Miscel.lania Zooloqica 22.2 (1999) 2 1 Ectoparasites on (Gamasida, Ixodida, Diptera) in Biscay (N lberian peninsula)

E. Imaz, J. R. Aihartza & M. J. Totorika

Imaz, E., Aihartza, J. R. & Totorika, M. J., 1999. Ectoparasites on bats (Gamasida, Ixodida, Diptera) in Biscay (N lberian peninsula). Misc. Zool., 22.2: 21-30.

Ectoparasites on bats (Gamasida, Ixodida, Diptera) in Biscay (N lberian peninsula).- A study on ectoparasites infesting Chiroptera in Biscay (N lberian peninsula) was carried out during a distribution survey of bats. 160 potential hosts were examined and 664 ectoparasites were found, collected manually from living bats by means of pointed tweezers. The ectoparasites belonged to 12 and 2 subspecies: 5 species and 2 subspecies of Gamasida, 2 species of lxodida and 5 species of Diptera. First records in the study area were obtained for Eyndhovenia euryalis euryalis, Eyndhovenia euryalis oudemansi, Argas vespertilionisa n d Penicillidia dufouri. plecotina on Rhinolophus ferrumequinum and Rhinolophus euryale and lxodes vespertilionis on Myotis nattereri are reported for the first time in the lberian peninsula; nattereri is new on Myotis nattereriin Biscay. Associations between parasites and hosts are also reported.

Key words: Chiroptera, Gamasida, Ixodida, Diptera, N lberian peninsula.

(Rebut: 13 X 98; Acceptació condicional: 2 11 99; Acc. definitiva: 2 1 XII 99)

E. Imaz, J. R. Aihartza & M. J. Totorika, Zoologia eta Animali Zelulen Dinamika Saila, Euskal Herriko Unibertsitatea, 644 p. k., E 48080, Bilbo, Espana (Spain).

O 1999 Museu de Zoologia 22 Imaz et al.

Introduction to the following families of Arthropoda: (, Gamasida), Most papers on ectoparasites are de- (Acari, Ixodida), (Acari, Ixodida) scriptive and about most groups little is and Nycteribiidae (Diptera). Different taxa known about their world-wide distribution found on bats and associations between and biology and/or ecology. The few studies parasites and hosts are reported. in the lberian peninsula are fragmentary and the only records on bat ectoparasites in Bis- cay were assembled during the 50s and 60s Material and methods by Dr. BALCELLS (1968), some of which were later reexamined by ESTRADA-PErZJA et al. (1988, During a distribution survey 12 bat species 1989, 1990, 1991). were examined for ectoparasites. Batflies This paper shows the results of a survey (Diptera) were collected from 160 individu- carried out in Biscay. Studied groups belong als from July 1994 to June 1995. (Acari,

Table 1, Number of examined bats and number of associated ectoparasites. Ectoparasites: EEE. E. e. euryalis; EEO. E. e. oudemansi; EES. E. e. subspp; SPL. S. plecotína; SMY. S. myotí; SPS. S. Psi; PRH. rhinolophinus; IVE. /. vespertilionis; AVE. A. vespertilionis; NBI. N. biarticulata; NSC. N. schmidii; BNA. 13. nana; BNT. B. nattereri; PDU. P dufouri. Hosts: Rhf. R. ferrumequinum; Rhe. R. euryale; Rhh. R. hipposíderos; Pp. P. pipistrellus; Pn. P. nathussil; Pk. P. kuhlii; Ms. M. screíbersii; Mm. M. myotis; Me. M. emarginatus; Mn. M. natteril; Paur. P. auritus; Paus. P. austriacus. Nb. Number of bats examined; Np. Number of parasites found. Número de murciélagos estudiados y número de ectoparésitos asociados. (Para las abreviaturas ver arriba.)

Ectoparasites Arachnida Insecta Gamasida Ixodida Diptera Hosts Nb Np EEE EEO EES SPL SMY SPS PRH IVE AVE NBI NSC BNA BNT PDU Rhf 24 65 5 30 3 9 0 0 7 4 0 7 0 0 0 0 Rhe 23 44 14 0 2 1 0 0 0 0 0 24 3 • 0 0 0 Rhh 50 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Pp 26 20 4 0 0 0 0 0 0 0 20 0 0 0 0 Pn 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Pk 19 11 0 0 0 0 0 0 0 0 11 0 0 0 0 0 Ms 33 273 2 0 0 0 0 251 0 0 0 0 19 0 0 1 Mm 2 34 0 0 0 0 33 0 0 0 0 0 0 0 0 1 Me 5 2 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Mn 4 6 0 0 0 0 0 Q 0 1 0 0 0 3 2 0 Paur 12 3 0 0 0 3 0 0 0 0 0 0 0 0 0 0 Paus 6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Total 160 458 21 30 5 13 33 251 7 5 31 31 22 3 2 4 Miscel.lania Zoologica 22.2 (1999)

Garnasida) and (Acari, Ixodida) were sampled in 135 potential hosts frorn January to June in 1995. When more than one bat was captured at the sarne time they were kept in separate bags. Ectoparasites were collected rnanually frorn living bats by rneans of pointed tweezers. Consequently, only the House; Mt. ~ihetunnel most conspicuous parasites on fur and the Ob. Other buildings. largest on the patagium were removed. Bats Localidades donde se were then released. Sarnples were stored in ethyl acetate and identified under rnicro- scope. Even if the sarnpling rnethod is biased an equal bias for al1 the species is accepted. Thus, we calculated associations between parasites and hosts frorn data in table 1, excluding bat species on which no parasites were found, by rnean of Pi-square rneasure to estimate similarity and the unweighted pair-group rnethod (UPGNA) for clustering Localitv Tv~e Townshi~ UTM (No~usis,1993).

Results

A total of 664 ectoparasites were collected, frorn which 12 species and 2 subspecies were identified: 5 species and 2 subspecies of Gamasida, 2 species of lxodida and 5 species of Diptera (table 1). Besides groups 0056 C Karrantza VN68 studied here, rnany rnacronyssids (Garnasida, Macronyssidae) and fleas (Siphonaptera, Ischnopsyllidae) were also collected but have not been included in this work. In- forrnation about each locality is shown in table 2.

Class Arachnida 0243 Ch Atxondo WN37 Order Parasitiforrnes Suborder Garnasida (= Mesostigrnata) Spinturnicidae

Genus Eyndhovenia Rudnick, 1960 0292 Ch Lernoiz WPOO Eyndhovenia euryalis (Canestrini, 1884)

A total of 56 individuals of this species was collected. Five of thern could not be identified at subspecies level, dueto hard asyrnrnetry on setae distribution and their poor preservation status. Three of them were collected from one Rhinolophus ferrumequinum (Schreber. 1774) in cave No 0028, where colonies of R. lmaz et al. ferrumequinum, Rhinolophus euryale Blasius, (genus Myotis) were exarnined, it was only 1853, Miniopterus schreibersii (Kuhl, 1817) found occurring on one Myotis myotis and Myotis emarginatus (E. Geoffroy, 1806) (Borkhausen, 1797). can be found. Two more were found on R. euryale in roost No 0006; this roost is used by Spinturnix psi Kolenati, 1856 small breeding colonies of R. ferrumequinum and M. emarginatus, but R. euryale is only an A total of 251 individuals were collected, al1 occasional visitor. occurring on Miniopterus schreibersii. All hosts were caught from two ternporary colo- Eyndhovenia euryalis euryalis (Canestrini, 1884) nies in caves No 0004 and 0031.

This subspecies is recorded for the first time in Biscay, where 21 individuals were found. Genus Paraperiglischrus Rudnick, 1960 R. euryale seems to be its main host (see table l), even though al1 the individuals car- Paraperiglischrus rhinolophinus Koch, 1841 rying this parasite were caught in the same temporary colony, in cave No 0004, which is It is recorded for the first time in Biscay. also used by R. ferrumequinum, M. schreibersii Seven specimens were collected, al1 from a and M. emarginatus. It was also collected single sample of the 44 R. ferrumequinum from R. ferrumequinum in cave No 0028, and host species checked. It was found in an old from M. schreibersii in cave No 0031, where farm house, No 0006, where srnall colonies of this species forms temporary groups. R. ferrumequinum and M. emarginatus breed.

Eyndhovenia euryalis oudemansi (Eyndhoven, 1941) Suborder lxodida (= Metastigmata) Family lxodidae This species is reported on R. ferrumequinum, on which it was found in 30% of checked Genus lxodes Latreille individuals. Recorded for the first time in Biscay, where it shows a high specificity for lxodes (Eschatocephalus) vespertilionis Koch, this bat species (table 1). Found in the attic 1844 of church No 0266, in cave No 0028 and in a small breeding colony in farm house No0006. This parasite was found occurring on R. ferrumequinum and Myotis nattereri (Kuhl, 1817). The latter is recorded for the first time Genus Spinturnix Von Heyden, 1826 in the lberian peninsula. All these records were obtained from hosts caught in under- Spinturnix plecotina Koch, 1839 ground roosts: caves No0295,0297 and 0028, and mine tunnel No 0055. This species was collected on Plecotus auritus (Linnaeus, 1758) in churches No 0243 and 0326, and on R. ferrumequinum in church Family Argasidae No0266, and from R. euryale in cave No 0004. Horseshoe bats are new hosts in the lberian Genus Argas Latreille, 1796 peninsula. Argas (Carios) vespertilionis (Latreille, 1802) Spinturnix myoti Kolenati, 1856 These parasites were collected frorn The myoti species group is conflictive within Pipistrellus pipistrellus (Schreber, 1774) and the Spinturnix genera (UCHIKAWAet al., 1994). Pipistrellus kuhlii (Kuhl, 1819). Hosts were The species was identified according to the caught both in roosts and foraging sites, description given by UCHIKAWAet al. (1994) localities No 0234, 0288, 0292, 0340 and for S. myoti. 0336 for P. pipistrellus, and No 0090 and Although sorne other mouse-eared bats 0358 for P. kuhlii. Miscel.lania Zoologica 22.2 (1 999) 2 5

Class lnsecta (= Hexapoda) caught in caves 0004 and 0051, and the latter Order Diptera in buildings number 0006 and 0071. Farnily Nycteribiidae

Genus Nycteribia Latreille, 1796 Affinities between species

Nycteribia biarticulata Herrnann, 1804 Figure 1 shows the affinities between para- site species depending on host species. They had been collected occurring on R. A preliminary view enables identification of ferrumequinum and R. euryale. For the forrner a large block of parasites which includes the host species, this parasite was found in first seven taxa arranged in the cluster. The 6 roosts, four caves and two breeding places first group is forrned by Eyndhovenia euryalis in houses, localities No 0006, 0028, 0066, 0272, oudemansi and Paraperiglischrus rhinolophinus, 0295, and 0297. It was also collected from R. with high affinity, followed by Spinturnix euryale in three caves and one house, plecotina and lxodes vespertilionic. The sec- No 0004,0006,0056 and 0066. ond group in this big block is formed by Eyndhovenia euryalis euryalis, Nycteribia Nycteribia (Listropodia) schmidli Scheiner, 1853 biarticulata and Eyndhovenia euryalis subspp. Rernaining parasite taxa do not seern to Almost al1 of thern were collected from M. be so clearly inter-related further than at level schreibersi. Three specirnens found occurring of small groups. Thus, Basilia nana and Basilia on one R. euryale can not be considered as nattereri appeared very close. Nycteribia valid data, as this bat was not isolated after schmidli is also linked to Spinturnix psi but capture, but was in contact with other the affinity is lower. Finally, Spinturnix myoti Schreiber bats. Thus it can be considered as and Penicillidia dufouri are not clearly in- a casual parasitisrn. M. schreibersii bearing cluded in any group, and Argas vespertilionis this species were caught in three temporary appears notably alone. roost caves, No 0004, 0028 and 0031. The Figure 2 shows association between host specimen of R. euryale was taken in cave species depending on their parasites. Affin- N00066, which is also used as a temporary ity is low except for P. pipistrellus and P. roost by M. schreibersii. kuhlii. The second group is constituted by R. ferrumequinum, P. auritus and R. euryale, and the third by M. myotis, M. emarginatus Genus Basilia Ribeiro, 1903 and M. schreibersii with lower affinity. Fi- nally, M. nattereri appears alone. Basilia nana Theodor & Moscona, 1954

These parasites were collected on M. nattereri Discussion in two artificial underground sites, No 0055 and 0073. used as occasional roosts. These data confirm the presence of five spe- cies and two subspecies of Garnrnasids in Basilia nattereri Kolenati, 1857 Biscay, of which one species and one sub- species are recorded for the first time. Arnong Collected frorn M. nattereri, this is a new thern, Eyndhovenia euryalis is a typical para- host for this batfly in the Biscay locality No site of horseshoe bats (genus Rhinolophus) 0055. in Europe, in which three subspecies have been described according to different rneas- urernents for different forrns and rnorpho- Genus Penicillidia Kolenati. 1863 logical aspects related with the idiosoma, dorsal shield, seta, peritrerna, tritosternum, Penicillidia dufouriWestwood, 1835 sternal shield, and legs (UCHIKAWA& DUSBABEK, 1978): E. euryalis euryalis, E. euryalis Collected on M. schreibersii, M. myotisand M. oudemansi, and E. euryalis cornuti. Only the emarginatus. The forrner two host species were first two were found in this work. Regarding 2 6 lrnaz et al.

O 5 10 15 20 25 Label+------+------+--I------.+-----.---+-.---.--- +

EEO PRH IVE SPL EEE NBI I EES

AVE

the first, our data agree with previous works (Schreber, 1774), Eptesicus nilssonii (Keyserling stating horseshoe bats are the usual host of et Blasius, 1839), Nyctalus noctula (Schreber, E. euryalis euryalis in other countries of Eu- 1774), Myotis nattereri, Myotis mystacinus rope and the lberian peninsula, where it (Kuhl, 1817), Myotis brandtii (Eversrnann, also occurs on M. schreibersii (DEUNFF,1977; 1845) and Myotis daubentonii (Kuhl, 1817) UCHIKAWA& DUSBABEK,1978; PERIBANEZ-L~PEZet (DEUNFF, 1977; ESTRADA-PENAet al., 1989; al., 1989). However, E. euryalis oudernansi STANJUKOVICH, 1990). Furthermore, Spinturnix was found only on R. ferrurnequinurn. This rnyoti was previously found in al1 checked also agrees with other authors'data for Eu- Myotis species in the lberian peninsula rope and the lberian peninsula (UCHIKAWA& (ESTRADA-PENAet al., 1991). as well as in gen- DUSBABEK,1978; PEREIRA,1989; ESTRADA-PENAet era Barbastella, Eptesicus, Vespertilio, al., 1991), and it differs considerably frorn Plecotus, Rhinolophus and Pipictrellus in the North African pattern, where E. e. other European areas (DEUNFF,1977; STANJU- oudernansi was also reported on R. mehelyi KOVICH, 1990). Lastly, Spinturnix psi, appears and R. euryale (ESTRADA-PENAet al., 1991). specific for M. schreiberssi, and even indi- Three species of genus Spinturnix were vidual~of R. euryale caught in the same confirmed in Biscay S. plecotina, S. rnyoti and roosts were free of it. This agrees with data S. psi, whose presence was unclear from the from other authors in Europe (RUDNICK,1960; bibliography available (ESTRADA-PENAet al., BERON,1971; DEUNFF,1977; ESTRADA-PENAet al., 1989; ESTRADA-PE~Aet al., 1991; CORDERODEL 1989, 1991; PERIBANEZ-LÓPEZet al., 1993), and CAMPILLO~~al., 1994). On the first hand, horse- although this parasite has been found on shoe bats are new hosts for S. plecotina in Pipistrellus, Rhinolophus and Myotis bat spe- the lberian peninsula. P. auritus is its rnost cies, these cases were considered occasional usual host in Europe, although it occurs on contarnination (DEUNFF,1977). R. ferrurnequinurn, Eptesicus serotinus The last garnrnasid, Paraperiglischrus Miscel.lania Zoologica 22.2 (1999) 27

Rf Paur I Re Mn

rhinolophinus, which was found only on R. Europe on M. schreibersii but is rare in the ferrumequinum during this work, has been lberian peninsula (ARTHUR,1956; BEAUCOURNU, reported showing high specificity as a para- 1967; ESTRADA-PENAet al., 1991).Argas (Carios) site of other horseshoe bats (DEUNFF,1977; vespertilionis, recorded for the first time in ESTRADA-PENAet al., 1989; CORDERODEL CAMPILLO Biscay, is a widely distributed species (Eu- et al., 1994). Some lack of observations about rope, Asia and Africa), which prefers this species could happen due to the more pipistrelle bats but can occur on al1 bat spe- inexhaustive examination methods of lat- cies except on horseshoe bats (BEAUCOURNU, eral and wing membranes. lndividuals of 1961; CORDERODEL CAMPILLOet al., 1994; this species, mainly females and nymphs, pre- DUSBABEK & ROSICKY,1976). It can also occur fer to be on tail membranes (DEUNFF& on people in human buildings when no bat BEAUCOURNU,1981). host is present (COLAS-BELCOUR,1933; HOOSTRAAL, Among Ixod ida, lxodes (Escha tocephalus) 1956). vespertilionis was reported previously occur- Concering batflies, the occurrence of ring on R. ferrumequinum and P. auritus in Nycteribia biarticulata on R. ferrumequinum Biscay (ESTRADA-PENAet al., 1989; CORDERODEL and R. euryale agrees with data from other CAMPILLOet al., 1994), where it does not seem countries, where this was also reported to be an abundant species. R. ferrumequinum on other horseshoe bats, but mainly on these has been considered its main host in Europe, two species (THEODOR& MOSCONA,1954; but it has been detected on several bat spe- BALCELLS,1968; KRISTOFIK,1982; ESTRADA-PENAet cies, showing high infestation level in some al., 1991; CORDERODEL CAMPILLOet al., 1984). In cases (ARTHUR,1963; BEAUCOURNU,1967; HUTSON, the same way, the high amount of Nycteribia 1971; ESTRADA-PENAet al., 1989; CORDERODEL schmidli found over M. schreibersii agrees CAMPILLOet al., 1994). Although it had also with the general view that considers this been considered as a "winter species" batfly as a typical constituent of this bat (BEAUCOURNU,1967) only five individuals were parasite fauna, even if occasionally small collected from February to June. No lxodes numbers are found on other species (THEODOR simples Neumann, 1906, was recorded. This & MOSCONA,1954; KRISTOFIK,1982; ESTRADA-PENA parasite is widespread in Central and Eastern et al., 1991). 28 Imaz et al.

The first record of Basilia nana in Biscay euryalis and Nycteribia biarticulata, which and its occurrence on M. nattereri, agrees following our data can be identified as the with previous knowledge about this parasite parasite group of R. euryale, though they in the lberian peninsula, where this bat spe- are also found also in other rhinolophids in cies is its main host. It has also been re- Europe (THEODOR MOSCONA, 1954; BALCELLS, corded occurring on Myotis bechsteini 1968; DEUNFF, 1977; UCHIKAWA DUSBABEK, 1978; (Bechsteins bat), Myotis dasycneme (pond KRISTOFIK, 1982; CORDERO DEL CAMPILLO et al., bat), Myotis blythi (lesser mouse-eared bat), 1984; PERIBANEZ-LOPEZ et al., 1989; ESTRADA-PENA Myotis myotis, Myotis emarginatus(Geoffroys et al., 1991). bat), R. ferrumequinum, P auritus as well as About the remainder, the only consistent on M. schreibersii (THEODOR MOSCONA, 1954; group is formed by Nycteribia schmidli and HUTSON, 1984). BEAUCOURNU (1961) noted M. Spinturnix psi, which are considered as the bechsteini as its main host in Western France. typical parasite fauna of M. schreibersii (THEODOR On the other hand, the occurrence of Basilia MOSCONA, 1954; KRISTOFIK, 1982; ESTRADA-PENA nattereritogether with B. nana on the same et al., 1991). The higher affinity of Basilia nana individual of M. nattereri shows an interme- and Basilia nattereri is explained by the scar- diate position between previous northern city of data. Even if they have only been found and southern records. In fact, M. nattereri is on M. nattereri during this work, they are a common host of Basilia nattereri in Central known to occur on several bat species in Eu- Europe where it also infests other small Myotis rope (THEODOR MOSCONA, 1954; HUTSON, 1984; bats and E. serotinus but on the lberian BEAUCOURNU, 1961). The same occurs with peninsula this bat is usually parasited by Spinturnix myoti and Penicillidia dufouri, which Basilia nana (ESTRADA-PENA et al., 1991). are not clearly included in any group. And the The last species, Penicillidia dufouri, is re- last case, Argas vespertilionis, which in this corded first on M. emarginatus in the lberian work was found only on Pipistrellus, can also peninsula. It seems to be a nonspecific parasite, be found on other vespertilionids but it shows since it was previously recorded mainly on M. preference for this bat genus (BEAUCOURNU, myotis and M. blythii, but also on other several 1961; CORDERO DEL CAMPILLO et al., 1994; DUSBABEK species such as M. nattereri, Myotis capaccini, R. ROSICKY, 1976). mehelyi, R. euryale, R. ferrumequinum, E. About the affinities between host spe- serotinus and M. schreibersii (FALCOZ, 1923; cies, the first group formed by P. pipistrellus BALCELLS, 1968; KRISTOFIK, 1982; ESTRADA-PENA et al., and P. kuhlii and the high affinity it shows 1991; CORDERO DEL CAMPILLO et al., 1994). is a consequence of the ixodid Argas It is difficult to draw conclusions concern- vespertilionis, which has a documented pref- ing the affinities between the parasite spe- erence for this host genus (BEAUCOURNU, 1961; cies and host species shown in figures 1 and CORDERO DEL CAMPILLO et al., 1994; DUSBABEK 2 as there is an insufficient quantity of data ROSICKY, 1976). The second group includes M. available for many parasite and host species. myotis, M. emarginatus and M. schreibersii, In figure 1, the large block formed by the but it could be the consequence of data first seven taxa is related to parasites of horse- scarcity, specially about the first two bat shoe bats (genus Rhinolophus), though species. In fact, the parasite species that can Spinturnix plecotina is known as parasite on explain this grouping in our data is P. several bat species, and its occurence in this dufouri, and many host species have been group can be an artifact (DEUNFF, 1977; recorded for it in previous works. Moreover, ESTRADA-PENA et al., 1989; STANJUKOVICH, 1990). despite occasional cases, M. schreibersii has The highest affinity between Eyndhovenia an own parasite fauna, including species euryalis oudemansi and Paraperiglischrus such as S. psi and N. schmidli. Thus, we can rhinolophinus reflects that they were re- suppose that more data about ectoparasites corded only on R. ferrumequinum, but only of M. myotis and M. emarginatus would the former has a high specifity for this host, change this grouping. The third group, while the latter is a typical parasite of horse- formed by R. ferrumequinum, P. auritus and shoe bats (DEUNFF, 1977; ESTRADA-PENA et al., R. euryale, to be more clear, being a conse- 1989; CORDERO DEL CAMPILLO et al., 1994). The quence of typical parasites of horseshoe bats same can be argued for Eyndhovenia euryalis such as N. biarticulata, E. euryalis euryalis, Miscel.lania Zooloqica 22.2 (1999) and S. plecotina, which is the link between biarticulata, N. schmidli, Basilia nana, B. rhinolophids and P. auritus. Finally, the se- nattereri observado por primera vez sobre clusion of M. nattereri must be explained in M. nattereri en Bizkaia, y Penicillidia dufouri the same way as for the other Myotis spe- primera cita en Bizkaia. También se comenta cies, as a consequence of the scarcity of la afinidad entre especies de parásitos data, and more information is needed be- dependiendo de las especies de murciélagos fore a clear view about their affinity pat- que infestan (fig. 1) y la afinidad entre tern is obtained. especies de murciélagos dependiendo de las especies de parásitos que soportan (fig. 2).

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