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<mods:title id="C357D0C24D42BEF20D9D8C042BD87F62">Taxonomy of North European Lumbricillus (Clitellata, Enchytraeidae)</mods:title>
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<treatment id="470CA82BA291357D92BC8D6861D65EE3" ID-GBIF-Taxon="134860662" LSID="urn:lsid:plazi:treatment:470CA82BA291357D92BC8D6861D65EE3" httpUri="http://treatment.plazi.org/id/470CA82BA291357D92BC8D6861D65EE3" lastPageId="11" lastPageNumber="26" pageId="9" pageNumber="24">
<subSubSection id="96EE51DBB0DD647EE907E9CB97309AE8" pageId="9" pageNumber="24" type="nomenclature">
<paragraph id="50BB47F3191378BD934797A9C5E7938B" pageId="9" pageNumber="24">
<taxonomicName id="17E7AAEA616880B69399D417D6058F6E" ID-CoL="4BRXM" authority="Claparede, 1861" authorityName="Claparede" authorityYear="1861" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus verrucosus" order="Enchytraeida" pageId="9" pageNumber="24" phylum="Annelida" rank="species" species="verrucosus">
Lumbricillus verrucosus (
<normalizedToken id="1FF1E034BF9098F2C74D6A842AF43FCB" originalValue="Claparède">Claparede</normalizedToken>
, 1861)
</taxonomicName>
Figs 3
<normalizedToken id="87CBA49477C2CD9FA1D2924589ECBAE9" originalValue="CD">C-D</normalizedToken>
, 6
</paragraph>
</subSubSection>
<subSubSection id="CECB17485162A18FD126A4D25453AC0E" pageId="9" pageNumber="24" type="reference_group">
<paragraph id="ABCF8EAA1C42B8CC6EF4E6CC5383F09D" pageId="9" pageNumber="24">
<taxonomicName id="59855BB8C067A3118120CB18E05ADD9D" class="Clitellata" family="Enchytraeidae" genus="Pachydrilus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pachydrilus verrucosus" order="Enchytraeida" pageId="9" pageNumber="24" phylum="Annelida" rank="species" species="verrucosus">Pachydrilus verrucosus</taxonomicName>
<normalizedToken id="16975A1FCDA4064A6058E49ABA715F5F" originalValue="Claparède">Claparede</normalizedToken>
, 1861: pp. 82-85, pl. I, figs 1-6;
</paragraph>
<paragraph id="DF093B2384B0CA7DAF08BBCE0FCDF315" pageId="9" pageNumber="24">
<taxonomicName id="F992014291A8D0A0E95B0DF70DA4559D" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus verrucosus" order="Enchytraeida" pageId="9" pageNumber="24" phylum="Annelida" rank="species" species="verrucosus">Lumbricillus verrucosus</taxonomicName>
;
<bibRefCitation id="8083B391E9826BB133D23C84CC26AFC9" author="Michaelsen, W" journalOrPublisher="Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten" pageId="47" pageNumber="62" url="https://doi.org/10.5962/bhl" year="1900">Michaelsen 1900</bibRefCitation>
: p. 80;
<bibRefCitation id="835D04BBE24B9049CA6D5D0B777B6152" author="Klinth, MJ" journalOrPublisher="Zoologica Scripta" pageId="46" pageNumber="61" pagination="96 - 110" title="Phylogeny and species delimitation of North European Lumbricillus (Clitellata, Enchytraeidae)." url="https://doi.org/10.1111/zsc.12187" volume="46" year="2017">Klinth et al. 2017</bibRefCitation>
.
</paragraph>
<paragraph id="8FBEFE51D5808FAADF167DCBA54B08FD" pageId="9" pageNumber="24">
<taxonomicName id="A56A458235180CC6AF0AEA8C885ED9C5" class="Clitellata" family="Enchytraeidae" form="verrucosus" genus="Pachydrilus" higherTaxonomySource="CoL" infraspecific-rank="forma" kingdom="Animalia" lsidName="Pachydrilus lineatus" order="Enchytraeida" pageId="9" pageNumber="24" phylum="Annelida" rank="form" species="lineatus">Pachydrilus lineatus forma verrucosus</taxonomicName>
;
<bibRefCitation id="ACC0C78D09DC50D230E69690D285F142" author="Cernosvitov, L" journalOrPublisher="Bulletin de l'Association Russe pour les Recherches Scientifiques a Prague" pageId="44" pageNumber="59" pagination="263 - 295" title="System der Enchytraeiden." volume="5" year="1937">
<normalizedToken id="CE37C195BD7C7889E2982915B6EBB8B4" originalValue="Černosvitov">Cernosvitov</normalizedToken>
1937
</bibRefCitation>
: p. 292.
</paragraph>
<paragraph id="3AC5EC3EB7D40BC041C90F614CAD51E8" pageId="9" pageNumber="24">
<taxonomicName id="5476646BEB8FBFA50436C1E9FBC02C1C" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="9" pageNumber="24" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
partim;
<bibRefCitation id="10679E78D98E125DD2D9C70B97AE28CA" pageId="9" pageNumber="24">Nielsen and Christensen 1959</bibRefCitation>
: pp. 100-102, figs 109-112.
</paragraph>
</subSubSection>
<subSubSection id="97A7B725B8C7BFC0BB107E032618266A" pageId="9" pageNumber="24" type="type material">
<paragraph id="998E674FC9524181315A76A99D037CE0" pageId="9" pageNumber="24">Type material.</paragraph>
<paragraph id="3562242FAA603DF9B7845E54476C9D89" pageId="9" pageNumber="24">
Typus amissus (Nomenclatura Oligochaetologica). Type locality: Sound of Sleat, Isle of Skye, Hebrides, United Kingdom (
<normalizedToken id="59652F4B497ED3210E96DDAA34DCAA52" originalValue="Claparède">Claparede</normalizedToken>
, 1861). We did not designate a neotype as we do not have material from the type locality.
</paragraph>
</subSubSection>
<subSubSection id="9C535B8078D2D9345ED0D43AB30B9FE2" pageId="9" pageNumber="24" type="materials_examined">
<paragraph id="46CED4E35699160518EC743F2B71312A" pageId="9" pageNumber="24">Material examined.</paragraph>
<paragraph id="73606257CFDFD4747EF971658778E9E3" pageId="9" pageNumber="24">SMNH 152826 (CE968), one mature specimen from Sweden, ZMBN 107919 (CE21479), ZMBN 107920 (CE21486), ZMBN 107921 (CE21490), ZMBN 107922 (CE21494), ZMBN 107924 (CE21811), ZMBN 107925 (CE21816) &amp; ZMBN 107926 (CE21821), seven mature specimens from Norway. For information on specimen collection localities and GenBank accession numbers see Appendix 1.</paragraph>
</subSubSection>
<subSubSection id="5817F0E216F18E919786F0E361E84A38" lastPageId="11" lastPageNumber="26" pageId="9" pageNumber="24" type="description">
<paragraph id="AF654A584FEE34B2649E96E2166FDE4C" pageId="9" pageNumber="24">Description.</paragraph>
<paragraph id="898C19486539CA0D42D1F5D41BCE4B63" lastPageId="10" lastPageNumber="25" pageId="9" pageNumber="24">
White to yellow worms. Length (fixed worms) more than 2.3-5.7 mm (amputated specimens), first 15 segments 2.3-3.4 mm long, width at clitellum 0.42-0.60 mm. More than 18-33 segments. Chaetae slightly sigmoid (Fig. 6A). Dorsal bundles
<pageBreakToken id="8B8900071DAFA2B96245D17D945846FB" pageId="10" pageNumber="25" start="start">with</pageBreakToken>
(2)3-5(6) chaetae anterior to clitellum, 2-4 chaetae in postclitellar segments. Ventral bundles with (2)3-6(7) chaetae anterior to clitellum, (2)3-4(5) chaetae posteriorly. Each
<normalizedToken id="0A36DD2F7F1A22B73D3CE0723380F159" originalValue="worms">worm's</normalizedToken>
longest measured chaetae 45-60
<normalizedToken id="6044DA54570D43B4241685A507EE25C1" originalValue="µm">µm</normalizedToken>
long, about 2.5
<normalizedToken id="FA8AB78CD576655856D81106FA1753A1" originalValue="µm">µm</normalizedToken>
wide. Clitellum extending over
<normalizedToken id="F25470DB04E8715BBAC6636387623E32" originalValue="XIIXIII">XII-XIII</normalizedToken>
. Head pore at 0/1. Epidermis with transverse rows of gland cells.
</paragraph>
<paragraph id="048BD3ED6DA26412E2C666982DADB8C1" pageId="10" pageNumber="25">
Coelomocytes in some specimens numerous, 10-25
<normalizedToken id="480115ED0F37E4EBF148E9F6DE1BBFEB" originalValue="µm">µm</normalizedToken>
long, round, oval or spindle-shaped, granulated. Paired pharyngeal glands present in IV, V and VI; each pair converging dorsally (Fig. 6B). Dorsal vessel originating in XIII. Nephridia observed in
<normalizedToken id="89E1F65F6884D31501493E087820AA48" originalValue="XIVXXV">XIV-XXV</normalizedToken>
, 75-120
<normalizedToken id="509C860DDFB776D3D4A942899CAAC8FC" originalValue="µm">µm</normalizedToken>
long. Anteseptale small, consisting of funnel only. Postseptale oval, tapering posteriorly into efferent duct. Brain twice as long as wide, with posterior incision.
</paragraph>
<paragraph id="82DF4DA33B9A28885BE3281BA8B6886A" pageId="10" pageNumber="25">
Male genitalia paired (Fig. 6D). Testes originating in XI, extending forwards into X, sometimes IX, with testis sacs forming regular club-shaped lobes. Sperm funnels in XI, 230-370
<normalizedToken id="8FBEE52408A16BC3E944DED864B0A010" originalValue="µm">µm</normalizedToken>
long, 125-175
<normalizedToken id="9F5C25883F21D3F0D5A7DD39D99718E9" originalValue="µm">µm</normalizedToken>
wide, making them about 1.5-2.5 times longer than wide, funnels tapering towards vasa deferentia. Most of vasa irregularly coiled in XII, 15-20
<normalizedToken id="88A8B2789629A177F27DC9C0115DA1FF" originalValue="µm">µm</normalizedToken>
wide. Penial bulbs round, 105-140
<normalizedToken id="E05363F4BECC144BE5B007FF1C2688D6" originalValue="µm">µm</normalizedToken>
in diameter. Ovaries in XII. One to five mature eggs present at a time.
</paragraph>
<paragraph id="D175BFA811D267C2DA03A32FA2F0D875" lastPageId="11" lastPageNumber="26" pageId="10" pageNumber="25">
Spermathecae (Figs 3
<normalizedToken id="186D3D56BD6C404B805088770D8C6C9F" originalValue="CD">C-D</normalizedToken>
, 6C) in V, spindle-shaped, without distinct ampulla. Ectal duct short, widening into ampulla. Ampulla with constriction midway to two thirds of the length, dividing it into two sections, the inner one of which connecting with oesophagus. Sperm filling lumen of ectal duct, heads of spermatozoa embedded in inner part of ampulla, sometimes also in outer part, forming aggregates. Spermathecae
<pageBreakToken id="86D5A3EB7584843A998C7AB779F0AECB" pageId="11" pageNumber="26" start="start">180</pageBreakToken>
-300
<normalizedToken id="CF3B27048EBD575865EAD98E0D63CB43" originalValue="µm">µm</normalizedToken>
long, 65-110
<normalizedToken id="A1A713F5CF92D238710DE133FAA3E529" originalValue="µm">µm</normalizedToken>
wide at widest part of ampulla. Gland cells surrounding ectal pore, forming compact mass, whole glandular body 95-180
<normalizedToken id="BDF8F5275A9E2B49D857A1960A0E2F0E" originalValue="µm">µm</normalizedToken>
in diameter at its widest part. One or two midventral subneural glands in
<normalizedToken id="8388DE47F0CF7683AFC4BDFA950BBECF" originalValue="XIIIXIV">XIII-XIV</normalizedToken>
, 90-125
<normalizedToken id="7FB00CC95AEBFF1D239E39EC136E85FF" originalValue="µm">µm</normalizedToken>
and 95-125
<normalizedToken id="5E76546F4359E28D2D78E8076CDD110C" originalValue="µm">µm</normalizedToken>
long, respectively; gland in XIV not observed in all specimens.
</paragraph>
<caption id="8EA41191DCF21F290900AB9162344C4A" pageId="11" pageNumber="26">
<paragraph id="56A76BE07D3393E6DB288A680A2DB42D" pageId="11" pageNumber="26">
Figure 6.
<taxonomicName id="8853DB4468C63BA032412C34629A245B" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus verrucosus" order="Enchytraeida" pageId="11" pageNumber="26" phylum="Annelida" rank="species" species="verrucosus">Lumbricillus verrucosus</taxonomicName>
. A Chaetal bundle B Anterior body C Spermatheca D Other genitalia. Abbreviations under general notes. Scale bars: 100
<normalizedToken id="F8B7554758F9BBE8C20DF9D017E7B3BF" originalValue="µm">µm</normalizedToken>
.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="4F8777AD3D021DC1FE2719ED1ED11D4C" pageId="11" pageNumber="26" type="distribution">
<paragraph id="FE638F8DE7F2CEF6C807507C8CCF27EC" pageId="11" pageNumber="26">Geographical distribution.</paragraph>
<paragraph id="DD17C4A385751CFCF5B80784A38BAFAA" pageId="11" pageNumber="26">
Originally described from the United Kingdom, now genetically identified from Norway and Sweden. The full extent of this
<normalizedToken id="68EDCE226AD47FD78020F378B447E583" originalValue="species">species'</normalizedToken>
distribution is difficult to ascertain since it was previously synonymized with
<taxonomicName id="9E4662391366F67BB792DA06C556E0D7" lsidName="L. lineatus" pageId="11" pageNumber="26" rank="species" species="lineatus">L. lineatus</taxonomicName>
, a species distributed worldwide. BIN-number BOLD:ACV7714.
</paragraph>
</subSubSection>
<subSubSection id="DF7BD9812100253B6274733F8DF8614F" pageId="11" pageNumber="26" type="remarks">
<paragraph id="4942FB08E240168A346BC2B2B38806CA" pageId="11" pageNumber="26">Remarks.</paragraph>
<paragraph id="B1AC50882868590A5D8552FB4185CCF9" pageId="11" pageNumber="26">
<taxonomicName id="3ED0B248BB3632DFEF4B795117A9F326" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus verrucosus" order="Enchytraeida" pageId="11" pageNumber="26" phylum="Annelida" rank="species" species="verrucosus">Lumbricillus verrucosus</taxonomicName>
was originally described by
<bibRefCitation id="2D3929A93B570D6B8371E74FC6411E2B" author="Claparede, E" journalOrPublisher="Memoires de la Societe de Physique et d'Histoire Naturelle de Geneve" pageId="45" pageNumber="60" pagination="71 - 164" title="Etudes anatomiques sur les Annelides, Turbellaries, Opalines et Gregarines observes dans les Hebrides." volume="16" year="1861">
<normalizedToken id="1B558380C81CA3038E0665C379A15BCD" originalValue="Claparède">Claparede</normalizedToken>
(1861)
</bibRefCitation>
and later synonymized with
<taxonomicName id="0A2C32C2E69E246BEDADCDE199D572FD" lsidName="L. lineatus" pageId="11" pageNumber="26" rank="species" species="lineatus">L. lineatus</taxonomicName>
by
<bibRefCitation id="6EF119025FAB7EB1ACAB1EF5BFFFEA84" pageId="11" pageNumber="26">Nielsen and Christensen (1959)</bibRefCitation>
, probably following
<bibRefCitation id="3339A074CD5ECFE1AFE31E3ABE1A635B" author="Cernosvitov, L" journalOrPublisher="Bulletin de l'Association Russe pour les Recherches Scientifiques a Prague" pageId="44" pageNumber="59" pagination="263 - 295" title="System der Enchytraeiden." volume="5" year="1937">
<normalizedToken id="115BE4AD4698CE498AFC3A7DC6498D75" originalValue="Černosvitov">Cernosvitov</normalizedToken>
(1937)
</bibRefCitation>
who had downgraded the former to a form of the latter. Specimens from the two species examined in this study are indeed very similar when considering the shape of the spermathecae (Fig. 3) and most body measurements, but they differ in body colour, chaetal number and the proportions of the sperm funnels: indeed
<taxonomicName id="430FF1F075B36169EF085484A9D09755" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus verrucosus" order="Enchytraeida" pageId="11" pageNumber="26" phylum="Annelida" rank="species" species="verrucosus">Lumbricillus verrucosus</taxonomicName>
was described as being pale yellow, having 3-5 chaetae and sperm funnels about three times longer than wide (
<bibRefCitation id="40387B78766AF66A2BD53C48517D5D18" author="Claparede, E" journalOrPublisher="Memoires de la Societe de Physique et d'Histoire Naturelle de Geneve" pageId="45" pageNumber="60" pagination="71 - 164" title="Etudes anatomiques sur les Annelides, Turbellaries, Opalines et Gregarines observes dans les Hebrides." volume="16" year="1861">
<normalizedToken id="592798F7937363D8EC3A18EF6C814822" originalValue="Claparède">Claparede</normalizedToken>
1861
</bibRefCitation>
), against
<taxonomicName id="22767317823817BE220EC6C39FA35F31" lsidName="L. lineatus" pageId="11" pageNumber="26" rank="species" species="lineatus">L. lineatus</taxonomicName>
being orange-red, having more chaetae and sperm funnels about five times longer than wide. Furthermore, the spermathecae of
<taxonomicName id="7360E59107AFBF8285AA4B288EC37F68" lsidName="L. verrucosus" pageId="11" pageNumber="26" rank="species" species="verrucosus">L. verrucosus</taxonomicName>
seem to have an ampulla that is slightly longer and wider in the part ectal to the constriction, but the importance of this character remains to be proved. Even though the two species may be very difficult to separate morphologically, except perhaps by body colour, they are supported as separate species molecularly and avesiculate specimens have been found by us only in
<taxonomicName id="1943F646FAF83960B91A53461D4B9768" lsidName="L. lineatus" pageId="11" pageNumber="26" rank="species" species="lineatus">L. lineatus</taxonomicName>
, not in
<taxonomicName id="552EDDB61346052BB096EA137097D0C5" lsidName="L. verrucosus" pageId="11" pageNumber="26" rank="species" species="verrucosus">L. verrucosus</taxonomicName>
.
</paragraph>
<paragraph id="8A062E0848AE7F0512E94640B7C27762" pageId="11" pageNumber="26">
Interestingly,
<taxonomicName id="E873CF9CEBB19DCF3D75E3A21989D34B" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus verrucosus" order="Enchytraeida" pageId="11" pageNumber="26" phylum="Annelida" rank="species" species="verrucosus">Lumbricillus verrucosus</taxonomicName>
is genetically most closely related to
<taxonomicName id="42496824737F981C64237374A2E2D9AD" lsidName="L. rivalis" pageId="11" pageNumber="26" rank="species" species="rivalis">L. rivalis</taxonomicName>
(Levinsen, 1883) and not to
<taxonomicName id="BA4031068F727E719AFDD5CA7D525AB2" lsidName="L. lineatus" pageId="11" pageNumber="26" rank="species" species="lineatus">L. lineatus</taxonomicName>
(Fig. 1). However, these three species are well supported as closely related to each other.
</paragraph>
</subSubSection>
</treatment>
</document>