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<mods:title id="19F9D8C43B31E8952DF132ACD542EDC9">The genital area of Halacaridae (Acari), life stages and development of morphological characters and implication on the classification</mods:title>
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<paragraph id="8BDE36F3FFFAFF83FF12A787FE2AFF34" blockId="17.[151,450,152,211]" box="[151,450,152,178]" pageId="17" pageNumber="218">
<heading id="D096819FFFFAFF83FF12A787FE2AFF34" bold="true" box="[151,450,152,178]" fontSize="11" level="1" pageId="17" pageNumber="218" reason="1">
<taxonomicName id="4C614D70FFFAFF83FF12A787FE2AFF34" ID-CoL="4SMF" ID-ENA="2060444" authority="Viets, 1927" authorityName="Viets" authorityYear="1927" box="[151,450,152,178]" class="Arachnida" family="Halacaridae" genus="Halacarellus" kingdom="Animalia" order="Trombidiformes" pageId="17" pageNumber="218" phylum="Arthropoda" rank="genus">
<emphasis id="B915EAE1FFFAFF83FF12A787FE2AFF34" bold="true" box="[151,450,152,178]" pageId="17" pageNumber="218">
<emphasis id="B915EAE1FFFAFF83FF12A787FEDBFF34" bold="true" box="[151,307,152,178]" italics="true" pageId="17" pageNumber="218">Halacarellus</emphasis>
<bibRefCitation id="EFF04B02FFFAFF83FEBFA787FE2AFF34" author="Viets" box="[314,450,152,178]" pageId="17" pageNumber="218" refString="Viets, K. (1927) Die Halacaridae der Nordsee. Zeitschrift fur Wissenschaftliche Zoologie, 130, 83 - 173." type="journal article" year="1927">Viets, 1927</bibRefCitation>
</emphasis>
</taxonomicName>
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</paragraph>
<paragraph id="8BDE36F3FFFAFF83FF12A7A5FECFFF55" blockId="17.[151,450,152,211]" box="[151,295,186,211]" pageId="17" pageNumber="218">(Figs 4446)</paragraph>
</subSubSection>
<subSubSection id="C37B6578FFFAFF83FF12A61DFB6EFB89" pageId="17" pageNumber="218" type="materials_examined">
<paragraph id="8BDE36F3FFFAFF83FF12A61DFD27FE9D" blockId="17.[151,1437,258,1147]" box="[151,719,258,283]" pageId="17" pageNumber="218">
<emphasis id="B915EAE1FFFAFF83FF12A61DFEDAFE9D" bold="true" box="[151,306,258,283]" pageId="17" pageNumber="218">
<typeStatus id="54DA8851FFFAFF83FF12A61DFF3AFE9D" box="[151,210,258,283]" pageId="17" pageNumber="218">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="4C614D70FFFAFF83FEBCA61DFD22FE9D" authority="Lohmann, 1889" authorityName="Lohmann" authorityYear="1889" box="[313,714,258,283]" class="Arachnida" family="Halacaridae" genus="Halacarus" kingdom="Animalia" order="Trombidiformes" pageId="17" pageNumber="218" phylum="Arthropoda" rank="species" species="balticus">
<emphasis id="B915EAE1FFFAFF83FEBCA61DFDE7FE9D" box="[313,527,258,283]" italics="true" pageId="17" pageNumber="218">Halacarus balticus</emphasis>
<bibRefCitation id="EFF04B02FFFAFF83FD93A61DFD22FE9D" author="Lohmann" box="[534,714,258,283]" pageId="17" pageNumber="218" refString="Lohmann, H. (1889) Die Unterfamilie der Halacaridae Murr. und die Meeresmilben der Ostsee. Zoologische Jahrbucher, Abteilung fur Systematik, Okologie und Geographie der Tiere, 4, 269 - 408." type="journal article" year="1889">Lohmann, 1889</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="8BDE36F3FFFAFF83FF42A639FBEBFD4C" blockId="17.[151,1437,258,1147]" pageId="17" pageNumber="218">
<emphasis id="B915EAE1FFFAFF83FF42A639FEF5FEB9" bold="true" box="[199,285,294,319]" pageId="17" pageNumber="218">Adults.</emphasis>
Female and male with GA. Female GA with two to four (rarely seven) pairs of pgs, genital sclerites with zero to five pairs of sgs. Three pairs of internal acetabula always distinct, in freshwater inhabiting species enlarged (
<bibRefCitation id="EFF04B02FFFAFF83FE8DA671FE7DFE01" author="Viets" box="[264,405,366,391]" pageId="17" pageNumber="218" refString="Viets, K. (1928 a) Wassermilben aus dem Schwarzen Meer, dem Kaspischen Meer und dem Aral-See. Abhandlungen. Naturwissenschaftlicher Verein zu Bremen, 27, 47 - 80." type="journal article" year="1928" yearSuffix="a">Viets 1928a</bibRefCitation>
: fig. 49;
<bibRefCitation id="EFF04B02FFFAFF83FD85A671FCC9FE01" author="Bartsch" box="[512,801,366,391]" pageId="17" pageNumber="218" refString="Bartsch, I. &amp; Gerecke, R. (2011) A new freshwater mite of the marine genus Halacarellus (Acari: Halacaridae) from the Austrian Alps (Styria, Gesause National Park): Description and reflections on its origin. Zoologischer Anzeiger, 250, 151 - 159. http: // dx. doi. org / 10.1016 / j. jcz. 2011.02.003" type="journal article" year="2011">Bartsch &amp; Gerecke 2011</bibRefCitation>
: fig. 1H). Acetabula arranged in line along genital slit, anterior pair anterior or posterior to level of mid-GO (Fig. 44;
<bibRefCitation id="EFF04B02FFFAFF83FC14A68CFBE4FE2D" author="Otto" box="[913,1036,403,427]" pageId="17" pageNumber="218" refString="Otto, J. C. (1994) New species of Halacaridae from Australia (Acarina: Prostigmata). Acarologia, 35, 31 - 48." type="journal article" year="1994">Otto 1994</bibRefCitation>
: fig. 27). Ovipositor at rest often extending beyond GO. Ovipositor with two basal and eight or ten apical genital spines. Basal pair of genital spines papilli- or setiform. Apical genital spines dissimilar in shape, robust and with tines, slender and spiniform, or reduced to small papillae (Figs 45, 46;
<bibRefCitation id="EFF04B02FFFAFF83FDC9A6E1FCE6FD91" author="Womersley" box="[588,782,510,535]" pageId="17" pageNumber="218" refString="Womersley, H. (1937) Acarina. Australasian Antarctic Expedition 1911 - 1914, Scientific Reports, Series C, 10 (6), 1 - 24." type="journal article" year="1937">Womersley 1937</bibRefCitation>
: pl. 6, fig. 12;
<bibRefCitation id="EFF04B02FFFAFF83FC34A6E1FBB0FD91" author="Bartsch" box="[945,1112,510,535]" pageId="17" pageNumber="218" refString="Bartsch, I. (1998 b) Halacarinae (Acari, Halacaroidea) from the northwestern Black Sea: A review. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 95, 143 - 178." type="journal article" year="1998" yearSuffix="b">Bartsch 1998b</bibRefCitation>
: figs 31, 61, 2007b: fig. 6G). Male GA with 1050 pairs of pgs and in general five pairs of sgs (rarely less). In the majority of species all three pairs of acetabula internal, arranged along genital slit; in a few species (of the
<taxonomicName id="4C614D70FFFAFF83FB93A558FB42FDD8" box="[1046,1194,583,606]" class="Arachnida" family="Halacaridae" genus="Halacarellus" kingdom="Animalia" order="Trombidiformes" pageId="17" pageNumber="218" phylum="Arthropoda" rank="species" species="capuzinus">
<emphasis id="B915EAE1FFFAFF83FB93A558FB42FDD8" box="[1046,1194,583,606]" italics="true" pageId="17" pageNumber="218">H. capuzinus</emphasis>
</taxonomicName>
group) three pairs of acetabula arranged in posterior part of GO, within a fovea that opens to the exterior (
<bibRefCitation id="EFF04B02FFFAFF83FBECA575FA98FD05" author="Bartsch" box="[1129,1392,618,643]" pageId="17" pageNumber="218" refString="Bartsch, I. (1998 b) Halacarinae (Acari, Halacaroidea) from the northwestern Black Sea: A review. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 95, 143 - 178." type="journal article" year="1998" yearSuffix="b">Bartsch 1998b: fig. 48</bibRefCitation>
). In freshwater species two pairs of gac posterior
<collectingCountry id="F3767663FFFAFF83FD19A590FD0DFD21" box="[668,741,655,679]" name="Togo" pageId="17" pageNumber="218">to GO</collectingCountry>
, enlarged and external, one pair of gac small, near middle of GO (
<bibRefCitation id="EFF04B02FFFAFF83FF4BA5ADFEB1FD4D" author="Viets" box="[206,345,690,715]" pageId="17" pageNumber="218" refString="Viets, K. (1928 a) Wassermilben aus dem Schwarzen Meer, dem Kaspischen Meer und dem Aral-See. Abhandlungen. Naturwissenschaftlicher Verein zu Bremen, 27, 47 - 80." type="journal article" year="1928" yearSuffix="a">Viets 1928a</bibRefCitation>
: fig. 48;
<bibRefCitation id="EFF04B02FFFAFF83FE3EA5ADFD8AFD4D" author="Bartsch" box="[443,610,690,715]" pageId="17" pageNumber="218" refString="Bartsch, I. (1998 b) Halacarinae (Acari, Halacaroidea) from the northwestern Black Sea: A review. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 95, 143 - 178." type="journal article" year="1998" yearSuffix="b">Bartsch 1998b</bibRefCitation>
: fig. 15). AE without epimeral pores.
</paragraph>
<paragraph id="8BDE36F3FFFAFF83FF42A5C9FB6EFB89" blockId="17.[151,1437,258,1147]" pageId="17" pageNumber="218">
<emphasis id="B915EAE1FFFAFF83FF42A5C9FED4FD69" bold="true" box="[199,316,726,751]" pageId="17" pageNumber="218">Juveniles.</emphasis>
With larva and two nymphal stages. Nymphs of northern hemisphere species with genital and anal plate fused to a GA (e.g. in the
<typeStatus id="54DA8851FFFAFF83FD90A5E4FDADFC95" box="[533,581,763,787]" pageId="17" pageNumber="218">type</typeStatus>
species
<taxonomicName id="4C614D70FFFAFF83FD29A5E3FCC3FC95" box="[684,811,762,787]" class="Arachnida" family="Halacaridae" genus="Halacarus" kingdom="Animalia" order="Trombidiformes" pageId="17" pageNumber="218" phylum="Arthropoda" rank="species" species="balticus">
<emphasis id="B915EAE1FFFAFF83FD29A5E3FCC3FC95" box="[684,811,762,787]" italics="true" pageId="17" pageNumber="218">H. balticus</emphasis>
</taxonomicName>
), most southern hemisphere species with GP and AP separated. Deutonymphal GA with two pairs of pgs, zero to two pairs of sgs and two pairs of gac (
<bibRefCitation id="EFF04B02FFFAFF83FA84A401FA71FCB1" author="Newell" box="[1281,1433,798,823]" pageId="17" pageNumber="218" refString="Newell, I. M. (1951) New species of Agaue and Thalassarachna from the Aleutians (Acari, Halacaridae). American Museum Novitates, 1489, 1 - 19." type="journal article" year="1951">Newell 1951</bibRefCitation>
: figs 3, 19;
<bibRefCitation id="EFF04B02FFFAFF83FE8BA45DFE4FFCDD" author="Bartsch" box="[270,423,834,859]" pageId="17" pageNumber="218" refString="Bartsch, I. (1972) Ein Beitrag zur Systematik, Biologie und Okologie der Halacaridae (Acari) aus dem Litoral der Nord- und Ostsee. I. Systematik und Biologie. Abhandlungen und Verhandlungen des Naturwissenschaftlichen Vereins zu Hamburg, Neue Folge, 16, 155 - 230." type="journal article" year="1972">Bartsch 1972</bibRefCitation>
: figs 25B, 28B, 1995a: fig. 19, 1998b: figs 24, 41, 63, 82, 101, 1990c: fig. 137, 1999c: fig. 3F). Acetabula of nymphs small and internal, large and internal or large and external (e.g. in
<taxonomicName id="4C614D70FFFAFF83FB45A478FF30FC25" authority="Viets, 1928" authorityName="Viets" authorityYear="1928" class="Arachnida" family="Halacaridae" genus="Halacarus" kingdom="Animalia" order="Trombidiformes" pageId="17" pageNumber="218" phylum="Arthropoda" rank="species" species="hyrcanus">
<emphasis id="B915EAE1FFFAFF83FB45A478FAA2FCF9" box="[1216,1354,870,895]" italics="true" pageId="17" pageNumber="218">H. hyrcanus</emphasis>
(Viets, 1928)
</taxonomicName>
and
<taxonomicName id="4C614D70FFFAFF83FE92A493FD3DFC25" authority="Bartsch &amp; Gerecke 2011" authorityName="Bartsch &amp; Gerecke" authorityYear="2011" box="[279,725,906,931]" class="Arachnida" family="Halacaridae" genus="Halacarus" kingdom="Animalia" order="Trombidiformes" pageId="17" pageNumber="218" phylum="Arthropoda" rank="species" species="fontinalis">
<emphasis id="B915EAE1FFFAFF83FE92A493FE40FC25" box="[279,424,906,931]" italics="true" pageId="17" pageNumber="218">H. fontinalis</emphasis>
<bibRefCitation id="EFF04B02FFFAFF83FE36A495FD3DFC25" author="Bartsch" box="[435,725,906,931]" pageId="17" pageNumber="218" refString="Bartsch, I. &amp; Gerecke, R. (2011) A new freshwater mite of the marine genus Halacarellus (Acari: Halacaridae) from the Austrian Alps (Styria, Gesause National Park): Description and reflections on its origin. Zoologischer Anzeiger, 250, 151 - 159. http: // dx. doi. org / 10.1016 / j. jcz. 2011.02.003" type="journal article" year="2011">Bartsch &amp; Gerecke 2011</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EFF04B02FFFAFF83FD62A495FC7CFC25" author="Bartsch" box="[743,916,906,931]" pageId="17" pageNumber="218" refString="Bartsch, I. (1998 b) Halacarinae (Acari, Halacaroidea) from the northwestern Black Sea: A review. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 95, 143 - 178." type="journal article" year="1998" yearSuffix="b">Bartsch 1998b</bibRefCitation>
: fig. 24;
<bibRefCitation id="EFF04B02FFFAFF83FB86A495FAC1FC25" author="Bartsch" box="[1027,1321,906,931]" pageId="17" pageNumber="218" refString="Bartsch, I. &amp; Gerecke, R. (2011) A new freshwater mite of the marine genus Halacarellus (Acari: Halacaridae) from the Austrian Alps (Styria, Gesause National Park): Description and reflections on its origin. Zoologischer Anzeiger, 250, 151 - 159. http: // dx. doi. org / 10.1016 / j. jcz. 2011.02.003" type="journal article" year="2011">Bartsch &amp; Gerecke 2011</bibRefCitation>
: fig. 2G). Protonymphal GA generally with pair of internal gac; pgs and sgs lacking (
<bibRefCitation id="EFF04B02FFFAFF83FC5FA4B1FB9CFC41" author="Bartsch" box="[986,1140,942,967]" pageId="17" pageNumber="218" refString="Bartsch, I. (1972) Ein Beitrag zur Systematik, Biologie und Okologie der Halacaridae (Acari) aus dem Litoral der Nord- und Ostsee. I. Systematik und Biologie. Abhandlungen und Verhandlungen des Naturwissenschaftlichen Vereins zu Hamburg, Neue Folge, 16, 155 - 230." type="journal article" year="1972">Bartsch 1972</bibRefCitation>
: figs 25D, 28F, 1995a: fig. 11, 1998b: figs 44, 85, 1999c: fig. 3G). Larva with pair of epimeral pores on AE (
<bibRefCitation id="EFF04B02FFFAFF83FBAAA4CDFA66FC6D" author="Bartsch" box="[1071,1422,978,1003]" pageId="17" pageNumber="218" refString="Bartsch, I. &amp; Panesar, A. R. (2000) Die Meeresmilbe Caspihalacarus hyrcanus in der Donau bei Wien, ein ponto-kaspisches Relikt. Natur und Museum, 130 (8), 258 - 263." type="journal article" year="2000">Bartsch &amp; Panesar 2000: fig. 3</bibRefCitation>
). Larval anal plate often with large ventral part (
<bibRefCitation id="EFF04B02FFFAFF83FD24A4E9FCD3FB89" author="Bartsch" box="[673,827,1014,1039]" pageId="17" pageNumber="218" refString="Bartsch, I. (1972) Ein Beitrag zur Systematik, Biologie und Okologie der Halacaridae (Acari) aus dem Litoral der Nord- und Ostsee. I. Systematik und Biologie. Abhandlungen und Verhandlungen des Naturwissenschaftlichen Vereins zu Hamburg, Neue Folge, 16, 155 - 230." type="journal article" year="1972">Bartsch 1972</bibRefCitation>
: fig. 25F, 1998b: figs 67, 88).
</paragraph>
</subSubSection>
<subSubSection id="C37B6578FFFAFF83FF42A305FC40FBFD" pageId="17" pageNumber="218" type="discussion">
<paragraph id="8BDE36F3FFFAFF83FF42A305FC40FBFD" blockId="17.[151,1437,258,1147]" pageId="17" pageNumber="218">
<emphasis id="B915EAE1FFFAFF83FF42A305FED3FBB5" bold="true" box="[199,315,1050,1075]" pageId="17" pageNumber="218">Remarks.</emphasis>
The genus, with about 50 species, is spread world-wide (
<bibRefCitation id="EFF04B02FFFAFF83FC3AA305FB8FFBB5" author="Bartsch" box="[959,1127,1050,1075]" pageId="17" pageNumber="218" 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">Bartsch 2009a</bibRefCitation>
). Several of the species can be assigned to distinct species groups (
<bibRefCitation id="EFF04B02FFFAFF83FDCBA321FD1EFBD1" author="Bartsch" box="[590,758,1086,1111]" pageId="17" pageNumber="218" refString="Bartsch, I. (1997 a) Thalassarachna and Halacarellus (Halacaridae: Acari): two separate genera. Journal of Natural History, 31, 1223 - 1236. http: // dx. doi. org / 10.1080 / 00222939700770661" type="journal article" year="1997" yearSuffix="a">Bartsch 1997a</bibRefCitation>
). Representatives of the northern Atlantic Ocean groups are not found on the southern hemisphere and vice versa (
<bibRefCitation id="EFF04B02FFFAFF83FD70A37DFC73FBFD" author="Bartsch" box="[757,923,1122,1147]" pageId="17" pageNumber="218" refString="Bartsch, I. (1997 a) Thalassarachna and Halacarellus (Halacaridae: Acari): two separate genera. Journal of Natural History, 31, 1223 - 1236. http: // dx. doi. org / 10.1080 / 00222939700770661" type="journal article" year="1997" yearSuffix="a">Bartsch 1997a</bibRefCitation>
).
</paragraph>
</subSubSection>
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