Search

The South Asian ticks Hyalomma (Hyalommina) brevipunctata Sharif, 1928, Hyalomma (Hyalommina) hussaini Sharif, 1928 and Hyalomma (Hyalommina) kumari Sharif, 1928, are morphologically fairly similar. The males can be distinguished from each other by size, color and their punctation patterns. The females are easily distinguished by the shape of the genital aperture and its preatrial fold, and they also differ in size, color and punctation patterns. The immature stages can be distinguished by the measurements of various characteristics and the ratios of these, the shape of the scutum, gnathosoma and hypostome, number of denticles on the hypostome and size of the spurs on coxae. All parasitic stages of the three species are illustrated and redescribed here. Their distributions are restricted to South Asia, although there are some records from Central Asia. Large and medium-sized ungulates are the principal hosts of the adults, whereas the immature stages infest various species of small mammals. All are three-host species. Data on their disease relationships are also provided.

Kluge, Nikita J., Godunko, Roman J., Apanaskevich, Dmitry A. (2013): Mayflies of the Caucasus Mountains. II. Description of the first representative of the subgenus Helvetoraeticus Bauernfeind & Soldán, 2012 (Heptageniidae: Ecdyonurus). Zootaxa 3608 (1): 51-66, DOI: http://dx.doi.org/10.11646/zootaxa.3608.1.3

Ecdyonurus (Helvetoraeticus) adjaricus sp. nov. Figures 1���41 = Ecdyonurus /fg 2 sp. H 5: Kluge 2004: 176. Types. HOLOTYPE: male imago (dried pinned specimen; right fore and hind wings, forelegs, parts of middle and hind legs, and cerci on slide), GEORGIA, Autonomous Republic of Adjara, Kobuleti District, Kintrishi State Nature Reserve (KSNR), meadow near Zeraboseli farm, middle part of Kintrishi River, approximately 41 �� 43,592 ���N 41 �� 49,096 ���E, 480 m a.s.l., 28.vi. 1974, leg. A. Zaguliaev. PARATYPES: female imago (reared from larva, with larval and subimaginal skins preserved), same locality as holotype, 27���31.vii. 1997, leg. D. Apanaskevich; 3 mature larvae (1 female, 2 males), ibid, 27���31.vii. 1997, leg. D. Apanaskevich; 4 larvae, Autonomous Republic of Adjara, Kobuleti District, 3 km downstream of Zeraboseli farm, border of KSNR near bridge, 41 �� 45,739 ���N 41 �� 58,582 ���E, 325 m a.s.l., 18.viii. 2009, leg. R.J. Godunko, T. Sold��n and P. Sroka; 1 larva, Kobuleti District, Chakhati village (centre), Kintrishi River, 41 �� 48,120 ���N 41 �� 55,545 ���E, 140 m a.s.l., 18.viii. 2009, leg. R.J. Godunko, T. Sold��n and P. Sroka; 1 larva, Kobuleti District, Khala village, Kemerkobis stream (bridge 100 m upstream from mouth), 41 �� 42,226 ���N 41 �� 47,525 ���E, 67 m a.s.l., 18.viii. 2009, leg. P. Sroka; 3 larvae, Kobuleti District, Khala village, Chakvistskhali River (mouth of Kemerkobis stream), 41 �� 42,226 ���N 41 �� 47,455 ���E, 60 m a.s.l., 19.viii. 2009, leg. R.J. Godunko; 14 larvae (6 of them were mounted on slides), Kobuleti District, 3 km upstream at Chakhati village, Kintrishi River, 41 �� 46,266 ���N 41 �� 58,325 ���E, 275, m a.s.l., 31.vii. 2010, leg. R.J. Godunko and N.O. Godunko; 5 larvae, Autonomous Republic of Adjara, Khelvachauri District, Tskhemlara [Tamara���] bridge near Chikuneti village, Machakhlistskali River, 41 �� 30,424 ���N 41 �� 49,215 ���E, 195 m a.s.l., 20.viii. 2009, leg. P. Sroka; 11 larvae, Khelvachauri District, 200 m downstream at Tskhemlara [Tamara���] bridge near Chikuneti village, small stream (left side tributary of Machakhlistskali River), 41 �� 30,345 ��� N 41 49 ��02,42���E, 185 m a.s.l., 20.viii. 2009, leg. R.J. Godunko and P. Sroka. Male imago (dried pinned specimen). Size: body length: 15.5 mm; forewings length: 17.0 mm; cerci lost. General body color distinctly dark, reddish-brown; few whitish spots on lateral sides of mesonotum. Head dirty brown to reddish-brown. Antennae blackish-brown, slightly paler distally. Ocelli dirty yellow apically. Eyes grayish, contiguous dorsally, with relatively narrow dark band basally. Thorax dark brown to reddish-brown, slightly paler ventrally. Forewings hyaline with yellowish area in basal part of wing, in anal and cubital fields, and between C and RA; C and Sc dark brown, other veins brown; pterostigmatic area opaque, dirty yellow, with 6 simple and 7���8 branched veins. Hind wings entirely yellowish; venation brown, darker distally. Forelegs distinctly darker than middle and hind ones, brown to reddish-brown; forefemora only slightly darker than tibiae and tarsi. Middle and hind legs brown; tarsi slightly darker than femora and tibiae. No maculation on femora surface. Abdomen dark brown to reddish-brown, sterna slightly paler. Lateral sides of terga II ���VIII with roundish or triangular spots surrounded with an indistinct light L -shape marking typical for Helvetoraeticus species. Cerci dark brown. Penis lobes yellowish-brown to brown. In dorsal and ventral view outline of penis lobes broadly rounded, not expanded laterally; lobes contiguous nearly over entire length (Fig. 1). Apical sclerite relatively massive and short, slightly curved, only slightly expanded laterally and bluntly pointed at apex; lack of conspicuous denticulation at medio-internal portion; several small spines at external margin near tip of titillators; apex of sclerite not projecting above the lobes; only a slight projection at inner border. Lateral sclerite not broad, relatively straight, with subparallel anterior and posterior margins. Basal sclerite relatively massive, with 2���3 small spines only, without conspicuous teeth. Penis stem without a hump. Titillators brown. Styliger intensively brown, with two distinct, slightly curved protuberances near gonostyli bases; gonostyli slightly darker; segment II without any hump at inner margin basally (Fig. 2). Female imago. Size: body length: 15.0 mm; forewings length: 20.0 mm; cerci length: 13.0 mm. General color of body similar to that in male imago, slightly paler; thoraxal pleurae with whitish blotches. Eyes with relatively narrow dark band basally. Fore and hind wings hyaline and transparent, uncolored; venation brown, darker distally. Legs light brown to brown, forelegs slightly paler. Abdominal pleurae with an indistinct light L - shape pattern. Cerci brown. Subgenital plate relatively wide; posterior margin slightly rounded. Subanal plate bluntly pointed apically. Female subimago. Measurements and body color similar to those in female imago; thorax and abdomen slightly paler. Forewings relatively dark, with well visible pattern formed by transversal grayish bands (zigzag type). Male subimago. Unknown. Mature larva. Size: body length: 12.0���16.0 mm; cerci length: 9.0��� 12.5 mm. General body color yellowish-olive, brownish-olive to brown. Head yellowish-olive to yellowish-brown with apparent light pattern, consisting of three blotches near fore margin of ocelli, and pair of spots near antennal bases; two small blotches near anterior margin of head; further large diffuse light spots situated laterally along posterior margin of head (Figs. 3, 5, 6, 8). Head capsule rounded anteriorly. Antennae with brown scape and pedicel; flagellum yellowish-white. Eyes and ocelli black. Labrum relatively narrow with slightly curved and short lateral lobes; ratio of total labrum width to its width at base 2.3 / 1; 8���11 relatively slender bristles arranged in a single row on ventral side of labrum (Fig. 17), diverging laterally into two irregular rows. Maxillae (n = 12): number of comb-shaped bristles (N_CBS) = 24���27 (mainly 26���27); number of teeth on 5 th comb-shaped bristle (N_TCB 5) = 11���14 (mainly 11���12); number of setae on inner side of first segment of maxillary palps (N_PLP)> 10; number of setae on outer margin of first segment of maxillary palps (N_PLS) = 22���38; number of hairs on dorsal upper side of galea-lacinia (N_DOR)> 15. Glossae stretched laterally, relatively widely spaced; glossae and paraglossae massive (Fig. 16). Superlinguae with pilosity typical for Helvetoraeticus (only few setae at tip). Pronotum yellowish-olive to light brown with several light spots centrally and laterally; lateral projections yellow to yellowish-white (Figs. 5, 6). Pronotum slightly extended laterally. Lateral projection large and long, relatively straight, only slightly asymmetrical, oriented parallel to body axis (Figs. 3, 5, 6, 8, 13, 14), or slightly outwards (Fig. 32); apex of lateral projection bluntly pointed; width/length ratio of semipronotum to caudal portion of projection is 1.95���2.30; length of caudal portion of projection is approximately half of total projection length. Meso- and metanotum yellowish-olive to light brown; mesonotum with longitudinal light lines and spots laterally from median longitudinal suture, interrupted by pale areas near fore and hind margins. Legs yellowish-olive and light brown to brown. Trochanters with 2 (rarely 3) irregular rows of bluntly pointed and slender spatulate bristles (Figs. 24, 25). Femora long and relatively slender; length/width ratio of metafemora is 2.75���3.35 (mean 3.0, �� = 0.21; n = 8); widest part of metafemur at its midlength. Dorsally femora with large yellowish-olive to light brown spots on yellowish to yellowish-white background; uniformly yellowish-white to whitish ventrally (general color pattern of dorsal side of femora as illustrated by Godunko & Kłonowska-Olejnik 2004: 17, fig. 10; 2008: 812, fig. 33). Dense rows of sharply pointed bristles at inner margin of femora (Fig. 19); outer margin of femur with row of long whip-like setae and dense rows of sharply pointed bristles (Figs. 18, 26, 27); same type of sharply pointed bristles at dorsal surface of femora (centrally and distally), occasionally more blunt in proximal part (Figs. 20, 21, 26, 27); a row of short pointed bristles on ventral side of posterior border of femora (Fig. 27). Tibiae yellowish-brown, with well visible darker smudge in centrally; dorsal surface with row of spatulate bristles (Figs. 28, 29). Tarsi brownish, with paler smudge centrally, indistinctly darker apically; sparse row of slender spatulate bristles at dorsal surface (Fig. 30). Claw brown with 2 denticles (Fig. 31). Abdominal terga with pronounced yellowish-brown to dark brown pattern and yellowish background (Figs. 3, 4, 7, 8): tergum I relatively pale, with broad diffuse spot medially and a pair of spots laterally; terga II and III with a pair of spots medially, and a pair of spots laterally; terga IV and V with V- (or U-) shaped spot medially, and a pair of broad spots laterally (occasionally lateral spots fused with central spot at tergum V); tergum VI with U- (or V-) shaped spot medially (consisting of three more or less separated small spots); terga VII and VIII with broad Vshaped spot medially, turning into light strip stretching along posterior margin and lateral sides of segment; tergum IX with oval median spot, occasionally a pair of small spots laterally; tergum X generally unicolorous brown, occasionally with a pair of unclear longitudinal paler strokes medially. Abdominal sterna yellowish-white, distinct neural ganglion in segment VII (occasionally also in segments IV ���VI); brown pattern of sterna mainly absent at segments I ���V (occasionally VI); sterna VII ���IX with pair of triangular brown spots laterally (in male) and with transversal brown stripes (in female); larval protogonostyli darker than abdominal sterna, light brown (Figs. 9���12). Posterior margin of terga with dense row of stout spines pointed apically, alternating with smaller ones (Figs. 15, 22); surface of terga with numerous hairs. Posterolateral extensions of abdominal sterna well developed, distinctly long, sharpened apically and directed outwards at different angles to the body axis (except for expansions of sterna 7���8 directed parallely to the body axis) (Figs. 9���12). Tergalii whitish to yellowish-gray, with distinct dark tracheae; tergalius I tongue-shaped, relatively long, moderately tapered distally, with plate longer than filamentous part (Fig. 33); tergalius IV relatively slender, moderately asymmetrical, only with slightly conspicuously convex outer margin, distal margin not blunt, length/width ratio 2.25���2.60 (mean 2.41, �� = 0.19; n = 8) (Fig. 34); tergalius VII broad and asymmetrical (widest part in 2 / 3 of tergalius length), without a tuft of tracheal filaments (Fig. 35). Caudalii yellowish-brown, paler distally; each segment with sharply pointed bristles alternating with hairs (Fig. 23). Egg. Measurements: length 180���200 ��m; width 130���155 ��m. Egg oval, slightly elongated (Figs. 36, 37). Chorion covered with attachment structures ��� knob-terminated coiled threads (KCTs), numerous small rounded tubercles and delicate granules (granular ground matrix) (Figs. 39, 40). Significant concentration of KCTs attachment structures (diameter 3.6���4.2 ��m) at one egg pole, where distance between them is 0.3���2.4 ��m (Figs. 38, 39); other egg surface with rather sparsely KCTs attachment structures (distance between them 2.0��� 12.5 ��m) (Fig. 40). Small rounded tubercles (1.4 ���2.0 ��m in diameter) are regularly distributed over whole chorion (0.1 ���3.0 ��m distance between them) (Figs. 39, 40). Two to three micropyles (13.3 ���15.0 ��m in length; 8.5���10.2 ��m in width) are visible in subequatorial region; micropylar rim thin, with a few sparsely distributed tubercles (Fig. 41). Etymology. The species is named after Adjara (autonomous republic within Georgia). Affinities. Ecdyonurus adjaricus sp. nov. holds a clearly isolated position within Helvetoraeticus. The shape of the male imago penis lobes is typical for the subgenus; at the same time apical sclerites are expanded only slightly laterally (in contrast to other Helvetoraeticus species with markedly laterally stretched apical sclerites). The larva of E. adjaricus sp. nov. can be recognized within the subgenus by the presence a well developed, relatively long posterolateral expansions of abdominal sterna; such size and shape of expansions is characteristic only for a few species of Ecdyonurus s. str. (Haybach 1999, figs. 2, 11). Additionally, several other characters of the larval mouthparts should be noted, e.g.: the single row of bristles on ventral side of labrum; a relatively small number of setae on the inner side of the first segment of the maxillary palps and the number of comb-shaped bristles on the maxilla (see Haybach 1999: 121, 137, 139). The placement of E. adjaricus sp. nov. within the subgenus Helvetoraeticus can be confirmed by the absence of long and dense setae on the distal portions of the superlinguae of the hypopharynx of larva, and the general shape and structure of male imago genitalia. The new species is very similar to E. krueperi (Stein, 1863) in both the adult and larval stages. These two species can be easily separated from other representatives of Helvetoraeticus by conspicuous wing coloration, especially that of the hind wings: yellowish in E. adjaricus sp. nov.; golden-brown in E. krueperi. At the same time, the hind wings of the new species are colored only in male imago, while they are colored in both sexes of E. krueperi (Puthz 1980: 349; Hefti & Tomka 1988: 332; Bauernfeind & Sold��n 2012: 277). Other differences in male imagoes are the following: (1) penis lobes are broadly rounded and not expanded laterally, with apically touching lobes in E. adjaricus sp. nov. (outline of penis lobes are distinctly triangularly rounded, more or less separated apically in E. krueperi; cf. Puthz 1980: 350, fig. 2; Hefti & Tomka 1988: 331���332, fig. 3; Hefti et al. 1989: 334, 338, fig. 10 d; Bauernfeind & Sold��n 2012: 277, 589, fig. 163); (2) apical sclerite only slightly curved, without conspicuous denticulation at medio-internal portion in E. adjaricus sp. nov. (apical sclerite strongly curved, with conspicuous denticulation medio-internally in E. krueperi; cf. Puthz 1980: 350, figs. 4, 5; Bauernfeind & Sold��n 2012: 277); (3) lateral sclerite not broad, relatively straight, with subparallel anterior and posterior margins, evenly sclerotized throughout its length in E. adjaricus sp. nov. (lateral sclerite almost square distally, broad, visibly sclerotized only in its central and distal part in E. krueperi; cf. Bauernfeind & Sold��n 2012: 277; for this reason some inconsistencies in the descriptions and figure were published by Hefti & Tomka 1988: 331���332, fig. 3 and Hefti et al. 1989: 334). Larvae of E. adjaricus sp. nov. and E. krueperi possess a similar shape of the lateral projection of the pronotum (relatively straight, only slightly asymmetrical, bluntly pointed at the tip, oriented almost parallel to body axis), and similar shape of femoral bristles (sharply pointed in both species). Larva of E. adjaricus sp. nov. can be separated from E. krueperi by the following features: (i) row of bristles on ventral side of labrum medially singular (in contrast to double in E. krueperi); (ii) glossae stretched laterally and relatively widely spaced (in contrast to elongated-quadrangular glossae in E. krueperi; cf. Hefti & Tomka 1988: 330, fig. 1 b; Hefti et al. 1989: 333, 335 [Table 1], fig. 6 b); (iii) pointed bristles on ventral side of posterior border of femora (lack of this type of bristles in E. krueperi; cf. Hefti et al. 1989: 333, 335 [Table 1]); (iv) posterolateral extensions of abdominal sterna well developed and distinctly long (in contrast to relatively short sternal expansions of E. krueperi, typical for all other Helvetoraeticus representatives); (v) shape of tergalius I (tongue-shaped, moderately tapered distally) and tergalius IV (relatively slender, moderately asymmetrical) (in contrast to broadly-shaped tergalius I and distinctly asymmetrical tergalius IV in E. krueperi; cf. Jacob & Braasch 1984: 60, fig. 27; Hefti et al. 1989, 335 [Table 1]). Additional differences between E. adjaricus sp. nov. and E. krueperi are found in the color of thoracic and abdominal segments of adults and larvae, i.e. the new species is generally more darkly colored. Only a few species of Helvetoraeticus (together with E. adjaricus sp. nov.) are characterized by a significant concentration of KCTs attachment structures on one egg pole, namely E. epeorides Demoulin, 1955, E. krueperi, E. picteti (Meyer-D��r, 1864), E. silvaegabretae Sold��n & Godunko, 2006 and E. siveci Jacob & Braasch, 1984 (see summarized data in Bauernfeind & Sold��n 2012). We are confident that the male imago (holotype, not reared from larva) and larvae (as well as reared female imago) described here belong to the same species. This is strongly supported by the fact that all material was collected in the same region (distance between the outermost studied localities is 30���35 km; 2���5 km between localities where holotype and paratypes were collected) and during the same summer period. The male imago body color is similar to that of the female. However the hind wings are colored in male and colorless in female. Distribution and biology. The known distribution of E. adjaricus sp. nov. is restricted to several rivers of south-western spurs of Meskheti [Adjar-Imereti] Range and Shavsheti Range within Georgia. In Meskheti Range all investigated rivers are located on northern slopes; Kintrishi and Chakvistskhali rivers flow directly into the Black Sea. The localities within the Machakhlistskali river-basin (a part of Chorokh [��oruh] river-basin) are situated on south slopes near the Turkey border. All habitats are located in the lowest parts of this area within altitudes 60���480 m a.s.l. There is a high probability that the new species can be found in other tributaries of Chorokh [��oruh] river-basin within Turkey (at least in rivers of Turkish part of Shavsheti Range, as well as rivers within Ka��kar Mountains [Ka��kar Daġları]). Information about distribution of Helvetoraeticus in Turkey have been earlier published by Kazancı (2001: 29, 50) and Tanatmış (2004: 235), as records of Ecdyonurus helveticus Eaton, 1883 from, Published as part of Kluge, Nikita J., Godunko, Roman J. & Apanaskevich, Dmitry A., 2013, Mayflies of the Caucasus Mountains. II. Description of the first representative of the subgenus Helvetoraeticus Bauernfeind & Sold��n, 2012 (Heptageniidae: Ecdyonurus), pp. 51-66 in Zootaxa 3608 (1) on pages 52-65, DOI: 10.11646/zootaxa.3608.1.3, http://zenodo.org/record/283261

The first collection of unengorged and fully engorged larvae of Nuttalliella sp. (N. namaqua?) from the murid rodents Micaelamys namaquensis, Aethomys chrysophilus and Acomys spinosissimus in Limpopo Province and from M. namaquensis in the Northern Cape Province, South Africa, is documented. A total of nine larvae were collected from two M. namaquensis in the Soutpansberg mountain range in the Limpopo Province during April 2009. During the last week of September 2011, 221 larvae were collected from rodents at the same locality and 10 of 48 M. namaquensis, 6 of 12 Ae. chrysophilus and 3 of 14 Ac. spinosissimus were infested. One of the M. namaquensis harboured 53 larvae. Five larvae were collected from two M. namaquensis in the Northern Cape Province. Total genomic DNA was extracted from two larvae and a region of the 18S rRNA gene was sequenced for these. BLASTn searches revealed similarity between these specimens and the Nuttalliella sequences published on GenBank.

Ixodidae We exclude the following names from our list of Ixodidae: (i) Amblyomma laticaudae Warburton, 1933 is considered valid in Camicas et al. (1998), Horak et al. (2002) and Barker and Murrell (2008); Guglielmone et al. (2009) do not include it in their list of controversial names, while Kolonin (2009) considers this name a synonym of Amblyomma nitidum Hirst and Hirst, 1910. In fact, Keirans, in Voltzit and Keirans (2002), determined that the types of these taxa are conspecific. Therefore, we agree with Kolonin (2009) that A. laticaudae is an invalid name. (ii) Bothriocroton decorosum (Koch, 1867) is included as a member of Aponomma by Kolonin (2009), but this name is a synonym of B. undatum, as explained in Santos Dias (1993). (iii) Haemaphysalis vietnamensis Hoogstraal and Wilson, 1966 is included as a valid name in Kolonin (2009), but it is a synonym of H. colasbelcouri (Guglielmone et al. 2009). (iv) Haemaphysalis xinjiangensis Teng, 1980 is not included as a valid species in Kolonin (2009). Teng and Jiang (1981) consider this name a synonym of H. danieli, and we accept their decision, since the senior author also described H. xinjiangensis. (v) Hyalomma erythraeum Tonelli-Rondelli, 1932 is included as a valid name in Kolonin (2009), but it was convincingly shown to be a synonym of H. impeltatum by Apanaskevich and Horak (2009). (vi) Rhipicephalus hoogstraali Kolonin, 2009 was described on the Internet, contrary to the rules of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 2009, Articles 8.6 and 9.8), and is therefore a nomen nudum. 1. Amblyomma albolimbatum Neumann, 1907 2. Amblyomma albopictum Neumann, 1899 3. Amblyomma americanum (Linnaeus, 1758) 4. Amblyomma antillorum Kohls, 1969 5. Amblyomma arcanum Karsch, 1879 (Aponomma) 6. Amblyomma argentinae Neumann, 1905. Guglielmone et al. (2003) stated that the year of description for A. argentinae is 1904, but this is in error because Neumann recognized 1905 as the correct year. Previously, this species was incorrectly named A. testudinis (Guglielmone et al. 2009). 7. Amblyomma astrion Dönitz, 1909 8. Amblyomma aureolatum (Pallas, 1772) 9. Amblyomma auricularium (Conil, 1878) 10. Amblyomma australiense Neumann, 1905. See A. echidnae. 11. Amblyomma babirussae Schulze, 1933 12. Amblyomma beaurepairei Vogelsang and Santos Dias, 1953. This species is close to A. inornatum and A. auricularium but is provisionally considered valid in Guglielmone et al. (2009). However, Kolonin (2009) considers it a synonym of A. auricularium. 13. Amblyomma boeroi Nava, Mangold, Mastropaolo, Venzal, Oscherov and Guglielmone, 2009 14. Amblyomma boulengeri Hirst and Hirst, 1910 15. Amblyomma brasiliense Aragão, 1908 16. Amblyomma breviscutatum Neumann, 1899. Until recently, this species was known as A. cyprium Neumann, 1899 (Guglielmone et al. 2009), a name still considered valid in Kolonin (2009), who does not justify his position. However, Keirans, in Voltzit and Keirans (2002), determined, as had Santos Dias (1956) before him, that the types of A. cyprium and A. breviscutatum are conspecific and that the latter name has page priority. 17. Amblyomma cajennense (Fabricius, 1787). This is the type species of the genus Amblyomma; it was originally named Acarus cajennensis. Beati, L. and Nava, S. (pers. comm.) believe that A. cajennense comprises a complex of sibling species. 18. Amblyomma calabyi Roberts, 1963 19. Amblyomma calcaratum Neumann, 1899 20. Amblyomma chabaudi Rageau, 1964 21. Amblyomma clypeolatum Neumann, 1899 22. Amblyomma coelebs Neumann, 1899 23. Amblyomma cohaerens Dönitz, 1909 24. Amblyomma compressum (Macalister, 1872) 25. Amblyomma cordiferum Neumann, 1899 26. Amblyomma crassipes (Neumann, 1901) (Aponomma). This species is similar to A. fuscolineatum but is provisionally considered valid because little material is available for examination and the types have not been compared (Guglielmone et al. 2009). Kolonin (2009), using the name Aponomma, considers A. crassipes invalid but does not include a comparison of types. 27. Amblyomma crassum Robinson, 1926 28. Amblyomma crenatum Neumann, 1899 29. Amblyomma cruciferum Neumann, 1901 30. Amblyomma darwini Hirst and Hirst, 1910 31. Amblyomma dissimile Koch, 1844. Guglielmone et al. (2009) argue in favor of this name instead of A. bibroni (Gervais, 1842) because of the confused history of the latter taxon. 32. Amblyomma dubitatum Neumann, 1899. Until recently, this species was known as A. cooperi Nuttall and Warburton, 1908 (Guglielmone et al. 2009). 33. Amblyomma eburneum Gerstäcker, 1873 34. Amblyomma echidnae Roberts, 1953. This species is very similar to A. australiense, but Guglielmone et al. (2009) consider A. echidnae provisionally valid until its status can be clarified. Kolonin (2009) does not include this species in his list of the Ixodidae of the world. 35. Amblyomma elaphense (Price, 1959) (Aponomma). See A. sphenodonti. 36. Amblyomma exornatum Koch, 1844 (Aponomma) 37. Amblyomma extraoculatum Neumann, 1899. See A. romitii. 38. Amblyomma falsomarmoreum Tonelli-Rondelli, 1935 39. Amblyomma fimbriatum Koch, 1844 (Aponomma) 40. Amblyomma flavomaculatum (Lucas, 1846) (Aponomma). See A. inopinatum. 41. Amblyomma fulvum Neumann, 1899 42. Amblyomma fuscolineatum (Lucas, 1847) (Aponomma). See A. crassipes and A. varanense. 43. Amblyomma fuscum Neumann, 1907. A valid taxon ignored in some lists of ticks (Guglielmone et al. 2009). 44. Amblyomma geayi Neumann, 1899. Guglielmone et al. (2009) discuss the lack of agreement concerning this name, because some authors regard A. geayi as a synonym of A. perpunctatum (Packard, 1869). However, the description of the latter taxon is inadequate, and we therefore consider the name A. geayi valid, pending type comparison. 45. Amblyomma gemma Dönitz, 1909 46. Amblyomma geochelone Durden, Keirans and Smith, 2002 47. Amblyomma geoemydae (Cantor, 1847) 48. Amblyomma gervaisi (Lucas, 1847) (Aponomma). This is the type species of the genus Aponomma; it was originally named Ixodes gervaisii. 49. Amblyomma glauerti Keirans, King and Sharrad, 1994 50. Amblyomma goeldii Neumann, 1899 51. Amblyomma hainanense Teng, 1981. Kolonin (2009) considers A. hainanense a synonym of A. helvolum, but no comparison of types has been attempted. Therefore, we treat A. hainanense as tentatively valid. 52. Amblyomma hebraeum Koch, 1844 53. Amblyomma helvolum Koch, 1844 54. Amblyomma hirtum Neumann, 1906 55. Amblyomma humerale Koch, 1844 56. Amblyomma imitator Kohls, 1958 57. Amblyomma incisum Neumann, 1906. See A. latepunctatum. 58. Amblyomma inopinatum (Santos Dias, 1989) (Aponomma). This species is not included in Kolonin (2009), who appears to consider it a synonym of A. flavomaculatum and places it in the genus Aponomma. 59. Amblyomma inornatum (Banks, 1909) 60. Amblyomma integrum Karsch, 1879 61. Amblyomma javanense (Supino, 1897) 62. Amblyomma komodoense (Oudemans, 1928) (Aponomma) 63. Amblyomma kraneveldi (Anastos, 1956) (Aponomma) 64. Amblyomma latepunctatum Tonelli-Rondelli, 1939. This species was previously confused with A. incisum and with A. scalpturatum (Guglielmone et al. 2009). 65. Amblyomma latum Koch, 1844 (Aponomma) 66. Amblyomma lepidum Dönitz, 1909 67. Amblyomma limbatum Neumann, 1899 68. Amblyomma loculosum Neumann, 1907 69. Amblyomma longirostre (Koch, 1844) 70. Amblyomma macfarlandi Keirans, Hoogstraal and Clifford, 1973 71. Amblyomma macropi Roberts, 1953 72. Amblyomma maculatum Koch, 1844 73. Amblyomma marmoreum Koch, 1844 74. Amblyomma moreliae (Koch, 1867) 75. Amblyomma moyi Roberts, 1953 76. Amblyomma multipunctum Neumann, 1899 77. Amblyomma naponense (Packard, 1869) 78. Amblyomma neumanni Ribaga, 1902. Nava et al. (2009 a) state that the description by Voltzit (2007) of A. neumanni actually applies to A. parvitarsum. 79. Amblyomma nitidum Hirst and Hirst, 1910. See paragraph “i” above our list of Ixodidae. 80. Amblyomma nodosum Neumann, 1899 81. Amblyomma nuttalli Dönitz, 1909 82. Amblyomma oblongoguttatum Koch, 1844 83. Amblyomma orlovi (Kolonin, 1992) (Aponomma). This species is considered tentatively valid in Guglielmone et al. (2009), who nonetheless strongly suggest that it is a synonym of A. transversale. Guglielmone et al. (2009) err in listing the date of description as 1995. 84. Amblyomma ovale Koch, 1844 85. Amblyomma pacae Aragão, 1911 86. Amblyomma papuanum Hirst, 1914 87. Amblyomma parkeri Fonseca and Aragão, 1952. Kolonin (2009) expresses doubt about the status of this species, but Labruna et al. (2009 a) present sound evidence for its validity. 88. Amblyomma parvitarsum Neumann, 1901. See A. neumanni. 89. Amblyomma parvum Aragão, 1908 90. Amblyomma pattoni (Neumann, 1910) (Aponomma) 91. Amblyomma paulopunctatum Neumann, 1899 92. Amblyomma pecarium Dunn, 1933 93. Amblyomma personatum Neumann, 1901 94. Amblyomma pictum Neumann, 1906 95. Amblyomma pilosum Neumann, 1899 96. Amblyomma pomposum Dönitz, 1909 97. Amblyomma postoculatum Neumann, 1899 98. Amblyomma pseudoconcolor Aragão, 1908 99. Amblyomma pseudoparvum Guglielmone, Mangold and Keirans, 1990 100. Amblyomma quadricavum (Schulze, 1941) 101. Amblyomma rhinocerotis (de Geer, 1778) 102. Amblyomma robinsoni Warburton, 1927 103. Amblyomma romitii Tonelli-Rondelli, 1939. This species was previously considered a synonym of A. extraoculatum (Guglielmone et al. 2009). 104. Amblyomma rotundatum Koch, 1844 105. Amblyomma sabanerae Stoll, 1894 106. Amblyomma scalpturatum Neumann, 1906. See A. latepunctatum. 107. Amblyomma scutatum Neumann, 1899 108. Amblyomma soembawense (Anastos, 1956) (Aponomma) 109. Amblyomma sparsum Neumann, 1899 110. Amblyomma sphenodonti (Dumbleton, 1943) (Aponomma). Miller et al. (2007) believe that this species, and possibly A. elaphense, should be classified in a genus other than Amblyomma. 111. Amblyomma splendidum Giebel, 1877 112. Amblyomma squamosum Kohls, 1953 113. Amblyomma supinoi Neumann, 1904 114. Amblyomma sylvaticum (de Geer, 1778) 115. Amblyomma tapirellum Dunn, 1933 116. Amblyomma testudinarium Koch, 1844 117. Amblyomma tholloni Neumann, 1899 118. Amblyomma tigrinum Koch, 1844 119. Amblyomma torrei Pérez Vigueras, 1934 120. Amblyomma transversale (Lucas, 1845) (Aponomma). Most authors consider 1844 the year of description of A. transversale, but the name Ixodes transversalis in Lucas (1844) lacks a description (i.e., it is a nomen nudum), and this situation was amended in Lucas (1845). See A. orlovi. 121. Amblyomma triguttatum Koch, 1844 122. Amblyomma trimaculatum (Lucas, 1878) (Aponomma) 123. Amblyomma triste Koch, 1844 124. Amblyomma tuberculatum Marx, 1894 125. Amblyomma usingeri Keirans, Hoogstraal and Clifford, 1973 126. Amblyomma varanense (Supino, 1897) (Aponomma). Kolonin (2009) considers A. varanense a synonym of A. fuscolineatum (both are treated as species of Aponomma) but without presenting evidence. We believe A. varanense to be valid. 127. Amblyomma variegatum (Fabricius, 1794) 128. Amblyomma varium Koch, 1844 129. Amblyomma vikirri Keirans, Bull and Duffield, 1996 130. Amblyomma williamsi Banks, 1924 1. Anomalohimalaya cricetuli Teng and Huang, 1981. Deng et al. (1999) consider this species a synonym of A. lotozkyi but without explanation. We therefore maintain the validity of A. cricetuli. 2. Anomalohimalaya lamai Hoogstraal, Kaiser and Mitchell, 1970. This is the type species of the genus Anomalohimalaya. 3. Anomalohimalaya lotozkyi Filippova and Panova, 1978. See A. cricetuli. 1. Bothriocroton auruginans (Schulze, 1936) (Aponomma) 2. Bothriocroton concolor (Neumann, 1899) (Aponomma). See B. oudemansi. 3. Bothriocroton glebopalma (Keirans, King and Sharrad, 1994) (Aponomma). This is the type species of the genus Bothriocroton, by original designation (Klompen et al. 2002); it was originally named Aponomma glebopalma. 4. Bothriocroton hydrosauri (Denny, 1843) (Aponomma). See B. tachyglossi. 5. Bothriocroton oudemansi (Neumann, 1910) (Aponomma). This species was considered a synonym of B. concolor, but there is now sound scientific evidence for its validity as a species (Guglielmone et al. 2009). 6. Bothriocroton tachyglossi (Roberts, 1953) (Aponomma). This species was considered a synonym of B. hydrosauri, but there is now convincing evidence for its validity (Guglielmone et al. 2009). 7. Bothriocroton undatum (Fabricius, 1775) (Aponomma). This name is considered invalid by Kolonin (2009). See B. decorosum, paragraph “ii” above our list of Ixodidae. 1. Compluriscutula vetulum Poinar and Buckley, 2008. This is the type species of the fossil genus Compluriscutula. It is not included in the list of Kolonin (2009). 1. Cornupalpatum burmanicum Poinar and Brown, 2003. This is the type species of the fossil genus Cornupalpatum. It is not included in the list of Kolonin (2009). 2. Cosmiomma hippopotamensis (Denny, 1843). This is the type species of the monotypic genus Cosmiomma. It was originally named Ixodes hippopotamensis. 1. Dermacentor abaensis Teng, 1963. This species is similar to D. everestianus but is provisionally considered valid in Guglielmone et al. (2009) because there has been no definitive study to support this synonymy. One of us (DAA) is currently comparing the types of D. abaensis, D. everestianus and D. birulai Olenev, 1927. Therefore, additional information on the specific status of these taxa should be forthcoming. 2. Dermacentor albipictus (Packard, 1869) 3. Dermacentor andersoni Stiles, 1908 4. Dermacentor asper Arthur, 1960. Kolonin (2009) considers D. asper a probable synonym of D. sinicus but presents no evidence for this statement. 5. Dermacentor atrosignatus Neumann, 1906 6. Dermacentor auratus Supino, 1897 7. Dermacentor circumguttatus Neumann, 1897 8. Dermacentor compactus Neumann, 1901 9. Dermacentor confragus (Schulze, 1933). Originally spelled confractus, the name was later corrected to confragus. The type specimens are currently being examined by one of us (TNP). We consider this name valid until studies of the type material permit a conclusive statement. Kolonin (2009) excludes this name from his list of ixodid ticks of the world. 10. Dermacentor dispar Cooley, 1937 11. Dermacentor dissimilis, Cooley 1947 12. Dermacentor everestianus Hirst, 1926. See D. abaensis. 13. Dermacentor halli McIntosh, 1931 14. Dermacentor hunteri Bishopp, 1912 15. Dermacentor imitans Warburton, 1933 16. Dermacentor latus Cooley, 1937 17. Dermacentor marginatus (Sulzer, 1776). See D. niveus and D. ushakovae. 18. Dermacentor montanus Filippova and Panova, 1974 19. Dermacentor nitens Neumann, 1897 (Anocentor). This is the type species of the genus Anocentor. 20. Dermacentor niveus Neumann, 1897. Kolonin (2009) lists this species as a synonym of D. daghestanicus Olenev, 1928, but, in fact, the latter name is a junior synonym of D. niveus (Guglielmone et al. 2009). Estrada-Peña and Estrada-Peña (1991) examined part of the syntype series of D. niveus (Cherestanek, Iran, ex Ovis aries, 1882, loaned by Prof. P.-C. Morel), concluding that this species is conspecific with D. marginatus, a view shared by Moshaverinia et al. (2009). A wider comparative study of D. marginatus, D. niveus and D. ushakovae appears to be needed to further demonstrate the validity of these taxa. We consider D. niveus provisionally valid while awaiting the results of this comparison. See also D. ushakovae. 21. Dermacentor nuttalli Olenev, 1928 22. Dermacentor occidentalis Marx, 1892 23. Dermacentor parumapertus Neumann, 1901 24. Dermacentor pavlovskyi Olenev, 1927 25. Dermacentor pomerantzevi Serdjukova, 1951 26. Dermacentor raskemensis Pomerantzev, 1946 27. Dermacentor reticulatus (Fabricius, 1794). This is the type species of the genus Dermacentor. It was originally named Acarus reticulatus. 28. Dermacentor rhinocerinus (Denny, 1843) 29. Dermacentor silvarum Olenev, 1931 30. Dermacentor sinicus Schulze, 1932. See D. asper. 31. Dermacentor

Argasidae 1. Antricola armasi de la Cruz and Estrada-Peña, 1995 (Carios) 2. Antricola centralis de la Cruz and Estrada-Peña, 1995 (Carios) 3. Antricola cernyi de la Cruz, 1978 (Carios) 4. Antricola coprophilus (McIntosh, 1935) (Carios). This is the type species of the genus Antricola; it was originally named Ornithodoros coprophilus. 5. Antricola delacruzi Estrada-Peña, Barros-Battesti and Venzal, 2004 (Carios) 6. Antricola granasi de la Cruz, 1973 (Carios) 7. Antricola guglielmonei Estrada-Peña, Barros-Battesti and Venzal, 2004 (Carios) 8. Antricola habanensis de la Cruz, 1976 (Carios) 9. Antricola hummelincki de la Cruz and Estrada-Peña, 1995 (Carios) 10. Antricola inexpectata Estrada-Peña, Barros-Battesti and Venzal, 2004 (Carios) 11. Antricola marginatus (Banks, 1910) (Carios, Parantricola). This is the type species of the genus Parantricola; it was originally named Ornithodoros marginatus. 12. Antricola martelorum de la Cruz, 1978 (Carios) 13. Antricola mexicanus Hoffmann, 1958 (Carios). Some authors have listed the year 1958 for the original description of Antricola mexicanus while others have used the year 1959. An inquiry to the U.S. Library of Congress has shown that Hoffmann (1958) was actually published in 1958. 14. Antricola naomiae de la Cruz, 1978 (Carios) 15. Antricola occidentalis de la Cruz, 1978 (Carios) 16. Antricola siboneyi de la Cruz and Estrada-Peña, 1995 (Carios) 17. Antricola silvai Č erný, 1967 (Carios) 1. Argas abdussalami Hoogstraal and McCarthy, 1965 2. Argas africolumbae Hoogstraal, Kaiser, Walker, Ledger, Converse and Rice, 1975 3. Argas arboreus Kaiser, Hoogstraal and Kohls, 1964 4. Argas assimilis Teng and Song, 1983 5. Argas australiensis Kohls and Hoogstraal, 1962 (Carios) 6. Argas beijingensis Teng, 1983 7. Argas beklemischevi Pospelova-Shtrom, Vasil´yeva and Semashko, 1963 8. Argas boueti Roubaud and Colas-Belcour, 1933 (Carios) 9. Argas brevipes Banks, 1908 10. Argas brumpti Neumann, 1907 (Ogadenus). This is the type species of the genus Ogadenus. 11. Argas bureschi Dryenski, 1957 12. Argas ceylonensis Hoogstraal and Kaiser, 1968 (Carios) 13. Argas confusus Hoogstraal, 1955 (Carios) 14. Argas cooleyi Kohls and Hoogstraal, 1960. If Ornithodoros cooleyi McIvor, 1941 belongs to Argas, it has priority here. 15. Argas cordiformis Hoogstraal and Kohls, 1967 (Carios) 16. Argas cucumerinus Neumann, 1901 17. Argas dalei Clifford, Keirans, Hoogstraal and Corwin, 1976 18. Argas daviesi Kaiser and Hoogstraal, 1973 (Carios) 19. Argas delicatus Neumann, 1910. This species is very close to A. vulgaris, as discussed in Guglielmone et al. (2009). We consider both names valid until the types can be compared. 20. Argas dewae Kaiser and Hoogstraal, 1974 (Carios) 21. Argas dulus Keirans, Clifford and Capriles, 1971 22. Argas echinops Hoogstraal, Uilenberg and Blanc, 1967 (Ogadenus) 23. Argas falco Kaiser and Hoogstraal, 1974 24. Argas giganteus Kohls and Clifford, 1968 25. Argas gilcolladoi Estrada-Peña, Lucientes and Sánchez, 1987 26. Argas hermanni Audouin, 1826 27. Argas himalayensis Hoogstraal and Kaiser, 1973 28. Argas hoogstraali Morel and Vassiliades, 1965 (Ogadenus) 29. Argas japonicus Yamaguti, Clifford and Tipton, 1968 30. Argas keiransi Estrada-Peña, Venzal and González-Acuña, 2003 31. Argas lagenoplastis Froggatt, 1906 32. Argas latus Filippova, 1961 33. Argas lowryae Kaiser and Hoogstraal, 1975 34. Argas macrodermae Hoogstraal, Moorhouse, Wolf and Wassef, 1977 (Carios) 35. Argas macrostigmatus Filippova, 1961 36. Argas magnus Neumann, 1896 37. Argas miniatus Koch, 1844 38. Argas monachus Keirans, Radovsky and Clifford, 1973 39. Argas monolakensis Schwan, Corwin and Brown, 1992 40. Argas moreli Keirans, Hoogstraal and Clifford, 1979 41. Argas neghmei Kohls and Hoogstraal, 1961 42. Argas nullarborensis Hoogstraal and Kaiser, 1973 43. Argas persicus (Oken, 1818) 44. Argas polonicus Siuda, Hoogstraal, Clifford and Wassef, 1979 45. Argas pusillus Kohls, 1950 (Carios) 46. Argas radiatus Railliet, 1893 47. Argas reflexus (Fabricius, 1794). This is the type species of the genus Argas; it was originally named Acarus reflexus. 48. Argas ricei Hoogstraal, Kaiser, Clifford and Keirans, 1975 49. Argas robertsi Hoogstraal, Kaiser and Kohls, 1968 50. Argas sanchezi Dugès, 1887 51. Argas sinensis Jeu and Zhu, 1982 (Carios) 52. Argas streptopelia Kaiser, Hoogstraal and Horner, 1970 53. Argas striatus Bedford, 1932 54. Argas theilerae Hoogstraal and Kaiser, 1970 55. Argas transgariepinus White, 1846 (Ogadenus) 56. Argas transversus Banks, 1902 (Microargas, Ornithodoros). This is the type species of the genus Microargas; it was originally named Argas transversa. 57. Argas tridentatus Filippova, 1961 58. Argas vespertilionis (Latreille, 1796) (Carios). This is the type species of the genus Carios; it was originally named Carios vespertilionis. Camicas et al. (1998) and Barker and Murrell (2008) consider Argas / Carios fischeri Audouin, 1826 a valid name, while Guglielmone et al. (2009) follow Hoogstraal (1958) and Filippova (1964) in considering this name a provisional synonym of A. vespertilionis. 59. Argas vulgaris Filippova, 1961. See A. delicatus. 60. Argas walkerae Kaiser and Hoogstraal, 1969 61. Argas zumpti Hoogstraal, Kaiser and Kohls, 1968 1. Nothoaspis reddelli Keirans and Clifford, 1975 (Carios). This is the type species of the genus Nothoaspis. 1. Ornithodoros acinus Whittick, 1938 (Alveonasus, Argas) 2. Ornithodoros alactagalis Issaakjan, 1936 (Alectorobius) 3. Ornithodoros amblus Chamberlin, 1920 (Alectorobius, Carios) 4. Ornithodoros antiquus Poinar, 1995. Fossil. 5. Ornithodoros apertus Walton, 1962 6. Ornithodoros aragaoi Fonseca, 1960 (Alectorobius, Carios). We consider this name to be provisionally valid, pending type comparison with O. rudis (Guglielmone et al. 2009). 7. Ornithodoros arenicolous Hoogstraal, 1953 (Alectorobius) 8. Ornithodoros asperus Warburton, 1918 (Alectorobius). See O. verrucosus. 9. Ornithodoros azteci Matheson, 1935 (Alectorobius, Carios) 10. Ornithodoros batuensis Hirst, 1929 (Alectorobius, Carios) 11. Ornithodoros brasiliensis Aragão, 1923 (Alectorobius) 12. Ornithodoros brodyi Matheson, 1935 (Alectorobius, Carios) 13. Ornithodoros camicasi (Sylla, Cornet and Marchand, 1997) (Alectorobius, Carios) 14. Ornithodoros canestrinii (Birula, 1895) (Argas, Alveonasus) 15. Ornithodoros capensis Neumann, 1901 (Alectorobius, Carios) 16. Ornithodoros casebeeri Jones and Clifford, 1972 (Alectorobius, Carios) 17. Ornithodoros cheikhi Vermeil, Marjolet and Vermeil, 1997 (Alectorobius, Carios) 18. Ornithodoros chironectes Jones and Clifford, 1972 (Alectorobius, Carios) 19. Ornithodoros chiropterphila Dhanda and Rajagopalan, 1971 (Alectorobius, Carios) 20. Ornithodoros cholodkovskyi Pavlovsky, 1930 (Alectorobius) 21. Ornithodoros clarki Jones and Clifford, 1972 (Carios) 22. Ornithodoros collocaliae Hoogstraal, Kadarsan, Kaiser and Van Peenen, 1974 (Alectorobius, Carios) 23. Ornithodoros compactus Walton, 1962 24. Ornithodoros concanensis Cooley and Kohls, 1941 (Alectorobius, Carios) 25. Ornithodoros coniceps (Canestrini, 1890) (Alectorobius, Carios) 26. Ornithodoros cooleyi McIvor, 1941 (Alveonasus, Argas). If this species belongs to Argas, it has priority over A. cooleyi Kohls and Hoogstraal, 1960. 27. Ornithodoros coriaceus Koch, 1844 28. Ornithodoros cyclurae de la Cruz, 1984 (Alectorobius, Carios) 29. Ornithodoros darwini Kohls, Clifford and Hoogstraal, 1969 (Alectorobius, Carios) 30. Ornithodoros delanoei Roubaud and Colas-Belcour, 1931 (Alveonasus, Argas) 31. Ornithodoros denmarki Kohls, Sonenshine and Clifford, 1965 (Alectorobius, Carios) 32. Ornithodoros dugesi Mazzotti, 1943 (Alectorobius, Carios). We consider this name to be provisionally valid (Guglielmone et al. 2009), although it is generally regarded as a synonym of O. talaje. 33. Ornithodoros dusbabeki Č erný, 1967 (Alectorobius, Carios) 34. Ornithodoros dyeri Cooley and Kohls, 1940 (Alectorobius, Carios) 35. Ornithodoros eboris Theiler, 1959 (Alveonasus, Argas) 36. Ornithodoros echimys Kohls, Clifford and Jones, 1969 (Alectorobius, Carios) 37. Ornithodoros elongatus Kohls, Sonenshine and Clifford, 1965 (Alectorobius, Carios) 38. Ornithodoros eptesicus Kohls, Clifford and Jones, 1969 (Alectorobius, Carios) 39. Ornithodoros eremicus Cooley and Kohls, 1941 40. Ornithodoros erraticus (Lucas, 1849) (Alectorobius) 41. Ornithodoros faini Hoogstraal, 1960 (Alectorobius, Carios) 42. Ornithodoros foleyi Parrot, 1928 (Alveonasus, Argas) 43. Ornithodoros fonsecai (Labruna and Venzal, 2009) (n. syn.) (Alectorobius, Carios). This species was recently described by Labruna and Venzal (2009), who placed it in the genus Carios. However, its morphology indicates that it should be included as a member of Ornithodoros (as in this classification) or Alectorobius. 44. Ornithodoros furcosus Neumann, 1908 (Alectorobius) 45. Ornithodoros galapagensis Kohls, Clifford and Hoogstraal, 1969 (Alectorobius, Carios) 46. Ornithodoros graingeri Heisch and Guggisberg, 1953 (Alectorobius) 47. Ornithodoros grenieri Klein, 1965 (Alectorobius) 48. Ornithodoros gurneyi Warburton, 1926 49. Ornithodoros hadiae (Klompen, Keirans and Durden, 1995) (Alectorobius, Carios) 50. Ornithodoros hasei (Schulze, 1935) (Alectorobius, Carios) 51. Ornithodoros hermsi Wheeler, Herms and Meyer, 1935 (Alectorobius) 52. Ornithodoros indica Rau and Rao, 1971 53. Ornithodoros jerseyi (Klompen and Grimaldi, 2001) (Carios). Fossil. 54. Ornithodoros jul Schulze, 1940 (Alectorobius, Carios) 55. Ornithodoros kelleyi Cooley and Kohls, 1941 (Alectorobius, Carios) 56. Ornithodoros knoxjonesi Jones and Clifford, 1972 (Alectorobius, Carios). We consider this name provisionally valid, pending comparison with O. dyeri (Guglielmone et al. 2009). 57. Ornithodoros kohlsi Guglielmone and Keirans, 2002 (Alectorobius, Carios). Originally named O. boliviensis Kohls and Clifford, 1964, which is preoccupied (Guglielmone et al. 2009). 58. Ornithodoros lahorensis Neumann, 1908 (Alveonasus, Argas). This is the type species of the genus Alveonasus. 59. Ornithodoros macmillani Hoogstraal and Kohls, 1966 (Alectorobius) 60. Ornithodoros madagascariensis Hoogstraal, 1962 (Alectorobius, Carios) 61. Ornithodoros marinkellei Kohls, Clifford and Jones, 1969 (Alectorobius, Carios) 62. Ornithodoros maritimus Vermeil and Marguet, 1967 (Alectorobius, Carios) 63. Ornithodoros marmosae Jones and Clifford, 1972 (Alectorobius, Carios) 64. Ornithodoros marocanus Velu, 1919. We consider this name provisionally valid, pending type comparison with O. erraticus (Guglielmone et al. 2009). 65. Ornithodoros mimon Kohls, Clifford and Jones, 1969 (Alectorobius, Carios) 66. Ornithodoros mormoops Kohls, Clifford and Jones, 1969 (Alectorobius, Carios) 67. Ornithodoros moubata (Murray, 1877) 68. Ornithodoros muesebecki Hoogstraal, 1969 (Alectorobius, Carios) 69. Ornithodoros multisetosus (Klompen, Keirans and Durden, 1995) (Alectorobius, Carios) 70. Ornithodoros natalinus Č erný and Dusbábek, 1967 (Alectorobius, Carios) 71. Ornithodoros nattereri Warburton, 1927. We consider this name provisionally valid, pending comparison with O. rostratus (Guglielmone et al. 2009). 72. Ornithodoros nicollei Mooser, 1932 73. Ornithodoros normandi Larrousse, 1923 (Alectorobius) 74. Ornithodoros papuensis (Klompen, Keirans and Durden, 1995) (Alectorobius, Carios) 75. Ornithodoros papillipes (Birula, 1895) (Alectorobius). This species is considered a synonym of O. tholozani by many Western scientists (Hoogstraal, 1985), but East European workers strongly defend the validity of this name with arguments that are scientifically sound (Filippova, 1966). The uncertain status of these taxa has led us to treat them both as provisionally valid. 76. Ornithodoros parkeri Cooley, 1936 (Alectorobius) 77. Ornithodoros peringueyi Bedford and Hewitt, 1925 (Argas, Ogadenus) 78. Ornithodoros peropteryx Kohls, Clifford and Jones, 1969 (Alectorobius, Carios) 79. Ornithodoros peruvianus Kohls, Clifford and Jones, 1969 (Alectorobius, Carios) 80. Ornithodoros peusi (Schulze, 1943) (Argas, Ogadenus) 81. Ornithodoros piriformis Warburton, 1918 (Alectorobius, Carios) 82. Ornithodoros porcinus Walton, 1962 83. Ornithodoros procaviae Theodor and Costa, 1960 (Alectorobius) 84. Ornithodoros puertoricensis Fox, 1947 (Alectorobius, Carios) 85. Ornithodoros rennellensis Clifford and Sonenshine, 1962 (Alectorobius, Carios) 86. Ornithodoros rioplatensis Venzal, Estrada-Peña and Mangold, 2008 (Alectorobius, Carios) 87. Ornithodoros rondoniensis (Labruna, Terassini, Camargo, Brandão, Ribeiro and Estrada-Peña, 2008) (Alectorobius, Carios). 88. Ornithodoros rossi Kohls, Sonenshine and Clifford, 1965 (Alectorobius, Carios) 89. Ornithodoros rostratus Aragão, 1911 90. Ornithodoros rudis Karsch, 1880 (Alectorobius, Carios) 91. Ornithodoros salahi Hoogstraal, 1953 (Alectorobius, Carios) 92. Ornithodoros savignyi (Audouin, 1826). This is the type species of the genus Ornithodoros; it was originally named Argas savignyi. There is discrepancy concerning the year of description, which some authors consider to be 1827; this discrepancy also includes the reference for the description. 93. Ornithodoros sawaii Kitaoka and Suzuki, 1973 (Alectorobius, Carios) 94. Ornithodoros setosus Kohls, Clifford and Jones, 1969 (Alectorobius, Carios) 95. Ornithodoros solomonis Dumbleton, 1959 (Alectorobius, Carios) 96. Ornithodoros sonrai Sautet and Witkowski, 1943 (Alectorobius) 97. Ornithodoros sparnus Kohls and Clifford, 1963 (Alectorobius, Otobius) 98. Ornithodoros spheniscus Hoogstraal, Wassef, Hays and Keirans, 1985 (Alectorobius, Carios) 99. Ornithodoros stageri Cooley and Kohls, 1941 (Alectorobius, Carios) 100. Ornithodoros tadaridae Č erný and Dusbábek, 1967 (Alectorobius, Carios) 101. Ornithodoros talaje (Guérin-Méneville, 1849) (Alectorobius, Carios). This is the type species of the genus Alectorobius; it was originally named Argas talaje. See also O. dugesi. 102. Ornithodoros tartakovskyi Olenev, 1931 (Alectorobius) 103. Ornithodoros tholozani (Laboulbène and Mégnin, 1882) (Alectorobius). See also O. papillipes. 104. Ornithodoros tiptoni Jones and Clifford, 1972 (Alectorobius, Carios) 105. Ornithodoros turicata (Dugès, 1876) (Alectorobius) 106. Ornithodoros tuttlei Jones and Clifford, 1972 (Alectorobius, Carios) 107. Ornithodoros vansomereni Keirans, Hoogstraal and Clifford, 1977 (Argas, Ogadenus) 108. Ornithodoros verrucosus Olenev, Zasukhin and Fenyuk, 1934 (Alectorobius) is considered a synonym of O. asperus by Western workers (Hoogstraal, 1985), but Russian scientists present sound evidence for its validity (Filippova, 1966). We tentatively accept both taxa, pending further studies. 109. Ornithodoros viguerasi Cooley and Kohls, 1941 (Alectorobius, Carios) 110. Ornithodoros yumatensis Cooley and Kohls, 1941 (Alectorobius, Carios) 111. Ornithodoros yunkeri Keirans, Clifford and Hoogstraal, 1984 (Alectorobius, Carios) 112. Ornithodoros zump

Nuttalliellidae 1. Nuttalliella namaqua Bedford, 1931. This is the type species of the genus Nuttalliella.

Hyalomma (Hyalommina) kumari Sharif, 1928 (Figs. 13���18) Type specimens. Syntypes (male and female; not quantified) ex tahr, Hemitragus hylocrius (Ogilby) from Parambikulam, Cochin State [now Kerala State, India]; deposited in the Indian Museum, reg. № 2173 / 17 (Kolkata, India) (p. 320, Sharif 1928). Some of syntypes (2 males, 2 females; Nuttall 3659) are housed in the Natural History Museum (London) (Keirans & Hillyard 2001). Later authors have added the following data for the syntypes: the date of the collection (16-24 Sept 1924) and the collector (F.A. Gravely). Material studied. A total of 40 males, 30 females, 17 nymphs and 30 larvae that originated from Afghanistan, India, Iran, Nepal, Pakistan and Tajikistan were examined in the present study. Description. Male (Figs. 13, 14) Conscutum (Fig. 13): narrow, length 2.28���3.22 (2.68 �� 0.25, n = 19), width 1.34���1.78 (1.52 �� 0.12, n = 19), ratio length:width 1.62���1.87 (1.76 �� 0.07, n = 19); usually yellow-brown; fairly sparse large, mediumsized and small punctations mainly on anterior part of conscutum, caudal and lateral fields. Spiracular plate (Fig. 14 C): perforated portion of prolongation very broad. Female (Figs. 15, 16) Scutum (Fig. 15): length 1.42���2.05 (1.74 �� 0.16, n = 10), width 1.29���1.85 (1.55 �� 0.16, n = 10), ratio length:width 1.09���1.20 (1.13 �� 0.03, n = 10); usually yellow-brown; large and medium-sized punctations moderately dense on lateral fields and anterior part of central fields; punctations sparse on cervical fields and on central and posterior parts of central field. Genital structures (Fig. 16 A): genital aperture narrow, arcuate (U-shaped); vestibular portion of vagina markedly bulging; preatrial fold of genital aperture bulging throughout its length (Fig. 16 B). Nymph (Fig. 17) Scutum (Fig. 17 A): length 461���556 (510 �� 22.94, n= 16), width 421���485 (463 �� 18.47, n= 16), ratio length:width 1.03���1.15 (1.10 �� 0.03, n= 16), distance between posterior margin of eyes and posterior margin of scutum 143���187 (165 �� 11.52, n= 16), width:length of posterior portion of scutum 2.55 ���3.00 (2.81 �� 0.13, n= 16); slight posterolateral depressions on either side of scutal extremity. Basis capituli (Figs. 17 D, E): length 340���400 (365 �� 17.37, n= 16); width 290���348 (316 �� 17.48, n= 16), ratio length:width 1.11���1.20 (1.16 �� 0.03, n= 16); ventrally lateral saliences slightly convex without spur. Palpi (segment II) (Figs. 17 D, E): length 146���180 (158 �� 9.03, n= 16), width 44���48 (46 �� 1.57, n= 16), ratio length:width 3.11���3.83 (3.42 �� 0.20, n= 16). Hypostome (Fig. 17 E): length 154���190 (170 �� 9.86, n= 16), width 38���50 (44 �� 3.39, n= 16), ratio length:width 3.24���4.10 (3.87 �� 0.20, n= 16); rounded at apex; 8 or 9 large denticles in median file. Coxae (Fig. 17 F): coxae II with moderate spur. Larva (Fig. 18) Scutum (Fig. 18 A): length 216���232 (225 �� 3.92, n= 30), width 314���332 (323 �� 4.54, n= 30), ratio length:width 0.67���0.72 (0.69 �� 0.01, n= 30), distance from posterior margin of eyes to posterior margin of scutum 48���56 (52 �� 2.22, n= 30), ratio width:length of posterior portion 5.71���6.92 (6.25 �� 0.29, n= 30). Portion of scutum posterior to eyes slightly less than 1 / 4 of scutal length. Basis capituli (Figs. 18 B, C): length 148���166 (159 �� 3.53, n= 30), width 152���162 (158 �� 2.19, n= 30), ratio length:width 0.95���1.06 (1.01 �� 0.02, n= 30). Palpi (segments II and III) (Figs. 18 B, C): length 108���114 (111 �� 1.58, n= 30), width 31���36 (32 �� 0.97, n= 30), ratio length:width 3.08���3.56 (3.42 �� 0.11, n= 30). Hypostome (Fig. 18 C): length 86���92 (90 �� 1.67, n= 30), width 21���22 (21 �� 0.51, n= 30), ratio length:width 4.00��� 4.38 (4.17 �� 0.11, n= 30); 7 or 8 large denticles in median file. Genu I: length 118���126 (121 �� 1.98, n= 30), width 42���44 (43 �� 0.92, n= 8), ratio length:width 2.73 ���3.00 (2.83 �� 0.08, n= 8). Hosts. The principal hosts of the adults are large and medium-sized domestic and wild ungulates: cattle, goats, buffaloes, sheep, pigs, camels, horses, donkeys, Indian muntjac, Muntiacus muntjak (Zimmermann), tahr, Hemitragus hylocrius (Ogilby), markhor, Capra falconeri (Wagner) and wild goat, Capra hircus Linnaeus. Adults have also been recorded from domestic dogs and tiger, P. tigris (our data; Sharif 1928; Starkov 1972; Miranpuri & Naithani 1978; Hoogstraal & Valdez 1980; Geevarghese & Dhanda 1987). The main hosts of the immature stages of H. kumari are various rodents and other small mammals. The immature stages have been recorded from Asian house shrew, S. murinus, Blanford���s rat, C. blanfordi, Cutch rat, C. cutchicus, house rat, R. rattus, soft-furred rat, M. meltada, Indian bush rat, G. ellioti, rock-loving mouse, M. saxicola, flat-haired mouse, M. platythrix, little Indian field mouse, M. booduga and Indian gerbil, T. indica (Singh & Dhanda 1965; Kaul et al. 1978, 1979; Geevarghese & Dhanda 1987). One larva has been recorded from a bird: munia, Lonchura sp. (Kaul et al. 1978). It is most probable that the records of the immature stages from larger domestic animals such as goats, buffaloes and dogs (Chaudhuri 1970; Gill & Gill 1977) are the result of confusing their identification with those of H. anatolicum or H. dromedarii Koch, 1844. These records and the identification of the immature stages need to be rechecked. Geographic distribution. Asia: Afghanistan, India, Iran, Nepal, Pakistan and Tajikistan (our data; Sharif 1928; Kaiser & Hoogstraal 1963, 1964; Starkov 1972; Hoogstraal & Valdez 1980; Geevarghese & Dhanda 1987). Upon revising the Hyalomma collection of the USNTC we found a collection lot containing H. kumari from Iran: RML 98817 (HH 54068), 2 females, Iran, Fars, 3.4 miles North of Bastak, ex Capra hircus aegagrus, 20 November 1968, R.W. Rust leg. Previously both females had been incorrectly identified as Hyalomma asiaticum Schulze & Schlottke, 1930; this is now the second record of H. kumari from Iran. Identification. The differential characters for identification of all stages of the life cycles of H. brevipunctata, H. hussaini and H. kumari are summarized in Table 1. Character Tick species and stage of development H. brevipunctata H. hussaini H. kumari Male 1. Conscutum ��� avg. length �� width (their ratio) 2.52 �� 1.48 (1.70) 2.65 �� 1.65 (1.61) 2.68 �� 1.52 (1.76) 2. Conscutum ��� width Relatively broad Relatively broad Narrow 3. Conscutum ��� color Red-brown Red-brown Yellow-brown 4. Punctations on conscutum Dense Sparse Moderately sparse Female 1. Scutum ��� avg. length �� width (their ratio) 1.78 �� 1.64 (1.08) 1.75 �� 1.69 (1.03) 1.74 �� 1.55 (1.13) 2. Conscutum ��� color Red-brown Red-brown Yellow-brown 3. Punctations of scutum Dense Sparse Moderately sparse 4. Genital operculum Narrow U-shaped Broad U-shaped Narrow U-shaped 5. Pre-atrial fold of genital operculum Humped anteriorly and Humped anteriorly and Humped throughout sloping posteriorly sloping posteriorly its length Nymph 1. Scutum ��� avg. length �� width (their ratio) 609 �� 512 (1.19) 487 �� 494 (0.99) 510 �� 463 (1.10) 2. Scutum ��� avg. length of posterior part 230 156 165 3. Scutum ��� avg. ratio width to length of posterior part 2.23 3.16 2.81 4. Gnathosoma ��� avg. length �� width (their ratio) 402 �� 346 (1.16) 351 �� 313 (1.12) 365 �� 316 (1.16) 5. Palpi ��� avg. length �� width (their ratio) 168 �� 47 (3.57) 158 �� 47 (3.37) 146 �� 46 (3.42) 6. Hypostome ��� avg. length �� width (their ratio) 182 �� 44 (4.14) 165 �� 52 (3.18) 170 �� 44 (3.87) 7. Scutum posteriolateral margin Without depressions With depressions With depressions 8. Lateral saliences of basis capituli ventrally Without spur With spur Without spur 9. Hypostome Tapering at apex Blunt at apex Blunt at apex 10. Number of denticles in median file of hypostome 7 7 or 8 8 or 9 11. Spur of coxae II Moderate Large Moderate Larva 1. Scutum ��� avg. length �� width (their ratio) 233 �� 317 (0.74) 223 �� 325 (0.69) 225 �� 323 (0.69) 2. Scutum ��� avg. length of posterior part 57 53 52 3. Scutum ��� avg. ratio width to length of posterior part 5.52 6.15 6.25 4. Gnathosoma ��� avg. length �� width (their ratio) 153 �� 157 (0.98) 147 �� 158 (0.93) 159 �� 158 (1.01) 5. Palpi ��� avg. length �� width (their ratio) 112 �� 36 (3.12) 109 �� 33 (3.26) 111 �� 32 (3.42) 6. Hypostome ��� avg. length �� width (their ratio) 86 �� 21 (4.12) 86 �� 22 (3.80) 90 �� 21 (4.17) 7. Genu ��� avg. length �� width (their ratio) 128 �� 44 (2.95) 120 �� 44 (2.69) 121 �� 43 (2.83) 8. Number of denticles in median file of hypostome 6 6 7 or 8, Published as part of Apanaskevich, Dmitry A., Horak, Ivan G. & Geevarghese, Geevarghese, 2009, The genus Hyalomma Koch, 1844. VIII. Redescription of three Hyalommina Schulze, 1919 species (Acari: Ixodidae) from South Asia with notes on their biology, pp. 31-55 in Zootaxa 2050 on pages 47-53, DOI: 10.5281/zenodo.186557, {"references":["Sharif, M. (1928) A revision of the Indian Ixodidae with special reference to the collection in the Indian Museum. Records of the Indian Museum, 30, 217 - 344.","Keirans, J. E. & Hillyard, P. D. (2001) A catalogue of the type specimens of Ixodida (Acari: Argasidae, Ixodidae) deposited in the Natural History Museum, London. Occasional Papers on Systematic Entomology, 13, 1 - 74.","Starkov, O. A. (1972) Hyalomma (Hyalommina) kumari Sharif, 1928 (Ixodoidea, Ixodidae) - a new species of bloodsucking tick for fauna of the USSR. Izvestiya Akademii Nauk Tadzhikskoi SSR, Otdelenie Biologicheskikh Nauk, 1, 73 - 75. (in Russian)","Miranpuri, G. S. & Naithani R. C. (1978) A check list of Indian ticks (Ixodoidea: Acarina). Indian Veterinary Research Institute, Izatnagar, India, 50 pp.","Hoogstraal, H. & Valdez, R. (1980) Ticks (Ixodoidea) from wild sheep and goats in Iran and medical and veterinary implications. Fieldiana Zoology, 6, 1 - 16.","Geevarghese, G. & Dhanda, V. (1987) The Indian Hyalomma ticks (Ixodoidea: Ixodidae). Indian Council of Agricultural Research, New Delhi, India, 119 pp.","Singh, K. R. P. & Dhanda, V. (1965) Description and keys of immature stages of some species of Indian Hyalomma Koch, 1844 (Ixodoidea, Ixodidae). Acarologia, 7, 636 - 651.","Kaul, H. N., Mishra, A. C., Dhanda, V., Kulkarni, S. M. & Guttikar, S. N. (1978) Ectoparasitic arthropods of birds and mammals from Rajasthan State, India. Indian Journal of Parasitology, 2, 19 - 25.","Kaul, H. N., Dhanda, V. & Mishra, A. C. (1979) A survey of ixodid ticks in Orissa State, India. Indian Journal of Animal Sciences, 49, 707 - 712.","Gill, H. S. & Gill, B. S. (1977) Ixodid ticks of domestic animals in the Punjab State. Punjab Agricultural University, Ludhiana, India, 69 pp.","Kaiser, M. N. and Hoogstraal, H. (1963) The Hyalomma ticks (Ixodoidea, Ixodidae) of Afghanistan. Journal of Parasitology, 49, 130 - 139.","Kaiser, M. N. and Hoogstraal, H. (1964) The Hyalomma ticks (Ixodoidea, Ixodidae) of Pakistan, India, and Ceylon, with keys to subgenera and species. Acarologia, 6, 257 - 286."]}

Hyalomma (Hyalommina) hussaini Sharif, 1928 (Figs. 7���12) Type specimens. Syntypes (sex not indicated; not quantified) ex cattle from Akola Town in the Central Provinces [now Maharashtra State, India]; deposited in the Indian Museum, reg. № 57 / 18 (Kolkata, India) (p. 317, Sharif 1928). Material studied. A total of 31 males, 26 females, 7 nymphs and 32 larvae that originated from India, Myanmar and Pakistan were examined in the current study. Synonym. Hyalomma hussaini forma typica Sharif, 1928 Description. Male (Figs. 7, 8) Conscutum (Fig. 7): length 2.21���2.93 (2.65 �� 0.17, n = 25), width 1.34���1.82 (1.65 �� 0.12, n = 25), ratio length:width 1.51���1.76 (1.61 �� 0.06, n = 25); dark red-brown; sparse large punctations mainly on anterior part of conscutum, lateral and caudal fields. Spiracular plate (Fig. 8 C): perforated portion of prolongation very broad. Female (Figs. 9, 10) Scutum (Fig. 9): length 1.62���1.88 (1.75 �� 0.08, n = 19), width 1.52���1.85 (1.69 �� 0.09, n = 19), ratio length:width 1.00��� 1.06 (1.03 �� 0.02, n = 19); usually dark red-brown; large, deep punctations sparse, evenly distributed over scutum. Genital structures (Fig. 10 A): genital aperture very wide, arcuate (U-shaped); vestibular portion of vagina markedly bulging; preatrial fold of genital aperture bulging anteriorly and sloping sharply posteriorly (Fig. 10 B). Nymph (Fig. 11) Scutum (Fig. 11 A): length 449���525 (487 �� 27.51, n= 6), width 453���533 (494 �� 27.59, n= 6), ratio length:width 0.88���1.06 (0.99 �� 0.07, n= 6), distance between posterior margin of eyes and posterior margin of scutum 151���175 (156 �� 9.63, n= 6), width:length of posterior portion of scutum 2.82���3.39 (3.16 �� 0.22, n= 6); slight posterolateral depressions on either side of scutal extremity. Basis capituli (Figs. 11 D, E): length 312���376 (351 �� 21.97, n= 6); width 292���332 (313 �� 16.33, n= 6), ratio length:width 1.07���1.15 (1.12 �� 0.03, n= 6); ventrally lateral saliences with moderate spur directed posteriorly. Palpi (segment II) (Figs. 11 D, E): length 140���176 (158 �� 11.48, n= 6), width 45���48 (47 �� 1.09, n= 6), ratio length:width 3.04���3.83 (3.37 �� 0.26, n= 6). Hypostome (Fig. 11 E): length 144���178 (165 �� 11.64, n= 6), width 50���54 (52 �� 1.26, n= 6), ratio length:width 2.77���3.42 (3.18 �� 0.23, n= 6); rounded at apex; 7 or 8 large denticles in median file. Coxae (Fig. 11 F): coxae II with large triangular spur. Larva (Fig. 12) Scutum (Fig. 12 A): length 208���232 (223 �� 5.69, n= 32), width 300���344 (325 �� 9.21, n= 32), ratio length:width 0.66���0.72 (0.69 �� 0.01, n= 32), distance from posterior margin of eyes to posterior margin of scutum 48���58 (53 �� 2.32, n= 32), ratio width:length of posterior portion 5.57���6.92 (6.15 �� 0.31, n= 32). Portion of scutum posterior to eyes nearly equal to 1 / 4 of scutal length. Basis capituli (Figs. 12 B, C): length 136���160 (147 �� 5.60, n= 32), width 150���164 (158 �� 3.42, n= 32), ratio length:width 0.89 ���1.00 (0.93 �� 0.03, n= 32). Palpi (segments II and III) (Figs. 12 B, C): length 104���114 (109 �� 2.18, n= 32), width 32���34 (33 �� 0.83, n= 32), ratio length:width 3.12���3.44 (3.26 �� 0.08, n= 32). Hypostome (Fig. 12 C): length 80���90 (86 �� 2.33, n= 32), width 22���24 (22 �� 0.62, n= 32), ratio length:width 3.58 ���4.00 (3.80 �� 0.10, n= 32); 6 large denticles in median file. Genu I: length 114���128 (120 �� 2.44, n= 32), width 42���46 (44 �� 1.40, n= 7), ratio length:width 2.56���2.86 (2.69 �� 0.10, n= 7). Hosts. The main hosts of the adults are large and medium-sized domestic and wild ungulates: cattle, buffaloes, camels, goats, sheep, pigs, horses and nilgai, B. tragocamelus. The adults have also been recorded from domestic dogs, a bear and humans (our data; Sharif 1928; Kaiser & Hoogstraal 1964; Miranpuri & Naithani 1978; Geevarghese & Dhanda 1987; Robbins et al. 2002). The chief hosts of the immature stages are small mammals such as rodents and shrews. Larvae and nymphs have been recorded from: Asian house shrew, S. murinus, house rat, R. rattus, Indian bush rat, Golunda ellioti Gray, Indian desert jird, Meriones hurrianae Jordon and rock-loving mouse, M. saxicola (Singh & Dhanda 1965; Wattal & Srivastva 1967; Geevarghese & Dhanda 1987). Geographic distribution. Asia: India, Myanmar and Pakistan (our data; Sharif 1928; Kaiser & Hoogstraal 1964; Geevarghese & Dhanda 1987; Robbins et al. 2002)., Published as part of Apanaskevich, Dmitry A., Horak, Ivan G. & Geevarghese, Geevarghese, 2009, The genus Hyalomma Koch, 1844. VIII. Redescription of three Hyalommina Schulze, 1919 species (Acari: Ixodidae) from South Asia with notes on their biology, pp. 31-55 in Zootaxa 2050 on pages 39-47, DOI: 10.5281/zenodo.186557, {"references":["Sharif, M. (1928) A revision of the Indian Ixodidae with special reference to the collection in the Indian Museum. Records of the Indian Museum, 30, 217 - 344.","Kaiser, M. N. and Hoogstraal, H. (1964) The Hyalomma ticks (Ixodoidea, Ixodidae) of Pakistan, India, and Ceylon, with keys to subgenera and species. Acarologia, 6, 257 - 286.","Miranpuri, G. S. & Naithani R. C. (1978) A check list of Indian ticks (Ixodoidea: Acarina). Indian Veterinary Research Institute, Izatnagar, India, 50 pp.","Geevarghese, G. & Dhanda, V. (1987) The Indian Hyalomma ticks (Ixodoidea: Ixodidae). Indian Council of Agricultural Research, New Delhi, India, 119 pp.","Robbins, R. G., Platt, S. G. & Keirans, J. E. (2002) First report of Hyalomma marginatum isaaci Sharif (Acari: Ixodida: Ixodidae) from the Union of Myanmar, with a concurrent collection of H. hussaini Sharif. Proceedings of the Entomological Society of Washington, 104, 1061 - 1063.","Singh, K. R. P. & Dhanda, V. (1965) Description and keys of immature stages of some species of Indian Hyalomma Koch, 1844 (Ixodoidea, Ixodidae). Acarologia, 7, 636 - 651.","Wattal, B. L. & Srivastva, S. P. (1967) Ectoparasite fauna of small mammals and domestic animals in the neighbourhood of Alwar city (Rajasthan). Bulletin of the Indian Society for Malaria and other Communicable Diseases, 4, 191 - 202."]}

Hyalomma (Hyalommina) kumari group (Figs. 1���18) Species composition. Hyalomma brevipunctata Sharif, 1928, H. hussaini Sharif, 1928 and H. kumari Sharif, 1928. Description. Male (Figs. 1, 2, 7, 8, 13, 14) Conscutum (Figs. 1, 7, 13): pale marbling absent; narrowly oval in shape; broadest posterior to midlength; slight narrowing in region of spiracular plates; cervical and lateral grooves shallow, up to 1 / 4 - 1 / 3 length of conscutum; marginal grooves long, almost reaching eyes; posteromedian groove does not reach median festoon (or may reach it in H. hussaini); paramedian grooves well defined, pit-like; caudal field inconspicuous; parma always absent; 11 distinct festoons. Genital structures (Figs. 2 A, 8 A, 14 A) as illustrated. Anal shields (Figs. 2 B, 8 B, 14 B): 2 pairs, subanal plates absent; adanal plates short, broad, lateral margin slightly convex or straight, anteromedian margin concave, median projection absent, posteromedian margin straight, posterior margin straight. Sclerotized plaques present ventrally on median and paramedian festoons. Spiracular plate (Figs. 2 C, 8 C, 14 C): dorsal prolongation long and not clearly separable from body of plate; perforated portion of prolongation straight, curving at its apex. Circumspiracular setae sparse. Basis capituli (Figs. 2 D, E, 8 D, E, 14 D, E): with prominent lateral projections dorsally and ventrally; dorsal posterior margin straight or slightly concave; cornua modest. Palpi (Figs. 2 F, 8 F, 14 F): segment I with 2 or 3 ventromedian setae. Hypostome (Figs. 2 G, 8 G, 14 G): club-shaped; denticulate portion nearly twice as long as denticle-free portion (small scale-like projections posterior to last large denticle are not considered denticles). Coxae (Figs. 2 H, 8 H, 14 H): posteromedian and posterolateral spurs of coxae I long, posterolateral spur slightly longer than posteromedian spur, juxtaposed, tapering to apices; median margin of coxae I without triangular projection; coxae II���IV each with distinct, subtriangular posterolateral spur, and with slight, broadly arcuate, posteromedian spur. Segments of legs uniformly yellow- or red-brown. Female (Figs. 3, 4, 9, 10, 15, 16) Scutum (Figs. 3, 9, 15): pale marbling absent; slightly longer than broad; posterolateral angles moderate; cervical and lateral grooves shallow, extending medially to posterior margin of scutum. Spiracular plates (Figs. 4 C, 10 C, 16 C): perforated portion of dorsal projection curved, broad. Circumspiracular setae sparse. Basis capituli (Figs. 4 D, E, 10 D, E, 16 D, E): with prominent dorsal and ventral lateral projections; dorsal posterior margin very slightly concave or straight; dorsal cornua inconspicuous. Palpi (Figs. 4 F, 10 F, 16 F): segment I with 3 or 4 ventromedian setae. Hypostome (Figs. 4 G, 10 G, 16 G): club-shaped; denticulate portion nearly twice as long as denticle-free portion. Coxae (Figs. 4 H, 10 H, 16 H): posteromedian and posterolateral spurs of coxa I long, posterolateral spur slightly longer than posteromedian spur, juxtaposed, tapering to apices; median margin of coxae I without triangular projection; coxae II���IV each with distinct, broadly triangular posterolateral spur, with rounded apex; and each with slight, broadly arcuate, posteromedian spur. Segments of legs uniformly yellow- or redbrown. Nymph (Figs. 5, 11, 17) Scutum (Figs. 5 A, 11 A, 17 A): posterior margin of scutum broadly rounded. Setae of alloscutum (Figs. 5 B, 11 B, 17 B): narrowing to denticulate apex. Spiracular plates (Figs. 5 C, 11 C, 17 C): oval; dorsal prolongation indistinct, short, broad, blunt at apex; submarginal row of perforations incomplete. Basis capituli (Figs. 5 D, E, 11 D, E, 17 D, E): subtriangular dorsally with long and acute lateral saliences. Palpi (segment II) (Figs. 5 D, E, 11 D, E, 17 D, E): palpal segment II proximally narrow, gradually expanding distally. Hypostome (Figs. 5 E, 11 E, 17 E): transition of denticulate portion to denticle-free portion abrupt; denticulate portion approximately 2 / 3 of hypostome length. Coxae (Figs. 5 F, 11 F, 17 F): coxa I with long, very broad, juxtaposed, subtriangular spurs nearly equal in length; coxae III and IV each with moderate spur, spurs conspicuously decrease in size from coxae II to IV; coxal pore present (at least on coxae I). Larva (Figs. 6, 12, 18) Scutum (Figs. 6 A, 12 A, 18 A): posterior margin of scutum broadly rounded; posterolateral depressions absent. Basis capituli (Figs. 6 B, C, 12 B, C, 18 B, C): subtriangular dorsally; apex of dorsolateral projections directed laterally; dorsolateral projections distinct and acute from ventral aspect. Hypostome (Figs. 6 C, 12 C, 18 C): transition of denticulate portion to denticle-free portion abrupt; denticulate portion approximately 2 / 3 of hypostome length. Coxae (Figs. 6 D, 12 D, 18 D): coxa I with large, long triangular spur tapering at apex directed posteriorly or medially; coxae II and III each with slight spur ��� arcuate on coxae II and triangular on coxae III. Hosts. All species of the H. kumari group are three-host ticks (Geevarghese & Dhanda 1987). The principal hosts of the adults are various wild and domestic ungulates. Carnivores are apparently secondary or occasional hosts. Larvae and nymphs are specific to various groups of small mammals (Geevarghese & Dhanda 1987; Kaiser & Hoogstraal 1963, 1964; Sharif 1928; Singh & Dhanda 1965). Geographic distribution. South Asia and neighboring regions of Central Asia. Disease relationships. Little is known about the vector status of the South Asian Hyalommina species, except that the organism causing Q-fever, Coxiella burnetti, has been isolated from H. hussaini in India (Geevarghese & Dhanda 1987)., Published as part of Apanaskevich, Dmitry A., Horak, Ivan G. & Geevarghese, Geevarghese, 2009, The genus Hyalomma Koch, 1844. VIII. Redescription of three Hyalommina Schulze, 1919 species (Acari: Ixodidae) from South Asia with notes on their biology, pp. 31-55 in Zootaxa 2050 on pages 32-34, DOI: 10.5281/zenodo.186557, {"references":["Sharif, M. (1928) A revision of the Indian Ixodidae with special reference to the collection in the Indian Museum. Records of the Indian Museum, 30, 217 - 344.","Geevarghese, G. & Dhanda, V. (1987) The Indian Hyalomma ticks (Ixodoidea: Ixodidae). Indian Council of Agricultural Research, New Delhi, India, 119 pp.","Kaiser, M. N. and Hoogstraal, H. (1963) The Hyalomma ticks (Ixodoidea, Ixodidae) of Afghanistan. Journal of Parasitology, 49, 130 - 139.","Kaiser, M. N. and Hoogstraal, H. (1964) The Hyalomma ticks (Ixodoidea, Ixodidae) of Pakistan, India, and Ceylon, with keys to subgenera and species. Acarologia, 6, 257 - 286.","Singh, K. R. P. & Dhanda, V. (1965) Description and keys of immature stages of some species of Indian Hyalomma Koch, 1844 (Ixodoidea, Ixodidae). Acarologia, 7, 636 - 651."]}

Hyalomma (Hyalommina) brevipunctata Sharif, 1928 (Figs. 1���6) Type specimens. Syntypes (male and female; not quantified) ex dog from Anantapur in the Madras Presidency [now Andhra Pradesh State, India]; deposited in the Indian Museum, reg. № 56 / 18 (Kolkata, India) (p. 318, Sharif 1928). Material studied. A total of 77 males, 46 females, 6 nymphs and 24 larvae that were collected in India, Myanmar, Nepal, Pakistan and Sri Lanka were examined in the present study. Synonym. Hyalomma hussaini var. brevipunctata Sharif, 1928 Description. Male (Figs. 1, 2) Conscutum (Fig. 1): length 2.16���2.98 (2.52 �� 0.24, n = 34), width 1.20���1.73 (1.48 �� 0.14, n = 34), ratio length:width 1.51���1.87 (1.70 �� 0.08, n = 34); red-brown; fairly dense large and medium-sized punctations mainly on anterior part of conscutum, and on caudal and lateral fields, sparse, fine punctations on central part of conscutum. Spiracular plate (Fig. 2 C): perforated portion of prolongation moderately broad. Female (Figs. 3, 4) Scutum (Fig. 3): length 1.45���2.05 (1.78 �� 0.14, n = 30), width 1.42���1.91 (1.64 �� 0.13, n = 30), ratio length:width 1.00��� 1.17 (1.08 �� 0.04, n = 30); red-brown; moderately dense large punctations mainly on cervical and anterior part of central fields; moderately dense medium-sized and small punctations evenly distributed on lateral and central fields. Genital structures (Fig. 4 A): genital aperture narrow, arcuate (Ushaped) with arcuate or straight posterior margin; vestibular portion of vagina markedly bulging; preatrial fold of genital aperture bulging anteriorly and sharply sloping posteriorly (Fig. 4 B). Nymph (Fig. 5) Scutum (Fig. 5 A): length 548���644 (609 �� 42.29, n= 5), width 477���533 (512 �� 21.48, n= 5), ratio length:width 1.14���1.23 (1.19 �� 0.04, n= 5), distance between posterior margin of eyes and posterior margin of scutum 215���238 (230 �� 10.67, n= 5), width:length of posterior portion of scutum 2.14���2.37 (2.23 �� 0.09, n= 5); posterolateral depressions on either side of scutal extremity absent. Basis capituli (Figs. 5 D, E): length 372���432 (402 �� 26.32, n= 5); width 328���362 (346 �� 14.79, n= 5), ratio length:width 1.13���1.19 (1.16 �� 0.03, n= 5); ventrally lateral saliences slightly convex without spur. Palpi (segment II) (Figs. 5 D, E): length 154���178 (168 �� 9.32, n= 5), width 45���48 (47 �� 1.41, n= 5), ratio length:width 3.21���3.87 (3.57 �� 0.29, n= 5). Hypostome (Fig. 5 E): length 168���196 (182 �� 13.04, n= 5), width 40���48 (44 �� 3.16, n= 5), ratio length:width 4.04���4.26 (4.14 �� 0.09, n= 5); tapering at apex; 7 large denticles in median file. Coxae (Fig. 5 F): coxae II with moderate spur. Larva (Fig. 6) Scutum (Fig. 6 A): length 220���240 (233 �� 4.81, n= 24), width 308���328 (317 �� 5.44, n= 24), ratio length:width 0.70���0.77 (0.74 �� 0.02, n= 24), distance from posterior margin of eyes to posterior margin of scutum 52���60 (57 �� 2.23, n= 24), ratio width:length of posterior portion 5.13���6.08 (5.52 �� 0.22, n= 24). Portion of scutum posterior to eyes nearly equal to 1 / 4 of scutal length. Basis capituli (Figs. 6 B, C): length 144���160 (153 �� 3.39, n= 24), width 152���162 (157 �� 2.91, n= 23), ratio length:width 0.94���1.01 (0.98 �� 0.02, n= 23). Palpi (segments II and III) (Figs. 6 B, C): length 109���114 (112 �� 1.57, n= 24), width 34���38 (36 �� 0.78, n= 24), ratio length:width 2.89���3.26 (3.12 �� 0.08, n= 24). Hypostome (Fig. 6 C): length 84���90 (86 �� 1.55, n= 24), width 20���22 (21 �� 0.93, n= 24), ratio length:width 3.82���4.40 (4.12 �� 0.18, n= 24); 6 large denticles in median file. Genu I: length 118���134 (128 �� 3.92, n= 24), width 42���48 (44 �� 1.68, n= 15), ratio length:width 2.50���3.19 (2.95 �� 0.17, n= 15). Hosts. The main hosts of the adults are large and medium-sized domestic and wild ungulates: cattle, buffaloes, goats, camels, sheep, horses, chital, Axis axis (Erxleben), nilgai, Boselaphus tragocamelus (Pallas), sambar, Cervus unicolor Kerr and an antelope. The adults have also been recorded from domestic dogs, tiger, Panthera tigris (Linnaeus) and humans (our data; Sharif 1928; Kaiser & Hoogstraal 1964; Miranpuri & Naithani 1978; Mitchell 1979; Geevarghese & Dhanda 1987). The main hosts of the immature stages of H. brevipunctata are various rodents and other small mammals. They have been recorded from Asian house shrew, Suncus murinus (Linnaeus), Blanford���s rat, Cremnomys blanfordi (Thomas), Cutch rat, Cremnomys cutchicus Wroughton, house rat, Rattus rattus (Linnaeus), Tanezumi rat, Rattus tanezumi Temminck, soft-furred rat, Millardia meltada (Gray), greater bandicoot rat, Bandicota indica (Bechstein), Asiatic long-tailed climbing mouse, Vandeleuria oleracea (Bennett), flat-haired mouse, Mus platythrix Bennett, little Indian field mouse, Mus booduga (Gray), rock-loving mouse, Mus saxicola Elliot, Indian gerbil, Tatera indica (Hardwicke) and jungle palm squirrel, Funambulus tristriatus (Waterhouse) (Singh & Dhanda 1965; Rebello & Reuben 1967; Kaul et al. 1979; Geevarghese & Dhanda 1987). Geographic distribution. Asia: India, Myanmar, Nepal, Pakistan and Sri Lanka (our data; Sharif 1928; Kaiser & Hoogstraal 1964; Mitchell 1979; Geevarghese & Dhanda 1987; Dilrukshi 2006). Collection lot of the USNTC [RML 98964 (HH 58217), consisting of 2 males and 3 females, Burma, Pegu, Rangoon abattoir, ex domestic goat, December 1975, San-San Lin leg.] previously identified as H. hussaini, but we have reidentified them as H. brevipunctata. This is the first record of this species from Myanmar., Published as part of Apanaskevich, Dmitry A., Horak, Ivan G. & Geevarghese, Geevarghese, 2009, The genus Hyalomma Koch, 1844. VIII. Redescription of three Hyalommina Schulze, 1919 species (Acari: Ixodidae) from South Asia with notes on their biology, pp. 31-55 in Zootaxa 2050 on pages 34-37, DOI: 10.5281/zenodo.186557, {"references":["Sharif, M. (1928) A revision of the Indian Ixodidae with special reference to the collection in the Indian Museum. Records of the Indian Museum, 30, 217 - 344.","Kaiser, M. N. and Hoogstraal, H. (1964) The Hyalomma ticks (Ixodoidea, Ixodidae) of Pakistan, India, and Ceylon, with keys to subgenera and species. Acarologia, 6, 257 - 286.","Miranpuri, G. S. & Naithani R. C. (1978) A check list of Indian ticks (Ixodoidea: Acarina). Indian Veterinary Research Institute, Izatnagar, India, 50 pp.","Mitchell, R. M. (1979) A list of ectoparasites from Nepalese mammals, collected during the Nepal ectoparasite program. Journal of Medical Entomology, 16, 227 - 233.","Geevarghese, G. & Dhanda, V. (1987) The Indian Hyalomma ticks (Ixodoidea: Ixodidae). Indian Council of Agricultural Research, New Delhi, India, 119 pp.","Singh, K. R. P. & Dhanda, V. (1965) Description and keys of immature stages of some species of Indian Hyalomma Koch, 1844 (Ixodoidea, Ixodidae). Acarologia, 7, 636 - 651.","Rebello, M. J. & Reuben, R. (1967) A report on ticks collected from birds and small mammals in North Arcot and Chittoor districts, South India. Journal of the Bombay Natural History Society, 63, 283 - 289.","Kaul, H. N., Dhanda, V. & Mishra, A. C. (1979) A survey of ixodid ticks in Orissa State, India. Indian Journal of Animal Sciences, 49, 707 - 712.","Dilrukshi, P. R. M. P. (2006) Taxonomic status of ticks in Sri Lanka. In: Bambaradeniya C. N. B. (Ed.), The Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation. The World Conservation Union, Colombo, Sri Lanka, pp. 65 - 69."]}