Fundarn. appl. Nernawl., 1997,20 (3),243-251

The genus Dorylaimoides Thorne & Swanger, 1936 (Nematoda: ). 1. and variability

Reyes PENA SA.NTIAGO* and Manuel PERALTA**

e' Universidad de Jacn. Deparlarnenlo de Biologia , Vegelal)' Ecologia. Paraje "Las Lagu.mllas "s/n, Edificio n" 5, 23071jaen, Spain and ,,;, 1. B. "Auringis ", Avda. de Andaluda s/n, 23005jaen, Spain. Accepted for publication ] 3 May] 996.

Summary -A general revision of differenr aspects of the morphology and taxonomy of the genus DOly/airnoides Thome & Swanger, 1936 is presenred. Variabiliry ofthe morphological features is described and illustrated, and their taxonomic value is briefly discussed. The taxonomical position of the genus, its relationships with other genera and its intrageneric taxonomy are likewise treated. A list of the species is also given.

Reswne - Le genre Dorylaimoides Thome & Swanger, 1936 (Nematoda: Dorylairnida). 1. Taxinornie et variabilite­ Une revision generale des differenrs aspects de la morphologie et de la taxonomie du genre Dorytairnoides Thome & Swanger, 1936 est presentee. La variabilite des caraeteristiques morphologiques est decrite et illustree, et !eur valeur taxonomique est discuree brievemenr. La position taxonomique du genre, ses relations avec d'autres genres et la taxonomie intragenerique sont ega!emenr traitees. Une liste des especes est donnee. Key words: D01ylairnoides, , taxonomy.

Dorylalmoldes Thorne & Swanger, 1936 is one espe­ body diameter at neck base; tapering also, towards the cially interesting dorylaimid genus due to its posterior extremity more or less according to the tail intricate taxonomy and its relatively wide morphological shape. In general, body slender since de Man's " a " variability. At present, it contains more than 60 species, ratio usually varies from 30 to 40 but may exceed the some of which have been described from a small num­ latter value. After fixation, habitus more or less ventrally ber of specimens or have been poorly illustrated; other curved to C- or ]-shaped; almost always the male pre­ species are known from several localities and present sents a more curved posterior body region than the fe­ significant intraspecific variability mainly affecting the male. morphology of the tail and the female genital system but CUllcle: Under optical microscopy it is possible to also other features. distinguish two layers: outer layer thin, constant in In Andalucia Oriental (Southeastern Spain) the genus thickness along the entire body; fine and transverse sur­ appears well distributed and diversified. Very recendy face striations normally present but some interspecific we have examined (Peralta & Pefia Santiago 1995a, b, c) variability exists since inconspicuous as well as clear some eighty populations collected from very different striation can occur; inner layer always thicker than the habitats (forest, brushwood, cultures, meadow, etc.), outer one, widening visibly at the caudal region. and including fourteen different species. In our opinion, this Spanish material constitutes a good representation Laleral chord and pores: A relatively narrow lateral of the genus due to the number of species, bur mosdy chord (in Spanish material varying from one-tenth to because of the morphological variability observed one-seventh of the midbody diameter) always obvious, among these species. The information so obtained has its margins appearing clear and including a granular or led us to undertake a more general revision of the mor­ amorphous content, although in some species the exist­ phology and the taxonomy of the genus, which is pre­ ence of glandular bodies is easily perceptible. Lateral sented below. pores fine and located in a simple row, frequendy very inconspicuous. Notes on general morphology Lip regIon: A wide interspecific varibility can be ob­ served (see Fig. 1 A-C) : the lip region is often some­ General aspecl : Body length varying strongly, ranging what angular and offset from the adjacent body by a from 0.7 to 3.0 mm but very rarely exceeding 2 mm; depression or a more or less deep constriction; however, frequendy around 1 mm. Shape cylindrical, tapering to practically continuous and more rounded in some spe­ the anterior end which becomes about one-third of the cies. Lips almost always amalgamated, with the outer

ISSN 1164-5571/97/03 S 7.00/ © Gawhier-Vi/tars - ORSTOM 243 R. Peiia Sanliago & M. Peralla

relatively robust and its length (ventral side) varies from half to scarcely more than the lip region width (Fig. 1 E­ A G). Aperture always well visible and occupying 25-33 % of the total stylet length. In lateral view, the ventral side of the odontostyle appears practically straight and with the anterior end dorsally bent; dorsal side somewhat sigmoid and in all cases longer than the ventral one. Odontophore presenting a typical ventral curvature and surrounded by a somewhat hyaline tissue; odontophore D always longer than the odontostyle but its precise length F(I difficult to establish since it joins the pharyngeal lining without a clear transition. Digestive system: Pharynx consisting ofa slender ante­ H I rior part and a basal bulb (Fig. 1 H); the former begins with a somewhat thickened portion which becomes more slender at level of the nerve ring, then widens slightly again until the junction with the bulb. Both parts of the pharynx clearly distinguishable since the anterior part expands always abruptly (Fig. 1 K, L) into the basal bulb and a more or less pronounced constric­ tion can mark the separation. Pharyngeal bulb typically cylindrical and relatively short, occupying one-fourth to one-third of the total neck length (in D. subhasi the bulb represents 43-45 % of the total neck length, an excep­ tional percentage in the genus). Nuclei and outlets of the r' pharyngeal glands always easily visible; DN and SzN in Fig. 1. A-C: Lip region in surface lateral view (A: Dorylai­ general clearer than S JN; these are situated somewhat moides rorundicephalus; B: D. hispanicus; C: D. grandis); D: Lip region in median lateral view (D. rotundicephalus); behind the middle of the DN-SzN distance. Cardia he­ E-G: OdonlOstyle (E: D. rorundicephalus; F: D. grandis; G: mispherical or rounded conoid (Fig. 1]), relatively small D. his panicus); H: Pharynx (D. striarus); I: Imestine-prerec­ and often surrounded by intestinal tissue. Intestine a LUmjunction (D. hispanicus); J: Cardia (D. rorundicephalus); simple tube without special modifications; cells number K, L: Pharyngeal basal bulb (K: D. striarus; L: D. reres). Jow; cells characterized by a granular aspect and promi­ nent nucleus. Transition between the intestine and the prerectum generally marked off by three guard cells part normally rounded but sometimes angular and the (Fig. 1 I). Prerectum length variable even in the same inner part frequently elevated and even forming perioral species or population and ranges from three to ten anal liplets. Labial and cephalic papillae with the typical body widths. Rectum (in females) a short tube equal to (6 + 6 + 4) distribution for dorylaims and easily percep­ or scarcely longer than the anal body diameter. tible by their often clear innervation since they are gen­ Female genital system: An interesting interspecific vari­ erally not raised above the head contour. ability can be noted in its morphology (Fig. 2 A, D) : Amphid: Amphid opening at the level of the cephalic two ovaries or only the posterior one, can exist. Didel­ depression or constriction (the base of the lip region), or phic species with two similar genital branches each con­ inmediately behind, and generally occupying more than sisting of ovary, oviduct, sphincter and uterus. In spe­ half of the corresponding body diameter. Fovea cup-like cies with only the posterior ovary developed, the and more rarely stirrup-like;jusus located near the odon­ posterior genital tract is similar to a genital branch of the tophore base, more clearly visible in dorsal or ventral didelphic species and the anterior one is normally re­ vIew. duced to a more or less developed uterine sac (mono­ delphic-opisthodelphic species sensu la to ). D. limnophi­ Cheilostome and guiding ring: Oral aperture followed lus is the only species with the anterior sac practically by a cylindrical or truncate conical cavity (cheilostome) absent (monodelphic-opisthodelphic species sensu stric­ whose walls appear sometimes slightly refractive and to). Moreover, in a few species, termed by us (Peralta & somewhat thickened in the perioral area; length of the Peria Santiago, 1995b) pseudodidelphic-opisthodel­ cavity 2-3 times its maximum width; guiding ring al­ phic, the anterior tract appears to be constituted by a ways simple and conspicuous since it is relatively thick sphincter, a vestigial oviduct with solid appearance, and and refractive. even a small cell mass resembling a rudimentary ovary; Stylet: The stylet morphology (Fig. 1 D) is one of the we prefer the terminology pseudodidelphic versus pseu­ characteristic features of the genus. The odontostyle is domonodelphic (cl Cohn & Sher, 1972) because these

244 Fundam. appl. NemalOl. Genus Dorylaimoides I. Taxonomy

species present both genital tracts (genital rubes) al­ though the anterior one is reduced or not functional. Ovary almost always medium sized, reaching the ovi­ duct-uterus junction but rarely continuing past it; 00­ cytes siruated first in two or more rows and then in a single row. Oviduct joining the ovary subterminally and consisting of a slender part with cubic or prismatic cells and a generally well developed pars dilatata. A promi­ nent sphincter, often with a more refractive inner part surrounded by the circular muscularure, separates ovi­ duct and uterus. Uterus a rube shorter and wider than the oviduct but without special modifications. Vagina cylindrical (Fig. 2 G) or somewhat pyriform and ex­ tending over half or scarcely less than the corresponding body diameter; its wall adjacent to the vulva composed of inner body cuticle extending inwards and widening in this region, the other part of the wall encircled by a more or less developed muscularure. Vulva, in all cases in which it was observed in frontal view, a transverse slit. Male genital system: Following the general dorylaim pattern and consisting oftwo opposed testes leading into a common vas deferens which joins the ejaculatory duct, the ejaculatory duct opens into the recrum and together they form the cloaca whose surface opening is the cloa­ cal opening. The system does not have special taxonom­ ic interest since it presents only very little morphological variation; however, the secondary sexual organs com­ posed of the copulatory appararus (spicules and lateral guiding pieces) and genital papillae (supplements), al­ though rather constant in the genus, are useful in dis­ tinguishing the species. Spicules more or less ventrally curved (Fig. 2 E, F) and consisting of a poorly devel­ oped head, a blade and a prominent median piece; main variation affects size (19-56 fLm) and the more or less slender aspect. Lateral guiding pieces relatively small, almost straight or sigmoid (Fig. 2 H, I), and with the tip acute or more frequently bifurcate. Supplements mam­ miform differing scarcely among the species; in addition to the adanal pair, a series of one to twelve regularly spaced ventromedian papillae is present beginning nor­ 'I mally at the level of the spicules, but with some in­ terspecific, or even intraspecific variability. J Tail: General morphology (size and shape mainly) extremely variable in the genus (Fig. 3). However, with­ in a particular species, both sexes have tails practically identical, except that the male tail is often somewhat shorter and more ventrally curved than in females. In­ traspecific variability of the caudal region in general small (it should be pointed out that most species are Fig. 2. A-D : Female genital sySlem. A : Didelphic species (Dory­ known from only one or a few populations and from a Iaimoides teres); B : Pseudodidelphzc-opislhodelphic species (D. small number of specimens), but interspecific variation rorundicephalus); c: Monodelphic-opislhodelphic species sensu is very large. Tail frequently filiform in the genus, practi­ lato (D. grandis); D : Monodelphic-opislhodelphic speCles sensu cally straight or slightly ventrally curved but in many stricto (D. limnophilus); E, F: Spicules (E: D. arcuaruSj F: D. cases tip dorsally recurved; from this filiform shape, aJJ hispanicus); G: Vagina (D. paraconfuslls); H, J: La/eralguid­ transitions can be found to a very short hemispherical ing pieces (H: D. paraconfusus; J: D. grandis). one.

Vol 20, n° 3 - 1997 245 R. Peria Samiago & M. Peralta

are rather close, varying from 3 fJ..m (minimum) to 11 fJ..m (maximum). The existence of a constriction bet\Veen the t\vo parts of the pharynx must be considered as the apomorphic state as compared to absence of constriction, in spite of the fact that the t\VO states can be found within the same species and even the same population. It seems that a tendency towards the reduction of the anterior genital branch exists inside the genus. Conse­ quently, a didelphic system must be considered plesio­ morphic with respect to the opisthodelphic system sensu stricto, which is considered apomorphic. In this context, the species presenting a pseudodidelphic-opisthodel­ phic or opisthodelphic sensu lato genital system can be interpreted as intermediate steps in the process of reduc­ tion. The fact that the pseudodidelphic-opisthodelphic condition is only rarely found can indicate (Coomans, in lill.) that it is a transitionary situation which is probably selected out rapidly; indeed, whereas such a long branch with" annex" does not seem to serve any useful func­ tion, a short uterine sac can store sperm and is important for ovejector formation. In any case, under a taxonom­ ical point of view, it is useful (cf Southey, 1973) to consider and distinguish the degree of reduction of the anterior genital tract with t\Vo clear possibilities: only uterine sac or uterus, sphincter and more or less devel­ Fig. 3. [mrageneric variability of the tail morphology in several oped oviduct and ovary. Spanish species of the genus DoryJaimoides. A: D. grandis; B: D. limnophilus; C: D. arcuarus; D: D. sp.; E: D. baeticus; F: It is possible to accept that the morphology of the tail D. rorundicephalus. presents a clear evolutive tendency from a more primi­ tive filiform shape towards an apomorphic state with short rounded aspect. However, bet\Veen these t\VO ex­ Taxonomical considerations tremes, we find almost all the intermediate possibilities of shape and size whose interpretation is more difficult EVOLUTIVE TRENDS fN THE GENUS AND INTRAGENER­ due to the numerous different cases and the intraspecific le TAXONOMY variability observed within each one of these cases. We Dorylaimoides seems to be a diversified genus with a propose a tentative hypothesis in which from the fLliform large amount of polymorphism in several morphological tail (plesiomorphic state) three basically different mod­ qualitative and quantitative features. To determine the els can be derived (short rounded, conical and regularly direction of changes produced during the evolutionary ventrally curved, and conical with the ventral side history is an intricate problem. However, the most im­ straight or dorsally bent at the tip) which are considered portant diagnostic features can be determined and their apomorphic states, and this evolutive pattern has repeat­ plesiomorphic and apomorphic character states can be ed itselfseveral times in different species and in different investigated. moments of the evolutionary history of the group. The A rounded lip region, continuous with the adjacent same apomorphic character state presented by different body (see D. parateres, D. bre'videns or D. constriclOides) species could be thus due to convergence. This would could be interpreted as an apomorphic state against the mean that similar species may have arisen from different angular and offset lip region (see D. grandis), which can ancestors. be considered as plesiomorphic, but several grades of A genus containing a high number of species poses a cephalic differentiation can be found making the in­ challenge to taxonomists who may try to split it in order terpretation more difficult. to make the identification and classification of the spe­ A slender and relatively long odontosryle (equal to or cies easier. However, in our opinion, species should be somewhat longer than lip region width; see D. parateres, clustered in natural groups (based on their evolutionary D. teres or D. ornatus) is surely plesiomorphic vs a broad history) which then can be identified as subgenera or and relatively short odonstyle (equal to or scarcely more other taxa. Other tentatives based on artificial and than half of the lip region width; see D. grandis or D. speculative characters should be avoided. angustus) but, here again, there is a continuous range of As mentioned above, we think that, at the present and possibilities bet\veen the two extremes which, moreover, except for a few cases which will be commented upon

246 Fundam. appl. NemaLO/. Genus Dorylaimoides 1. Taxonomy below, the species included in Dorylaimoides constitute a clearly represents an apomorphic state, in our opinion, natural and rather well defined group whose general such a unique character does not justify the proposal ofa evolutionary trends can be established. The relation­ different genus; in fact, several dorylaimid (and not be­ ships among these species nevertheless remain obscure londirid) genera as Nygolaimus (see, for instance, firstly because of poor information available for a num­ Heyns, 1968) and Careharolaimus (see Pefia Santiago & ber ofthem, secondly and more important, because apo­ Liebanas, 1994) include one or more species having a morphic states of several morphological features have sheath around the bulb. However, we have not exam­ originated independently in different species and at dif­ ined any material of iVfydonomus and, at this moment, ferent moments of the evolutionary history of the group. we prefer not to propose the corresponding synonymy. It means that a morphological similarity do not necessari­ The unique differential feature between Dorylai­ ly represent a recent relationship. So, we cannot support moides and Morasia is the tail that is morphologicallly the proposal of]airajpuri and Ahmad (1992) who have similar or dissimilar (filiform or elongated in female, divided D01ylaimoides into six subgenera (Dorylai­ rounded in male) in both sexes, respectively. It repre­ moides, Digidorylaimoides, Longidmylaimoides, Shamimo­ sents an evolutionary trend repeated in other dorylaimid nema, Areidorylaimoides and Tmjania), mainly on the groups without a common recent history as, e.g., Prodo­ basis ofthe morphology ofthe female genital system and rylaimuslLaimydorus, ThornenemalSieaguttur or Tra­ the tail. We prefer not to recognize or to define any ehaetinolairnuslAetinolaimus, which has been discussed, supraspecific taxa in the genus. among others, by Loof (1983, 1990) and Coomans and Carbonell (1988). The existence of dissimilar tails in T AXONOMICAL POSITION OF THE GENUS AND RELA­ both sexes represents an apomorphic state derived from TIONSHIPS the corrresponding plesiomorphic one present in Dory­ Species classified under Dorylaimoides certainly form laimoides species. In our opinion it is possible that the a natural (monophyletic) group because of several species presenting this derived state do not share a very shared derived characters, of which the most significant recent ancestor and, as a consequence, do not constitute are the morphology of the odonstosryle, the odonto­ a natural group, and therefore must be included in Dory­ phore and the pharyngeal bulb. lairnoides. However, taking into consideration that this The taxonomical position of the genus has been the idea is purely speculative since we cannot bring any object of controversy: Thorne (1939), Goseco et al. evidence supporting it and, so, it has the same scientific (1976) and ]airajpuri and Ahmad (1992) considered it value as the alternative argumentation, we maintain the as rather close to leptonchid nematodes including it into species having dissimilar tail as a separate group under the same superfarnily (Leptonchoidea or Tylencholai­ Morasia. Those species in which males are not known or moidea), while Andnissy (1976) classified the genus have not been found must be provisionally retained un­ under in Dorylaimoidea. We support in der Dorylaimoides. part this last proposal in the sense that the genus must be For the present we accept, according to ]airajpuri and included in the superfamily Dorylaimoidea since several Ahmad (1992), the inclusion of Dorylaimoides in the features, such as the morphology of the stylet and the subfamily Mydonominae Thorne, 1964 in Mydonomi­ cuticle, clearly separates Dorylaimoides from members of dae Thorne, 1964, and support their classification of the the superfarnily Tylencholaimoidea with which it shares six genera in two subfamilies. However, we consider that the short pharyngeal bulb; however, this bulb, even Mydonomidae must be classified under the superfamily when it is relatively short in relation to the neck length, is Dorylaimoidea, not in Tylencholaimoidea; the relation­ longer than in leptonchs. Moreover, it is known that the ships of Mydonomidae with other families of Dorylai­ reduction of the bulb length is a trend which has surely moidea must be discussed in a more general revision of occurred several times and in different groups in the this group. evolutionary history of dorylaims. The oligocytous in­ testine is a feature also shared with leptonchs but this fact is surely a homoplasy. Dorylairnoides Thorne & Swanger, 1936 The relationship between Dorylaimoides and two oth­ er genera, viz. lvfydonomus Thorne, 1964 and Morasia = Tmjania Brzeski & Szczygiel, 1961 Baqri & ]airajpuri, 1969, is obvious and merits some = LepLOnema ]airajpuri, 1964 nee LeplOnema Guerin­ comments (see inrnediately below). With other genera Meneville, 1843 as Calolaimus Timm, 1964, Timmus Goseco, Ferris & =Sharnimonema Chawla, Khan & Prasad, 1965 Ferris, 1976 and Miranema Thorne, 1939, Dorylai­ =Dorylaimoides (Dorylairnoides) Thorne & Swanger, moides presents important and significant differences 1936 which cannot be analyzed here. = Dorylaimoides (Digzdorylaimoides) Jairajpuri & Ah­ Mydonomus is distinguished from Dorylaimoides by mad,1992 the presence (vs absence) of a muscular sheath sur­ = Dorylaimoides (Longidorylaimoides) Jairajpuri & Ah­ rounding the pharyngeal bulb. Although this feature mad, 1992

Vol. 20, n° 3 - 1997 247 R. Peiia Santiago & M. Peralca

= Dorylaimoides (Shamimonema) Jairajpuri & Ahmad, D. charnoliensis Ahmad & Jairajpuri, 1983 1992 =D. (Dorylaimoides) charnoliensis Ahmad & Jairaj­ =Dorylaimoides (Arcldorylaimoides) Jairajpuri & Ah­ puri, 1983 mad, 1992 D. chathami Yeates, 1979 =Dorylaimoides (Tarjania) Jairajpuri & Ahmad, 1992 = D. (D01ylaimoides) chatharni Yeates, 1979 D. confusus Peralta & Pefia Santiago, 1995 DIAGNOSIS D. constrietOldes Goseco, Ferris & Ferris, 1976 Slender nematodes of medium size, almost always =D. (Tarjania) constrietoides Goseco, Ferris & Fer­ 1-2 mm long. Habitus ventrally curved, to C-shaped. ris, 1976 Outer cuticle relatively thin and with fine transverse D. constrietus Baqri & Jairajpuri, 1969 striations. Inner cuticle thicker than the outer layer. Lat­ =D. (Arcidorylairnoides) constn'ctus Baqri & Jairaj- eral chord very narrow. Odontostyle asymmetrical, ven­ puri, 1969 tral side shorter than the dorsal one and practically D. conurus Thorne, 1939 straight but dorsally bent at its anterior end; dorsal side D. cylindricaudatus Peralta & Pefia Santiago, 1991 longer and somewhat sigmoid. Odontophore usually ar­ D. dactylurus Heyns, 1963 cuate or angular, involved by the pharynx. Guiding ring = D. (Sharnirnonema) daetylurus Heyns, 1963 simple. Pharynx consisting of a slender and weakly mus­ D. elaboratus Siddiqi, 1965 cular anterior part expanding more or less abruptly into = D. (Longidorylaimoides) elaboratus Siddiqi, 1965 a basal bulb. Pharyngeal bulb cylindrical and occupying D. elegans (de Man, 1880) Thorne & Swanger, 1936 one-fourth to one-third of the total neck length. Female =Dorylairnus elegans (de Man, 1880) genital system didelphic, pseudodidelphic-opisthodel­ =D. (Digidorylaimoides) elegans (de Man, 1880) phic or opisthodelphic. Vulva transverse. Male genital Thorne & Swanger, 1936 system diorchic. Spicules dorylaimoid, 19-56 /-lm long. D. elongatus Husain & Khan, 1968 Lateral guiding pieces small. One to twelve ventrome­ =D. (Arcidorylairnoides) elongatus Husain & Khan, dian supplements regularly spaced and beginning usu­ 1968 ally into the range of the spicule. Tail similar in both D. enodis Goseco, Ferris & Ferris, 1976 sexes: fliiform, conical-elongated, conical, conoid or he­ =D. (Dorylaimoides) enodis Goseco, Ferris & Ferris, mispherical. 1976 D. filicaudatus Jana & Baqri, 1981 TYPE SPECIES =D. (Longidorylairnoides) filicaudatus Jana & Baqri, D. teres Thorne & Swanger, 1936 1981 =D. (Dorylaimoides) teres Thorne & Swanger, 1936 D. grandis Peralta & Pefia Santiago, 1995 D. hispanicus Peralta & Pefia Santiago, 1995 OTHER SPECIES D. ilyasi Ahmad & Jairajpuri, 1980 D. akon Goseco, Ferris & Ferris, 1976 =D. (Tarjania) ilyasi Ahmad & Jairajpuri, 1980 =D. (Longidorylaimoides) akon Goseco, Ferris & D. indicus Jairajpuri, 1965 Ferris, 1976 =D. (Dorylazrnoides) indicus Jairajpuri, 1965 D. angustus Sauer, 1967 D. kalingus Ahmad & Jairajpuri, 1983 =D. (Sharnimonema) angustus Sauer, 1967 = D. (Sharnirnonema) kalingus Ahmad & Jairajpuri, D. arcuatus Siddiqi, 1964 1983 =D. (Arcidorylairnoides) arcuatus Siddiqi, 1964 D. lepidus Timm, 1964 =D. rusticus Timm, 1964 =D. (Longidorylairnoides) lepidus Timm, 1964 =D. intermedius Thorne, 1964 D. leptura Siddiqi, 1965 D. arcuicaudatus Baqri & Jairajpuri, 1969 =D. (Longidorylaimoides) leptura Siddiqi, 1965 =D. (Dorylairnoides) arcuicaudatus Baqri & Jairaj- D. leptus Husain & Khan, 1968 puri, 1969 =D. (Digidorylairnoides) leptus Husain & Khan, D. ariasae Loof, 1990 1968 D. baeticus Peralta & Pefia Santiago, 1991 D. lzmnophilus (de Man, 1880) Loof, 1964 D. brevidens Thorne, 1964 = lirnnophilus de Man, 1880 = D. (Tarjania) brevidens Thorne, 1964 = Thornenema lzrnnophllum (de Man, 1880) An­ D. buccinator Sauer, 1967 drassy, 1959 = D. (Dorylairnoides) buccinator Sauer, 1967 =D. (Tarjania) lirnnophilus (de Man, 1880) Loof, D. bulbosus (Brzeski & Szczygiel, 1961) Szczygiel, 1964 1965 =Dorylairnoldes riparius Andrassy, 1962 = Tarjania bulbosa Brzeski & Szczygiel, 1961 =D. (Tarjama) riparius Andrassy, 1962 =D. (Tarjania) bulbosa (Brzeski & Szczygiel, 1961) D. longicaudatus (Imamura, 1931) Thorne & Swan­ Szczygiel, 1965 ger, 1936

248 Fundam. ap-pl. NemalOl. Genus Dorylaimoides 1. Taxonomy

= Dorylaimus elegans var. longicaudaws Imamura, SPECIES lNQUIRENDAE 1931 D. microdenlalus Altherr, 1968 [probably not Dorylai­ = D. (Longidorylaimoides) longicaudalUs (Imamura, moides] 1931) Thome & Swanger, 1936 D. pan' Maha;an, 1973 [a prodelpruc species] D. longiurus Siddiqi, 1965 D. stenodorus Altherr, 1953 [described in base to a = D. (Tarjania) longiurus Siddiqi, 1965 single male] D. loofi Baqri & Khera, 1979 D. thienemanni (Schneider, 1937) Jairajpuri, Ahmad D. malabaricus Ahmad & Jairajpuri, 1982 & Dhanachand, 1980 = D. (Longidorylaimoides) malaban'cus Ahmad & Jai­ = Dorylaimus thienemanni Schneider, 1937 rajpuri, 1982 = Thomenema thienemanni (Schneider, 1937) An­ D. micoletzkyi (de Man, 1921) Thorne & Swanger, drassy, 1959 1936 = D. (Tarjania) thienemanni (Schneider, 1937) Jai­ = Dorylaimus micoletzkyi de Man, 1921 rajpuri, Ahmad & Dhanachand, 1980 =D. (Digidorylaimoides) micoletzkyi (de Man, 1921) Thorne & Swanger, 1936 AcknowledgeITlents = D. pakislanensis Siddiqi, 1964 Authors thank Prof. A. Coomans and Dr. P.A.A. Loof for = D. (Digidorylaimoides) pakislanensis Siddiqi, 1964 the critical revision of the first manuscript of the paper and D. mitis Sauer, 1967 their interesting suggestions which have greatly improved the =D. (Shamimonema) mitis Sauer, 1967 final text and content. The first author is also grateful for the D. modeslUs Siddiqi, 1965 financial support received from the project entitled" Fauna = D. (Tarjania) modeslUs Siddiqi, 1965 Ibhica Ill" (DGICYT PB92-0121). D. omalUs Peralta & Pena Santiago, 1995 References D. paraconfusus Peralta & Pena Santiago, 1995 AHMAO, W. &JAlRAJPURI, M.S. (1980). Four new species of D. paraleres Siddiqi, 1964 Leptonchoidea (Nematoda: Dorylaimida). Indian]. Nema­ = D. (Dorylaimoides) paratms Siddiqi, 1964 LOl.,9 (1979): 125-135. D. paruus Thorne & Swanger, 1936 =D. (Longidorylaimoides) paruus Thorne & Swan­ AHMAO, W. & JAlRAJPURl, M.S. (1982). Three new and two ger, 1936 known species of dorylaim nematodes with proposal ofAee­ = D. n"pan'us apud Loof, 1964 phalodorylalmus n. gen. NemaLOloglea, 28 : 233-246. D. paulbuchnen' Meyl, 1956 AHMAO, W. & JAlRAJPURI, M.S. (1983). Descriptions of new = D. (Longidorylaimoides) paulbuchnen Meyl, 1956 species of Dorylalmoldes and Calolaimus (Dorylaimida) D. prelOn'ensis Heyns, 1963 from India. Revue NemaLOl., 6: 65-72. = D. (Shamimonema) prelOn'ensis Heyns, 1963 ALTHERR, E. (1953). Nematodes du sol du Jura vaudois et D. reversus Thorne, 1964 francais (1). Bull. Soc. Vaud. Sa. naL. (284),65: 429-460. = D. (Tarjania) reversus Thorne, 1964 ALTHERR, E. (1968). Nematodes de la nappe phreatique du D. TOtlmdiaphalus Peralta & Pena Santiago, 1995 reseau fluvial de la Saale (Thuringe) et psammniques du D. sauen' Baqri & Jairajpuri, 1969 Lac Stechlin (Brandebourg du nord). Limnologiw, 6: 247­ = D. (Arcidorylaimoides) sat/en' Baqri & Jairajpuri, 320. 1969 ANORAssY,1. (1959a). Neue und wenig bekannte Nematoden D. siddiqii Baqri & Khera, 1979 aus Jugoslawien. Annls hisl.-naL. Mus. nam. hung., 51 : 259­ D. similis Thorne, 1964 275. = D. (Longidorylaimoides) similis Thorne, 1964 ANoRAssY,1. (1959b). Taxonomische Obersicht der Dorylai­ D. sln'atus Peralta & Pena Santiago, 1995 men (Nematoda) 1. ACla zool. hung., 5 : 191-240. D. subhasi Jana & Baqri, 1981 = D. (Tarjania) subhasi Jana & Baqri, 1982 ANORAssy, 1. (1962). Zwei neue Nematoden-Arten aus dem D. thecolaimus Heyns, 1963 Oberschwemmungsgebiet der Donau (Danubiala Hungar­ = D. (Dorylaimoides) thecolaimus Heyns, 1963 ica, XlII). Opuse. zool., Bpesl., 4: 3-8. D. thomei Qairajpuri, 1964) Siddiqi, 1969 ANORAssy, 1. (1976). EvolUllon as a basis for lhe syslemallza­ = Leptonema thornei Jairajpuri, 1964 lion of nemaLOdes. London, UK, Pitman Publishing, 287 p. = Shamimonema thomei Qairajpuri, 1964) Chawla, BAQRI, Q.H. &JA1RAjPURI, M.S. (1969). Morasia n. gen., and Khan & Prasad, 1965 three new species ofDorylaimoides Thorne & Swanger, 1936 = D. (Shamimonema) thomez Qairajpuri, 1964) Sid­ (Nematoda: Dorylaimoidea) from India. NemaLOlogica, 15 : diqi, 1969 408-424. D. venustus Andnissy, 1959 BAQRI, Q.H. & KHERA, S. (1979). Nematodes from West = D. (Tarjania) venuslUS Andnissy, 1959 Bengal (India) IV. Three known and two new species of the D. websten' Sauer, 1967 genus Dorylaimoides Thorne & Swanger, 1936 (Leptonchi­ = D. (Dorylaimoides) websten' Sauer, 1967 dae: Dorylairnida). Ree. zool. Surv. India, 75 : 247-254.

Vol. 20, n° 3 - 1997 249 R. Peiia Santiago & M. Peralta

BRZESKJ, H.W. & SZCZYGIEL, A. (1961). Two new species of LOOF, P.A.A. (1990). Systematic observations on some spe­ the subfamily DoryJaiminae (Nematoda, Dorylaimidae). cies of Dorylaimoididae (Nemata : Dorylaimida). Revue Ne­ Bull. Acad. Polon. Sci., 9: 511-514. matol., 13: 161-170. CHAWLA, M.L., KHAN, E. & PRr\SAD, S.K. (1965). Shamimo­ MAHA.lAN, R. (1973). Two new nematodes, Tylenchus kash­ nema nom. novo for Leptonema Jairajpuri, 1964. Labdev]. mirensis n. sp. (Tylenchidae) and D01ylaimoides pan' n. sp. Sci. Technol., 3 : 143. (Dorylaimoididae) from Kashmir, India. Riv. Parassit., 34 : 201-204. COHN, E. & SHER, S.A. (1972). A contribution to the taxono­ my of the genus Cobb, 1913. J. Nematol., 4 : DE MAN, ].G. (1880). Die einheimischen, frei in der reinen 36-65. Erde und im silssen Wasser lebenden Nematoden. Tijdschr. Ned. Dierk. Vereen., 5 : 1-104. COOMANS, A & CARBONELL, E. (1988). The status of the family Thornenematidae Siddiqi, 1969 (Nematoda: Dory­ DE MAN, JG. (1921). 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(I 991). Two new spe­ HEYNS,J (1963). New species of the superfamily Dorylaimoi­ cies of the genus D01ylaimoides Thorne et Swanger, 1936 dea (Nemata) from South African soils, whith a description (Nematoda: Dorylaimida) from Spain. Nematologica, 37 : of a new genus Kochinema. S. Afr. J. agric. Sci., 6 : 289-302. 382-394. HEYNS, J (1968). A monographic study of the nematode fami­ PERALTA, M. & PENA SANTiAGO, R. (I 995a). Nematodes of lies and NygoJaimeJlidae. Ent. Mem. Dep. the order Dorylaimida from Andalucia Oriental, Spain. The Agn·c. Techn. Serv., Repub. S. Afr., 19 : 1-144. genus Dorylaimoides Thorne & Swanger, 1936. I. Didelphic HUSAlN, S.l. & KHAN, AM. (1968). BasirotyleplUs modestus n. species. Fundam. appl. NemalOl., 18 : 35-53. sp. and rwo new species of Dorylaimoides Thorne & Swan­ PERALTA, M. & PENA SA.1'JTIAGO, R. (I 995b). Nematodes of ger, 1936 from India. Nematologica, 14: 362-368. the order Dorylaimida from Andalucia Oriental, Spain. The IMM-IURA, S. (1931). Nematodes in the paddy field, with genus Dorylaim01'des Thorne & Swanger, 1936.2. Pseudodi­ notes on their population before and after irrigation. J. Coli. delphic-opisthodelphic species. Fundam. appl. NemalOl., Agric. Imp. Univ. Tokyo, 11 : 193-240. 18: 167-180. JAlRAJPURI, M.S. (1964). Leptonema thomei n. gen., n. sp. PERALTA, M. & PENA SANTiAGO, R. (1995c). Nematodes of (Nematoda : Dorylaimoidea) from India. Nematologica, 10 : the order Dorylaimida from Andalucia Oriental, Spain. The 399-402. genus Dorylaimoides Thorne & Swanger, 1936. 3. Opistho­ delphic species. Fundam. appl. NemalOl., 18: 315-327. JAlRAJPURl, M.S. (1965). D01ylaimoides indicus n. sp. (Nema­ SAUER, M.R. (I967). Four new species of DOlylaimoides. Ne­ toda : Dorylaimoidea) from India. Labdev. J. Sci. Technol., malOlogica, 12 : 523-529. 3: 124-125. SCHNEIDER, W. (1937). Freilebenden Nematoden der deut­ JAlRAJPURI, M.S. & AHMAD, W. (1992). DOlylaimida. Free­ schen limnologischen Sunda Expedition nach Sumarra, Java living, predaceous and plant-parasitic Nematodes. Leiden, The und Bali. Arch. Hydrobiol., 15 : 30-108. Netherlands, EJBrill : 458 p. SIDDIQI, M.R. (1964). Three new species of Dorylaimoides JAlRAJPURl, M.S. AHMAD, M. & DHANACHAND, CH. (1980). Thome & Swanger, 1936 with a description of Xiphinema NomencJatorial notes on Tharnenema thienemanni with a key orbum n. sp. (Nematoda: Dorylaimoidea). NemalOlogica, 9 : to species of Thomenema (Nematoda: Dorylaimida). Ne­ 626-634. matologica, 26: 182-186. SIDDIQI, M.R. (1965a). Seven new species of Dorylaimoidea Jt\NA, A. & BAQRl, Q.H. (1981) : Nematodes from west Bengal (Nematoda) from India, with descriptions of Lenonchium n. (India) Xl. Studies on the species of the superfamily Lep­ gen. and Galophinema n. gen. Proc. helminth. Soc. Wash., 32 : tonchoidea (Dorylaimida). J. zool. Soc. India, 33 : 1-24. 81-90. LOOF, P.A.A. (1964). Free-living and plant-parasitic nema­ SIDDIQI, M.R. (1965b). Five new species of soil nematodes in todes from Venezuela. NemalOlogica, 10 : 201-300. the genera Dorylaimoides Thorne & Swanger, 1936, and LOOF, P.AA. (1983). Taxonomic problems in long-tailed do­ Discolaimium Thorne, 1939 from India. Nematologica, 11 : rylaims. In: Stone, AR., Plan, H.M. & Khalil, L.F. (Eds). 100-108. Concepts in nematode systematics, London, UK, Academic SIDDIQ1, M.R. (1969). Crateronema n. gen. (Crateronemati­ Press: 175-186. dae n. fam.), Poronemella n. gen. (LordeUonematinae n. sub-

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fam.) and Chrysonemoides n. gen. (Chrysonematidae n. THORNE, G. (1964). Nematodes of Puerto Rico. Belondiroi­ fam.), with a revised classification of Dorylaimoidea (Ne­ dea new superfamily, Thome, 1935, and Be­ matoda). NemalOwgica, 15: 81-100. lonenchidae new family (Nematoda, Adenophorea, Dory­ laimida). Techn. Pap. Univ. P. Rico agric. Exp. Sm, 39 : 1-51. SOUTHEY, IF. (1973). Terminology of the female reproduc­ tive organs in nematodes with special reference to Xiphine­ THORNE, G. & SWANGER, H.H. (1936). A monograph of the ma. NemalOlogica, 19: 405. nematode genera Dorylaimus Dujardin, Aporcelaimus n. gen.) Dorylaimoides n. gen., Pungenlus n. gen. Capita Zoo!., SZCZYGIEL, A. (1965). Taxonomic starus of Tarjania Brzeski 6: 1-223. & Szczygiel and redescription of Dorylaimoides bulbosus TINLVI, R.W. (1964). Nematodes of the superfamily Dorylai­ (Brzeski & Szczygiel, 1961) n. comb. (Nematoda: Lep­ moidea from East Pakistan. Proc. helminth. Soc. Wash., 31 : tonchidae). Nematologica, 1I : 409-412. 144-153. THORNE, G. (1939). A monograph of the nematodes of the YEATES, G. W. (1979). Nine new DOl}'laimida (Nematoda) superfamily Dorylaimoidea. Capita Zoo!., 8 : 1-261. fron the New Zealand region. Nemalologica, 25 : 419-438.

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