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Chordodes combiareolatus n. sp. (Figures 1, 2) Holotype. One female from the type locality, deposited in the Zoological Survey of India, North Eastern Regional Centre, Shillong under the accession number IV/NERC- 308. Etymology. The name ��� combiareolatus ��� refers to a unique type of areoles, in which characteristics of two areoles are combined. Type locality. Terrace paddy field in Tsupo, Viswema, Kohima, Nagaland, India (25 �� 34 ��� 50.59 ���N / 94 ��09��� 56.89 ���E), elevation: 4175 feet. Description of female. The specimen is 210 mm long and has a width of 1.9 mm in the midbody region. The anterior end is tapered; the anterior most tip is white. The main body colour is moderately dark brown with darker patches forming the so-called ���leopard pattern���. The posterior end is distinctly swollen (Fig. 2 C), the cloacal opening is terminal. In the midbody region, all 6 types of areoles known from the genus Chordodes are present. Areoles cover the body with moderate density, meaning that regions of non-areolar cuticular surface can be seen between the areoles (Figs. 1 A, 1 C). This surface is structured into parallel superficial folds (Fig. 1 A). The most abundant type of areoles is simple areoles (Figs. 1 A, 1 C). They are of differing height and smooth except for an apical tiny tuft of bristles. Tubercle areoles are present among simple areoles (Fig. 1 A). They appear as a mixture between simple and tubercle areoles by combining characters of both these areolar types. Tubercle areoles carry a tuft of small bristles on their apical surface and have a finger-like tubercle, which is rarely placed centrally, but more often shifted towards the edge of the areole or even originating from the side of the areole (Figs. 1 A, 1 B). There are also tubercle areoles that correspond to the ���typical��� form (i.e. without bristles and with a more or less central tubercle) (e.g. Fig. 1 C), but the ���combined type ��� occurs more often. Thorn areoles are present, and the thorns have a broad basis and are slightly curved (Fig. 1 C). Clusters of crowned areoles are present. On the lateral sides of the body, these clusters are composed of two central crowned areoles and 8���10 surrounding circumcluster areoles (Figs. 1 D, 1 E). The circumcluster areoles appear as elongated simple areoles, they also carry a tuft of bristles on top (Fig. 1 E). The crowned areoles have a flat apical surface, from which filaments of moderate length (10���15 ��m) extend laterally (Fig. 1 E). Some of these filaments divide into several branches. We observed in several cases that in one of the two crowned areoles the filaments branch off from the entire surface, while in the second areole the side facing the first areole remains free of filaments (Fig. 1 E). We could not observe a tubercle between the crowned areole, a structure which is present in several other Chordodes species. Along the midline (it was not possible to determine if this was the ventral or the dorsal one) crowned areoles occur on both sides next to the midline and have distinctly longer apical filaments (Fig. 1 D). The length of the filaments is maximally around 150 ��m. We could also observe a number of arrangements of areoles, in which areoles resembling circumcluster areoles surround a central flat region with roughly structures surface (Figs. 1 F, 1 G). In the anterior end, the shape of the areoles changes strongly (Fig. 2 A). The dominating type of areoles has a dense tuft of apical filaments on top (Fig. 2 B). These areoles are not arranged in particular patterns with respect to each other, but cover the cuticle densely. Between these areoles some long tubercles appear, but the origin of these tubercles was unclear, i.e. whether they originated directly from the cuticular surface or from an areole (Fig. 2 B). In the anterior-most tip of the animal the areoles still have a tuft of apical filaments, but not as dense as in the areoles described previously. Additionally, the apical surface is depressed (Fig. 2 A). In the posterior end the terminal cloacal opening is surrounded by a region with smooth cuticle with a radius of approximately 200 ��m (Figs. 2 C, 2 D). In the margin of this smooth circular region of the cuticle some bristles are present as the cuticle transitions into the typical areolar pattern (Fig. 2 E). Areoles transform from simple elevations of different shape to the areolar pattern described above. Crowned areole clusters start about 600 ��m from the margin between smooth and areolated cuticle. The circular appearance of the posterior end is only interrupted by one slight groove that runs to the margin of the terminal smooth region (Fig. 2 D). In this part, areoles are present a bit further inside the smooth region than in other places. Taxonomic comments. The specimen described above shows several characters well distributed among species of the genus Chordodes. Simple areoles are the most abundant type of areole, and tubercle and thorn areoles are also widely distributed types. Crowned areoles are characteristic for the genus Chordodes and in many species they occur along the ventral and sometimes also along the dorsal midline with very long bristles. The character that separates the new species from the previous descriptions is that the tubercle areoles include the tuft of bristles present in simple areoles. Usually, tubercle areoles have a smooth surface and the tubercle is positioned apically. Some exceptions are known. In the African species C. digitatus Von Linstow, 1901 the basal areolar part is horseshoe-shaped and surrounds the tubercle (Fig. 5 A, 5 B in De Villalobos et al. 2007). In the African species C. tuberculatus Von Linstow, 1901 tubercles appear to be displaced from areoles or attach laterally on areoles (see De Villalobos et al. 2007). In the Japanese species C. japonensis Inoue, 1951, tubercle areoles are usually of the ���standard pattern���, but one broad tubercle areole is documented, which carries a tubercle and a small tuft of bristles (Fig. 2 C in Schmidt-Rhaesa 2004). None of these cases looks exactly like in the new species, but this shows that too little attention has been paid to the exact shape of certain areolar types. The crowned areoles of the new species show (at least in several clusters) an asymmetrical pair of crowned areoles, where apical filaments originate from all sides in one areole, but not from all sides in the other areole. Such a pattern has not been described yet and may be another diagnostic character of the new species. The regions, in which some longer areoles surround a circular region of flat surface are interpreted by us as crowned areole clusters (circumcluster areoles), where the central crowned areoles are broken off. The reasons for this interpretation are that the regions with an ���empty center��� are distributed in the same pattern as otherwise the crowned areole clusters. The circular ring of elevated areoles resembles circumcluster areoles and the rough surface in the centre does not appear to be the regular, undamaged cuticular surface. Crowned areoles are prominent elevated structures that might be prone to mechanical damage and abrasion under some circumstances. The anterior end of the specimen described above shows a dramatic change in areoles and the areolar pattern in the anterior end differs completely from the one in midbody. This shows the importance of comparing cuticular sections from comparable body regions. The cuticular structure is rarely described in detail. Areoles with a dense tuft of bristles are described also from C. moutoni (see below) and from C. fomosanus Chiu, Huang, Wu & Shiao, 2011 (Chiu et al. 2011). Therefore the change of the areolar pattern may be further distributed than currently documented and we hesitate to include this character as species specific for C. combiareolatus. The posterior end in general corresponds to the female posterior ends that have been described for the genus Chordodes. It is usually swollen, in contrast to the posterior ends in other genera (see Schmidt-Rhaesa 2012) and the terminal cloacal opening is in the center of a flat or even concave terminal disc., Published as part of Schmidt-Rhaesa, Andreas & Yadav, Arun K., 2015, Chordodes combiareolatus, a new species of horsehair worms (Nematomorpha) from Nagaland, India, with further comments on Chordodes moutoni, pp. 202-210 in Zootaxa 3925 (2) on pages 203-206, DOI: 10.11646/zootaxa.3925.2.3, http://zenodo.org/record/237357, {"references":["Linstow, O von. (1901) Helminthen von den Ufern des Nyassa-Sees. Jenaische Zeitschrift fur Naturwissenschaften, 35, 409 - 428.","De Villalobos, C., Zanca, F. & Schmidt-Rhaesa, A. (2007) Revision of the genus Chordodes (Gordiida, Nematomorpha) from Africa-III: ultrastructural redescription of Chordodes capensis Camerano, 1895, C. clavatus Linstow, 1906, C. digitatus Linstow, 1901, C. tuberculatus Linstow, 1901, and reinterpretation of C. ibembensis Sciacchitano, 1958 and C. uncinatus Sciacchitano, 1958. Journal of Natural History, 41 (45 - 48), 2775 - 2788. http: // dx. doi. org / 10.1080 / 00222930701754855","Inoue, I. (1951) An additional new species of Gordiacean parasitizing praying mantids, and the domestic and overseas distribution of the genus Chordodes. Zoological Magazine (Tokyo), 60, 32.","Schmidt-Rhaesa, A. (2004) Japanese Horsehair Worms (Nematomorpha) from the Lake Biwa Museum. Species Diversity, 9, 97 - 107.","Chiu, M. C., Huang, C. G., Wu, W. J. & Shiao, S. F. (2011) A new horsehair worm, Chordodes formosanus sp. n. (Nematomorpha, Gordiida) from Hierodula mantids of Taiwan and Japan with redescription of a closely related species, Chordodes japonensis. ZooKeys, 160, 1 - 22. http: // dx. doi. org / 10.3897 / zookeys. 160.2290"]}

Chordodes moutoni Camerano, 1895 (Figure 3) Material examined. From same location as holotype of C. combiareolatus, but collected in a small time difference on same date: Terrace paddy field in Tsupo, Viswema, Kohima, Nagaland, India (25 �� 34 ��� 50.59 ���N / 94 ��09��� 56.89 ���E), elevation: 4175 feet. Description of female. The specimen is 200 mm long and has a width of 1.6 mm in the midbody region. The anterior end is tapered, the anterior-most tip is white. The main body colour is moderately dark brown with darker patches forming the so-called ���leopard pattern���. The posterior end is distinctly swollen, the cloacal opening is terminal. In the midbody region, all 6 types of areoles known from the genus Chordodes are present. Some surface areas between areoles are visible and show the parallel superficial folds, which in some regions form slight elevations (Fig. 3 A). Such elevations are low compared to areoles, so it is not clear whether they represent areoles. The most abundant type of areoles is simple areoles (Fig. 3 A). They are of differing height and have an apical tuft of bristles. Tubercle areoles are present among simple areoles (Fig. 3 A). They have a smooth surface, bristles could not be observed. The tubercle arises more or less from the center of the areoles. Thorn areoles were rarely observed, but are present (not figured here). Clusters of crowned areoles are present in the same distribution as in the previous species. On the lateral sides of the body the clusters are composed of two central crowned areoles and 10���12 surrounding circumcluster areoles, although the number is hard to count because circumcluster areoles decrease gradually in size and then cannot be distinguished from simple areoles. The circumcluster areoles are slender and elongated and have a tuft of bristles apically. The crowned areoles do not have a flat apical surface, the apical bristles form a dense tuft of thin filaments. Their length is moderate (10���15 ��m). Division of filaments could not be observed, but cannot be excluded. A tubercle between the crowned areoles was not observed. Along the midline (ventral and/or dorsal could not be decided) crowned areoles occur on both sides next to the midline and have distinctly longer apical filaments (Fig. 3 B). The length of the filaments is about 300 ��m. The arrangements of areoles, in which areoles resembling circumcluster areoles surround a central flat region in which the cuticular surface is rough could also be observed (Figs. 3 C, 3 D). In the anterior end, the structure of the cuticle resembles the one described above for C. combiareolatus. The most abundant type of areoles has a large tuft of apical filaments on top (Fig. 3 E). Between these areoles are tubercle areoles and thorn areoles. The thorn areoles carry a thin, apically curved thorn (Fig. 3 E). As far as could be observed, the anterior-most areoles have no depressed surface. In the posterior end (Fig. 3 F) the terminal cloacal opening is surrounded by a region with finely striped cuticle. In the margin of this smooth circular region are some bristles and the transition to areoles. Taxonomic comments. Exact species determinations are often difficult in Nematomorpha, because the fine structural details are in some cases difficult to compare between older descriptions including drawings and newer descriptions including SEM images. Additionally, many specimens deviate from each other by very small differences and it is not known whether such differences can be due to intraspecific variation or are a document of interspecific variation. species C. combiareolatus C. moutoni Nagaland C. moutoni China, C. moutoni Malaysia C. moutoni Shillong lectotype source this investigation this investigation Zanca & De Villalobos Schmidt-Rhaesa & Brune Schmidt-Rhaesa & Yadav 2005 2008 2013 simple areoles with bristles with bristles rough surface, some with bristles with bristles areoles (described as type 3) with bristles In this case the specimens resembles in many details C. moutoni, a species first described from China (Camerano 1895), later from Malaysia (Camerano 1899, 1901, 1903, Schmidt-Rhaesa & Brune 2008) and again from China (Wu & Tang 1933). The original material consisted of two males and two females, which were distributed among two museums, Paris and Torino. The male and the female specimen from the Mus��um National d��Histoire Naturelle in Paris were reinvestigated by Zanca & De Villalobos (2005) and designated as lectotype (male) and paralectotype (female). Our specimen from Nagaland resembles the female paralectotype in many respects, but differs in the following characters. For reasons of simplicity, the specimens are in the following named the Nagaland specimen and the China specimen. The simple areoles of the China specimen have a rough surface and no clearly visible bristles on their apical surface. However, one type of areoles is described, which has some apical bristles, but in the images bristles appear more or less attached to the surface and not clearly visible (see Zanca & De Villalobos 2005). Thorn areoles are not described for the China specimen, but are present in the Nagaland specimen. As thorns are generally rare structures, they may be overlooked, especially when only small pieces of cuticle are investigated. The posterior end of the China specimen is slightly invaginated, so a proper comparison to the posterior end of the Nagaland specimen is not possible. The structure of the anterior end is not described for the China specimen. Two further specimens have been documented by SEM. One male specimen collected in Malaysia (see Schmidt-Rhaesa & Brune 2008) and one female from Shillong, North-Eastern India (Schmidt-Rhaesa & Yadav 2013) correspond in most characters with the Nagaland specimen. One interesting difference is the shape of crowned areoles. While in the Nagaland specimen and in the Malaysia specimen apical filaments originate as a kind of unordered tuft from the apical surface of areoles, the apical surface of the China specimen and the Shillong specimen is more or less flat and filaments originate to the side. Asymmetrical patterns as in C. combiareolatus were not observed, but in the Shillong specimen filaments branch from both crowned areoles in all directions (see Schmidt-Rhaesa & Yadav 2013) and in the China specimen filaments do not originate from the sides where crowned areoles face each other (see Zanca & De Villalobos 2005). Until now, almost no attention has been paid to this character and these differences should be kept in mind for further investigations. With the mentioned exceptions, specimens described as C. moutoni appear to belong to the same species. The small differences in comparison with the paralectotype are regarded by us as not fundamental enough to describe a new species, but this possibility should be kept in mind. For a comparison of characters among specimens documented by SEM see Tab. 1. Chordodes moutoni and C. combiareolatus were found in the same location. They show some correspondences and some differences. Most characters are common, widespread characters typical for Chordodes. Simple areoles are rarely described with apical bristles and a clear tuft of bristles was even used by Schmidt-Rhaesa et al. (2008) in a key of the genus Chordodes to separate two species, C. villalobi Schmidt-Rhaesa & Brune, 2008 from Malaysia and C. corderoi Carvalho, 1946 from Venezuela from all other species. It is suspected that tiny bristles are more commonly distributed than previously recorded, but this character can only be documented reliably with SEM. The special form of the tubercle areoles forms the most prominent difference between C. combiareolatus and C. moutoni, as the Nagaland specimen of C. moutoni has tubercle areoles with a smooth surface, as described for almost all other Chordodes species. One further difference is the shape of the crowned areoles. In C. combiareolatus they have a flat surface and filaments extend only to the sides, while in C. moutoni filaments extend like a tuft in all directions. Both characters were already described or figured for other Chordodes species (flat surface e.g. in C. mizoramensis Schmidt-Rhaesa & Lalramliana, 2011 and C. parabipilus Kintsurashvili, Schmidt-Rhaesa & Gorgadze, 2011, tuft of bristles e.g. in C. kenyaensis Bolek, Szmygiel, Kubat, Schmidt-Rhaesa & Hanelt, 2013; see Kintsurashvili et al. 2011, Schmidt-Rhaesa & Lalramliana 2011, Bolek et al. 2013), but this difference has not received much attention until now. The shape of crowned areoles should be described in more detail than has been done before., particularly the origin of the apical filaments, branching patterns of the apical filaments and their fine structure. The specimen determined here as C. moutoni corresponds with C. combiareolatus in two special characters. First, the cuticle contains the circular arrangements of areoles around an ���empty center���. As described above this is interpreted here as an artifact, but as this phenomenon has not been documented from other species, it remains unknown why two specimens of two different species should both show this kind of cuticular destruction. The second character is the areolar pattern in the anterior end. As described above, the anterior end is rarely investigated in detail and it remains to be shown whether the composition of areoles in the anterior end is particular or widely distributed among Chordodes., Published as part of Schmidt-Rhaesa, Andreas & Yadav, Arun K., 2015, Chordodes combiareolatus, a new species of horsehair worms (Nematomorpha) from Nagaland, India, with further comments on Chordodes moutoni, pp. 202-210 in Zootaxa 3925 (2) on pages 206-209, DOI: 10.11646/zootaxa.3925.2.3, http://zenodo.org/record/237357, {"references":["Camerano, L. (1895) Description d´une nouvelle especie de Gordien de la Chine. Bulletin de la Societe Zoologique de France, 20, 99 - 100.","Camerano, L. (1899) Gordii della Malesia e del Messico. Atti delle Reale Accademia delle Science di Torino, 34, 460 - 469.","Camerano, L. (1901) Gordii raccolti dalla spedizione \" Skeat \" nella Penisola Malese 1899 - 1900. Bollettino dei Musei di Zoologia ed Anatomia Comparata della Reale Universita di Torino, 16, 1 - 2.","Camerano, L. (1903) On the entomo-parasites of the \" Skeat expedition \". Proceedings of Zoological Society of London, 2, 152 - 153.","Schmidt-Rhaesa, A. & Brune, S. (2008) Description of one known and three new Chordodes species (Nematomorpha) parasitizing praying mantids (Mantoptera) in Malaysia, with a discussion of sexual dimorphism in Chordodes. Zoosystematics and Evolution, 84, 57 - 66. http: // dx. doi. org / 10.1002 / zoos. 200700014","Wu, H. W. & Tang, S. F. (1933) Notes on the Nematomorpha of China. Sinensia, 3, 173 - 178.","Zanca, F. & De Villalobos, C. (2005) Scanning electron microscopy of Chordodes moutoni Camerano, 1895 (Gordiida, Nematomorpha). Zootaxa, 1082, 37 - 44.","Schmidt-Rhaesa, A. & Yadav, A. K. (2013) One new species and a new record of the genus Chordodes. Zootaxa, 3693 (2), 293 - 300. http: // dx. doi. org / 10.11646 / zootaxa. 3693.2.10","Carvalho, J. C. M. (1946) Gordiaceos du Museo de Historia Natural de Montevido. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo, 2, 1 - 8.","Kintsurashvili, N., Schmidt-Rhaesa, A. & Gorgadze, O. (2011) Chordodes parabipilus (Nematomorpha: Gordiida), a new species of horsehair worms from Georgia. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg, 46, 235 - 241.","Bolek, M. G., Szmygiel, C., Kubat, A., Schmidt-Rhaesa, A. & Hanelt, B. (2013) Novel techniques for biodiversity studies of gordiids and description of a new species of Chordodes (Gordiida, Nematomorpha) from Kenya, Africa. Zootaxa, 3717 (1), 23 - 38. http: // dx. doi. org / 10.11646 / zootaxa. 3717.1.2"]}

Chordodes lasuboni n. sp. (Figures 1 B, 4, 5) Type locality. Lasubon Hostel, North-Eastern Hill University campus, Shillong, Meghalaya, India, 25 ��. 57 'N, 91 ��. 77 'E. Collected August 6, 2012 by Ms. Luiluile Lungalang. Holotype: A single male specimen from the type locality emerged from Hierodula sp.; alcohol preserved pieces and SEM stubs of holotype deposited in the Museum of Department of Zoology, NEHU, Shillong with the accession no. MDOZ/NEHU/INV/ 112. Paratype: Nil. Host. Holotype emerged from one specimen of Hierodula sp. (Mantodea) (Fig. 1 B). Etymology. The species is named after its type locality, the Lasubon Hostel on the campus of the North- Eastern Hill University. Description. The holotype is 195 mm long, with a diameter of 1.2 mm in mid-body region. The specimen is dark brown in colour. The anterior end is distinctly tapered (Fig. 5 E), the posterior end is curved ventrally. The cuticle contains six types of areoles (terminology of Schmidt-Rhaesa et al. 2008). Most abundant are simple areoles. They are variable in size and shape, but are usually rounded structures with few tiny bristles on their apical surface (Figs. 4 D, 4 E, 4 F, 5 B, 5 C). Tubercle areoles are present, and seem to be more abundant at the anterior end than at mid-body and at the posterior end (Figs. 4 D, 5 B). Thorn areoles are present, but rare. Of the pieces investigated here, few thorn areoles were found at the anterior end (Fig. 5 C) and one at the posterior end (Fig. 5 B). Thorns are slender and hook-like (Fig. 5 C). Slender bulging areoles are abundant over the entire body (Fig. 4 C). They occur as single areoles or in small groups. Clusters of crowned areoles and circumcluster areoles are present over the entire cuticle (Figs. 4 A, 4 B, 4 C). Crowned areoles occur in pairs, and in one case a larger cluster was observed which may be due to a close proximity of two smaller clusters (Fig. 5 D). The apical filaments of the crowned areoles are moderately long and extend slightly over the borders of the entire cluster (Fig. 4 D, 4 E, 4 F). Some filaments are branched. The circumcluster areoles are slender areoles with a tuft of tiny bristles at the tip (Fig. 4 C, 4 D, 4 E, 4 F). Crowned areoles with very long apical filaments are present in two rows next to the ventral midline (Figs. 4 B, 5 A, 5 B). Length of the apical filaments is between 150 and 200 ��m. A different cuticular structure is present in the anterior 200 ��m of the body (Fig. 5 E). In this part the simple areoles carry several hair-like structures (Fig. 5 F). Single hairs are about 5 ��m long and occur over the entire surface of the single areoles, but not in great density. A considerable number of tubercle areoles is present, the areolar base of these is similar to that of the simple areoles (Fig. 5 F). About 200 ��m posterior to the anterior tip, the cuticular pattern as described in the previous paragraph starts. The posterior end cannot be described here in detail due to shrinkage during preparation for SEM, which did not allow close observation. Taxonomic remarks. This specimen is unique in having many slender bulging areoles, hooked thorn areoles and a distinctly different cuticular pattern at the anterior end. Abundant slender bulging areoles are typical for two neotropical species, Chordodes brasiliensis Janda 1894 from Argentina, Brazil and Uruguay (see De Villalobos et al. 2004) and C. polytuberculatus Schmidt-Rhaesa & Menzel 2005 from Costa Rica (see Schmidt-Rhaesa & Menzel 2005). For ���old world��� species, this feature is new. Thorn areoles were not observed in a number of species. Although they are evident structures, they are usually rare and might have been absent on the piece of cuticle investigated. Our observations suggest that they might also be distributed unequally along the body. In C. lasuboni n. sp. , thorn areoles seemed to be present only at the anterior end or at least to be rare in the mid-body region and posterior end. The slender and distinctively hook-shaped form is also a peculiar feature of the species, as thorn areoles are usually solid and slightly curved (see, e.g., Schmidt-Rhaesa et al. 2008). The different cuticular structure at the anterior end has not been described for other species, but this may be because the structure of the anterior end is rarely included in descriptions., Published as part of Schmidt-Rhaesa, Andreas & Yadav, Arun K., 2013, One new species and a new record of the genus Chordodes (Nematomorpha: Gordiida) from North-East India, pp. 293-300 in Zootaxa 3693 (2) on pages 297-299, DOI: 10.11646/zootaxa.3693.2.10, http://zenodo.org/record/218970

Schmidt-Rhaesa, Andreas, Yadav, Arun K. (2013): One new species and a new record of the genus Chordodes (Nematomorpha: Gordiida) from North-East India. Zootaxa 3693 (2): 293-300, DOI: http://dx.doi.org/10.11646/zootaxa.3693.2.10