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<mods:title id="A356BEDF09D0A8A91F14F5FBEDEB3666">Spionidae (Annelida: Polychaeta: Canalipalpata) from Lizard Island, Great Barrier Reef, Australia: the genera Malacoceros, Scolelepis, Spio, Microspio, and Spiophanes</mods:title>
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<emphasis id="2DC168058931FFAADFCD95C9FE3263E9" bold="true" box="[151,401,943,969]" italics="true" pageId="5" pageNumber="383">Malacoceros indicus</emphasis>
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<paragraph id="1F0AB4178931FFAADFCD95B4FF4863CB" blockId="5.[151,585,943,1003]" box="[151,235,978,1003]" pageId="5" pageNumber="383">(Fig. 1)</paragraph>
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<emphasis id="2DC168058931FFAADFCD927CFEE56410" box="[151,326,1050,1072]" italics="true" pageId="5" pageNumber="383">Scolelepis indica</emphasis>
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, figs
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m.—
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: 35
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, figs
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m;
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: 313
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314, figs
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m;
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: 25
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;
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: 21
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;
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: 277
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.
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<emphasis id="2DC168058931FFAADFCD923EFECF644E" box="[151,364,1112,1134]" italics="true" pageId="5" pageNumber="383">Malacoceros indicus</emphasis>
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.—
<treatmentCitation id="9E1492068931FFAADECE923EFDF8644E" author="Pettibone" box="[404,603,1112,1134]" page="99" pageId="5" pageNumber="383" year="1963">
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: 99
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;
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<bibRefCitation id="7B24C9E68931FFAADD32923FFD72644E" author="Day" box="[616,721,1112,1134]" pageId="5" pageNumber="383" refString="Day, J. H. (1967) A monograph on the polychaeta of southern Africa. Part 2. Sedentaria. Trustees of the British Museum (Natural History), London, 383 pp. [pp. 459 - 841]" type="book chapter" year="1967">Day 1967</bibRefCitation>
: 477
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, figs 18.5.pu;
<treatmentCitation id="9E1492068931FFAADCF0923EFB78644E" author="Blake" box="[938,1243,1112,1134]" page="195" pageId="5" pageNumber="383" year="1978">
<bibRefCitation id="7B24C9E68931FFAADCF0923EFB05644E" author="Blake" box="[938,1190,1112,1134]" pageId="5" pageNumber="383" refString="Blake, J. A. &amp; Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280." type="journal article" year="1978">Blake &amp; Kudenov 1978</bibRefCitation>
: 195
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;
<treatmentCitation id="9E1492068931FFAADBB2923EFA3B644E" author="Blake" box="[1256,1432,1112,1134]" page="219" pageId="5" pageNumber="383" year="1983">
<bibRefCitation id="7B24C9E68931FFAADBB2923EFAC1644E" author="Blake" box="[1256,1378,1112,1134]" pageId="5" pageNumber="383" refString="Blake, J. A. (1983) Polychaetes of the family Spionidae from South America, Antarctica, and adjacent seas and islands. Biology of the Antarctic Seas XIV, 39, 205 - 288. http: // dx. doi. org / 10.1029 / AR 039 p 0205" type="journal article" year="1983">Blake 1983</bibRefCitation>
: 219
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;
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<bibRefCitation id="7B24C9E68931FFAADF9D9210FE9D64AC" author="Blake" box="[199,318,1142,1164]" pageId="5" pageNumber="383" refString="Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. &amp; Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, pp. 81 - 223." type="book chapter" year="1996">Blake 1996</bibRefCitation>
: 105
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107, fig. 4.4;
<treatmentCitation id="9E1492068931FFAADD599210FCB364AC" author="Ben-Eliahu" box="[515,784,1142,1165]" page="96" pageId="5" pageNumber="383" year="1984">
<bibRefCitation id="7B24C9E68931FFAADD599210FD4964AC" author="Ben-Eliahu" box="[515,746,1142,1165]" pageId="5" pageNumber="383" refString="Ben-Eliahu, M. N., Hutchings, P. A. &amp; Glasby, C. J. (1984) Ceratonereis lizardensis n. sp. (Polychaeta; Nereididae) and Malacoceros indicus (Spionidae) from a mangrove habitat at Lizard Island, North Queensland. Proceedings of the first international polychaete conference, Sydney, Linnean Society of New South Wales, 1984, 91 - 97." type="proceedings paper" year="1984">
Ben-Eliahu
<emphasis id="2DC168058931FFAADD26921EFD0C64AC" box="[636,687,1142,1165]" italics="true" pageId="5" pageNumber="383">et al.</emphasis>
1984
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: 96
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;
<treatmentCitation id="9E1492068931FFAADC419211FB8464AC" author="Dauer" box="[795,1063,1142,1164]" page="395" pageId="5" pageNumber="383" year="1991">
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: 395
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400, fig. 1;
<treatmentCitation id="9E1492068931FFAADBF39210FAF364AC" author="Imajima" box="[1193,1360,1142,1164]" page="6" pageId="5" pageNumber="383" year="1991">
<bibRefCitation id="7B24C9E68931FFAADBF39210FA9A64AC" author="Imajima" box="[1193,1337,1142,1164]" pageId="5" pageNumber="383" refString="Imajima, M. (1991) Spionidae (Annelida, Polychaeta) from Japan. VI. The genera Malacoceros and Rhynchospio. Bulletin of the National Science Museum, Tokyo, Series A, 17, 5 - 17." type="journal article" year="1991">Imajima 1991</bibRefCitation>
: 6
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9, figs 2ag, 3aj.
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<taxonomicName id="D8B5CF948931FFAADFCD92D2FDA764EA" ID-CoL="7353S" box="[151,516,1204,1226]" class="Polychaeta" family="Spionidae" genus="Malacoceros" kingdom="Animalia" order="Spionida" pageId="5" pageNumber="383" phylum="Annelida" rank="species" species="indicus" subGenus="Malacoceros">
<emphasis id="2DC168058931FFAADFCD92D2FEB964EA" box="[151,282,1204,1226]" italics="true" pageId="5" pageNumber="383">Malacoceros</emphasis>
(
<emphasis id="2DC168058931FFAADE7392D2FE0F64EA" box="[297,428,1204,1226]" italics="true" pageId="5" pageNumber="383">Malacoceros</emphasis>
)
<emphasis id="2DC168058931FFAADEE192D2FDA764EA" box="[443,516,1204,1226]" italics="true" pageId="5" pageNumber="383">indicus</emphasis>
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.—
<treatmentCitation id="9E1492068931FFAADD7092D3FD7564EA" author="Foster" box="[554,726,1204,1226]" page="50" pageId="5" pageNumber="383" year="1971">
<bibRefCitation id="7B24C9E68931FFAADD7092D3FD1264EA" author="Foster" box="[554,689,1204,1226]" pageId="5" pageNumber="383" refString="Foster, N. M. (1971 a) Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea. Studies on the fauna of Curacao and other Caribbean Islands, 36, 1 - 150." type="journal article" year="1971" yearSuffix="a">Foster 1971a</bibRefCitation>
: 50
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53, figs 9399;
<treatmentCitation id="9E1492068931FFAADC2592D3FBE664EA" author="Foster" box="[895,1093,1204,1226]" page="1455" pageId="5" pageNumber="383" year="1971">
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:
<date id="6B0B92D78931FFAADB4892D2FBE664EA" box="[1042,1093,1204,1226]" pageId="5" pageNumber="383" value="1455" valueMax="1457">1455</date>
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1457, figs 16.
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<taxonomicName id="D8B5CF948931FFAADFCD92B5FE8164C8" ID-CoL="4Z67R" box="[151,290,1235,1256]" class="Polychaeta" family="Spionidae" genus="Spio" kingdom="Animalia" order="Spionida" pageId="5" pageNumber="383" phylum="Annelida" rank="species" species="punctata">
<emphasis id="2DC168058931FFAADFCD92B5FE8164C8" box="[151,290,1235,1256]" italics="true" pageId="5" pageNumber="383">Spio punctata</emphasis>
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.—
<treatmentCitation id="9E1492068931FFAADE1392B5FDA264C8" author="Hartman" box="[329,513,1234,1256]" page="89" pageId="5" pageNumber="383" year="1961">
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90, plate 11, figs13;
<treatmentCitation id="9E1492068931FFAADDB392B5FC1564C8" author="Hartman" box="[745,950,1234,1256]" page="175" pageId="5" pageNumber="383" year="1969">
<bibRefCitation id="7B24C9E68931FFAADDB392B5FC2664C8" author="Hartman" box="[745,901,1234,1256]" pageId="5" pageNumber="383" refString="Hartman, O. (1969) Atlas of the sedentariate polychaetous annelids from California. Allan Hancock Foundation, University of Southern California, Los Angeles, 812 pp." type="book" year="1969">Hartman, 1969</bibRefCitation>
: 175
</treatmentCitation>
, figs 13;
<emphasis id="2DC168058931FFAADB7B92B4FBE464C8" box="[1057,1095,1234,1256]" italics="true" pageId="5" pageNumber="383">fide</emphasis>
<bibRefCitation id="7B24C9E68931FFAADB1792B4FB6864C8" author="Blake" box="[1101,1227,1234,1256]" pageId="5" pageNumber="383" refString="Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. &amp; Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, pp. 81 - 223." type="book chapter" year="1996">Blake, 1996</bibRefCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph id="1F0AB4178931FFAADFCD9297FE7E6527" blockId="5.[151,1436,1050,1287]" box="[151,477,1265,1287]" pageId="5" pageNumber="383">
[Synonymy
<emphasis id="2DC168058931FFAADE499297FE9A6527" box="[275,313,1265,1287]" italics="true" pageId="5" pageNumber="383">fide</emphasis>
<bibRefCitation id="7B24C9E68931FFAADE1A9297FE746527" author="Williams" box="[320,471,1265,1287]" pageId="5" pageNumber="383" refString="Williams, J. D. (2007) New records and description of four new species of spionids (Annelida: Polychaeta: Spionidae) from the Philippines: the genera Dispio, Malacoceros, Polydora, and Scolelepis, with notes on palp ciliation patterns of the genus Scolelepis. Zootaxa, 1459, 1 - 35." type="journal article" year="2007">Williams 2007</bibRefCitation>
]
</paragraph>
</subSubSection>
<subSubSection id="57AFE79C8931FFAADFCD9352FECA6620" pageId="5" pageNumber="383" type="materials_examined">
<paragraph id="1F0AB4178931FFAADFCD9352FECA6620" blockId="5.[151,1437,1332,2005]" pageId="5" pageNumber="383">
<emphasis id="2DC168058931FFAADFCD9352FE21656D" bold="true" box="[151,386,1332,1357]" pageId="5" pageNumber="383">Material examined.</emphasis>
AM W.44376, MI QLD 2376 (2 af), formalin; AM W.44384, MI QLD 2376, af, formalin; AM W.44377, MI QLD 2376, af, formalin; AM W.44378, MI QLD 2376, af, mf, palps, 96% ethanol; AM W.44565, MI QLD 2422, af, formalin; AM W.44835, MI QLD 2422, af, formalin; AM W.44834, MI QLD 2422, af, formalin; AM W.44838, MI QLD 2422, af, formalin; AM W.44834.001, MI QLD 2422, mf, no longer extant; AM W.44858, MI QLD 2439, af, formalin; AM W.44858.001, MI QLD 2439, mf, ethanol; AM W.44859, MI QLD 2438, af, formalin.
</paragraph>
</subSubSection>
<subSubSection id="57AFE79C8931FFAADF9D906AFD2366DD" pageId="5" pageNumber="383" type="diagnosis">
<paragraph id="1F0AB4178931FFAADF9D906AFD2366DD" blockId="5.[151,1437,1332,2005]" pageId="5" pageNumber="383">
<emphasis id="2DC168058931FFAADF9D906AFEE26605" bold="true" box="[199,321,1548,1573]" pageId="5" pageNumber="383">Diagnosis.</emphasis>
Prostomium with distinct anterolateral horns or with anterolateral projections; prostomium posteriorly extended into short caruncle which terminates at the end of chaetiger 1. Branchiae present from chaetiger 1 for most of body length; cirriform with elongate, slender tip, separate from notopodial lamellae; first pair of branchiae usually about same length or longer than notopodial lamellae; branchiae often overlapping at midline in anterior mid-body region. Neuropodial postchaetal lamellae rounded, with small medial nipple-like projection in chaetigers of middle body region. Neuropodial bi-, tri-, or quadridentate hooded hooks present from chaetigers 28115, 311 hooks per fascicle.
</paragraph>
</subSubSection>
<subSubSection id="57AFE79C8931FFA8DF9D9161FEF963A0" lastPageId="7" lastPageNumber="385" pageId="5" pageNumber="383" type="description">
<paragraph id="1F0AB4178931FFA9DF9D9161FC956160" blockId="5.[151,1437,1332,2005]" lastBlockId="6.[151,1437,151,320]" lastPageId="6" lastPageNumber="384" pageId="5" pageNumber="383">
<emphasis id="2DC168058931FFAADF9D9161FEFB6700" bold="true" box="[199,344,1799,1824]" pageId="5" pageNumber="383">Description.</emphasis>
(based on specimens examined in the course of the present study). All specimens incomplete; longest specimen 154 chaetigers,
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wide and
<quantity id="D84D19F28931FFAADDBC914AFC946764" box="[742,823,1836,1860]" metricMagnitude="-2" metricUnit="m" metricValue="4.1" pageId="5" pageNumber="383" unit="mm" value="41.0">41 mm</quantity>
long; other specimens
<quantity id="D84D19F28931FFAADB1B914AFB696764" box="[1089,1226,1836,1860]" metricMagnitude="-3" metricUnit="m" metricValue="2.3" metricValueMax="3.8" metricValueMin="0.8" pageId="5" pageNumber="383" unit="mm" value="2.3" valueMax="3.8" valueMin="0.8">0.83.8 mm</quantity>
wide and
<quantity id="D84D19F28931FFAADA18914AFF606747" metricMagnitude="-2" metricUnit="m" metricValue="1.91" metricValueMax="2.7" metricValueMin="1.1199999999999999" pageId="5" pageNumber="383" unit="mm" value="19.1" valueMax="27.0" valueMin="11.2">11.227 mm</quantity>
long. Prostomium bell-shaped to sub-triangular, larger specimens with distinct anterolateral horns (Fig. 1A, B), smaller specimens with anterolateral projections; prostomium posteriorly extended into short protuberant caruncle which terminates at the end of chaetiger 1 (Fig. 1B). Prostomium usually with two groups of black eyes positioned on posterior half at maximal width of prostomium is reached, up to six eyes in one group, arranged in irregular patches or vertical rows (Fig. 1B). Occipital antenna absent. Peristomium not well developed. Palps detached, but scars of palp insertion discernable laterally to beginning of caruncle anterior to first chaetiger. Nuchal organs not unambiguously discernable, probably as small elongate grooves laterally to the posterior tip of the prostomium. Metameric dorsal ciliated organs not discernable. Dorsal crests absent but transverse ciliary bands across dorsum in well-preserved specimens present until posterior chaetigers.
</paragraph>
<paragraph id="1F0AB4178932FFA9DFCD93F3FC306608" blockId="6.[151,1436,1429,1576]" pageId="6" pageNumber="384">
<emphasis id="2DC168058932FFA9DFCD93F3FEB0658A" bold="true" box="[151,275,1429,1451]" pageId="6" pageNumber="384">FIGURE 1.</emphasis>
<taxonomicName id="D8B5CF948932FFA9DE4393F3FD29658B" authority="Fauvel, 1928" authorityName="Fauvel" authorityYear="1928" box="[281,650,1429,1451]" class="Polychaeta" family="Spionidae" genus="Malacoceros" kingdom="Animalia" order="Spionida" pageId="6" pageNumber="384" phylum="Annelida" rank="species" species="indicus">
<emphasis id="2DC168058932FFA9DE4393F3FE4F658B" box="[281,492,1429,1451]" italics="true" pageId="6" pageNumber="384">Malacoceros indicus</emphasis>
(
<bibRefCitation id="7B24C9E68932FFA9DEA093F3FD27658B" author="Fauvel" box="[506,644,1429,1451]" pageId="6" pageNumber="384" refString="Fauvel, P. (1928) Annelides Polychetes nouvelles de l'Inde. Bulletin du Museum National D'Histoire Naturelle, 34, 90 - 96." type="journal article" year="1928">Fauvel, 1928</bibRefCitation>
)
</taxonomicName>
, MI QLD 2429 (A, B); AM W.44838, MI QLD 2422 (C). A. Anterior end of live specimen, palps removed; B. Enlargement of detail from A, prostomium and anteriormost chaetigers, note protuberant caruncle and anterolateral horns; C. Left parapodium from 23rd chaetiger; note rounded shape of neuropodial postchaetal lamella with small (very indistinct) pointed tip and interramal bulge indicated by arrow. Scale bars: A, B no scale available, specimens are at least
<quantity id="D84D19F28932FFA9DE239074FE106608" box="[377,435,1554,1576]" metricMagnitude="-3" metricUnit="m" metricValue="3.0" pageId="6" pageNumber="384" unit="mm" value="3.0">3 mm</quantity>
wide, C = 100 µm. Photo: A, B—A. Semenov.
</paragraph>
<paragraph id="1F0AB4178932FFA9DF9D900DFDEB6718" blockId="6.[151,1437,1643,2028]" pageId="6" pageNumber="384">Dorsal branchiae present from chaetiger 1 until the end of fragment; cirriform with elongate, slender tip, separate from notopodial lamellae throughout (Fig. 1AC); first branchiae either slightly shorter, about same length or even longer than notopodial lamellae, from about third chaetiger reaching full length and often reaching dorsal midline or branchiae from both sides overlapping at midline (Fig. 1A); after first third of fragment (from about chaetiger 50) branchiae becoming thinner and shorter, but still distinctly longer than notopodial lamellae. Interparapodial lateral pouches absent.</paragraph>
<paragraph id="1F0AB4178932FFA8DF9D9125FE1260F5" blockId="6.[151,1437,1643,2028]" lastBlockId="7.[151,1437,151,2015]" lastPageId="7" lastPageNumber="385" pageId="6" pageNumber="384">Parapodia on chaetiger 1 positioned slightly more dorsally than on following chaetigers, there in lateral position. Notopodial postchaetal lamellae lanceolate, slender, becoming shorter posteriorly; neuropodial postchaetal lamellae of first and also second chaetiger elongate and tapered, then becoming rounded (Fig. 1C), often with small pointed tip on outer margin, but nipple-like projection rarely accentuated; more posteriorly neuropodial lamellae lower. Both parapodial rami without prominent prechaetal lamellae prechaetal thickened swelling (Fig. 1C) developing from chaetiger 3, present in near interramal position until about chaetigers 3040, thereafter inconspicuous.</paragraph>
<paragraph id="1F0AB4178933FFA8DF9D96B9FCB962CF" blockId="7.[151,1437,151,2015]" pageId="7" pageNumber="385">
Chaetae in anterior and middle chaetigers capillaries; in neuropodia with very fine granulations near tip and with narrow sheaths, arranged in two rows, chaetae in anterior row shorter than in posterior row; in notopodia chaetae alimbate and rather smooth, arranged in two rows, chaetae in anterior row shorter than in posterior row; few very long smooth capillaries present in uppermost position in notopodia. From middle body region capillaries and hooks present; in specimens up to
<quantity id="D84D19F28933FFA8DD0A9716FD0B61A7" box="[592,680,368,392]" metricMagnitude="-3" metricUnit="m" metricValue="1.4" pageId="7" pageNumber="385" unit="mm" value="1.4">1.4 mm</quantity>
wide neuropodial hooks in neuropodia present from chaetiger 31 39, in two very large specimens (anterior fragment
<quantity id="D84D19F28933FFA8DDB597F2FCEB618C" box="[751,840,404,428]" metricMagnitude="-3" metricUnit="m" metricValue="2.2" pageId="7" pageNumber="385" unit="mm" value="2.2">2.2 mm</quantity>
wide with 154 chaetigers or
<quantity id="D84D19F28933FFA8DBFA97F2FB59618C" box="[1184,1274,404,428]" metricMagnitude="-3" metricUnit="m" metricValue="3.8" pageId="7" pageNumber="385" unit="mm" value="3.8">3.8 mm</quantity>
wide with 83 chaetigers) hooks either present from chaetiger 121 or absent; hooks hooded with three apical teeth above main fang, consisting of one pair of teeth and additional minute apical tooth (difficult to observe due to size); 36 hooks present per fascicle; hooks accompanied by 13 thin smooth capillaries in superior position; notopodia with smooth capillaries without sheaths, first arranged in two irregular rows, more posteriorly bundle of capillaries of different length; very long capillaries present in notopodia in superiormost position. Capillaries in inferiormost position in neuropodia from chaetiger 2, numbering up to five (in very large specimen up to 15), chaetae arranged in a bundle; these capillaries slightly stronger and shorter than other capillaries in neuropodium; hook-bearing chaetigers with few (up to five) slightly granulated sabre chaetae with narrow sheath in inferiormost position; in more posterior chaetigers only 13 sabre chaetae present.
</paragraph>
<paragraph id="1F0AB4178933FFA8DF9D949AFCAB6335" blockId="7.[151,1437,151,2015]" box="[199,776,764,789]" pageId="7" pageNumber="385">Pygidium not observed (all specimens incomplete).</paragraph>
<paragraph id="1F0AB4178933FFA8DF9D9579FBCD6318" blockId="7.[151,1437,151,2015]" box="[199,1134,799,824]" pageId="7" pageNumber="385">
<emphasis id="2DC168058933FFA8DF9D9579FECD6318" bold="true" box="[199,366,799,824]" pageId="7" pageNumber="385">Pigmentation.</emphasis>
Pigment neither discernable in live or formalin preserved specimen.
</paragraph>
<paragraph id="1F0AB4178933FFA8DF9D9522FEF963A0" blockId="7.[151,1437,151,2015]" pageId="7" pageNumber="385">
<emphasis id="2DC168058933FFA8DF9D9522FDEE637D" bold="true" box="[199,589,836,861]" pageId="7" pageNumber="385">Methyl green staining pattern.</emphasis>
Inconspicuous. Prostomium, branchiae and postchaetal lamellae most intensely stained.
</paragraph>
</subSubSection>
<subSubSection id="57AFE79C8933FFA8DF9D95EAFD5E67FF" pageId="7" pageNumber="385" type="discussion">
<paragraph id="1F0AB4178933FFA8DF9D95EAFDFE6513" blockId="7.[151,1437,151,2015]" pageId="7" pageNumber="385">
<emphasis id="2DC168058933FFA8DF9D95EAFE986385" bold="true" box="[199,315,908,933]" pageId="7" pageNumber="385">Remarks.</emphasis>
<taxonomicName id="D8B5CF948933FFA8DE1D95EAFD956385" box="[327,566,908,933]" class="Polychaeta" family="Spionidae" genus="Malacoceros" kingdom="Animalia" order="Spionida" pageId="7" pageNumber="385" phylum="Annelida" rank="species" species="indicus">
<emphasis id="2DC168058933FFA8DE1D95EAFD956385" box="[327,566,908,933]" italics="true" pageId="7" pageNumber="385">Malacoceros indicus</emphasis>
</taxonomicName>
is, based on current knowledge, a very widespread species. Intraspecific variability is acknowledged to be large. For example, the number of secondary teeth in neuropodial hooks are observed to range from two (
<bibRefCitation id="7B24C9E68933FFA8DEB995B2FDD963CD" author="Blake" box="[483,634,980,1005]" pageId="7" pageNumber="385" refString="Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. &amp; Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, pp. 81 - 223." type="book chapter" year="1996">Blake (1996)</bibRefCitation>
for specimens from California), three (
<bibRefCitation id="7B24C9E68933FFA8DB6195B2FB5663CC" author="Williams" box="[1083,1269,980,1005]" pageId="7" pageNumber="385" refString="Williams, J. D. (2007) New records and description of four new species of spionids (Annelida: Polychaeta: Spionidae) from the Philippines: the genera Dispio, Malacoceros, Polydora, and Scolelepis, with notes on palp ciliation patterns of the genus Scolelepis. Zootaxa, 1459, 1 - 35." type="journal article" year="2007">Williams (2007)</bibRefCitation>
for specimens from the
<collectingCountry id="67A2F4878933FFA8DE599591FE276430" box="[259,388,1015,1040]" name="Philippines" pageId="7" pageNumber="385">Philippines</collectingCountry>
, also this paper for Lizard Island specimens), to four (
<bibRefCitation id="7B24C9E68933FFA8DCA49591FB116430" author="Imajima" box="[1022,1202,1015,1040]" pageId="7" pageNumber="385" refString="Imajima, M. (1991) Spionidae (Annelida, Polychaeta) from Japan. VI. The genera Malacoceros and Rhynchospio. Bulletin of the National Science Museum, Tokyo, Series A, 17, 5 - 17." type="journal article" year="1991">Imajima (1991)</bibRefCitation>
for specimens from
<collectingCountry id="67A2F4878933FFA8DFCD927AFF7F6414" box="[151,220,1052,1076]" name="Japan" pageId="7" pageNumber="385">Japan</collectingCountry>
).
<bibRefCitation id="7B24C9E68933FFA8DFAF927BFE016415" author="Foster" box="[245,418,1052,1077]" pageId="7" pageNumber="385" refString="Foster, N. M. (1971 b) Redescription of the spionid polychaete Malacoceros (Malacoceros) indicus (Fauvel 1928). Journal of the Fisheries Research Board of Canada, 28, 1455 - 1457. http: // dx. doi. org / 10.1139 / f 71 - 224" type="journal article" year="1971" yearSuffix="b">Foster (1971b)</bibRefCitation>
found tri- and quadridentate hooks in the same parapodium in specimens from the Carribean. Also the number of hooks per fascicle varies: our specimens have 36 hooks per fascicle,
<bibRefCitation id="7B24C9E68933FFA8DA6D9259FF42645C" author="Williams" pageId="7" pageNumber="385" refString="Williams, J. D. (2007) New records and description of four new species of spionids (Annelida: Polychaeta: Spionidae) from the Philippines: the genera Dispio, Malacoceros, Polydora, and Scolelepis, with notes on palp ciliation patterns of the genus Scolelepis. Zootaxa, 1459, 1 - 35." type="journal article" year="2007">Williams (2007)</bibRefCitation>
reports up to 7 hooks,
<bibRefCitation id="7B24C9E68933FFA8DEAB9203FD3B645D" author="Foster" box="[497,664,1124,1149]" pageId="7" pageNumber="385" refString="Foster, N. M. (1971 b) Redescription of the spionid polychaete Malacoceros (Malacoceros) indicus (Fauvel 1928). Journal of the Fisheries Research Board of Canada, 28, 1455 - 1457. http: // dx. doi. org / 10.1139 / f 71 - 224" type="journal article" year="1971" yearSuffix="b">Foster (1971b)</bibRefCitation>
78 hooks,
<bibRefCitation id="7B24C9E68933FFA8DC739202FC1D645D" author="Blake" box="[809,958,1124,1149]" pageId="7" pageNumber="385" refString="Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. &amp; Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, pp. 81 - 223." type="book chapter" year="1996">Blake (1996)</bibRefCitation>
710,
<bibRefCitation id="7B24C9E68933FFA8DB4A9202FB62645C" author="Imajima" box="[1040,1217,1124,1149]" pageId="7" pageNumber="385" refString="Imajima, M. (1991) Spionidae (Annelida, Polychaeta) from Japan. VI. The genera Malacoceros and Rhynchospio. Bulletin of the National Science Museum, Tokyo, Series A, 17, 5 - 17." type="journal article" year="1991">Imajima (1991)</bibRefCitation>
up to 11. The first presence of neuropodial hooks was observed in chaetigers 31121 (Lizard Island, present paper), chaetigers 2862 (
<bibRefCitation id="7B24C9E68933FFA8DFC492C8FEED64E7" author="Williams" box="[158,334,1198,1223]" pageId="7" pageNumber="385" refString="Williams, J. D. (2007) New records and description of four new species of spionids (Annelida: Polychaeta: Spionidae) from the Philippines: the genera Dispio, Malacoceros, Polydora, and Scolelepis, with notes on palp ciliation patterns of the genus Scolelepis. Zootaxa, 1459, 1 - 35." type="journal article" year="2007">Williams 2007</bibRefCitation>
), 3057 (
<bibRefCitation id="7B24C9E68933FFA8DE9B92C8FDCB64E7" author="Imajima" box="[449,616,1198,1223]" pageId="7" pageNumber="385" refString="Imajima, M. (1991) Spionidae (Annelida, Polychaeta) from Japan. VI. The genera Malacoceros and Rhynchospio. Bulletin of the National Science Museum, Tokyo, Series A, 17, 5 - 17." type="journal article" year="1991">Imajima 1991</bibRefCitation>
), 3035 (
<bibRefCitation id="7B24C9E68933FFA8DD8692C8FCC464E7" author="Blake" box="[732,871,1198,1223]" pageId="7" pageNumber="385" refString="Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. &amp; Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, pp. 81 - 223." type="book chapter" year="1996">Blake 1996</bibRefCitation>
), and at the 37th chaetiger (
<bibRefCitation id="7B24C9E68933FFA8DBE992C9FAF264E7" author="Foster" box="[1203,1361,1198,1223]" pageId="7" pageNumber="385" refString="Foster, N. M. (1971 b) Redescription of the spionid polychaete Malacoceros (Malacoceros) indicus (Fauvel 1928). Journal of the Fisheries Research Board of Canada, 28, 1455 - 1457. http: // dx. doi. org / 10.1139 / f 71 - 224" type="journal article" year="1971" yearSuffix="b">Foster 1971b</bibRefCitation>
). This character seems to be size-dependent in the specimens collected at Lizard Island since only the largest two individuals exhibit a very late start of neuropodial hooks (see description above). Hooks might be continously lost in middle chaetigers during ontogenesis.
</paragraph>
<paragraph id="1F0AB4178933FFA8DF9D9358FDB66673" blockId="7.[151,1437,151,2015]" pageId="7" pageNumber="385">
The prostomial shape varies with the development of anterolateral horns. Among specimens from Lizard Island only the two large individuals (2.2 and
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wide) have anterolateral horns whereas all other specimens (
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wide) have anterolateral projections. In the literature the species is usually illustrated having anterolateral projections and rarely short anterolateral horns (compare
<bibRefCitation id="7B24C9E68933FFA8DCE493CDFBF565E3" author="Foster" box="[958,1110,1450,1475]" pageId="7" pageNumber="385" refString="Foster, N. M. (1971 b) Redescription of the spionid polychaete Malacoceros (Malacoceros) indicus (Fauvel 1928). Journal of the Fisheries Research Board of Canada, 28, 1455 - 1457. http: // dx. doi. org / 10.1139 / f 71 - 224" type="journal article" year="1971" yearSuffix="b">Foster 1971b</bibRefCitation>
,
<bibRefCitation id="7B24C9E68933FFA8DB3E93CCFAA765E3" author="Imajima" box="[1124,1284,1450,1475]" pageId="7" pageNumber="385" refString="Imajima, M. (1991) Spionidae (Annelida, Polychaeta) from Japan. VI. The genera Malacoceros and Rhynchospio. Bulletin of the National Science Museum, Tokyo, Series A, 17, 5 - 17." type="journal article" year="1991">Imajima 1991</bibRefCitation>
,
<bibRefCitation id="7B24C9E68933FFA8DA4893CCFA3B65E3" author="Blake" box="[1298,1432,1450,1475]" pageId="7" pageNumber="385" refString="Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. &amp; Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, pp. 81 - 223." type="book chapter" year="1996">Blake 1996</bibRefCitation>
,
<bibRefCitation id="7B24C9E68933FFA8DFCD93A8FEE265C7" author="Williams" box="[151,321,1486,1511]" pageId="7" pageNumber="385" refString="Williams, J. D. (2007) New records and description of four new species of spionids (Annelida: Polychaeta: Spionidae) from the Philippines: the genera Dispio, Malacoceros, Polydora, and Scolelepis, with notes on palp ciliation patterns of the genus Scolelepis. Zootaxa, 1459, 1 - 35." type="journal article" year="2007">Williams 2007</bibRefCitation>
). We noticed that specimens from Lizard Island have a protuberant caruncle. This is obviously also the case in specimens from the
<collectingCountry id="67A2F4878933FFA8DD5B9394FDDC662B" box="[513,639,1522,1547]" name="Philippines" pageId="7" pageNumber="385">Philippines</collectingCountry>
(compare illustrations provided by
<bibRefCitation id="7B24C9E68933FFA8DB429394FB7D662B" author="Williams" box="[1048,1246,1522,1547]" pageId="7" pageNumber="385" refString="Williams, J. D. (2007) New records and description of four new species of spionids (Annelida: Polychaeta: Spionidae) from the Philippines: the genera Dispio, Malacoceros, Polydora, and Scolelepis, with notes on palp ciliation patterns of the genus Scolelepis. Zootaxa, 1459, 1 - 35." type="journal article" year="2007">Williams (2007))</bibRefCitation>
. All other above mentioned authors have provided illustrations displaying a differently shaped caruncle and did not mention such a development in their descriptions.
</paragraph>
<paragraph id="1F0AB4178933FFA8DF9D9038FC0966C2" blockId="7.[151,1437,151,2015]" pageId="7" pageNumber="385">
Information on three different molecular markers is provided for
<taxonomicName id="D8B5CF948933FFA8DCC69006FBB16657" box="[924,1042,1630,1655]" class="Polychaeta" family="Spionidae" genus="Malacoceros" kingdom="Animalia" order="Spionida" pageId="7" pageNumber="385" phylum="Annelida" rank="species" species="indicus">
<emphasis id="2DC168058933FFA8DCC69006FBB16657" box="[924,1042,1630,1655]" italics="true" pageId="7" pageNumber="385">M. indicus</emphasis>
</taxonomicName>
from Lizard Island in the course of the present study. Since data from other localities are not available from sources like GenBank or BOLD a comparative analysis could not be undertaken. The inclusion of both molecular and morphological data should be aspired too for future studies to verify the species status of
<taxonomicName id="D8B5CF948933FFA8DC7690AAFC0066C3" box="[812,931,1738,1763]" class="Polychaeta" family="Spionidae" genus="Malacoceros" kingdom="Animalia" order="Spionida" pageId="7" pageNumber="385" phylum="Annelida" rank="species" species="indicus">
<emphasis id="2DC168058933FFA8DC7690AAFC0066C3" box="[812,931,1738,1763]" italics="true" pageId="7" pageNumber="385">M. indicus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="1F0AB4178933FFA8DF9D9088FD5E67FF" blockId="7.[151,1437,151,2015]" pageId="7" pageNumber="385">
<emphasis id="2DC168058933FFA8DF9D9088FE3F6727" bold="true" box="[199,412,1774,1799]" pageId="7" pageNumber="385">Habitat / Ecology.</emphasis>
In the Lizard Island area the species was found in shallow water on sandy beaches, between seagrass and mangroves. The species was also found from sandy beaches, in the shallow subtidal (&lt;
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) in Boracay of the Aklan province in the
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(
<bibRefCitation id="7B24C9E68933FFA8DD8A9150FCDF676F" author="Williams" box="[720,892,1846,1871]" pageId="7" pageNumber="385" refString="Williams, J. D. (2007) New records and description of four new species of spionids (Annelida: Polychaeta: Spionidae) from the Philippines: the genera Dispio, Malacoceros, Polydora, and Scolelepis, with notes on palp ciliation patterns of the genus Scolelepis. Zootaxa, 1459, 1 - 35." type="journal article" year="2007">Williams 2007</bibRefCitation>
). Records from
<collectingCountry id="67A2F4878933FFA8DB6B915EFBD2676F" box="[1073,1137,1848,1871]" name="Japan" pageId="7" pageNumber="385">Japan</collectingCountry>
are from shallow water to
<quantity id="D84D19F28933FFA8DFCD913CFF7C6752" box="[151,223,1882,1907]" metricMagnitude="2" metricUnit="m" metricValue="1.59" pageId="7" pageNumber="385" unit="m" value="159.0">159 m</quantity>
water depth (
<bibRefCitation id="7B24C9E68933FFA8DE26913CFD836753" author="Imajima" box="[380,544,1882,1907]" pageId="7" pageNumber="385" refString="Imajima, M. (1991) Spionidae (Annelida, Polychaeta) from Japan. VI. The genera Malacoceros and Rhynchospio. Bulletin of the National Science Museum, Tokyo, Series A, 17, 5 - 17." type="journal article" year="1991">Imajima 1991</bibRefCitation>
).
<bibRefCitation id="7B24C9E68933FFA8DD6F913DFD796753" author="Foster" box="[565,730,1883,1907]" pageId="7" pageNumber="385" refString="Foster, N. M. (1971 a) Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea. Studies on the fauna of Curacao and other Caribbean Islands, 36, 1 - 150." type="journal article" year="1971" yearSuffix="a">Foster (1971a)</bibRefCitation>
reported the species from
<collectingCountry id="67A2F4878933FFA8DB56913CFB3D6753" box="[1036,1182,1882,1907]" name="Bahamas" pageId="7" pageNumber="385">the Bahamas</collectingCountry>
and
<collectingCountry id="67A2F4878933FFA8DB8D913DFAFE6753" box="[1239,1373,1882,1907]" name="Puerto Rico" pageId="7" pageNumber="385">Puerto Rico</collectingCountry>
from shallow water from sandy bottom and
<taxonomicName id="D8B5CF948933FFA8DD0C9118FD6067B7" box="[598,707,1918,1943]" class="Liliopsida" family="Hydrocharitaceae" genus="Thalassia" kingdom="Animalia" order="Alismatales" pageId="7" pageNumber="385" phylum="Tracheophyta" rank="genus">
<emphasis id="2DC168058933FFA8DD0C9118FD6067B7" box="[598,707,1918,1943]" italics="true" pageId="7" pageNumber="385">Thalassia</emphasis>
</taxonomicName>
beds. Records from
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(Golfo de Nicoya) account for subtidal habitats (
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water depth) with muddy sand or sand (
<bibRefCitation id="7B24C9E68933FFA8DCD591C5FBB2679B" author="Dean" box="[911,1041,1955,1979]" pageId="7" pageNumber="385" refString="Dean, D. (2004) Marine biodiversity of Costa Rica: Class Polychaeta (Annelida). Revista de Biologia Tropical, 52, 1 - 69." type="journal article" year="2004">Dean 2004</bibRefCitation>
).
<bibRefCitation id="7B24C9E68933FFA8DB7E91C5FA89679B" author="Dauer" box="[1060,1322,1954,1979]" pageId="7" pageNumber="385" refString="Dauer, D. M. &amp; Ewing, R. M. (1991) Functional morphology and feeding behavior of Malacoceros indicus (Polychaeta: Spionidae). Bulletin of Marine Science, 48, 395 - 400." type="journal article" year="1991">Dauer &amp; Ewing (1991)</bibRefCitation>
classified the species as an indiscriminate surface deposit-feeder.
</paragraph>
</subSubSection>
<subSubSection id="57AFE79C893CFFA7DF9D96F1FB3E60F5" pageId="8" pageNumber="386" type="distribution">
<paragraph id="1F0AB417893CFFA7DF9D96F1FB3E60F5" blockId="8.[151,1436,151,213]" pageId="8" pageNumber="386">
<emphasis id="2DC16805893CFFA7DF9D96F1FEC36090" bold="true" box="[199,352,151,176]" pageId="8" pageNumber="386">Distribution.</emphasis>
<collectingCountry id="67A2F487893CFFA7DE3C96F1FE6E6090" box="[358,461,151,176]" name="Australia" pageId="8" pageNumber="386">Australia</collectingCountry>
(Queensland, NSW),
<collectingCountry id="67A2F487893CFFA7DD9996FEFCD16090" box="[707,882,151,176]" name="New Caledonia" pageId="8" pageNumber="386">New Caledonia</collectingCountry>
,
<collectingCountry id="67A2F487893CFFA7DC2796F1FC5D6090" box="[893,1022,151,176]" name="Philippines" pageId="8" pageNumber="386">Philippines</collectingCountry>
,
<collectingCountry id="67A2F487893CFFA7DB5396FEFBEF608F" box="[1033,1100,152,175]" name="Japan" pageId="8" pageNumber="386">Japan</collectingCountry>
,
<collectingCountry id="67A2F487893CFFA7DB0C96F1FB316090" box="[1110,1170,151,176]" name="India" pageId="8" pageNumber="386">India</collectingCountry>
, SW Africa, Caribbean,
<collectingCountry id="67A2F487893CFFA7DFCD96DAFF7460F5" box="[151,215,188,213]" name="Chile" pageId="8" pageNumber="386">Chile</collectingCountry>
,
<collectingCountry id="67A2F487893CFFA7DFB896DAFEB960F4" box="[226,282,188,212]" name="United States of America" pageId="8" pageNumber="386">USA</collectingCountry>
(southern California, Massachusetts to
<collectingCountry id="67A2F487893CFFA7DD8C96DAFC9760F5" box="[726,820,188,213]" name="Georgia" pageId="8" pageNumber="386">Georgia</collectingCountry>
),
<collectingCountry id="67A2F487893CFFA7DC1D96DAFC6260F5" box="[839,961,188,213]" name="Costa Rica" pageId="8" pageNumber="386">Costa Rica</collectingCountry>
(Golfo de Nicoya).
</paragraph>
</subSubSection>
</treatment>
</document>