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Diagnostic and Phylogenetic Significance of Lateral Field Incisures

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Diagnostic and Phylogenetic Significance of Lateral Field Incisures Russian Journal of Nematology, 2018, 26 (1), 1 – 27 Diagnostic and phylogenetic significance of lateral field incisures, phasmids and other morphological characters of second-stage juveniles and males of Heteroderidae (Nematoda, Tylenchida), with notes on hosts and phylogeography Dieter Sturhan Arnethstr. 13D, 48159, Münster, Germany e-mail: [email protected] Accepted for publication 20 October 2017 Summary. Data of the current knowledge about the number of incisures in the lateral fields of second- stage juveniles and males, shape of phasmids and tip of spicules of the 158 recognised species in the 19 genera in the family Heteroderidae are reviewed and compiled. All characters are of diagnostic value in distinguishing genera and intrageneric species groups, but more detailed studies are needed. Differences in the number of incisures or in the size of phasmids among species of Cryphodera, Meloidodera and Atalodera are considered as an indication of paraphyly. In agreement with results of molecular analyses, three additional species groups are proposed for Heterodera species with Poales as hosts (Cardiolata, Koreana and Orientalis groups), and in Globodera designation of Rostochiensis, Millefolii and Zelandica groups. Information on host ranges and host reaction to parasitism has great relevance for the study of phylogenetic relationships and estimation of the age of taxa. Evaluation of biogeographical data restricted to ‘natural’ occurrence as well as consideration of geological and palaeoclimatic events are essential for the recovery of putative ancestral areas of origin of taxa. Heteroderid populations tentatively identified as Meloidodera, Hylonema and Sarisodera are briefly characterised from Africa (the first two genera) and Vietnam (the third genus). Other populations of unidentified species or representatives of still unknown genera are reported from Borneo, the Canary Islands, the Dominican Republic, New Zealand, Thailand, Tunisia and Vietnam. Key words: Ataloderinae, Cryphodera, distribution, Globodera, Heterodera, Heteroderinae, Hylonema, Meloidodera, Meloidoderinae, morphology, plant hosts, Punctoderinae, Sarisodera, Verutinae, Verutus. The family Heteroderidae is among the best to non-acceptance of genera such as Afenestrata, studied taxa of tylenchid nematodes, with currently Bidera, Ephippiodera and others and the transfer of 158 recognised species in a total of 19 genera. Many individual species to other genera or to different species are of economic significance as plant species groups within a genus, particularly in the parasites and require precise identification for their genus Heterodera. Species with descriptions management in agriculture. The genera and species exclusively based on cysts characteristics or lacking placed in the cyst-forming subfamily Heteroderinae data on second-stage juveniles (J2) and males were and in the non-cyst forming subfamilies commonly considered as species inquirendae. Meloidoderinae, Ataloderinae and Verutinae are Morphological characters of males and J2 were mainly distinguished and characterised by mostly considered of secondary significance, but J2 morphological characters of cysts or females. More features are presently more often used for distinction recent integrated morphological and molecular and identification of species. Differences in host studies have shown that the significance of cyst preferences were other important characters used for characters has often been overestimated, which led changes in classification. For instance, the number © Russian Society of Nematologists, 2018; doi:10.24411/0869-6918-2018-10001 1 D. Sturhan of incisures in the lateral fields of J2 and the size of essential for identification of genera, species groups phasmids were considered decisive for excluding or individual species. These data are very useful for Heterodera bifenestra from the H. avenae species the elucidation and validation of potential group, phasmid size and host preference for phylogenetic relationships, which are so far mainly excluding H. turcomanica from the same group based on molecular studies. Moreover, another goal (Wouts & Sturhan, 1995), and the number of lateral of compiling, discussing and reviewing the incisures for separating Betulodera betulae from the mentioned morphological characters and other data genus Cactodera (Sturhan, 2002). is to indicate the gaps in current knowledge and to In the present paper, data of our current provide a useful basis for further studies on knowledge on the number of longitudinal incisures Heteroderidae, which will hopefully stimulate future in the lateral fields of J2 and males, on presence and research. size of phasmids in both developmental stages and on the shape of the tip of spicules are compiled for MATERIALS AND METHODS all genera and intrageneric groups currently placed in the family Heteroderidae. The number of lateral Original descriptions of the valid species in incisures is commonly used in Tylenchida as a Heteroderidae and subsequent publications significant diagnostic character for genera, in supplementing morphological data of individual particular in Telotylenchidae and Merliniidae. In species were critically reviewed and evaluated for Heteroderidae the number of lateral field incisures information on the selected morphological (‘lateral lines’) has already often been used for characters of J2 and males. These characters include distinction of species and occasionally also for the numbers of incisures in the lateral fields, distinction of species groups within the genus presence and size of phasmids and the shape of the Heterodera (see Subbotin et al., 2010a). Data on spicule tip. Some previous data of such characters presence or absence of phasmids in males of proved to be incorrect and had subsequently already Heteroderidae as potential diagnostic character for been corrected in several publications; other distinction of subfamilies have recently been characters were used as described or depicted in the compiled (Sturhan, 2016). original descriptions or in subsequent emending A compilation of known data on a few selected publications. The books published by Subbotin et al. characters of motile life stages of heteroderids and (2010a, b) served in particular as valuable sources of their critical evaluation are presented to support and information for the cyst-forming Heteroderinae. facilitate identification, based on some Published data on lateral fields and the number morphological characters that can generally be seen of incisures in J2 are available for almost all valid using light-microscopical examination of J2, which heteroderid species, but not so in males of many are often the only life stages available for species. Phasmids were often not precisely identification. Thus, data on lip region characters, described or depicted. Mostly, no distinction was which were also used for the distinction of made between phasmidial opening to the outside heteroderid taxa but difficult to discern in routine (indistinct, small, distinct) and the phasmidial duct microscopical studies and only known for a leading through the cuticle to the receptor cavity comparatively low number of taxa, were not (narrow, with small lens-like extension, with large included in this paper. lens-like ampulla). Likewise, the shape of the Another objective of this publication is to spicules and spicule tips have often not been confirm or to question congruence of the few carefully or correctly described or depicted (pointed, selected morphological characters with available bluntly rounded, notched, dentate, bidentate, molecular data and eventually to indicate tridentate). The significance of these characters is relationships or paraphylies that had not been thus sometimes of only limited value and more demonstrated by previous molecular or biochemical detailed studies are required. analyses. The total number of species included into In the present paper, supplemental information molecular studies to date is still relatively low. on certain morphological characters is sometimes However, a high number of species is (still) placed complemented by microscopical studies conducted into selected genera or species groups, such as by the author of type and other reliably identified Heteroderinae, exclusively on the basis of their specimens available in the German Nematode morphological characters, especially those of cysts. Collection (DNST), microscopical slides or other Data on host plants, host reaction to parasitism and material obtained from several nematode collections native geographical distribution are briefly added in or directly studied in other nematological research the present paper, because such information is often centres. Several presumably undescribed heteroderid 2 Diagnostics and phylogeography of Heteroderidae species and representatives of still unknown genera classification for the plant orders and families is were included in the present studies. Second-stage used (Angiosperm Phylogeny Group, 2016). In juveniles were mostly available for such light presenting data on geographical distribution, a microscope studies, males only exceptionally. distinction is made or attempted between supposed The classification used in the present paper is, to ‘native’ or ‘natural’ occurrence and recent dispersal. a large extent, the same as that used by Sturhan Geographical records, which are most likely due to (2016). In this classification, three subfamilies: human activities, were not taken into account. Ataloderinae, Heteroderinae and
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