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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">On the species status of the root-knot nematode <italic>Meloidogyne ulmi</italic>
Palmisano & Ambrogioni, 2000 (Nematoda, Meloidogynidae)</title>
<author><name sortKey="Ahmed, Mohammed" sort="Ahmed, Mohammed" uniqKey="Ahmed M" first="Mohammed" last="Ahmed">Mohammed Ahmed</name>
<affiliation><nlm:aff id="A1">National Plant Protection Organization, Wageningen Nematode Collection, P.O. Box 9102, 6700 HC Wageningen, The Netherlands</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Ghent University, Department of Biology, Ledeganckstraat 35, 9000 Ghent, Belgium</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Van De Vossenberg, Bart Tlh" sort="Van De Vossenberg, Bart Tlh" uniqKey="Van De Vossenberg B" first="Bart Tlh" last="Van De Vossenberg">Bart Tlh Van De Vossenberg</name>
<affiliation><nlm:aff id="A1">National Plant Protection Organization, Wageningen Nematode Collection, P.O. Box 9102, 6700 HC Wageningen, The Netherlands</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cornelisse, Chris" sort="Cornelisse, Chris" uniqKey="Cornelisse C" first="Chris" last="Cornelisse">Chris Cornelisse</name>
<affiliation><nlm:aff id="A1">National Plant Protection Organization, Wageningen Nematode Collection, P.O. Box 9102, 6700 HC Wageningen, The Netherlands</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Karssen, Gerrit" sort="Karssen, Gerrit" uniqKey="Karssen G" first="Gerrit" last="Karssen">Gerrit Karssen</name>
<affiliation><nlm:aff id="A1">National Plant Protection Organization, Wageningen Nematode Collection, P.O. Box 9102, 6700 HC Wageningen, The Netherlands</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Ghent University, Department of Biology, Ledeganckstraat 35, 9000 Ghent, Belgium</nlm:aff>
</affiliation>
</author>
</titleStmt>
<publicationStmt><idno type="wicri:source">PMC</idno>
<idno type="pmid">24363598</idno>
<idno type="pmc">3867104</idno>
<idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3867104</idno>
<idno type="RBID">PMC:3867104</idno>
<idno type="doi">10.3897/zookeys.362.6352</idno>
<date when="2013">2013</date>
<idno type="wicri:Area/Pmc/Corpus">000271</idno>
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</publicationStmt>
<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">On the species status of the root-knot nematode <italic>Meloidogyne ulmi</italic>
Palmisano & Ambrogioni, 2000 (Nematoda, Meloidogynidae)</title>
<author><name sortKey="Ahmed, Mohammed" sort="Ahmed, Mohammed" uniqKey="Ahmed M" first="Mohammed" last="Ahmed">Mohammed Ahmed</name>
<affiliation><nlm:aff id="A1">National Plant Protection Organization, Wageningen Nematode Collection, P.O. Box 9102, 6700 HC Wageningen, The Netherlands</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Ghent University, Department of Biology, Ledeganckstraat 35, 9000 Ghent, Belgium</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Van De Vossenberg, Bart Tlh" sort="Van De Vossenberg, Bart Tlh" uniqKey="Van De Vossenberg B" first="Bart Tlh" last="Van De Vossenberg">Bart Tlh Van De Vossenberg</name>
<affiliation><nlm:aff id="A1">National Plant Protection Organization, Wageningen Nematode Collection, P.O. Box 9102, 6700 HC Wageningen, The Netherlands</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cornelisse, Chris" sort="Cornelisse, Chris" uniqKey="Cornelisse C" first="Chris" last="Cornelisse">Chris Cornelisse</name>
<affiliation><nlm:aff id="A1">National Plant Protection Organization, Wageningen Nematode Collection, P.O. Box 9102, 6700 HC Wageningen, The Netherlands</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Karssen, Gerrit" sort="Karssen, Gerrit" uniqKey="Karssen G" first="Gerrit" last="Karssen">Gerrit Karssen</name>
<affiliation><nlm:aff id="A1">National Plant Protection Organization, Wageningen Nematode Collection, P.O. Box 9102, 6700 HC Wageningen, The Netherlands</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Ghent University, Department of Biology, Ledeganckstraat 35, 9000 Ghent, Belgium</nlm:aff>
</affiliation>
</author>
</analytic>
<series><title level="j">ZooKeys</title>
<idno type="ISSN">1313-2989</idno>
<idno type="eISSN">1313-2970</idno>
<imprint><date when="2013">2013</date>
</imprint>
</series>
</biblStruct>
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<front><div type="abstract" xml:lang="en"><label>Abstract</label>
<p>The root-knot nematode <italic>Meloidogyne ulmi</italic>
is synonymised with <italic>Meloidogyne mali</italic>
based on morphological and morphometric similarities, common hosts, as well as biochemical similarities at both protein and DNA levels. <italic>M. mali</italic>
was first described in Japan on <italic>Malus prunifolia</italic>
Borkh.; and <italic>M. ulmi</italic>
in Italy on <italic>Ulmus chenmoui</italic>
W.C. Cheng. Morphological and morphometric studies of their holo- and paratypes revealed important similarities in the major characters as well as some general variability in a few others. Host test also showed that besides the two species being able to parasitize the type hosts of the other, they share some other common hosts. Our study of the esterase and malate dehydrogenase isozyme phenotypes of some <italic>M. ulmi</italic>
populations gave a perfectly comparable result to that already known for <italic>M. mali</italic>
. Finally, phylogenetic studies of their SSU and LSU rDNA sequence data revealed that the two are not distinguishable at DNA level. All these put together, leave strong evidences to support the fact that <italic>M. ulmi</italic>
is not a valid species, but a junior synonym of <italic>M. mali</italic>
. Brief discussion on the biology and life cycle of <italic>M. mali</italic>
is given. An overview of all known hosts and the possible distribution of <italic>M. mali</italic>
in Europe are also presented.</p>
</div>
</front>
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</back>
</TEI>
<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">24363598</article-id>
<article-id pub-id-type="pmc">3867104</article-id>
<article-id pub-id-type="doi">10.3897/zookeys.362.6352</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Article</subject>
</subj-group>
</article-categories>
<title-group><article-title>On the species status of the root-knot nematode <italic>Meloidogyne ulmi</italic>
Palmisano & Ambrogioni, 2000 (Nematoda, Meloidogynidae)</article-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Ahmed</surname>
<given-names>Mohammed</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<xref ref-type="aff" rid="A2">2</xref>
</contrib>
<contrib contrib-type="author"><name><surname>van de Vossenberg</surname>
<given-names>Bart TLH</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Cornelisse</surname>
<given-names>Chris</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Karssen</surname>
<given-names>Gerrit</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<xref ref-type="aff" rid="A2">2</xref>
</contrib>
</contrib-group>
<aff id="A1"><label>1</label>
National Plant Protection Organization, Wageningen Nematode Collection, P.O. Box 9102, 6700 HC Wageningen, The Netherlands</aff>
<aff id="A2"><label>2</label>
Ghent University, Department of Biology, Ledeganckstraat 35, 9000 Ghent, Belgium</aff>
<author-notes><corresp>Corresponding author: Gerrit Karssen (<email xlink:type="simple">g.karssen@minlnv.nl</email>
)</corresp>
<fn fn-type="edited-by"><p>Academic editor: S. Subbotin</p>
</fn>
</author-notes>
<pub-date pub-type="collection"><year>2013</year>
</pub-date>
<pub-date pub-type="epub"><day>13</day>
<month>12</month>
<year>2013</year>
</pub-date>
<issue>362</issue>
<fpage>1</fpage>
<lpage>27</lpage>
<history><date date-type="received"><day>30</day>
<month>9</month>
<year>2013</year>
</date>
<date date-type="accepted"><day>2</day>
<month>12</month>
<year>2013</year>
</date>
</history>
<permissions><copyright-statement>Mohammed Ahmed, Bart TLH van de Vossenberg, Chris Cornelisse, Gerrit Karssen</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>
<abstract><label>Abstract</label>
<p>The root-knot nematode <italic>Meloidogyne ulmi</italic>
is synonymised with <italic>Meloidogyne mali</italic>
based on morphological and morphometric similarities, common hosts, as well as biochemical similarities at both protein and DNA levels. <italic>M. mali</italic>
was first described in Japan on <italic>Malus prunifolia</italic>
Borkh.; and <italic>M. ulmi</italic>
in Italy on <italic>Ulmus chenmoui</italic>
W.C. Cheng. Morphological and morphometric studies of their holo- and paratypes revealed important similarities in the major characters as well as some general variability in a few others. Host test also showed that besides the two species being able to parasitize the type hosts of the other, they share some other common hosts. Our study of the esterase and malate dehydrogenase isozyme phenotypes of some <italic>M. ulmi</italic>
populations gave a perfectly comparable result to that already known for <italic>M. mali</italic>
. Finally, phylogenetic studies of their SSU and LSU rDNA sequence data revealed that the two are not distinguishable at DNA level. All these put together, leave strong evidences to support the fact that <italic>M. ulmi</italic>
is not a valid species, but a junior synonym of <italic>M. mali</italic>
. Brief discussion on the biology and life cycle of <italic>M. mali</italic>
is given. An overview of all known hosts and the possible distribution of <italic>M. mali</italic>
in Europe are also presented.</p>
</abstract>
<kwd-group><label>Keywords</label>
<kwd>Morphological</kwd>
<kwd>morphometrics</kwd>
<kwd>esterase</kwd>
<kwd>malate dehydrogenase</kwd>
<kwd>Japan</kwd>
<kwd>Italy</kwd>
<kwd>Mierenbos</kwd>
<kwd><italic>Malus prunifolia</italic>
</kwd>
<kwd><italic>Ulmus chenmoui</italic>
</kwd>
<kwd>SSU rDNA</kwd>
<kwd>LSU rDNA</kwd>
</kwd-group>
</article-meta>
</front>
<body><sec><title>Introduction</title>
<p>The genus <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
comprises all root-knot nematodes. It contains over 100 described species (<xref ref-type="bibr" rid="B19">Karssen et al. 2013</xref>
). Its members are without a doubt the most widely distributed of all plant-parasitic nematodes (<xref ref-type="bibr" rid="B31">Sasser 1977</xref>
). This widespread distribution and their economic importance are primarily the reasons why the genus has been the subject of more research than any other plant-parasitic nematodes, including the cyst-forming nematodes (<xref ref-type="bibr" rid="B32">Sasser and Carter 1982</xref>
). Despite the numerous studies about their biology and taxonomy, their identification to the species level still pose a huge challenge to many diagnosticians (<xref ref-type="bibr" rid="B1">Blok and Powers 2009</xref>
) mostly because of their very small inter-specific morphological variation (<xref ref-type="bibr" rid="B18">Jepson 1987</xref>
).</p>
<p>In 2000, Palmisano and Ambrogioni described from Italy the root-knot nematode, <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
, from <italic><named-content content-type="taxon-name">Ulmus chenmoui</named-content>
</italic>
W.C. Chengon which it was found to induce large galls. For many years, elm remained the only known host of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
. According to the authors, the tree at the type locality was introduced from the Netherlands as part of a breeding programme focussed on resistance to the Dutch elm disease. The Netherlands, like many other countries in Europe and North America, as well as New Zealand, has for years been battling against the notorious Dutch elm disease. It was for this reason that the former Dutch phytopathological laboratory “Willie Commelin Scholten” (WCS), based in Baarn, was mandated with the research on Dutch elm disease. This breeding programme was later on moved to Wageningen at the former institute the Dorschkamp Research Institute for Forestry & Landscape Planning; and in this programme, elm trees from all over the world were tested. Trial field “Mierenbos”, a part of the Dorschkamp Research Institute for Forestry & Landscape Planning, was used for growing and improving resistant elm cultivars. It was from this trial field that resistant elm seedlings were sent to ten other European countries at the end of the breeding programme, among others, Italy in 1992 (<xref ref-type="bibr" rid="B10">Heybroek 1993</xref>
).</p>
<p>The first observation of galls on elm trees was already in 1960 at Baarn, and the associating nematode was diagnosed as <italic><named-content content-type="taxon-name">Meloidogyne arenaria</named-content>
</italic>
(Neal, 1889) Chitwood, 1949 by the National Plant Protection Organization (<xref ref-type="bibr" rid="B26">Oostenbrink 1961</xref>
). Interestingly, about that same period, a <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
species found parasitizing apple trees in Japan was also inadvertently misidentified as <italic><named-content content-type="taxon-name">Meloidogyne arenaria</named-content>
</italic>
, because this nematode species, like the one on elm bore some resemblance to <italic><named-content content-type="taxon-name">Meloidogyne arenaria</named-content>
</italic>
perineal patterns (<xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
). This Japanese species would later be described and named <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
by <xref ref-type="bibr" rid="B17">Itoh et al. (1969)</xref>
. In his comprehensive study on the host range of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
, <xref ref-type="bibr" rid="B36">Toida (1979)</xref>
associated this species with several other plant species belonging to different families, particularly the <named-content content-type="taxon-name">Rosaceae</named-content>
. Following its description, several studies have also been conducted on its taxonomy, ecology, damage and control (<xref ref-type="bibr" rid="B16">Inagaki 1978</xref>
), SEM studies of male and second-stage juvenile head morphology, and morphological variability of its different populations (<xref ref-type="bibr" rid="B25">Okamoto et al. 1983</xref>
). At the trial field “Mierenbos”, the first report of galling symptoms on <italic><named-content content-type="taxon-name">Ulmus</named-content>
</italic>
trees was in 1979 (<xref ref-type="bibr" rid="B2">Brinkman 1980</xref>
). Presently, all the <italic><named-content content-type="taxon-name">Ulmus</named-content>
</italic>
trees there are infected with <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
and are showing severe symptoms of root galling (<xref ref-type="bibr" rid="B22">Karssen et al. 2008</xref>
and <xref ref-type="bibr" rid="B21">Karssen 2009</xref>
).</p>
<p>In 2006, root samples of the dying type host apple containing <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
from the type locality in Japan sent by Dr. Takayuki Mizukubo were received at the Dutch National Plant Protection organization. About the same time, galled root samples of the type host of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
were obtained from Italy. They were propagated and maintained on the <italic><named-content content-type="taxon-name">Ulmus</named-content>
</italic>
× <italic>hollandica</italic>
variety “Wredei”. Juveniles isolated from the Japanese apple root samples were used for sequencing and the resulting SSU rRNA sequence was discovered to be almost identical to that of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
(<xref ref-type="bibr" rid="B12">Holterman et al. 2009</xref>
). Additionally, isozyme phenotypes of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
population from the trial field in Wageningen were also compared to that of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
from Japan (Karssen unpublished; <xref ref-type="bibr" rid="B29">Sakai and Mizukubo 2009</xref>
). Those also revealed similar patterns of esterase and malate dehydrogenase to that obtained for <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
(<xref ref-type="bibr" rid="B29">Sakai and Mizukubo 2009</xref>
). With these observed similarities, a closer look needed to be taken into these two species. Based on the evidences available to us now, we hypothesize that <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
probably entered Europe as <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
with elm rootstocks imported from Japan. Supporting this is the report that <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
, in addition to its numerous hosts, can also infect <italic><named-content content-type="taxon-name">Ulmus davidiana</named-content>
</italic>
var. <italic>japonica</italic>
(<xref ref-type="bibr" rid="B36">Toida 1979</xref>
).</p>
<p>The original description of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
differentiates it from <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
on the basis of characters that generally show high intraspecific variations. With the original description being the only paper written about <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
, all the known features so far are ones from the original description. On <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
, however, there have been quite a lot of research on the hosts, life cycle, ecology, detailed morphology, as well as their variations within species (<xref ref-type="bibr" rid="B36">Toida 1979</xref>
; <xref ref-type="bibr" rid="B16">Inagaki 1978</xref>
; <xref ref-type="bibr" rid="B25">Okamoto et al. 1983</xref>
).</p>
<p>The objectives of this current research, therefore, are:</p>
<p>i) to evaluate the morphological similarities between <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
.</p>
<p>ii) to search for other host plants, than <italic><named-content content-type="taxon-name">Ulmus</named-content>
</italic>
sp. present at the trail field “Mierenbos”.</p>
<p>iii) to test <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
on selected host plants on which <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
is already known to reproduce.</p>
<p>iv) analyze their biochemical similarities, at the protein and DNA levels.</p>
</sec>
<sec sec-type="materials|methods"><title>Materials and methods</title>
<sec sec-type="Morphology and morphometrics"><title>Morphology and morphometrics</title>
<p>Paratype slides of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
used for morphological and morphometric studies were obtained from Dr. Z.A. Handoo of the USDA Nematode Collection. In addition to these, we obtained <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
specimens on slides taken to the USDA by Dr. Ichinohe during his visit in 1958 as well as additional specimens of males, second-stage juveniles and females stored in formalin that were only recently isolated from root samples sent to USDA by Ichinohe during that same visit. Also, by courtesy of Dr. Hiromichi Sakai and Shigeyuki Sekimoto, paratypes that were deposited at the National Agriculture and Food Research Organization, Agricultural Research Center (Kannondai, Tsukuba, Ibaraki, Japan), the then Central Agricultural Experiment Station were also obtained. All slides were observed using compound light microscope (DM 2500, LEICA) equipped with differential interference contrast (DIC), and camera (DC 300F, LEICA) for taking images. Comparisons of morphological and morphometric characters were based on the most differential characters, previously used by <xref ref-type="bibr" rid="B20">Karssen (2002)</xref>
.</p>
</sec>
<sec sec-type="Host test"><title>Host test</title>
<p>This is a combination of sampling undertaken in 2011 and 2012 at the former trial field “Mierenbos” on several plant species and a subsequent greenhouse experiment involving some important plant species already associated with <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
in previous studies (<xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
; <xref ref-type="bibr" rid="B36">Toida 1979</xref>
). Host herein is defined as a plant on which the nematode can reproduce, after a successful penetration.</p>
</sec>
<sec sec-type="Isozyme analysis"><title>Isozyme analysis</title>
<p>Esterase and malate dehydrogenase isozymes were analysed for <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
sampled at “Mierenbos”, following the method described by <xref ref-type="bibr" rid="B23">Karssen et al. (1995)</xref>
. In summary, young females were isolated from roots into an isotonic (0.9%) salt solution. This was followed by a desalting step which involved transfer of the females from the NaCl solution to a reagent-grade water on ice for few minutes. Females were then singly transferred into sample wells containing 0.6 µl extraction buffer. With the aid of a small glass rod, the females in the wells were crushed; and the macerated females were then loaded into sample applicators (0.3 µl per well). All twelve wells, with the exception of 6 and 7, were loaded with our test samples of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
. <italic><named-content content-type="taxon-name">Meloidogyne javanica</named-content>
</italic>
was used as reference in wells 6 and 7.</p>
<p>Electrophoresis was run using the PhastSystem (Pharmacia Ltd, Uppsala, Sweden) and the gels were subsequently stained in a Petri dish and placed in an incubator at 37 °C. Staining for non-specific esterase (EST; EC 3.1.1.1) was allowed to stand for 60 minutes whiles that for malate dehydrogenase (MDH; EC 1.1.1.37) stayed for 5 minutes.</p>
<p>Following staining, the gels were rinsed with distilled water and fixed for 5 minutes in a 10% acetic acid / 10 % glycerol/ 80 % distilled water solution. Pictures of the gels were taken by placing them on a glass surface illuminated from below.</p>
</sec>
<sec sec-type="Molecular analysis"><title>Molecular analysis</title>
<p>Already published sequences of both <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
(<xref ref-type="bibr" rid="B12">Holterman et al. 2009</xref>
) together with our own sequence of the latter were included in this analysis.</p>
</sec>
<sec sec-type="DNA extraction"><title>DNA extraction</title>
<p>Nucleic acids were isolated from single male or second-stage juveniles of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
populations taken from “Mierenbos” and type populations kept in culture at the Dutch National Plant Protection Organization on an elm tree (<italic><named-content content-type="taxon-name">Ulmus</named-content>
</italic>
× <italic>hollandica</italic>
Mill “Wredei”). Genomic DNAs were isolated from these samples using High Pure PCR Template Preparation Kit (<ext-link ext-link-type="uri" xlink:href="http://www.roche-applied-science.com">www.roche-applied-science.com</ext-link>
, Cat. No. 11796828001, Version 16.0) protocol for isolation of nucleic acids from Mammalian Tissue with slight modification in the first step to suit nematode DNA isolation (150 µl tissue lysis buffer added to 50 µl sterile water containing nematodes, minimum protease incubation time of 16 hours and elution volume of 50 µl).</p>
</sec>
<sec sec-type="PCR and sequencing"><title>PCR and sequencing</title>
<p>Amplification of 1000 base pairs (bp) of the large ribosomal subunit (LSU) (28S) was performed using primer set 28–81for (forward) 5’-TTAAGCATATCATTTAGCGGAGGAA-3’ and 28–1006rev (reverse) 5’-GTTCGATTAGTCTTTCGCCCCT-3’ described by <xref ref-type="bibr" rid="B13">Holterman et al. (2008)</xref>
.</p>
<p>To amplify the nearly full length sequence of the small ribosomal subunit (SSU) (18S), two partially overlapping fragments were generated using three universal primers and one nematode-specific primer (1912R) described by <xref ref-type="bibr" rid="B14">Holterman et al. (2006)</xref>
. The latter’s inclusion was to avoid amplification of non-target eukaryotic SSU rDNA, for example from fungal spores attached to the nematode cuticle. The primers 988F (forward) (5’-CTCAAAGATTAAGCCATGC-3’) and 1912R (reverse) (5’-TTTACGGTCAGAACTAGGG-3’) were used to amplify the first fragment. The second fragment was amplified with primers 1813F (foward) (5’-CTGCGTGAGAGGTGAAAT-3’) and 2646R (reverse) (5’-GCTACCTTGTTACGACTTTT-3). Each PCR reaction mixture contained Molecular Grade Water (MGW)–DNase RNase free water (Sigma-Aldrich, Saint Louis, USA), 1x PCR buffer (incl. 2.0 mM MgCl<sub>2</sub>
, Roche), dNTPs (0.2 mM each), 0.24 µM of each primer, FastStart Taq DNA polymerase (1 U/µl, Roche) and 1 µl of the template DNA. The final reaction volume was 25 µl. PCR was performed in C1000 touch thermal cycler (Bio-Rad) with the following amplification condition: 15 min at 95°C; 5 cycles of 30 sec at 94°C, 30 sec at 45°C and 30 sec at 72°C; 35 cycles of 30 sec at 94°C, 30 sec at 54°C and 30 sec at 72°C; final extension for 5 min at 72°C. To test for amplification and the quality of PCR products, 5 μl of the PCR products mixed with 1 µl 6x Bromophenol Blue Loading solution (Promega, Madison, USA) were subjected to electrophoresis and SYBR safe (Invitrogen, Carlsbad, USA) staining on a 1.5 % agarose gel by standard methods (<xref ref-type="bibr" rid="B30">Sambrook et al. 1989</xref>
) along with a 1kb-plus DNA ladder (Invitrogen, Carlsbad, USA) to size fragments. PCR products were imaged under UV light using a GeneGenious gel imaging system (Syngene, Cambridge, United Kingdom).</p>
<p>PCR products were purified after amplification using QIAquick PCR Purification Kit (Qiagen), and the genomic DNA concentration measured using a ND1000 spectrophotometer (NanoDrop). This was followed by a cycle sequencing reaction in a final volume of 20 µl (molecular grade water (Sigma-Aldrich, Saint Louis, USA), BigDye Terminator v1.1, 1x sequencing buffer, purified PCR product and 0.5 µM template-specific forward or reverse primers). Cycling reactions were carried out separately for each of forward and reverse primers. The reaction programme was set for 1 min at 96°C, 25x (10 sec 96°C, 5 sec 50°C, 2.5 min 60°C), 1 min 20°C. The cycle sequence products were cleaned up using DyeEx 2.0 Spin Kit (Qiagen) and run on a multi-capillary 3500 Genetic Analyzer DNA sequencer (Applied Biosystems, Carlsbad, USA).</p>
</sec>
<sec sec-type="Sequence alignment and phylogenetic analyses"><title>Sequence alignment and phylogenetic analyses</title>
<p>Trace files of D2–D3 expansion segments of 28S and 18S-rRNA genes were assembled into contigs and amplification primer sequences trimmed using Geneouis 6.1.6 (Biomatters, New Zealand). Additional trimming was performed when needed, to obtain high quality consensus sequence data. Conflicts in the consensus sequence were assessed visually and corrected where possible. The sequences were aligned with selected sequences of other species from GeneBank using MAFFT alignment (<xref ref-type="bibr" rid="B24">Katoh et al. 2002</xref>
) within the programme Geneious 6.1.6 (Biomatters New Zealand) for both 28S and 18S-rRNA. Alignments were improved manually. Analysis of phylogeny of the sequence data set was performed with Bayesian inference (BI) using MrBayes 3.2.1 (<xref ref-type="bibr" rid="B15">Huelsenback and Ronquist 2001</xref>
). The optimal model for nucleotide substitution was obtained using JModelTest ver. 2.1.3 (<xref ref-type="bibr" rid="B5">Darriba et al. 2012</xref>
) with AIC, AICc, BIC and DT defaulted in JModelTest. For SSU sequence, analysis of Bayesian inference was performed with a random starting tree and four Markov chains for 1 × 10<sup>6</sup>
under the model TYMef + I. Trees were sampled at interval of 100 generations. Two independent runs were performed for each analysis. The first 100,000 generations were discarded as burn-ins, and the remaining trees combined to generate 50% majority rule consensus tree which represent posterior probabilities. The same parameter settings were used for LSU phylogenetic analysis, but under the model TYM + G.</p>
</sec>
</sec>
<sec sec-type="Results"><title>Results</title>
<sec sec-type="Morphology"><title>Morphology</title>
<p>The following are the observations made on selected features considered to be the most differential for species discrimination among members of the genus <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
(<xref ref-type="bibr" rid="B18">Jepson 1987</xref>
; <xref ref-type="bibr" rid="B20">Karssen 2002</xref>
). <xref ref-type="table" rid="T3">Table 3</xref>
shows a comparison of our observations of the most important characters with the ones mentioned in the original descriptions.</p>
<table-wrap id="T1" orientation="portrait" position="float"><label>Table 1.</label>
<caption><p>The various forms of type specimens of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
studied and their sources.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="2" colspan="1">Form</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
</th>
</tr>
<tr><th rowspan="1" colspan="1">Sex/stage</th>
<th rowspan="1" colspan="1">Source</th>
<th rowspan="1" colspan="1">Sex/stage</th>
<th rowspan="1" colspan="1">Source</th>
</tr>
<tr><td rowspan="1" colspan="1">Holotype</td>
<td rowspan="1" colspan="1">1 female</td>
<td rowspan="1" colspan="1">Shigeyuki Sekimoto</td>
<td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">–</td>
</tr>
<tr><td rowspan="1" colspan="1">Allotype<xref ref-type="fn" rid="T1TN1">*</xref>
</td>
<td rowspan="1" colspan="1">1 male</td>
<td rowspan="1" colspan="1">Shigeyuki Sekimoto</td>
<td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">–</td>
</tr>
<tr><td rowspan="1" colspan="1">Paratype</td>
<td rowspan="1" colspan="1">17 perineal patterns</td>
<td rowspan="1" colspan="1">Dr. Zafar A. Handoo</td>
<td rowspan="1" colspan="1">2 PP & 1 anterior part</td>
<td rowspan="1" colspan="1">Dr. Zafar A. Handoo</td>
</tr>
<tr><td rowspan="1" colspan="1">Paratype</td>
<td rowspan="1" colspan="1">3 males</td>
<td rowspan="1" colspan="1">Dr. Zafar A. Handoo</td>
<td rowspan="1" colspan="1">2 males</td>
<td rowspan="1" colspan="1">Dr. Zafar A. Handoo</td>
</tr>
<tr><td rowspan="1" colspan="1">Paratype</td>
<td rowspan="1" colspan="1">4 juveniles</td>
<td rowspan="1" colspan="1">Dr. Zafar A. Handoo</td>
<td rowspan="1" colspan="1">3 juveniles</td>
<td rowspan="1" colspan="1">Dr. Zafar A. Handoo</td>
</tr>
<tr><td rowspan="1" colspan="1">Paratype</td>
<td rowspan="1" colspan="1">1 male</td>
<td rowspan="1" colspan="1">Dr. Hiromichi Sakai</td>
<td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">–</td>
</tr>
<tr><td rowspan="1" colspan="1">Paratype</td>
<td rowspan="1" colspan="1">1 juvenile</td>
<td rowspan="1" colspan="1">Dr. Hiromichi Sakai</td>
<td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">–</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="T1TN1"><p>*Slide marked as allotype, however at present not recognized by the ICZN, i.e. it is a paratype.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T2" orientation="portrait" position="float"><label>Table 2.</label>
<caption><p>Plant species included in the host plant test with <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
in the greenhouse.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Family</th>
<th rowspan="1" colspan="1">Plant species</th>
</tr>
<tr><td rowspan="2" colspan="1"><named-content content-type="taxon-name">Brassicaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Brassica oleracea</named-content>
</italic>
L. var. Gemmifera (cabbage)</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Brassica pekinensis</named-content>
</italic>
(Lour.) Rupr. (celery cabbage)</td>
</tr>
<tr><td rowspan="2" colspan="1"><named-content content-type="taxon-name">Rosaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Malus pumila</named-content>
</italic>
“M9” (apple)</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Rosa hybrida</named-content>
</italic>
L. (rose)</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Fabaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Trifolium repens</named-content>
</italic>
L. (white clover)</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Solanaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Solanum lycopersicum</named-content>
</italic>
L. (tomato)</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Ulmaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Ulmus glabra</named-content>
</italic>
Huds. (wych elm)</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T3" orientation="portrait" position="float"><label>Table 3.</label>
<caption><p>Observations of the differential characteristics of female, male and second-stage juveniles of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
types in comparison with their interpretation in the original description.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Species</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
</th>
</tr>
<tr><th rowspan="1" colspan="1">Character</th>
<th rowspan="1" colspan="1">Described</th>
<th rowspan="1" colspan="1">Observed</th>
<th rowspan="1" colspan="1">Described</th>
<th rowspan="1" colspan="1">Observed</th>
</tr>
<tr><td rowspan="1" colspan="5"><bold>Female</bold>
</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet knobs</td>
<td rowspan="1" colspan="1">Well developed knobs that tend to slope backward or forward in the ratio of 16 to 8</td>
<td rowspan="1" colspan="1">Rounded to pear-shaped knobs, set off and slightly anteriorly concave to backwardly sloping</td>
<td rowspan="1" colspan="1">Knobs rounded to transversely ovoid, slightly concave anteriorly</td>
<td rowspan="1" colspan="1">Rounded knobs that are slightly anteriorly concave and offset</td>
</tr>
<tr><td rowspan="1" colspan="1">Perineal pattern</td>
<td rowspan="1" colspan="1">Oval, made up of smooth striae, finely spaced, dorsal arch low and flat. Phasmids large, lateral field clearly marked with single or double incisures</td>
<td rowspan="1" colspan="1">Oval, dorsal arch low to slightly high, rounded to square shaped.
Phasmids distinct. Lateral field marked by breaks in the striae or showing indistinct lateral lines</td>
<td rowspan="1" colspan="1">Oval, dorsal arch flattened to medium high, rounded or somewhat square, phasmids conspicuous, lateral field indistinct or marked by folds, sometimes by lateral lines on one or both sides</td>
<td rowspan="1" colspan="1">Oval, dorsal arch low to slightly high, rounded to square shaped.
Phasmids distinct. Lateral field marked by breaks in the striae or showing indistinct lateral lines</td>
</tr>
<tr><td rowspan="1" colspan="5"><bold>Male</bold>
</td>
</tr>
<tr><td rowspan="1" colspan="1">Head shape</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">Head weakly offset, head cap low and slightly narrower than the postlabial region. Postlabial incisures absent</td>
<td rowspan="1" colspan="1">Head slightly set off, labial cap shallowly rounded, one-fifth to one-fourth as high as postlabial region</td>
<td rowspan="1" colspan="1">Head weakly offset, head cap low and slightly narrower than the postlabial region. Postlabial incisures absent</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet knobs</td>
<td rowspan="1" colspan="1">Knobs rounded</td>
<td rowspan="1" colspan="1">Backwardly sloping with rounded to pear shaped knobs</td>
<td rowspan="1" colspan="1">Knobs rounded to pear shaped more or less backwardly sloping</td>
<td rowspan="1" colspan="1">Backwardly sloping with rounded to pear shaped knobs</td>
</tr>
<tr><td rowspan="1" colspan="5"><bold>Second-stage juvenile</bold>
</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet knobs</td>
<td rowspan="1" colspan="1">Knobs backwardly sloped</td>
<td rowspan="1" colspan="1">Small rounded knobs, slightly backwardly sloping</td>
<td rowspan="1" colspan="1">Knobs rounded and set off from shaft</td>
<td rowspan="1" colspan="1">Small rounded knobs, backwardly sloping</td>
</tr>
<tr><td rowspan="1" colspan="1">Tail shape</td>
<td rowspan="1" colspan="1">Short</td>
<td rowspan="1" colspan="1">Conical with a broad to finely pointed tip</td>
<td rowspan="1" colspan="1">Conical, tapering to a finely rounded almost pointed terminus or broader and rounded at the tip</td>
<td rowspan="1" colspan="1">Conical and tapers to a broadly or finely pointed tip</td>
</tr>
<tr><td rowspan="1" colspan="1">Hyaline tail part</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">Constrictions present along hyaline part, length short or long. Anterior part clearly delimitated</td>
<td rowspan="1" colspan="1">Cuticular constrictions present along hyaline tail terminus, variable in length</td>
<td rowspan="1" colspan="1">Constrictions present along hyaline part, anterior part clearly delimitated</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec sec-type="Female"><title>Female</title>
<sec sec-type="Perineal pattern"><title>Perineal pattern</title>
<p>The general shape of the perineal pattern in both species studied ranged from low rounded to oval. The dorsal arch of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
was mostly low rounded with very few instances where some specimens showed relatively high square patterns. Lateral field was marked by change in direction or breaks in striae resulting in what would appear as weak lateral lines. The double lateral lines mentioned in the description of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
were not observed in the studied specimens. The interphasmidial distance in both species was about the same as their corresponding vulva slit lengths. As mentioned in the description, the phasmids were distinct but did not appear large when observed at the correct focus. However, attempting to observe them at the same (relatively deeper) focus as the vulva slit makes them look larger and even farther apart than they really are, due to the diagonally sloping phasmid canals.</p>
</sec>
<sec sec-type="Stylet"><title>Stylet</title>
<p>Same variations in stylet knobs shape as described in <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
were observed for <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
paratypes i.e. slightly backwardly sloping to anteriorly concave, with the former being the more frequent. Such variations, however, cannot be mentioned about the <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
paratype since there was only a single anterior part of the female on the slides we obtained. We therefore supplemented it with specimens taken from samples from the “Mierenbos”, where the type host originated from. This population showed similar variation as described for <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
, but not reported in <xref ref-type="table" rid="T3">Table 3</xref>
. Our observation of the shape of the stylet itself was typical of the genus, i.e. straight shaft with a slightly dorsally curved cone.</p>
</sec>
<sec sec-type="Secretory-excretory pore"><title>Secretory-excretory pore</title>
<p>S-E pore position measured from the anterior end showed quite some variations. Nevertheless, all measurements taken for both species fell within the range described for <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
. This character in <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
description was measured on the basis of the number of annuli counted from the anterior end to the one bearing the S-E pore.</p>
</sec>
</sec>
<sec sec-type="Male"><title>Male</title>
<sec sec-type="Head region"><title>Head region</title>
<p>Under light microscope, both species have the same head outline. This was already illustrated in the descriptions of the two species (<xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
; <xref ref-type="bibr" rid="B27">Palmisano and Ambrogioni 2000</xref>
). The head cap in both species is low. The presence of lip annuli mentioned in <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
was observed on some of the paratypes studied as well as in some of the additional specimens included later from “Mierenbos”. The post-labial cephalic region slightly set off from the remainder of the body. SEM observation of the <italic>en face</italic>
view of the lip region was not part of this study. Nevertheless, this will be discussed further on in this work based on previous study conducted by <xref ref-type="bibr" rid="B38">Yaegashi and Okamoto (1981)</xref>
as well as the original description of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
.</p>
</sec>
<sec sec-type="Stylet"><title>Stylet</title>
<p>The stylet moderately slender. Conus with bluntly pointed tip. The shaft width the same along its entire length, although in some specimens it appeared to be broader close to the junction with the knobs. Individual knobs rounded to pear shaped. Knobs backwardly sloping in both species.</p>
</sec>
<sec sec-type="Lateral field"><title>Lateral field</title>
<p>The lateral field marked by four incisures. In most of the specimens studied, the outer lines appeared areolated along most part of the body. No difference in the number of lateral incisures was observed along the body, except at the anterior part where it reduces to two and gradually fades out further anterior.</p>
</sec>
<sec sec-type="Hemizonid position relative to S-E pore"><title>Hemizonid position relative to S-E pore</title>
<p>Although not considered to be of any diagnostic significance, this character remained fairly consistent in all specimens studied. The hemizonid always occurred anterior to the S-E pore, at slightly varying distances.</p>
</sec>
<sec sec-type="Second-stage juveniles"><title>Second-stage juveniles</title>
<p>Examination of the second-stage juvenile characters was based on six <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
type specimens and two of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
.</p>
</sec>
<sec sec-type="Head region"><title>Head region</title>
<p>Head slightly set off from the rest of the body, with a low lip. Post-labial region lacking any annule.</p>
</sec>
<sec sec-type="Stylet"><title>Stylet</title>
<p>Stylet somewhat slender, with conus terminating in a fine tip, in both species. Stylet knobs small and rounded; slightly backwardly sloping.</p>
</sec>
<sec sec-type="Hemizonid position relative to S-E pore"><title>Hemizonid position relative to S-E pore</title>
<p>Contrary to the condition in males, hemizonid always located behind the S-E pore in second-stage juveniles. However, the exact position is variable.</p>
</sec>
<sec sec-type="Tail"><title>Tail</title>
<p>Tail mostly straight, ranging from short to medium; with a fine to bluntly rounded tip. Hyaline tail terminus with varying length, anterior part distinctly delimitated.</p>
</sec>
</sec>
<sec sec-type="Morphometrics"><title>Morphometrics</title>
<sec sec-type="Females"><title>Females</title>
<p>Almost all our average measurements were within the range of those in the original descriptions (<xref ref-type="table" rid="T4">Tables 4</xref>
<xref ref-type="table" rid="T5">–</xref>
<xref ref-type="table" rid="T6">6</xref>
). In the case of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
, measured values of the female anterior part are based only on a single paratype specimen. Useful differential characters like the stylet length, stylet knob widths and stylet knob heights showed great similarities. From the perineal patterns, measurements of all the known important features also gave comparable values with those in the descriptions. Interphasmidial distance and the vulva slit were in most cases similar, rarely significantly different. In <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
, these two measurements were almost identical. There was however, a slight difference in these two measurements from <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
types (<xref ref-type="table" rid="T4">Table 4</xref>
), probably because only two perineal patterns were studied.</p>
<table-wrap id="T4" orientation="portrait" position="float"><label>Table 4.</label>
<caption><p>Morphometrics of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
females in comparison with the original descriptions. All measurements are in µm and in the form: mean ± sd. (range).</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Species</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
</th>
</tr>
<tr><th rowspan="1" colspan="1">Character</th>
<th rowspan="1" colspan="1">Described</th>
<th rowspan="1" colspan="1">Observed</th>
<th rowspan="1" colspan="1">Described</th>
<th rowspan="1" colspan="1">Observed</th>
</tr>
<tr><td rowspan="1" colspan="1">N</td>
<td rowspan="1" colspan="1">25</td>
<td rowspan="1" colspan="1">17</td>
<td rowspan="1" colspan="1">30</td>
<td rowspan="1" colspan="1">2<xref ref-type="fn" rid="T4TN1">*</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1">Body length</td>
<td rowspan="1" colspan="1">847
(684–1044)</td>
<td rowspan="1" colspan="1">762 ± 115
(608–890)</td>
<td rowspan="1" colspan="1">771 ± 140
(568–1043)</td>
<td rowspan="1" colspan="1">_</td>
</tr>
<tr><td rowspan="1" colspan="1">Body width</td>
<td rowspan="1" colspan="1">660
(540–864)</td>
<td rowspan="1" colspan="1">570 ± 122
(372–700)</td>
<td rowspan="1" colspan="1">618 ± 152
(357–1007)</td>
<td rowspan="1" colspan="1">_</td>
</tr>
<tr><td rowspan="1" colspan="1">Neck length</td>
<td rowspan="1" colspan="1">166 ± 43.7
(90–252)</td>
<td rowspan="1" colspan="1">165 ± 62
(60–265)</td>
<td rowspan="1" colspan="1">165 ± 67
(58–382)</td>
<td rowspan="1" colspan="1">205</td>
</tr>
<tr><td rowspan="1" colspan="1">Neck diameter</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">100 ± 34.3
(48–160)</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">152</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet length</td>
<td rowspan="1" colspan="1">15
(13–17)</td>
<td rowspan="1" colspan="1">11.9 ± 1.8
(7.7–15.4)</td>
<td rowspan="1" colspan="1">14.2 ± 1.0
(12.0–15.7)</td>
<td rowspan="1" colspan="1">13.4</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet knob height</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">1.6 ± 0.3
(1–2.2)</td>
<td rowspan="1" colspan="1">1.8 ± 0.6
(1.1–3.9)</td>
<td rowspan="1" colspan="1">1.6</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet knob width</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">3.2 ± 0.4
(2.6–3.8)</td>
<td rowspan="1" colspan="1">3.5 ± 0.7
(2.6–5.2)</td>
<td rowspan="1" colspan="1">3.2</td>
</tr>
<tr><td rowspan="1" colspan="1">DGO</td>
<td rowspan="1" colspan="1">5.5
(4–7)</td>
<td rowspan="1" colspan="1">4.3 ± 1.5
(2.2–6.7)</td>
<td rowspan="1" colspan="1">4.6 ± 0.8
(3.3–6.5)</td>
<td rowspan="1" colspan="1">3.9</td>
</tr>
<tr><td rowspan="1" colspan="1">S-E pore</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">32.8 ± 5.5
(25–43.5)</td>
<td rowspan="1" colspan="1">32.3 ± 7.8
(15.7–45.1)</td>
<td rowspan="1" colspan="1">36.5</td>
</tr>
<tr><td rowspan="1" colspan="1">Metacorpus</td>
<td rowspan="1" colspan="1">110
(90–147)</td>
<td rowspan="1" colspan="1">103 ± 7.9
(90–117)</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">92.8</td>
</tr>
<tr><td rowspan="1" colspan="1">Metacorpus length</td>
<td rowspan="1" colspan="1">39
(32–44)</td>
<td rowspan="1" colspan="1">40.4 ± 4.5
(32–50)</td>
<td rowspan="1" colspan="1">42.6 ± 6.5
(32.7–58.8)</td>
<td rowspan="1" colspan="1">39.7</td>
</tr>
<tr><td rowspan="1" colspan="1">Metacorpus diameter</td>
<td rowspan="1" colspan="1">49
(40–73)</td>
<td rowspan="1" colspan="1">39.7 ± 6.5
(29–47)</td>
<td rowspan="1" colspan="1">40.9 ± 7.0
(31.3–59.0)</td>
<td rowspan="1" colspan="1">36.5</td>
</tr>
<tr><td rowspan="1" colspan="1">Metacorpus valve length</td>
<td rowspan="1" colspan="1">12
(11–13)</td>
<td rowspan="1" colspan="1">13.0 ± 1.1
(11.5–15.4)</td>
<td rowspan="1" colspan="1">12.4 ± 1.0
(11.1–14.4)</td>
<td rowspan="1" colspan="1">17.9</td>
</tr>
<tr><td rowspan="1" colspan="1">Metacorpus valve width</td>
<td rowspan="1" colspan="1">10
(9–11)</td>
<td rowspan="1" colspan="1">9.5 ± 1.3
(7–11.2)</td>
<td rowspan="1" colspan="1">9.7 ± 1.4
(7.2–12.4)</td>
<td rowspan="1" colspan="1">10.2</td>
</tr>
<tr><td rowspan="1" colspan="1">Vulva – anus distance</td>
<td rowspan="1" colspan="1">17 ± 1.8
(14–22)</td>
<td rowspan="1" colspan="1">19.1 ± 2.6
(12.8–22.4)</td>
<td rowspan="1" colspan="1">19.0 ± 1.9
(15.0–22.2)</td>
<td rowspan="1" colspan="1">19.2 ± 0.8
(18.8–19.8)</td>
</tr>
<tr><td rowspan="1" colspan="1">Interphasmidial distance</td>
<td rowspan="1" colspan="1">22 ± 3.5
(17–29)</td>
<td rowspan="1" colspan="1">24.8 ± 4.7
(17.6–35.2)</td>
<td rowspan="1" colspan="1">19.2 ± 3.8
(13.7–28.9)</td>
<td rowspan="1" colspan="1">22.4 ± 4.5
(19.2–25.6)</td>
</tr>
<tr><td rowspan="1" colspan="1">Level of phasmids to vulva</td>
<td rowspan="1" colspan="1">25 ± 2.4
(19–31)</td>
<td rowspan="1" colspan="1">27.4 ± 2.8
(24–33.9)</td>
<td rowspan="1" colspan="1">25.1 ± 4.2
(15.7–39.2)</td>
<td rowspan="1" colspan="1">27.6 ± 0.9
(26.9–28.2)</td>
</tr>
<tr><td rowspan="1" colspan="1">Level of phasmids to anus</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">8.1 ± 2.5
(5.1–12.8)</td>
<td rowspan="1" colspan="1">6.9 ± 2.2
(2.6–15.7)</td>
<td rowspan="1" colspan="1">8.7 ± 1.3
(7.7–9.6)</td>
</tr>
<tr><td rowspan="1" colspan="1">Vulva slit length</td>
<td rowspan="1" colspan="1">18 ± 2.5
(12–24)</td>
<td rowspan="1" colspan="1">24.4 ± 3.3
(16–28.2)</td>
<td rowspan="1" colspan="1">22.0 ± 2.9
(17–28.7)</td>
<td rowspan="1" colspan="1">24.5 ± 1.1
(23.7–25.3)</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="T4TN1"><p>*Two perineal patterns and a single anterior part.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T5" orientation="portrait" position="float"><label>Table 5.</label>
<caption><p>Morphometrics of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
males in comparison with the original descriptions. All measurements are in µm and in the form: mean ± sd. (range).</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Species</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
</th>
</tr>
<tr><th rowspan="1" colspan="1">Character</th>
<th rowspan="1" colspan="1">Described</th>
<th rowspan="1" colspan="1">Observed</th>
<th rowspan="1" colspan="1">Described</th>
<th rowspan="1" colspan="1">Observed</th>
</tr>
<tr><td rowspan="1" colspan="1">N</td>
<td rowspan="1" colspan="1">25</td>
<td rowspan="1" colspan="1">3</td>
<td rowspan="1" colspan="1">30</td>
<td rowspan="1" colspan="1">2</td>
</tr>
<tr><td rowspan="1" colspan="1">Body length</td>
<td rowspan="1" colspan="1">1447
(1270–1630)</td>
<td rowspan="1" colspan="1">1428 ± 41.0
(1380–1452)</td>
<td rowspan="1" colspan="1">1462 ± 190
(1053–1776)</td>
<td rowspan="1" colspan="1">1455 ± 64
(1410–15.0)</td>
</tr>
<tr><td rowspan="1" colspan="1">Body width</td>
<td rowspan="1" colspan="1">38
(30–47)</td>
<td rowspan="1" colspan="1">34.8 ± 9.6
(28.0–41.8)</td>
<td rowspan="1" colspan="1">36.9 ± 4.3
(26.6–48.4)</td>
<td rowspan="1" colspan="1">40.5 ± 0.7
(40.0–41.0)</td>
</tr>
<tr><td rowspan="1" colspan="1">Body width at stylet knobs</td>
<td rowspan="1" colspan="1">_
</td>
<td rowspan="1" colspan="1">15.7 ± 0.4
(15.4–16.0)</td>
<td rowspan="1" colspan="1">16.7 ± 1.3
(14.5–19.4)</td>
<td rowspan="1" colspan="1">17.6 ± 1.1
(16.8–18.4)</td>
</tr>
<tr><td rowspan="1" colspan="1">Body width at S.E pore</td>
<td rowspan="1" colspan="1">_
</td>
<td rowspan="1" colspan="1">24.5 ± 6.1
(20.2–28.8)</td>
<td rowspan="1" colspan="1">27.6 ± 2.6
(23.0–31.5</td>
<td rowspan="1" colspan="1">28.4 ± 0.6
(28.0–28.8)</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet length</td>
<td rowspan="1" colspan="1">20
18–22</td>
<td rowspan="1" colspan="1">19.9 ± 1.8
(18.6–21.1)</td>
<td rowspan="1" colspan="1">19.4 ± 1.2
(17.5–22.9)</td>
<td rowspan="1" colspan="1">19.9 ± 0.9
(19.2–20.5)</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet knob height</td>
<td rowspan="1" colspan="1">_
</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">2.4 ± 0.1
(2.0–3.0)</td>
<td rowspan="1" colspan="1">_</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet knob width</td>
<td rowspan="1" colspan="1">_
</td>
<td rowspan="1" colspan="1">_
</td>
<td rowspan="1" colspan="1">3.9 ± 0.4
(3.0–4.8)</td>
<td rowspan="1" colspan="1">_</td>
</tr>
<tr><td rowspan="1" colspan="1">DGO</td>
<td rowspan="1" colspan="1">8
(6–13)</td>
<td rowspan="1" colspan="1">9
</td>
<td rowspan="1" colspan="1">6.3 ± 0.8
(4.8–8.5)</td>
<td rowspan="1" colspan="1">7.4 ± 0.5
(7.0–7.7)</td>
</tr>
<tr><td rowspan="1" colspan="1">S–E pore<xref ref-type="fn" rid="T5TN1">*</xref>
</td>
<td rowspan="1" colspan="1">_
</td>
<td rowspan="1" colspan="1">135.2
</td>
<td rowspan="1" colspan="1">147 ± 18.8
(97–187)</td>
<td rowspan="1" colspan="1">139 ± 7.1
134–144</td>
</tr>
<tr><td rowspan="1" colspan="1">Metacorpus<xref ref-type="fn" rid="T5TN2">**</xref>
</td>
<td rowspan="1" colspan="1">_
</td>
<td rowspan="1" colspan="1">98 ± 21.5
(83–114)</td>
<td rowspan="1" colspan="1">99 ± 9.8
(76–119)</td>
<td rowspan="1" colspan="1">83 ± 12.7
(134–144)</td>
</tr>
<tr><td rowspan="1" colspan="1">Spicule</td>
<td rowspan="1" colspan="1">32
(28–35)</td>
<td rowspan="1" colspan="1">28.1 ± 10.3
(20.8–35.3)</td>
<td rowspan="1" colspan="1">33.8 ± 1.9
(30.0–37.5)</td>
<td rowspan="1" colspan="1">29.8</td>
</tr>
<tr><td rowspan="1" colspan="1">Gubernaculum</td>
<td rowspan="1" colspan="1">8.5
(7–10)</td>
<td rowspan="1" colspan="1">10.1
</td>
<td rowspan="1" colspan="1">9.0 ± 0.9
(7.3–9.8)</td>
<td rowspan="1" colspan="1">8.4 ± 0.4
(8.1–8.7)</td>
</tr>
<tr><td rowspan="1" colspan="1">Testis length</td>
<td rowspan="1" colspan="1">788
(540–970)</td>
<td rowspan="1" colspan="1">803 ± 125.9
(714–892)</td>
<td rowspan="1" colspan="1">716 ± 167
(324–977)</td>
<td rowspan="1" colspan="1">752 ± 79
(696–808)</td>
</tr>
<tr><td rowspan="1" colspan="1">T</td>
<td rowspan="1" colspan="1">55
(34–65)</td>
<td rowspan="1" colspan="1">55.3 ± 8.6
(49.2–61.4)</td>
<td rowspan="1" colspan="1">48.7 ± 9.7
(27.9–71)</td>
<td rowspan="1" colspan="1">52 ± 3.2
(49–54)</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="T5TN1"><p>* Distance from anterior end to S-E pore.</p>
</fn>
<fn id="T5TN2"><p>** Distance from anterior end to valve plate of median bulb.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T6" orientation="portrait" position="float"><label>Table 6.</label>
<caption><p>Morphometrics of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
second-stage juveniles in comparison with the original descriptions. All measurements are in µm and in the form: mean ± sd. (range).</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Species</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
</th>
<th rowspan="1" colspan="2"><italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
</th>
</tr>
<tr><th rowspan="1" colspan="1">Character</th>
<th rowspan="1" colspan="1">Described</th>
<th rowspan="1" colspan="1">Observed</th>
<th rowspan="1" colspan="1">Described</th>
<th rowspan="1" colspan="1">Observed</th>
</tr>
<tr><td rowspan="1" colspan="1">N</td>
<td rowspan="1" colspan="1">25</td>
<td rowspan="1" colspan="1">5</td>
<td rowspan="1" colspan="1">30</td>
<td rowspan="1" colspan="1">3</td>
</tr>
<tr><td rowspan="1" colspan="1">Body length</td>
<td rowspan="1" colspan="1">418
(390–450)</td>
<td rowspan="1" colspan="1">420 ± 21.7
(390–446)</td>
<td rowspan="1" colspan="1">413 ± 20.6
(373–460)</td>
<td rowspan="1" colspan="1">384 ± 9.5
(374–394)</td>
</tr>
<tr><td rowspan="1" colspan="1">Body width</td>
<td rowspan="1" colspan="1">14.5
(14–16)</td>
<td rowspan="1" colspan="1">14.0 ± 1.1
(12.2–15.2)</td>
<td rowspan="1" colspan="1">14.2 ± 1.8
(12.1–18.2)</td>
<td rowspan="1" colspan="1">12.7 ± 3.8
(8.6–16.9)</td>
</tr>
<tr><td rowspan="1" colspan="1">Body diameter at anus</td>
<td rowspan="1" colspan="1">8.5
(7–9)</td>
<td rowspan="1" colspan="1">9.4 ± 1.8
(8.3–11.5)</td>
<td rowspan="1" colspan="1">8.4 ± 1.0
(7.3–10.9)</td>
<td rowspan="1" colspan="1">6.5 ± 0.7
(6.0–7.0)</td>
</tr>
<tr><td rowspan="1" colspan="1">Stylet length</td>
<td rowspan="1" colspan="1">14
(12–15)</td>
<td rowspan="1" colspan="1">12.1 ± 1.5
(10.9–13.8)</td>
<td rowspan="1" colspan="1">10.0 ± 0.8
(8.5–11.1)</td>
<td rowspan="1" colspan="1">11.1 ± 0.6
10.6–11.5</td>
</tr>
<tr><td rowspan="1" colspan="1">Tail length</td>
<td rowspan="1" colspan="1">31
(30–34)</td>
<td rowspan="1" colspan="1">30.2 ± 4.3
(24.3–33.9)</td>
<td rowspan="1" colspan="1">31.3 ± 3.1
(24.2–37.5)</td>
<td rowspan="1" colspan="1">24.2 ± 0.8
(23.4–25.0)</td>
</tr>
<tr><td rowspan="1" colspan="1">Tail terminus length</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">7.0 ± 2.1
(5.1–9.8)</td>
<td rowspan="1" colspan="1">8.2 ± 1.8
(4.8–12.7)</td>
<td rowspan="1" colspan="1">5.7 ± 1.1
(4.5–6.7)</td>
</tr>
<tr><td rowspan="1" colspan="1">Anus–primordium</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">139 ± 11.4
(125–152)</td>
<td rowspan="1" colspan="1">_</td>
<td rowspan="1" colspan="1">126 ± 21.8
(111–151)</td>
</tr>
<tr><td rowspan="1" colspan="1">a</td>
<td rowspan="1" colspan="1">28.5
(27–31)</td>
<td rowspan="1" colspan="1">30.2 ± 3.2
(27.1–34.8)</td>
<td rowspan="1" colspan="1">29.5 ± 3.4
(22.3–35.5)</td>
<td rowspan="1" colspan="1">32.5 ± 10.9
(23.4–44.7)</td>
</tr>
<tr><td rowspan="1" colspan="1">c</td>
<td rowspan="1" colspan="1">13.3
(12–15)</td>
<td rowspan="1" colspan="1">14.4 ± 2.3
(12.5–17.4)</td>
<td rowspan="1" colspan="1">13.3 ± 1.2
(11.5–16.6)</td>
<td rowspan="1" colspan="1">16.3 ± 0.7
(15.8–16.8)</td>
</tr>
<tr><td rowspan="1" colspan="1">c’</td>
<td rowspan="1" colspan="1">3.7
(3–5)</td>
<td rowspan="1" colspan="1">3.3 ± 0.7
(2.5–3.9)</td>
<td rowspan="1" colspan="1">3.7 ± 0.5
(2.5–4.7)</td>
<td rowspan="1" colspan="1">3.7 ± 0.5
(3.3–4.1)</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec sec-type="Males"><title>Males</title>
<p>Three male paratypes of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and two of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
were measured. Some of the studied characters were only visible enough for measurement on single specimens, and therefore for such characters absolute values were taken rather than their averages. The stylet knobs widths and heights were examples of characters for which measurements were not taken on either species (<xref ref-type="table" rid="T5">Table 5</xref>
) due to the fact that they appeared slightly degenerated on all slides, and so may give false measurements. Nevertheless there were still some outstanding similarities in the stylet length, spicule length and DGO between the observed and the described values.</p>
</sec>
<sec sec-type="Second-stage juveniles"><title>Second-stage juveniles</title>
<p>Similar to the observations made in the females and the males, the second-stage juvenile morphometrics was very comparable in many features between the two species studied. There was, however an unaccountable difference between stylet length as described for <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
(14µm (12–15µm)) and that which was measured (12.1 ± 1.5µm (10.9–13.8µm)). Values of body width at anus level between the two descriptions were very similar. Somemeasurements taken from <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
, likewise were quite similar to those in the original descriptions, particularly, the Demanian ratios a and c’, while others such as stylet lengths showed slight differences (<xref ref-type="table" rid="T6">Table 6</xref>
).</p>
</sec>
<sec sec-type="Host test"><title>Host test</title>
<p>The ability of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
to reproduce on various plant species was examined under greenhouse conditions. Host statuses of the various plants used in the greenhouse test are presented in <xref ref-type="table" rid="T7">Table 7</xref>
. <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
population from “Mierenbos” used as inoculum was able to induce galls and reproduce on both <italic><named-content content-type="taxon-name">Ulmus glabra</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Ulmus hollandica</named-content>
</italic>
‘belgica’. The apple ‘M9’ also had galls which contained egg-laying females. Although galls were induced by <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
on <italic><named-content content-type="taxon-name">Brassica oleracea</named-content>
</italic>
var. <italic>gemmifera</italic>
, most of these galls contained small non-gravid females whose development seemed to have ceased at some point. Therefore, it is herein not considered as a host. There were no galls on <italic><named-content content-type="taxon-name">Rosa hybrida</named-content>
</italic>
and the other cabbage species, <italic><named-content content-type="taxon-name">Brassica pekinensis</named-content>
</italic>
.</p>
<table-wrap id="T7" orientation="portrait" position="float"><label>Table 7.</label>
<caption><p>Plants species identified as host of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
from the green house experiments and field survey at “Mierenbos”.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Family</th>
<th rowspan="1" colspan="1">Plant species</th>
</tr>
<tr><td rowspan="1" colspan="2"><bold>Greenhouse test</bold>
</td>
</tr>
<tr><td rowspan="2" colspan="1"><named-content content-type="taxon-name">Ulmaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Ulmus glabra</named-content>
</italic>
Huds.</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Ulmus hollandica</named-content>
</italic>
‘belgica’</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Rosaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Malus pumila</named-content>
</italic>
‘M9’</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Solanaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Solanum lycopersicum</named-content>
</italic>
L.</td>
</tr>
<tr><td rowspan="1" colspan="2"><bold>Field hosts</bold>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Sapindaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Acer pseudoplatanus</named-content>
</italic>
L.</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Balsaminaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Impatiens parviflora</named-content>
</italic>
DC.</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Asteraceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Taraxacum officinale</named-content>
</italic>
F.H. Wigg.</td>
</tr>
<tr><td rowspan="2" colspan="1"><named-content content-type="taxon-name">Dyopteridaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Dryopteris filix-mas</named-content>
</italic>
(L.) Schott</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Dryopteris carthusiana</named-content>
</italic>
(Vill.) H.P. Fuchs</td>
</tr>
<tr><td rowspan="2" colspan="1"><named-content content-type="taxon-name">Fagaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Fagus sylvatica</named-content>
</italic>
L.</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Quercus robur</named-content>
</italic>
L.</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Geraniaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Geranium robertianum</named-content>
</italic>
L.</td>
</tr>
<tr><td rowspan="3" colspan="1"><named-content content-type="taxon-name">Rosaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Geum coccineum</named-content>
</italic>
Lindl.</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Rubus idaeus</named-content>
</italic>
L.</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Sorbus aucuparia</named-content>
</italic>
L.</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Taxaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Taxus baccata</named-content>
</italic>
L.</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Ulmaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Ulmus davidiana</named-content>
</italic>
var. <italic>japonica</italic>
Rehder</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Urticaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Urtica dioica</named-content>
</italic>
L.</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T8" orientation="portrait" position="float"><label>Table 8.</label>
<caption><p>A compilation of all known host plants of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
to date.</p>
</caption>
<table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Family</th>
<th rowspan="1" colspan="1">Plant species</th>
<th rowspan="1" colspan="1">Reference</th>
</tr>
<tr><td rowspan="10" colspan="1"><named-content content-type="taxon-name">Rosaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Malus pumila</named-content>
</italic>
Mill.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Malus prunifolia</named-content>
</italic>
Borkh.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Malus sieboldii</named-content>
</italic>
Rehd.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Malus pumila</named-content>
</italic>
“M9”</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Prunus yedoensis</named-content>
</italic>
Matsum</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Rosa hybrida</named-content>
</italic>
Hort.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Geum coccineum</named-content>
</italic>
Lindl.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Vitis vinifera</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Rubus idaeus</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Sorbus aucuparia</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="5" colspan="1"><named-content content-type="taxon-name">Moraceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Morus bombycis</named-content>
</italic>
Koidz.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Ficus carica</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Maclura tricuspidata</named-content>
</italic>
(Carriere) Bureau</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Broussonetia papyrifera</named-content>
</italic>
(L.) Vent</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Broussonetia kazinoki</named-content>
</italic>
Seibold.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="3" colspan="1"><named-content content-type="taxon-name">Fagaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Castanea crenata</named-content>
</italic>
Seib. Et Zucc</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Fagus sylvatica</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Quercus robur</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="4" colspan="1"><named-content content-type="taxon-name">Ulmaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Ulmus davidiana</named-content>
</italic>
var. <italic>japonica</italic>
</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Ulmus chenmoui</named-content>
</italic>
W.C. Cheng</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B27">Palmisano and Ambrogioni 2000</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Ulmus glabra</named-content>
</italic>
Hud.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B27">Palmisano and Ambrogioni 2000</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Ulmus</named-content>
</italic>
× <italic>hollandica</italic>
“belgica”</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="3" colspan="1"><named-content content-type="taxon-name">Sapindaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Acer palmatum</named-content>
</italic>
Thunb.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Acer pseudoplatanus</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Trifolium repens</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Taxaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Taxus baccata</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Fabaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Impatiens parviflora</named-content>
</italic>
DC.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="3" colspan="1"><named-content content-type="taxon-name">Solanaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Solanum lycopersicum</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Solanum melongena</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Capsicum annuum</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="3" colspan="1"><named-content content-type="taxon-name">Cucurbitaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Cucumis sativus</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Cucurbita</named-content>
</italic>
spp.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Citrillus vulgaris</named-content>
</italic>
Schrad. Ex Eckl. & Zeyh.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="3" colspan="1"><named-content content-type="taxon-name">Cruciferae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Brassica pekinensis</named-content>
</italic>
Rupy.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Brassica oleracea</named-content>
</italic>
var. <italic>capitata</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Brassica napus</named-content>
</italic>
var. <italic>oleifera</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="2" colspan="1"><named-content content-type="taxon-name">Compositae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Arcutium lappa</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Taraxacum officinale</named-content>
</italic>
F.H. Wigg.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Umbelliferae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Daucus carota</named-content>
</italic>
var. <italic>sativa</italic>
L.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Leguminaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Glycine max</named-content>
</italic>
(L.) Merr.</td>
<td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B36">Toida 1979</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Urticaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Urtica dioica</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="2" colspan="1"><named-content content-type="taxon-name">Dryopteridaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Dryopteris filix-mas</named-content>
</italic>
(L.) Schott</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Dryopteris carthusiana</named-content>
</italic>
(Vill.) H.P. Fuchs</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
<tr><td rowspan="1" colspan="1"><named-content content-type="taxon-name">Geraniaceae</named-content>
</td>
<td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Geranium robertianum</named-content>
</italic>
L.</td>
<td rowspan="1" colspan="1">Current work</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>Additionally, samples collected during 2011 and 2012 revealed that <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
is able to parasitize one or more species of <italic><named-content content-type="taxon-name">Acer</named-content>
</italic>
(<named-content content-type="taxon-name">Aceraceae</named-content>
), <italic><named-content content-type="taxon-name">Impatiens</named-content>
</italic>
(<named-content content-type="taxon-name">Balsaminaceae</named-content>
), <italic><named-content content-type="taxon-name">Taraxacum</named-content>
</italic>
(<named-content content-type="taxon-name">Compositae</named-content>
), <italic><named-content content-type="taxon-name">Dryopteris</named-content>
</italic>
(<named-content content-type="taxon-name">Dryopteridaceae</named-content>
), <italic><named-content content-type="taxon-name">Fagus</named-content>
</italic>
(<named-content content-type="taxon-name">Fagaceae</named-content>
), <italic><named-content content-type="taxon-name">Quercus</named-content>
</italic>
(<named-content content-type="taxon-name">Fagaceae</named-content>
), <italic><named-content content-type="taxon-name">Geranium</named-content>
</italic>
(<named-content content-type="taxon-name">Geraniaceae</named-content>
), <italic><named-content content-type="taxon-name">Geum</named-content>
</italic>
(<named-content content-type="taxon-name">Rosaceae</named-content>
), <italic><named-content content-type="taxon-name">Rubus</named-content>
</italic>
(<named-content content-type="taxon-name">Rosaceae</named-content>
), <italic><named-content content-type="taxon-name">Sorbus</named-content>
</italic>
(<named-content content-type="taxon-name">Rosaceae</named-content>
), <italic><named-content content-type="taxon-name">Taxus</named-content>
</italic>
(<named-content content-type="taxon-name">Taxaceae</named-content>
), <italic><named-content content-type="taxon-name">Urtica</named-content>
</italic>
(<named-content content-type="taxon-name">Urticaceae</named-content>
), as shown in <xref ref-type="table" rid="T7">Table 7</xref>
.</p>
</sec>
<sec sec-type="Isozyme analysis"><title>Isozyme analysis</title>
<p>Samples taken from the trial field “Mierenbos” all gave the same type of esterase isozyme pattern of weak single bands, corresponding to the VS1 type (<xref ref-type="bibr" rid="B8">Esbenshade and Triantaphyllou 1985</xref>
). When analysed for MDH, some individuals gave single-banded patterns of the H1 type (<xref ref-type="bibr" rid="B8">Esbenshade and Triantaphyllou 1985</xref>
), while others revealed a three-banded pattern, herein designated H3. Usually, the H1 type had two additional weaker bands at the same level as the upper two H3 bands. There was also an additional observation in the types of single bands some of the specimens produced (<xref ref-type="fig" rid="F3">Fig. 3</xref>
). These single bands were positioned at the same level as the upper H3 band, which herein are given the name H1a.</p>
<fig id="F1" orientation="portrait" position="float"><label>Figure 1.</label>
<caption><p>LM photograph of perineal patterns and anterior parts of female <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
(<bold>A, C, E, G</bold>
) and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
(<bold>B, D, F</bold>
), bar = 10 µm.</p>
</caption>
<graphic xlink:href="ZooKeys-362-001-g001"></graphic>
</fig>
<fig id="F2" orientation="portrait" position="float"><label>Figure 2.</label>
<caption><p>LM photographs of males anterior part and second- stage juvenile tails of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
(<bold>A, C, E</bold>
) and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
(<bold>B, D, F</bold>
), bar = 10 µm.</p>
</caption>
<graphic xlink:href="ZooKeys-362-001-g002"></graphic>
</fig>
<fig id="F3" orientation="portrait" position="float"><label>Figure 3.</label>
<caption><p>Isozyme phenotypes from ten individual females of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
from “Mierenbos”. <bold>A</bold>
Esterase <bold>B</bold>
Malate dehydrogenase. <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
(1–5 and 8–12); <italic><named-content content-type="taxon-name">Meloidogyne javanica</named-content>
</italic>
(6 and 7) as reference marker.</p>
</caption>
<graphic xlink:href="ZooKeys-362-001-g003"></graphic>
</fig>
</sec>
</sec>
<sec sec-type="Phylogenetic relationship between Meloidogyne mali and M. ulmi"><title>Phylogenetic relationship between <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
</title>
<p>The obtained SSU rDNA and LSU rDNA sequence lengths for <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
were 781bp (including gaps) and 698bp (including gaps) respectively. In addition to our four SSU rDNA sequences of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
“Mierenbos” (<ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/nuccore/KF895397">KF895397</ext-link>
, <ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/nuccore/KF895398">KF895398</ext-link>
, <ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/nuccore/KF895399">KF895399</ext-link>
and <ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/nuccore/KF895400">KF895400</ext-link>
), 69 accessions belonging to other species of <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
from GeneBank were included in the local alignment (781 aligned positions, including gaps). For LSU rDNA, we had only one sequence of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
“Mierenbos” (<ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/nuccore/KF895396">KF895396</ext-link>
) due to poor data, resulting in lack of consensus sequence. Therefore the multiple sequence alignment included this sequence and 69 GeneBank accessions from other species of <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
. <italic><named-content content-type="taxon-name">Pratylenchus vulnus</named-content>
</italic>
Allen & Jensen, 1951 was selected as outgroup for constructing gene trees using Bayesian inference from both SSU rDNA and LSU rDNA sequences. SSU rDNA-based phylogenetic analysis put all sequences of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
obtained together with those of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
from GeneBank in one strongly supported polytomous branch. Despite the relatively short sequence length of SSU rDNA, the tree was able to resolve relationship between certain species in a way comparable with that of <xref ref-type="bibr" rid="B12">Holterman et al. (2009)</xref>
. LSU rDNA-based Bayesian analysis revealed higher resolution within the group containing our sequence and sequences of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
from GeneBank. Our sequence of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
was positioned in a branch that contained three other sequences of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
, forming a sister group to another branch composed also of two sequences of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
and one of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
. As would be expected, there was higher resolution in the overall topology of LSU rDNA-based tree than that of SSU rDNA. The sequence identity between the closest sequences of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
was 98.8 % with one gap. Interestingly, the same percentage was obtained between two sequences of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
from GeneBank (<ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/nuccore/JX978226">JX978226</ext-link>
and <ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/nuccore/JX978227">JX978227</ext-link>
).</p>
</sec>
</sec>
<sec sec-type="Discussion"><title>Discussion</title>
<p>Type specimens representing holotypes and paratypes of the two <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
species were analysed in order to demonstrate the morphological similarities that existed between them. Most of the slides we received were in good conditions except for some few individuals that showed some signs of deterioration, either due to long period of storage or poor conditions at the time they were prepared. Nevertheless, the general states of the important characters were still maintained.</p>
<p>On the morphology, an important character like the occurrence of double lateral lines mentioned in the description and by <xref ref-type="bibr" rid="B18">Jepson (1987)</xref>
was not observed in all specimens. In fact this area was marked by breaks in the striae on some of the specimens studied. Morphometric values of some characters for the two species fell within the range of values reported in the original description of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
(<xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
), for example position of DGO in females, as well as in males, vulva-anus distance, level of phasmids to vulva, vulva slit length, male stylet length, testis length, spicule length gubernaculum length, J2 body length, and J2 c’ values (<xref ref-type="table" rid="T4">Tables 4</xref>
<xref ref-type="table" rid="T5">–</xref>
<xref ref-type="table" rid="T6">6</xref>
). For some characters, however, morphometric values recorded agreed more with those reported for <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
. For example, the stylet length measured in females and J2s was significantly lower than the values given for <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
, but comparable with that of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
. A possible explanation might be the fact that stylet in juveniles and sometimes in females appear less visible, causing the anterior end to be mistaken for the tip of the conus. This leads to misleadingly higher values for the stylet length.</p>
<p>Already in the original description of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
, only a few differences could be found to separate it from <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
. And in some cases, the differences emanated from some apparent mistakes in the original description of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
. An example is the use of male tail length of the two species to draw differences. Tail length in males of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
was given as ranging between 28 and 44 µm, making it extremely longer than that of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
10.9 µm. However, it is important to mention that tail length values as long as the range of between 28 and 44 µm never exists among males of species of the genus <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
.</p>
<p>The DGO position in males with reference to the stylet knobs according to <xref ref-type="bibr" rid="B18">Jepson (1987)</xref>
bears some broad interspecific variation, making it very useful for species discrimination. <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
is by far the species with the most farther DGO position (6–13 µm) within the genus <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
. The observation of similar values for both species studied here therefore separates the two from all other species that have relatively shorter DGO position. Additional <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
specimens studied also gave DGO position values averaging 8 µm (data not shown).</p>
<p><xref ref-type="bibr" rid="B6">Eisenback and Hirschmann (1981)</xref>
highlighted the significance of SEM studies of male head shapes in <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
taxonomy, outlining the role SEM has played in raising the value of males for use in comparison of species. Head and stylet shape morphologies of males and juveniles are the most useful supplemental taxonomic characters that SEM studies have given new insights into (<xref ref-type="bibr" rid="B7">Eisenback and Hirschmann 1979</xref>
; <xref ref-type="bibr" rid="B6">1981</xref>
). It is not surprising that a number of variability in these characters were outlined to separate <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
from <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
by <xref ref-type="bibr" rid="B27">Palmisano and Ambriogioni (2000)</xref>
. It was mentioned (p. 288) that “under SEM lateral lips absent or vestigial (in <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
lateral lips apparent)”. Interestingly, this contradicted the comparison made by <xref ref-type="bibr" rid="B35">Toida and Yaegashi (1984)</xref>
when they attempted to point out the differentiating characters between <italic><named-content content-type="taxon-name">Meloidogyne suginamiensis</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
. In their comparison, the <italic>enface</italic>
view of the lip region of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
was mentioned as having no or obscure lateral lips to separate it from <italic><named-content content-type="taxon-name">Meloidogyne suginamiensis</named-content>
</italic>
in which lateral lips were described as clear. One would not expect such contradicting accounts especially with the possibility that both works referred to the same publication (<xref ref-type="bibr" rid="B25">Okamoto et al. 1983</xref>
). Referring to a separate work (<xref ref-type="bibr" rid="B38">Yeagashi and Okamoto 1981</xref>
), the account given by <xref ref-type="bibr" rid="B35">Toida and Yaegashi (1984)</xref>
seems to us more probable, the first reason being that they published the same work which is being referred to. Therefore they understand the details of their results more. And the second reason is that our observations of SEM images of (<xref ref-type="bibr" rid="B38">Yeagashi and Okamoto 1981</xref>
) agree more with the account that lateral lips were vestigial and not apparent in <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
.</p>
<p>Both apple and elm trees supported <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
reproduction. This does not only provide an additional support for the synonymization of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
with <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
, but represents the first and only test involving the former on an apple plant. In principle, however, the first actual report was the description of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
on apple in Japan (<xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
). Contrary to the finding in the original description, the status of white clover as host to <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
could not be confirmed with <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
. Again, although representatives of the family <named-content content-type="taxon-name">Rosaceae</named-content>
form the larger part of the plants <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
parasitizes (<xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
; <xref ref-type="bibr" rid="B36">Toida 1979</xref>
), rose (<italic><named-content content-type="taxon-name">Rosa hybrida</named-content>
</italic>
) could not support the reproduction of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
. This contradicts earlier finding, <xref ref-type="bibr" rid="B17">Itoh et al. 1969</xref>
, who identified rose as host. <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
was also able to induce galls on <italic><named-content content-type="taxon-name">Prunus yodoensis</named-content>
</italic>
grown in the field, confirming the earlier report of the latter’s status as a host for <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
by <xref ref-type="bibr" rid="B36">Toida (1979)</xref>
. Results of the sampling have also revealed new natural hosts for <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
like <italic><named-content content-type="taxon-name">Acer pseudoplatanus</named-content>
</italic>
L., <italic><named-content content-type="taxon-name">Fagus sylvatica</named-content>
</italic>
L., <italic><named-content content-type="taxon-name">Quercus robur</named-content>
</italic>
L., <italic><named-content content-type="taxon-name">Rubus idaeus</named-content>
</italic>
L., <italic><named-content content-type="taxon-name">Sorbus aucuparia</named-content>
</italic>
L., <italic><named-content content-type="taxon-name">Taxus baccata</named-content>
</italic>
L., <italic><named-content content-type="taxon-name">Dryopteris filix-mas</named-content>
</italic>
(L.) Schott, <italic><named-content content-type="taxon-name">Dryopteris carthusiana</named-content>
</italic>
(Vill.) Fuchs, <italic><named-content content-type="taxon-name">Geranium robertianum</named-content>
</italic>
L., <italic><named-content content-type="taxon-name">Urtica dioica</named-content>
</italic>
Rehder., <italic><named-content content-type="taxon-name">Impatiens parviflora</named-content>
</italic>
DC., <italic><named-content content-type="taxon-name">Taraxacum officinale</named-content>
</italic>
F.H. Wigg., and <italic><named-content content-type="taxon-name">Geum coccineum</named-content>
</italic>
Lindl. The most relevant evidence deduced from this host plant test is the ability of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
to reproduce on apple.</p>
<p>It is interesting to mention that the observed variability of the MDH isozyme phenotypes among the different specimens was similar to the findings of <xref ref-type="bibr" rid="B29">Sakai and Mizukubo (2009)</xref>
when they studied two populations of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
from Hokkaida on apple and Saitama on cherry in Japan. The populations from Hokkaida gave phenotypes with single MDH bands whereas those from Saitama on cherry produced variable patterns with single and triple bands. PCR-RFLP of D2/D3 expansion segment of 28S rDNA and mtDNA intergenic region with <italic>Alu</italic>
I was able to confirm that population from Saitama were all identical, despite their expression of variable MDH isozyme phenotypes. Similar observations of intraspecific phenotype variations were made by <xref ref-type="bibr" rid="B4">Dalmasso and Bergé (1978)</xref>
among a certain <italic><named-content content-type="taxon-name">Meloidogyne arenaria</named-content>
</italic>
population where there were three MDH bands instead of two. Such type of variable isozyme patterns were also observed within one population of the sexually reproducing species <italic><named-content content-type="taxon-name">Meloidogyne microtyla</named-content>
</italic>
Mulvey, Townshend & Potter, 1975 (Karssen unpublished). This indicates that <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
could also be a sexually reproducing species or a meiotic parthenogenetic one, a claim which is further supported by the frequency at which males are encountered in galled root samples—at least one male per female in a gall. Meanwhile, the esterase phenotypes were rather stable across all studied specimens and were characterised by weak indistinct single bands.</p>
<p>Trimming the SSU and LSU datasets to high quality sequence data may have caused a loss in phylogenetic signal. For SSU rDNA, over half of the target sequence length was trimmed out because of the poor quality of the dataset obtained. Although not ideal for reconstruction of phylogeny, it was still sufficient to resolve the taxa on a species level. Moreover, it has to be emphasized that the purpose here is not to reconstruct any formal phylogeny of <italic><named-content content-type="taxon-name">Meloidogyne</named-content>
</italic>
, a subject which is well covered already in previous studies (<xref ref-type="bibr" rid="B34">Tandingan De Ley et al. 2002</xref>
; <xref ref-type="bibr" rid="B12">Holterman et al. 2009</xref>
), but only to demonstrate that <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
are highly similar at the molecular level and belong to the same clade. A recent phylogenetic analyses involving these two species has already pointed to the fact that the two can not be separated based on their SSU rDNA sequences (<xref ref-type="bibr" rid="B28">Rybarczyk-Mydłowska et al. in press</xref>
). The SSU rDNA sequence once again gave resolution till the species level, confirming previous proposition that SSU rDNA sequence signatures can be defined at species level for a wide range of parasitic and non-parasitic nematodes (<xref ref-type="bibr" rid="B14">Holterman et al. 2006</xref>
). As was expected, LSU rDNA-based analysis gave even higher resolution and more clearly defined the relationship between <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
. On the SSU rDNA based tree, it is unquestionable that our sequence of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
with all the other sequences of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
are the same (<xref ref-type="fig" rid="F4">Fig. 4</xref>
). The clustering of our sequence of LSU rDNA for <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
with those for <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
may be an indication that the branching could only be due to intraspecific sequence variation.</p>
<fig id="F4" orientation="portrait" position="float"><label>Figure 4.</label>
<caption><p>Bayesian tree inferred from part of 18S rRNA using TYMef + I model. Sequences were aligned with MAFFT alignment. Numbers near the nodes indicate posterior probabilities. NCBI accession numbers are listed with the species names.</p>
</caption>
<graphic xlink:href="ZooKeys-362-001-g004"></graphic>
</fig>
</sec>
<sec sec-type="Conclusion"><title>Conclusion</title>
<p>In conclusion, the evidence from morphological and morphometrical studies of holo– and paratype materials of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
and <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
as well as host plant studies, isozyme analysis and DNA analysis all confirm the status of <italic><named-content content-type="taxon-name">Meloidogyne ulmi</named-content>
</italic>
as a junior synonym of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
.</p>
<sec sec-type="Notes on the life cycle and biology of Meloidogyne mali on Ulmus spp"><title>Notes on the life cycle and biology of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
on <italic><named-content content-type="taxon-name">Ulmus</named-content>
</italic>
spp.</title>
<p>The life cycle of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
is in many respects typical of the genus. <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
requires 18–22 weeks to complete one full generation on apple and does so only once in a year (<xref ref-type="bibr" rid="B16">Inagaki 1978</xref>
). The study also reported that adult males and females first were observed after twelve weeks and continued to increase till the twentieth, when egg masses began to appear. There was also some reports on the distribution of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
in the field, both vertically and horizontally. However, nothing is known so far about its survival on apple or any other plant during frost conditions of winter. Regarding this, a very interesting observation was made during early spring of 2013 at the trial field “Mierenbos”. Egg-laying females were already found in most galls that were examined, a rare phenomenon known to occur only in <italic><named-content content-type="taxon-name">Meloidogyne ardenensis</named-content>
</italic>
(<xref ref-type="bibr" rid="B33">Stephan and Trudgill 1982</xref>
). The only plausible explanation to why egg-laying females can be observed so early in the year is that, like reported for <italic><named-content content-type="taxon-name">Meloidogyne ardenensis</named-content>
</italic>
, the nematodes overwintered in the roots. Additional observations need to be made to find out exactly what stage in the development overwinters in the root.</p>
</sec>
<sec sec-type="Host plants and distribution of M. mali in Europe"><title>Host plants and distribution of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
in Europe</title>
<p><italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
induces a similar type of galls as do <italic><named-content content-type="taxon-name">Meloidogyne arenaria</named-content>
</italic>
on tomatoes, a type of gall commonly referred to as bead-like galls (<xref ref-type="fig" rid="F6">Fig. 6</xref>
). Concerning the current distribution of the nematode in Europe, no study has yet been done to investigate this. However, it would be rational to speculate that <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
may be found in all the ten European countries to which rooted seedlings were sent after the breeding programme. These countries include Belgium, England, France, Ireland, Italy, Spain, Denmark, Germany, Slovakia and Romania (<xref ref-type="bibr" rid="B10">Heybroek 1993</xref>
). Elsewhere in Asia, it has also been found in <italic><named-content content-type="taxon-name">Acer palmatum</named-content>
</italic>
trees from Japan that were intercepted in China (Gu unpublished). Sequence data from these were also included in the analysis.</p>
<fig id="F5" orientation="portrait" position="float"><label>Figure 5.</label>
<caption><p>Bayesian tree inferred from part of D2 D3 of 28S rRNA using TYM + G model. Sequences were aligned with MAFFT alignment. Numbers near the nodes indicate posterior probabilities. NCBI accession numbers are listed with the species names.</p>
</caption>
<graphic xlink:href="ZooKeys-362-001-g005"></graphic>
</fig>
<fig id="F6" orientation="portrait" position="float"><label>Figure 6.</label>
<caption><p>Root gall symptoms of <italic><named-content content-type="taxon-name">Meloidogyne mali</named-content>
</italic>
infection on (<bold>A, B</bold>
) <italic><named-content content-type="taxon-name">Maluspumila</named-content>
</italic>
“M9” (<bold>C, D</bold>
) <italic><named-content content-type="taxon-name">Ulmus davidiana</named-content>
</italic>
var. <italic>japonica</italic>
and (<bold>E, F</bold>
) <italic><named-content content-type="taxon-name">Solanum lycopersicum</named-content>
</italic>
.</p>
</caption>
<graphic xlink:href="ZooKeys-362-001-g006"></graphic>
</fig>
</sec>
</sec>
</body>
<back><ack><title>Acknowledgement</title>
<p>The authors are grateful to the following people for their help with providing type slides for study: Dr. Z.A. Handoo, Shigeyuki Sekimoto and Dr. Hiromichi Sakai. They also wish to thank Takayuki Mizukubo for providing root sample from the type apple tree, and to Tiziana Irdani and Beatrice Carletti for Elm root materials from the type tree. Evelyn van Heese for helping with slides preparation and electrophoresis, Anne-Sophie van Bruggen and Johan van Woggelum for help with field sampling and host plant tests. They are also grateful to Dr. M. Heybroek, formally of the Dorschkamp Research Institute for Forestry & Landscape Planning, for providing essential information about the <italic><named-content content-type="taxon-name">Ulmus</named-content>
</italic>
breeding programme. Very special thanks also to Eveline Metz-Verschure and Dr. Marcel Westenberg for their help with molecular analysis.</p>
</ack>
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