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<document id="2CC3688766F4D83654A16B4CF5BCEFC4" ID-ISSN="1175-5326" IM.materialsCitations_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1629618962334" checkinUser="felipe" docAuthor="Fend, En. V." docDate="2009" docId="F44687BC04376261FF70EA59FB8FFC74" docLanguage="en" docName="zt02077p030.pdf" docOrigin="Zootaxa 2077" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Kincaidiana hexatheca Altman 1936" docType="treatment" docVersion="2" lastPageNumber="6" masterDocId="087FFFC404356264FFE7EE0FFF82FFAA" masterDocTitle="An evaluation of the genus Kincaidiana Altman, 1936, with the designation of Altmanella n. gen. (Annelida, Clitellata, Lumbriculidae)" masterLastPageNumber="30" masterPageNumber="1" pageNumber="3" updateTime="1699084233931" updateUser="plazi" zenodo-license-document="CLOSED">
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<mods:title id="222D5014A24445245E43515BBA311ECB">An evaluation of the genus Kincaidiana Altman, 1936, with the designation of Altmanella n. gen. (Annelida, Clitellata, Lumbriculidae)</mods:title>
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<mods:namePart id="4513629795C9BC1991C09B81AD1D5602">Fend, En. V.</mods:namePart>
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<treatment id="F44687BC04376261FF70EA59FB8FFC74" LSID="urn:lsid:plazi:treatment:F44687BC04376261FF70EA59FB8FFC74" httpUri="http://treatment.plazi.org/id/F44687BC04376261FF70EA59FB8FFC74" lastPageId="5" lastPageNumber="6" pageId="2" pageNumber="3">
<subSubSection id="34F5652104376266FF70EA59FDDEFBDA" box="[151,604,1110,1136]" pageId="2" pageNumber="3" type="nomenclature">
<paragraph id="7C5036AA04376266FF70EA59FDDEFBDA" blockId="2.[151,604,1110,1171]" box="[151,604,1110,1136]" pageId="2" pageNumber="3">
<heading id="271881C604376266FF70EA59FDDEFBDA" bold="true" box="[151,604,1110,1136]" fontSize="11" level="1" pageId="2" pageNumber="3" reason="1">
<taxonomicName id="BBEF4D2904376266FF70EA59FDDEFBDA" ID-CoL="3R7V9" authority="Altman, 1936" authorityName="Altman" authorityYear="1936" box="[151,604,1110,1136]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="2" pageNumber="3" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804376266FF70EA59FDDEFBDA" bold="true" box="[151,604,1110,1136]" pageId="2" pageNumber="3">
<emphasis id="4E9BEAB804376266FF70EA59FE2CFBDA" bold="true" box="[151,430,1110,1136]" italics="true" pageId="2" pageNumber="3">Kincaidiana hexatheca</emphasis>
<bibRefCitation id="187E4B5B04376266FE52EA59FDDEFBDA" author="Altman" box="[437,604,1110,1136]" firstAuthor="Altman" pageId="2" pageNumber="3" pagination="1 - 137" refId="ref15043" refString="Altman L. C. (1936) Oligochaeta of Washington. University of Washington Publications in Biology, 4, 1 - 137." type="journal article" year="1936">Altman, 1936</bibRefCitation>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="34F5652104376266FF70EA76FEA0FB39" box="[151,290,1145,1171]" pageId="2" pageNumber="3" type="description">
<paragraph id="7C5036AA04376266FF70EA76FEA0FB39" blockId="2.[151,604,1110,1171]" box="[151,290,1145,1171]" pageId="2" pageNumber="3">
<figureCitation id="E4D42A2F04376266FF70EA76FEA0FB39" box="[151,290,1145,1171]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStartId-0="3.[151,256,1641,1665]" captionStartId-1="4.[151,257,1679,1703]" captionTargetBox-0="[157,1430,187,1610]" captionTargetBox-1="[154,1432,380,1658]" captionTargetId-0="figure-182@3.[151,1436,187,1613]" captionTargetId-1="figure-260@4.[151,1436,376,1664]" captionTargetPageId-0="3" captionTargetPageId-1="4" captionText-0="FIGURE 1. Kincaidiana hexatheca. AD from the Hoh River, Washington; E from Inglenook Fen, California. A, an unmounted specimen in ventral view, showing male and spermathecal pores. B, representative chaetae: from top to bottom, a ventral chaeta in II, a dorsal pair in VIII, a ventral in VI, a ventral in X, and a ventral from mid-body. C, lateral blood vessels in two posterior segments. D, nephridium. E, reproductive organs. Scale bars: A 2mm, B 200 µm, C 1 mm, D 200 µm, E 300 µm." captionText-1="FIGURE 2. Kincaidiana hexatheca. AC, F and IJ from the Hoh River, Washington; DE and GH from the type series, Washington. AB, whole worms, with A. freidris specimens (Owyhee River, Oregon) for size comparison. C, sagittal section in an anterior segment, showing transverse (circular) muscle layer of body wall. D, head of a dissected worm. E, spermatheca from a dissected worm. F, spermatheca from a sectioned worm, showing sperm lined up along ectal end of ampulla. GH, male pores and atria of a dissected and a sectioned worm. IJ, sections of proximal and distal parts of vas deferens; proximal to the atrium, the vas has a thick muscle coat. Scale bars: A 10 mm, B 5 mm, C 50 µm, EH 200 µm, IJ 50 µm." pageId="2" pageNumber="3">Figures 12</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="34F5652104376266FF70EAC9FD11F91A" pageId="2" pageNumber="3" type="materials_examined">
<paragraph id="7C5036AA04376266FF70EAC9FD46FA84" blockId="2.[151,1436,1222,1983]" pageId="2" pageNumber="3">
<emphasis id="4E9BEAB804376266FF70EAC9FE19FB4A" bold="true" box="[151,411,1222,1248]" pageId="2" pageNumber="3">Material examined:</emphasis>
<materialsCitation id="CC873CF704376266FE42EAC9FD42FA84" collectionCode="II, IV, V, VI, VII, VIII, IX, XII, XIII" collectorName="L. C. Altman" county="Pacific Co." location="Loomis Lake" pageId="2" pageNumber="3" specimenCount="1" stateProvince="Washington" typeStatus="holotype">
<emphasis id="4E9BEAB804376266FE42EAC9FDBAFB4A" bold="true" box="[421,568,1222,1248]" pageId="2" pageNumber="3">
<typeStatus id="A354880804376266FE42EAC9FE5FFB4A" box="[421,477,1222,1248]" pageId="2" pageNumber="3">type</typeStatus>
series.
</emphasis>
<collectingRegion id="BE2BF84804376266FDA5EAC9FD58FB4A" box="[578,730,1222,1248]" country="United States of America" name="Washington" pageId="2" pageNumber="3">Washington</collectingRegion>
:
<collectingCounty id="95314E2604376266FD00EAC9FCF4FB4A" box="[743,886,1222,1248]" pageId="2" pageNumber="3">Pacific Co.</collectingCounty>
:
<location id="7930607104376266FC63EAC9FBA9FB4A" LSID="urn:lsid:plazi:treatment:F44687BC04376261FF70EA59FB8FFC74:7930607104376266FC63EAC9FBA9FB4A" box="[900,1067,1222,1248]" county="Pacific Co." name="Loomis Lake" pageId="2" pageNumber="3" stateProvince="Washington">Loomis Lake</location>
, collected in 1931 or 1932, 6 dissected, slide mounted worms; sagittally sectioned worms
<collectionCode id="1AFEAE6F04376266FC84EAE2FCFBFAAD" box="[867,889,1261,1287]" pageId="2" pageNumber="3">II</collectionCode>
,
<collectionCode id="1AFEAE6F04376266FC60EAE2FC27FAAD" box="[903,933,1261,1287]" pageId="2" pageNumber="3">IV</collectionCode>
,
<collectionCode id="1AFEAE6F04376266FC56EAE2FC47FAAD" box="[945,965,1261,1287]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="2" pageNumber="3" type="Museum">V</collectionCode>
,
<collectionCode id="1AFEAE6F04376266FC35EAE2FC76FAAD" box="[978,1012,1261,1287]" country="Norway" name="Mykotektet, National Veterinary Institute" pageId="2" pageNumber="3">VI</collectionCode>
,
<collectionCode id="1AFEAE6F04376266FBE6EAE2FBAEFAAD" box="[1025,1068,1261,1287]" pageId="2" pageNumber="3">VII</collectionCode>
,
<collectionCode id="1AFEAE6F04376266FBDEEAE2FBECFAAD" box="[1081,1134,1261,1287]" pageId="2" pageNumber="3">VIII</collectionCode>
,
<collectionCode id="1AFEAE6F04376266FB9CEAE2FB1FFAAD" box="[1147,1181,1261,1287]" pageId="2" pageNumber="3">IX</collectionCode>
,
<collectionCode id="1AFEAE6F04376266FB4DEAE2FB54FAAD" box="[1194,1238,1261,1287]" pageId="2" pageNumber="3">XII</collectionCode>
,
<collectionCode id="1AFEAE6F04376266FB05EAE2FA9AFAAD" box="[1250,1304,1261,1287]" pageId="2" pageNumber="3">XIII</collectionCode>
; transverse sections X, XI, XIV. Collected by
<collectorName id="D11A537C04376266FDCEEB1BFD42FA84" box="[553,704,1300,1326]" pageId="2" pageNumber="3">L.C. Altman</collectorName>
</materialsCitation>
.
</paragraph>
<paragraph id="7C5036AA04376266FF21EB35FD11F91A" blockId="2.[151,1436,1222,1983]" pageId="2" pageNumber="3">
<emphasis id="4E9BEAB804376266FF21EB35FE08FAFE" bold="true" box="[198,394,1338,1364]" pageId="2" pageNumber="3">Other material.</emphasis>
<collectingRegion id="BE2BF84804376266FE75EB35FD8FFAFE" box="[402,525,1338,1364]" country="United States of America" name="California" pageId="2" pageNumber="3">California</collectingRegion>
: Mendocino Co.: Inglenook Fen at MacKerricher State Park near Fort Bragg, 1-Jul-05.
<collectingRegion id="BE2BF84804376266FEEDEB6EFEEAFAD1" box="[266,360,1377,1403]" country="United States of America" name="Oregon" pageId="2" pageNumber="3">Oregon</collectingRegion>
: Curry Co.: Rogue River at Quosatana Campground, 8-Jun-03. Douglas Co.: Cow Creek, 28-Apr-04. Unnamed cold spring, 7-Jun-03. Josephine Co.:
<taxonomicName id="BBEF4D2904376266FC63EB87FBA7FA0B" box="[900,1061,1416,1441]" class="Magnoliopsida" family="Sarraceniaceae" genus="Darlingtonia" kingdom="Plantae" order="Ericales" pageId="2" pageNumber="3" phylum="Tracheophyta" rank="genus">
<emphasis id="4E9BEAB804376266FC63EB87FBA7FA0B" box="[900,1061,1416,1441]" italics="true" pageId="2" pageNumber="3">Darlingtonia</emphasis>
</taxonomicName>
bog near O'Brien, 26-Oct-99.
<collectingRegion id="BE2BF84804376266FF70EBA1FF68FA62" box="[151,234,1454,1480]" country="United States of America" name="Illinois" pageId="2" pageNumber="3">Illinois</collectingRegion>
River near Sixmile Creek, 25-Oct-99. Lane Co.: Spring at mouth of Tenmile Creek, 1-May-99. Spring/ bog at Big Creek, 11-May-01. Marsh at base of Mt. Pisgah, near Eugene, 29-Jan-00. Multnomah Co.: Oneonta Creek, Columbia Gorge, 4-Jun-03. Tillamook Co.: small spring on Nestucca
<collectionCode id="1AFEAE6F04376266FBB8EBF3FBF5F9BC" box="[1119,1143,1532,1558]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="2" pageNumber="3">R</collectionCode>
. near Blaine, 4-Jun-03. Yamhill Co: Seep on east side of McGuire Reservoir near McMinnville, 4-Jun-03.
<collectingRegion id="BE2BF84804376266FB8DE82DFB78F996" box="[1130,1274,1570,1596]" country="United States of America" name="Washington" pageId="2" pageNumber="3">Washington</collectingRegion>
: Clallam Co.: pool along Bogachiel River on Undie Road, 26-Apr-04. Jefferson Co.: seeps along Hoh River Road, 29-Apr- 99. 26-Apr-04. Roadside ditch on Clearwater Road, near Hoh River, 3-Jun-03. Pacific Co.: Spring on Naselle River, 30-Apr-99. All collected by S. Fend.
</paragraph>
</subSubSection>
<subSubSection id="34F5652104376261FF21E8B2FB8FFC74" lastPageId="5" lastPageNumber="6" pageId="2" pageNumber="3" type="discussion">
<paragraph id="7C5036AA04376266FF21E8B2FCDCF815" blockId="2.[151,1436,1222,1983]" pageId="2" pageNumber="3">
<emphasis id="4E9BEAB804376266FF21E8B2FEBDF97D" bold="true" box="[198,319,1725,1751]" pageId="2" pageNumber="3">Remarks.</emphasis>
The emended diagnosis limits the genus to worms resembling
<taxonomicName id="BBEF4D2904376266FBDBE8B2FB55F97C" authorityName="Altman" authorityYear="1936" box="[1084,1239,1725,1750]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="2" pageNumber="3" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804376266FBDBE8B2FB55F97C" box="[1084,1239,1725,1750]" italics="true" pageId="2" pageNumber="3">K. hexatheca</emphasis>
</taxonomicName>
and its possible undescribed relatives, rather than the broadly inclusive definition given by
<bibRefCitation id="187E4B5B04376266FBB6E8EBFB76F954" author="Cook" box="[1105,1268,1764,1790]" firstAuthor="Cook" pageId="2" pageNumber="3" pagination="200 - 285" refId="ref15259" refString="Cook, D. G. (1971) Family Lumbriculidae. In: Brinkhurst R. O. &amp; Jamieson B. G. M. Aquatic Oligochaeta of the World. Oliver and Boyd, Edinburgh, pp. 200 - 285." type="book chapter" year="1971">Cook (1971)</bibRefCitation>
. The original description of
<taxonomicName id="BBEF4D2904376266FED8E904FE5AF88E" authorityName="Altman" authorityYear="1936" box="[319,472,1803,1828]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="2" pageNumber="3" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804376266FED8E904FE5AF88E" box="[319,472,1803,1828]" italics="true" pageId="2" pageNumber="3">K. hexatheca</emphasis>
</taxonomicName>
is relatively complete, and new collections from additional sites confirm
<bibRefCitation id="187E4B5B04376266FAD7E905FF64F8E1" author="Altman's" firstAuthor="Altman's" pageId="2" pageNumber="3" pagination="1 - 137" refId="ref15043" refString="Altman L. C. (1936) Oligochaeta of Washington. University of Washington Publications in Biology, 4, 1 - 137." type="journal article" year="1936">Altmans (1936)</bibRefCitation>
observations, as well as the above diagnosis. The combination of a very elongate proboscis, unusual anterior chaetae, a forward shift of reproductive organs by one segment (see
<bibRefCitation id="187E4B5B04376266FBF8E957FB76F8D8" author="Brinkhurst" box="[1055,1268,1880,1906]" firstAuthor="Brinkhurst" pageId="2" pageNumber="3" pagination="2731 - 2739" refId="ref15068" refString="Brinkhurst R. O. (1989) A phylogenetic analysis of the Lumbriculidae (Annelida, Oligochaeta). Canadian Journal of Zoology, 67, 2731 - 2739." type="journal article" year="1989">Brinkhurst [1989]</bibRefCitation>
), prosoporous male ducts, and the hexathecate condition with the first pair of spermathecae in the atrial segment easily distinguishes
<taxonomicName id="BBEF4D2904376266FED0E9AAFE4AF814" authorityName="Altman" authorityYear="1936" box="[311,456,1957,1982]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="2" pageNumber="3" phylum="Annelida" rank="genus">
<emphasis id="4E9BEAB804376266FED0E9AAFE4AF814" box="[311,456,1957,1982]" italics="true" pageId="2" pageNumber="3">Kincaidiana</emphasis>
</taxonomicName>
from all other lumbriculid genera.
</paragraph>
<caption id="2890662204366267FF70E866FEFDF8A1" pageId="3" pageNumber="4" startId="3.[151,256,1641,1665]" targetBox="[157,1430,187,1610]" targetPageId="3">
<paragraph id="7C5036AA04366267FF70E866FEFDF8A1" blockId="3.[151,1436,1641,1803]" pageId="3" pageNumber="4">
<emphasis id="4E9BEAB804366267FF70E866FE9EF92B" bold="true" box="[151,284,1641,1665]" pageId="3" pageNumber="4">FIGURE 1.</emphasis>
<taxonomicName id="BBEF4D2904366267FEC3E865FDA2F92B" authorityName="Altman" authorityYear="1936" box="[292,544,1642,1665]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="3" pageNumber="4" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804366267FEC3E865FDA2F92B" box="[292,544,1642,1665]" italics="true" pageId="3" pageNumber="4">Kincaidiana hexatheca</emphasis>
</taxonomicName>
. AD from the Hoh River, Washington; E from Inglenook Fen, California. A, an unmounted specimen in ventral view, showing male and spermathecal pores. B, representative chaetae: from top to bottom, a ventral chaeta in II, a dorsal pair in VIII, a ventral in VI, a ventral in X, and a ventral from mid-body. C, lateral blood vessels in two posterior segments. D, nephridium. E, reproductive organs. Scale bars: A 2mm, B 200 µm, C 1 mm, D 200 µm, E 300 µm.
</paragraph>
</caption>
<paragraph id="7C5036AA04366260FF21E932FAB8FEE6" blockId="3.[151,1436,1853,2034]" lastBlockId="4.[151,1436,152,332]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
Additional reproductive characters distinguishing
<taxonomicName id="BBEF4D2904366267FCD3E931FC47F8FD" authorityName="Altman" authorityYear="1936" box="[820,965,1854,1879]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="3" pageNumber="4" phylum="Annelida" rank="genus">
<emphasis id="4E9BEAB804366267FCD3E931FC47F8FD" box="[820,965,1854,1879]" italics="true" pageId="3" pageNumber="4">Kincaidiana</emphasis>
</taxonomicName>
include the penial and spermathecal morphology, the posterior location of the male funnels, and vasa deferentia with a partial muscle coat. The short, pendant penes are within eversible sacs, and these in turn are subtended by conical, possibly telescoping porophores (
<figureCitation id="E4D42A2F04366267FECCE9BEFE2DF861" box="[299,431,1969,1995]" captionStart="FIGURE 2" captionStartId="4.[151,257,1679,1703]" captionTargetBox="[154,1432,380,1658]" captionTargetId="figure-260@4.[151,1436,376,1664]" captionTargetPageId="4" captionText="FIGURE 2. Kincaidiana hexatheca. AC, F and IJ from the Hoh River, Washington; DE and GH from the type series, Washington. AB, whole worms, with A. freidris specimens (Owyhee River, Oregon) for size comparison. C, sagittal section in an anterior segment, showing transverse (circular) muscle layer of body wall. D, head of a dissected worm. E, spermatheca from a dissected worm. F, spermatheca from a sectioned worm, showing sperm lined up along ectal end of ampulla. GH, male pores and atria of a dissected and a sectioned worm. IJ, sections of proximal and distal parts of vas deferens; proximal to the atrium, the vas has a thick muscle coat. Scale bars: A 10 mm, B 5 mm, C 50 µm, EH 200 µm, IJ 50 µm." pageId="3" pageNumber="4">Fig. 2GH</figureCitation>
). The spermathecal ampullae are unusually elongate, with most of the sperm lined up along the epithelium, in an expansion near the ectal duct (
<figureCitation id="E4D42A2F04366267FCDBE9D7FC22F858" box="[828,928,2008,2034]" captionStart="FIGURE 1" captionStartId="3.[151,256,1641,1665]" captionTargetBox="[157,1430,187,1610]" captionTargetId="figure-182@3.[151,1436,187,1613]" captionTargetPageId="3" captionText="FIGURE 1. Kincaidiana hexatheca. AD from the Hoh River, Washington; E from Inglenook Fen, California. A, an unmounted specimen in ventral view, showing male and spermathecal pores. B, representative chaetae: from top to bottom, a ventral chaeta in II, a dorsal pair in VIII, a ventral in VI, a ventral in X, and a ventral from mid-body. C, lateral blood vessels in two posterior segments. D, nephridium. E, reproductive organs. Scale bars: A 2mm, B 200 µm, C 1 mm, D 200 µm, E 300 µm." pageId="3" pageNumber="4">Figs. 1E</figureCitation>
,
<figureCitation id="E4D42A2F04366267FC4AE9D7FC70F858" box="[941,1010,2008,2034]" captionStart="FIGURE 2" captionStartId="4.[151,257,1679,1703]" captionTargetBox="[154,1432,380,1658]" captionTargetId="figure-260@4.[151,1436,376,1664]" captionTargetPageId="4" captionText="FIGURE 2. Kincaidiana hexatheca. AC, F and IJ from the Hoh River, Washington; DE and GH from the type series, Washington. AB, whole worms, with A. freidris specimens (Owyhee River, Oregon) for size comparison. C, sagittal section in an anterior segment, showing transverse (circular) muscle layer of body wall. D, head of a dissected worm. E, spermatheca from a dissected worm. F, spermatheca from a sectioned worm, showing sperm lined up along ectal end of ampulla. GH, male pores and atria of a dissected and a sectioned worm. IJ, sections of proximal and distal parts of vas deferens; proximal to the atrium, the vas has a thick muscle coat. Scale bars: A 10 mm, B 5 mm, C 50 µm, EH 200 µm, IJ 50 µm." pageId="3" pageNumber="4">2EF</figureCitation>
); the large, thick spermathecal ducts are also unusual. The muscle layer extending onto the proximal part of the vas deferens (
<figureCitation id="E4D42A2F04316260FB59EE97FA8DFF18" box="[1214,1295,152,178]" captionStart="FIGURE 2" captionStartId="4.[151,257,1679,1703]" captionTargetBox="[154,1432,380,1658]" captionTargetId="figure-260@4.[151,1436,376,1664]" captionTargetPageId="4" captionText="FIGURE 2. Kincaidiana hexatheca. AC, F and IJ from the Hoh River, Washington; DE and GH from the type series, Washington. AB, whole worms, with A. freidris specimens (Owyhee River, Oregon) for size comparison. C, sagittal section in an anterior segment, showing transverse (circular) muscle layer of body wall. D, head of a dissected worm. E, spermatheca from a dissected worm. F, spermatheca from a sectioned worm, showing sperm lined up along ectal end of ampulla. GH, male pores and atria of a dissected and a sectioned worm. IJ, sections of proximal and distal parts of vas deferens; proximal to the atrium, the vas has a thick muscle coat. Scale bars: A 10 mm, B 5 mm, C 50 µm, EH 200 µm, IJ 50 µm." pageId="4" pageNumber="5">Fig. 2I</figureCitation>
vs. 2J) was noted by
<bibRefCitation id="187E4B5B04316260FEEAEEB1FE2CFF72" author="Cook" box="[269,430,190,216]" firstAuthor="Cook" pageId="4" pageNumber="5" pagination="200 - 285" refId="ref15259" refString="Cook, D. G. (1971) Family Lumbriculidae. In: Brinkhurst R. O. &amp; Jamieson B. G. M. Aquatic Oligochaeta of the World. Oliver and Boyd, Edinburgh, pp. 200 - 285." type="book chapter" year="1971">Cook (1971)</bibRefCitation>
; this character may have been overlooked in other lumbriculid descriptions, but otherwise has only been noted in
<taxonomicName id="BBEF4D2904316260FDC0EEEAFC1EFF55" authority="Fend &amp; Lenat. The" authorityName="Fend &amp; Lenat. The" authorityYear="2007" box="[551,924,229,255]" class="Clitellata" family="Lumbriculidae" genus="Pilaridrilus" kingdom="Animalia" order="Lumbriculida" pageId="4" pageNumber="5" phylum="Annelida" rank="genus">
<emphasis id="4E9BEAB804316260FDC0EEEAFD33FF54" box="[551,689,229,254]" italics="true" pageId="4" pageNumber="5">Pilaridrilus</emphasis>
Fend &amp; Lenat. The
</taxonomicName>
(posterior) male funnels fold back into the sperm sacs in many lumbriculids, but the posterior displacement by one or more segments (
<figureCitation id="E4D42A2F04316260FAF1EF03FAF6FE8C" box="[1302,1396,268,294]" captionStart="FIGURE 1" captionStartId="3.[151,256,1641,1665]" captionTargetBox="[157,1430,187,1610]" captionTargetId="figure-182@3.[151,1436,187,1613]" captionTargetPageId="3" captionText="FIGURE 1. Kincaidiana hexatheca. AD from the Hoh River, Washington; E from Inglenook Fen, California. A, an unmounted specimen in ventral view, showing male and spermathecal pores. B, representative chaetae: from top to bottom, a ventral chaeta in II, a dorsal pair in VIII, a ventral in VI, a ventral in X, and a ventral from mid-body. C, lateral blood vessels in two posterior segments. D, nephridium. E, reproductive organs. Scale bars: A 2mm, B 200 µm, C 1 mm, D 200 µm, E 300 µm." pageId="4" pageNumber="5">Fig. 1E</figureCitation>
) is unusual; it has, however, been illustrated for the Palearctic
<taxonomicName id="BBEF4D2904316260FCABEF3DFAB4FEE6" authority="(Michaelsen, 1933)" baseAuthorityName="Michaelsen" baseAuthorityYear="1933" box="[844,1334,306,332]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" kingdom="Animalia" order="Lumbriculida" pageId="4" pageNumber="5" phylum="Annelida" rank="species" species="wiardi">
<emphasis id="4E9BEAB804316260FCABEF3DFBC5FEE1" box="[844,1095,306,331]" italics="true" pageId="4" pageNumber="5">Guestphalinus wiardi</emphasis>
(
<bibRefCitation id="187E4B5B04316260FBB0EF3DFAAFFEE6" author="Michaelsen" box="[1111,1325,306,332]" firstAuthor="Michaelsen" pageId="4" pageNumber="5" pagination="1 - 19" refId="ref15621" refString="Michaelsen, W. (1933) Uber Hohlen Oligochaten. Mitteilungen Hohlen-Universitat Karstforsch, Berlin, 1, 1 - 19." type="journal article" year="1933">Michaelsen, 1933</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<caption id="2890662204316260FF70E880FE22F8DC" pageId="4" pageNumber="5" startId="4.[151,257,1679,1703]" targetBox="[154,1432,380,1658]" targetPageId="4">
<paragraph id="7C5036AA04316260FF70E880FE22F8DC" blockId="4.[151,1436,1679,1911]" pageId="4" pageNumber="5">
<emphasis id="4E9BEAB804316260FF70E880FE9EF90D" bold="true" box="[151,284,1679,1703]" pageId="4" pageNumber="5">FIGURE 2.</emphasis>
<taxonomicName id="BBEF4D2904316260FEC2E89FFDA6F90D" authorityName="Altman" authorityYear="1936" box="[293,548,1680,1703]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="4" pageNumber="5" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804316260FEC2E89FFDA6F90D" box="[293,548,1680,1703]" italics="true" pageId="4" pageNumber="5">Kincaidiana hexatheca</emphasis>
</taxonomicName>
. AC, F and IJ from the Hoh River, Washington; DE and GH from the type series, Washington. AB, whole worms, with
<taxonomicName id="BBEF4D2904316260FD75E8BDFC83F963" baseAuthorityName="Cook" baseAuthorityYear="1966" box="[658,769,1714,1737]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="4" pageNumber="5" phylum="Annelida" rank="species" species="freidris">
<emphasis id="4E9BEAB804316260FD75E8BDFC83F963" box="[658,769,1714,1737]" italics="true" pageId="4" pageNumber="5">A. freidris</emphasis>
</taxonomicName>
specimens (Owyhee River, Oregon) for size comparison. C, sagittal section in an anterior segment, showing transverse (circular) muscle layer of body wall. D, head of a dissected worm. E, spermatheca from a dissected worm. F, spermatheca from a sectioned worm, showing sperm lined up along ectal end of ampulla. GH, male pores and atria of a dissected and a sectioned worm. IJ, sections of proximal and distal parts of vas deferens; proximal to the atrium, the vas has a thick muscle coat. Scale bars: A 10 mm, B 5 mm, C 50 µm, EH 200 µm, IJ 50 µm.
</paragraph>
</caption>
<paragraph id="7C5036AA04316261FF22E9A6FE7CFCB6" blockId="4.[151,1436,1961,2025]" lastBlockId="5.[151,1436,152,990]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
Internal somatic characters have generally not been emphasized in lumbriculid descriptions. Nephridia are difficult to observe, and thus poorly described in most lumbriculids; however,
<taxonomicName id="BBEF4D2904316260FBCBE9DFFB46F843" authorityName="Altman" authorityYear="1936" box="[1068,1220,2000,2025]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="4" pageNumber="5" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804316260FBCBE9DFFB46F843" box="[1068,1220,2000,2025]" italics="true" pageId="4" pageNumber="5">K. hexatheca</emphasis>
</taxonomicName>
nephridia have an unusual form and vertical orientation. The long, narrow postseptal expansion and duct typically follow a lateral blood vessel dorsally, similar to nephridia of
<taxonomicName id="BBEF4D2904306261FCE5EEB0FBADFF72" authority="Hoffmeister" authorityName="Hoffmeister" authorityYear="1843" box="[770,1071,190,216]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" kingdom="Animalia" order="Lumbriculida" pageId="5" pageNumber="6" phylum="Annelida" rank="genus">
<emphasis id="4E9BEAB804306261FCE5EEB0FC1AFF72" box="[770,920,191,216]" italics="true" pageId="5" pageNumber="6">Rhynchelmis</emphasis>
Hoffmeister
</taxonomicName>
(cf. Fend &amp;
<bibRefCitation id="187E4B5B04306261FB2EEEB1FA0CFF72" author="Brinkhurst" box="[1225,1422,190,216]" firstAuthor="Brinkhurst" pageId="5" pageNumber="6" pagination="2731 - 2739" refId="ref15068" refString="Brinkhurst R. O. (1989) A phylogenetic analysis of the Lumbriculidae (Annelida, Oligochaeta). Canadian Journal of Zoology, 67, 2731 - 2739." type="journal article" year="1989">Brinkhurst 1989</bibRefCitation>
); the dorsal loop then usually extends posteriorad (ventromedially) before looping anteriad to the duct and nephropore (
<figureCitation id="E4D42A2F04306261FEC8EF03FE0CFE8C" box="[303,398,268,294]" captionStart="FIGURE 1" captionStartId="3.[151,256,1641,1665]" captionTargetBox="[157,1430,187,1610]" captionTargetId="figure-182@3.[151,1436,187,1613]" captionTargetPageId="3" captionText="FIGURE 1. Kincaidiana hexatheca. AD from the Hoh River, Washington; E from Inglenook Fen, California. A, an unmounted specimen in ventral view, showing male and spermathecal pores. B, representative chaetae: from top to bottom, a ventral chaeta in II, a dorsal pair in VIII, a ventral in VI, a ventral in X, and a ventral from mid-body. C, lateral blood vessels in two posterior segments. D, nephridium. E, reproductive organs. Scale bars: A 2mm, B 200 µm, C 1 mm, D 200 µm, E 300 µm." pageId="5" pageNumber="6">Fig. 1D</figureCitation>
). Pre-clitellar nephridia were not observed in
<taxonomicName id="BBEF4D2904306261FC51EF03FBD0FE8F" authorityName="Altman" authorityYear="1936" box="[950,1106,268,293]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="5" pageNumber="6" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804306261FC51EF03FBD0FE8F" box="[950,1106,268,293]" italics="true" pageId="5" pageNumber="6">K. hexatheca</emphasis>
</taxonomicName>
; instead, the first nephridia are on the first septum behind the female pores. This appears to be the most common arrangement in the
<taxonomicName id="BBEF4D2904306261FF70EF56FEC5FED9" box="[151,327,345,371]" class="Clitellata" family="Lumbriculidae" kingdom="Animalia" order="Lumbriculida" pageId="5" pageNumber="6" phylum="Annelida" rank="family">Lumbriculidae</taxonomicName>
, although due to the anterior shift in reproductive organs (
<emphasis id="4E9BEAB804306261FC10EF56FBBAFED8" box="[1015,1080,345,370]" italics="true" pageId="5" pageNumber="6">sensu</emphasis>
<bibRefCitation id="187E4B5B04306261FBA7EF56FA81FED9" author="Brinkhurst" box="[1088,1283,345,371]" firstAuthor="Brinkhurst" pageId="5" pageNumber="6" pagination="2731 - 2739" refId="ref15068" refString="Brinkhurst R. O. (1989) A phylogenetic analysis of the Lumbriculidae (Annelida, Oligochaeta). Canadian Journal of Zoology, 67, 2731 - 2739." type="journal article" year="1989">Brinkhurst 1989</bibRefCitation>
), they are on 11/12, rather than the usual 12/13. Highly branched blood vessels in posterior segments are not unusual in the larger lumbriculids, although the pattern may be distinctive at the genus or species level. Details of the vascular system are difficult to see in many preparations, but the new
<taxonomicName id="BBEF4D2904306261FC2DEFC2FBE1FE4C" authorityName="Altman" authorityYear="1936" box="[970,1123,461,486]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="5" pageNumber="6" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804306261FC2DEFC2FBE1FE4C" box="[970,1123,461,486]" italics="true" pageId="5" pageNumber="6">K. hexatheca</emphasis>
</taxonomicName>
material differs somewhat from the original description. In the new material, branched lateral vessels begin after about XX; one or two may be present at the dorsal vessel, with the anterior larger, and a branch terminates in the ventral vessel and/ or (more commonly) in the perivisceral sinus (
<figureCitation id="E4D42A2F04306261FD2FEC4EFCA4FDF1" box="[712,806,577,603]" captionStart="FIGURE 1" captionStartId="3.[151,256,1641,1665]" captionTargetBox="[157,1430,187,1610]" captionTargetId="figure-182@3.[151,1436,187,1613]" captionTargetPageId="3" captionText="FIGURE 1. Kincaidiana hexatheca. AD from the Hoh River, Washington; E from Inglenook Fen, California. A, an unmounted specimen in ventral view, showing male and spermathecal pores. B, representative chaetae: from top to bottom, a ventral chaeta in II, a dorsal pair in VIII, a ventral in VI, a ventral in X, and a ventral from mid-body. C, lateral blood vessels in two posterior segments. D, nephridium. E, reproductive organs. Scale bars: A 2mm, B 200 µm, C 1 mm, D 200 µm, E 300 µm." pageId="5" pageNumber="6">Fig. 1C</figureCitation>
). This arrangement differs slightly from other large, proboscis-bearing, Nearctic lumbriculids such as
<taxonomicName id="BBEF4D2904306261FD12EC67FB2CFD28" authority="Wassell" authorityName="Wassell" box="[757,1198,616,642]" class="Clitellata" family="Lumbriculidae" genus="Eclipidrilus" kingdom="Animalia" order="Lumbriculida" pageId="5" pageNumber="6" phylum="Annelida" rank="subGenus" subGenus="Premnodrilus">
<emphasis id="4E9BEAB804306261FD12EC67FC06FD2B" box="[757,900,616,641]" italics="true" pageId="5" pageNumber="6">Eclipidrilus</emphasis>
(
<emphasis id="4E9BEAB804306261FC7FEC67FBBFFD2B" box="[920,1085,616,641]" italics="true" pageId="5" pageNumber="6">Premnodrilus</emphasis>
) Wassell
</taxonomicName>
or
<taxonomicName id="BBEF4D2904306261FB3EEC67FAF1FD2B" authorityName="Hoffmeister" authorityYear="1843" box="[1241,1395,616,641]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" kingdom="Animalia" order="Lumbriculida" pageId="5" pageNumber="6" phylum="Annelida" rank="genus">
<emphasis id="4E9BEAB804306261FB3EEC67FAF1FD2B" box="[1241,1395,616,641]" italics="true" pageId="5" pageNumber="6">Rhynchelmis</emphasis>
</taxonomicName>
, in which branched lateral vessels (if present) are two pairs per segment, beginning near X (
<bibRefCitation id="187E4B5B04306261FB55EC81FAB7FD02" author="Fend" box="[1202,1333,654,680]" firstAuthor="Fend" pageId="5" pageNumber="6" pagination="1 - 42" refId="ref15339" refString="Fend, S. V. (2005) A review of the genus Eclipidrilus (Annelida: Clitellata: Lumbriculidae), with description of a new species from western North America. Zootaxa, 969, 1 - 42." type="journal article" year="2005">Fend 2005</bibRefCitation>
,
<bibRefCitation id="187E4B5B04306261FAA5EC81FEE5FD65" author="Fend &amp; Brinkhurst" firstAuthor="Fend" pageId="5" pageNumber="6" pagination="1 - 59" refId="ref15381" refString="Fend, S. V. &amp; Brinkhurst, R. O. (2000) New species of Rhynchelmis (Clitellata, Lumbriculidae), with observations on the Nearctic species. Hydrobiologia, 428, 1 - 59." type="journal article" year="2000">Fend &amp; Brinkhurst 2000</bibRefCitation>
). Body wall musculature in
<taxonomicName id="BBEF4D2904306261FD2EECBAFCE9FD64" authorityName="Altman" authorityYear="1936" box="[713,875,693,718]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="5" pageNumber="6" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804306261FD2EECBAFCE9FD64" box="[713,875,693,718]" italics="true" pageId="5" pageNumber="6">K. hexatheca</emphasis>
</taxonomicName>
may also be unusual, pending more detailed descriptions of other genera: the circular muscle fibers are arranged in regular, transverse arches or cylinders in anterior segments (
<figureCitation id="E4D42A2F04306261FE72ED0DFE73FCB6" box="[405,497,770,796]" captionStart="FIGURE 2" captionStartId="4.[151,257,1679,1703]" captionTargetBox="[154,1432,380,1658]" captionTargetId="figure-260@4.[151,1436,376,1664]" captionTargetPageId="4" captionText="FIGURE 2. Kincaidiana hexatheca. AC, F and IJ from the Hoh River, Washington; DE and GH from the type series, Washington. AB, whole worms, with A. freidris specimens (Owyhee River, Oregon) for size comparison. C, sagittal section in an anterior segment, showing transverse (circular) muscle layer of body wall. D, head of a dissected worm. E, spermatheca from a dissected worm. F, spermatheca from a sectioned worm, showing sperm lined up along ectal end of ampulla. GH, male pores and atria of a dissected and a sectioned worm. IJ, sections of proximal and distal parts of vas deferens; proximal to the atrium, the vas has a thick muscle coat. Scale bars: A 10 mm, B 5 mm, C 50 µm, EH 200 µm, IJ 50 µm." pageId="5" pageNumber="6">Fig. 2C</figureCitation>
).
</paragraph>
<paragraph id="7C5036AA04306261FF22ED26FB8FFC74" blockId="5.[151,1436,152,990]" pageId="5" pageNumber="6">
Many of these characters are likely to be apomorphic according to the analysis of
<bibRefCitation id="187E4B5B04306261FB68ED26FAE7FCE9" author="Brinkhurst" box="[1167,1381,809,835]" firstAuthor="Brinkhurst" pageId="5" pageNumber="6" pagination="2731 - 2739" refId="ref15068" refString="Brinkhurst R. O. (1989) A phylogenetic analysis of the Lumbriculidae (Annelida, Oligochaeta). Canadian Journal of Zoology, 67, 2731 - 2739." type="journal article" year="1989">Brinkhurst (1989)</bibRefCitation>
, and some are unusual (although not unique) within the family. Although morphological details vary both within and among populations, in most cases there do not appear to be regional morphotypes that would indicate additional species. Additional details on
<taxonomicName id="BBEF4D2904306261FD6EED92FCA5FC1C" authorityName="Altman" authorityYear="1936" box="[649,807,925,950]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" kingdom="Animalia" order="Lumbriculida" pageId="5" pageNumber="6" phylum="Annelida" rank="species" species="hexatheca">
<emphasis id="4E9BEAB804306261FD6EED92FCA5FC1C" box="[649,807,925,950]" italics="true" pageId="5" pageNumber="6">K. hexatheca</emphasis>
</taxonomicName>
morphology and population differences, and a new species from northern
<collectingRegion id="BE2BF84804306261FE7AEDCBFD96FC74" box="[413,532,964,990]" country="United States of America" name="California" pageId="5" pageNumber="6">California</collectingRegion>
will be reported in a separate contribution.
</paragraph>
</subSubSection>
</treatment>
</document>