CheneBelgiqueV2, Pmc, Corpus, bibRecord, 000273

Calcaridorylaimus castaneae sp. n. (Nematoda, Dorylaimidae) from Bulgaria with an identification key to the species of the genus

Identifieur interne : 000273 ( Pmc/Corpus ); précédent : 000272; suivant : 000274

Calcaridorylaimus castaneae sp. n. (Nematoda, Dorylaimidae) from Bulgaria with an identification key to the species of the genus

Auteurs : Sevdan Nedelchev ; Milka Elshishka ; Stela Lazarova ; Georgi Radoslavov ; Peter Hristov ; Vlada Peneva

Source :

ZooKeys [ 1313-2989 ] ; 2014.

RBID : PMC:4042707

Abstract

An unknown species belonging to the genusPageBreakCalcaridorylaimus Andrássy, 1986 was collected from the litter of broadleaf forests dominated by Castanea sativa Mill. and mixed with Quercus daleshampii Ten. and Fagus sylvatica L. on Belasitsa Mountain, south-western Bulgaria. Calcaridorylaimus castaneaesp. n. is characterised by its long body (1.4–2.1 mm), lip region practically not offset, vulva transverse, short odontostyle (14.5–16 μm) and tail (75.5–110.5 μm, c=14.7–23.6; c’=2.9–4.4) in females and 38–46 μm long spicules with small spur before their distant end in males. It is most similar to C. andrassyi Ahmad & Shaheen, 2004, but differs in having transverse vs pore-like vulva and shorter spicules (38–46 μm vs 52–57 μm). An identification key to the species of the genus Calcaridorylaimus is proposed. Phylogenetic analyses were performed on 18S and D2-D3 expansion domains of 28S rRNA genes by Neighbor-Joining, Maximum Likelihood and Bayesian Inference methods. The phylograms inferred from 18S sequences showed closest relationships of the new species with some species belonging to the genus Mesodorylaimus. However, insufficient molecular data for members of both genera do not allow the phylogenetic relationships of Calcaridorylaimus and the new species described herein to be elucidated.

Url:

http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4042707

DOI: 10.3897/zookeys.410.6955
PubMed: 24899849
PubMed Central: 4042707

Links to Exploration step

PMC:4042707

Le document en format XML

<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en"><italic>Calcaridorylaimus castaneae</italic>
 sp. n. (Nematoda, Dorylaimidae) from Bulgaria with an identification key to the species of the genus</title>
<author><name sortKey="Nedelchev, Sevdan" sort="Nedelchev, Sevdan" uniqKey="Nedelchev S" first="Sevdan" last="Nedelchev">Sevdan Nedelchev</name>
<affiliation><nlm:aff id="A1">Faculty of Biology, University of Sofia, Bd. “Dragan Tzankov” 8, 1421 Sofia, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Elshishka, Milka" sort="Elshishka, Milka" uniqKey="Elshishka M" first="Milka" last="Elshishka">Milka Elshishka</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
</affiliation>
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<author><name sortKey="Lazarova, Stela" sort="Lazarova, Stela" uniqKey="Lazarova S" first="Stela" last="Lazarova">Stela Lazarova</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
</affiliation>
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<author><name sortKey="Radoslavov, Georgi" sort="Radoslavov, Georgi" uniqKey="Radoslavov G" first="Georgi" last="Radoslavov">Georgi Radoslavov</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hristov, Peter" sort="Hristov, Peter" uniqKey="Hristov P" first="Peter" last="Hristov">Peter Hristov</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Peneva, Vlada" sort="Peneva, Vlada" uniqKey="Peneva V" first="Vlada" last="Peneva">Vlada Peneva</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
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<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main"><italic>Calcaridorylaimus castaneae</italic>
 sp. n. (Nematoda, Dorylaimidae) from Bulgaria with an identification key to the species of the genus</title>
<author><name sortKey="Nedelchev, Sevdan" sort="Nedelchev, Sevdan" uniqKey="Nedelchev S" first="Sevdan" last="Nedelchev">Sevdan Nedelchev</name>
<affiliation><nlm:aff id="A1">Faculty of Biology, University of Sofia, Bd. “Dragan Tzankov” 8, 1421 Sofia, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Elshishka, Milka" sort="Elshishka, Milka" uniqKey="Elshishka M" first="Milka" last="Elshishka">Milka Elshishka</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Lazarova, Stela" sort="Lazarova, Stela" uniqKey="Lazarova S" first="Stela" last="Lazarova">Stela Lazarova</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Radoslavov, Georgi" sort="Radoslavov, Georgi" uniqKey="Radoslavov G" first="Georgi" last="Radoslavov">Georgi Radoslavov</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hristov, Peter" sort="Hristov, Peter" uniqKey="Hristov P" first="Peter" last="Hristov">Peter Hristov</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Peneva, Vlada" sort="Peneva, Vlada" uniqKey="Peneva V" first="Vlada" last="Peneva">Vlada Peneva</name>
<affiliation><nlm:aff id="A2">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</nlm:aff>
</affiliation>
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<series><title level="j">ZooKeys</title>
<idno type="ISSN">1313-2989</idno>
<idno type="eISSN">1313-2970</idno>
<imprint><date when="2014">2014</date>
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<front><div type="abstract" xml:lang="en"><label>Abstract</label>
<p>An unknown species belonging to the genus<pmc-comment>PageBreak</pmc-comment>
<italic>Calcaridorylaimus</italic>
 Andrássy, 1986 was collected from the litter of broadleaf forests dominated by <italic>Castanea sativa</italic>
 Mill. and mixed with <italic>Quercus daleshampii</italic>
 Ten. and <italic>Fagus sylvatica</italic>
 L. on Belasitsa Mountain, south-western Bulgaria. <italic>Calcaridorylaimus castaneae</italic>
<bold>sp. n.</bold>
 is characterised by its long body (1.4–2.1 mm), lip region practically not offset, vulva transverse, short odontostyle (14.5–16 μm) and tail (75.5–110.5 μm, c=14.7–23.6; c’=2.9–4.4) in females and 38–46 μm long spicules with small spur before their distant end in males. It is most similar to <italic>C. andrassyi</italic>
 Ahmad & Shaheen, 2004, but differs in having transverse vs pore-like vulva and shorter spicules (38–46 μm vs 52–57 μm). An identification key to the species of the genus <italic>Calcaridorylaimus</italic>
 is proposed. Phylogenetic analyses were performed on 18S and D2-D3 expansion domains of 28S rRNA genes by Neighbor-Joining, Maximum Likelihood and Bayesian Inference methods. The phylograms inferred from 18S sequences showed closest relationships of the new species with some species belonging to the genus <italic>Mesodorylaimus</italic>
. However, insufficient molecular data for members of both genera do not allow the phylogenetic relationships of <italic>Calcaridorylaimus</italic>
 and the new species described herein to be elucidated.</p>
</div>
</front>
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<pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
  <front><journal-meta><journal-id journal-id-type="nlm-ta">Zookeys</journal-id>
<journal-id journal-id-type="iso-abbrev">Zookeys</journal-id>
<journal-id journal-id-type="publisher-id">ZooKeys</journal-id>
<journal-title-group><journal-title>ZooKeys</journal-title>
</journal-title-group>
<issn pub-type="ppub">1313-2989</issn>
<issn pub-type="epub">1313-2970</issn>
<publisher><publisher-name>Pensoft Publishers</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">24899849</article-id>
<article-id pub-id-type="pmc">4042707</article-id>
<article-id pub-id-type="doi">10.3897/zookeys.410.6955</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject>
</subj-group>
</article-categories>
<title-group><article-title><italic>Calcaridorylaimus castaneae</italic>
 sp. n. (Nematoda, Dorylaimidae) from Bulgaria with an identification key to the species of the genus</article-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Nedelchev</surname>
<given-names>Sevdan</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/9BDC2CA1-9183-40A5-94D0-F7FD7CD16275">http://zoobank.org/9BDC2CA1-9183-40A5-94D0-F7FD7CD16275</uri>
</contrib>
<contrib contrib-type="author"><name><surname>Elshishka</surname>
<given-names>Milka</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
<uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/79DCEFF6-47EB-41DF-ABE3-69213F54806B">http://zoobank.org/79DCEFF6-47EB-41DF-ABE3-69213F54806B</uri>
</contrib>
<contrib contrib-type="author"><name><surname>Lazarova</surname>
<given-names>Stela</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
<uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/83C802AD-6631-4008-8E2E-3B8F3ACA1F5C">http://zoobank.org/83C802AD-6631-4008-8E2E-3B8F3ACA1F5C</uri>
</contrib>
<contrib contrib-type="author"><name><surname>Radoslavov</surname>
<given-names>Georgi</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
<uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/415009FF-DE2E-4A02-83D4-66125561AFA7">http://zoobank.org/415009FF-DE2E-4A02-83D4-66125561AFA7</uri>
</contrib>
<contrib contrib-type="author"><name><surname>Hristov</surname>
<given-names>Peter</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
<uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/4C865CC6-DED3-4DCE-AAA1-71A51922752B">http://zoobank.org/4C865CC6-DED3-4DCE-AAA1-71A51922752B</uri>
</contrib>
<contrib contrib-type="author"><name><surname>Peneva</surname>
<given-names>Vlada</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
<uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/D066DD81-0D99-4117-835C-34CD35BE6F41">http://zoobank.org/D066DD81-0D99-4117-835C-34CD35BE6F41</uri>
</contrib>
</contrib-group>
<aff id="A1"><label>1</label>
Faculty of Biology, University of Sofia, Bd. “Dragan Tzankov” 8, 1421 Sofia, Bulgaria</aff>
<aff id="A2"><label>2</label>
Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria</aff>
<author-notes><corresp>Corresponding author: Vlada Peneva (<email xlink:type="simple">vpeneva@ecolab.bas.bg</email>
)</corresp>
<fn fn-type="edited-by"><p>Academic editor: S. Subbotin</p>
</fn>
</author-notes>
<pub-date pub-type="collection"><year>2014</year>
</pub-date>
<pub-date pub-type="epub"><day>20</day>
<month>5</month>
<year>2014</year>
</pub-date>
<issue>410</issue>
<fpage>41</fpage>
<lpage>61</lpage>
<history><date date-type="received"><day>3</day>
<month>1</month>
<year>2014</year>
</date>
<date date-type="accepted"><day>17</day>
<month>4</month>
<year>2014</year>
</date>
</history>
<permissions><copyright-statement>Sevdan Nedelchev, Milka Elshishka, Stela Lazarova, Georgi Radoslavov, Peter Hristov, Vlada Peneva</copyright-statement>
<license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0"><license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
</license>
</permissions>
<self-uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/501921E5-E350-470F-A472-6D65E737B133">http://zoobank.org/501921E5-E350-470F-A472-6D65E737B133</self-uri>
<abstract><label>Abstract</label>
<p>An unknown species belonging to the genus<pmc-comment>PageBreak</pmc-comment>
<italic>Calcaridorylaimus</italic>
 Andrássy, 1986 was collected from the litter of broadleaf forests dominated by <italic>Castanea sativa</italic>
 Mill. and mixed with <italic>Quercus daleshampii</italic>
 Ten. and <italic>Fagus sylvatica</italic>
 L. on Belasitsa Mountain, south-western Bulgaria. <italic>Calcaridorylaimus castaneae</italic>
<bold>sp. n.</bold>
 is characterised by its long body (1.4–2.1 mm), lip region practically not offset, vulva transverse, short odontostyle (14.5–16 μm) and tail (75.5–110.5 μm, c=14.7–23.6; c’=2.9–4.4) in females and 38–46 μm long spicules with small spur before their distant end in males. It is most similar to <italic>C. andrassyi</italic>
 Ahmad & Shaheen, 2004, but differs in having transverse vs pore-like vulva and shorter spicules (38–46 μm vs 52–57 μm). An identification key to the species of the genus <italic>Calcaridorylaimus</italic>
 is proposed. Phylogenetic analyses were performed on 18S and D2-D3 expansion domains of 28S rRNA genes by Neighbor-Joining, Maximum Likelihood and Bayesian Inference methods. The phylograms inferred from 18S sequences showed closest relationships of the new species with some species belonging to the genus <italic>Mesodorylaimus</italic>
. However, insufficient molecular data for members of both genera do not allow the phylogenetic relationships of <italic>Calcaridorylaimus</italic>
 and the new species described herein to be elucidated.</p>
</abstract>
<kwd-group><label>Keywords</label>
<kwd>Taxonomy</kwd>
<kwd>morphology</kwd>
<kwd>18S and D2-D3 rRNA genes</kwd>
<kwd>compendium</kwd>
</kwd-group>
</article-meta>
<notes><sec sec-type="Citation"><title>Citation</title>
<p>Nedelchev S, Elshishka M, Lazarova S, Radoslavov G, Hristov P, Peneva V (2014) <italic>Calcaridorylaimus castaneae</italic>
 sp. n. (Nematoda, Dorylaimidae) from Bulgaria with an identification key to the species of the genus. ZooKeys 410: 41–61. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.3897/zookeys.410.6955">10.3897/zookeys.410.6955</ext-link>
</p>
</sec>
</notes>
</front>
<body><sec sec-type="Introduction"><title>Introduction</title>
<p>During an ecological study of chestnut forests on Belasitsa Mountain (2003-2005) an undescribed species belonging to the genus <italic><named-content content-type="taxon-name">Calcaridorylaimus</named-content>
</italic>
 Andrássy, 1986 was recovered. The genus <italic><named-content content-type="taxon-name">Calcaridorylaimus</named-content>
</italic>
 is represented by nine species worldwide: <italic><named-content content-type="taxon-name">Calcaridorylaimus calcarifer</named-content>
</italic>
 Andrássy, 1986, <italic><named-content content-type="taxon-name">Calcaridorylaimus promissus</named-content>
</italic>
 Andrássy, 1986, <italic><named-content content-type="taxon-name">Calcaridorylaimus ruwenzorii</named-content>
</italic>
 (De Coninck, 1935) Andrássy, 1986, <italic><named-content content-type="taxon-name">Calcaridorylaimus signatus</named-content>
</italic>
 (Loof, 1975) Andrássy, 1986, <italic><named-content content-type="taxon-name">Calcaridorylaimus simillimus</named-content>
</italic>
 Andrássy, 1986, <italic><named-content content-type="taxon-name">Calcaridorylaimus sirgeli</named-content>
</italic>
 Heyns & Meyer, 1995, <italic><named-content content-type="taxon-name">Calcaridorylaimus arcticus</named-content>
</italic>
 Gagarin, 1997, <italic><named-content content-type="taxon-name">Calcaridorylaimus andrassyi</named-content>
</italic>
 Ahmad & Shaheen, 2004 and <italic><named-content content-type="taxon-name">Calcaridorylaimus beatus</named-content>
</italic>
 Andrássy, 2011. The genus is distributed mainly in the southern hemisphere: three species occur in Africa, two in South America and one each in Antarctic, Central America, and Europe, and <italic><named-content content-type="taxon-name">Calcaridorylaimus promissus</named-content>
</italic>
 was recorded from Australia and Alaska, North America (<xref rid="B6" ref-type="bibr">Andrássy 1986</xref>
, <xref rid="B7" ref-type="bibr">2003</xref>
) (<xref ref-type="fig" rid="F1">Fig. 1</xref>
). The most characteristic features of these species are the shapes and structures of the spicules which are provided with a small spur before the distal tip. The new species is described based on both morphological and molecular data.</p>
<fig id="F1" orientation="portrait" position="float"><label>Figure 1.</label>
<caption><p>Distribution of <italic><named-content content-type="taxon-name">Calcaridorylaimus</named-content>
</italic>
 spp. (■ <italic><named-content content-type="taxon-name">Calcaridorylaimus calcarifer</named-content>
</italic>
, ● <italic><named-content content-type="taxon-name">Calcaridorylaimus andrassyi</named-content>
</italic>
, ♦ <italic><named-content content-type="taxon-name">Calcaridorylaimus arcticus</named-content>
</italic>
, □ <italic><named-content content-type="taxon-name">Calcaridorylaimus promissus</named-content>
</italic>
, ▲ <italic><named-content content-type="taxon-name">Calcaridorylaimus signatus</named-content>
</italic>
, ○ <italic><named-content content-type="taxon-name">Calcaridorylaimus simillimus</named-content>
</italic>
, ♢ <italic><named-content content-type="taxon-name">Calcaridorylaimus sirgeli</named-content>
</italic>
, △ <italic><named-content content-type="taxon-name">Calcaridorylaimus ruwenzorii</named-content>
</italic>
, ◙ <italic><named-content content-type="taxon-name">Calcaridorylaimus beatus</named-content>
</italic>
, ▼ <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n.).</p>
</caption>
<graphic xlink:href="zookeys-410-041-g001"></graphic>
</fig>
</sec>
<sec sec-type="materials|methods"><title>Materials and methods</title>
<sec sec-type="Sample collection"><title>Sample collection</title>
<p>The litter samples were collected in 2003 by the last author (VP) from three sites on Belasitsa Mountain representing different types of broadleaf forests dominated by <pmc-comment>PageBreak</pmc-comment>
<italic><named-content content-type="taxon-name">Castanea sativa</named-content>
</italic>
 Mill. mixed with <italic><named-content content-type="taxon-name">Quercus daleshampii</named-content>
</italic>
 Ten. and <italic><named-content content-type="taxon-name">Fagus sylvatica</named-content>
</italic>
 L. (Forest Management Plan database, sub-compartments 104g, 140b and 146a). Subsequently, on 17.10.2012 new litter samples were collected by Dr Michaela Ilieva from one of these sites, sub-compartment 140b, in order to obtain fresh material for molecular studies. Nematodes were recovered from the litter using the Baermann funnel method. They were killed by heat (65 °C), fixed in TAF (Triethanolamine-formalin, <xref rid="B9" ref-type="bibr">Courtney et al. 1955</xref>
), and processed to anhydrous glycerine (<xref rid="B29" ref-type="bibr">Seinhorst 1959</xref>
). Drawings were prepared using an Amplival 30-G048b and a drawing tube РА-6У42. Photographs were taken using an Axio Imager M2-Carl Zeiss compound microscope equipped with a digital camera (ProgRes C7) and specialised software (CapturePro Software 2.8). Measurements were made using an Olympus BX41 light microscope, a digitising tablet (CalComp Drawing Board III, GTCO CalCom Peripherals, Scottsdale, AZ, USA), and computer programme Digitrak 1.0f (Philip Smith, Scottish Crop Research Institute, Dundee, UK).</p>
</sec>
<sec sec-type="DNA extraction, amplification and sequencing "><title>DNA extraction, amplification and sequencing</title>
<p>Genomic DNA was extracted from two female and two male worms using a standard nematode digestion protocol (<xref rid="B18" ref-type="bibr">Holterman et al. 2006</xref>
). Two overlapping fragments of 18S rRNA genes (~1600 bp) were amplified from each specimen using primer sets 988F (5`-CTC AAA GAT TAA GCC ATG C-3`) and 1912R (5`-TTT ACG GTC AGA ACT AGG G-3)` for the first fragment, and 1813F (5`-CTG CGT GAG AGG TGA AAT-3`, 2646R 5`-GCT ACC TTG TTA CGA CTT TT-3`) for the second fragment (<xref rid="B18" ref-type="bibr">Holterman et al. 2006</xref>
). The D2/D3 expansion segments of the 28S rRNA gene (~900 bp) were additionally amplified from all specimens using the primers D2A (5`-ACA AGT ACC GTG AGG GAA AGT TG-3`) and D3B (5`-TCG GAA GGA ACC AGC TAC TA-3`) (<xref rid="B12" ref-type="bibr">De Ley et al. 1999</xref>
). Each PCR reaction was performed under the following conditions: initial denaturation 94 °C for 5 min; 40 cycles (denaturation 94 °C for 30 sec; primer annealing 50 °C for 30 sec; extension 72 °C for 1 min), and final extension 72 °C for 10 min. PCR products were visualized on 1% agarose gel with GreenSafe (NZYtech) under visible and UV light. Fragment size was determined using GeneRuler™ 100 bp Ladder Plus (Ferments, Thermo Scientific). The amplified products were sequenced by Eurofins MWG Operon.</p>
<pmc-comment>PageBreak</pmc-comment>
      </sec>
<sec sec-type="Sequence and phylogenetic analysis"><title>Sequence and phylogenetic analysis</title>
<p>The sequences of the new species have been deposited in GenBank with the accession numbers <ext-link ext-link-type="gen" xlink:href="KF717497">KF717497</ext-link>
 and <ext-link ext-link-type="gen" xlink:href="KF717498">KF717498</ext-link>
 for the 18S and the D2-D3 rRNA genes, respectively. A BLAST (Basic Local Alignment Search Tool) search at NCBI (National Center for Biotechnology Information) was performed using the obtained sequences as queries to confirm their nematode origin and to identify the most closely related nematode sequences. The sequences revealing a similarity up to 97% and 85% with nematodes from various Dorylaimida families were included in the phylogenetic analyses of 18S and D2-D3 regions, respectively (<xref rid="B15" ref-type="bibr">Griffiths et al. 2006</xref>
; <xref rid="B18" ref-type="bibr">Holterman et al. 2006</xref>
; <xref rid="B24" ref-type="bibr">Meldal et al. 2007</xref>
; <xref rid="B21" ref-type="bibr">Lesaulnier et al. 2008</xref>
; <xref rid="B26" ref-type="bibr">Pedram et al. 2010</xref>
; <xref rid="B27" ref-type="bibr">Pedram et al. 2011</xref>
; <xref rid="B3" ref-type="bibr">Álvarez-Ortega and Peña-Santiago 2012a</xref>
; <xref rid="B4" ref-type="bibr">2012b</xref>
; <xref rid="B13" ref-type="bibr">Donn et al. 2012</xref>
; <xref rid="B5" ref-type="bibr">Álvarez-Ortega et al. 2013</xref>
). The Multiple Sequence Alignments (MSA) of both datasets were performed using the Clustal Omega tool (<xref rid="B30" ref-type="bibr">Sievers et al. 2011</xref>
) via the EBI webserver: <ext-link ext-link-type="uri" xlink:href="http://www.ebi.ac.uk/Tools/msa/clustalw2/">http://www.ebi.ac.uk/Tools/msa/clustalw2/</ext-link>
. Subsequently, the MSAs were manually optimised and trimmed using MEGA 5 (<xref rid="B32" ref-type="bibr">Tamura et al. 2011</xref>
). <italic><named-content content-type="taxon-name">Eudorylaimus</named-content>
</italic>
 sp. (family <named-content content-type="taxon-name">Qudsianematidae</named-content>
) was used as an outgroup taxon for both 18S and D2-D3 rDNA sequence datasets (accession numbers <ext-link ext-link-type="gen" xlink:href="AY284800">AY284800</ext-link>
 and <ext-link ext-link-type="gen" xlink:href="AY593037">AY593037</ext-link>
, respectively; <xref rid="B19" ref-type="bibr">Holterman et al. 2008</xref>
). The phylogenetic reconstructions of three datasets D2-D3, complete and partial 18S rDNA were performed using Neighbor Joining (NJ), Maximum Likelihood (ML) and the Bayesian Inference (BI) algorithms and implemented in MEGA 5.0 and MrBayes v3.2.1 (<xref rid="B20" ref-type="bibr">Huelsenbeck and Ronquist 2001</xref>
, <xref rid="B32" ref-type="bibr">Tamura et al. 2011</xref>
, <xref rid="B28" ref-type="bibr">Ronquist et al. 2012</xref>
). The NJ phylogenetic inferences were performed under the following settings: Maximum Composite Likelihood method for computing evolutionary distances; Gamma distributed rates among sites, estimated values set up to 0.3429 (D2-D3) and 0.05 (18S rDNA); 2000 bootstrap replications. A total of 755 and 1593 positions in the final datasets were used for both analyses, respectively. General Time Reversible model (GTR) plus Gamma distribution rates (G) and 1000 bootstrap replications were used as ML analyses settings for all datasets. The Bayesian MCMC tree searches were conducted using MrBayes 3.2.1. Each analysis was run for 10, 000, 000 generations with a sample frequency of 1000 generations. The first 25% of the chains discarded as burning and the remaining 75% trees kept to summarise the tree topology, branch lengths, and posterior probabilities (PP) of branch support. The evolutionary models for nucleotide substitutions were set up as for ML analyses. Convergence diagnostic values were calculated every 1000 generations with a predefined stop value equal to 0.01. A single strict consensus tree was visualised using FigTree v1.4.0 graphical viewer (<ext-link ext-link-type="uri" xlink:href="http://tree.bio.ed.ac.uk/software/figtree/">http://tree.bio.ed.ac.uk/software/figtree/</ext-link>
). Posterior probabilities values of ≥0.80 (BI) and bootstrap values of ≥70 (NJ and ML) were considered as credible support values for nodes.</p>
<pmc-comment>PageBreak</pmc-comment>
      </sec>
</sec>
<sec><title>Taxonomy</title>
<sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Calcaridorylaimus</named-content>
<named-content content-type="species">castaneae</named-content>
</named-content>
<named-content content-type="taxon-status">sp. n.</named-content>
</title>
<p>http://zoobank.org/9BF1D302-1986-47C5-B0D8-2F5DC75BD6D2</p>
<p>http://species-id.net/wiki/Calcaridorylaimus_castaneae</p>
<p><xref ref-type="fig" rid="F2">Figs 2</xref>
<xref ref-type="fig" rid="F3"></xref>
<xref ref-type="fig" rid="F4"></xref>
<xref ref-type="fig" rid="F5"></xref>
<xref ref-type="fig" rid="F6">–6</xref>
</p>
<sec sec-type="treatment-Measurements"><title>Measurements.</title>
<p>See <xref ref-type="table" rid="T2">Table 1</xref>
.</p>
<table-wrap id="T2" orientation="portrait" position="float"><label>Table 1.</label>
<caption><p>Morphometrics of <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n., females and males, from Belasitsa Mountain. All measurements, unless indicated otherwise, in μm and in the form: mean ± SD (range).</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Habitat</th>
<th rowspan="1" colspan="5">Mixed broadleaf forest</th>
<th rowspan="1" colspan="2">Chestnut forest</th>
</tr>
<tr><th rowspan="1" colspan="1">Locality and year of collection</th>
<th rowspan="1" colspan="3">140b 2003</th>
<th rowspan="1" colspan="2">140b 2012</th>
<th rowspan="1" colspan="2">104g 2003</th>
</tr>
<tr><th rowspan="1" colspan="1">Characters</th>
<th rowspan="1" colspan="1">Holotype</th>
<th rowspan="1" colspan="6">Paratypes</th>
</tr>
<tr><th rowspan="1" colspan="1"></th>
<th rowspan="1" colspan="1"></th>
<th rowspan="1" colspan="1">Females</th>
<th rowspan="1" colspan="1">Males</th>
<th rowspan="1" colspan="1">Females</th>
<th rowspan="1" colspan="1">Males</th>
<th rowspan="1" colspan="1">Females</th>
<th rowspan="1" colspan="1">Males</th>
</tr>
<tr><th rowspan="1" colspan="1">n</th>
<th rowspan="1" colspan="1"></th>
<th rowspan="1" colspan="1">40</th>
<th rowspan="1" colspan="1">20</th>
<th rowspan="1" colspan="1">10</th>
<th rowspan="1" colspan="1">10</th>
<th rowspan="1" colspan="1">18</th>
<th rowspan="1" colspan="1">12</th>
</tr>
<tr><td rowspan="1" colspan="1">L (mm)</td>
<td rowspan="1" colspan="1">1.7</td>
<td rowspan="1" colspan="1">1.7±0.11 <break></break>
 (1.5–2.0)</td>
<td rowspan="1" colspan="1">1.6±0.08 <break></break>
 (1.4–1.8)</td>
<td rowspan="1" colspan="1">1.6±0.2 <break></break>
 (1.4–1.8)</td>
<td rowspan="1" colspan="1">1.6±0.1 <break></break>
 (1.4–1.8)</td>
<td rowspan="1" colspan="1">1.6±0.15 <break></break>
 (1.4–2.1)</td>
<td rowspan="1" colspan="1">1.5±0.06 <break></break>
 (1.4–1.6)</td>
</tr>
<tr><td rowspan="1" colspan="1">a</td>
<td rowspan="1" colspan="1">35.7</td>
<td rowspan="1" colspan="1">36.6±2.4 <break></break>
 (32.5–42.7)</td>
<td rowspan="1" colspan="1">37.9±2.3 <break></break>
 (33.3–42.2)</td>
<td rowspan="1" colspan="1">35.7±2.1 <break></break>
 (32.2–38.7)</td>
<td rowspan="1" colspan="1">40.0±2.3 <break></break>
 (37.0–44.3)</td>
<td rowspan="1" colspan="1">38.4±1.4 <break></break>
 (35.7–40.6)</td>
<td rowspan="1" colspan="1">38.1±2.0 <break></break>
 (34.0–41.6)</td>
</tr>
<tr><td rowspan="1" colspan="1">b</td>
<td rowspan="1" colspan="1">4.9</td>
<td rowspan="1" colspan="1">5.0±0.3 <break></break>
 (4.5–5.7)</td>
<td rowspan="1" colspan="1">4.7±0.2 <break></break>
 (4.4–5.1)</td>
<td rowspan="1" colspan="1">5.1±0.2 <break></break>
 (4.8–5.6)</td>
<td rowspan="1" colspan="1">5.1±0.4 <break></break>
 (4.7–6.0)</td>
<td rowspan="1" colspan="1">4.9±0.3 <break></break>
 (4.4–5.5)</td>
<td rowspan="1" colspan="1">4.6±0.2 <break></break>
 (4.3–5.0)</td>
</tr>
<tr><td rowspan="1" colspan="1">c</td>
<td rowspan="1" colspan="1">19.4</td>
<td rowspan="1" colspan="1">18.8±0.4 <break></break>
 (16.0–22.0)</td>
<td rowspan="1" colspan="1">74.4±7.4 <break></break>
 (63.8–89.7)</td>
<td rowspan="1" colspan="1">18.3±1.4 <break></break>
 (16.8–21.0)</td>
<td rowspan="1" colspan="1">66.7±6.9 <break></break>
 (53.7–73.2)</td>
<td rowspan="1" colspan="1">18.5±2.3 <break></break>
 (14.7–23.6)</td>
<td rowspan="1" colspan="1">72.2±7.0 <break></break>
 (61.4–83.5)</td>
</tr>
<tr><td rowspan="1" colspan="1">c‘</td>
<td rowspan="1" colspan="1">3.9</td>
<td rowspan="1" colspan="1">3.6±0.4 <break></break>
 (2.9–4.4)</td>
<td rowspan="1" colspan="1">0.76±0.06 <break></break>
 (0.67–0.84)</td>
<td rowspan="1" colspan="1">3.5±0.4 <break></break>
 (3.0–4.0)</td>
<td rowspan="1" colspan="1">0.84±0.06 <break></break>
 (0.77–0.95)</td>
<td rowspan="1" colspan="1">3.6±0.3 <break></break>
 (2.9–4.2)</td>
<td rowspan="1" colspan="1">0.76±0.06 <break></break>
 (0.68–0.87)</td>
</tr>
<tr><td rowspan="1" colspan="1">V/T %</td>
<td rowspan="1" colspan="1">50.7</td>
<td rowspan="1" colspan="1">51.6±1.7 <break></break>
 (47.9–54.8)</td>
<td rowspan="1" colspan="1">66.3±3.2 <break></break>
 (60–72)</td>
<td rowspan="1" colspan="1">51.2±1.6 <break></break>
 (49–54)</td>
<td rowspan="1" colspan="1">59.5±2.5 <break></break>
 (56–62.5)</td>
<td rowspan="1" colspan="1">51.0±2.3 <break></break>
 (46.5–54.5)</td>
<td rowspan="1" colspan="1">62.8±4.3 <break></break>
 (54–70)</td>
</tr>
<tr><td rowspan="1" colspan="1">G1%</td>
<td rowspan="1" colspan="1">17</td>
<td rowspan="1" colspan="1">18.4±1.7 <break></break>
 (14-22.6)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">19.1±1.0 <break></break>
 (18.1–20.7)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">17.8±2.1 <break></break>
 (13.8–21.7)</td>
<td rowspan="1" colspan="1">-</td>
</tr>
<tr><td rowspan="1" colspan="1">G2%</td>
<td rowspan="1" colspan="1">16</td>
<td rowspan="1" colspan="1">18.3±2.3 <break></break>
 (14.1–24.4)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">18.2±1.0 <break></break>
 (16.6-19.1)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">18.7±3 <break></break>
 (14.5-26.5)</td>
<td rowspan="1" colspan="1">-</td>
</tr>
<tr><td rowspan="1" colspan="1">Odontostyle</td>
<td rowspan="1" colspan="1">14.5</td>
<td rowspan="1" colspan="1">15.0±0.4 <break></break>
 (14.5–16)</td>
<td rowspan="1" colspan="1">15.1±0.25 <break></break>
 (14.5–16)</td>
<td rowspan="1" colspan="1">15.1±0.2 <break></break>
 (14.9–15.4)</td>
<td rowspan="1" colspan="1">14.9±0.4 <break></break>
 (14–15)</td>
<td rowspan="1" colspan="1">15.0±0.2 <break></break>
 (14.5–15)</td>
<td rowspan="1" colspan="1">15±0.3 <break></break>
 (14.5–16)</td>
</tr>
<tr><td rowspan="1" colspan="1">Odontophore</td>
<td rowspan="1" colspan="1">26</td>
<td rowspan="1" colspan="1">24.8±2.2 <break></break>
 (20–29)</td>
<td rowspan="1" colspan="1">22.9±1.5 <break></break>
 (21–27)</td>
<td rowspan="1" colspan="1">21.6±2.1 <break></break>
 (20–26)</td>
<td rowspan="1" colspan="1">23.7±4.0 <break></break>
 (20–28)</td>
<td rowspan="1" colspan="1">27.2±1.6 <break></break>
 (24–30)</td>
<td rowspan="1" colspan="1">25.6±1.7 <break></break>
 (23–28)</td>
</tr>
<tr><td rowspan="1" colspan="1">Spear</td>
<td rowspan="1" colspan="1">41</td>
<td rowspan="1" colspan="1">39.8±2.2 <break></break>
 (35–43)</td>
<td rowspan="1" colspan="1">37.7±1.2 <break></break>
 (35.5–41.5)</td>
<td rowspan="1" colspan="1">36.8±2.0 <break></break>
 (35–41)</td>
<td rowspan="1" colspan="1">38.6±4.2 <break></break>
 (34–43)</td>
<td rowspan="1" colspan="1">41±1.4 <break></break>
 (38–44)</td>
<td rowspan="1" colspan="1">40.7±1.6 <break></break>
 (38–43)</td>
</tr>
<tr><td rowspan="1" colspan="1">Neck length</td>
<td rowspan="1" colspan="1">346</td>
<td rowspan="1" colspan="1">342±13.2 <break></break>
 (310–366)</td>
<td rowspan="1" colspan="1">338.6±8.8 <break></break>
 (318–352)</td>
<td rowspan="1" colspan="1">319.3±27.3 <break></break>
 (272.5–352)</td>
<td rowspan="1" colspan="1">315±31.4 <break></break>
 (242–340)</td>
<td rowspan="1" colspan="1">335.8±17 <break></break>
 (306–376)</td>
<td rowspan="1" colspan="1">336±11.9 <break></break>
 (310–352)</td>
</tr>
<tr><td rowspan="1" colspan="1">Cardia length</td>
<td rowspan="1" colspan="1">19</td>
<td rowspan="1" colspan="1">19±1.8 <break></break>
 (16–24)</td>
<td rowspan="1" colspan="1">19±2.1 <break></break>
 (16–24)</td>
<td rowspan="1" colspan="1">33.3±7.0 <break></break>
 (22-41)</td>
<td rowspan="1" colspan="1">29.6±5.8 <break></break>
 (24–39)</td>
<td rowspan="1" colspan="1">18.5±2.4 <break></break>
 (15–25)</td>
<td rowspan="1" colspan="1">19.9±3.8 <break></break>
 (14.5–26)</td>
</tr>
<tr><td rowspan="1" colspan="1">Body width at: <break></break>
 lip region</td>
<td rowspan="1" colspan="1">12</td>
<td rowspan="1" colspan="1">12.3±0.3 <break></break>
 (12–12.5)</td>
<td rowspan="1" colspan="1">12.3±0.3 <break></break>
 (12–12.5)</td>
<td rowspan="1" colspan="1">12.4±0.5 <break></break>
 (12–13)</td>
<td rowspan="1" colspan="1">12.5±0.5 <break></break>
 (12–13)</td>
<td rowspan="1" colspan="1">12.3±0.4 <break></break>
 (12–13)</td>
<td rowspan="1" colspan="1">12.5±0.19 <break></break>
 (12–13)</td>
</tr>
<tr><td rowspan="1" colspan="1">mid-body</td>
<td rowspan="1" colspan="1">48</td>
<td rowspan="1" colspan="1">46.8±4.7 <break></break>
 (39.5–55)</td>
<td rowspan="1" colspan="1">42.8±2.3 <break></break>
 (37–47)</td>
<td rowspan="1" colspan="1">45.2±2.2 <break></break>
 (41–48)</td>
<td rowspan="1" colspan="1">40.1±2.7 <break></break>
 (35–44.5)</td>
<td rowspan="1" colspan="1">42±4.7 <break></break>
 (37–58)</td>
<td rowspan="1" colspan="1">40.7±2.2 <break></break>
 (37–45)</td>
</tr>
<tr><td rowspan="1" colspan="1">anus</td>
<td rowspan="1" colspan="1">22</td>
<td rowspan="1" colspan="1">24.9±1.8 <break></break>
 (22–30)</td>
<td rowspan="1" colspan="1">28.8±0.9 <break></break>
 (27–30)</td>
<td rowspan="1" colspan="1">25.7±1.1 <break></break>
 (24.5–28.5)</td>
<td rowspan="1" colspan="1">28.7±1.7 <break></break>
 (25–30.5)</td>
<td rowspan="1" colspan="1">24.8±1.3 <break></break>
 (23–27)</td>
<td rowspan="1" colspan="1">28.5±1.0 <break></break>
 (27–30)</td>
</tr>
<tr><td rowspan="1" colspan="1">Lateral chord</td>
<td rowspan="1" colspan="1">12</td>
<td rowspan="1" colspan="1">12.7±1.8 <break></break>
 (10.5–18)</td>
<td rowspan="1" colspan="1">10.1±1.3 <break></break>
 (8–12.5)</td>
<td rowspan="1" colspan="1">12.8±1.1 <break></break>
 (11–15)</td>
<td rowspan="1" colspan="1">10.4±1.4 <break></break>
 (9-13)</td>
<td rowspan="1" colspan="1">12.2±1.2 <break></break>
 (10.5–16)</td>
<td rowspan="1" colspan="1">10±0.9 <break></break>
 (8.5–12)</td>
</tr>
<tr><td rowspan="1" colspan="1">Prerectum</td>
<td rowspan="1" colspan="1">89</td>
<td rowspan="1" colspan="1">89.4±11 <break></break>
 (68–117)</td>
<td rowspan="1" colspan="1">184±17.2 <break></break>
 (132–207)</td>
<td rowspan="1" colspan="1">89.5±19.1 <break></break>
 (64–104.5)</td>
<td rowspan="1" colspan="1">155.5±24.6 <break></break>
 (130-200)</td>
<td rowspan="1" colspan="1">76.6±9.3 <break></break>
 (60–96)</td>
<td rowspan="1" colspan="1">165.4±10 <break></break>
 (145–176)</td>
</tr>
<tr><td rowspan="1" colspan="1">Rectum</td>
<td rowspan="1" colspan="1">41</td>
<td rowspan="1" colspan="1">37±2.4 <break></break>
 (30–41)</td>
<td rowspan="1" colspan="1">41±2.1 <break></break>
 (37.5–44)</td>
<td rowspan="1" colspan="1">34.5±4.2 <break></break>
 (30–40.5)</td>
<td rowspan="1" colspan="1">40.3±3.3 <break></break>
 (34-43)</td>
<td rowspan="1" colspan="1">39.6±1.4 <break></break>
 (37–41)</td>
<td rowspan="1" colspan="1">43.8±1.5 <break></break>
 (41–46)</td>
</tr>
<tr><td rowspan="1" colspan="1">Tail</td>
<td rowspan="1" colspan="1">88</td>
<td rowspan="1" colspan="1">90.9±7.1 <break></break>
 (76–110)</td>
<td rowspan="1" colspan="1">22.1±1.8 <break></break>
 (19–25)</td>
<td rowspan="1" colspan="1">88.7±8.5 <break></break>
 (75.5–103)</td>
<td rowspan="1" colspan="1">24.3±2.6 <break></break>
 (20–28)</td>
<td rowspan="1" colspan="1">90.1±9 <break></break>
 (78–110.5)</td>
<td rowspan="1" colspan="1">21.6±2.3 <break></break>
 (18–26)</td>
</tr>
<tr><td rowspan="1" colspan="1">Spicules</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">43±1 <break></break>
 (41–45)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">43.6±2.9 <break></break>
 (38–46)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">43.5±2 <break></break>
 (39.5–45)</td>
</tr>
<tr><td rowspan="1" colspan="1">T-distance anterior end of anterior testis-cloaca (mm)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">1.1±0.1 <break></break>
 (0.9–1.3)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">0.96±0.11 <break></break>
 (0.86-1.11)</td>
<td rowspan="1" colspan="1">-</td>
<td rowspan="1" colspan="1">1.0±0.1 <break></break>
 (0.8–1.15)</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec sec-type="treatment-Description"><title>Description.</title>
<p><italic>Female.</italic>
 Body slender, more or less curved ventrally. Cuticle ca 2 μm at anterior part of neck, 3 μm thick at midbody, 4.5–5.5 μm at postanal region; outer layer with fine transverse striae. Lateral chord ca 1/4 of body width. Three dorsal, two ventral and three lateral pores are observed in the spear area. Lip region practically not offset, lip region width ca 20% of its height. Lips partly fused, labial papilae slightly protruding. Body at proximal end of pharynx 3–4 times the width of the lip region diameter. Amphidial aperture 5–6 μm wide or about half the lip region width. Odontostyle 1.2–1.3 times the lip region diameter, aperture occupying 35–40% of its length. Odontophore simple 1.3–1.7 times odontostyle length. Guiding ring at 8.5–9 μm from anterior end. Nerve ring surrounding the pharynx at 36–39% of neck length from head end. Hemizonid and conspicuous excretory pore observed in the nerve ring region. Pharyngeal characters (five females and five males): pharynx beginning to widen at 56–60% and attaining its full width at 61–64% of neck length from anterior end. DO (for terminology see <xref rid="B23" ref-type="bibr">Loof and Coomans 1970</xref>
) lying near the point where the pharynx attains its full width; DO–DN 8–13 μm. The two S<sub>1</sub>
N lying at a small distance behind the middle of the distance DN–S<sub>2</sub>
N, the anterior one (S<sub>1</sub>
N<sub>1</sub>
) smaller, ca 2 μm diam; S<sub>1</sub>
N<sub>2</sub>
 comparatively large and distinct, 3–4 μm diam. DN nucleolus 4–4.5 μm diam.; S<sub>2</sub>
N 2–3 μm diam. (DN>S<sub>1</sub>
N<sub>2</sub>
>S<sub>2</sub>
N). Locations (%):</p>
<table-wrap orientation="portrait" id="d36e1211" position="anchor"><table frame="hsides" rules="groups" id="T1"><tbody><tr><td rowspan="1" colspan="1">DO=60–64</td>
<td rowspan="1" colspan="1">S<sub>1</sub>
N<sub>1</sub>
=78-81</td>
<td rowspan="1" colspan="1">S<sub>2</sub>
N=91–92</td>
<td rowspan="1" colspan="1">K=73–84</td>
</tr>
<tr><td rowspan="1" colspan="1">DN=62–66</td>
<td rowspan="1" colspan="1">S<sub>1</sub>
N<sub>2</sub>
=82–85</td>
<td rowspan="1" colspan="1">S<sub>2</sub>
O=93–95</td>
<td rowspan="1" colspan="1">K’=78–85</td>
</tr>
<tr><td rowspan="1" colspan="1">DO–DN=2.2–3.8</td>
<td rowspan="1" colspan="1">S<sub>1</sub>
N<sub>1</sub>
–S<sub>1</sub>
N<sub>2</sub>
=2.9–4.8.</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
</tbody>
</table>
</table-wrap>
<pmc-comment>PageBreak</pmc-comment>
          <pmc-comment>PageBreak</pmc-comment>
          <pmc-comment>PageBreak</pmc-comment>
          <pmc-comment>PageBreak</pmc-comment>
          <pmc-comment>PageBreak</pmc-comment>
          <pmc-comment>PageBreak</pmc-comment>
          <p>Cardia conoid, variable in length, microvilli visible only here. Genital system didelphic-amphidelphic, ovaries reflexed, reaching rarely the vulva level. Oviducts and ovaries very long compared to uteri. Uteri not differentiated, short, anterior 96-125 μm and posterior 95-135 μm long, in one female with no sperm inside the genital system uteri shorter, 83 μm each, sphincter between uterus and <italic>pars dilatata oviductus</italic>
 well developed. Sperm present in uteri and <italic>pars dilatata oviductus</italic>
. Synchronous uterine eggs 1–3, measuring 72–78 × 32–44 μm. Vulva transverse. Vagina extending to 60–70% inwards: <italic>pars proximalis</italic>
 9–13 μm wide, 14.5–19.5 μm long, <italic>pars refringens</italic>
 more or less rounded trapezoid, 10–12.5 μm wide, 4–6 μm deep; <italic>pars distalis</italic>
 approx. 1.5 μm (terminology following <xref rid="B11" ref-type="bibr">De Ley et al. 1993</xref>
). Peculiar tongue-like valve present at intestine-prerectum junction. Rectum 1.3–1.5, prerectum 3–4 times anal body width long, respectively. Tail first conoid, then more or less uniformly tapering to a narrowly rounded terminus. Posterior part of tail usually slightly curved dorsally. Two pairs of caudal pores, one subventral, other subdorsal.</p>
<pmc-comment>PageBreak</pmc-comment>
          <p><italic>Male.</italic>
 General morphology similar to that of female, body curved ventrally in J-shape when fixed. Genital system diorchic, testes opposed, well developed. Spicules dorylaimoid, with double contour on dorsal arm, 1.3–1.6 times the corresponding body diameter long; ventral arm smaller than dorsal. A spur present dorsally before the distal tip, distinctly visible in extruded spicules. Lateral guiding pieces 9-11 μm long or ca. 23 % spicule length. In addition to adcloacal pair seven to twelve (mostly nine or ten), regularly spaced ventromedian supplements present (9 supplements in 8 specimens; 10 suppl. – in 7, and 7, 8 and 12 each in one specimen). Prerectum 4.5–7.5 times the corresponding body diameter long, extending 0.7–1.8 body widths anterior to the supplement series. Tail dorsally conoid and broadly rounded. One subdorsal and one subterminal pair of caudal pores.</p>
</sec>
<sec sec-type="treatment-Differential diagnosis and relationships"><title>Differential diagnosis and relationships.</title>
<p><italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. differs from all species in the genus by a combination of the following characters: long body (1.4–2.1 mm), lip region practically not offset, short odontostyle (14.5–16 μm in females and 14-16 μm in males) and short female tail (75.5–110.5 μm, c=14.7–23.6; c’=2.9–4.4); vulva transverse, 38–46 μm long spicules with spur before its distal end. The new species is most similar to <italic><named-content content-type="taxon-name">Calcaridorylaimus andrassyi</named-content>
</italic>
 from which it can be differentiated by having transverse vulva vs pore-like and shorter spicules (38–46 μm vs 52–57 μm). Further, <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 differs from <italic><named-content content-type="taxon-name">Calcaridorylaimus ruwenzorii</named-content>
</italic>
 by having shorter odontostyle (14.5-16 μm vs 19.5-25 μm), and tail (75.5-110.5 μm vs 160 μm, higher c (c=14.7-23.6 vs c=10) and lower c’ (c’=2.9–4.4 vs 7) values in females. It can be differentiated from <italic><named-content content-type="taxon-name">Calcaridorylaimus arcticus</named-content>
</italic>
, <italic><named-content content-type="taxon-name">Calcaridorylaimus beatus</named-content>
</italic>
, <italic><named-content content-type="taxon-name">Calcaridorylaimus calcarifer</named-content>
</italic>
 and <italic><named-content content-type="taxon-name">Calcaridorylaimus signatus</named-content>
</italic>
 by having different vulva shape (transverse vs longitudinal) and shorter spicules (38–46 μm vs 57–67 μm; 48–55 μm; 52–54 μm and 72 μm). From <italic><named-content content-type="taxon-name">Calcaridorylaimus sirgeli</named-content>
</italic>
 it differs by having transverse vulva vs pore-like, higher c (c=14.7–23.6 vs c=9.7–11.3) and lower c’ (c’=2.9–4.4 vs c’=5.3–6.7) values. Finally, <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 differs from <italic><named-content content-type="taxon-name">Calcaridorylaimus promissus</named-content>
</italic>
 and <italic><named-content content-type="taxon-name">Calcaridorylaimus simillimus</named-content>
</italic>
 by having longer odontostyle (14.5–16 μm vs 13 μm and 11 μm) and shorter tail (75.5–110.5 μm vs 158–178 μm and 175 μm).</p>
<p>Presence of the conspicuous excretory pore observed in <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 is unusual for members of Dorylaimida, however, similar structure has been mentioned only for two <italic><named-content content-type="taxon-name">Longidorus</named-content>
</italic>
 species, namely <italic><named-content content-type="taxon-name">Longidorus macrosoma</named-content>
</italic>
 Hooper, 1961 and <italic><named-content content-type="taxon-name">Longidorus carniolensis</named-content>
</italic>
 Širca et al., 2011 (<xref rid="B1" ref-type="bibr">Aboul-Eid 1969</xref>
, <xref rid="B31" ref-type="bibr">Širca et al. 2011</xref>
) and for several <italic><named-content content-type="taxon-name">Mesodorylaimus</named-content>
</italic>
 species origination from Antarctica – <italic><named-content content-type="taxon-name">Mesodorylaimus chipevi</named-content>
</italic>
 Nedelchev & Peneva, 2000, <italic><named-content content-type="taxon-name">Mesodorylaimus antarcticus</named-content>
</italic>
 Nedelchev & Peneva, 2000, <italic><named-content content-type="taxon-name">Mesodorylaimus masleni</named-content>
</italic>
 Nedelchev & Peneva, 2000, <italic><named-content content-type="taxon-name">Mesodorylaimus imperator</named-content>
</italic>
 Loof, 1975 (<xref rid="B25" ref-type="bibr">Nedelchev and Peneva 2000</xref>
).</p>
</sec>
<sec sec-type="treatment-Type locality and plant association"><title>Type locality and plant association.</title>
<p>Belasitsa Mountain, south-western Bulgaria, litter from old broadleaf forest (100-140 years) dominated by <italic><named-content content-type="taxon-name">Castanea sativa</named-content>
</italic>
, mixed with <italic><named-content content-type="taxon-name">Quercus daleshampii</named-content>
</italic>
 and <italic><named-content content-type="taxon-name">Fagus sylvatica</named-content>
</italic>
. Site is located in the vicinity of Belasitsa hut, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="dwc:verbatimCoordinates">41°22'12"N, 23°11'12"E</named-content>
</named-content>
 (sub-compartment 140b). Second locality: young sweet chestnut forest (30-40 years old) near Belasitsa village (sub-compartment 104g).</p>
</sec>
<sec sec-type="treatment-Type material"><title>Type material.</title>
<p>Holotype and 80 paratype females and 47 males deposited in the nematode collection of the Institute of Biodiversity and Ecosystem Research, Sofia, Bulgaria. Other paratypes deposited as follows: four females and two males in the Nematode Collection of the Foodland Environment Research Agency, Sand Hutton, UK (former Rothamsted Nematode Collection); three females and three males in the USDA Nematode Collection, Beltsville, Maryland, USA; two females and two males in the Riverside Nematode Collection, University of California, Riverside, USA; four females, and four males in the Wageningen Nematode Collection (WANECO), Wageningen, the Netherlands; four females and three males in the Nematode Collection of the Zoology Museum of the Ghent University, Belgium.</p>
</sec>
<sec sec-type="treatment-Etymology"><title>Etymology.</title>
<p>The scientific name is derived from the generic name of dominant tree species, the sweet chestnut tree (<italic><named-content content-type="taxon-name">Castanea</named-content>
</italic>
) in the forest where this nematode was found.</p>
<fig id="F2" orientation="portrait" position="float"><label>Figure 2.</label>
<caption><p><italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. <italic>Female</italic>
: <bold>A</bold>
 Pharyngeal gland nuclei <bold>B</bold>
 Anterior region <bold>C</bold>
 Pharyngeal region <bold>D</bold>
 Genital system <bold>E</bold>
 Vulval region <bold>F</bold>
 Sperm cells in uterus. Scale bars: <bold>A, B, E, F</bold>
 – 30 μm; <bold>C, D</bold>
 – 50 μm.</p>
</caption>
<graphic xlink:href="zookeys-410-041-g002"></graphic>
</fig>
<fig id="F3" orientation="portrait" position="float"><label>Figure 3.</label>
<caption><p><italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. <italic>Female</italic>
: <bold>A</bold>
 Posterior region. <italic>Male</italic>
: <bold>B</bold>
, <bold>C</bold>
 Extruded spicules with supplements <bold>D</bold>
 Posterior region. Scale bar: <bold>A–D</bold>
 – 30 μm.</p>
</caption>
<graphic xlink:href="zookeys-410-041-g003"></graphic>
</fig>
<fig id="F4" orientation="portrait" position="float"><label>Figure 4.</label>
<caption><p><italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. <italic>Female</italic>
: <bold>A</bold>
 Anterior end <bold>C</bold>
 Amphid <bold>F–I</bold>
 Tail shapes <italic>Male</italic>
<bold>B</bold>
 Anterior end <bold>D</bold>
 Posterior end with extruded spicules, arrow indicating the spur <bold>E</bold>
 Posterior end. Scale bars: <bold>A, B, D–I</bold>
 – 20 μm; <bold>C</bold>
 – 6 μm.</p>
</caption>
<graphic xlink:href="zookeys-410-041-g004"></graphic>
</fig>
<fig id="F5" orientation="portrait" position="float"><label>Figure 5.</label>
<caption><p><italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. <italic>Female</italic>
: <bold>A</bold>
 Entire body <bold>B</bold>
 Lip region <bold>D</bold>
 Prerectum, arrow pointing tongue-like valve <bold>E</bold>
 Pharyngeal bulb <bold>F</bold>
 Vulval region with posterior uterus <bold>G</bold>
 Vulval region with egg in posterior uterus <bold>H, I</bold>
 Vulval region <bold>J</bold>
 Cardia <bold>L</bold>
 Lateral field. <italic>Male</italic>
: <bold>A</bold>
 Entire body <bold>C</bold>
 Lip region <bold>K</bold>
 Cardia <bold>M</bold>
 Supplements. Scale bars: <bold>A</bold>
 – 200 μm; <bold>B, C, M</bold>
 – 6 μm; <bold>E–L</bold>
 – 20 μm.</p>
</caption>
<graphic xlink:href="zookeys-410-041-g005"></graphic>
</fig>
<fig id="F6" orientation="portrait" position="float"><label>Figure 6.</label>
<caption><p><italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. <italic>Male</italic>
: <bold>A</bold>
 Posterior end <bold>B</bold>
 Sperm cells in testis <bold>C–F</bold>
 Spicular region <bold>C</bold>
 Lateral piece of spicules <bold>D, E</bold>
 Extruded spicules, arrows pointing the spur <bold>F</bold>
 Spicules in the body. Scale bars: <bold>A, B</bold>
 – 20 μm; <bold>C–F</bold>
 – 18 μm.</p>
</caption>
<graphic xlink:href="zookeys-410-041-g006"></graphic>
</fig>
<pmc-comment>PageBreak</pmc-comment>
          <pmc-comment>PageBreak</pmc-comment>
        </sec>
</sec>
<sec sec-type="Key to species of Calcaridorylaimus"><title>Key to species of <italic><named-content content-type="taxon-name">Calcaridorylaimus</named-content>
</italic>
</title>
<table-wrap content-type="key" orientation="portrait" id="d36e1663" position="anchor"><table frame="hsides" rules="groups"><tbody><tr><td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">Odontostyle 19.5–25 µm long</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus ruwenzorii</named-content>
</italic>
</td>
</tr>
<tr><td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Odontostyle shorter, ≤17 µm</td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr><td rowspan="1" colspan="1">2</td>
<td rowspan="1" colspan="1">c’=11</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus simillimus</named-content>
</italic>
</td>
</tr>
<tr><td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">c’<9</td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr><td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">c’=2.2–2.4</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus beatus</named-content>
</italic>
</td>
</tr>
<tr><td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">c’≥ 2.9</td>
<td rowspan="1" colspan="1">4</td>
</tr>
<tr><td rowspan="1" colspan="1">4</td>
<td rowspan="1" colspan="1">c≤11</td>
<td rowspan="1" colspan="1">5</td>
</tr>
<tr><td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">c>11</td>
<td rowspan="1" colspan="1">7</td>
</tr>
<tr><td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">V=45–47, supplements 10–13</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus promissus</named-content>
</italic>
</td>
</tr>
<tr><td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">V>49, supplements 6–9</td>
<td rowspan="1" colspan="1">6</td>
</tr>
<tr><td rowspan="1" colspan="1">6</td>
<td rowspan="1" colspan="1">Vulva longitudinal, spicules 52–54 µm</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus calcarifer</named-content>
</italic>
</td>
</tr>
<tr><td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Vulva pore-like, spicules 39–45 µm</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus sirgeli</named-content>
</italic>
</td>
</tr>
<tr><td rowspan="1" colspan="1">7</td>
<td rowspan="1" colspan="1">Lip region 14–17 µm wide, vulva longitudinal</td>
<td rowspan="1" colspan="1">8</td>
</tr>
<tr><td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Lip region 11–13 µm wide, vulva not longitudinal</td>
<td rowspan="1" colspan="1">9</td>
</tr>
<tr><td rowspan="1" colspan="1">8</td>
<td rowspan="1" colspan="1">Tail 120–171 µm long, L=1.6–2.27, spicules 57–67</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus arcticus</named-content>
</italic>
</td>
</tr>
<tr><td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Tail 100 µm long, L=1.3–1.7, spicules 72</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus signatus</named-content>
</italic>
</td>
</tr>
<tr><td rowspan="1" colspan="1">9</td>
<td rowspan="1" colspan="1">Vulva pore-like, spicules 52–57</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus andrassyi</named-content>
</italic>
</td>
</tr>
<tr><td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Vulva transverse, spicules 38–46</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T3" orientation="portrait" position="float"><label>Table 2.</label>
<caption><p>Main morphological and morphometrical data of <italic><named-content content-type="taxon-name">Calcaridorylaimus</named-content>
</italic>
 species, habitat type and distribution.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Species</th>
<th rowspan="1" colspan="1">Body length (mm)</th>
<th rowspan="1" colspan="1">a</th>
<th rowspan="1" colspan="1">c</th>
<th rowspan="1" colspan="1">c’</th>
<th rowspan="1" colspan="1">V%</th>
<th rowspan="1" colspan="1">Odontostyle</th>
<th rowspan="1" colspan="1">Lip region width</th>
<th rowspan="1" colspan="1">Tail</th>
<th rowspan="1" colspan="1">Vulva shape</th>
<th rowspan="1" colspan="1">Spicule length</th>
<th rowspan="1" colspan="1">Supple-ments</th>
<th rowspan="1" colspan="1">Habitat</th>
<th rowspan="1" colspan="1">Distribution</th>
<th rowspan="1" colspan="1">References</th>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus andrassyi</named-content>
</italic>
<break></break>
 ♀ <break></break>
 ♂</td>
<td rowspan="1" colspan="1">1.8 <break></break>
 (1.7–1.9) <break></break>
 1.5 <break></break>
 (1.5–1.6)</td>
<td rowspan="1" colspan="1">44 <break></break>
 (42–46) <break></break>
 39 <break></break>
 (34–45)</td>
<td rowspan="1" colspan="1">15 <break></break>
 (14–16) <break></break>
 78 <break></break>
 (76–81)</td>
<td rowspan="1" colspan="1">4.4 <break></break>
 (4.1–4.7) <break></break>
 0.66 <break></break>
 (0.6–0.7)</td>
<td rowspan="1" colspan="1">49 <break></break>
 (48–52)</td>
<td rowspan="1" colspan="1">15 <break></break>
 (15–16) <break></break>
 15 <break></break>
</td>
<td rowspan="1" colspan="1">12 <break></break>
 (11.5–12) <break></break>
 12 <break></break>
 (11.5–12.5)</td>
<td rowspan="1" colspan="1">113 <break></break>
 (105–120) <break></break>
 20 <break></break>
 (18.5–21.5)</td>
<td rowspan="1" colspan="1">P</td>
<td rowspan="1" colspan="1"><break></break>
<break></break>
 54 <break></break>
 (52–57)</td>
<td rowspan="1" colspan="1"><break></break>
<break></break>
 10</td>
<td rowspan="1" colspan="1">forest</td>
<td rowspan="1" colspan="1">Costa Rica</td>
<td rowspan="1" colspan="1"><xref rid="B2" ref-type="bibr">Ahmad and Shaheen 2004</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus arcticus</named-content>
</italic>
</td>
<td rowspan="1" colspan="1">1.6–2.3 <break></break>
<break></break>
 1.84–2.24</td>
<td rowspan="1" colspan="1">24–34 <break></break>
<break></break>
 24–34</td>
<td rowspan="1" colspan="1">13–16 <break></break>
<break></break>
 53–75</td>
<td rowspan="1" colspan="1">3.5–5.8 <break></break>
<break></break>
 0.6–0.7</td>
<td rowspan="1" colspan="1">49–53.4</td>
<td rowspan="1" colspan="1">14–17 <break></break>
<break></break>
 14–17</td>
<td rowspan="1" colspan="1">15–17 <break></break>
<break></break>
 15–17</td>
<td rowspan="1" colspan="1">139 <break></break>
 (120–171) <break></break>
 36 <break></break>
 (29–48)</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"><break></break>
<break></break>
 63 <break></break>
 (57–67)</td>
<td rowspan="1" colspan="1"><break></break>
<break></break>
 10–12</td>
<td rowspan="1" colspan="1">aquatic, fresh-water</td>
<td rowspan="1" colspan="1">Russian Arctic</td>
<td rowspan="1" colspan="1"><xref rid="B14" ref-type="bibr">Gagarin 1997</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus beatus</named-content>
</italic>
</td>
<td rowspan="1" colspan="1">1.28–1.30 <break></break>
 1.36–1.48</td>
<td rowspan="1" colspan="1">24–29 <break></break>
 28–33</td>
<td rowspan="1" colspan="1">19–23 <break></break>
 57–72</td>
<td rowspan="1" colspan="1">2.2–2.4 <break></break>
 0.6–0.8</td>
<td rowspan="1" colspan="1">51–52</td>
<td rowspan="1" colspan="1">15–16 <break></break>
</td>
<td rowspan="1" colspan="1">14–15</td>
<td rowspan="1" colspan="1">60–68</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1">48–55</td>
<td rowspan="1" colspan="1"><break></break>
 13–14</td>
<td rowspan="1" colspan="1">Moist grassy soil, 4500 m asl</td>
<td rowspan="1" colspan="1">Chile</td>
<td rowspan="1" colspan="1"><xref rid="B8" ref-type="bibr">Andrássy 2011</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus calcarifer</named-content>
</italic>
</td>
<td rowspan="1" colspan="1">1.18–1.30 <break></break>
 0.95–1.07</td>
<td rowspan="1" colspan="1">30–32 <break></break>
 26–28</td>
<td rowspan="1" colspan="1">7.8–10.3 <break></break>
 45–46</td>
<td rowspan="1" colspan="1">6–8 <break></break>
 0.7–0.8</td>
<td rowspan="1" colspan="1">49–52</td>
<td rowspan="1" colspan="1">13–14</td>
<td rowspan="1" colspan="1">10–12</td>
<td rowspan="1" colspan="1">115–167 <break></break>
 21–23</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"><break></break>
 52–54</td>
<td rowspan="1" colspan="1"><break></break>
 8–9</td>
<td rowspan="1" colspan="1">rain forest soil</td>
<td rowspan="1" colspan="1">Republic of Congo</td>
<td rowspan="1" colspan="1"><xref rid="B6" ref-type="bibr">Andrássy 1986</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n.</td>
<td rowspan="1" colspan="1">1.4–2.1 <break></break>
 1.4–1.8</td>
<td rowspan="1" colspan="1">32.2–42.7 <break></break>
 33.3–44.3</td>
<td rowspan="1" colspan="1">16.0–23.6 <break></break>
 53.7–89.7</td>
<td rowspan="1" colspan="1">2.9–4.4 <break></break>
 0.7–0.95</td>
<td rowspan="1" colspan="1">46.5–54.8</td>
<td rowspan="1" colspan="1">14.5–16 <break></break>
 14–16</td>
<td rowspan="1" colspan="1">12–13 <break></break>
 12–13</td>
<td rowspan="1" colspan="1">75.5–110.5 <break></break>
 18–28</td>
<td rowspan="1" colspan="1">T</td>
<td rowspan="1" colspan="1"><break></break>
 38–46</td>
<td rowspan="1" colspan="1"><break></break>
 7–12</td>
<td rowspan="1" colspan="1">broadleaf forest, litter</td>
<td rowspan="1" colspan="1">Bulgaria</td>
<td rowspan="1" colspan="1">Present study</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus promissus</named-content>
</italic>
</td>
<td rowspan="1" colspan="1">1.28–1.37 <break></break>
 1.02–1.10</td>
<td rowspan="1" colspan="1">36–38 <break></break>
 28–30</td>
<td rowspan="1" colspan="1">7.7–8.4 <break></break>
 47–57</td>
<td rowspan="1" colspan="1">8.4–9.0 <break></break>
 0.8–0.9</td>
<td rowspan="1" colspan="1">45–47 <break></break>
</td>
<td rowspan="1" colspan="1">13 <break></break>
</td>
<td rowspan="1" colspan="1">10–11 <break></break>
 -</td>
<td rowspan="1" colspan="1">158–178 <break></break>
 18–22</td>
<td rowspan="1" colspan="1">T</td>
<td rowspan="1" colspan="1"><break></break>
 50–53</td>
<td rowspan="1" colspan="1"><break></break>
 10–13</td>
<td rowspan="1" colspan="1">wet moss from a rock, forest</td>
<td rowspan="1" colspan="1">Australia</td>
<td rowspan="1" colspan="1"><xref rid="B6" ref-type="bibr">Andrássy 1986</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1">**<italic><named-content content-type="taxon-name">Calcaridorylaimus promissus</named-content>
</italic>
</td>
<td rowspan="1" colspan="1">1.6–1.65 <break></break>
 1.55</td>
<td rowspan="1" colspan="1">34–37 <break></break>
 35</td>
<td rowspan="1" colspan="1">9–11 <break></break>
 90</td>
<td rowspan="1" colspan="1">7–8 <break></break>
 ?</td>
<td rowspan="1" colspan="1">50–51</td>
<td rowspan="1" colspan="1">14–15</td>
<td rowspan="1" colspan="1">?-</td>
<td rowspan="1" colspan="1">150–160 <break></break>
 ?</td>
<td rowspan="1" colspan="1">?</td>
<td rowspan="1" colspan="1"><break></break>
 50</td>
<td rowspan="1" colspan="1"><break></break>
 9–11</td>
<td rowspan="1" colspan="1">moss from swampy soil</td>
<td rowspan="1" colspan="1">Alaska</td>
<td rowspan="1" colspan="1"><xref rid="B7" ref-type="bibr">Andrássy 2003</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus ruwenzorii</named-content>
</italic>
</td>
<td rowspan="1" colspan="1">1.6 <break></break>
 1.3–1.5</td>
<td rowspan="1" colspan="1">32 <break></break>
 35–40</td>
<td rowspan="1" colspan="1">10 <break></break>
 44–74</td>
<td rowspan="1" colspan="1">7</td>
<td rowspan="1" colspan="1">47</td>
<td rowspan="1" colspan="1">19.5–25</td>
<td rowspan="1" colspan="1">12–14</td>
<td rowspan="1" colspan="1">160</td>
<td rowspan="1" colspan="1">?</td>
<td rowspan="1" colspan="1"><break></break>
 44</td>
<td rowspan="1" colspan="1"><break></break>
 7–8</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1">Republic of Congo</td>
<td rowspan="1" colspan="1"><xref rid="B10" ref-type="bibr">de Coninck 1935</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus signatus</named-content>
</italic>
</td>
<td rowspan="1" colspan="1">1.3–1.7 <break></break>
 1.7</td>
<td rowspan="1" colspan="1">25–33 <break></break>
 29</td>
<td rowspan="1" colspan="1">12.0–18.0 <break></break>
 61</td>
<td rowspan="1" colspan="1">2.9–4.2 <break></break>
 0.6*</td>
<td rowspan="1" colspan="1">49–56 <break></break>
 -</td>
<td rowspan="1" colspan="1">16–18 <break></break>
 17</td>
<td rowspan="1" colspan="1">14* <break></break>
 -</td>
<td rowspan="1" colspan="1">100* <break></break>
 24*</td>
<td rowspan="1" colspan="1">L</td>
<td rowspan="1" colspan="1"><break></break>
 72</td>
<td rowspan="1" colspan="1"><break></break>
 12</td>
<td rowspan="1" colspan="1">wet moss</td>
<td rowspan="1" colspan="1">Antarctic</td>
<td rowspan="1" colspan="1"><xref rid="B22" ref-type="bibr">Loof 1975</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus simillimus</named-content>
</italic>
</td>
<td rowspan="1" colspan="1">1.3 <break></break>
 1.14</td>
<td rowspan="1" colspan="1">43 <break></break>
 33</td>
<td rowspan="1" colspan="1">7.4 <break></break>
 90</td>
<td rowspan="1" colspan="1">11 <break></break>
 0.7</td>
<td rowspan="1" colspan="1">50</td>
<td rowspan="1" colspan="1">11.5</td>
<td rowspan="1" colspan="1">9.5–10</td>
<td rowspan="1" colspan="1">175 <break></break>
 12</td>
<td rowspan="1" colspan="1">T</td>
<td rowspan="1" colspan="1"><break></break>
 45</td>
<td rowspan="1" colspan="1"><break></break>
 11</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Stipa</named-content>
</italic>
 sp., near lake Titikaka</td>
<td rowspan="1" colspan="1">Bolivia</td>
<td rowspan="1" colspan="1"><xref rid="B6" ref-type="bibr">Andrássy 1986</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Calcaridorylaimus sirgeli</named-content>
</italic>
</td>
<td rowspan="1" colspan="1">1.34 <break></break>
 (1.1–1.5) <break></break>
 1.36 <break></break>
 (1.2–1.5)</td>
<td rowspan="1" colspan="1">33.2 <break></break>
 (30–36) <break></break>
 37.7 <break></break>
 (35–38)</td>
<td rowspan="1" colspan="1">10.6 <break></break>
 (9.7–11.3) <break></break>
 77 <break></break>
 (63–85)</td>
<td rowspan="1" colspan="1">6.1 <break></break>
 (5.3–6.7) <break></break>
 0.7 <break></break>
 (0.6-.8)</td>
<td rowspan="1" colspan="1">53.4 <break></break>
 (52–56)</td>
<td rowspan="1" colspan="1">13.6 <break></break>
 (12–15) <break></break>
 13.1 <break></break>
 (12.5–14)</td>
<td rowspan="1" colspan="1">11.5 <break></break>
 (10–12.5) <break></break>
 11.3 <break></break>
 (10–12.5)</td>
<td rowspan="1" colspan="1">127 <break></break>
 (105–148) <break></break>
 17.9 <break></break>
 (15–22)</td>
<td rowspan="1" colspan="1">P</td>
<td rowspan="1" colspan="1"><break></break>
<break></break>
 42.4 <break></break>
 (39–45)</td>
<td rowspan="1" colspan="1"><break></break>
<break></break>
 6–9</td>
<td rowspan="1" colspan="1">moist soil under fynbos and mosses</td>
<td rowspan="1" colspan="1">South Africa</td>
<td rowspan="1" colspan="1"><xref rid="B16" ref-type="bibr">Heyns and Meyer 1995</xref>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn><p>Shape of vulva: <bold>L</bold>
 – longitudinal; <bold>P</bold>
 – pore like; <bold>T</bold>
 – transverse; when average values are present ranges in parentheses; * from the drawing; ** <italic><named-content content-type="taxon-name">Calcaridorylaimus promissus</named-content>
</italic>
 from Alaska (<xref rid="B7" ref-type="bibr">Andrássy 2003</xref>
) was not included in a subsequent paper by the same author (<xref rid="B8" ref-type="bibr">Andrássy 2011</xref>
); since most of the characters deviate substantially from the original description, probably this population belongs to another species.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec sec-type="Phylogenetic relationships of Calcaridorylaimus castaneae based on partial 28S and 18S rDNA sequences"><title>Phylogenetic relationships of <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 based on partial 28S and 18S rDNA sequences</title>
<p>The sequences of D2-D3 expansion domains of 28S and 18S ribosomal DNA from two female and two male nematodes have been processed. No inter-individual variability within both domains have been observed, thus only two consensus sequences for each of the genes (787 and 1699 bp long) have been submitted to GenBank (accession numbers <ext-link ext-link-type="gen" xlink:href="KF717498">KF717498</ext-link>
 and <ext-link ext-link-type="gen" xlink:href="KF717497">KF717497</ext-link>
). A BLAST search for D2-D3 region showed highest similarity (88%) to the sequence of <italic><named-content content-type="taxon-name">Labronema vulvapapillatum</named-content>
</italic>
 clone 2 (<ext-link ext-link-type="gen" xlink:href="AY592997">AY592997</ext-link>
, <xref rid="B19" ref-type="bibr">Holterman et al. 2008</xref>
) while the 18S rDNA showed 99% similarity (4-6 nucleotide differences) to the sequences of <italic><named-content content-type="taxon-name">Mesodorylaimus bastiani</named-content>
</italic>
 (<ext-link ext-link-type="gen" xlink:href="AJ966488">AJ966488</ext-link>
) from France (<xref rid="B24" ref-type="bibr">Meldal et al. 2007</xref>
) and two sequences of unidentified species from environmental samples from Scotland (<ext-link ext-link-type="gen" xlink:href="AJ875133">AJ875133</ext-link>
 and <ext-link ext-link-type="gen" xlink:href="JN049666">JN049666</ext-link>
) (<xref rid="B15" ref-type="bibr">Griffiths et al. 2006</xref>
, <xref rid="B13" ref-type="bibr">Donn et al. 2012</xref>
).</p>
<p>The phylogenetic analyses based on 18S rDNA and D2-D3 of 28S rDNA sequences from various dorylaimid species with the highest matches of the BLAST search (up to 97% and 85%, respectively) were aligned along with our sequence. The phylograms obtained by NJ, ML and BI methods showed similar topology and differed only in the positions of poorly supported clades. The BI trees (<xref ref-type="fig" rid="F7">Figs 7</xref>
 and <xref ref-type="fig" rid="F8">8</xref>
) with posterior probabilities higher than 0.8 and NJ-ML trees with bootstrap values above 70% are presented (<xref ref-type="fig" rid="F9">Figs 9</xref>
 and <xref ref-type="fig" rid="F10">10</xref>
). The new species has clustered in a well-supported <pmc-comment>PageBreak</pmc-comment>
group with several <italic><named-content content-type="taxon-name">Mesodorylaimus</named-content>
</italic>
 spp. (<italic><named-content content-type="taxon-name">Mesodorylaimus bastiani</named-content>
</italic>
 (Bütschli, 1873), <italic><named-content content-type="taxon-name">Mesodorylaimus japonicus</named-content>
</italic>
 (Cobb in Thorne and Swanger, 1936) and <italic><named-content content-type="taxon-name">Mesodorylaimus</named-content>
</italic>
 cf. <italic>nigritulus</italic>
 (Schneider, 1937)) and two unidentified nematodes from environmental samples in the 18S rDNA phylogenetic reconstructions (<xref ref-type="fig" rid="F7">Figs 7</xref>
 and <xref ref-type="fig" rid="F9">9</xref>
); and to more distantly related species from various dorylaimid genera and families (<italic><named-content content-type="taxon-name">Prodorylaimus</named-content>
</italic>
, <italic><named-content content-type="taxon-name">Labronema</named-content>
</italic>
, <italic><named-content content-type="taxon-name">Nevadanema</named-content>
</italic>
, <italic><named-content content-type="taxon-name">Paractinolaimus</named-content>
</italic>
) in the partial D2-D3 LSU reconstructions (<xref ref-type="fig" rid="F8">Figs 8</xref>
 and <xref ref-type="fig" rid="F10">10</xref>
).</p>
<fig id="F7" orientation="portrait" position="float"><label>Figure 7.</label>
<caption><p>Phylogenetic relationships of <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. and its closest species for 18S rRNA gene. Bayesian Inference strict consensus tree acquired under GTR+G model. Posterior probabilities higher than 0.8 are presented.</p>
</caption>
<graphic xlink:href="zookeys-410-041-g007"></graphic>
</fig>
<fig id="F8" orientation="portrait" position="float"><label>Figure 8.</label>
<caption><p>Phylogenetic relationships of <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. and its closest species for D2-D3 expansion segments of the 28S rRNA gene. Bayesian Inference strict consensus tree acquired under GTR+G model. Posterior probabilities higher than 0.8 are presented.</p>
</caption>
<graphic xlink:href="zookeys-410-041-g008"></graphic>
</fig>
<fig id="F9" orientation="portrait" position="float"><label>Figure 9.</label>
<caption><p>Phylogenetic relationships of <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. and its closest species for 18S rRNA gene. Tree acquired with Neighbor Joining and Maximum Likelihood (GTR+G model) methods. Bootstrap values higher than 70% are presented.</p>
</caption>
<graphic xlink:href="zookeys-410-041-g009"></graphic>
</fig>
<fig id="F10" orientation="portrait" position="float"><label>Figure 10.</label>
<caption><p>Phylogenetic relationships of <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 sp. n. and its closest species for D2-D3 expansion segments of the 28S rRNA gene. Tree acquired with Neighbor Joining and Maximum Likelihood (GTR+G model) methods. Bootstrap values higher than 70% are presented.</p>
</caption>
<graphic xlink:href="zookeys-410-041-g010"></graphic>
</fig>
<p>The genus <pmc-comment>PageBreak</pmc-comment>
<pmc-comment>PageBreak</pmc-comment>
<pmc-comment>PageBreak</pmc-comment>
<italic><named-content content-type="taxon-name">Calcaridorylaimus</named-content>
</italic>
 was erected by <xref rid="B6" ref-type="bibr">Andrássy (1986)</xref>
 to accommodate a few species having different shapes and structures of spicules from those of <italic><named-content content-type="taxon-name">Mesodorylaimus</named-content>
</italic>
, with the males and females being practically indistinguishable. The phylograms inferred from 18S sequences showed the closest relationships of <italic><named-content content-type="taxon-name">Calcaridorylaimus castaneae</named-content>
</italic>
 with some members of the latter genus; however, the insufficiency of molecular data complementary to detailed morphological studies of species belonging to both genera does not allow the elucidation of evolutionary relationships among them and the position of the new species herein described.</p>
<pmc-comment>PageBreak</pmc-comment>
      </sec>
</sec>
<sec sec-type="supplementary-material"><title>Supplementary Material</title>
<supplementary-material id="zookeys.410.6955-treatment1" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Calcaridorylaimus</named-content>
<named-content content-type="species">castaneae</named-content>
</title>
</caption>
<media xlink:href="zookeys.410.6955-treatment1.xml" mimetype="text" mime-subtype="xml"></media>
</supplementary-material>
</sec>
</body>
<back><ack><title>Acknowledgements</title>
<p>The authors thanks Dr Michaela Ilieva from Institute of Biodiversity and Ecosystem Research for collecting additional litter samples from a sweet chestnut forest. This study was partly funded by the project ANIDIV, supported by the Bulgarian Academy of Sciences and project <italic>Assessment of the dynamics of structural and functional parameters of chestnut ecosystems in Belasitsa Mountain</italic>
; funded by Ministry of Agriculture and Forests, Bulgaria. The authors are thankful to Dr Nathalie Yonow from Swansea University, Wales, UK for critical reading of the manuscript and helpful suggestions.</p>
</ack>
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Calcaridorylaimus castaneae sp. n. (Nematoda, Dorylaimidae) from Bulgaria with an identification key to the species of the genus

Calcaridorylaimus castaneae sp. n. (Nematoda, Dorylaimidae) from Bulgaria with an identification key to the species of the genus

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