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<mods:title id="B09B1897A3D71E5FF068478139F4EBF2">Trematocranuspachychilus, a new endemic cichlid from Lake Malawi (Teleostei, Cichlidae)</mods:title>
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<mods:namePart id="164E2E47D1846A49755A4AE3E8EE7653">Dierickx, Katrien</mods:namePart>
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<mods:namePart id="86B1973A6E8C8DF075B7FE64B99A358A">Hanssens, Mark</mods:namePart>
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<treatment id="1D4F155A974554AA3D5E3D99A91C42DF" ID-GBIF-Taxon="142283967" LSID="urn:lsid:zoobank.org:act:50FF20F1-74E0-4C11-90AC-37FAB2121378" httpUri="http://treatment.plazi.org/id/1D4F155A974554AA3D5E3D99A91C42DF" lastPageId="6" lastPageNumber="158" pageId="3" pageNumber="156">
<subSubSection id="BBF4BF31C13EE9405DD6A1861EB8C877" pageId="3" pageNumber="156" type="nomenclature">
<paragraph id="0B72F6B670BEE00AF45718FFA3FFA701" pageId="3" pageNumber="156">
<taxonomicName id="F064B755A02F1AD6A11EFF5EFC34A3F9" LSID="http://zoobank.org/50FF20F1-74E0-4C11-90AC-37FAB2121378" family="Cichlidae" genus="Trematocranus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trematocranus pachychilus" order="Perciformes" pageId="3" pageNumber="156" phylum="Chordata" rank="species" species="pachychilus">Trematocranus pachychilus</taxonomicName>
<taxonomicNameLabel id="8ABF150A466C6815D75060EAE0272365" pageId="3" pageNumber="156">sp. n.</taxonomicNameLabel>
</paragraph>
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<subSubSection id="6884491188023E5D9505F1529E6F3F2E" pageId="3" pageNumber="156" type="type material">
<paragraph id="E9F02E93E436DE6F978042726E3BED0C" pageId="3" pageNumber="156">Type material.</paragraph>
<paragraph id="FCA1A8EA6403E8DBDEAAC9434A7FB213" pageId="3" pageNumber="156">Holotype: MRAC 99-041-P-4781, 154.6 mm SL, Jafua Bay, Lake Malawi, Mozambique, 12°07.13'S, 34°45.89'E, Coll. Snoeks &amp; Hanssens (4 April 1998). Paratypes: MRAC 99-041-P-4782, 151.9 mm SL, same data as holotype; MRAC 99-041-P-5037-5041 (5), 117.7 - 154.5 mm SL, same data as holotype.</paragraph>
</subSubSection>
<subSubSection id="65048F9F8AD80C4E827F70F45B4395CE" lastPageId="5" lastPageNumber="158" pageId="3" pageNumber="156" type="differential diagnosis">
<paragraph id="10971F572E1DA83282E740C051D525D9" pageId="3" pageNumber="156">Differential diagnosis.</paragraph>
<paragraph id="C393CED37A5D8F8D7B6C6244832CDE47" pageId="3" pageNumber="156">
<taxonomicName id="C2FB90E3D7138523FBDFB1639162AFC7" family="Cichlidae" genus="Trematocranus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trematocranus pachychilus" order="Perciformes" pageId="3" pageNumber="156" phylum="Chordata" rank="species" species="pachychilus">Trematocranus pachychilus</taxonomicName>
is clearly distinct from all other known species of the genus by its thick lips. It can further be distinguished from
<taxonomicName id="3B85E1F8FCCC99C24CE494A5F6F17D51" lsidName="T. labifer" pageId="3" pageNumber="156" rank="species" species="labifer">T. labifer</taxonomicName>
by its molariform pharyngeal dentition while the latter has slender teeth on the lower pharyngeal jaw (
<bibRefCitation id="38731BB265A3B2657C2A81F8F8629043" author="Eccles, DH" journalOrPublisher="Lake Fish Movies, West Germany" pageId="8" pageNumber="161" title="Malawian cichlid fishes: the classification of some haplochromine genera." year="1989">Eccles and Trewavas 1989</bibRefCitation>
).
</paragraph>
<paragraph id="33D9FEA765406F307730C6ADD6002719" lastPageId="4" lastPageNumber="157" pageId="3" pageNumber="156">
In
<taxonomicName id="E6E64A855B18624902B3B55E0B0F5B35" lsidName="T. placodon" pageId="3" pageNumber="156" rank="species" species="placodon">T. placodon</taxonomicName>
(Fig. 5C, D) lips are usually very small. While
<taxonomicName id="6E247E6EDADD1F675B2401A7C8BF7BC6" lsidName="T. pachychilus" pageId="3" pageNumber="156" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
has a rather concave head, as in
<taxonomicName id="772C9962180DC86373D04843B5FF662F" lsidName="T. microstoma" pageId="3" pageNumber="156" rank="species" species="microstoma">T. microstoma</taxonomicName>
,
<taxonomicName id="8041A6B9E31D511F129BDE0B42175B50" lsidName="T. placodon" pageId="3" pageNumber="156" rank="species" species="placodon">T. placodon</taxonomicName>
has a more distinct convex
<pageBreakToken id="2E71CB2B804DDD5ACB938856E1DA098B" pageId="4" pageNumber="157" start="start">head</pageBreakToken>
. In both
<taxonomicName id="9237D91F9EF1FE531A5FDC4298E747F3" lsidName="T. pachychilus" pageId="4" pageNumber="157" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
and
<taxonomicName id="995BAE9C8306A4E5AFEC1A0D55EF0289" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
, the smaller specimens have a straighter head profile. Coincidentally, in two large specimens of
<taxonomicName id="47825FE806C167BFBBD7FA37201B2D2A" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
, found at Jafua Bay, the head is slightly concave. The teeth of
<taxonomicName id="55AB958AD9AF55A09C170859A0791100" lsidName="T. pachychilus" pageId="4" pageNumber="157" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
are less broad, more densely spaced and slightly less numerous (36-47 vs. 41-57) than in
<taxonomicName id="0EE9D59B460D86B75C1900C9172EA204" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
.
<taxonomicName id="0A549142A22FE64201D70E23BB57B30E" family="Cichlidae" genus="Trematocranus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trematocranus pachychilus" order="Perciformes" pageId="4" pageNumber="157" phylum="Chordata" rank="species" species="pachychilus">Trematocranus pachychilus</taxonomicName>
has more upper lateral line scales (22-24 vs. 19-22) and more dorsal fin rays (11-12 vs. 9-11) than
<taxonomicName id="F6D02395ABF51FE6E12140ACC1719E09" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
. The pharyngeal bones of
<taxonomicName id="238AE4DF294003960043C2A4C3451347" lsidName="T. pachychilus" pageId="4" pageNumber="157" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
are less developed than in
<taxonomicName id="B03FB54C686AF32ED8D48C6D845702FF" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
. They are shallower (9.4-10.4% HL vs 13.8-16.8% HL), narrower (29.7-33.2% HL vs. 36.1-48.7% HL) and shorter (21.8-30.8% HL vs. 29.8-35.9% HL) than those of
<taxonomicName id="9048A1F27BC52CA8FFDF9BA74DD7D3E7" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
.
</paragraph>
<caption id="CB6387484B1903FD8045F0235D1D59FB" pageId="4" pageNumber="157">
<paragraph id="C8D8F866F43FCBAF0ED3728615D2A402" pageId="4" pageNumber="157">
Figure 5. Photographs of preserved specimens: A
<taxonomicName id="4E714E8C4FAE6EB0213183FB9950DDFA" family="Cichlidae" genus="Trematocranus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trematocranus pachychilus" order="Perciformes" pageId="4" pageNumber="157" phylum="Chordata" rank="species" species="pachychilus">Trematocranus pachychilus</taxonomicName>
, holotype, MRAC 99-041-P-4781, adult male, 154.6 mm SL, Lake Malawi, Jafua Bay B
<taxonomicName id="B714F24F1115EC21F1D4B08BC3498F50" lsidName="T. microstoma" pageId="4" pageNumber="157" rank="species" species="microstoma">T. microstoma</taxonomicName>
, MRAC 99-041-P-4787, 143.7 mm SL, Lake Malawi, Mazinzi Bay C
<taxonomicName id="49C2FD9C5E868854053B3F50CAE90C42" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
, MRAC 99-041-P-4776, 153.8 mm SL, Lake Malawi, Jafua Bay D
<taxonomicName id="4E241D3F530717398D3F735942A33690" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
, MRAC 99-041-P-4798, 149.3 mm SL, Lake Malawi, Senga Bay. Scale bar: 2 cm.
</paragraph>
</caption>
<paragraph id="A6F3A167FB6C8722D072AE86AD6117F8" pageId="4" pageNumber="157">
The dentigerous area is also narrower than in
<taxonomicName id="2B649A152584433F1A9B2D9EC1E98179" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
(21.9-24.4% HL vs. 24.2-27.5% HL). Both species have large, molariform pharyngeal teeth, although in
<taxonomicName id="6F22D23515F22A26B06BD1197696B27B" lsidName="T. pachychilus" pageId="4" pageNumber="157" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
the lateral teeth are smaller and more numerous than in
<taxonomicName id="88228BE479A53D2D0DB336573BD57772" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
. The number of teeth on the posterior pharyngeal row in
<taxonomicName id="20495B677A2F7614DFF5226DB4C610BD" lsidName="T. pachychilus" pageId="4" pageNumber="157" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
is higher than in
<taxonomicName id="CF99F3F679DA0C306501F970A3167E2E" lsidName="T. placodon" pageId="4" pageNumber="157" rank="species" species="placodon">T. placodon</taxonomicName>
(23-27 vs.11-23).
</paragraph>
<paragraph id="C3A975124E511FA95904DB4B7B96AC30" lastPageId="5" lastPageNumber="158" pageId="4" pageNumber="157">
In
<taxonomicName id="1D105BAE40FE03545CDBCE87C8583730" lsidName="T. microstoma" pageId="4" pageNumber="157" rank="species" species="microstoma">T. microstoma</taxonomicName>
(Fig. 5B) the lips are intermediate except for some specimens of Mazinzi Bay (see discussion). The snout of
<taxonomicName id="7FF0A8821F4AC66E422C5DBD4423AB74" lsidName="T. pachychilus" pageId="4" pageNumber="157" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
is shorter than that of
<taxonomicName id="D245805C29EEF533EC0F7794D6C23047" lsidName="T. microstoma" pageId="4" pageNumber="157" rank="species" species="microstoma">T. microstoma</taxonomicName>
(34.8-40.9% HL vs. 38.0-42.4% HL).
<taxonomicName id="A7418DF4AA9F963B3BCD6C0FC84EE570" family="Cichlidae" genus="Trematocranus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trematocranus pachychilus" order="Perciformes" pageId="4" pageNumber="157" phylum="Chordata" rank="species" species="pachychilus">Trematocranus pachychilus</taxonomicName>
has bicuspid oral teeth on the outer rows while
<taxonomicName id="97F1E31B569CEF6A91E1EE74DB5DCA05" lsidName="T. microstoma" pageId="4" pageNumber="157" rank="species" species="microstoma">T. microstoma</taxonomicName>
has slender unicuspid teeth. It has fewer inner teeth rows (3-5 vs. 6-8) on the lower jaw than
<taxonomicName id="71780AD6BC1581F45AC0CEEDAC5C924A" lsidName="T. microstoma" pageId="4" pageNumber="157" rank="species" species="microstoma">T. microstoma</taxonomicName>
. The lachrymal depth is clearly shorter than in
<taxonomicName id="4A0669DCBE95570616EDEB0B7C76BECC" lsidName="T. microstoma" pageId="4" pageNumber="157" rank="species" species="microstoma">T. microstoma</taxonomicName>
(22.9-29.7% HL vs. 29.3-32.0% HL), while the eye diameter is larger (25.2-29.5% HL vs. 21.2-25.5% HL). The pharyngeal bones (Fig. 6) are more developed in
<taxonomicName id="E795D0783D7BB013A9499830E2FB6789" lsidName="T. pachychilus" pageId="4" pageNumber="157" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
than in
<taxonomicName id="237060CF3F676DC2F6B10AD8204B6E4F" lsidName="T. microstoma" pageId="4" pageNumber="157" rank="species" species="microstoma">T. microstoma</taxonomicName>
.
<pageBreakToken id="35752C883DB6C08621D57DC90D430B30" pageId="5" pageNumber="158" start="start">The</pageBreakToken>
dentigerous area of
<taxonomicName id="D88BD6E3D87CB1D42CF03A9D4B7A8524" lsidName="T. pachychilus" pageId="5" pageNumber="158" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
is wider than in
<taxonomicName id="AFABFCB47490DD7B0E228CEEC9DD3963" lsidName="T. microstoma" pageId="5" pageNumber="158" rank="species" species="microstoma">T. microstoma</taxonomicName>
(21.9-24.4% HL vs. 18.8-23.6% HL). The dentition of the lower pharyngeal bone is similar in both species, with the median teeth enlarged and the lateral teeth small and numerous.
</paragraph>
<caption id="64293831D3AB5FBC5ADCB2DC2366520D" pageId="5" pageNumber="158">
<paragraph id="8A84FC860E209EDA5AB1FB1C1C4776BD" pageId="5" pageNumber="158">
Figure 6. Lower pharyngeal bones, dorsal view (left) and lateral view (right): A.
<taxonomicName id="FC1BD70E572B3ED89C8B6259A8BE7365" family="Cichlidae" genus="Trematocranus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trematocranus pachychilus" order="Perciformes" pageId="5" pageNumber="158" phylum="Chordata" rank="species" species="pachychilus">Trematocranus pachychilus</taxonomicName>
, paratype, MRAC 99-041-P-4782, 151.9 mm SL, Lake Malawi, Jafua Bay; B.
<taxonomicName id="128E1F2629FE827C6825F1750BF8FFA9" lsidName="T. microstoma" pageId="5" pageNumber="158" rank="species" species="microstoma">T. microstoma</taxonomicName>
, MRAC 99-041-P-4787, 143.7 mm SL; C.
<taxonomicName id="7567F11CEDD21B83B21C52FCDCE43D31" lsidName="T. placodon" pageId="5" pageNumber="158" rank="species" species="placodon">T. placodon</taxonomicName>
, MRAC 99-041-P-4776, 153.8 mm SL. Scale bar: 5 mm.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="1DA2C27548490B33120E1240F25941AC" pageId="5" pageNumber="158" type="description">
<paragraph id="C72F982311F507C23130C785F66200E3" pageId="5" pageNumber="158">Description.</paragraph>
<paragraph id="DFCAB0ECEDE0B1F815E52D112C8728D6" pageId="5" pageNumber="158">
Based on the holotype and six paratypes examined. See Figures 5A and 6A. Qualitative observations are made in the context of Lake Malawi haplochromine cichlids as was done by
<bibRefCitation id="672A52CD3E83409C33E44C1B42236859" pageId="5" pageNumber="158">Snoeks (2004)</bibRefCitation>
.
</paragraph>
<paragraph id="45D532A874906987C2E3789172150C30" pageId="5" pageNumber="158">Body deep and laterally compressed. Head steep. Most specimens with a slight concavity at eye level; some, especially smaller specimens, with a merely straight head profile. Snout pointed. Mouth terminal. Lips very thick and equally developed in most specimens. Larger specimens with thicker lips than small specimens; smaller specimens often with a clear boundary between lower lip and chin. Maxilla does not extend to level of eye.</paragraph>
<paragraph id="A4BC8651E9539ED1E6CDA30B66D25845" pageId="5" pageNumber="158">Teeth not readily observable, being to a large extent or fully covered by the fleshy gums. Outer row of teeth on upper and lower jaws with mainly unequally bicuspid and some unicuspid teeth in the postero-lateral parts; teeth slender, generally straight and slightly curved inwards; crown slightly wider than base. Anterior bicuspid teeth more pointed and sharp; lateral bicuspid teeth often rounded. Inner rows mostly with unicuspid teeth. Larger specimens with more bicuspid teeth anteriorly in inner rows. Inner teeth rows difficult to observe; 3-4 inner rows on upper jaw (counted in four specimens); 3-5 on lower jaw (six specimens).</paragraph>
<paragraph id="A640F09F4CB4425281CD413E0099DBA5" pageId="5" pageNumber="158">Pectoral fins implanted slightly behind level of dorsal fin origin. Pelvic fin origin positioned slightly more backwards. Anal fin implanted anterior to level of first soft dorsal fin ray.</paragraph>
<paragraph id="A193D079D7DCE855F14A75EB50CAF83A" pageId="5" pageNumber="158">Lower pharyngeal bone triangular, large, wider than long, and deep. Teeth molariform; central teeth much larger than lateral teeth. Outermost teeth on the posterior rows very small and numerous.</paragraph>
</subSubSection>
<subSubSection id="4182BBF7B57E02A336C0615D5B661753" pageId="5" pageNumber="158" type="colour pattern in preservation">
<paragraph id="27A5307550C88CBE4D80C3686FAA43E7" pageId="5" pageNumber="158">Colour pattern in preservation.</paragraph>
<paragraph id="F4EBC0B87BF5926EAE5457E628D4239F" pageId="5" pageNumber="158">Body generally brown; some specimens slightly more greyish. Dorsum darker than belly. Very dark-coloured on the dorsal parts of head and body contiguous with dorsal fin base in most specimens. Some large darker spots usually on operculum, supra-pectoral in front of dorsal fin origin, dorso-lateral above and on upper lateral line in middle of flank, supra-anal near end of upper lateral line, and caudal peduncle. Clear dark maculae on spiny part of dorsal fin; sometimes also on soft dorsal fin part and caudal fin. Pelvic and anal fins yellowish brown in females; fin base and distal part clearly darker in males. One specimen with a vague brown midlateral stripe along flank.</paragraph>
</subSubSection>
<subSubSection id="D8954A0F9F7FA990420296C1680E87A9" pageId="5" pageNumber="158" type="etymology">
<paragraph id="0688F97B62FEB82143BDC07AD87A0E45" pageId="5" pageNumber="158">Etymology.</paragraph>
<paragraph id="B61129DB80A29F97D4706035CE5C8E1F" pageId="5" pageNumber="158">
The specific name,
<taxonomicName id="8A03641DB459A6773FF1AC5192754469" lsidName="T pachychilus" pageId="5" pageNumber="158" rank="species" species="pachychilus">pachychilus</taxonomicName>
, means thick-lip and is derived from the Greek words
<normalizedToken id="4B3BCF1C7273FF7644FD86F06C23B574" originalValue="παχυς">παxυς</normalizedToken>
(fat, adipose, plump) and
<normalizedToken id="72C45DBDFA6255B3F47EBD7192341B49" originalValue="χειλος">ιλος</normalizedToken>
(lip, edge) with reference to the diagnostic character, the thick lips.
</paragraph>
</subSubSection>
<subSubSection id="DE8777757AA4DB4A71C15C30EFABB9AD" pageId="5" pageNumber="158" type="distribution">
<paragraph id="7EFD9A8B0CE861ACA39E0914849A6D4E" pageId="5" pageNumber="158">Geographical distribution.</paragraph>
<paragraph id="C5A05A00E45ECF657F06DB59FB08003E" pageId="5" pageNumber="158">
The specimens of
<taxonomicName id="F77E4457F6096D854C95AC00BAF59AA8" lsidName="T. pachychilus" pageId="5" pageNumber="158" rank="species" species="pachychilus">T. pachychilus</taxonomicName>
were only found at Jafua Bay, north of Cobue in Lake Malawi, Mozambique. All specimens were caught in a single haul of a bottom trawl at a depth of 10.8-12.8 m (
<geoCoordinate id="B4D9F44D5C497FF20CF42CF7B5793290" direction="south" orientation="latitude" precision="9" value="-12.118834">12°07.13'S</geoCoordinate>
,
<geoCoordinate id="488AC5087E15E0CB4AA5293BEB48F7FD" direction="east" orientation="longitude" precision="9" value="34.76483">34°45.89'E</geoCoordinate>
).
</paragraph>
</subSubSection>
<subSubSection id="468FE8A3C4F7386EA1B74BC06507F016" lastPageId="6" lastPageNumber="159" pageId="5" pageNumber="158" type="ecology">
<paragraph id="DA3B41387C69DBAFC2E2D453643F1157" pageId="5" pageNumber="158">Ecology.</paragraph>
<paragraph id="11D6EF1816DE2787B0802B1DC5356DB6" lastPageId="6" lastPageNumber="159" pageId="5" pageNumber="158">
Shell fragments of snails were observed in the stomachs of some specimens on X-rays. Stomach analyses were not performed so as to not damage the type
<pageBreakToken id="E3BD4CFD93A1E16514363A15A235B871" pageId="6" pageNumber="159" start="start">series</pageBreakToken>
. Thick and fleshy lips are associated with rocky habitats in cichlids, since fish feeding between rocks use their lips as a seal to be able to suck out their prey from crevices and as a protection against sharp rocks (
<bibRefCitation id="590DBC8842C2E1B5493A35B985142D52" author="Machado-Schiaffino, G" journalOrPublisher="Evolution" pageId="9" pageNumber="162" pagination="2086 - 2091" title="Species-specific differences in adaptive phenotypic plasticity in an ecologically relevant trophic trait: Hyperthrophic lips in Midas cichlid fishes." url="https://doi.org/10.1111/evo.12367" volume="68" year="2014">Machado-Schiaffino et al. 2014</bibRefCitation>
). Since all specimens were found in bottom trawls over soft substrates, we are unsure about the association of the new species with rocky habitats.
</paragraph>
</subSubSection>
</treatment>
</document>