treatments-xml/data/03/AE/62/03AE6204FFE5F928FEB5FEFCFCF9C127.xml

<document id="8DF0BF3B77E18834693B9551143A7078" ID-CLB-Dataset="51249" ID-DOI="10.5281/zenodo.156662" ID-GBIF-Dataset="5900719a-6bf2-4955-9f84-3e84e1a7ab1b" ID-ISSN="1175­5326" ID-Zenodo-Dep="156662" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1459637569271" checkinUser="plazi" docAuthor="Leonard, William P., Chichester, Lyle, Baugh, Jim &amp; Wilke, Thomas" docDate="2003" docId="03AE6204FFE5F928FEB5FEFCFCF9C127" docLanguage="en" docName="zt00355.pdf" docOrigin="Zootaxa 355" docStyle="DocumentStyle:FA7E419B012A62B0FC3AC15A186C3DAF.3:Zootaxa.2001-2006.journal_article" docStyleId="FA7E419B012A62B0FC3AC15A186C3DAF" docStyleName="Zootaxa.2001-2006.journal_article" docStyleVersion="3" docTitle="Kootenaia burkei Leonard, Chichester, Baugh &amp; Wilke, 2003, new species" docType="treatment" docVersion="11" lastPageNumber="12" masterDocId="FF971A7CFFE3F923FFBDFFE0FFC1C642" masterDocTitle="Kootenaia burkei, a new genus and species of slug from northern Idaho, United States (Gastropoda: Pulmonata: Arionidae)" masterLastPageNumber="16" masterPageNumber="1" pageNumber="7" updateTime="1698205722074" updateUser="ExternalLinkService">
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<mods:title id="16E4A048A22B5B16E85D91764BBC3A83">Kootenaia burkei, a new genus and species of slug from northern Idaho, United States (Gastropoda: Pulmonata: Arionidae)</mods:title>
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<mods:namePart id="58FD1D9C14D8C5EF4F232D840799755A">Leonard, William P.</mods:namePart>
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<mods:namePart id="BB2BF59B940C9EA1296768165402F306">Chichester, Lyle</mods:namePart>
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<mods:namePart id="67DAD673239750C53E6274B24B31538B">Wilke, Thomas</mods:namePart>
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<paragraph id="8BB8D312FFE5F925FEB5FEFCFDB0C777" blockId="6.[264,625,283,345]" box="[264,625,283,310]" pageId="6" pageNumber="7">
<heading id="D0F0647EFFE5F925FEB5FEFCFDB0C777" bold="true" box="[264,625,283,310]" fontSize="11" level="1" pageId="6" pageNumber="7" reason="1">
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<taxonomicName id="4C07A891FFE5F925FEB5FEFCFE18C774" ID-CoL="7RSHL" box="[264,473,284,310]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="7" phylum="Mollusca" rank="species" species="burkei" status="sp. nov.">
<emphasis id="B9730F00FFE5F925FEB5FEFCFE18C774" bold="true" box="[264,473,284,310]" italics="true" pageId="6" pageNumber="7">Kootenaia burkei</emphasis>
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<figureCitation id="133CCF97FFE5F925FEB5FEDFFE99C71B" box="[264,344,319,345]" captionStart="FIGURE 1" captionStartId="2.[264,367,1455,1479]" captionTargetBox="[267,1319,315,1431]" captionTargetId="figure@2.[267,1320,315,1431]" captionTargetPageId="2" captionText="FIGURE 1: Relief map of Idaho showing the localities of Kootenaia burkei new species. 1) Little Bumblebee Creek, Shoshone County (type locality), 2) Trestle Creek, Bonner County, 3) Beauty Creek, Kootenai County, 4) Heyburn State Park, Benewah County, and 5) East side of Harrison, Kootenai County. For detailed locality information see the Appendix. The lower photograph shows paratype CM 66247." httpUri="https://zenodo.org/record/168834/files/figure.png" pageId="6" pageNumber="7">Figs. 1</figureCitation>
, 
<figureCitation id="133CCF97FFE5F925FEDBFEDFFEB4C71B" box="[358,373,319,345]" captionStart="FIGURE 2" captionStartId="7.[264,367,1444,1468]" captionTargetBox="[278,1308,435,1428]" captionTargetId="figure@7.[278,1309,429,1437]" captionTargetPageId="7" captionText="FIGURE 2. Dissected holotype of Kootenaia burkei from Shoshone County, Idaho, showing the reproductive and digestive systems. Dorsal view of reproductive system on the left and ventral view of digestive system on the right. Total body length in alcohol = 10 mm." httpUri="https://zenodo.org/record/156664/files/figure.png" pageId="6" pageNumber="7">2</figureCitation>
</paragraph>
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<paragraph id="8BB8D312FFE5F925FEB5FE6FFE2DC483" blockId="6.[264,1325,399,1105]" pageId="6" pageNumber="7">
<emphasis id="B9730F00FFE5F925FEB5FE6FFE93C7EA" box="[264,338,399,424]" italics="true" pageId="6" pageNumber="7">
<typeStatus id="54BC6DB0FFE5F925FEB5FE6FFE89C7EA" box="[264,328,399,424]" pageId="6" pageNumber="7">Types</typeStatus>
:
</emphasis>
The 
<typeStatus id="54BC6DB0FFE5F925FE29FE6FFE3BC7EB" box="[404,506,399,425]" pageId="6" pageNumber="7" type="holotype">holotype</typeStatus>
(Carnegie Museum [CM] 66404), collected on 
<date id="FFB9F5D2FFE5F925FB8BFE6FFB32C7EB" box="[1078,1267,399,425]" pageId="6" pageNumber="7" value="2002-10-03">3 October 2002</date>
, and one 
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(CM 66247), collected on 
<date id="FFB9F5D2FFE5F925FD53FE57FC5AC793" box="[750,923,439,465]" pageId="6" pageNumber="7" value="2003-04-10">10 April 2003</date>
, were taken by J. Baugh and W. Leonard on woody debris and among black cottonwood (
<taxonomicName id="4C07A891FFE5F925FC00FE3FFE4FC463" authority="Torrey &amp; Gray, 1852" authorityName="Torrey &amp; Gray" authorityYear="1852" class="Magnoliopsida" family="Salicaceae" genus="Populus" kingdom="Plantae" order="Malpighiales" pageId="6" pageNumber="7" phylum="Tracheophyta" rank="species" species="trichocarpa">
<emphasis id="B9730F00FFE5F925FC00FE3FFB70C7BA" box="[957,1201,479,504]" italics="true" pageId="6" pageNumber="7">Populus trichocarpa</emphasis>
Torrey &amp; Gray, 1852
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) leaves in riparian forest along Little Bumblebee Creek, Panhandle National Forest, Shoshone County, Idaho at an elevation of 
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above sea level (asl). Additional 
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were collected by J. Baugh and W. Leonard in riparian forest along Trestle Creek, Kootenai County, Idaho (CM 63987, 66246) and Chatcolet Lake, Benewah County, Idaho (CM 64703).
</paragraph>
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<subSubSection id="C31D8099FFE5F925FE85FD2FFCB4C57B" pageId="6" pageNumber="7" type="etymology">
<paragraph id="8BB8D312FFE5F925FE85FD2FFCB4C57B" blockId="6.[264,1325,399,1105]" pageId="6" pageNumber="7">
<emphasis id="B9730F00FFE5F925FE85FD2FFE7FC4AA" box="[312,446,719,744]" italics="true" pageId="6" pageNumber="7">Etymology:</emphasis>
The specific name 
<taxonomicName id="4C07A891FFE5F925FD16FD2FFD37C4AA" box="[683,758,719,744]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="7" phylum="Mollusca" rank="species" species="burkei">
<emphasis id="B9730F00FFE5F925FD16FD2FFD37C4AA" box="[683,758,719,744]" italics="true" pageId="6" pageNumber="7">burkei</emphasis>
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honors naturalist Thomas E. Burke in recognition of over three decades of study of Pacific Northwest mollusks, and for teaching WL and JB most everything they know about land snails.
</paragraph>
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<subSubSection id="C31D8099FFE5F925FE85FCA7FAE1C213" pageId="6" pageNumber="7" type="distribution">
<paragraph id="8BB8D312FFE5F925FE85FCA7FAE1C213" blockId="6.[264,1325,399,1105]" pageId="6" pageNumber="7">
<emphasis id="B9730F00FFE5F925FE85FCA7FE10C522" box="[312,465,839,864]" italics="true" pageId="6" pageNumber="7">Distribution:</emphasis>
All known sites are located in either the Lake Pend Oreille or Coeur d’Alene Lake watersheds in the “panhandle” of northern Idaho (
<figureCitation id="133CCF97FFE5F925FBA5FC8FFBA0C5CB" box="[1048,1121,879,905]" captionStart="FIGURE 1" captionStartId="2.[264,367,1455,1479]" captionTargetBox="[267,1319,315,1431]" captionTargetId="figure@2.[267,1320,315,1431]" captionTargetPageId="2" captionText="FIGURE 1: Relief map of Idaho showing the localities of Kootenaia burkei new species. 1) Little Bumblebee Creek, Shoshone County (type locality), 2) Trestle Creek, Bonner County, 3) Beauty Creek, Kootenai County, 4) Heyburn State Park, Benewah County, and 5) East side of Harrison, Kootenai County. For detailed locality information see the Appendix. The lower photograph shows paratype CM 66247." httpUri="https://zenodo.org/record/168834/files/figure.png" pageId="6" pageNumber="7">Fig. 1</figureCitation>
). Specimens are known from the following locations in addition to the 
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locality: 2) Trestle Creek, Panhandle National Forest, Bonner County, Idaho, elevation 
<quantity id="4CFF7EF7FFE5F925FC76FC5FFBDBC59B" box="[971,1050,959,985]" metricMagnitude="2" metricUnit="m" metricValue="7.0" pageId="6" pageNumber="7" unit="m" value="700.0">700 m</quantity>
asl, 3) Beauty Creek, Idaho Panhandle National Forest, Kootenai County, Idaho, elevation 
<quantity id="4CFF7EF7FFE5F925FBFAFC07FB55C243" box="[1095,1172,999,1025]" metricMagnitude="2" metricUnit="m" metricValue="6.4" pageId="6" pageNumber="7" unit="m" value="640.0">640 m</quantity>
asl, 4) Heyburn State Park, Chatcolet Lake, Benewah County, Idaho, elevation 
<quantity id="4CFF7EF7FFE5F925FB95FBEFFBB3C26B" box="[1064,1138,1039,1065]" metricMagnitude="2" metricUnit="m" metricValue="6.5" pageId="6" pageNumber="7" unit="m" value="650.0">650 m</quantity>
asl, and 5) East side of Harrison, Kootenai County, Idaho, elevation 
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asl (for details see Appendix).
</paragraph>
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<subSubSection id="C31D8099FFE5F928FEB5FB67FCF9C127" lastPageId="11" lastPageNumber="12" pageId="6" pageNumber="7" type="description">
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<emphasis id="B9730F00FFE5F925FEB5FB67FE53C2E2" box="[264,402,1159,1184]" italics="true" pageId="6" pageNumber="7">Description</emphasis>
</paragraph>
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<emphasis id="B9730F00FFE5F925FE85FB4FFEB0C28A" box="[312,369,1199,1224]" italics="true" pageId="6" pageNumber="7">Size:</emphasis>
Very small slug, adults from 
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extended length while in movement.
</paragraph>
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<emphasis id="B9730F00FFE5F925FE85FB37FDD9C2B2" box="[312,536,1239,1264]" italics="true" pageId="6" pageNumber="7">External Features:</emphasis>
Head extending relatively short distance beyond mantle. Mantle elliptical, approximately half as long as extended body length; surface with lumps or wrinkles oriented roughly perpendicular to longitudinal axis. Tail rounded dorsally with no trace of mid­dorsal keel; with series of shallow parallel, longitudinal and oblique grooves; without line of abscission; lacking caudal mucus pore. Pneumostome positioned slightly posterior to middle of right side of mantle. Sole undivided, moderately wide. Pedal groove present above foot margin. Foot fringe moderately wide.
</paragraph>
<paragraph id="8BB8D312FFE5F925FE85FA0FFAE2C0BB" blockId="6.[264,1322,1159,1785]" pageId="6" pageNumber="7">
<emphasis id="B9730F00FFE5F925FE85FA0FFE03C04A" box="[312,450,1519,1544]" italics="true" pageId="6" pageNumber="7">Coloration:</emphasis>
Mantle pale gray or tan, with pale blue flecking, with dark gray or brown spots, blotches (
<figureCitation id="133CCF97FFE5F925FE77F9F7FDD0C073" box="[458,529,1559,1585]" captionStart="FIGURE 1" captionStartId="2.[264,367,1455,1479]" captionTargetBox="[267,1319,315,1431]" captionTargetId="figure@2.[267,1320,315,1431]" captionTargetPageId="2" captionText="FIGURE 1: Relief map of Idaho showing the localities of Kootenaia burkei new species. 1) Little Bumblebee Creek, Shoshone County (type locality), 2) Trestle Creek, Bonner County, 3) Beauty Creek, Kootenai County, 4) Heyburn State Park, Benewah County, and 5) East side of Harrison, Kootenai County. For detailed locality information see the Appendix. The lower photograph shows paratype CM 66247." httpUri="https://zenodo.org/record/168834/files/figure.png" pageId="6" pageNumber="7">Fig. 1</figureCitation>
). Tail light gray or tan with pale blue flecking and series of dark gray or brown, parallel, longitudinal stripes (centered on grooves); on dorsal surface two or more stripes converge approximately 
<quantity id="4CFF7EF7FFE5F925FD58F987FCEBC0C3" box="[741,810,1639,1665]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="6" pageNumber="7" unit="mm" value="1.0">1 mm</quantity>
prior to reaching posterior end of tail, then continue to pedal furrow; along sides stripes run transversely without merging. Sole light gray with whitish flecks. Mucus clear. Occasionally, specimens lack dark gray pigmentation on mantle, tail; other specimens with intense blue flecking covering both mantle, tail.
</paragraph>
<paragraph id="8BB8D312FFE4F924FEB5FEFCFE0DC777" blockId="7.[264,1006,284,390]" box="[264,460,284,309]" pageId="7" pageNumber="8">
<emphasis id="B9730F00FFE4F924FEB5FEFCFE0DC777" box="[264,460,284,309]" italics="true" pageId="7" pageNumber="8">Internal features</emphasis>
</paragraph>
<paragraph id="8BB8D312FFE4F924FE85FEA4FC2FC7C4" blockId="7.[264,1006,284,390]" pageId="7" pageNumber="8">
<emphasis id="B9730F00FFE4F924FE85FEA4FEBDC71F" box="[312,380,324,349]" italics="true" pageId="7" pageNumber="8">Shell:</emphasis>
Calcareous plate with slight anterior apex. 
<emphasis id="B9730F00FFE4F924FE85FE8CFEB1C7C7" box="[312,368,364,389]" italics="true" pageId="7" pageNumber="8">Jaw:</emphasis>
Ribbed, with as few as five to more than thirteen ribs.
</paragraph>
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<paragraph id="8BB8D312FFE4F924FEB5FA44FBC1C043" blockId="7.[264,1323,1444,1537]" pageId="7" pageNumber="8">
<emphasis id="B9730F00FFE4F924FEB5FA44FE44C3FE" bold="true" box="[264,389,1444,1468]" pageId="7" pageNumber="8">FIGURE 2</emphasis>
. Dissected holotype of 
<taxonomicName id="4C07A891FFE4F924FD31FA45FC86C3FE" box="[652,839,1445,1468]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="7" pageNumber="8" phylum="Mollusca" rank="species" species="burkei">
<emphasis id="B9730F00FFE4F924FD31FA45FC86C3FE" box="[652,839,1445,1468]" italics="true" pageId="7" pageNumber="8">Kootenaia burkei</emphasis>
</taxonomicName>
from Shoshone County, Idaho, showing the reproductive and digestive systems. Dorsal view of reproductive system on the left and ventral view of digestive system on the right. Total body length in alcohol = 10 mm.
</paragraph>
</caption>
<paragraph id="8BB8D312FFE4F92BFE85F9B5FC98C7BC" blockId="7.[264,1325,1621,1887]" lastBlockId="8.[264,1324,284,1190]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
<emphasis id="B9730F00FFE4F924FE85F9B5FDF9C02C" box="[312,568,1621,1646]" italics="true" pageId="7" pageNumber="8">Reproductive System:</emphasis>
Ovotestis comprises small number of lobules (as few as five); ovotestis partially embedded in posterior digestive gland (
<figureCitation id="133CCF97FFE4F924FC70F99DFBD8C0D5" box="[973,1049,1661,1687]" captionStart="FIGURE 2" captionStartId="7.[264,367,1444,1468]" captionTargetBox="[278,1308,435,1428]" captionTargetId="figure@7.[278,1309,429,1437]" captionTargetPageId="7" captionText="FIGURE 2. Dissected holotype of Kootenaia burkei from Shoshone County, Idaho, showing the reproductive and digestive systems. Dorsal view of reproductive system on the left and ventral view of digestive system on the right. Total body length in alcohol = 10 mm." httpUri="https://zenodo.org/record/156664/files/figure.png" pageId="7" pageNumber="8">Fig. 2</figureCitation>
). Hermaphroditic duct long, straight, not convoluted. Albumen gland large, extending to posterior limit of body cavity. Common duct long, occupying much of body cavity in mature individuals. No free epiphallus or vas deferens. Male component of reproductive system reduced to penial loop. Penial retractor muscle originates at anterior edge of diaphragm, inserts at apex of the penial loop. Penial loop consists of slender ascending duct, joining, at apex, a broader descending duct, the penis. On one stained slide mount, penis extended well into atrium. Atrium relatively large, barrel shaped, thin walled except for thickenings at genital pore. Duct of seminal receptacle arises from common duct at same level as penial loop but on opposite side of common duct. Duct of seminal receptacle, seminal receptacle loosely adhere to common duct, can be difficult to see in young animals. Duct and seminal receptacle enlarged, easily seen in fully mature animals.
</paragraph>
<paragraph id="8BB8D312FFEBF92BFE85FDECFB24C40C" blockId="8.[264,1324,284,1190]" pageId="8" pageNumber="9">
<emphasis id="B9730F00FFEBF92BFE85FDECFCEDC467" box="[312,812,524,549]" italics="true" pageId="8" pageNumber="9">Buccal and Tentacular Retractor Muscles:</emphasis>
Both buccal, tentacular retractors have origins widely separated at posterior margin of diaphragm. No retentor muscle present.
</paragraph>
<paragraph id="8BB8D312FFEBF92BFE85FDBCFEB7C2E4" blockId="8.[264,1324,284,1190]" pageId="8" pageNumber="9">
<emphasis id="B9730F00FFEBF92BFE85FDBCFDCAC437" box="[312,523,604,629]" italics="true" pageId="8" pageNumber="9">Digestive System:</emphasis>
Buccal mass large but otherwise unremarkable. Slender esophagus emerges dorsally, posteriorly to join crop. Crop distinctive: cylindrical shape modified by several circumferential constrictions that partially divide crop lumen into series of tandem compartments. Because constrictions are not quite perpendicular to long axis of crop, effect is to produce a spiral pathway for crop contents moving toward the stomach. Inner lining of crop consists of myriad of small, uniformly sized, spindle­shaped, opaque tissue patches. Patches arranged to reflect the spiral nature of space they help enclose. Pattern of patches changes at posterior end of crop to become longitudinal one just before crop joins stomach. Stomach forms right angle with long axis of crop. Anterior and posterior digestive glands connected to stomach by pair of ducts. Stomach leads to intestine, a somewhat flattened tube of uniform width that makes three bends before arriving at anus in wall of pneumostome. Aside from partially embedded ovotestis and minor adhesions between posterior digestive gland and albumen gland, the digestive and reproductive systems not intimately intertwined. 
<figureCitation id="133CCF97FFEBF92BFDA2FB84FDA7C23C" box="[543,614,1124,1150]" captionStart="FIGURE 2" captionStartId="7.[264,367,1444,1468]" captionTargetBox="[278,1308,435,1428]" captionTargetId="figure@7.[278,1309,429,1437]" captionTargetPageId="7" captionText="FIGURE 2. Dissected holotype of Kootenaia burkei from Shoshone County, Idaho, showing the reproductive and digestive systems. Dorsal view of reproductive system on the left and ventral view of digestive system on the right. Total body length in alcohol = 10 mm." httpUri="https://zenodo.org/record/156664/files/figure.png" pageId="8" pageNumber="9">Fig. 2</figureCitation>
shows major anatomical features revealed by dissection of 
<typeStatus id="54BC6DB0FFEBF92BFEB5FB6CFEB3C2E4" box="[264,370,1164,1190]" pageId="8" pageNumber="9" type="holotype">holotype</typeStatus>
.
</paragraph>
<paragraph id="8BB8D312FFEBF92BFEB5FB3CFE7AC2B7" blockId="8.[264,1323,1244,1870]" box="[264,443,1244,1269]" pageId="8" pageNumber="9">
<emphasis id="B9730F00FFEBF92BFEB5FB3CFE7AC2B7" box="[264,443,1244,1269]" italics="true" pageId="8" pageNumber="9">Natural history</emphasis>
</paragraph>
<paragraph id="8BB8D312FFEBF92BFE85FAE4FC9FC10C" blockId="8.[264,1323,1244,1870]" pageId="8" pageNumber="9">
All known sites are forested and adjacent to a perennial water body, which is probably necessary for maintaining high soil moisture and moderating ambient temperature. Vegetation at the sites was composed of western hemlock (
<taxonomicName id="4C07A891FFEBF92BFC1EFAB4FE97C3D4" authority="Rafinesque, 1832" authorityName="Rafinesque" authorityYear="1832" class="Pinopsida" family="Pinaceae" genus="Tsuga" kingdom="Plantae" order="Pinales" pageId="8" pageNumber="9" phylum="Tracheophyta" rank="species" species="heterophylla">
<emphasis id="B9730F00FFEBF92BFC1EFAB4FB49C32F" box="[931,1160,1364,1389]" italics="true" pageId="8" pageNumber="9">Tsuga heterophylla</emphasis>
(Rafinesque, 1832))
</taxonomicName>
forest, with scattered western redcedar (
<taxonomicName id="4C07A891FFEBF92BFC82FA9CFBB9C3D4" authority="Donn, 1824" authorityName="Donn" authorityYear="1824" box="[831,1144,1404,1430]" class="Pinopsida" family="Cupressaceae" genus="Thuja" kingdom="Plantae" order="Pinales" pageId="8" pageNumber="9" phylum="Tracheophyta" rank="species" species="plicata">
<emphasis id="B9730F00FFEBF92BFC82FA9CFC1FC3D7" box="[831,990,1404,1429]" italics="true" pageId="8" pageNumber="9">Thuja plicata</emphasis>
Donn, 1824
</taxonomicName>
), black cottonwood (
<taxonomicName id="4C07A891FFEBF92BFEE6FA44FC93C3FC" authority="Torrey &amp; Gray, 1852" authorityName="Torrey &amp; Gray" authorityYear="1852" box="[347,850,1444,1470]" class="Magnoliopsida" family="Salicaceae" genus="Populus" kingdom="Plantae" order="Malpighiales" pageId="8" pageNumber="9" phylum="Tracheophyta" rank="species" species="trichocarpa">
<emphasis id="B9730F00FFEBF92BFEE6FA44FD8FC3FF" box="[347,590,1444,1469]" italics="true" pageId="8" pageNumber="9">Populus trichocarpa</emphasis>
Torrey &amp; Gray, 1852
</taxonomicName>
), paper birch (
<taxonomicName id="4C07A891FFEBF92BFC42FA44FE87C3A4" authority="Marsh, 1785" authorityName="Marsh" authorityYear="1785" class="Magnoliopsida" family="Betulaceae" genus="Betula" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="9" phylum="Tracheophyta" rank="species" species="papyrifera">
<emphasis id="B9730F00FFEBF92BFC42FA44FB11C3FF" box="[1023,1232,1444,1469]" italics="true" pageId="8" pageNumber="9">Betula papyrifera</emphasis>
Marsh, 1785
</taxonomicName>
), chokecherry (
<taxonomicName id="4C07A891FFEBF92BFDBBFA2CFC69C3A4" authority="Linnaeus, 1753" authorityName="Linnaeus" authorityYear="1753" box="[518,936,1484,1510]" class="Magnoliopsida" family="Rosaceae" genus="Prunus" kingdom="Plantae" order="Rosales" pageId="8" pageNumber="9" phylum="Tracheophyta" rank="species" species="virginiana">
<emphasis id="B9730F00FFEBF92BFDBBFA2CFD21C3A7" box="[518,736,1484,1509]" italics="true" pageId="8" pageNumber="9">Prunus virginiana</emphasis>
Linnaeus, 1753
</taxonomicName>
), and/or red alder (
<taxonomicName id="4C07A891FFEBF92BFB24FA2CFE7FC04C" authority="Bongard, 1832" authorityName="Bongard" authorityYear="1832" class="Magnoliopsida" family="Betulaceae" genus="Alnus" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="9" phylum="Tracheophyta" rank="species" species="rubra">
<emphasis id="B9730F00FFEBF92BFB24FA2CFAEAC3A7" box="[1177,1323,1484,1509]" italics="true" pageId="8" pageNumber="9">Alnus rubra</emphasis>
Bongard, 1832
</taxonomicName>
). Most specimens were collected on the forest floor, either on or under woody debris, mats of moss, or deciduous tree leaves; two specimens were collected approximately 
<quantity id="4CFF7EF7FFEBF92BFE03F9A4FDC6C01C" box="[446,519,1604,1630]" metricMagnitude="-1" metricUnit="m" metricValue="2.0" pageId="8" pageNumber="9" unit="m" value="0.2">0.2 m</quantity>
aboveground on a moss­covered tree trunk along the edge of a stream. In the field we have observed specimens resting on and, apparently, eating lichens that were growing on woody debris. Five specimens from Trestle Creek, maintained in captivity between 
<date id="FFB9F5D2FFEBF92BFE60F95CFD52C094" box="[477,659,1724,1750]" pageId="8" pageNumber="9" value="2002-10-05">5 October 2002</date>
and 
<date id="FFB9F5D2FFEBF92BFD77F95CFCFEC094" box="[714,831,1724,1750]" pageId="8" pageNumber="9" value="2003-07">July 2003</date>
, ate carrot, lettuce, sweet potato, yam, dry dog food, and goldfish flakes; one of these specimens (
<quantity id="4CFF7EF7FFEBF92BFC16F904FC32C0BC" box="[939,1011,1764,1790]" metricMagnitude="-3" metricUnit="m" metricValue="9.0" pageId="8" pageNumber="9" unit="mm" value="9.0">9 mm</quantity>
extended length while in movement) laid 3 oval eggs on 
<date id="FFB9F5D2FFEBF92BFDC1F8ECFCD6C164" box="[636,791,1804,1830]" pageId="8" pageNumber="9" value="2003-07-17">17 July 2003</date>
. These eggs (
<emphasis id="B9730F00FFEBF92BFC04F8ECFC15C167" box="[953,980,1804,1829]" italics="true" pageId="8" pageNumber="9">ca</emphasis>
. 
<date id="FFB9F5D2FFEBF92BFC59F8ECFBCBC164" box="[996,1034,1804,1830]" pageId="8" pageNumber="9">1 x</date>
<quantity id="4CFF7EF7FFEBF92BFBACF8ECFBAFC164" box="[1041,1134,1804,1830]" metricMagnitude="-3" metricUnit="m" metricValue="1.8" pageId="8" pageNumber="9" unit="mm" value="1.8">1.8 mm</quantity>
) were all viable and near to hatching on 
<date id="FFB9F5D2FFEBF92BFD9FF8D4FD39C10C" box="[546,760,1844,1870]" pageId="8" pageNumber="9" value="2003-09-08">8 September 2003</date>
(
<figureCitation id="133CCF97FFEBF92BFCBAF8D4FC8FC10C" box="[775,846,1844,1870]" captionStart="FIGURE 3" captionStartId="9.[264,367,710,734]" captionTargetBox="[491,1095,313,699]" captionTargetId="figure@9.[469,1118,312,702]" captionTargetPageId="9" captionText="FIGURE 3. Eggs of Kootenaia burkei photographed on 10 September 2003, 62 days after being laid in the laboratory." httpUri="https://zenodo.org/record/168835/files/figure.png" pageId="8" pageNumber="9">Fig. 3</figureCitation>
).
</paragraph>
<caption id="DF78839AFFEAF92AFEB5FD26FE2CC542" httpUri="https://zenodo.org/record/168835/files/figure.png" pageId="9" pageNumber="10" targetBox="[491,1095,313,699]" targetPageId="9">
<paragraph id="8BB8D312FFEAF92AFEB5FD26FE2CC542" blockId="9.[264,1322,710,768]" pageId="9" pageNumber="10">
<emphasis id="B9730F00FFEAF92AFEB5FD26FE44C49C" bold="true" box="[264,389,710,734]" pageId="9" pageNumber="10">FIGURE 3</emphasis>
. Eggs of 
<taxonomicName id="4C07A891FFEAF92AFE4FFD27FD6FC49C" box="[498,686,711,734]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="burkei">
<emphasis id="B9730F00FFEAF92AFE4FFD27FD6FC49C" box="[498,686,711,734]" italics="true" pageId="9" pageNumber="10">Kootenaia burkei</emphasis>
</taxonomicName>
photographed on 10 September 2003, 62 days after being laid in the laboratory.
</paragraph>
</caption>
<paragraph id="8BB8D312FFEAF92AFEB5FCBCFE02C584" pageId="9" pageNumber="10">
<table id="F90721B2FFEA06DCFEB5FCBCFAE8C584" box="[264,1321,860,966]" gridcols="7" gridrows="3" pageId="9" pageNumber="10">
<tr id="3537D150FFEA06DCFEB5FCBCFAE8C537" box="[264,1321,860,885]" gridrow="0" pageId="9" pageNumber="10" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1">
<th id="76E6B82CFFEA06DCFEB5FCBCFD95C537" box="[264,596,860,885]" gridcol="0" gridrow="0" pageId="9" pageNumber="10">
<emphasis id="B9730F00FFEAF92AFEB5FCBCFDC1C537" box="[264,512,860,885]" italics="true" pageId="9" pageNumber="10">Additional comments</emphasis>
</th>
</tr>
<tr id="3537D150FFEA06DCFEB5FC64FAE8C5DC" box="[264,1321,900,926]" gridrow="1" pageId="9" pageNumber="10">
<th id="76E6B82CFFEA06DCFEB5FC64FD95C5DC" box="[264,596,900,926]" gridcol="0" gridrow="1" pageId="9" pageNumber="10">We suggest the common</th>
<td id="76E6B82CFFEA06DCFDE6FC64FD3EC5DC" box="[603,767,900,926]" gridcol="1" gridrow="1" pageId="9" pageNumber="10">name “pygmy</td>
<td id="76E6B82CFFEA06DCFCBBFC64FC8AC5DC" box="[774,843,900,926]" gridcol="2" gridrow="1" pageId="9" pageNumber="10">slug,”</td>
<td id="76E6B82CFFEA06DCFCEFFC64FCA8C5DC" box="[850,873,900,926]" gridcol="3" gridrow="1" pageId="9" pageNumber="10">in</td>
<td id="76E6B82CFFEA06DCFCCDFC64FBD5C5DC" box="[880,1044,900,926]" gridcol="4" gridrow="1" pageId="9" pageNumber="10">recognition of</td>
<td id="76E6B82CFFEA06DCFBA6FC64FB62C5DC" box="[1051,1187,900,926]" gridcol="5" gridrow="1" pageId="9" pageNumber="10">the species’</td>
<td id="76E6B82CFFEA06DCFB17FC64FAE8C5DC" box="[1194,1321,900,926]" gridcol="6" gridrow="1" pageId="9" pageNumber="10">diminutive</td>
</tr>
<tr id="3537D150FFEA06DCFEB5FC4CFAE8C584" box="[264,1321,940,966]" gridrow="2" pageId="9" pageNumber="10" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1">
<th id="76E6B82CFFEA06DCFEB5FC4CFD95C584" box="[264,596,940,966]" gridcol="0" gridrow="2" pageId="9" pageNumber="10">size at maturity.</th>
</tr>
</table>
</paragraph>
<paragraph id="8BB8D312FFEAF92AFEB5FC1CFDCCC257" blockId="9.[264,1323,1020,1686]" box="[264,525,1020,1045]" pageId="9" pageNumber="10">
<emphasis id="B9730F00FFEAF92AFEB5FC1CFDCCC257" box="[264,525,1020,1045]" italics="true" pageId="9" pageNumber="10">Comparative anatomy</emphasis>
</paragraph>
<paragraph id="8BB8D312FFEAF92AFE85FBC4FEF9C0D4" blockId="9.[264,1323,1020,1686]" pageId="9" pageNumber="10">
<taxonomicName id="4C07A891FFEAF92AFE85FBC4FDC5C27F" box="[312,516,1060,1085]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="burkei">
<emphasis id="B9730F00FFEAF92AFE85FBC4FDC5C27F" box="[312,516,1060,1085]" italics="true" pageId="9" pageNumber="10">Kootenaia burkei</emphasis>
</taxonomicName>
differs from all other known arionids by lacking a free epiphallus or vas deferens; to our knowledge, no other arionid has such a drastically reduced male component of the reproductive system. The only character that it shares with all other arionids is a ribbed jaw. We place 
<taxonomicName id="4C07A891FFEAF92AFD83FB7CFD6CC2F7" box="[574,685,1180,1205]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="burkei">
<emphasis id="B9730F00FFEAF92AFD83FB7CFD6CC2F7" box="[574,685,1180,1205]" italics="true" pageId="9" pageNumber="10">K. burkei</emphasis>
</taxonomicName>
in the 
<taxonomicName id="4C07A891FFEAF92AFCBEFB7CFCB9C2F4" box="[771,888,1180,1206]" class="Gastropoda" family="Arionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="family">Arionidae</taxonomicName>
based on the presence of the ribbed jaw and the widely separated origins of the buccal and tentacular retractors, a condition found in several arionid genera including 
<taxonomicName id="4C07A891FFEAF92AFD49FB0CFC40C347" box="[756,897,1260,1285]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFD49FB0CFC40C347" box="[756,897,1260,1285]" italics="true" pageId="9" pageNumber="10">Prophysaon</emphasis>
</taxonomicName>
. It differs from 
<taxonomicName id="4C07A891FFEAF92AFB80FB0CFB0BC347" box="[1085,1226,1260,1285]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFB80FB0CFB0BC347" box="[1085,1226,1260,1285]" italics="true" pageId="9" pageNumber="10">Prophysaon</emphasis>
</taxonomicName>
in lacking a line of abscission and in possessing a penial retractor. It differs from 
<taxonomicName id="4C07A891FFEAF92AFBCBFAF4FB21C36F" box="[1142,1248,1300,1325]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFBCBFAF4FB21C36F" box="[1142,1248,1300,1325]" italics="true" pageId="9" pageNumber="10">Zacoleus</emphasis>
</taxonomicName>
in the widely separated attachments of the buccal and tentacular retractors, in possessing a penial retractor, and in having an undivided sole. It differs from 
<taxonomicName id="4C07A891FFEAF92AFC12FA84FBF2C33F" box="[943,1075,1380,1405]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFC12FA84FBF2C33F" box="[943,1075,1380,1405]" italics="true" pageId="9" pageNumber="10">Hemphillia</emphasis>
</taxonomicName>
in possessing a fully embedded, calcareous shell; in the widely separated attachments of the buccal and tentacular retractors; in lacking a retentor muscle and in having the visceral cavity extend to the tip of the tail (see 
<tableCitation id="C685E6A9FFEAF92AFE5EFA3CFDFAC3B4" box="[483,571,1500,1526]" captionStart="TABLE 1" captionStartId="5.[264,352,443,467]" captionTargetBox="[267,1291,591,1776]" captionTargetPageId="5" captionText="TABLE 1. Comparison of selected characters and their states in Kootenaia burkei, Hemphillia camelus, H. dromedarius, H. glandulosa, Prophysaon coeruleum, P. dubium, P. humile, P. vanattae, and Zacoleus idahoensis." httpUri="http://table.plazi.org/id/DF78839AFFE6F926FEB5FE5BFDD4C463" pageId="9" pageNumber="10" tableUuid="DF78839AFFE6F926FEB5FE5BFDD4C463">Table 1</tableCitation>
). A maximum parsimony tree, based on 15 anatomical characters for taxa of 
<emphasis id="B9730F00FFEAF92AFE7AF9E4FCB4C05F" box="[455,885,1540,1565]" italics="true" pageId="9" pageNumber="10">
<taxonomicName id="4C07A891FFEAF92AFE7AF9E4FD8FC05F" box="[455,590,1540,1565]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">Hemphillia</taxonomicName>
, 
<taxonomicName id="4C07A891FFEAF92AFDE0F9E4FD1BC05F" box="[605,730,1540,1565]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">Kootenaia</taxonomicName>
, 
<taxonomicName id="4C07A891FFEAF92AFD54F9E4FCB4C05F" box="[745,885,1540,1565]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">Prophysaon</taxonomicName>
</emphasis>
, and 
<taxonomicName id="4C07A891FFEAF92AFC06F9E4FBE4C05F" box="[955,1061,1540,1565]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFC06F9E4FBE4C05F" box="[955,1061,1540,1565]" italics="true" pageId="9" pageNumber="10">Zacoleus</emphasis>
</taxonomicName>
(see 
<figureCitation id="133CCF97FFEAF92AFBD4F9E4FB0BC05C" box="[1129,1226,1540,1566]" captionStart="FIGURE 4" captionStartId="10.[264,367,1244,1268]" captionTargetBox="[268,1318,449,1218]" captionTargetId="figure@10.[268,1318,449,1218]" captionTargetPageId="10" captionText="FIGURE 4: A. Maximum parsimony tree for representatives of the genera Hemphillia, Kootenaia, Prophysaon, and Zacoleus based on 15 anatomical characters. Bootstrap values (10,000 replicates) are given at the nodes. The scale bar indicated the number of changes. B. Bayesian phylogram for the same species based on combined fragments of mitochondrial COI DNA and LSU rRNA showing the 50 % majority­rule consensus of topologies sampled during the Bayesian search. The scale bar indicates the substitution rate according to the model of sequence evolution applied. The numbers at the branches are the percentages that the clade occurs among all sampled trees; i. e. the posterior probability of that clade." httpUri="https://zenodo.org/record/156666/files/figure.png" pageId="9" pageNumber="10">Fig. 4A</figureCitation>
), shows that our new taxon clusters distinctly apart from the four species of 
<taxonomicName id="4C07A891FFEAF92AFB82F9CCFB0DC007" box="[1087,1228,1580,1605]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFB82F9CCFB0DC007" box="[1087,1228,1580,1605]" italics="true" pageId="9" pageNumber="10">Prophysaon</emphasis>
</taxonomicName>
studied here. The individuals of 
<taxonomicName id="4C07A891FFEAF92AFD8DF9B4FD6BC02F" box="[560,682,1620,1645]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFD8DF9B4FD6BC02F" box="[560,682,1620,1645]" italics="true" pageId="9" pageNumber="10">Kootenaia</emphasis>
</taxonomicName>
also cluster apart from 
<taxonomicName id="4C07A891FFEAF92AFC76F9B4FBA8C02F" box="[971,1129,1620,1645]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="idahoensis">
<emphasis id="B9730F00FFEAF92AFC76F9B4FBA8C02F" box="[971,1129,1620,1645]" italics="true" pageId="9" pageNumber="10">Z. idahoensis</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C07A891FFEAF92AFB1AF9B4FAEAC02F" box="[1191,1323,1620,1645]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFB1AF9B4FAEAC02F" box="[1191,1323,1620,1645]" italics="true" pageId="9" pageNumber="10">Hemphillia</emphasis>
</taxonomicName>
spp.
</paragraph>
<paragraph id="8BB8D312FFEAF92AFEB5F92CFD47C0A7" blockId="9.[264,1323,1740,1886]" box="[264,646,1740,1765]" pageId="9" pageNumber="10">
<emphasis id="B9730F00FFEAF92AFEB5F92CFD47C0A7" box="[264,646,1740,1765]" italics="true" pageId="9" pageNumber="10">Comparative molecular genetics</emphasis>
</paragraph>
<paragraph id="8BB8D312FFEAF929FE85F914FB9EC71C" blockId="9.[264,1323,1740,1886]" lastBlockId="10.[264,1323,284,350]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
The average among­group divergence based on K2P­distances ranges from 0.184 0.012 (between 
<taxonomicName id="4C07A891FFEAF92AFDA7F8FCFD45C177" box="[538,644,1820,1845]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFDA7F8FCFD45C177" box="[538,644,1820,1845]" italics="true" pageId="9" pageNumber="10">Zacoleus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C07A891FFEAF92AFD7CF8FCFC84C177" box="[705,837,1820,1845]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFD7CF8FCFC84C177" box="[705,837,1820,1845]" italics="true" pageId="9" pageNumber="10">Hemphillia</emphasis>
</taxonomicName>
) and 0.227±0.012 (between 
<taxonomicName id="4C07A891FFEAF92AFB23F8FCFAEBC177" box="[1182,1322,1820,1845]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFB23F8FCFAEBC177" box="[1182,1322,1820,1845]" italics="true" pageId="9" pageNumber="10">Prophysaon</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C07A891FFEAF92AFE86F8A4FE7EC11F" box="[315,447,1860,1885]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFE86F8A4FE7EC11F" box="[315,447,1860,1885]" italics="true" pageId="9" pageNumber="10">Hemphillia</emphasis>
</taxonomicName>
) (see 
<tableCitation id="C685E6A9FFEAF92AFDB4F8A4FDA2C11C" box="[521,611,1860,1886]" captionStart="TABLE 2" captionStartId="11.[264,352,403,427]" captionTargetBox="[264,1265,543,858]" captionTargetPageId="11" captionText="TABLE 2. Pairwise K 2 P­distances among the genera Kootenaia, Zacoleus, Prophysaon, and Hemphillia for the combined COI and LSU rRNA sequences. Standard errors (based on 1000 bootstrap replications) are given in parenthesis." httpUri="http://table.plazi.org/id/DF78839AFFE8F928FEB5FE73FD13C7BB" pageId="9" pageNumber="10" tableUuid="DF78839AFFE8F928FEB5FE73FD13C7BB">Table 2</tableCitation>
). 
<taxonomicName id="4C07A891FFEAF92AFDC6F8A4FD34C11F" box="[635,757,1860,1885]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFDC6F8A4FD34C11F" box="[635,757,1860,1885]" italics="true" pageId="9" pageNumber="10">Kootenaia</emphasis>
</taxonomicName>
appears to be most similar to 
<taxonomicName id="4C07A891FFEAF92AFBE2F8A4FB08C11F" box="[1119,1225,1860,1885]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFEAF92AFBE2F8A4FB08C11F" box="[1119,1225,1860,1885]" italics="true" pageId="9" pageNumber="10">Zacoleus</emphasis>
</taxonomicName>
with an average divergence of 0.185±0.014, though the divergences with 
<taxonomicName id="4C07A891FFE9F929FB23FEFCFAEAC777" box="[1182,1323,284,309]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FB23FEFCFAEAC777" box="[1182,1323,284,309]" italics="true" pageId="10" pageNumber="11">Prophysaon</emphasis>
</taxonomicName>
(0.204±0.013) and 
<taxonomicName id="4C07A891FFE9F929FE54FEA4FDACC71F" box="[489,621,324,349]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FE54FEA4FDACC71F" box="[489,621,324,349]" italics="true" pageId="10" pageNumber="11">Hemphillia</emphasis>
</taxonomicName>
(0.206±0.012) are not significantly larger.
</paragraph>
<caption id="DF78839AFFE9F929FEB5FB3CFD8EC3A5" httpUri="https://zenodo.org/record/156666/files/figure.png" pageId="10" pageNumber="11" targetBox="[268,1318,449,1218]" targetPageId="10">
<paragraph id="8BB8D312FFE9F929FEB5FB3CFD8EC3A5" blockId="10.[264,1323,1244,1511]" pageId="10" pageNumber="11">
<emphasis id="B9730F00FFE9F929FEB5FB3CFE42C2B6" bold="true" box="[264,387,1244,1268]" pageId="10" pageNumber="11">FIGURE 4</emphasis>
: A. Maximum parsimony tree for representatives of the genera 
<taxonomicName id="4C07A891FFE9F929FB92FB3DFB66C2B6" box="[1071,1191,1245,1268]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FB92FB3DFB66C2B6" box="[1071,1191,1245,1268]" italics="true" pageId="10" pageNumber="11">Hemphillia</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C07A891FFE9F929FB08FB3DFAE5C2B6" box="[1205,1316,1245,1268]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FB08FB3DFAE5C2B6" box="[1205,1316,1245,1268]" italics="true" pageId="10" pageNumber="11">Kootenaia</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C07A891FFE9F929FEB5FAE0FE49C355" box="[264,392,1280,1303]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FEB5FAE0FE49C355" box="[264,392,1280,1303]" italics="true" pageId="10" pageNumber="11">Prophysaon</emphasis>
</taxonomicName>
, and 
<taxonomicName id="4C07A891FFE9F929FE79FAE0FDE5C355" box="[452,548,1280,1303]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FE79FAE0FDE5C355" box="[452,548,1280,1303]" italics="true" pageId="10" pageNumber="11">Zacoleus</emphasis>
</taxonomicName>
based on 15 anatomical characters. Bootstrap values (10,000 replicates) are given at the nodes. The scale bar indicated the number of changes. B. Bayesian phylogram for the same species based on combined fragments of mitochondrial COI DNA and LSU rRNA showing the 50% majority­rule consensus of topologies sampled during the Bayesian search. The scale bar indicates the substitution rate according to the model of sequence evolution applied. The numbers at the branches are the percentages that the clade occurs among all sampled trees; 
<emphasis id="B9730F00FFE9F929FB10FA4DFB0CC386" box="[1197,1229,1453,1476]" italics="true" pageId="10" pageNumber="11">i.e.</emphasis>
the posterior probability of that clade.
</paragraph>
</caption>
<paragraph id="8BB8D312FFE9F929FE85F9A2FCD8C096" blockId="10.[264,1323,1602,1868]" pageId="10" pageNumber="11">
A Bayesian phylogram of the combined COI/LSU rRNA data set reflecting the 50% majority­rule consensus of topologies sampled during the Bayesian search is shown in 
<figureCitation id="133CCF97FFE9F929FB44F98AFED8C0EE" captionStart="FIGURE 4" captionStartId="10.[264,367,1244,1268]" captionTargetBox="[268,1318,449,1218]" captionTargetId="figure@10.[268,1318,449,1218]" captionTargetPageId="10" captionText="FIGURE 4: A. Maximum parsimony tree for representatives of the genera Hemphillia, Kootenaia, Prophysaon, and Zacoleus based on 15 anatomical characters. Bootstrap values (10,000 replicates) are given at the nodes. The scale bar indicated the number of changes. B. Bayesian phylogram for the same species based on combined fragments of mitochondrial COI DNA and LSU rRNA showing the 50 % majority­rule consensus of topologies sampled during the Bayesian search. The scale bar indicates the substitution rate according to the model of sequence evolution applied. The numbers at the branches are the percentages that the clade occurs among all sampled trees; i. e. the posterior probability of that clade." httpUri="https://zenodo.org/record/156666/files/figure.png" pageId="10" pageNumber="11">Fig. 4</figureCitation>
B. The tree comprises the ‘outgroup’ taxa 
<taxonomicName id="4C07A891FFE9F929FCBBF973FC63C0EE" box="[774,930,1683,1708]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="idahoensis">
<emphasis id="B9730F00FFE9F929FCBBF973FC63C0EE" box="[774,930,1683,1708]" italics="true" pageId="10" pageNumber="11">Z. idahoensis</emphasis>
</taxonomicName>
and three major clades: 
<taxonomicName id="4C07A891FFE9F929FB7DF973FEF5C096" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FB7DF973FEF5C096" italics="true" pageId="10" pageNumber="11">Prophysaon</emphasis>
</taxonomicName>
spp., 
<taxonomicName id="4C07A891FFE9F929FEC7F95BFE35C096" box="[378,500,1723,1748]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FEC7F95BFE35C096" box="[378,500,1723,1748]" italics="true" pageId="10" pageNumber="11">Kootenaia</emphasis>
</taxonomicName>
sp., and 
<taxonomicName id="4C07A891FFE9F929FDE0F95BFD20C096" box="[605,737,1723,1748]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FDE0F95BFD20C096" box="[605,737,1723,1748]" italics="true" pageId="10" pageNumber="11">Hemphillia</emphasis>
</taxonomicName>
spp.
</paragraph>
<paragraph id="8BB8D312FFE9F928FE85F903FE72C71C" blockId="10.[264,1323,1602,1868]" lastBlockId="11.[264,1322,284,350]" lastPageId="11" lastPageNumber="12" pageId="10" pageNumber="11">
<taxonomicName id="4C07A891FFE9F929FE85F903FE73C0BE" box="[312,434,1763,1788]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FE85F903FE73C0BE" box="[312,434,1763,1788]" italics="true" pageId="10" pageNumber="11">Kootenaia</emphasis>
</taxonomicName>
clusters distinctly apart from all other taxa studied here and both the 
<taxonomicName id="4C07A891FFE9F929FB4BF903FEABC166" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FB4BF903FEABC166" italics="true" pageId="10" pageNumber="11">Prophysaon</emphasis>
</taxonomicName>
and the 
<taxonomicName id="4C07A891FFE9F929FE6EF8EBFD8CC166" box="[467,589,1803,1828]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE9F929FE6EF8EBFD8CC166" box="[467,589,1803,1828]" italics="true" pageId="10" pageNumber="11">Kootenaia</emphasis>
</taxonomicName>
clades have posterior probabilities of 1.00. Within 
<taxonomicName id="4C07A891FFE9F929FB09F8EBFAE2C166" box="[1204,1315,1803,1828]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="burkei">
<emphasis id="B9730F00FFE9F929FB09F8EBFAE2C166" box="[1204,1315,1803,1828]" italics="true" pageId="10" pageNumber="11">K. burkei</emphasis>
</taxonomicName>
, the two specimens for each of the two populations studied are homogeneous and the average divergence between the two populations is very low with a K2P­distance of 0.0044±0.002.
</paragraph>
<caption id="DF78839AFFE8F928FEB5FE73FD13C7BB" ID-Table-UUID="DF78839AFFE8F928FEB5FE73FD13C7BB" httpUri="http://table.plazi.org/id/DF78839AFFE8F928FEB5FE73FD13C7BB" pageId="11" pageNumber="12" targetBox="[264,1265,543,858]" targetIsTable="true" targetPageId="11">
<paragraph id="8BB8D312FFE8F928FEB5FE73FD13C7BB" blockId="11.[264,1323,403,505]" pageId="11" pageNumber="12">
<emphasis id="B9730F00FFE8F928FEB5FE73FE40C7E9" bold="true" box="[264,385,403,427]" pageId="11" pageNumber="12">TABLE 2.</emphasis>
Pairwise K2P­distances among the genera 
<taxonomicName id="4C07A891FFE8F928FCC4FE74FC29C7E9" box="[889,1000,404,427]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FCC4FE74FC29C7E9" box="[889,1000,404,427]" italics="true" pageId="11" pageNumber="12">Kootenaia</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C07A891FFE8F928FC41FE74FB9DC7E9" box="[1020,1116,404,427]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FC41FE74FB9DC7E9" box="[1020,1116,404,427]" italics="true" pageId="11" pageNumber="12">Zacoleus</emphasis>
</taxonomicName>
, 
<emphasis id="B9730F00FFE8F928FBCDFE74FB36C7E9" box="[1136,1271,404,427]" italics="true" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FBCDFE74FB32C7E9" box="[1136,1267,404,427]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">Prophysaon</taxonomicName>
,
</emphasis>
and 
<taxonomicName id="4C07A891FFE8F928FEB5FE5AFE41C793" box="[264,384,442,465]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FEB5FE5AFE41C793" box="[264,384,442,465]" italics="true" pageId="11" pageNumber="12">Hemphillia</emphasis>
</taxonomicName>
for the combined COI and LSU rRNA sequences. Standard errors (based on 1000 bootstrap replications) are given in parenthesis.
</paragraph>
</caption>
<paragraph id="8BB8D312FFE8F928FDB3FDFFFE4DC518" pageId="11" pageNumber="12">
<table id="F90721B2FFE806DCFEB5FDFFFB30C518" box="[264,1265,543,858]" gridcols="5" gridrows="6" pageId="11" pageNumber="12">
<tr id="3537D150FFE806DCFEB5FDFFFB30C476" box="[264,1265,543,564]" gridrow="0" pageId="11" pageNumber="12" rowspan-0="1">
<th id="76E6B82CFFE806DCFE42FDFFFD43C476" box="[511,642,543,564]" gridcol="1" gridrow="0" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FDB3FDFFFDB3C476" box="[526,626,543,564]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FDB3FDFFFDB3C476" box="[526,626,543,564]" italics="true" pageId="11" pageNumber="12">Kootenaia</emphasis>
</taxonomicName>
</th>
<th id="76E6B82CFFE806DCFD6EFDFFFC94C476" box="[723,853,543,564]" gridcol="2" gridrow="0" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FD54FDFFFC81C476" box="[745,832,543,564]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FD54FDFFFC81C476" box="[745,832,543,564]" italics="true" pageId="11" pageNumber="12">Zacoleus</emphasis>
</taxonomicName>
</th>
<th id="76E6B82CFFE806DCFC1AFDFFFBE8C476" box="[935,1065,543,564]" gridcol="3" gridrow="0" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FC13FDFFFBE0C476" box="[942,1057,543,564]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FC13FDFFFBE0C476" box="[942,1057,543,564]" italics="true" pageId="11" pageNumber="12">Prophysaon</emphasis>
</taxonomicName>
</th>
<th id="76E6B82CFFE806DCFB38FDFFFB30C476" box="[1157,1265,543,564]" gridcol="4" gridrow="0" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FB38FDFFFB30C476" box="[1157,1265,543,564]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FB38FDFFFB30C476" box="[1157,1265,543,564]" italics="true" pageId="11" pageNumber="12">Hemphillia</emphasis>
</taxonomicName>
</th>
</tr>
<tr id="3537D150FFE806DCFEB5FDA9FB30C41C" box="[264,1265,585,606]" gridrow="1" pageId="11" pageNumber="12" rowspan-2="1" rowspan-3="1" rowspan-4="1">
<th id="76E6B82CFFE806DCFEB5FDA9FE4DC41C" box="[264,396,585,606]" gridcol="0" gridrow="1" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FEB6FDA9FEAEC41C" box="[267,367,585,606]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FEB6FDA9FEAEC41C" box="[267,367,585,606]" italics="true" pageId="11" pageNumber="12">Kootenaia</emphasis>
</taxonomicName>
</th>
<td id="76E6B82CFFE806DCFE42FDA9FD43C41C" box="[511,642,585,606]" gridcol="1" gridrow="1" pageId="11" pageNumber="12">
<emphasis id="B9730F00FFE8F928FD81FDA9FD85C41C" box="[572,580,585,606]" italics="true" pageId="11" pageNumber="12">­</emphasis>
</td>
</tr>
<tr id="3537D150FFE806DCFEB5FD94FB30C4CB" box="[264,1265,628,649]" gridrow="2" pageId="11" pageNumber="12" rowspan-3="1" rowspan-4="1">
<th id="76E6B82CFFE806DCFEB5FD94FE4DC4CB" box="[264,396,628,649]" gridcol="0" gridrow="2" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FEB6FD94FEA3C4CB" box="[267,354,628,649]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FEB6FD94FEA3C4CB" box="[267,354,628,649]" italics="true" pageId="11" pageNumber="12">Zacoleus</emphasis>
</taxonomicName>
</th>
<td id="76E6B82CFFE806DCFE42FD94FD43C4CB" box="[511,642,628,649]" gridcol="1" gridrow="2" pageId="11" pageNumber="12">0.185 (0.014)</td>
<td id="76E6B82CFFE806DCFD6EFD94FC94C4CB" box="[723,853,628,649]" gridcol="2" gridrow="2" pageId="11" pageNumber="12">­</td>
</tr>
<tr id="3537D150FFE806DCFEB5FD7EFB30C4F1" box="[264,1265,670,691]" gridrow="3" pageId="11" pageNumber="12" rowspan-4="1">
<th id="76E6B82CFFE806DCFEB5FD7EFE4DC4F1" box="[264,396,670,691]" gridcol="0" gridrow="3" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FEB6FD7EFEBFC4F1" box="[267,382,670,691]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FEB6FD7EFEBFC4F1" box="[267,382,670,691]" italics="true" pageId="11" pageNumber="12">Prophysaon</emphasis>
</taxonomicName>
</th>
<td id="76E6B82CFFE806DCFE42FD7EFD43C4F1" box="[511,642,670,691]" gridcol="1" gridrow="3" pageId="11" pageNumber="12">0.204 (0.013)</td>
<td id="76E6B82CFFE806DCFD6EFD7EFC94C4F1" box="[723,853,670,691]" gridcol="2" gridrow="3" pageId="11" pageNumber="12">0.203 (0.013)</td>
<td id="76E6B82CFFE806DCFC1AFD7EFBE8C4F1" box="[935,1065,670,691]" gridcol="3" gridrow="3" pageId="11" pageNumber="12">­</td>
</tr>
<tr id="3537D150FFE806DCFEB5FD28FB30C49C" box="[264,1265,712,734]" gridrow="4" pageId="11" pageNumber="12">
<th id="76E6B82CFFE806DCFEB5FD28FE4DC49C" box="[264,396,712,734]" gridcol="0" gridrow="4" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FEB6FD29FEB6C49C" box="[267,375,713,734]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FEB6FD29FEB6C49C" box="[267,375,713,734]" italics="true" pageId="11" pageNumber="12">Hemphillia</emphasis>
</taxonomicName>
</th>
<td id="76E6B82CFFE806DCFE42FD28FD43C49C" box="[511,642,712,734]" gridcol="1" gridrow="4" pageId="11" pageNumber="12">0.206 (0.012)</td>
<td id="76E6B82CFFE806DCFD6EFD28FC94C49C" box="[723,853,712,734]" gridcol="2" gridrow="4" pageId="11" pageNumber="12">0.184 (0.012)</td>
<td id="76E6B82CFFE806DCFC1AFD28FBE8C49C" box="[935,1065,712,734]" gridcol="3" gridrow="4" pageId="11" pageNumber="12">0.227 (0.012)</td>
<td id="76E6B82CFFE806DCFB38FD28FB30C49C" box="[1157,1265,712,734]" gridcol="4" gridrow="4" pageId="11" pageNumber="12">­</td>
</tr>
<tr id="3537D150FFE806DCFEB5FCA0FB30C518" box="[264,1265,832,858]" gridrow="5" pageId="11" pageNumber="12" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1">
<th id="76E6B82CFFE806DCFEB5FCA0FE4DC518" box="[264,396,832,858]" gridcol="0" gridrow="5" pageId="11" pageNumber="12">Discussion</th>
</tr>
</table>
</paragraph>
<paragraph id="8BB8D312FFE8F928FEB5FC6BFB71C2D7" blockId="11.[264,1323,907,1893]" pageId="11" pageNumber="12">
The new taxon deserves both new genus and new species designations on the basis of the absence of a free epiphallus alone. No other arionid even approaches this condition. Based upon the well­separated buccal and tentacular retractors and the general body appearance, 
<taxonomicName id="4C07A891FFE8F928FEB5FBE3FE43C25E" box="[264,386,1027,1052]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FEB5FBE3FE43C25E" box="[264,386,1027,1052]" italics="true" pageId="11" pageNumber="12">Kootenaia</emphasis>
</taxonomicName>
appears to be more closely allied with 
<taxonomicName id="4C07A891FFE8F928FCC8FBE3FBC3C25E" box="[885,1026,1027,1052]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FCC8FBE3FBC3C25E" box="[885,1026,1027,1052]" italics="true" pageId="11" pageNumber="12">Prophysaon</emphasis>
</taxonomicName>
than either 
<taxonomicName id="4C07A891FFE8F928FB21FBE3FAC7C25E" box="[1180,1286,1027,1052]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FB21FBE3FAC7C25E" box="[1180,1286,1027,1052]" italics="true" pageId="11" pageNumber="12">Zacoleus</emphasis>
</taxonomicName>
or 
<taxonomicName id="4C07A891FFE8F928FEB5FBCBFE4DC206" box="[264,396,1067,1092]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FEB5FBCBFE4DC206" box="[264,396,1067,1092]" italics="true" pageId="11" pageNumber="12">Hemphillia</emphasis>
</taxonomicName>
. In many 
<taxonomicName id="4C07A891FFE8F928FDB4FBCBFD57C206" box="[521,662,1067,1092]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FDB4FBCBFD57C206" box="[521,662,1067,1092]" italics="true" pageId="11" pageNumber="12">Prophysaon</emphasis>
</taxonomicName>
species, the penis is reduced to a penial loop but none possesses a penial retractor muscle and all possess a well­developed epiphallus suggesting that the relationship between 
<taxonomicName id="4C07A891FFE8F928FDDCFB9BFD1AC2D6" box="[609,731,1147,1172]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FDDCFB9BFD1AC2D6" box="[609,731,1147,1172]" italics="true" pageId="11" pageNumber="12">Kootenaia</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C07A891FFE8F928FCA9FB9BFC61C2D6" box="[788,928,1147,1172]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FCA9FB9BFC61C2D6" box="[788,928,1147,1172]" italics="true" pageId="11" pageNumber="12">Prophysaon</emphasis>
</taxonomicName>
is not especially close.
</paragraph>
<paragraph id="8BB8D312FFE8F928FE85FB43FC1FC3EF" blockId="11.[264,1323,907,1893]" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FE85FB43FE63C2FE" box="[312,418,1187,1212]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FE85FB43FE63C2FE" box="[312,418,1187,1212]" italics="true" pageId="11" pageNumber="12">Zacoleus</emphasis>
</taxonomicName>
possesses converging retractors, a well­developed penis without a penial retractor, and a very large spermathecal duct. When these characteristics are considered together, they suggest that 
<taxonomicName id="4C07A891FFE8F928FDF8FB13FD6EC34E" box="[581,687,1267,1292]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FDF8FB13FD6EC34E" box="[581,687,1267,1292]" italics="true" pageId="11" pageNumber="12">Zacoleus</emphasis>
</taxonomicName>
is even more distantly related to 
<taxonomicName id="4C07A891FFE8F928FB84FB13FB72C34E" box="[1081,1203,1267,1292]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FB84FB13FB72C34E" box="[1081,1203,1267,1292]" italics="true" pageId="11" pageNumber="12">Kootenaia</emphasis>
</taxonomicName>
than 
<emphasis id="B9730F00FFE8F928FB4BFB13FEB3C376" italics="true" pageId="11" pageNumber="12">
<taxonomicName id="4C07A891FFE8F928FB4BFB13FEAFC376" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">Prophysaon</taxonomicName>
.
</emphasis>
Even more distantly related is 
<taxonomicName id="4C07A891FFE8F928FD57FAFBFCAFC376" box="[746,878,1307,1332]" class="Gastropoda" family="Arionidae" genus="Hemphillia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FD57FAFBFCAFC376" box="[746,878,1307,1332]" italics="true" pageId="11" pageNumber="12">Hemphillia</emphasis>
</taxonomicName>
with its exposed horny shell, its visceral hump, its converging buccal and tentacular retractors, its retentor muscle and its well­developed penis. 
<bibRefCitation id="EF96AEE3FFE8F928FDAAFA8BFD02C3C7" author="Pilsbry" box="[535,707,1387,1413]" pageId="11" pageNumber="14" refString="Pilsbry, H. A. (1948) Land Mollusca of North America (north of Mexico), volume 2 part 2. Monographs of the Academy of Natural Sciences of Philadelphia (3), Philadelphia, 640 pp." type="book" year="1948">Pilsbry (1948)</bibRefCitation>
subdivided the 
<taxonomicName id="4C07A891FFE8F928FC3BFA8BFC3AC3C7" box="[902,1019,1387,1413]" class="Gastropoda" family="Arionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="family">Arionidae</taxonomicName>
into four subfamilies. In this scheme, 
<taxonomicName id="4C07A891FFE8F928FE1DFA73FDDBC3EE" box="[416,538,1427,1452]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FE1DFA73FDDBC3EE" box="[416,538,1427,1452]" italics="true" pageId="11" pageNumber="12">Kootenaia</emphasis>
</taxonomicName>
fits best in the subfamily Anadeninae.
</paragraph>
<paragraph id="8BB8D312FFE8F928FE85FA5BFCF9C127" blockId="11.[264,1323,907,1893]" pageId="11" pageNumber="12">
The molecular analyses using two mitochondrial genes and the anatomical data produce congruent topologies. Our new taxon does not fit into any of the genera studied here but represents a distinct group. In fact, the two trees based on anatomical and molecular data presented in 
<figureCitation id="133CCF97FFE8F928FE67F9D3FDE0C00F" box="[474,545,1587,1613]" captionStart="FIGURE 4" captionStartId="10.[264,367,1244,1268]" captionTargetBox="[268,1318,449,1218]" captionTargetId="figure@10.[268,1318,449,1218]" captionTargetPageId="10" captionText="FIGURE 4: A. Maximum parsimony tree for representatives of the genera Hemphillia, Kootenaia, Prophysaon, and Zacoleus based on 15 anatomical characters. Bootstrap values (10,000 replicates) are given at the nodes. The scale bar indicated the number of changes. B. Bayesian phylogram for the same species based on combined fragments of mitochondrial COI DNA and LSU rRNA showing the 50 % majority­rule consensus of topologies sampled during the Bayesian search. The scale bar indicates the substitution rate according to the model of sequence evolution applied. The numbers at the branches are the percentages that the clade occurs among all sampled trees; i. e. the posterior probability of that clade." httpUri="https://zenodo.org/record/156666/files/figure.png" pageId="11" pageNumber="12">Fig. 4</figureCitation>
show a very high degree of concordance, which is rarely seen in similar studies. However, as with the anatomical data, the molecular analyses cannot unambiguously solve the problem of sister group relationships. When using 
<taxonomicName id="4C07A891FFE8F928FB23F964FAC9C0DF" box="[1182,1288,1668,1693]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FB23F964FAC9C0DF" box="[1182,1288,1668,1693]" italics="true" pageId="11" pageNumber="12">Zacoleus</emphasis>
</taxonomicName>
as outgroup, Bayesian inference suggests that the genus 
<taxonomicName id="4C07A891FFE8F928FC28F94CFBE3C087" box="[917,1058,1708,1733]" class="Gastropoda" family="Arionidae" genus="Prophysaon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FC28F94CFBE3C087" box="[917,1058,1708,1733]" italics="true" pageId="11" pageNumber="12">Prophysaon</emphasis>
</taxonomicName>
is the sister group of 
<taxonomicName id="4C07A891FFE8F928FEB5F934FE43C0AF" box="[264,386,1748,1773]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FEB5F934FE43C0AF" box="[264,386,1748,1773]" italics="true" pageId="11" pageNumber="12">Kootenaia</emphasis>
</taxonomicName>
(see 
<figureCitation id="133CCF97FFE8F928FE7FF933FDC8C0AF" box="[450,521,1747,1773]" captionStart="FIGURE 4" captionStartId="10.[264,367,1244,1268]" captionTargetBox="[268,1318,449,1218]" captionTargetId="figure@10.[268,1318,449,1218]" captionTargetPageId="10" captionText="FIGURE 4: A. Maximum parsimony tree for representatives of the genera Hemphillia, Kootenaia, Prophysaon, and Zacoleus based on 15 anatomical characters. Bootstrap values (10,000 replicates) are given at the nodes. The scale bar indicated the number of changes. B. Bayesian phylogram for the same species based on combined fragments of mitochondrial COI DNA and LSU rRNA showing the 50 % majority­rule consensus of topologies sampled during the Bayesian search. The scale bar indicates the substitution rate according to the model of sequence evolution applied. The numbers at the branches are the percentages that the clade occurs among all sampled trees; i. e. the posterior probability of that clade." httpUri="https://zenodo.org/record/156666/files/figure.png" pageId="11" pageNumber="12">Fig. 4</figureCitation>
B). But based on distance data, the genus 
<taxonomicName id="4C07A891FFE8F928FC4AF934FBB0C0AF" box="[1015,1137,1748,1773]" class="Gastropoda" family="Arionidae" genus="Kootenaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FC4AF934FBB0C0AF" box="[1015,1137,1748,1773]" italics="true" pageId="11" pageNumber="12">Kootenaia</emphasis>
</taxonomicName>
would be more similar to 
<taxonomicName id="4C07A891FFE8F928FE39F91CFE2FC157" box="[388,494,1788,1813]" class="Gastropoda" family="Arionidae" genus="Zacoleus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="12" phylum="Mollusca" rank="genus">
<emphasis id="B9730F00FFE8F928FE39F91CFE2FC157" box="[388,494,1788,1813]" italics="true" pageId="11" pageNumber="12">Zacoleus</emphasis>
</taxonomicName>
(
<tableCitation id="C685E6A9FFE8F928FE42F91BFD9DC157" box="[511,604,1787,1813]" captionStart="TABLE 2" captionStartId="11.[264,352,403,427]" captionTargetBox="[264,1265,543,858]" captionTargetPageId="11" captionText="TABLE 2. Pairwise K 2 P­distances among the genera Kootenaia, Zacoleus, Prophysaon, and Hemphillia for the combined COI and LSU rRNA sequences. Standard errors (based on 1000 bootstrap replications) are given in parenthesis." httpUri="http://table.plazi.org/id/DF78839AFFE8F928FEB5FE73FD13C7BB" pageId="11" pageNumber="12" tableUuid="DF78839AFFE8F928FEB5FE73FD13C7BB">Table 2</tableCitation>
). It appears as if future studies with more data and/or more taxa would be necessary to better address the problem of phylogenetic relationships among arionid taxa from the Pacific Northwest.
</paragraph>
</subSubSection>
</treatment>
</document>