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<mods:title id="8159461F97410EB5B29A0F195A88E170">The genital area of Halacaridae (Acari), life stages and development of morphological characters and implication on the classification</mods:title>
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<subSubSection id="C37B6578FFE4FF9DFF12A637FECFFEE5" pageId="15" pageNumber="216" type="nomenclature">
<paragraph id="8BDE36F3FFE4FF9DFF12A637FDCEFEC4" blockId="15.[151,550,296,355]" box="[151,550,296,322]" pageId="15" pageNumber="216">
<heading id="D096819FFFE4FF9DFF12A637FDCEFEC4" bold="true" box="[151,550,296,322]" fontSize="11" level="1" pageId="15" pageNumber="216" reason="1">
<taxonomicName id="4C614D70FFE4FF9DFF12A637FDCEFEC4" ID-CoL="3TPJ" ID-ENA="1027280" authority="Trouessart, 1888" authorityName="Trouessart" authorityYear="1888" box="[151,550,296,322]" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="genus">
<emphasis id="B915EAE1FFE4FF9DFF12A637FDCEFEC4" bold="true" box="[151,550,296,322]" pageId="15" pageNumber="216">
<emphasis id="B915EAE1FFE4FF9DFF12A637FEA7FEC4" bold="true" box="[151,335,296,322]" italics="true" pageId="15" pageNumber="216">Copidognathus</emphasis>
<bibRefCitation id="EFF04B02FFE4FF9DFED3A637FDCEFEC4" author="Trouessart" box="[342,550,296,322]" pageId="15" pageNumber="216" refString="Trouessart, E. L. (1888) Note sur les acariens marins recueillis par M. Giard au laboratoire maritime de Wimereux. Comptes Rendus de l'Academie des Sciences, Paris, 107, 753 - 755." type="journal article" year="1888">Trouessart, 1888</bibRefCitation>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="8BDE36F3FFE4FF9DFF12A655FECFFEE5" blockId="15.[151,550,296,355]" box="[151,295,330,355]" pageId="15" pageNumber="216">(Figs 3740)</paragraph>
</subSubSection>
<subSubSection id="C37B6578FFE4FF9DFF12A68DFEFAFB19" pageId="15" pageNumber="216" type="materials_examined">
<paragraph id="8BDE36F3FFE4FF9DFF12A68DFCAAFE2D" blockId="15.[151,1437,402,1255]" box="[151,834,402,427]" pageId="15" pageNumber="216">
<emphasis id="B915EAE1FFE4FF9DFF12A68DFEDAFE2D" bold="true" box="[151,306,402,427]" pageId="15" pageNumber="216">
<typeStatus id="54DA8851FFE4FF9DFF12A68DFF3AFE2D" box="[151,210,402,427]" pageId="15" pageNumber="216">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="4C614D70FFE4FF9DFEBCA68DFCD6FE2D" authority="Trouessart, 1888" authorityName="Trouessart" authorityYear="1888" box="[313,830,402,427]" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="species" species="glyptoderma">
<emphasis id="B915EAE1FFE4FF9DFEBCA68DFD90FE2D" box="[313,632,402,427]" italics="true" pageId="15" pageNumber="216">Copidognathus glyptoderma</emphasis>
<bibRefCitation id="EFF04B02FFE4FF9DFDFAA68CFCD6FE2D" author="Trouessart" box="[639,830,403,427]" pageId="15" pageNumber="216" refString="Trouessart, E. L. (1888) Note sur les acariens marins recueillis par M. Giard au laboratoire maritime de Wimereux. Comptes Rendus de l'Academie des Sciences, Paris, 107, 753 - 755." type="journal article" year="1888">Trouessart, 1888</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="8BDE36F3FFE4FF9DFF42A6A9FB6FFB89" blockId="15.[151,1437,402,1255]" pageId="15" pageNumber="216">
<emphasis id="B915EAE1FFE4FF9DFF42A6A9FEF5FE49" bold="true" box="[199,285,438,463]" pageId="15" pageNumber="216">Adults.</emphasis>
In both sexes GP and AP fused. Female GO close to anal cone, in a few mounted (slide) specimens partly overlapping basis of anal sclerites. Female with single pair of internal acetabula in about middle of GO (Figs 37, 38;
<bibRefCitation id="EFF04B02FFE4FF9DFF70A6E1FE66FD91" author="Newell" box="[245,398,510,535]" pageId="15" pageNumber="216" refString="Newell, I. M. (1947) A systematic and ecological study of the Halacaridae of eastern North America. Bulletin of the Bingham Oceanographic Collection, 10, 1 - 232." type="journal article" year="1947">Newell 1947</bibRefCitation>
: figs 198, 224, 1971: fig. 24;
<bibRefCitation id="EFF04B02FFE4FF9DFD7FA6E1FC4CFD91" author="Bartsch" box="[762,932,510,535]" pageId="15" pageNumber="216" refString="Bartsch, I. (2003 a) Mangrove halacarid fauna (Halacaridae, Acari) of the Dampier region, Western Australia, with description of five new species. Journal of Natural History, 37, 1855 - 1877. http: // dx. doi. org / 10.1080 / 00222930110089184" type="journal article" year="2003" yearSuffix="a">Bartsch 2003a</bibRefCitation>
: fig. 8A, 2006a: fig. 22), in a few species acetabula somewhat or greatly enlarged, namely in
<taxonomicName id="4C614D70FFE4FF9DFD5CA53CFC14FDBD" authority="Bartsch 1986" authorityName="Bartsch" authorityYear="1986" box="[729,1020,546,571]" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="species" species="acnemus">
<emphasis id="B915EAE1FFE4FF9DFD5CA53CFCB4FDBD" box="[729,860,547,571]" italics="true" pageId="15" pageNumber="216">C. acnemus</emphasis>
Bartsch 1986
</taxonomicName>
,
<taxonomicName id="4C614D70FFE4FF9DFB82A53CFA9FFDBD" authority="Bartsch 1992" authorityName="Bartsch" authorityYear="1992" box="[1031,1399,546,571]" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="species" species="cephalocanthus">
<emphasis id="B915EAE1FFE4FF9DFB82A53CFB30FDBD" box="[1031,1240,546,571]" italics="true" pageId="15" pageNumber="216">C. cephalocanthus</emphasis>
Bartsch 1992
</taxonomicName>
,
<taxonomicName id="4C614D70FFE4FF9DFA07A53CFE57FDD9" authority="Bartsch 1993" authorityName="Bartsch" authorityYear="1993" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="species" species="laeviusculus">
<emphasis id="B915EAE1FFE4FF9DFA07A53CFECAFDD9" italics="true" pageId="15" pageNumber="216">C. laeviusculus</emphasis>
Bartsch 1993
</taxonomicName>
,
<taxonomicName id="4C614D70FFE4FF9DFE4FA558FCE8FDD9" authority="Bartsch, 1993" authorityName="Bartsch" authorityYear="1993" box="[458,768,582,607]" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="species" species="majorinus">
<emphasis id="B915EAE1FFE4FF9DFE4FA558FE35FDD8" box="[458,477,583,606]" italics="true" pageId="15" pageNumber="216">C</emphasis>
.
<emphasis id="B915EAE1FFE4FF9DFE6FA558FDB3FDD8" box="[490,603,583,606]" italics="true" pageId="15" pageNumber="216">majorinus</emphasis>
Bartsch, 1993
</taxonomicName>
,
<taxonomicName id="4C614D70FFE4FF9DFC8EA558FBE5FDD9" authority="Bartsch, 1984" authorityName="Bartsch" authorityYear="1984" box="[779,1037,582,607]" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="species" species="mirus">
<emphasis id="B915EAE1FFE4FF9DFC8EA558FC84FDD8" box="[779,876,583,606]" italics="true" pageId="15" pageNumber="216">C. mirus</emphasis>
Bartsch, 1984
</taxonomicName>
(
<bibRefCitation id="EFF04B02FFE4FF9DFB9EA559FB2AFDD9" author="Bartsch" box="[1051,1218,582,607]" pageId="15" pageNumber="216" refString="Bartsch, I. (1984 a) Three new psammobiont species of Copidognathus (Acari, Halacaridae) from the Philippines. The Philippine Journal of Science, 113, 201 - 214." type="journal article" year="1984" yearSuffix="a">Bartsch 1984a</bibRefCitation>
: fig. 27, 1986a: fig. 6, 1992b: fig. 26, 1993c: fig. 8B, 9G). GA with (2)3(5) pairs of pgs, genital sclerites with single pair of small sgs. Ovipositor at rest extending from slightly to far beyond anterior margin of GO. When at rest, zero to one pair of basal genital spines or papillae and two pairs of apical spines visible through genital sclerites (Figs 37, 38;
<bibRefCitation id="EFF04B02FFE4FF9DFAC2A5ADFF09FD69" author="Bartsch" pageId="15" pageNumber="216" refString="Bartsch, I. (2013 d) A new species, genus Copidognathus (Acari: Halacaridae), from a hydrothermal vent area of the East Pacific Rise. Cahiers de Biologie Marine, 54, 239 - 246." type="journal article" year="2013" yearSuffix="d">Bartsch 2013d</bibRefCitation>
: fig. 5). Everted ovipositor apically with four anterior (lateral) and seven posterior spines; their shape spiniform to claw-like and palmate (Figs 39, 40;
<bibRefCitation id="EFF04B02FFE4FF9DFD55A5E5FC90FC95" author="Bartsch" box="[720,888,762,787]" pageId="15" pageNumber="216" refString="Bartsch, I. (1977 a) Erganzung zur Halacariden-Fauna (Halacaridae, Acari) der Bretagne-Kuste. Zur Gattung Arhodeoporus und Copidognathus. Acarologia, 18, 626 - 641." type="journal article" year="1977" yearSuffix="a">Bartsch 1977a</bibRefCitation>
: figs 32, 48). Male GA with 440 pairs of pgs. Male GO similar-sized or somewhat smaller than female GO and in a more ventral position. Genital sclerites with (2)34 pairs of sgs; often one or two pairs of spur-like sgs in posterior half of sclerites, the other sgs setiform. Single pair of gac in posterior half of GO, similar-sized or slightly smaller than those of female (
<bibRefCitation id="EFF04B02FFE4FF9DFB50A479FA83FCF9" author="Newell" box="[1237,1387,870,895]" pageId="15" pageNumber="216" refString="Newell, I. M. (1971) Halacaridae (Acari) collected during cruise 17 of the R / V Anton Bruun, in the southeastern Pacific Ocean. Anton Bruun Report, 8, 1 - 58." type="journal article" year="1971">Newell 1971</bibRefCitation>
: fig. 55;
<bibRefCitation id="EFF04B02FFE4FF9DFF41A495FE86FC25" author="Bartsch" box="[196,366,906,931]" pageId="15" pageNumber="216" refString="Bartsch, I. (1977 b) Interstitielle Fauna von Galapagos. XX. Halacaridae (Acari). Mikrofauna des Meeresbodens, 65, 1 - 108." type="journal article" year="1977" yearSuffix="b">Bartsch 1977b</bibRefCitation>
: fig. 133, 2003a: fig. 7C, 2006a: fig. 34). AE of both female and male with epimeral pores. Species living in freshwater often with enlarged pores (if compared with that size in congeners from other habitats) (
<bibRefCitation id="EFF04B02FFE4FF9DFF1BA4CDFEAFFC6D" author="Bartsch" box="[158,327,978,1003]" pageId="15" pageNumber="216" refString="Bartsch, I. (1979 b) Halacaridae (Acari) von der Atlantikkuste Nordamerikas. Beschreibung der Arten. Mikrofauna des Meeresbodens, 79, 1 - 62." type="journal article" year="1979" yearSuffix="b">Bartsch 1979b</bibRefCitation>
: fig. 141, 1999d: fig. 3), a few inhabitants of marine sandy deposits with enlarged internal lumina, examples are
<taxonomicName id="4C614D70FFE4FF9DFEB4A4E9FD85FB89" box="[305,621,1014,1039]" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="species" species="laeviusculus">
<emphasis id="B915EAE1FFE4FF9DFEB4A4E9FD85FB89" box="[305,621,1014,1039]" italics="true" pageId="15" pageNumber="216">Copidognathus laeviusculus</emphasis>
</taxonomicName>
and C.
<emphasis id="B915EAE1FFE4FF9DFD41A4E8FCC3FB88" box="[708,811,1015,1038]" italics="true" pageId="15" pageNumber="216">majorinu</emphasis>
s (
<bibRefCitation id="EFF04B02FFE4FF9DFCC1A4E9FB92FB89" author="Bartsch" box="[836,1146,1014,1039]" pageId="15" pageNumber="216" refString="Bartsch, I. (1993 c) Arenicolous Halacaridae (Acari) from south-western Australia. In: Wells, F. E., Walker, D. I., Kirkman, H. &amp; Lethbridge, R. (Eds.), The Marine Flora and Fauna of Rottnest Island, Western Australia. Western Australian Museum, Perth, pp. 73 - 103." type="book chapter" year="1993" yearSuffix="c">Bartsch 1993c: figs 8B, 9B</bibRefCitation>
).
</paragraph>
<paragraph id="8BDE36F3FFE4FF9DFF42A305FEFAFB19" blockId="15.[151,1437,402,1255]" pageId="15" pageNumber="216">
<emphasis id="B915EAE1FFE4FF9DFF42A305FED4FBB5" bold="true" box="[199,316,1050,1075]" pageId="15" pageNumber="216">Juveniles.</emphasis>
A larval and a single nymphal stage (protonymph) known. In both stages AE with pair of epimeral pores. In protonymph GP and AP separated, an exception, the fused plates in
<taxonomicName id="4C614D70FFE4FF9DFB91A320FADCFBD1" authority="Bartsch, 1984" authorityName="Bartsch" authorityYear="1984" box="[1044,1332,1086,1111]" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="species" species="bunofer">
<emphasis id="B915EAE1FFE4FF9DFB91A320FB65FBD1" box="[1044,1165,1086,1111]" italics="true" pageId="15" pageNumber="216">C. bunofer</emphasis>
Bartsch, 1984
</taxonomicName>
(
<bibRefCitation id="EFF04B02FFE4FF9DFAC1A321FED8FBFD" author="Bartsch" pageId="15" pageNumber="216" refString="Bartsch, I. (1984 a) Three new psammobiont species of Copidognathus (Acari, Halacaridae) from the Philippines. The Philippine Journal of Science, 113, 201 - 214." type="journal article" year="1984" yearSuffix="a">Bartsch 1984a: fig. 8</bibRefCitation>
); this may be an anomaly. GP of protonymph with single pair of internal gac; pgs and sgs not developed.
</paragraph>
</subSubSection>
<subSubSection id="C37B6578FFE4FF9DFF42A3B5FC29FB61" pageId="15" pageNumber="216" type="discussion">
<paragraph id="8BDE36F3FFE4FF9DFF42A3B5FC29FB61" blockId="15.[151,1437,402,1255]" pageId="15" pageNumber="216">
<emphasis id="B915EAE1FFE4FF9DFF42A3B5FED3FB45" bold="true" box="[199,315,1194,1219]" pageId="15" pageNumber="216">Remarks.</emphasis>
<taxonomicName id="4C614D70FFE4FF9DFEC7A3B5FE05FB45" box="[322,493,1194,1219]" class="Arachnida" family="Halacaridae" genus="Copidognathus" kingdom="Animalia" order="Trombidiformes" pageId="15" pageNumber="216" phylum="Arthropoda" rank="genus">
<emphasis id="B915EAE1FFE4FF9DFEC7A3B5FE05FB45" box="[322,493,1194,1219]" italics="true" pageId="15" pageNumber="216">Copidognathus</emphasis>
</taxonomicName>
is spread world-wide, in all climatic and depth zones. The genus is the most diverse halacarid taxon, it holds one-third of all species (
<bibRefCitation id="EFF04B02FFE4FF9DFD3CA3D1FC88FB61" author="Bartsch" box="[697,864,1230,1255]" pageId="15" pageNumber="216" refString="Bartsch, I. (2004 b) Geographical and ecological distribution of marine halacarid genera and species (Acari: Halacaridae). Experimental &amp; Applied Acarology, 34, 37 - 58. http: // dx. doi. org / 10.1023 / B: APPA. 0000044438.32992.35" type="journal article" year="2004" yearSuffix="b">Bartsch 2004b</bibRefCitation>
,
<bibRefCitation id="EFF04B02FFE4FF9DFCE9A3D0FC5DFB61" author="Bartsch" box="[876,949,1231,1255]" pageId="15" pageNumber="216" refString="Bartsch, I. (2009 a) Checklist of marine and freshwater halacarid mite genera and species (Halacaridae: Acari) with notes on synonyms, habitats, distribution and descriptions of the taxa. Zootaxa, 1998, 1 - 170." type="journal article" year="2009" yearSuffix="a">2009a</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
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