J. Med. Entomol. Vol. 21, no. 2: 169-173 30 March 1984

DERMACENTOR (INDOCENTOR) AURATUS (ACARI: IXODOIDEA: IXODIDAE): IDENTITY OF MALE AND FEMALE12

Hilda Y. Wassef and Harry Hoogstraal3

Abstract. Dermacentor auratus, the type-species of the sub- Dermacentor auratus Supino. Neumann (1902: 126-27) dis- genus Indocentor, has long been confused taxonomically in much puted the name of this taxon (aurea = gold; Latin) because gold- Downloaded from https://academic.oup.com/jme/article/21/2/169/2220238 by guest on 25 September 2021 literature on ticks, viruses, and rickettsiae from India to the en markings are absent on 2 specimens identified by Supino; Pacific coasts of Southeast Asia and outlying islands. Dermacentor also stated that D. auratus is "very close" to D. compactus Neu- (I.) auratus is easily recognized as a distinctive species by the mann. large, broadly rounded female external genitalia, and by having Dermacentor auratus compactus Neumann. Neumann (1905: more than 2 (3-6) spurs on the greatly enlarged male coxa IV; 236-37) new combination. in both sexes, coxa I spurs are distinctly but not widely separated Dermacentor auratus Supino. Neumann (1911: 103) briefly de- and the scutal ornamentation pattern is distinctive. The male scribed 2 subspecies of D. auratus: D. a. auratus (6, 2) from and female are redescribed from study of numerous specimens Borneo, Java, Sumatra, and "Africa" and D. a. compactus (2) and the type materials in the Natural History Museum, Genoa, from Burma. Italy. Dermacentor auratus Supino. Sharif (1928: 291; fig. 31, 32; pi. VIII, fig. 6) concluded that only D. auratus (and D. everestianus As a step in revising the Oriental subgenus In- Hirst) can be recognized in India; redescribed 6 and 2 and de- docentor Schulze of the tick genus Dermacentor Koch,scribed nymph. Dermacentor auratus Supino. Krijgsman & Ponto (1932: 26, we redescribe the adults of the type-species of the 27; fig. 41, 42) redescribed 6 and 2 "Z). auratus" (from Neth- subgenus, D. (I.) auratus Supino. This is the only erlands East Indies) but illustrations show a different species of Dermacentor species in the southern half of India. the D. auratus group. However, northward into the Himalayas and east- Indocentor auratus (Supino). Schulze (1933: 417-18) briefly redescribed 8 and 2 from Burma and India and designated au- ward to the Pacific and on islands of Southeast Asia, ratus as the type species of the new genus Indocentor. D. (I.) auratus has been confused with other species, Indocentor auratus (Supino). Schulze (1939: 722-28; fig. 2) and literature on Dermacentor and viruses and rick- stated that a 2 collected in Burma entirely agrees with that described by Sharif from India. He proposed the subspecies ettsiae has been chaotic (Wassef & Hoogstraal 1983, sumatranus for Sumatran specimens differing slightly from those Hoogstraal & Wassef 1984). Dermacentor (I.) au- from Burma. ratus host and distribution data will be reported Dermacentor auratus Supino. Toumanoff (1941: 502-04; 1944: separately (Hoogstraal & Wassef, in prep.). 92-94) redescribed 6 and 2 from Indochina. Dermacentor auratus Supino. Anastos (1950: 51-59; fig. 10- Dermacentor (Indocentor) auratus Supino 12) redescribed <5 and 2 from various sources and considered Indocentor Schulze to be a junior synonym of Dermacentor; 32 FIG. 1-17 from Sumatra and 12 from Java were considered to represent Oriental Dermacentor "variations" of auratus. Dermacentor auratus Supino. Domrow & Nadchatram (1963: Notes and synonymy 157) stated that 2 "forms" of D. auratus can be distinguished Dermacentor auratus. Supino (1897: 27-28, pi. VIII, fig. 1- in Malaysia but that D. auratus is the only species represented there. 10) described $ and 9 from Ursus torquatus and Sus cristatus in Carin Cheba, 900-1000 m, and Mooleyit, 400-500 m, Te- Dermacentor auratus Supino. Seneviratna (1965: 38) stated that nasserim, Burma. auratus is the only species of Dermacentor found in Ceylon (Sri Dermacentor auratus Supino. Neumann (1897: 382) briefly re- Lanka). described 2 from Carin Cheba and Mooleyit, Burma (Fea). Dermacentor auratus Supino. Arthur (1960: 153-59; fig.3 18— 30) redescribed 8 and 2 from unstated sources and suggested that D. atrosignatus Neumann should be placed in synonymy 'From Research Project 3M161102BS10.AD.424, Naval under D. auratus. Medical Research and Development Command, National Naval Medical Center, Bethesda, Maryland. The opinions and asser- Supino Collection in Museum Civico di tions contained herein are the private ones of the authors and Storia Naturale, Genoa are not to be construed as official or as reflecting the views of The following data were recorded by one of us (HYW) in the Department of the Navy or of the naval service at large. September 1977 during a study of the tick collection in the 2 Request reprints from Medical Zoology Department, NAM- Genoa Natural History Museum, Italy. RU-3, FPO New York 09527, USA. Dermacentor auratus: 2 "type" (=holotype) (blue label) (Genoa s Medical Zoology Department, United States Naval Medical 45)(HH79,614). From "Ursus torquatus Wagner and Sus cristatus Research Unit Number Three (NAMRU-3), American Embas- Wagner, Carin Cheba, 900-1000 m alt., M. Mooleyit, Tenas- sy, Cairo, Arab Republic of Egypt. serim." In the same vial, another 2 that does not conform to 170 J. Med. Entomol. Vol. 21, no. 2 the Supino description of D. auratus is probably D. atrosignatus. dorsal shield (Fig. 1) enamelled, broadly triangular; trochanter Genitalia of both specimens were previously cut but the slides II—IV ventral spurs faintly developed or obsolete. Tarsi (Fig. are missing. A label "G.H.F. Nuttall and C. Warburton det." 10) stout, II to IV dorsal surfaces abruptly tapering distally; is also in the vial. each with 2 stout ventral spurs and a strong ventroapical hook. 5 and 9 cotypes (1 each) (in a separate vial with blue label) (L. Claws large, subequal. Pulvilli short, reaching only to midlength Fea)(Birmania 10a; HH79.615). 12Haemaphysalis hystricis in the of claws. same vial was identified by Nuttall and Warburton as a nymph of D. auratus and corrected as H. hystricis by T. Santos Dias in 2 (Fig. 3, 4, 11-16). Differs from 6 chiefly in sexual char- 1956. acters. Length 5.24 to 5.98 (avg. of 20 unfed specimens measured 5.71) mm; breadth 3.42 to 4.00 (avg. 3.8) mm. Redescription Capitulum (Fig. 11, 12). Basis capituli dorsal outline and cornua similar to those of The ca. 1850 adult D. auratus examined for this study are <5; porose areas circular, small, deep, widely spaced (as illustrated). Basis capituli ventrally as illustrated; setae listed by Hoogstraal & Wassef (in prep.). 2 pairs posteroexternally. Palpi similar to those of 6 but larger; Downloaded from https://academic.oup.com/jme/article/21/2/169/2220238 by guest on 25 September 2021 6 (Fig. 1,2,5-10). Length from palpal apices to scutal margin setae numbers differing from those of <3 as follows: segment 2 4.33 to 6.72 (avg. of 20 specimens measured 5.66) mm, breadth setae number 6 ventrally, 1 laterally. Segment 3 setae number 3.42 to 4.85 (avg. 3.9) mm. Color. Ivory enamelling covering 6 dorsally, 4 ventrally, 3 ventrointernally, 2 ventroanteriorly. most of scutum; absent on dark brown integumental elevations Hypostome (Fig. 12) essentially as in 3; corona ca. V-, as long as (median line from intercervical area to median festoon, 2 par- denticle files. amedian lines, entire pseudoscutum border), median festoon, Scutum (Fig. 3) 1.99 to 2.57 (avg. 2.19) mm long; 2.22 to 2.68 festoons 3 and 4 (in part), narrow intercervical area (variable), (avg. 2.42) mm broad, ca. 1.1 times as broad as long. Anterior cervical groove borders (variable), and irregular marginal area emargination deep, broad; scapulae blunt; margins gradually between eyes and festoons. (Color intensity variable in pre- diverging to level of eyes, thence gradually converging, pos- served specimens but usually brighter when specimens are wet.) teriorly broadly rounded. Eyes flat,oval , bordered by 2-3 punc- Capitulum (Fig. 5, 6). Basis capituli dorsally enamelled, ca. 1.8 tations. Color as in <3; extensive ivory enamelling except in brown times as broad as long (including cornua), central setae minute, area bordering eyes, a narrow line peripherally posterior to number 2-3 pairs; external margins straight; cornua short, eyes, in median line and along cervical grooves. Cervical pits as broadly triangular, apices rounded; basis capituli ventrally as in 6. Cervical grooves represented only by a slightly depressed illustrated, setae number 2-3 posteroexternal pairs. Palpi. Seg- area usually marked by a narrow brown line. Punctations mod- ment 1 a short, narrow pedicle; setae dorsally number 2 short, erately numerous, irregularly scattered in enamelled areas, size ventrally number 1 short posteriorly, 3-4 long internally. Seg- large to moderate; rare or absent in brown areas. ment 2 ca. 1.1 times as broad as long; setae number 8 dorsally, Dorsum (Fig. 3); punctations numerous, large, irregularly scat- 5 ventrally, 2 laterally; dorsointernal and ventrointernal setae tered, each with a single moderately long white seta; several number 2 and 4-5, respectively. Segment 3 ca. 0.9 times as present along lateral grooves, few on lateral margin and fes- long as 2; setae number 8 dorsally, 1 laterally, 2 ventrally, 3 toons. Color brown except in a short, pale yellowish marginal ventrointernally, 2 ventroanteriorly. Hypostome (Fig. 6) ca. 2 band slightly posterior to body midlength; middle festoon yel- lowish, other festoons partly yellowish (clear only in wet spec- times as long as broad, apex bluntly rounded, corona ca. V6 as long as denticle files, dental formula 3/3, denticles in files of imens). Venter (Fig. 4) as illustrated. Genital operculum (Fig. 13, 8-10, small crenulations in several rows posteriorly. 17) large, very short and broad. Spiracular plates (Fig. 14) as Scutum (Fig. 1) broadly oval, ca. 1.3 times as long as broad; illustrated. (for ornamentation, see Color, preceding); anterior emargina- Legs (Fig. 3, 4, 15, 16). Coxae (Fig. 15) I—III similar to those tion deep, broad; scapulae broad, margins gradually diverging of 6. Coxa IV not greatly enlarged, with only 2 spurs, internal to level of coxa IV, thence converging to broadly rounded spur usually narrower and sharper than external spur. Tro- posterior margin; spiracular plates visible dorsally. Eyes flat, chanter I dorsal shield (Fig. 3) and trochanters ventrally as in 6. oval, rarely bordered by 2-3 punctations. Cervical pits short, Claws and pulvilli as in <$. dark, deep, narrowly oval (dark posterior extension of pits not Diagnosis (adults) grooved). Cervical grooves obsolete. Lateral grooves each repre- sented by a line of contiguous or noncontiguous deep, large A moderately large Dermacentor (subgenus Indocentor) [total and medium-sized punctations extending to or beyond scutal length (mm): $ avg. 5.66; 2 avg. 5.71]. Basis capituli cornua short, midlength. Punctations number moderate to many, size large to porose areas (2) small, widely spaced. Scutum (<3) ca. 1.3 times as medium posteriorly, chiefly small anteriorly, irregularly scat- long as broad. Lateral grooves each a line of contiguous or non- tered in enamelled areas, absent on brown areas, few rarely contiguous deep, large and medium-sized punctations. Color (6). present posteriorly on pseudoscutum (punctation numbers and Ivory enamelling covering most of scutum; absent on dark brown distribution highly variable). Festoons number 11. integumental elevations (median line from intercervical area to Venter (Fig. 2, 7-10). Integument (Fig. 2) smooth; setae small, median festoon, 2 paramedian lines, entire border of pseudo- inconspicuous. Genital area (Fig. 7) as illustrated. Spiracular platesscutum), median festoon, festoons 3 and 4 (in part), narrow (Fig. 8) irregularly pyriform; dorsal projection bluntly rounded intercervical areas (variable), borders of cervical grooves (vari- apically; goblets minute, numerous. Legs (Fig. 1, 2, 9, 10) large, able), and irregular marginal area between eyes and festoons. robust, ivory-enamelled dorsally. Coxa I (Fig. 9) spurs short (as Scutum (2) ca. 1.1 times as broad as long, posteriorly broadly illustrated), length subequal, distinctly but not widely separated, rounded; extensive ivory enamelling (except in brown area bor- external spur narrower than broadly triangular internal spur. dering eyes, a narrow line peripherally posterior to eyes, in II and III spurs each number 2, small; external spur triangular, median line, and along cervical grooves). Dorsum integument (2) internal spur broader, less pointed. Coxa IV (Fig. 9) (except in punctations numerous, large, each with a single, moderately unusually small or teneral specimens) greatly enlarged, ca. 1.5 long white seta; several along lateral grooves, few on lateral times as broad as long; internal margin generally straight; spurs margin and festoons. Color (2) brown except in a short, pale number 3-6, sizes and outlines variable, those between internal yellowish marginal band slightly posterior to body midlength; and external spurs more or less closely spaced. Trochanter I middle festoon yellowish, other festoons partly yellowish (colors 1984 Wassef & Hoogstraal: Dermacentor {Indocentor) auratus identity 171 Downloaded from https://academic.oup.com/jme/article/21/2/169/2220238 by guest on 25 September 2021

FIG. 1-4. Dermacentor {Indocentor) auratus, Malaya (HH77.470): 1, 6 dorsal view; 2, $ ventral view; 3, 2 dorsal view; 4, 2 ventral view. 172 J. Med. Entomol. Vol. 21, no. 2 Downloaded from https://academic.oup.com/jme/article/21/2/169/2220238 by guest on 25 September 2021

13 15 14 1984 Wassef & Hoogstraal: Dermacentor (Indocentor) auratus identity 173

spurs number more than 2 (3-6); and ornamen- tation pattern bright in each sex. These critical differential characters are constant and clearly dis- cernible in well-preserved specimens that are not abnormally small.

Acknowledgments. For facilities to study D. (I.) auratus type materials, we are grateful to Professor Enrico Tortonese (ex Director) and to Dr L. Capocaccia (Director in Charge) of the Museo Civico di Storia Naturale "Giacomo Doria," Genoa. For the SEM illustration (Fig. 17) we are grateful to Professor M.A. Roshdy, Department of Biology, Faculty of Science, King Ab- dulaziz University, Jeddah, Saudi Arabia. Downloaded from https://academic.oup.com/jme/article/21/2/169/2220238 by guest on 25 September 2021

LITERATURE CITED Anastos, G. 1950. The scutate ticks, or Ixodidae, of Indonesia. Entomol. Am. n.s. 30: 1-144. Arthur, D.R. 1960. Ticks. A monograph of the Ixodoidea. Part V. On the genera Dermacentor, Anocentor, Cosmiomma, Boophilus and Margaropus. Cambridge University Press, New York. 251 p. Domrow, R. & M. Nadchatram. 1963. Two field collections of Malayan ticks and mites. Malay. Nat. J. 17: 145-64. Hoogstraal, H. & H.Y. Wassef. 1984. Dermacentor {Indocentor) compactus (Acari: Ixodoidea: Ixodidae): wild pigs and other hosts and distribution in Malaysia, Indonesia, and Borneo. J. Med. Entomol. 21: 174-78. Krijgsman, B.J. & S.A.S. Ponto. 1932. De teken van den Oost- Indischen Archipel. Veeartsenijk. Meded. Dep. Landb. Ned.- Indie,Java, No. 79, 62 p. Neumann, L.G. 1897. Revision de la famille des Ixodides (2e memoire). Mem. Soc. Zool. Fr. 10: 324-420. FIG. 17. Dermacentor {Indocentor) compactus, Malaya 1902. Notes sur les ixodides. Arch. Parasitol. 6: 109-28. (HH78.578). Scanning electron microscope view of 9 external 1905. Notes sur les ixodides. III. Arch. Parasitol. 9: 225-41. genitalia and coxae I to IV. 70 x. 1911. Ixodidae. No. 26. Tierreich, Berlin. 169 p. Schulze, P. 1933. Die Arten der Zeckengattung Dermacentor s. 1. aus Europa, Asien, und Neu-Guinea. Z. Parasitenkd. 6: clear only in wet specimens). Coxa I (<5, 2) spurs short, subequal, 416-31. distinctly but not widely separated, external spur narrower than 1939. Zur Zeckenfauna Burmas. Z. Parasitenkd. 10: 722-28. broadly triangular internal spur. II and III spurs (and 9 IV spur) Seneviratna, P. 1965. The Ixodoidea (ticks) of Ceylon. Parts each number 2, shorter than I spurs. Coxa IV {$) greatly en- II and III. Ceylon Vet. J. 13: 28-54. larged, ca. 1.5 times as broad as long; spurs number 3-6, size Sharif, M. 1928. A revision of the Indian Ixodidae with special and outline variable, inner spurs more or less closely spaced reference to the collection in the Indian Museum. Rec. In- between internal and external spurs. External genitalia (9) large, dian Mus. 30:217-344. very short and broad. Supino, F. 1897. Nuovi Ixodes della Birmania. R. Stabilimento P. Prosperini, Padova. 38 p., pi. 1-12. DISCUSSION AND CONCLUSIONS Toumanoff, C. 1941. Etude descriptive et iconographique de quelques tiques de l'lndochine. Rev. Med. Fr. Extreme-Orient Dermacentor (Indocentor) auratus is a distinctive 19: 478-505. species easily recognized by the following combi- 1944. Les tiques (Ixodoidea) de l'lndochine. Instituts Pasteur de nation of characters: female external genitalia large, l'lndochine, Saigon. 220 p. Wassef, H.Y. & H. Hoogstraal. 1983. Dermacentor {Indocentor) very short, broadly rounded; coxa I spurs short, compactus (Acari: Ixodoidea: Ixodidae): identity of male and distinctly but not widely separated; male coxa IV female./ Med. Entomol. 20: 648-52.

FIG. 5-16. Dermacentor {Indocentor) auratus, Malaya (HH77.470): 5, 6 capitulum dorsal view; 6, 6 capitulum ventral view; 7, 6 genital area; 8, $ spiracular plate (A = anterior, D = dorsal); 9, <5 coxae and trochanters I-IV; 10, 6 tarsi I-IV external view; 11, 2 capitulum dorsal view; 12, 9 capitulum ventral view; 13, 2 genital area; 14, 2 spiracular plate; 15, 2 coxae and trochanters I-IV; 16, 9 tarsi I-IV external view.