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<mods:title id="AC9979B11155453BAF57310D8A625998">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="8BDE36F3FFF8FF81FF12A364FE53FB13" blockId="19.[151,443,1147,1206]" box="[151,443,1147,1173]" pageId="19" pageNumber="220">
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<taxonomicName id="4C614D70FFF8FF81FF12A364FE53FB13" authority="Newell, 1947" authorityName="Newell" authorityYear="1947" box="[151,443,1147,1173]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="19" pageNumber="220" phylum="Arthropoda" rank="genus">
<emphasis id="B915EAE1FFF8FF81FF12A364FE53FB13" bold="true" box="[151,443,1147,1173]" pageId="19" pageNumber="220">
<emphasis id="B915EAE1FFF8FF81FF12A364FEFEFB13" bold="true" box="[151,278,1147,1173]" italics="true" pageId="19" pageNumber="220">Isobactrus</emphasis>
<bibRefCitation id="EFF04B02FFF8FF81FE98A364FE53FB13" author="Newell" box="[285,443,1147,1173]" pageId="19" pageNumber="220" 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>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="8BDE36F3FFF8FF81FF12A382FECFFB30" blockId="19.[151,443,1147,1206]" box="[151,295,1181,1206]" pageId="19" pageNumber="220">
(
<figureCitation id="135A2A76FFF8FF81FF1AA382FF1AFB30" box="[159,242,1181,1206]" captionStart="FIGURE 53" captionStartId="20.[151,250,1357,1379]" captionTargetBox="[166,1416,186,1320]" captionTargetId="figure@20.[151,1436,179,1336]" captionTargetPageId="20" captionText="FIGURE 53. Isobactrus hartmanni Bartsch, 1972, genital region, female. FIGURES 54 55. Isobactrus obesus Bartsch, 1992, 54. genital plate, female; 55. genital region, tritonymph. FIGURES 56 59. Isobactrus aspidotus Bartsch, 2009, 56. genital opening, male; 57. genital region, tritonymph; 58. genital region, deutonymph; 59. genital region, protonymph. FIGURE 60. Isobactrus uniscutatus (Viets, 1939), genital plate, male. FIGURE 61. Lohmannella falcata (Hodge, 1863), genital plate, protonymph. FIGURES 62 63. Makarovana spinosa (Bartsch, 1978), female, 62. posterior part of genitoanal plate; 63. genital opening with everted ovipositor and two genital spines (enlarged), lateral aspect. (bgsp, basal genital spine; pgs, perigenital setae; pogs, postgenital sclerite; prgs, pregenital sclerite) Scale = 50 µm." httpUri="https://zenodo.org/record/245251/files/figure.png" pageId="19" pageNumber="220">Figs 53</figureCitation>
60)
</paragraph>
</subSubSection>
<subSubSection id="C37B6578FFF8FF86FF12A3FAFA83F86A" lastPageId="20" lastPageNumber="221" pageId="19" pageNumber="220" type="materials_examined">
<paragraph id="8BDE36F3FFF8FF81FF12A3FAFD7DFB78" blockId="19.[151,1437,1253,2034]" box="[151,661,1253,1278]" pageId="19" pageNumber="220">
<emphasis id="B915EAE1FFF8FF81FF12A3FAFEDAFB78" bold="true" box="[151,306,1253,1278]" pageId="19" pageNumber="220">
<typeStatus id="54DA8851FFF8FF81FF12A3FAFF3AFB78" box="[151,210,1253,1278]" pageId="19" pageNumber="220">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="4C614D70FFF8FF81FEBCA3FAFD78FB78" authority="Lohmann, 1889" authorityName="Lohmann" authorityYear="1889" box="[313,656,1253,1278]" class="Magnoliopsida" family="Apiaceae" genus="Aletes" kingdom="Animalia" order="Apiales" pageId="19" pageNumber="220" phylum="Tracheophyta" rank="species" species="setosus">
<emphasis id="B915EAE1FFF8FF81FEBCA3FAFE3DFB7B" box="[313,469,1253,1278]" italics="true" pageId="19" pageNumber="220">Aletes setosus</emphasis>
<bibRefCitation id="EFF04B02FFF8FF81FE59A3FAFD78FB78" author="Lohmann" box="[476,656,1253,1278]" pageId="19" pageNumber="220" 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="8BDE36F3FFF8FF81FF42A216FDA6F874" blockId="19.[151,1437,1253,2034]" pageId="19" pageNumber="220">
<emphasis id="B915EAE1FFF8FF81FF42A216FEF5FAA4" bold="true" box="[199,285,1289,1314]" pageId="19" pageNumber="220">Adults.</emphasis>
Female GP delicate and often in form of pair of crescentic plates on either side of GO, not or only slightly longer than GO and not distinctly delimited from striated integument. Female genital area with three (rarely four) pairs of pgs, anterior pair in striated integument, anterior
<collectingCountry id="F3767663FFF8FF81FC39A24DFBEDFAEC" box="[956,1029,1362,1386]" name="Togo" pageId="19" pageNumber="220">to GO</collectingCountry>
, following two pairs level with and posterior
<collectingCountry id="F3767663FFF8FF81FE83A269FEBBFA08" box="[262,339,1398,1422]" name="Togo" pageId="19" pageNumber="220">to GO</collectingCountry>
, respectively, in margin or outside GP. Anterior and posterior edge of genital foramen often enforced by internal pregenital and postgenital sclerites (
<figureCitation id="135A2A76FFF8FF81FCA1A286FC91FA34" box="[804,889,1433,1458]" captionStart="FIGURE 53" captionStartId="20.[151,250,1357,1379]" captionTargetBox="[166,1416,186,1320]" captionTargetId="figure@20.[151,1436,179,1336]" captionTargetPageId="20" captionText="FIGURE 53. Isobactrus hartmanni Bartsch, 1972, genital region, female. FIGURES 54 55. Isobactrus obesus Bartsch, 1992, 54. genital plate, female; 55. genital region, tritonymph. FIGURES 56 59. Isobactrus aspidotus Bartsch, 2009, 56. genital opening, male; 57. genital region, tritonymph; 58. genital region, deutonymph; 59. genital region, protonymph. FIGURE 60. Isobactrus uniscutatus (Viets, 1939), genital plate, male. FIGURE 61. Lohmannella falcata (Hodge, 1863), genital plate, protonymph. FIGURES 62 63. Makarovana spinosa (Bartsch, 1978), female, 62. posterior part of genitoanal plate; 63. genital opening with everted ovipositor and two genital spines (enlarged), lateral aspect. (bgsp, basal genital spine; pgs, perigenital setae; pogs, postgenital sclerite; prgs, pregenital sclerite) Scale = 50 µm." httpUri="https://zenodo.org/record/245251/files/figure.png" pageId="19" pageNumber="220">Fig. 53</figureCitation>
). Genital sclerites with one to two pairs of sgs. Female with two or three almost equal-sized internal tube-like gac (Figs 54, 56), or two pairs of distinct, tube-like and one pair of small gac (
<figureCitation id="135A2A76FFF8FF81FE4AA2FEFDCDFA7C" box="[463,549,1505,1530]" captionStart="FIGURE 53" captionStartId="20.[151,250,1357,1379]" captionTargetBox="[166,1416,186,1320]" captionTargetId="figure@20.[151,1436,179,1336]" captionTargetPageId="20" captionText="FIGURE 53. Isobactrus hartmanni Bartsch, 1972, genital region, female. FIGURES 54 55. Isobactrus obesus Bartsch, 1992, 54. genital plate, female; 55. genital region, tritonymph. FIGURES 56 59. Isobactrus aspidotus Bartsch, 2009, 56. genital opening, male; 57. genital region, tritonymph; 58. genital region, deutonymph; 59. genital region, protonymph. FIGURE 60. Isobactrus uniscutatus (Viets, 1939), genital plate, male. FIGURE 61. Lohmannella falcata (Hodge, 1863), genital plate, protonymph. FIGURES 62 63. Makarovana spinosa (Bartsch, 1978), female, 62. posterior part of genitoanal plate; 63. genital opening with everted ovipositor and two genital spines (enlarged), lateral aspect. (bgsp, basal genital spine; pgs, perigenital setae; pogs, postgenital sclerite; prgs, pregenital sclerite) Scale = 50 µm." httpUri="https://zenodo.org/record/245251/files/figure.png" pageId="19" pageNumber="220">Fig. 53</figureCitation>
;
<bibRefCitation id="EFF04B02FFF8FF81FDB6A2FEFD28FA7C" author="Viets" box="[563,704,1505,1530]" pageId="19" pageNumber="220" refString="Viets, K. (1939 b) Meeresmilben aus der Adria (Halacaridae und Hydrachnellae, Acari). Archiv fur Naturgeschichte, (Neue Folge) 8, 518 - 550." type="journal article" year="1939" yearSuffix="b">Viets 1939b</bibRefCitation>
: fig. 21;
<bibRefCitation id="EFF04B02FFF8FF81FCAFA2FEFC28FA7C" author="Newell" box="[810,960,1505,1530]" pageId="19" pageNumber="220" 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 55, 80;
<bibRefCitation id="EFF04B02FFF8FF81FBDFA2FEFAECFA7C" author="Bartsch" box="[1114,1284,1505,1530]" pageId="19" pageNumber="220" refString="Bartsch, I. (1974 b) Uber das Auftreten von Epimeralporen besonders bei den Rhombognathinae (Halacaridae, Acari). Zoologischer Anzeiger, 193, 266 - 268." type="journal article" year="1974" yearSuffix="b">Bartsch 1974b</bibRefCitation>
: fig. 2b;
<bibRefCitation id="EFF04B02FFF8FF81FAE8A2FEFF37F998" author="Abe" pageId="19" pageNumber="220" refString="Abe, H. (1996 a) Rhombognathine mites (Acari: Halacaridae) from Hokkaido, Northern Japan. Publications of the Seto Marine Biological Laboratory, 37, 63 - 166." type="journal article" year="1996" yearSuffix="a">Abé 1996a</bibRefCitation>
: figs 4A, 8A, 16A, 20A, 1996b: fig. 2A) (the posterior markings illustrated in
<bibRefCitation id="EFF04B02FFF8FF81FBE4A11AFB1EF998" author="Newell" box="[1121,1270,1541,1566]" pageId="19" pageNumber="220" 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>
: fig. 59 are no acetabula). Ovipositor short, both at rest and when everted. Genital spines well-sclerotized, rather wide and palmate, generally ending with several tines; 1011 spines present, four in anterior and six to seven in posterior position (
<bibRefCitation id="EFF04B02FFF8FF81FE85A16EFE4FF90C" author="Bartsch" box="[256,423,1649,1674]" pageId="19" pageNumber="220" refString="Bartsch, I. (1975 a) Ein Beitrag zum System der Rhombognathinen (Halacaridae, Acari). Zur Morphologie der Tarsalregion und des Ovipositors. Zoologischer Anzeiger, 194, 193 - 200." type="journal article" year="1975" yearSuffix="a">Bartsch 1975a</bibRefCitation>
: figs 10b, 11, 1975b: fig 103, 2000: fig. 1D;
<bibRefCitation id="EFF04B02FFF8FF81FC1BA16EFBDAF90C" author="Newell" box="[926,1074,1649,1674]" pageId="19" pageNumber="220" refString="Newell, I. M. (1984) Antarctic Halacaroidea. Antarctic Research Series, 40, 1 - 284. http: // dx. doi. org / 10.1029 / AR 040" type="journal article" year="1984">Newell 1984</bibRefCitation>
: fig. 42). GP in male larger than in female, with about 30100 perigenital setae, these often arranged in an inner and outer ring around GO and inserted on GP, rarely in striated integument outside GP. Setae generally slender and smooth, in a few species some of anterior setae of inner ring short, bristle-like or spiniform, namely in
<taxonomicName id="4C614D70FFF8FF81FB85A1C1FAA2F970" authority="Newell, 1947" authorityName="Newell" authorityYear="1947" box="[1024,1354,1757,1782]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="19" pageNumber="220" phylum="Arthropoda" rank="species" species="hutchinsoni">
<emphasis id="B915EAE1FFF8FF81FB85A1C1FB48F970" box="[1024,1184,1757,1782]" italics="true" pageId="19" pageNumber="220">I. hutchinsoni</emphasis>
<bibRefCitation id="EFF04B02FFF8FF81FB28A1C2FAA2F970" author="Newell" box="[1197,1354,1757,1782]" pageId="19" pageNumber="220" 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>
</taxonomicName>
and
<taxonomicName id="4C614D70FFF8FF81FA09A1C1FE46F89C" authority="Viets, 1939" authorityName="Viets" authorityYear="1939" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="19" pageNumber="220" phylum="Arthropoda" rank="species" species="uniscutatus">
<emphasis id="B915EAE1FFF8FF81FA09A1C1FEFFF89F" italics="true" pageId="19" pageNumber="220">I. uniscutatus</emphasis>
(Viets, 1939)
</taxonomicName>
(Fig. 60;
<bibRefCitation id="EFF04B02FFF8FF81FD9FA01EFD46F89C" author="Newell" box="[538,686,1793,1818]" pageId="19" pageNumber="220" 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>
: fig. 76). Genital sclerites with four (rarely three) pairs of sgs, these setiform, rarely flattened and foliate or spur-like. Acetabula generally slightly smaller than in females and situated in posterior part of GO or even posterior
<collectingCountry id="F3767663FFF8FF81FDECA055FD58F8E4" box="[617,688,1866,1890]" name="Togo" pageId="19" pageNumber="220">to GO</collectingCountry>
(Fig. 60). As in female, male GO with either two to three pairs of large almost equal-sized gac or two large pairs and one anterior small pair (Fig. 56;
<bibRefCitation id="EFF04B02FFF8FF81FBDDA072FB0CF800" author="Viets" box="[1112,1252,1901,1926]" pageId="19" pageNumber="220" refString="Viets, K. (1939 b) Meeresmilben aus der Adria (Halacaridae und Hydrachnellae, Acari). Archiv fur Naturgeschichte, (Neue Folge) 8, 518 - 550." type="journal article" year="1939" yearSuffix="b">Viets 1939b</bibRefCitation>
: fig. 21;
<bibRefCitation id="EFF04B02FFF8FF81FACEA072FF3BF82C" author="Newell" pageId="19" pageNumber="220" 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>
: fig. 76, 1984: fig. 37;
<bibRefCitation id="EFF04B02FFF8FF81FE55A08EFDA6F82C" author="Abe" box="[464,590,1937,1962]" pageId="19" pageNumber="220" refString="Abe, H. (1996 a) Rhombognathine mites (Acari: Halacaridae) from Hokkaido, Northern Japan. Publications of the Seto Marine Biological Laboratory, 37, 63 - 166." type="journal article" year="1996" yearSuffix="a">Abé 1996a</bibRefCitation>
: figs 4B, 16B; 1996b: fig. 2B;
<bibRefCitation id="EFF04B02FFF8FF81FC26A08EFBA0F82C" author="Bartsch" box="[931,1096,1937,1962]" pageId="19" pageNumber="220" 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. 2M). Acetabula generally internal, in two species (
<taxonomicName id="4C614D70FFF8FF81FE23A0A9FDA8F848" box="[422,576,1973,1998]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="19" pageNumber="220" phylum="Arthropoda" rank="species" species="hutchinsoni">
<emphasis id="B915EAE1FFF8FF81FE23A0A9FDA8F848" box="[422,576,1973,1998]" italics="true" pageId="19" pageNumber="220">I. hutchinsoni</emphasis>
</taxonomicName>
,
<taxonomicName id="4C614D70FFF8FF81FDC8A0A9FD0CF84B" box="[589,740,1974,1997]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="19" pageNumber="220" phylum="Arthropoda" rank="species" species="uniscutatus">
<emphasis id="B915EAE1FFF8FF81FDC8A0A9FDBEF84B" box="[589,598,1974,1997]" italics="true" pageId="19" pageNumber="220">I</emphasis>
.
<emphasis id="B915EAE1FFF8FF81FDE1A0A9FD0CF84B" box="[612,740,1974,1997]" italics="true" pageId="19" pageNumber="220">uniscutatus</emphasis>
</taxonomicName>
) two pairs of external acetabula present (
<bibRefCitation id="EFF04B02FFF8FF81FB28A0AAFAAAF848" author="Newell" box="[1197,1346,1973,1998]" pageId="19" pageNumber="220" 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>
: fig. 76;
<bibRefCitation id="EFF04B02FFF8FF81FF12A0C6FED5F874" author="Bartsch" box="[151,317,2009,2034]" pageId="19" pageNumber="220" 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. 13, 2003c: fig. 3F).
</paragraph>
<caption id="DF1E667BFFFFFF86FF12A252FC9DF99D" httpUri="https://zenodo.org/record/245251/files/figure.png" pageId="20" pageNumber="221" targetBox="[166,1416,186,1320]" targetPageId="20">
<paragraph id="8BDE36F3FFFFFF86FF12A252FC9DF99D" blockId="20.[151,1436,1357,1563]" pageId="20" pageNumber="221">
<emphasis id="B915EAE1FFFFFF86FF12A252FEC8FAE5" bold="true" box="[151,288,1357,1379]" pageId="20" pageNumber="221">FIGURE 53.</emphasis>
<taxonomicName id="4C614D70FFFFFF86FEA0A252FD7DFAE5" authority="Bartsch, 1972" authorityName="Bartsch" authorityYear="1972" box="[293,661,1357,1379]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="20" pageNumber="221" phylum="Arthropoda" rank="species" species="hartmanni">
<emphasis id="B915EAE1FFFFFF86FEA0A252FDE8FAE5" box="[293,512,1357,1379]" italics="true" pageId="20" pageNumber="221">Isobactrus hartmanni</emphasis>
Bartsch, 1972
</taxonomicName>
, genital region, female.
<emphasis id="B915EAE1FFFFFF86FC0FA252FBAEFAE5" bold="true" box="[906,1094,1357,1379]" pageId="20" pageNumber="221">FIGURES 5455.</emphasis>
<taxonomicName id="4C614D70FFFFFF86FBC9A252FA70FAE5" authority="Bartsch, 1992" authorityName="Bartsch" authorityYear="1992" box="[1100,1432,1357,1379]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="20" pageNumber="221" phylum="Arthropoda" rank="species" species="obesus">
<emphasis id="B915EAE1FFFFFF86FBC9A252FAEBFAE5" box="[1100,1283,1357,1379]" italics="true" pageId="20" pageNumber="221">Isobactrus obesus</emphasis>
Bartsch, 1992
</taxonomicName>
, 54. genital plate, female; 55. genital region, tritonymph.
<emphasis id="B915EAE1FFFFFF86FD6CA273FC41FA04" bold="true" box="[745,937,1388,1410]" pageId="20" pageNumber="221">FIGURES 5659.</emphasis>
<taxonomicName id="4C614D70FFFFFF86FC37A273FAC9FA04" authority="Bartsch, 2009" authorityName="Bartsch" authorityYear="2009" box="[946,1313,1388,1410]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="20" pageNumber="221" phylum="Arthropoda" rank="species" species="aspidotus">
<emphasis id="B915EAE1FFFFFF86FC37A273FB6EFA04" box="[946,1158,1388,1410]" italics="true" pageId="20" pageNumber="221">Isobactrus aspidotus</emphasis>
Bartsch, 2009
</taxonomicName>
, 56. genital opening, male; 57. genital region, tritonymph; 58. genital region, deutonymph; 59. genital region, protonymph.
<emphasis id="B915EAE1FFFFFF86FA97A294FA74FA27" bold="true" box="[1298,1436,1419,1441]" pageId="20" pageNumber="221">FIGURE 60.</emphasis>
<taxonomicName id="4C614D70FFFFFF86FF12A2B6FDFBFA39" authority="Viets, 1939" authorityName="Viets" authorityYear="1939" box="[151,531,1449,1471]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="20" pageNumber="221" phylum="Arthropoda" rank="species" species="uniscutatus">
<emphasis id="B915EAE1FFFFFF86FF12A2B6FE68FA39" box="[151,384,1449,1471]" italics="true" pageId="20" pageNumber="221">Isobactrus uniscutatus</emphasis>
(Viets, 1939)
</taxonomicName>
, genital plate, male.
<emphasis id="B915EAE1FFFFFF86FD70A2B6FC6AFA39" bold="true" box="[757,898,1449,1471]" pageId="20" pageNumber="221">FIGURE 61.</emphasis>
<taxonomicName id="4C614D70FFFFFF86FC09A2B6FAEFFA39" authority="Hodge, 1863" authorityName="Hodge" authorityYear="1863" box="[908,1287,1449,1471]" class="Arachnida" family="Halacaridae" genus="Lohmannella" kingdom="Animalia" order="Trombidiformes" pageId="20" pageNumber="221" phylum="Arthropoda" rank="species" species="falcata">
<emphasis id="B915EAE1FFFFFF86FC09A2B6FB8AFA39" box="[908,1122,1449,1471]" italics="true" pageId="20" pageNumber="221">Lohmannella falcata</emphasis>
(Hodge, 1863)
</taxonomicName>
, genital plate, protonymph.
<emphasis id="B915EAE1FFFFFF86FE9AA2D7FE33FA58" bold="true" box="[287,475,1480,1502]" pageId="20" pageNumber="221">FIGURES 6263.</emphasis>
<taxonomicName id="4C614D70FFFFFF86FE64A2D7FCB3FA58" authority="Bartsch, 1978" authorityName="Bartsch" authorityYear="1978" box="[481,859,1480,1502]" class="Arachnida" family="Halacaridae" genus="Makarovana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Trombidiformes" pageId="20" pageNumber="221" phylum="Arthropoda" rank="species" species="spinosa">
<emphasis id="B915EAE1FFFFFF86FE64A2D7FD5CFA58" box="[481,692,1480,1502]" italics="true" pageId="20" pageNumber="221">Makarovana spinosa</emphasis>
(Bartsch, 1978)
</taxonomicName>
, female, 62. posterior part of genitoanal plate; 63. genital opening with everted ovipositor and two genital spines (enlarged), lateral aspect. (bgsp, basal genital spine; pgs, perigenital setae; pogs, postgenital sclerite; prgs, pregenital sclerite) Scale = 50 µm.
</paragraph>
</caption>
<paragraph id="8BDE36F3FFFFFF86FF42A157FA83F86A" blockId="20.[151,1437,1608,2029]" pageId="20" pageNumber="221">
<emphasis id="B915EAE1FFFFFF86FF42A157FED4F9E7" bold="true" box="[199,316,1608,1633]" pageId="20" pageNumber="221">Juveniles.</emphasis>
With four juvenile stages, one larva and three nymphs (proto-, deuto-, tritonymph). Nymphal genital and anal plate in some species separated in others fused. Tritonymph with a pair of small sgs and two pairs of pgs, one pair in striated integument anterior to GP, one pair on or close to GP. Genital region with two or three pairs of almost equal-sized acetabula (Fig. 55, 57;
<bibRefCitation id="EFF04B02FFFFFF86FD65A1ABFC9EF94A" author="Newell" box="[736,886,1716,1741]" pageId="20" pageNumber="221" 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>
: fig. 81;
<bibRefCitation id="EFF04B02FFFFFF86FC65A1ABFB92F94A" author="Bartsch" box="[992,1146,1716,1741]" pageId="20" pageNumber="221" 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. 18G, 2005c: fig. 14, 2009b: figs 5F, 7G;
<bibRefCitation id="EFF04B02FFFFFF86FEFFA1C7FE13F977" author="Abe" box="[378,507,1752,1777]" pageId="20" pageNumber="221" refString="Abe, H. (1996 b) A new species of the genus Isobactrus (Acari: Halacaridae) from Ponape, Micronesia, with special reference to intrageneric character state distribution. Species Diversity, 1, 17 - 29." type="journal article" year="1996" yearSuffix="b">Abé 1996b</bibRefCitation>
: fig. 2C) or with pair of large anterior acetabula, small, short middle pair, more or less adjacent to anterior pair of gac, and slender posterior pair (
<bibRefCitation id="EFF04B02FFFFFF86FCE2A1E3FBEBF892" author="Bartsch" box="[871,1027,1788,1813]" pageId="20" pageNumber="221" 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. 21D, 1975b: figs 81, 100, 121;
<bibRefCitation id="EFF04B02FFFFFF86FF12A03FFEFDF8BF" author="Abe" box="[151,277,1824,1849]" pageId="20" pageNumber="221" refString="Abe, H. (1996 a) Rhombognathine mites (Acari: Halacaridae) from Hokkaido, Northern Japan. Publications of the Seto Marine Biological Laboratory, 37, 63 - 166." type="journal article" year="1996" yearSuffix="a">Abé 1996a</bibRefCitation>
: figs 16C, 23A). Deuto- and protonymph without sgs. Deutonymph with single pair of pgs situated level with gac and on or in margin of GP. Majority of deutonymphs with two pairs of gac, either equal-sized (
<bibRefCitation id="EFF04B02FFFFFF86FA9DA05BFF07F806" author="Abe" pageId="20" pageNumber="221" refString="Abe, H. (1996 b) A new species of the genus Isobactrus (Acari: Halacaridae) from Ponape, Micronesia, with special reference to intrageneric character state distribution. Species Diversity, 1, 17 - 29." type="journal article" year="1996" yearSuffix="b">
Abé 1996b:
<figureCitation id="135A2A76FFFFFF86FF12A077FF32F806" box="[151,218,1896,1921]" captionStart="FIGURES 1 2" captionStartId="8.[151,264,1654,1676]" captionTargetBox="[163,1424,208,1620]" captionTargetId="figure@8.[151,1436,195,1630]" captionTargetPageId="8" captionText="FIGURES 1 2. Acanthohalacarus reticulatus Bartsch, 2001, 1. genital opening, female; 2. genital opening, male. FIGURES 3 4. Acarochelopodia biunguis Bartsch, 1988, 3. genitoanal plate, male; 4. genital plate, deutonymph. FIGURES 5 6. Acaromantis armatus Bartsch, 1977, 5. genital opening, female; 6. genital plate, protonymph. FIGURE 7. Actacarus spinosus Otto, 2000, posterior part of genitoanal plate, female. FIGURE 8. Agaue galatea Otto, 1999, genital plate, deutonymph. FIGURES 9 10. Agauopsis arborea Bartsch, 2003, 9. genital opening, female; 10. genital opening, male. FIGURE 11. Agauopsis arabia Bartsch &amp; Chatterjee, 2001, everted ovipositor and two genital spines (enlarged), ventral aspect, female. FIGURE 12. Agauopsis mokari Otto, 1994, genital opening, male. FIGURE 13. Agauopsis sp. (A. furcata group), genital opening, female. FIGURES 14 15. Agauopsis moorea Bartsch, 1992, 14. genital opening, female; 15. genital opening, male. (agsp, apical genital spine; bgsp, basal genital spine; gac, genital acetabula; GF, genital foramen; gsc, genital sclerite; gsp, genital spines; pgs, perigenital setae; sgs, subgenital seta) Scale = 50 µm" httpUri="https://zenodo.org/record/245247/files/figure.png" pageId="20" pageNumber="221">Fig. 2</figureCitation>
D
</bibRefCitation>
) or with small anterior and large posterior pair of gac (
<bibRefCitation id="EFF04B02FFFFFF86FCE2A077FBEBF806" author="Bartsch" box="[871,1027,1896,1921]" pageId="20" pageNumber="221" 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. 21C, 1975b: figs 82, 101, 120;
<bibRefCitation id="EFF04B02FFFFFF86FF12A093FEFDF823" author="Abe" box="[151,277,1932,1957]" pageId="20" pageNumber="221" refString="Abe, H. (1996 a) Rhombognathine mites (Acari: Halacaridae) from Hokkaido, Northern Japan. Publications of the Seto Marine Biological Laboratory, 37, 63 - 166." type="journal article" year="1996" yearSuffix="a">Abé 1996a</bibRefCitation>
: figs 4C, 23B), or with single pair of gac (Fig. 58;
<bibRefCitation id="EFF04B02FFFFFF86FCD5A093FC0CF822" author="Newell" box="[848,996,1932,1957]" pageId="20" pageNumber="221" 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>
: p. 80;
<bibRefCitation id="EFF04B02FFFFFF86FBB3A093FB27F822" author="Bartsch" box="[1078,1231,1932,1957]" pageId="20" pageNumber="221" 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>
: p. 191, fig. 18D). Protonymph with single pair of internal gac; pgs and sgs lacking (Fig. 59;
<bibRefCitation id="EFF04B02FFFFFF86FC67A0AFFB93F84E" author="Bartsch" box="[994,1147,1968,1993]" pageId="20" pageNumber="221" 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. 21B, 1975b: figs 83, 102, 119;
<bibRefCitation id="EFF04B02FFFFFF86FE83A0CBFE6DF86B" author="Abe" box="[262,389,2004,2029]" pageId="20" pageNumber="221" refString="Abe, H. (1996 a) Rhombognathine mites (Acari: Halacaridae) from Hokkaido, Northern Japan. Publications of the Seto Marine Biological Laboratory, 37, 63 - 166." type="journal article" year="1996" yearSuffix="a">Abé 1996a</bibRefCitation>
: fig. 23C, 1996b: fig. 2E). Larva with pair of epimeral pores (
<bibRefCitation id="EFF04B02FFFFFF86FBBDA0CBFB08F86B" author="Bartsch" box="[1080,1248,2004,2029]" pageId="20" pageNumber="221" refString="Bartsch, I. (1974 b) Uber das Auftreten von Epimeralporen besonders bei den Rhombognathinae (Halacaridae, Acari). Zoologischer Anzeiger, 193, 266 - 268." type="journal article" year="1974" yearSuffix="b">Bartsch 1974b</bibRefCitation>
: fig. 1a, 2c).
</paragraph>
</subSubSection>
<subSubSection id="C37B6578FFFEFF87FF42A788FEE0FDE6" pageId="21" pageNumber="222" type="discussion">
<paragraph id="8BDE36F3FFFEFF87FF42A788FEE0FDE6" blockId="21.[151,1437,151,608]" pageId="21" pageNumber="222">
<emphasis id="B915EAE1FFFEFF87FF42A788FED3FF36" bold="true" box="[199,315,151,176]" pageId="21" pageNumber="222">Remarks.</emphasis>
At present 29 species are known. The species live in a zone more or less regularly emerged; all feed on algae. The genus is spread world-wide but three different lineages have been separated on the basis of their character combinations (
<bibRefCitation id="EFF04B02FFFEFF87FE3CA7C0FDDBFF7E" author="Abe" box="[441,563,223,248]" pageId="21" pageNumber="222" refString="Abe, H. (2001) Phylogeny and character evolution of the marine genus Isobactrus (Acari: Halacaridae). Journal of Natural History, 35, 617 - 625. http: // dx. doi. org / 10.1080 / 00222930151098279" type="journal article" year="2001">Abé 2001</bibRefCitation>
;
<bibRefCitation id="EFF04B02FFFEFF87FDC3A7C0FD1BFF7E" author="Bartsch" box="[582,755,223,248]" pageId="21" pageNumber="222" refString="Bartsch, I. (2010 a) Reflections on distribution and origin of the halacarid genus Isobactrus (Acari: Halacaridae). Marine Biodiversity, 40, 17 - 32. http: // dx. doi. org / 10.1007 / s 12526 - 009 - 0030 - 7" type="journal article" year="2010" yearSuffix="a">Bartsch 2010a</bibRefCitation>
). The species of each lineage are concentrated within a geographical area which roughly can be attributed to the temperate northern Atlantic and Pacific Ocean (ca 75°N to 25°N), the tropics (ca 25°N to 25°S) and the temperate southern Pacific and Indian Ocean (ca 25°S to 55°S). The different number and shape of the genital acetabula are in concordance with these three groups. Adults and tritonymphs from the southern temperate zone have two pairs of acetabula, those from the tropics three pairs, and those from the northern temperate zone generally two pairs of large-sized and one pair of small acetabula. Exceptions are the North Atlantic species
<taxonomicName id="4C614D70FFFEFF87FD01A6A6FCF7FE56" box="[644,799,440,464]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="21" pageNumber="222" phylum="Arthropoda" rank="species" species="uniscutatus">
<emphasis id="B915EAE1FFFEFF87FD01A6A6FD65FE56" box="[644,653,441,464]" italics="true" pageId="21" pageNumber="222">I</emphasis>
.
<emphasis id="B915EAE1FFFEFF87FD1AA6A6FCF7FE56" box="[671,799,441,464]" italics="true" pageId="21" pageNumber="222">uniscutatus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C614D70FFFEFF87FCD9A6A6FC12FE56" box="[860,1018,439,464]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="21" pageNumber="222" phylum="Arthropoda" rank="species" species="hutchinsoni">
<emphasis id="B915EAE1FFFEFF87FCD9A6A6FC12FE56" box="[860,1018,439,464]" italics="true" pageId="21" pageNumber="222">I. hutchinsoni</emphasis>
</taxonomicName>
which have two pairs of acetabula; populations of both can survive in low saline brackish water. The deutonymphs of the southern temperate zone have a single pair of gac, those from the tropics two equal-sized pairs and species from the northern temperate zone a large and a minute pair of acetabula, again with the exception of
<taxonomicName id="4C614D70FFFEFF87FC35A53AFBA3FDBA" box="[944,1099,549,572]" class="Arachnida" family="Halacaridae" genus="Isobactrus" kingdom="Animalia" order="Trombidiformes" pageId="21" pageNumber="222" phylum="Arthropoda" rank="species" species="uniscutatus">
<emphasis id="B915EAE1FFFEFF87FC35A53AFC51FDBA" box="[944,953,549,572]" italics="true" pageId="21" pageNumber="222">I</emphasis>
.
<emphasis id="B915EAE1FFFEFF87FC4EA53AFBA3FDBA" box="[971,1099,549,572]" italics="true" pageId="21" pageNumber="222">uniscutatus</emphasis>
</taxonomicName>
which bears a single pair of acetabula.
</paragraph>
</subSubSection>
</treatment>
</document>