108;
<bibRefCitation author="Niemantsverdriet, P" journalOrPublisher="Cichlidae" pageId="24" pageNumber="25" pagination="150 - 156" title="Victoriacichliden houden en kweken in het laboratorium." volume="31" year="2005">Niemantsverdriet 2005</bibRefCitation>
: 151, 155;
<bibRefCitation author="Witte, F" journalOrPublisher="Transactions of the American Fisheries Society" pageId="26" pageNumber="27" pagination="1146 - 1159" title="Species distinction and the biodiversity crisis in Lake Victoria." url="10.1577/T05-179.1" volume="136" year="2007 a">Witte et al. 2007a</bibRefCitation>
: 1153.
</paragraph>
<paragraph pageId="5" pageNumber="6">
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis" order="Perciformes" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Haplochromis</taxonomicName>
sp.
<normalizedToken originalValue="“argens”">&quot;argens&quot;</normalizedToken>
;
<bibRefCitation author="Mizoiri, S" journalOrPublisher="Kougakutosho Ltd., Tokyo" pageId="24" pageNumber="25" title="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)." year="2008">Mizoiri et al. 2008</bibRefCitation>
: 228, 241, 254.
</paragraph>
<paragraph pageId="5" pageNumber="6">
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Yssichromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yssichromis argens" order="Perciformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="argens">Yssichromis argens</taxonomicName>
;
<bibRefCitation author="van Staaden, MJ" journalOrPublisher="Zoology" pageId="25" pageNumber="26" pagination="165 - 178" title="Brain evolution in cichlids of the African Great Lakes: brain and body size, general patterns, and evolutionary trends." volume="98" year="1995">van Staaden et al. 1995</bibRefCitation>
: 168;
<bibRefCitation author="Chapman, LJ" journalOrPublisher="Conservation Biology" pageId="23" pageNumber="24" pagination="1274 - 1287" title="Hypoxia tolerance in twelve species of East African cichlids: potential for low oxygen refugia in Lake Victoria." url="10.1046/j.1523-1739.1995.9051262.x-i1" volume="9" year="1995">Chapman et al. 1995</bibRefCitation>
: 1277, 1279-1282, 1285;
<bibRefCitation author="Huber, R" journalOrPublisher="Brain, Behaviour and Evolution" pageId="24" pageNumber="25" pagination="167 - 182" title="Microhabitat use, trophic patterns, and the evolution of brain structure in African cichlids." url="10.1159/000113330" volume="50" year="1997">Huber et al. 1997</bibRefCitation>
: 170;
<bibRefCitation author="Rosenberger, AE" journalOrPublisher="Journal of Fish Biology" pageId="25" pageNumber="26" pagination="483 - 501" title="Respiratory characters of three species of haplochromine cichlids: Implications for use of wetland refugia." url="10.1111/j.1095-8649.2000.tb02187.x" volume="57" year="2000">Rosenberger and Chapman 2000</bibRefCitation>
: 498;
<bibRefCitation author="Melnychuk, MC" journalOrPublisher="Environmental Biology of Fishes" pageId="24" pageNumber="25" pagination="99 - 110" title="Hypoxia tolerance of two haplochromine cichlids: swamp leakage and potential for interlacustrine dispersal." url="10.1023/A:1019602403098" volume="65" year="2002">Melnychuk and Chapman 2002</bibRefCitation>
: 107.
</paragraph>
<paragraph pageId="5" pageNumber="6">
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Yssichromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yssichromis" order="Perciformes" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Yssichromis</taxonomicName>
sp.
<normalizedToken originalValue="“argens”">&quot;argens&quot;</normalizedToken>
;
<bibRefCitation author="Mizoiri, S" journalOrPublisher="Kougakutosho Ltd., Tokyo" pageId="24" pageNumber="25" title="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)." year="2008">Mizoiri et al. 2008</bibRefCitation>
: 228.
</paragraph>
<paragraph pageId="5" pageNumber="6">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).</paragraph>
<paragraph pageId="5" pageNumber="6">Holotype. RMNH.PISC.835884, ♂, 67.3 mm SL, Tanzania, Lake Victoria, Mwanza Gulf, 8.iii.1979, HEST.</paragraph>
<paragraph lastPageId="7" lastPageNumber="8" pageId="5" pageNumber="6">
Paratypes. All type specimens collected by
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis" order="Perciformes" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Haplochromis</taxonomicName>
Ecology Survey Team (HEST) in Mwanza Gulf, Tanzania, Lake Victoria, except where noted otherwise.
<pageBreakToken pageId="6" pageNumber="7" start="start">Size</pageBreakToken>
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
<pageBreakToken pageId="7" pageNumber="8" start="start">,</pageBreakToken>
♂, 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.
</paragraph>
<paragraph pageId="7" pageNumber="8">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.</paragraph>
</subSubSection>
<subSubSection lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8" type="diagnosis">
<paragraph pageId="7" pageNumber="8">Diagnosis.</paragraph>
<paragraph lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
Small sized (&lt;8 cm SL), slender (BD &lt;31% SL), micrognathic (LJL &lt;45% HL) zooplanktivorous
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis" order="Perciformes" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Haplochromis</taxonomicName>
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
<pageBreakToken pageId="8" pageNumber="9" start="start">males</pageBreakToken>
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.
</paragraph>
</subSubSection>
<subSubSection lastPageId="10" lastPageNumber="11" pageId="8" pageNumber="9" type="description">
<paragraph pageId="8" pageNumber="9">Description.</paragraph>
<paragraph pageId="8" pageNumber="9">Proportional measurements of type material given in Table 1.</paragraph>
<paragraph pageId="8" pageNumber="9">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.</paragraph>
<caption pageId="8" pageNumber="9">
<paragraph pageId="8" pageNumber="9">
Figure 2. Habitus of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
sp. n. ♂ (holotype, RMNH.PISC.83588). Scale bar equals 10 mm. Drawing by I. Westbroek, missing (= dotted) scales added by M. van Oijen.
</paragraph>
</caption>
<paragraph pageId="8" pageNumber="9">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.</paragraph>
<paragraph pageId="8" pageNumber="9">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.</paragraph>
<paragraph pageId="8" pageNumber="9">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.</paragraph>
<paragraph pageId="8" pageNumber="9">Viscera. Ratio between intestine length and SL: 1.0-1.4 (n = 25).</paragraph>
<paragraph pageId="8" pageNumber="9">
Oral jaws. (Fig. 3
<normalizedToken originalValue="AC">A-C</normalizedToken>
) 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.
</paragraph>
<caption pageId="8" pageNumber="9">
<paragraph pageId="8" pageNumber="9">
Figure 3. Skeletal elements of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
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.
</paragraph>
</caption>
<paragraph lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
Oral teeth shape. (Fig. 3
<normalizedToken originalValue="AC">A-C</normalizedToken>
) 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
<pageBreakToken pageId="9" pageNumber="10" start="start">view</pageBreakToken>
, crown compressed. Outer-row teeth in both premaxilla and lower jaw recurved. Inner rows in both jaws with mainly tricuspid or weakly tricuspid teeth.
</paragraph>
<paragraph pageId="9" pageNumber="10">Oral teeth size. Outer-row teeth relatively long and slender, gradually decreasing in size from rostral to caudal.</paragraph>
<paragraph pageId="9" pageNumber="10">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.</paragraph>
<paragraph pageId="9" pageNumber="10">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.</paragraph>
<paragraph pageId="9" pageNumber="10">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.</paragraph>
<paragraph pageId="9" pageNumber="10">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.</paragraph>
<paragraph pageId="9" pageNumber="10">Pharyngeal teethcounts. Caudal-most transverse row with about 30-38 teeth, medial longitudinal rows with eight to 11 teeth.</paragraph>
<paragraph pageId="9" pageNumber="10">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.</paragraph>
<paragraph pageId="9" pageNumber="10">Vertebrae. Total number of vertebrae in 57 specimens: 30 (12), 31 (39) or 32 (6), comprising 13-14 abdominal and 16-19 caudal vertebrae.</paragraph>
<paragraph lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
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
<pageBreakToken pageId="10" pageNumber="11" start="start">flank</pageBreakToken>
silvery-grey, dorsum with bluish to purplish sheen, flank with yellow to greenish sheen. Chest, belly and ventral side silvery-white.
</paragraph>
<caption pageId="10" pageNumber="11">
<paragraph pageId="10" pageNumber="11">
Figure 4. Live colours of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
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.
</paragraph>
</caption>
<paragraph pageId="10" pageNumber="11">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.</paragraph>
<paragraph pageId="10" pageNumber="11">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.</paragraph>
<paragraph pageId="10" pageNumber="11">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.</paragraph>
<paragraph pageId="10" pageNumber="11">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.</paragraph>
<caption pageId="10" pageNumber="11">
<paragraph pageId="10" pageNumber="11">
Figure 5. Preserved colours of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
sp. n. ♂ 67.3 mm SL (holotype, RMNH.PISC.83588). Scale bar equals 10 mm.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="10" pageNumber="11" type="distribution">
<paragraph pageId="10" pageNumber="11">Distribution.</paragraph>
<paragraph pageId="10" pageNumber="11">
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
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).
</paragraph>
</subSubSection>
<subSubSection pageId="10" pageNumber="11" type="habitat">
<paragraph pageId="10" pageNumber="11">Habitat.</paragraph>
<paragraph pageId="10" pageNumber="11">
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
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 (
<bibRefCitation pageId="10" pageNumber="11">Witte 1984b</bibRefCitation>
,
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Oikos" pageId="24" pageNumber="25" pagination="343 - 355" title="Ecological segregation of zooplanktivorous haplochromine species (Pisces, Cichlidae) from Lake Victoria." url="10.2307/3545226" volume="58" year="1990">Goldschmidt et al. 1990</bibRefCitation>
). 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 (
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Oikos" pageId="24" pageNumber="25" pagination="343 - 355" title="Ecological segregation of zooplanktivorous haplochromine species (Pisces, Cichlidae) from Lake Victoria." url="10.2307/3545226" volume="58" year="1990">Goldschmidt et al. 1990</bibRefCitation>
,
<bibRefCitation author="Witte, F" journalOrPublisher="Environmental Biology of Fishes" pageId="26" pageNumber="27" pagination="1 - 28" title="The destruction of an endemic species flock: quantitative data on the decline of the haplochromine cichlids of Lake Victoria." url="10.1007/BF00004782" volume="34" year="1992 b">Witte et al. 1992b</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="11" lastPageNumber="12" pageId="10" pageNumber="11" type="abundance">
<paragraph pageId="10" pageNumber="11">Abundance.</paragraph>
<paragraph pageId="10" pageNumber="11">
During 1979-1982,
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
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
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
. From 2001 to 2011 more than 150 bottom-trawl tows contained about 15 individuals of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
, 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.
</paragraph>
<caption pageId="11" pageNumber="12">
<paragraph pageId="11" pageNumber="12">
<pageBreakToken pageId="11" pageNumber="12" start="start">Table</pageBreakToken>
2. Frequencies of occurrence (Foo) in trawl tows and mean numbers (
<normalizedToken originalValue="±">+/-</normalizedToken>
SD) per tow of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
. 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.
</paragraph>
</caption>
<paragraph pageId="11" pageNumber="12">
<table pageId="11" pageNumber="12">
<tr pageId="11" pageNumber="12">
<th colspan="1" pageId="11" pageNumber="12" rowspan="1">Station</th>
<th colspan="1" pageId="11" pageNumber="12" rowspan="1">Bottom trawl day</th>
<th colspan="1" pageId="11" pageNumber="12" rowspan="1">
1979-1982
<sup pageId="11" pageNumber="12"></sup>
n = 104
</th>
<th colspan="1" pageId="11" pageNumber="12" rowspan="1">
1987-1988
<sup pageId="11" pageNumber="12"></sup>
n = 29
</th>
<th colspan="1" pageId="11" pageNumber="12" rowspan="1">
1990-1999
<sup pageId="11" pageNumber="12"></sup>
n = 34
</th>
<th colspan="1" pageId="11" pageNumber="12" rowspan="1">
2001-2011
<sup pageId="11" pageNumber="12">§</sup>
n&gt; 150
</th>
</tr>
<tr pageId="11" pageNumber="12">
<td colspan="1" pageId="11" pageNumber="12" rowspan="1">Surface trawl night</td>
<td colspan="1" pageId="11" pageNumber="12" rowspan="1">1981/1982 n = 8</td>
<td colspan="1" pageId="11" pageNumber="12" rowspan="1">1987-19888| n = 26</td>
<td colspan="1" pageId="11" pageNumber="12" rowspan="1">-</td>
<td colspan="1" pageId="11" pageNumber="12" rowspan="1">2001-20111§ n = 15</td>
</tr>
</table>
</paragraph>
<paragraph pageId="11" pageNumber="12">
Food. Before the ecological changes in Lake Victoria,the diet of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
comprised mainly zooplankton during the day;
<taxonomicName class="Insecta" family="Chaoboridae" genus="Chaoborus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chaoborus" order="Diptera" pageId="11" pageNumber="12" phylum="Arthropoda" rank="genus">Chaoborus</taxonomicName>
larvae were important at night (
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Oikos" pageId="24" pageNumber="25" pagination="343 - 355" title="Ecological segregation of zooplanktivorous haplochromine species (Pisces, Cichlidae) from Lake Victoria." url="10.2307/3545226" volume="58" year="1990">Goldschmidt et al. 1990</bibRefCitation>
). The current diet is unknown, but all the studied resurgent species in the Mwanza Gulf changed their diet (e.g.
<bibRefCitation author="van Oijen, MJP" journalOrPublisher="Zoologische Verhandelingen Leiden" pageId="25" pageNumber="26" pagination="1 - 56" title="Taxonomical and ecological description of a species complex of zooplanktivorous and insectivorous cichlids from Lake Victoria." volume="302" year="1996">van Oijen and Witte 1996</bibRefCitation>
,
<bibRefCitation author="Katunzi, EFB" journalOrPublisher="Ecology of Freshwater Fish" pageId="24" pageNumber="25" pagination="237 - 240" title="Lost zooplanktivorous cichlid from Lake Victoria reappears with a new trade." url="10.1046/j.1600-0633.2003.00023.x" volume="12" year="2003">Katunzi et al. 2003</bibRefCitation>
,
<bibRefCitation author="Kishe-Machumu, M" journalOrPublisher="Animal Biology" pageId="24" pageNumber="25" pagination="401 - 417" title="Dietary shift in benthivorous cichlids after the ecological changes in Lake Victoria." url="10.1163/157075608X383700" volume="58" year="2008">Kishe-Machumu et al. 2008</bibRefCitation>
).
</paragraph>
<paragraph pageId="11" pageNumber="12">
Breeding.
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
is a female mouth brooder. Spawning sites are located at depths &lt;9 m (
<bibRefCitation author="Goldschmidt, T" journalOrPublisher="Oikos" pageId="23" pageNumber="24" pagination="356 - 368" title="Reproductive strategies of zooplanktivorous haplochromine species (Pisces, Cichlidae) from Lake Victoria before the Nile perch boom." url="10.2307/3545227" volume="58" year="1990">Goldschmidt and Witte 1990</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="11" pageNumber="12" type="etymology">
<paragraph pageId="11" pageNumber="12">Etymology.</paragraph>
<paragraph pageId="11" pageNumber="12">
In reference to the silver male colouration,
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
was given the nickname
<normalizedToken originalValue="“argens”">&quot;argens&quot;</normalizedToken>
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
<normalizedToken originalValue="species">species'</normalizedToken>
epithethon.
</paragraph>
</subSubSection>
<subSubSection lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12" type="comparisons">
<paragraph pageId="11" pageNumber="12">Comparisons.</paragraph>
<paragraph lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
The zooplanktivorous species
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis (Yssichromis) laparogramma" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="laparogramma" subGenus="Yssichromis">Haplochromis (Yssichromis) laparogramma</taxonomicName>
Greenwood &amp; Gee, 1969,
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis (Yssichromis) pyrrhocephalus" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="pyrrhocephalus" subGenus="Yssichromis">Haplochromis (Yssichromis) pyrrhocephalus</taxonomicName>
Witte &amp; Witte-Maas, 1987 and
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis (Yssichromis) heusinkveldi" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="heusinkveldi" subGenus="Yssichromis">Haplochromis (Yssichromis) heusinkveldi</taxonomicName>
, Witte &amp; Witte-Maas, 1987, have shorter bicuspid teeth in the oral jaws than
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
, 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
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis tanaos" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="tanaos">Haplochromis tanaos</taxonomicName>
van Oijen &amp; Witte, 1996 and
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis thereuterion" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="thereuterion">Haplochromis thereuterion</taxonomicName>
van Oijen &amp; Witte, 1996, are more or less similar to those of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
, but the former two species have more unicuspids.
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
is further distinguished from these and other species by its colouration. Sexually active males of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis tanaos" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="tanaos">Haplochromis tanaos</taxonomicName>
are dark blue, the females silvery with a distinct mid-lateral band and slightly less distinct dorso-lateral band. Sexually active males of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis thereuterion" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="thereuterion">Haplochromis thereuterion</taxonomicName>
are black, the females coloured like females of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis tanaos" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="tanaos">Haplochromis tanaos</taxonomicName>
(
<bibRefCitation author="van Oijen, MJP" journalOrPublisher="Zoologische Verhandelingen Leiden" pageId="25" pageNumber="26" pagination="1 - 56" title="Taxonomical and ecological description of a species complex of zooplanktivorous and insectivorous cichlids from Lake Victoria." volume="302" year="1996">van Oijen and Witte 1996</bibRefCitation>
). The body of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
is less slender (BD 26.0-30.7% of SL, mean 28.2%, Table 1) than that of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis tanaos" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="tanaos">Haplochromis tanaos</taxonomicName>
and
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis thereuterion" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="thereuterion">Haplochromis thereuterion</taxonomicName>
(22.1-27.1% and 24.4-27.6% of SL, respectively; Tables 3 and 7 in
<bibRefCitation author="van Oijen, MJP" journalOrPublisher="Zoologische Verhandelingen Leiden" pageId="25" pageNumber="26" pagination="1 - 56" title="Taxonomical and ecological description of a species complex of zooplanktivorous and insectivorous cichlids from Lake Victoria." volume="302" year="1996">van Oijen and Witte 1996</bibRefCitation>
). Live
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
is similar to (juvenile)
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis cassius" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="cassius">Haplochromis cassius</taxonomicName>
in colouration and general habitus. However,
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis cassius" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="cassius">Haplochromis cassius</taxonomicName>
has a broad, well defined mid-lateral band (
<bibRefCitation pageId="11" pageNumber="12">Greenwood and Barel 1978</bibRefCitation>
), more and distinctly longer unicuspid teeth, more
<pageBreakToken pageId="12" pageNumber="13" start="start">curved</pageBreakToken>
and more widely set teeth, and thicker lips than
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
. The maximum size of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis cassius" order="Perciformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="cassius">Haplochromis cassius</taxonomicName>
(99.0 mm SL) is larger than that of
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
, but when comparing similar size ranges (see material and methods)
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis argens" order="Perciformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="argens">Haplochromis argens</taxonomicName>
has: a smaller head (ratio HL/SL: 31.0 - 35.6%, mean 32.9% versus 35.0 - 36.0%, mean 35.5% in
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis cassius" order="Perciformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="cassius">Haplochromis cassius</taxonomicName>
); a shorter snout (ratio SnL/HL: 23.6 - 29.7%, mean 26.6% versus 29.8 - 32.6%, mean 31.2% in
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis cassius" order="Perciformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="cassius">Haplochromis cassius</taxonomicName>
); larger eyes (ratio EyL/HL: 30.9 - 39.0%, mean 35.7% versus 26.8 - 28.8%, mean 28.3% in
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis cassius" order="Perciformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="cassius">Haplochromis cassius</taxonomicName>
). For comparison with
<taxonomicName class="Actinopterygii" family="Cichlidae" genus="Haplochromis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Haplochromis goldschmidti" order="Perciformes" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="goldschmidti">Haplochromis goldschmidti</taxonomicName>
sp. n., see below.
</paragraph>
</subSubSection>
</treatment>
</document>