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<mods:title id="B5F3DB78FA69DA778F54C513211B707D">The unseen diversity of the semi-aquatic earthworms of the genus Sparganophilus (Oligochaeta: Sparganophilidae) from the Southeastern Appalachian Piedmont</mods:title>
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<mods:namePart id="AA3CA252C0059994634BD941F69E0170">Carrera-Martínez, Roberto</mods:namePart>
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<mods:affiliation id="55A57B9292074C247E517B2FB6AC7190">Warnell School of Forestry and Natural Resources, University of Georgia, Athens, GA, USA & Current address: Department of Integrative Biology, University of Wisconsin-Madison, Madison, WI, USA</mods:affiliation>
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<mods:namePart id="1ACC15A8BF9F5AE83F16CCD7294469F3">Taylor, Melanie K.</mods:namePart>
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<mods:affiliation id="C847C95DEEC63E0F8500BB9ABA48B858">Southern Research Station, US Forest Service, Athens, GA, USA & Odum School of Ecology, University of Georgia, Athens, GA, USA</mods:affiliation>
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<mods:namePart id="1E48905FD205AE14C104AFD796EA9F97">Jones, Daniel</mods:namePart>
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<mods:affiliation id="010ACFCE88375D4A25C0BE901BEACA07">Biological and Environmental Sciences, Georgia College & State University, Milledgeville, GA, USA</mods:affiliation>
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<mods:namePart id="63D54EA66333AD65F1900454D088BB18">Schoville, Sean D.</mods:namePart>
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<mods:affiliation id="7A71A2696890205A486568D0F21A9127">Department of Entomology, University of Wisconsin-Madison, Madison, WI, USA</mods:affiliation>
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<mods:namePart id="7A450A07C1A7E18F7AC8F6B835DC5046">Snyder, Bruce A.</mods:namePart>
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<mods:affiliation id="D8B2939EFE43E43D6B4CF70DEDE05E54">Biological and Environmental Sciences, Georgia College & State University, Milledgeville, GA, USA</mods:affiliation>
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<mods:namePart id="FF19D69A5D2235811C93FE40DAAE60D1">Callaham, Mac A.</mods:namePart>
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<mods:affiliation id="C80132150FED9A75394DFEA84F970316">Southern Research Station, US Forest Service, Athens, GA, USA</mods:affiliation>
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<mods:date id="B242C10519E13DE7FD5F1F87F23B1D5B">2025</mods:date>
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<mods:number id="8FE7DD5FAE3A51327F61C2BF0C8B5C73">2025-02-19</mods:number>
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<paragraph id="8BB336EAB042A648FF20B0ADE069B1A1" blockId="15.[151,956,474,573]" box="[151,794,474,501]" pageId="15" pageNumber="397">
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<taxonomicName id="4C0C4D69B042A648FF20B0ADE1C5B1A1" authority="Carrera-Martinez" authorityName="Carrera-Martinez" authorityYear="2025" box="[151,694,474,501]" class="Clitellata" family="Sparganophilidae" genus="Sparganophilus" kingdom="Animalia" order="Crassiclitellata" pageId="15" pageNumber="397" phylum="Annelida" rank="species" species="borgesae" status="sp. nov.">
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<emphasis id="B978EAF8B042A648FF20B0ADE2B8B1A1" bold="true" box="[151,459,474,501]" italics="true" pageId="15" pageNumber="397">Sparganophilus borgesae</emphasis>
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Carrera-Martínez
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<taxonomicNameLabel id="A24B5783B042A648FD0AB0ACE069B1A1" box="[701,794,475,501]" pageId="15" pageNumber="397" rank="species">sp. nov.</taxonomicNameLabel>
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</emphasis>
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<figureCitation id="13372A6FB042A648FF20B354E246B269" box="[151,309,547,573]" captionStart="FIGURE 7" captionStartId="16.[152,255,1265,1289]" captionTargetBox="[817,1080,396,1176]" captionTargetId="graphics-165@16.[832,1079,396,1159]" captionTargetPageId="16" captionText="FIGURE 7. Sparganophilus borgesae sp. nov. (A) external ventral view, (B) spermathecae, (C) setal formula. Sparganophilus sp02. (D) external ventral view, (E) spermathecae, (F) setal formula. Abbreviations and symbols points to segments with, ♀: ovipores, P: prostatic pores, ♂: male pores, N: segment with first nephridiopore, GP: genital pad, GT: genital tumescences." figureDoi="http://doi.org/10.5281/zenodo.14897294" httpUri="https://zenodo.org/record/14897294/files/figure.png" pageId="15" pageNumber="397">Figures 7A–C</figureCitation>
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<subSubSection id="C3166561B042A648FF20B31CE6FEB2FD" pageId="15" pageNumber="397" type="materials_examined">
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<paragraph id="8BB336EAB042A648FF20B31CE6FEB2FD" blockId="15.[151,1437,618,2013]" pageId="15" pageNumber="397">
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<materialsCitation id="3B643CB7B042A648FF20B31CE6FEB2FD" collectingDate="2017-07-14" collectionCode="R" collectorName="Carrera-Martinez & M. K. Taylor" country="Georgia" county="Greene Co." latitude="33.74306" location="Falling Creek" longLatPrecision="1" longitude="-83.2523" municipality="Scull Shoals Experimental Forest" pageId="15" pageNumber="397" specimenCount="1" specimenCount-adult="1" typeStatus="holotype">
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<emphasis id="B978EAF8B042A648FF20B31CE2C3B2D0" bold="true" box="[151,432,618,645]" pageId="15" pageNumber="397">
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<typeStatus id="54B78848B042A648FF20B31CE27BB2D1" box="[151,264,619,645]" pageId="15" pageNumber="397" type="holotype">Holotype</typeStatus>
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: GTIC-11490;
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</emphasis>
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<specimenCount id="9D0AFD63B042A648FE02B31CE150B2D1" box="[437,547,619,645]" count="1" pageId="15" pageNumber="397" type="adult">One adult</specimenCount>
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, unnamed first order tributary to
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,
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<collectingMunicipality id="6BD7AC90B042A648FB90B31CE6EBB2D1" box="[1063,1432,619,645]" pageId="15" pageNumber="397">Scull Shoals Experimental Forest</collectingMunicipality>
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,
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<collectingCounty id="62D24E66B042A648FF20B3F8E26AB2FD" box="[151,281,655,681]" pageId="15" pageNumber="397">Greene Co.</collectingCounty>
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,
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<collectingCountry id="F31B767AB042A648FE92B3F8E2F1B2FD" box="[293,386,655,681]" name="Georgia" pageId="15" pageNumber="397">Georgia</collectingCountry>
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, USA. Coll.
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<collectionCode id="ED1DAE2FB042A648FDA6B3F8E154B2FD" box="[529,551,655,681]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="15" pageNumber="397">R</collectionCode>
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.
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<collectorName id="26F9533CB042A648FD86B3F8E185B2FD" box="[561,758,655,681]" pageId="15" pageNumber="397">Carrera-Martínez</collectorName>
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, &
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<collectorName id="26F9533CB042A648FCA9B3F8E0D8B2FD" box="[798,939,655,681]" pageId="15" pageNumber="397">M.K. Taylor</collectorName>
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,
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<collectingDate id="EFF6E9C2B042A648FC01B3F8E738B2FD" box="[950,1099,655,681]" pageId="15" pageNumber="397" value="2017-07-14">14.JUL.2017</collectingDate>
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. (
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,
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<geoCoordinate id="EE38502DB042A648FB41B3F9E6F7B2FD" box="[1270,1412,654,681]" degrees="83.25230" direction="west" orientation="longitude" pageId="15" pageNumber="397" precision="1" value="-83.2523">83.25230°W</geoCoordinate>
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)
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<subSubSection id="C3166561B042A648FF70B3C4E2E8B2A5" pageId="15" pageNumber="397" type="etymology">
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<paragraph id="8BB336EAB042A648FF70B3C4E2E8B2A5" blockId="15.[151,1437,618,2013]" pageId="15" pageNumber="397">
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<emphasis id="B978EAF8B042A648FF70B3C4E23BB299" bold="true" box="[199,328,691,717]" pageId="15" pageNumber="397">Etymology</emphasis>
|
||||
: Named after Dr. Sonia Borges, pioneer of Caribbean earthworm taxonomy and who introduced the first author to the field.
|
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</paragraph>
|
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</subSubSection>
|
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<subSubSection id="C3166561B042A648FF70B38DE39EB445" pageId="15" pageNumber="397" type="diagnosis">
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<paragraph id="8BB336EAB042A648FF70B38DE39EB445" blockId="15.[151,1437,618,2013]" pageId="15" pageNumber="397">
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<emphasis id="B978EAF8B042A648FF70B38DE249B340" bold="true" box="[199,314,762,788]" pageId="15" pageNumber="397">Diagnosis</emphasis>
|
||||
: Length
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<quantity id="4CF49B0FB042A648FE14B38DE285B341" box="[419,502,762,789]" metricMagnitude="-2" metricUnit="m" metricValue="7.5" pageId="15" pageNumber="397" unit="mm" value="75.0">75 mm</quantity>
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. Dorsal pores rudimentary. Clitellum saddle-shaped in xv–xxvi, ventral limit in
|
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<emphasis id="B978EAF8B042A648FA30B38CE6E6B341" box="[1415,1429,763,789]" italics="true" pageId="15" pageNumber="397">b</emphasis>
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. Tubercula pubertatis continuous in xvii–xxii, band-shaped, ventral limit dorsal to
|
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<emphasis id="B978EAF8B042A648FB85B268E734B36D" box="[1074,1095,799,825]" italics="true" pageId="15" pageNumber="397">b.</emphasis>
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Setae closely paired:
|
||||
<emphasis id="B978EAF8B042A648FA88B268E628B36D" box="[1343,1371,799,825]" italics="true" pageId="15" pageNumber="397">ab</emphasis>
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=
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<emphasis id="B978EAF8B042A648FACCB268E6E6B36D" box="[1403,1429,799,825]" italics="true" pageId="15" pageNumber="397">cd</emphasis>
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,
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<emphasis id="B978EAF8B042A648FF20B234E3C0B308" box="[151,179,835,860]" italics="true" pageId="15" pageNumber="397">aa</emphasis>
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<
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<emphasis id="B978EAF8B042A648FF66B234E398B309" box="[209,235,835,861]" italics="true" pageId="15" pageNumber="397">bc</emphasis>
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<
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||||
<emphasis id="B978EAF8B042A648FEBFB234E257B309" box="[264,292,835,861]" italics="true" pageId="15" pageNumber="397">dd</emphasis>
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||||
. Spermathecal pores in 6/7–8/9, inconspicuous. Pores of prostate-like glands in xxiii–xxiv. Secondary annulation absent. Genital tumescences surrounding
|
||||
<emphasis id="B978EAF8B042A648FD44B210E07CB3D5" box="[755,783,871,897]" italics="true" pageId="15" pageNumber="397">ab</emphasis>
|
||||
weakly developed in xvii–xxv. Genital pad band-shaped in xxvi. Intestinal origin in ix. One pair of spermathecae per segment in vii–ix, spermathecal structure moderately differentiated into ampulla and duct, smooth, duct tubular, three times as long as the globular ampulla. Prostate-like glands in xxiii–xxiv and auxiliary prostate-like glands absent. First nephridia in xiii, tubular bladder, wide u-shaped vesicle.
|
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</paragraph>
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</subSubSection>
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<subSubSection id="C3166561B042A648FF70B56DE1A9B7C1" pageId="15" pageNumber="397" type="description">
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<paragraph id="8BB336EAB042A648FF70B56DE660B595" blockId="15.[151,1437,618,2013]" pageId="15" pageNumber="397">
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<emphasis id="B978EAF8B042A648FF70B56DE222B460" bold="true" box="[199,337,1050,1076]" pageId="15" pageNumber="397">Description</emphasis>
|
||||
: Length of unamputated preserved specimen
|
||||
<quantity id="4CF49B0FB042A648FCDFB56DE0CDB461" box="[872,958,1050,1077]" metricMagnitude="-2" metricUnit="m" metricValue="7.5" pageId="15" pageNumber="397" unit="mm" value="75.0">75 mm</quantity>
|
||||
. Diameter in x
|
||||
<quantity id="4CF49B0FB042A648FBCFB56DE7A6B461" box="[1144,1237,1050,1077]" metricMagnitude="-3" metricUnit="m" metricValue="2.5" pageId="15" pageNumber="397" unit="mm" value="2.5">2.5 mm</quantity>
|
||||
; in the clitellum,
|
||||
<quantity id="4CF49B0FB042A648FF20B548E387B40D" box="[151,244,1087,1113]" metricMagnitude="-3" metricUnit="m" metricValue="3.3" pageId="15" pageNumber="397" unit="mm" value="3.3">3.3 mm</quantity>
|
||||
. Number of segments, 184. Body ovoid in cross-section anteriorly, ventrally flattened at the clitellum, quadrangular in cross-section posterior to the clitellum. Preserved specimens with a pale whitish coloration, clitellum pale. Cuticle with a green and blue iridescence. Prostomium zygolobic. Dorsal pores rudimentary, discontinuous throughout. Spermathecal pores in intersegmental furrows 6/7–8/9, inconspicuous. Female pores in xiv, anterior to
|
||||
<emphasis id="B978EAF8B042A648FF20B5B8E3C0B4BD" box="[151,179,1231,1257]" italics="true" pageId="15" pageNumber="397">ab</emphasis>
|
||||
, minute but visible. Male pores lateral to
|
||||
<emphasis id="B978EAF8B042A648FDC9B5B8E1FFB4BD" box="[638,652,1231,1257]" italics="true" pageId="15" pageNumber="397">b</emphasis>
|
||||
in xix, pre-setal, at the dorsal margin of tubercula pubertatis. Clitellum in xv–xxvi saddle-shaped, ventral limit in
|
||||
<emphasis id="B978EAF8B042A648FDC2B584E1F0B559" box="[629,643,1267,1293]" italics="true" pageId="15" pageNumber="397">b</emphasis>
|
||||
. Tubercula pubertatis continuous xvii–xxii, band-shaped, ventral limit dorsal to setae line
|
||||
<emphasis id="B978EAF8B042A648FEDDB460E20CB565" box="[362,383,1303,1329]" italics="true" pageId="15" pageNumber="397">b.</emphasis>
|
||||
Prostate-like pores in xxiii–xxiv next to
|
||||
<emphasis id="B978EAF8B042A648FC88B460E027B565" box="[831,852,1303,1329]" italics="true" pageId="15" pageNumber="397">b.</emphasis>
|
||||
Nephridiopores anterior to
|
||||
<emphasis id="B978EAF8B042A648FB31B460E7D1B565" box="[1158,1186,1303,1329]" italics="true" pageId="15" pageNumber="397">ab</emphasis>
|
||||
, closer to
|
||||
<emphasis id="B978EAF8B042A648FAA5B460E653B564" box="[1298,1312,1303,1328]" italics="true" pageId="15" pageNumber="397">a</emphasis>
|
||||
, starting in segment xiii, minute but visible. First setae in ii, lumbricine, setae closely paired. In x
|
||||
<emphasis id="B978EAF8B042A648FBE5B44CE672B501" box="[1106,1281,1339,1365]" italics="true" pageId="15" pageNumber="397">aa:ab:bc:cd:dd</emphasis>
|
||||
= 6.50:1.00:7.50:1.00:10.00, in xxx = 6.50:1.00:8.00:1.00:10.00. Secondary annulation absent. Genital tumescences surrounding
|
||||
<emphasis id="B978EAF8B042A648FF20B4F4E3C0B5C9" box="[151,179,1411,1437]" italics="true" pageId="15" pageNumber="397">ab</emphasis>
|
||||
in xvii–xxv, weakly developed. Genital pad band-shaped in xxvi extending to setal line
|
||||
<emphasis id="B978EAF8B042A648FB2FB4F4E7D5B5C9" box="[1176,1190,1411,1437]" italics="true" pageId="15" pageNumber="397">b</emphasis>
|
||||
. Post-setal dark spots visible just ventral to
|
||||
<emphasis id="B978EAF8B042A648FE3DB4D0E2E5B594" box="[394,406,1447,1472]" italics="true" pageId="15" pageNumber="397">c</emphasis>
|
||||
, in a regular line in i–xiv. One dorsal depression in the equator of peristomium.
|
||||
</paragraph>
|
||||
<paragraph id="8BB336EAB042A648FF70B4BCE1A9B7C1" blockId="15.[151,1437,618,2013]" pageId="15" pageNumber="397">
|
||||
Septa 6/7–8/9 only slightly thickened. Pharynx ends in segment iv, joining a small esophagus. Intestine originates abruptly in ix. Intestinal caeca, typhlosole, calciferous glands, and gizzard absent. One pair of spermathecae in vii–ix, without diverticulum. Spermathecal structure moderately differentiated into ampulla and duct, all similar size. Duct about half the length of the ampulla, tubular, wide. Ampulla elongated, globular, rugose laterally, tip digitiform. Spermathecae located between a weak muscular fiber originated anteriorly in the intersegmental section of the esophagus and connecting posteriorly to the intersegmental-septum joint. Ovaries in xiii, free, string-like. Oviduct connects in segment xiv, anterior to
|
||||
<emphasis id="B978EAF8B042A648FD2BB7D4E1C2B6E8" box="[668,689,1699,1724]" italics="true" pageId="15" pageNumber="397">a.</emphasis>
|
||||
Testes free in x and xi. Epididymis small. Seminal vesicles, free in xi, and xii, large and restricted to their segment. Vas deferens within muscular body wall. Tubular, prostate-like glands in xxiii–xxiv, long, slender, and coiled. Auxiliary prostate-like glands absent. One pair of holonephridia per segment, starting at xiii. Holonephridia tubular, bladder composed of highly coiled tubules dorsally, ventrally with a wide u-shaped vesicle, all same size. Holonephridia of xiv attached to 13/14 septa, without vesicle. Paired moniliform hearts in vii–xi; vii smaller than the rest. Dorsal and ventral vessel complete, and simple. Paired lateral vessel present, joining dorsal vessel in xiv, filiform.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
<subSubSection id="C3166561B042A657FF70B6E8E231B486" lastPageId="16" lastPageNumber="398" pageId="15" pageNumber="397" type="discussion">
|
||||
<paragraph id="8BB336EAB042A657FF70B6E8E203B131" blockId="15.[151,1437,618,2013]" lastBlockId="16.[151,1437,151,357]" lastPageId="16" lastPageNumber="398" pageId="15" pageNumber="397">
|
||||
<emphasis id="B978EAF8B042A648FF70B6E8E247B7ED" bold="true" box="[199,308,1951,1977]" pageId="15" pageNumber="397">Remarks</emphasis>
|
||||
:
|
||||
<taxonomicName id="4C0C4D69B042A648FEF4B6E8E113B7ED" authorityName="Carrera-Martínez & Taylor & Jones & Schoville & Snyder & Callaham" authorityYear="2025" box="[323,608,1951,1977]" class="Clitellata" family="Sparganophilidae" genus="Sparganophilus" kingdom="Animalia" order="Crassiclitellata" pageId="15" pageNumber="397" phylum="Annelida" rank="species" species="borgesae" status="sp. nov.">
|
||||
<emphasis id="B978EAF8B042A648FEF4B6E8E113B7ED" box="[323,608,1951,1977]" italics="true" pageId="15" pageNumber="397">Sparganophilus borgesae</emphasis>
|
||||
</taxonomicName>
|
||||
<emphasis id="B978EAF8B042A648FDD0B6E8E1B2B7ED" bold="true" box="[615,705,1951,1977]" pageId="15" pageNumber="397">
|
||||
<taxonomicNameLabel id="A24B5783B042A648FDD0B6E8E1B2B7ED" box="[615,705,1951,1977]" pageId="15" pageNumber="397" rank="species">sp. nov.</taxonomicNameLabel>
|
||||
</emphasis>
|
||||
differs from all other known
|
||||
<taxonomicName id="4C0C4D69B042A648FBBBB6E8E7CFB7ED" authorityName="Benham" authorityYear="1892" box="[1036,1212,1951,1977]" class="Clitellata" family="Sparganophilidae" genus="Sparganophilus" kingdom="Animalia" order="Crassiclitellata" pageId="15" pageNumber="397" phylum="Annelida" rank="genus">
|
||||
<emphasis id="B978EAF8B042A648FBBBB6E8E7CFB7ED" box="[1036,1212,1951,1977]" italics="true" pageId="15" pageNumber="397">Sparganophilus</emphasis>
|
||||
</taxonomicName>
|
||||
by having only two pairs of well-developed prostates in xxiii–xxiv, rudimentary dorsal pores, presence of a genital pad, and a dorsal pigmented pit on the equator of the prostomium. We decided to describe and name this species, despite having only a single specimen after a year of collection at this site. Furthermore, previous sampling efforts by
|
||||
<bibRefCitation id="EF9D4B1BB05DA657FB67B1CCE6EFB081" author="Ikeda, H. & Callaham, M. A. & Shefferson, R. P. & Wenk, E. S. & Fragoso, C." box="[1232,1436,187,213]" pageId="16" pageNumber="398" pagination="1373 - 1382" refId="ref19465" refString="Ikeda, H., Callaham, M. A., Shefferson, R. P., Wenk, E. S. & Fragoso, C. (2020) A comparison of latitudinal species diversity patterns between riverine and terrestrial earthworms from the North American temperate zone. Journal of Biogeography, 47, 1373-1382. https://doi.org/10.1111/jbi.13826" type="journal article" year="2020">
|
||||
Ikeda
|
||||
<emphasis id="B978EAF8B05DA657FAA3B1CCE636B081" box="[1300,1349,187,213]" italics="true" pageId="16" pageNumber="398">et al</emphasis>
|
||||
. (2020)
|
||||
</bibRefCitation>
|
||||
in and around this area did not yield any match to this species, suggesting that
|
||||
<taxonomicName id="4C0C4D69B05DA657FBBCB1A8E7FEB0AD" authorityName="Carrera-Martínez & Taylor & Jones & Schoville & Snyder & Callaham" authorityYear="2025" box="[1035,1165,223,249]" class="Clitellata" family="Sparganophilidae" genus="Sparganophilus" kingdom="Animalia" order="Crassiclitellata" pageId="16" pageNumber="398" phylum="Annelida" rank="species" species="borgesae">
|
||||
<emphasis id="B978EAF8B05DA657FBBCB1A8E7FEB0AD" box="[1035,1165,223,249]" italics="true" pageId="16" pageNumber="398">S. borgesae</emphasis>
|
||||
</taxonomicName>
|
||||
might be a rare species. Given that no additional specimens were found, that 28S, 16S, and COI genetic distance are large and having distinct morphological differences, we followed
|
||||
<bibRefCitation id="EF9D4B1BB05DA657FD02B050E0B7B115" author="Fragoso, C. & Rojas, P." box="[693,964,295,321]" pageId="16" pageNumber="398" pagination="483 - 502" refId="ref19282" refString="Fragoso, C. & Rojas, P. (2019) More new Diplotrema earthworm species from southern Mexico (Annelida, Crassiclitellata, Acanthodrilidae, Acanthodrilinae). Zootaxa, 4688 (4), 483-502. https://doi.org/10.11646/zootaxa.4688.4.2" type="journal article" year="2019">Fragoso & Rojas (2019)</bibRefCitation>
|
||||
recommendations on describing singleton earthworm species.
|
||||
</paragraph>
|
||||
<paragraph id="8BB336EAB05DA657FB85B36DE231B486" blockId="16.[163,1393,400,1234]" pageId="16" pageNumber="398">
|
||||
v Exv GT C xv F d xx c c xx xxv a xxv a b xxx
|
||||
<quantity id="4CF49B0FB05DA657FC29B5D2E231B486" metricMagnitude="-3" metricUnit="m" metricValue="2.0" pageId="16" pageNumber="398" unit="mm" value="2.0">2 mm 2 mm</quantity>
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
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