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<mods:title id="0C6732C312C46D93C503D60798A8CA03">Molecular and morphological characterisation of Longidoruspolyae sp. n. and L. pisi Edward, Misra &amp; Singh, 1964 (Dorylaimida, Longidoridae) from Bulgaria</mods:title>
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<treatment id="61A5877479C8113EF656B144B1EB0E2D" ID-GBIF-Taxon="156200902" LSID="urn:lsid:plazi:treatment:61A5877479C8113EF656B144B1EB0E2D" httpUri="http://treatment.plazi.org/id/61A5877479C8113EF656B144B1EB0E2D" lastPageId="9" lastPageNumber="84" pageId="7" pageNumber="82">
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<taxonomicName id="85891BE7D7358314158A340837150B89" ID-CoL="9LRJG" authority="Edward, Misra &amp; Singh, 1964" authorityName="Edward, Misra &amp; Singh" authorityYear="1964" class="Enoplea" family="Longidoridae" genus="Longidorus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Longidorus pisi" order="Dorylaimida" pageId="7" pageNumber="82" phylum="Nematoda" rank="species" species="pisi">Longidorus pisi Edward, Misra &amp; Singh, 1964</taxonomicName>
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<paragraph id="228ED5D7FEA4C8D31D4C9BC2EC5C6E1A" pageId="7" pageNumber="82">
= 
<taxonomicName id="39CA1C03B8FE564156768EBBA6183404" class="Enoplea" family="Longidoridae" genus="Longidorus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Longidorus latocephalus" order="Dorylaimida" pageId="7" pageNumber="82" phylum="Nematoda" rank="species" species="latocephalus">Longidorus latocephalus</taxonomicName>
Lamberti, Choleva &amp; Agostinelli, 1983
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<paragraph id="53466E73F00C98E790253F2ADB769D5D" pageId="7" pageNumber="82">Notes.</paragraph>
<paragraph id="ED9E0B70216B706D72C10D5766122F37" lastPageId="9" lastPageNumber="84" pageId="7" pageNumber="82">
Morphological and morphometric data for females and juvenile stages are presented in Table 4 and in Figure 8. The morphometric data obtained in this study agreed with those of 
<taxonomicName id="9740349769DABA7B267EC5D775200B78" lsidName="L. latocephalus" pageId="7" pageNumber="82" rank="species" species="latocephalus">L. latocephalus</taxonomicName>
from its type locality (
<bibRefCitation id="6B01D3A23C1E3B173409B2B731DCBCF4" author="Lamberti, F" journalOrPublisher="Nematologia Mediterranea" pageId="11" pageNumber="86" pagination="49 - 72" title="Longidoridae from Bulgaria (Nematoda, Dorylaimida) with description of three new species of Longidorus and two new species of Xiphinema." volume="11" year="1983">Lamberti et al. 1983</bibRefCitation>
) and several additional populations studied by 
<bibRefCitation id="D5207EF3F786F53BCBF2FE633F4694E8" author="Lamberti, F" journalOrPublisher="Nematologia Mediterranea" pageId="11" pageNumber="86" pagination="213 - 237" title="Morphometric variation and juvenile stages of some longidorid nematodes from Bulgaria with comments on the number of juvenile stages of Longidorusafricanus, L. closelongatus and Xiphinemasantos." volume="25" year="1997">Lamberti et al. (1997)</bibRefCitation>
. Furthermore, when compared to the type population of 
<taxonomicName id="EFA4702D79F483B71F1CD12602FFBF77" lsidName="L. pisi" pageId="7" pageNumber="82" rank="species" species="pisi">L. pisi</taxonomicName>
(
<bibRefCitation id="128305F61FC175F8849E9704F520CF6D" author="Edward, JC" journalOrPublisher="Japanese Journal of Applied Entomology and Zoology" pageId="11" pageNumber="86" pagination="310 - 312" title="Longidoruspisi n. sp. (Nematoda, Dorylaimoidea) associated with the rhizosphere of Pisumsativum, from Uttar Pradesh, India." url="https://doi.org/10.1303/jjaez.8.310" volume="8" year="1964">Edward et al. 1964</bibRefCitation>
), specimens from our populations 
<pageBreakToken id="6336DB944411A7BED5A586ED08276D91" pageId="8" pageNumber="83" start="start">revealed</pageBreakToken>
longer odontostyle (average 75 (72-79) and average 76 (74-78) vs 58 (56-61) 
<normalizedToken id="38F30B3BBEAEEDE6AE46321E53538A25" originalValue="μm">μm</normalizedToken>
) and odontophore (average 50 (45-53) and average 49 (46-53) vs average 42.7 (35-43) 
<normalizedToken id="C2572FE52F9F2E76C3728E2515B7DBD4" originalValue="μm">μm</normalizedToken>
); longer distances anterior end to guide (average 44 (42-46) vs average 32 (31-35) 
<normalizedToken id="549FBD518E73C1C1DCBBFE80171033D4" originalValue="μm">μm</normalizedToken>
) and nerve ring (average 147 (138-151) and 141 (132-150) vs average 133 
<normalizedToken id="3C4B7BD924B38C867DA6B2B3F39E611A" originalValue="μm">μm</normalizedToken>
); wider (average 10.6 vs 7.5 
<normalizedToken id="88E85CF0FD9FA664A8E7A7831E2C4A9F" originalValue="μm">μm</normalizedToken>
) and higher (5-6 vs 3.5 
<normalizedToken id="40C648CBD186E1F8753CB83329E6A9C9" originalValue="μm">μm</normalizedToken>
) lip region and lower 
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<normalizedToken id="83B68DAE9C5F90170985BC1D257F76CC" originalValue="c’">c'</normalizedToken>
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ratio (average 1.9 (1.5-2.0) and average 1.7 (1.5-1.8) vs average 2.5 (2.4-2.6). The morphometrics of our populations were more similar to the Iranian population of 
<taxonomicName id="4748C4B346D59FF3F8766EF9415D6FBD" lsidName="L. pisi" pageId="9" pageNumber="84" rank="species" species="pisi">L. pisi</taxonomicName>
(Saveh, Markazi province) for which a D2-D3 expansion domain of 28S rRNA gene sequence identical to ours is available (
<bibRefCitation id="91C5FFF7E9ABD66F9BA5300AB0653806" author="Pedram, M" journalOrPublisher="Nematology" pageId="11" pageNumber="86" pagination="949 - 960" title="Molecular and morphological characterisation of Xiphinemagranatum n. sp. and Longidoruspisi Edward, Misra &amp; Singh, 1964 (Dorylaimida: Longidoridae) from Iran." url="https://doi.org/10.1163/156854112X638406" volume="14" year="2012">Pedram et al. 2012</bibRefCitation>
). Differences in a few characters were observed, e.g. smaller a (average 125.2 (117.2-132.1) and 127.3 (119.8-132.9) vs 139.4 (134.8-144.6) and 
<normalizedToken id="DE5B909933C5CFB3354027A0867F64CE" originalValue="c’">c'</normalizedToken>
(average 1.9 (1.5-2.0) and average 1.7 (1.5-1.8) vs average 2.5 (2.3-2.9) values, larger diameter at anus level (average 22.9 (21-24.5) and average 20.5 (20-22) vs average 18.1 (16-19) 
<normalizedToken id="9D1F0C9B3EC5252CC786B98E1BF8DEB7" originalValue="μm">μm</normalizedToken>
) and slightly shorter tail (average 38.9 (36-43) and average 39.1 (37-42) vs average 45 (43-47) 
<normalizedToken id="E76FB847F146DBE1F366F181AFF331A4" originalValue="μm">μm</normalizedToken>
) compared to the latter population.
</paragraph>
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<paragraph id="A0978CB3F9A1A724DAC66392C06B0B17" pageId="9" pageNumber="84">
Table 4. Measurements of adults and juveniles of 
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(mean 
<normalizedToken id="861C29A100EFF2439660F2F7C6D8A277" originalValue="±">+/-</normalizedToken>
standard deviation, with range) from different crops in Bulgaria. All measurements in micrometers except body length (mm).
</paragraph>
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<th id="F16A67DED9C690A9D2E69DC8A16F3CA1" colspan="1" pageId="9" pageNumber="84" rowspan="4">Host Character</th>
<th id="4DB2C77347F7F5798165E7AB4BF0F20E" colspan="1" pageId="9" pageNumber="84" rowspan="1">Petrich</th>
<th id="CEE1E7AAE84A9A9C624D60766D920F8C" colspan="1" pageId="9" pageNumber="84" rowspan="1">Sandanski</th>
<th id="95264A019B3A61DE66656C51E4069B90" colspan="1" pageId="9" pageNumber="84" rowspan="1">Kromidovo</th>
<th id="52E7F53D18EF60B32C90A3644E0DA3FF" colspan="3" pageId="9" pageNumber="84" rowspan="1">Petrich</th>
</tr>
<tr id="2AD28D88ED5BA409538DAB67D3622893" pageId="9" pageNumber="84">
<th id="8780EBB20226C175F1B4D0D5EEDF9B1E" colspan="1" pageId="9" pageNumber="84" rowspan="1">
<taxonomicName id="496831938BB5D90AA378844F4826AE98" class="Magnoliopsida" family="Solanaceae" genus="Nicotiana" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Nicotiana tabacum" order="Solanales" pageId="9" pageNumber="84" phylum="Tracheophyta" rank="species" species="tabacum">Nicotiana tabacum</taxonomicName>
</th>
<th id="C2EA4AF524BCA954BF835843A0FAAAEC" colspan="2" pageId="9" pageNumber="84" rowspan="1">
<taxonomicName id="D80E05BD8763E80B2461524FF232A290" class="Magnoliopsida" family="Vitaceae" genus="Vitis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Vitis vinifera" order="Vitales" pageId="9" pageNumber="84" phylum="Tracheophyta" rank="species" species="vinifera">Vitis vinifera</taxonomicName>
</th>
<th id="386E2BB8E3A729CA5B37058BEF355F05" colspan="3" pageId="9" pageNumber="84" rowspan="1">
<taxonomicName id="19D5911E28506D327B86C6AB0CF12EC4" class="Magnoliopsida" family="Solanaceae" genus="Nicotiana" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Nicotiana tabacum" order="Solanales" pageId="9" pageNumber="84" phylum="Tracheophyta" rank="species" species="tabacum">Nicotiana tabacum</taxonomicName>
</th>
</tr>
<tr id="439382B2EDA1884C7AA8775147CCDFF7" pageId="9" pageNumber="84">
<th id="5577ABAFA1D59180BE93CDA53C9BFF62" colspan="1" pageId="9" pageNumber="84" rowspan="1">female</th>
<th id="AA5D817FAFDDED851322BDE4E7C8D1DA" colspan="1" pageId="9" pageNumber="84" rowspan="1">female</th>
<th id="EA54759C9AD02BD96740D9EF7558D7DE" colspan="1" pageId="9" pageNumber="84" rowspan="1">female</th>
<th id="F80256A3990B4D7F3DA2C6DB35EBBA34" colspan="1" pageId="9" pageNumber="84" rowspan="1">J1</th>
<th id="E88C485BF36CF401FED680ED983551FD" colspan="1" pageId="9" pageNumber="84" rowspan="1">J2</th>
<th id="12081FEFDF4992D32E54E447B1BB1BAD" colspan="1" pageId="9" pageNumber="84" rowspan="1">J3</th>
</tr>
<tr id="D97575440C30041B4B9CEB2BAFD095E6" pageId="9" pageNumber="84">
<th id="2933FA735E6921E2684B52F0C9DF4D36" colspan="1" pageId="9" pageNumber="84" rowspan="1">n = 9</th>
<th id="B902EA6F8F3D44A1BC3E988D8DEF95BC" colspan="1" pageId="9" pageNumber="84" rowspan="1">n = 6</th>
<th id="90819FC0AC01C83AC0B57DD6A07133AC" colspan="1" pageId="9" pageNumber="84" rowspan="1">n = 2</th>
<th id="6D35A475ACE4F84C9B788851485273BF" colspan="1" pageId="9" pageNumber="84" rowspan="1">n = 2</th>
<th id="F513CC289C8104F4E3082AB7604F747B" colspan="1" pageId="9" pageNumber="84" rowspan="1">n = 9</th>
<th id="DE8E29350936E0F004CA8A138689A16D" colspan="1" pageId="9" pageNumber="84" rowspan="1">n = 2</th>
</tr>
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</paragraph>
<caption id="7727EE05B9402A1BD63D0F8CA36BA13D" pageId="9" pageNumber="84">
<paragraph id="31828B91DA977D631A2E37CC20C037A6" pageId="9" pageNumber="84">
Figure 8. Bulgarian populations of 
<taxonomicName id="24039DF62793D70902F8510F6A21DC5D" class="Enoplea" family="Longidoridae" genus="Longidorus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Longidorus pisi" order="Dorylaimida" pageId="9" pageNumber="84" phylum="Nematoda" rank="species" species="pisi">Longidorus pisi</taxonomicName>
Edward, Misra &amp; Singh, 1964, anterior end of female (A) and male (B) female C, D vagina and part of reproductive system (different populations) 
<normalizedToken id="341A996930454801E78DA3A1D8F9BF31" originalValue="E–G">E-G</normalizedToken>
pre-rectum inclusions H labial region I vagina 
<normalizedToken id="5092EE57FFF96B646DAB77F587943AC7" originalValue="J–M">J-M</normalizedToken>
variations in tail shape, female (J, L, M) and male (K). Populations: A, L Petrich C, 
<normalizedToken id="3F34B90E44AFE01B0F11B47BBAEA1743" originalValue="E–G">E-G</normalizedToken>
, M Sandanski D, H, I, J Kromidovo B, K male specimens from a tobacco field, Petrich region. Scale bars: 20 
<normalizedToken id="819DC00DC36DC3D3B8BB6528C472BAF3" originalValue="μm">μm</normalizedToken>
(
<normalizedToken id="3E499EBB0FD6143C1F5B285FB753EF7C" originalValue="A–G">A-G</normalizedToken>
, 
<normalizedToken id="9E00889EA520B76C236BFE9C10200BD4" originalValue="J–M">J-M</normalizedToken>
); 12 
<normalizedToken id="A8CF81B372D7A7A9A1448FB604D8CA18" originalValue="μm">μm</normalizedToken>
(H, I).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="796FCB59A01EA05AFA856DECACEAE5E4" pageId="9" pageNumber="84" type="sequence and phylogenetic analyses">
<paragraph id="57182E7D1B9715854038FCCC46BD0650" pageId="9" pageNumber="84">Sequence and phylogenetic analyses.</paragraph>
<paragraph id="B60C2F7C9DC9607D9AFD262F45B5ABBC" pageId="9" pageNumber="84">
Three ribosomal DNA regions (D2-D3 expansion segments of 28S rRNA gene, 18S rRNA gene, and ITS1-5.8S-ITS2 regions) of 
<taxonomicName id="309A4C07457B93D0503A0AC7FD4475C5" lsidName="L. pisi" pageId="9" pageNumber="84" rank="species" species="pisi">L. pisi</taxonomicName>
were amplified and sequenced. The D2-D3 expansion segments of 28S rRNA gene sequences from all populations were identical to that of the Iranian population (JQ240274, 
<bibRefCitation id="C07BD3CA7D6A1146C31AC1C4FED9948C" author="Pedram, M" journalOrPublisher="Nematology" pageId="11" pageNumber="86" pagination="949 - 960" title="Molecular and morphological characterisation of Xiphinemagranatum n. sp. and Longidoruspisi Edward, Misra &amp; Singh, 1964 (Dorylaimida: Longidoridae) from Iran." url="https://doi.org/10.1163/156854112X638406" volume="14" year="2012">Pedram et al. 2012</bibRefCitation>
) and differed slightly (1 and 3 bp) from those of other populations (Greece (AY601569) and South Africa (AY601568), respectively) identified as 
<taxonomicName id="3F0BE07444E6582224EFBA71BB02736A" lsidName="L. latocephalus" pageId="9" pageNumber="84" rank="species" species="latocephalus">L. latocephalus</taxonomicName>
(
<bibRefCitation id="0D7A9EE649C3015077C979BA34840235" author="He, Y" journalOrPublisher="Nematology" pageId="11" pageNumber="86" pagination="111 - 124" title="A molecular phylogenetic approach to Longidoridae (Nematoda: Dorylaimida)." url="https://doi.org/10.1163/1568541054192108" volume="7" year="2005">He et al. 2005</bibRefCitation>
). In the phylogenetic analysis, all aforementioned sequences formed a clade with maximal Bayesian posterior probability (1.0) and showed a close relationship with 
<taxonomicName id="896F3D362A383A6BC9D06F0C0F41FC77" lsidName="L. mindanaoensis" pageId="9" pageNumber="84" rank="species" species="mindanaoensis">L. mindanaoensis</taxonomicName>
. The BLASTn search using 18S rRNA gene sequence revealed highest identity (99%) with five accessions (two 
<taxonomicName id="935247D1C16DAA146B03EB012E07C19A" class="Enoplea" family="Longidoridae" genus="Longidorus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Longidorus" order="Dorylaimida" pageId="9" pageNumber="84" phylum="Nematoda" rank="genus">Longidorus</taxonomicName>
(HQ735099 
<taxonomicName id="12BEF31BA77ED0E0DD90B3A72120052D" lsidName="L. mindanaoensis" pageId="9" pageNumber="84" rank="species" species="mindanaoensis">L. mindanaoensis</taxonomicName>
Coomans, Tandingan De Ley, Angsinco Jimenez &amp; De Ley, 2012 and AY283163 
<taxonomicName id="11F4D5B9E33C43899A479ED3AB621426" lsidName="L. ferrisi" pageId="9" pageNumber="84" rank="species" species="ferrisi">L. ferrisi</taxonomicName>
Robbins, Ye &amp; Pedram, 2009) and three 
<taxonomicName id="2545F2D53BBE2BA21D00D89B4C6FE336" class="Enoplea" family="Longidoridae" genus="Paralongidorus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paralongidorus" order="Dorylaimida" pageId="9" pageNumber="84" phylum="Nematoda" rank="genus">Paralongidorus</taxonomicName>
species (JN032586 
<taxonomicName id="6BE6D79D04AC1C7C58AF4631755D9C02" lsidName="P. bikanerensis" pageId="9" pageNumber="84" rank="species" species="bikanerensis">P. bikanerensis</taxonomicName>
(Lal &amp; Mathur, 1987) Siddiqi, Baujard &amp; Mounport, 1993; AJ875152 
<taxonomicName id="67BF429809CD17ED0AADCDF86DAC9F4A" lsidName="P. maximus" pageId="9" pageNumber="84" rank="species" species="maximus">P. maximus</taxonomicName>
(
<normalizedToken id="8B809CE7565CE395B63D96D5162D8A50" originalValue="Bütschli">Buetschli</normalizedToken>
, 1874) Siddiqi, 1964 and KJ427794 
<taxonomicName id="DA72CF47BB454CF0A056C303AE68A447" lsidName="P. rex" pageId="9" pageNumber="84" rank="species" species="rex">P. rex</taxonomicName>
<normalizedToken id="6E9811BA3A2D03C0C3C665FFBB08030F" originalValue="Andrássy">Andrassy</normalizedToken>
, 1986). A pairwise comparison of 
<taxonomicName id="7553ACEF834DEEC585DF91CC2FB12861" lsidName="L. pisi" pageId="9" pageNumber="84" rank="species" species="pisi">L. pisi</taxonomicName>
sequence with the closest sequences (AY283163, HQ735099, JN032586, AJ875152 and KJ427794) revealed 14-22 different nucleotides. The 18S rDNA phylogenetic tree is presented in Figure 10.
</paragraph>
<caption id="9163B3975CE61B13BF79CA098FA01CA0" pageId="9" pageNumber="84">
<paragraph id="F8621B4A95DA04259A3D0ADAB4F237F0" pageId="9" pageNumber="84">Figure 9. Phylogenetic tree using D2-D3 expansion segments of the 28S rRNA gene inferred from a Bayesian analysis with GTR+I+G model. Numbers represent the Bayesian posterior probabilities.</paragraph>
</caption>
<caption id="373CBB796F708320210FDF0E154A8E55" pageId="9" pageNumber="84">
<paragraph id="F914A0849BE8512F6950D9E4EEFB5100" pageId="9" pageNumber="84">Figure 10. Phylogenetic tree using 18S rRNA gene inferred from a Bayesian analysis with GTR+I+G model. Numbers represent the Bayesian posterior probabilities.</paragraph>
</caption>
</subSubSection>
</treatment>
</document>