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Systematic & Applied Acarology (2005) 10, 67 74. ISSN 1362-1971 Psorergatoides nyctali (Prostigmata: Psorergatidae), a new mite species parasitizing the bat Nyctalus noctula (Mammalia: Chiroptera) in the British Isles ANNE S. BAKER Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK; e-mail: asb@nhm.ac.uk Abstract Psorergatoides nyctali sp. nov. is described from the wing membrane of a captive adult female Nyctalus noctula originating from Devon, England. Key words: Psorergatoides nyctali, new species, British Isles, bat, Nyctalus noctula Introduction Members of the genus Psorergatoides Fain (Prostigmata: Psorergatidae) are exclusively parasites of bats (Mammalia: Chiroptera), and are distinguished from other psorergatids by the presence of four pairs of minute lateral setae on the dorsal shield (Fain 1959a,b). Of the 17 species described previously, one has been recorded from each of the host families Desmodontidae (Lukoschus et al. 1979), Hipposideridae (Fain 1959b), Megadermatidae (Fain 1959a), Nycteridae (Fain 1959a) and Rhinolophidae (Fain 1959a,b), two from Molossidae (Lukoschus et al. 1973, Giesen et al. 1982b) and Vespertilionidae (Giesen et al. 1982a), four from Emballonuridae (Fain 1959b, Lukoschus et al. 1973, Lukoschus et al. 1979) and five from Phyllostomidae (Fain 1959b, Lukoschus et al. 1973, Lukoschus et al. 1979). Only one species, P. lonchorhina Fain, has been found on members of more than one host family (Fain 1959b, Giesen 1990). Psorergatoides species have been collected in the Australian, Oriental, Palaearctic, Ethiopian and Neotropical regions, with the majority of records emanating from the latter two. Psorergatoides kerivoulae Fain and P. rhinolophi Fain are the only Palaearctic representatives, respectively identified from Plecotus auritus (Linnaeus) in Belgium (Fain 1959b), and Rhinolophus species in Belgium, France, Italy and Spain (Fain 1959b, Lukoschus et al. 1973). In October, 2002, the carer of a captive adult female Nyctalus noctula (Schreber) (Chiroptera: Vespertilionidae) noticed unpigmented patches developing on both wings of the bat (Fig. 1). The patches slowly spread, and were observed to incorporate small cysts, and the finger joints became inflamed. The host was apparently not distressed by its condition and irritation did not seem to be a problem. Such symptoms were not exhibited by two other noctules sharing the same enclosure as the affected bat. Using a six millimetre skin biopsy punch, a cyst was removed from the anaesthetized host by a veterinary surgeon and examined. The skin showed marked epidermal hyperplasia and mild lymphoplasmacytic superficial dermatitis, with abundant intra-epithelial cross sections of mites (P. Briggs, pers. comm.). Two other cysts removed by the same method were examined by the present author and found to contain an unnamed species of Psorergatoides (Fig. 2). Psorergatoides nyctali sp. nov. is here described. Observations were made on specimens temporarily mounted in 60% lactic acid. Measurements are given in micrometres (µm) and as a mean followed by the range or as a range only. Because of 2005 Systematic & Applied Acarology Society 67

the difficulty of measuring the long setae due to their curvature and finely tapering ends, values given for these are somewhat approximate. Morphological terminology follows that of Giesen (1990). The description of the male includes only those character states that differ from the female. FIGURE 1. Psorergatoides nyctali sp. nov. Extended wing of host (Nyctalus noctula) showing unpigmented areas of infestation. FIGURE 2. Psorergatoides nyctali sp. nov. Dissected cyst, mite displaced from indentation arrowed in black, separated mite arrowed in white. Psorergatoides nyctali sp. nov. (Figs. 3 17) Psorergatoides sp. A, Baker & Craven, 2003: 13. Material examined Holotype female: ENGLAND, Hertfordshire, Bushey, captive adult female Nyctalus noctula (Chiroptera: Vespertilionidae), ex cyst in wing membrane (host rescued on 14.viii.1997 at 68 SYSTEMATIC & APPLIED ACAROLOGY VOL. 10

Dartington Hall, near Totnes, Devon, England), coll. P. Briggs, x.2002. Paratypes (20 females, 5 males, 2 deutonymphs, 7 protonymphs): same data as holotype. [Specimens mounted in Hoyer s medium. Deposited in the Natural History Museum, London; acc. no. BMNH(E)2005-143.] FIGURES 3 7. Psorergatoides nyctali sp. nov. (female). 3, dorsal view; 4, ventral view; 5, gnathosoma, dorsal view; 6, fixed digit of chelicera, lateral view; 7, palp tibiotarsal setae, lateral view. al = adanal lobe, go=genital opening. 2005 BAKER: PSORERGATOIDES NYCTALI SP. NOV. PARASITIZING BAT NYCTALUS NOCTULA 69

Diagnosis The female of Psorergatoides nyctali sp. nov. is distinguished from that of the other members of the genus by the presence of two setae on femora I III, and usually on at least one femur IV, in combination with terminal setae h1 2 being longer than 75µm and the tarsal spine being pointed. Males differ because they lack leg trochanteral and femoral setae. FEMALE (10 SPECIMENS MEASURED) Total length (idiosoma and gnathosoma) 151(140 170); maximum idiosomal width (ca. at level of legs III) 141(133 150). Idiosoma dorsum (Fig. 3). Integument surrounding dorsal shield striated. Shield punctate with irregular outline, median crease extends from posterior margin almost to level of setae d; median length 116(111 123), greatest width (ca. at level of d) 121(113 128); 5 pairs minute setae present (ve, sc i, c, d, e), ve located slightly posteromedially to lateral seta sc i. Idiosoma ventral (Fig. 4). Integument smooth. Epimera I curve outwards distally and with circular sclerotization associated with curved part, epimera II IV more or less straight. Ventral setae (ic III) 6 8 long, inserted 17 23 apart. Genital opening (go) just anterior to adanal lobes (al). Adanal lobes each bear 2 smooth, subequal tapering setae (h1 & 2), 90 112 long. Gnathosoma (Figs 5 7). Internal subcapitular setae (pmc 1) minute. Supracoxal seta (ep) twosegmented, ends in 3 prongs, ca. 6 long. Palp: dorsal setae ats spiniform, ca. 2 long, dg a broad spine, vg minute, df serrate, 21 24 long; two ventro-apical tibiotarsal setae claw-like and bifid, one spiniform (Fig. 7). Chelicera: fixed digit with 7 teeth, proximal 1 and distal 2 larger than median 4 (Fig. 6). Legs (Figs 8 11). Trochantera with one anterobasal and one small posterodistal spur ventrally, femora with posterobasal spur. Setal formulae (trochanter-tarsus) (solenidion in round, variations in square brackets): I 1-2[1]-1-2-4(1); II 1-2-1-2-4(1); III 1-2[1]-1-2-3; IV 1-2[1,0]-1-2-2; in 19 specimens examined, femoral setal variations on legs I, III and IV occurred as follows [I/I, III/III, IV/IV (number of specimens)]: 2/2, 2/2, 2/1 (5); 2/1, 2/2, 2/1 (4); 2/2, 2/2, 2/2 (3), 2/2, 2/2, 1/1 (2); 2/1, 2/2, 2/0 (1); 2/1, 2/2, 1/0 (1); 1/1, 2/2, 1/1 (1); 2/1, 2/2, 1/1 (1); 2/2, 2/1, 2/1 (1). Trochanteral seta 11 15 long. Proximal femoral seta ca. 0.7 0.8 length distal, respectively 14 18 and 17 22 long. Posterolateral genual seta I III spiniform, 2 3 long, that of IV slender, 17 19 long. Ventrolateral tibial seta robust, spiniform, mediodorsal seta slender, 20 30 long. Tarsi with ventral robust spiniform seta; I III also with 2 slender dorsolateral setae (tc', tc"), 15 23 long on I III; tc' of IV 14 17 long, tc" absent; I and II with enveloped seta (p"@) and 1 solenidion (ω). Apotele with two subequal pointed claws and membranous empodium, comprising small dorsal and large ventral lobe. MALE (5 SPECIMENS MEASURED) Total length 121(118 125), maximum width 114(108 120). Idiosoma dorsal (Fig. 12). Shield 93(89 95) long, greatest width 102(100 108), without median crease, bears 6 pairs minute setae (ve, ps, sc i, c, d, e); ve located slightly posterior to genital opening, ps near base of aedeagus (ae); distance ve ve and ps ps respectively 19 22 and 11 14. Aedeagus 18 21 long. Idiosoma ventral (Fig. 13). Ventral setae 5 6 long, inserted 19 22 apart; adanal lobes and h setae absent. Gnathosoma. Supracoxal seta 3 4 long. Palp dorsal seta ats ca. 1 long, df 18 20 long. Fixed cheliceral digit appears to have only 3 small median teeth in addition to larger proximal and distal ones. Legs (Figs 12 13). Setal formulae: I 0-0-0-1-4(1); II 0-0-0-1[2]-4(1); III 0-0-0-1[2]-2; IV 0-0- 0-1[1]-2. Mediodorsal tibial seta seen on a single leg in some specimens (present on leg II and III in 70 SYSTEMATIC & APPLIED ACAROLOGY VOL. 10

2 and 1 paratypes respectively, and on leg IV in 1, subsequently lost, specimen), 15 18 long. Dorsolateral tarsal setae (tc', tc") I III 15 18 long, tc' IV 10 14. DEUTONYMPH (2 SPECIMENS MEASURED) Total length 148 150, maximum width 135 143. Idiosoma (Fig.14). Lacks setae. Gnathosoma (Fig. 15). Setae like those of adults in shape. Legs (Figs 16 17). Two-segmented; dorsal setae absent; ventrally, basal segment with proximal spur, distal segment with 2 trifid claws, I and II also with two structures here hypothesized as being homologous to solenidion ω (distal) and enveloped seta p"@ (proximal). FIGURES 8 11. Psorergatoides nyctali sp. nov. (female), legs I IV, ventral view. 8, I, with detail of dorsal view of tarsus; 9, II; 10, III; 11, IV, with detail of femur of opposite leg. 2005 BAKER: PSORERGATOIDES NYCTALI SP. NOV. PARASITIZING BAT NYCTALUS NOCTULA 71

FIGURES 12 13. Psorergatoides nyctali sp. nov. (male). 12, dorsal view; 13, ventral view. ae=aedeagus, go=genital opening. 72 SYSTEMATIC & APPLIED ACAROLOGY VOL. 10

FIGURES 14 17. Psorergatoides nyctali sp. nov. (deutonymph). 14, ventral view; 15, gnathosoma, dorsal view; 16, leg I, ventral view; 17, leg III, ventral view. PROTONYMPH (7 SPECIMENS MEASURED) Total length 107(100 115), maximum width 113(103 120). Otherwise, as for deutonymph. Etymology The species name is derived from that of the host genus. Remarks Psorergatoides nyctali sp. nov. possesses a number of attributes that are unique to the family Psorergatidae and to the genus Psorergatoides. This is the first psorergatid described that, in either sex, lacks leg trochanteral and femoral setae, and in which the mediodorsal tibial seta is only rarely present. Some members of Psorergates Tyrrell and Psorobia Fain have two setae on femur IV, but this character state is not seen in any other member of Psorergatoides. The absence of genual setae is an unusual occurrence within the family, having been recorded in one species of Psorobia and six of Psorergatoides. Of these, only three of the latter (P. emballonurae Fain, P. hipposideros Fain and P. lonchorhina) lack genual setae in the male, but not in the female. Based on Giesen s (1990) phylogeny, P. nyctali sp. nov. is most closely related to P. australiensis Giesen, Lukoschus & Fain, P. kerivoulae, P. nycteris Fain and P. tadaridae Giesen, Lukoschus & Nadchatram, which are the only other members of the genus to have two setae (rather than one) on femora I III in the female. Furthermore, h setae are absent in the male of at least three of these four (P. nycteris males are 2005 BAKER: PSORERGATOIDES NYCTALI SP. NOV. PARASITIZING BAT NYCTALUS NOCTULA 73

unknown), genual seta IV is much longer than that of I III in P. australiensis, P. kerivoulae and P. tadaridae, while P. australiensis and P. kerivoulae were collected from hosts belonging to the family Vespertilionidae (respectively from four species in two genera and six species in three genera (Giesen 1990)) and have h setae that are over 75µm in length. In addition to the unique character states described above, P. nyctali sp. nov. differs from these four species in possessing a tapering, rather than bifid, ventral tarsal spine. No interspecific differences have been identified in Psorergatoides immatures, and protonymphs and deutonymphs are distinguished only according to their size (Giesen, 1990). The two structures on tarsi I and II of nymphal P. nyctali sp. nov., here proposed as solenidion ω and enveloped seta p"@, have been illustrated for other species (e.g., Lukoschus et al. 1973, Lukoschus et al. 1979), but both are referred to as solenidia. Psorergatoides species are evidently highly host specific. Only one of the previously described 17 parasitize more than one bat family, while 13 have been found on a single host genus, three on two genera and one on three. It is likely, therefore, that P. nyctali sp. nov. will be confined to the Vespertilionidae and possibly to Nyctalus. As reported for other members of the genus (Giesen 1990), P. nyctali sp. nov. individuals each lies in an impression in the epidermis (Fig. 2), and adults and immatures occur together in several layers. Acknowledgements Particular thanks go to Patty Briggs, Bushey, Hertfordshire, UK, for sending the cysts containing P. nyctali sp. nov., providing information about their finding, and for allowing the use of her photograph (Fig. 1). I am also grateful to Harry Taylor, Department of Communications & Development, The Natural History Museum, London, UK, for producing the photomacrographic images of the cysts (Fig. 2). References Fain, A. (1959a) Les acariens psoriques parasites des Chauves-souris. III. Le genre Psorergates Tyrell (Trombidiformes Psorergatidae). Bulletin et Annales de la Société Royale Entomologique de Belgique, 95, 54 69. Fain, A. (1959b) Les acariens psoriques parasites des Chauves-souris. IX. Nouvelles observations sur le genre Psorergates Tyrell. Bulletin et Annales de la Société Royale Entomologique de Belgique, 95, 232 248. Giesen, K.M.T. (1990) A review of the parasitic mite family Psorergatidae (Cheyletoidea: Prostigmata: Acari) with hypotheses on the phylogenetic relationships of species and species groups. Zoologische Verhandelingen, 259, 1 69. Giesen, K.M.T., Lukoschus, F.S. & Fain, A. (1982a) Parasites of Western Australia: 15. Psorergatoides australiensis (Acarina: Psorergatidae) from Australian bats. Records of the Western Australian Museum, 9, 315 323. Giesen, K.M.T., Lukoschus, F.S. & Nadchatram, M. (1982b) Three new itch mites of the family Psorergatidae (Acari, Prostigmata) from Malaysian small mammals. Malayan Nature Journal, 35, 315 328. Lukoschus, F.S., Rosmalen, P.G. & Fain, A. (1973) Parasitic mites of Surinam XI. Four new species of the genus Psorergatoides Fain, 1959, (Psorergatidae: Trombidiformes). Tijdschrift voor Entomologie, 116, 63 81. Lukoschus, F.S., Louppen, J.M.W. & Fauran, P. (1979) Parasitic mites of Surinam. 14. New observations on the genus Psorergatoides Fain, 1959 (Psorergatidae: Trombidiformes) with a key to the known species. International Journal of Acarology, 5, 311 324. Accepted: 30 April 2005 74 SYSTEMATIC & APPLIED ACAROLOGY VOL. 10