Orlova M.V. Hystrix It. J. Mamm. (n.s.) 22(1) 2011: 111-127 (Acarina, Gamasides). Gas. Gsl. Spol. ta) parasitic on bats. Univ. Calif. Publ. ent 11: 77-125 Entom 46. A CRITICAL ASSESSMENT OF THE PRESENCE OF Farafonova G.V., Masing, M.V. 1985. Rudnick A. 1960. A revision of the mites BARBASTELLA BARBASTELLUS AND NYCTALUS NOCTULA Findings of nycteribiid flies in the Pri- of family Spinturnicidae. Univ. Cali- baltic, Parazitologiya 19 (4): 317-318 fornia Publ. Entomologia, 17: 157- IN IRELAND WITH A DESCRIPTION OF N. LEISLERI [In Russian]. 283. ECHOLOCATION CALLS FROM IRELAND Medvedev S.G. 1996. Fleas of the fam. Stanyukovich M.K. 1997. Keys to gamasid Ischnopsyllidae (Siphonaptera) on the mites (Acari, Parasitiformes, Mesos- 1# 2#* fauna of Russia and adjacent countries. tigmata, Macronyssoidea et Laelaptoi- DANIEL J. BUCKLEY , SEBASTIEN J. PUECHMAILLE , 3 1-2 Entomol. Rev. 75(2): 438-454 [In Rus- dae) parasitizing bats (Mammalia, Chi- NIAMH ROCHE , EMMA C. TEELING sian]. roptera) from Russia and adjacent Medvedev S.G., Stanyukovich M.K., Ti- countries. Rudolstadter naturhisto- 1Centre for Irish Bat Research, School of Biology and Environmental Science, Science unov M.P., Faraphonova G.V. 1991. rische Schriften 7: 13-46. Centre West, University College Dublin, Belfield, Dublin 4, Ireland Ectoparasite of bats from Russian Far Stanyukovich M.K. 1990. The gamasid 2School of Biology and Environmental Science, Science Centre West, University College East. Parasitologiya 25(1): 27-37 [In mite and argasid ticks of bats from Dublin, Belfield, Dublin 4, Ireland Russian]. Baltic States and Leningrad district. *Corresponding author, E-mail: [email protected] Radovsky F.J. 1967. The Macronyssidae Parasitologiya 24(3): 193-200 [In Rus- 3Bat Conservation Ireland, Ulex House, Drumheel, Lisduff, Virginia, Co. Cavan and Laelapidae (Acarina: Mesostigma- sian]. #These authors contributed equally to this work
Received 25 August 2010; accepted 1 March 2011
ABSTRACT - Nyctalus noctula and Barbastella barbastellus were first reported from Ire- land in 1997, however these reports were based solely on echolocation call data. Since then, neither species have been reported again or confirmed as a resident species in Ireland. In this study the status of these two species in Ireland was assessed. For B. barbastellus, the woodlands in the area where it was previously reported from, in the Lough Derg region, were surveyed by walked transects using Pettersson D1000X bat detectors and through pas- sive monitoring using the SD1 Anabat detector and the Pettersson D1000X over three nights. Out of 1011 recordings, no calls of B. barbastellus were encountered. For N. noc- tula, 98 Nyctalus sp. calls recorded from five squares (30 km2) on the east coast of Ireland, during a car based monitoring scheme were analysed (peak frequency and call duration). These were compared to 220 reference calls of N. leisleri from Dartry and Phoenix Park, County Dublin, Ireland and published data on N. leisleri and N. noctula calls from Britain. All Irish calls recorded from Dartry Park, Phoenix Park and the car transect squares fell within the known parameters range of N. leisleri but also overlapped with the higher fre- quency and shortest duration calls of N. noctula. However, no Irish calls overlapped with the lower frequency range and longest call duration of N. noctula, indicating that this latter species was probably not recorded in the Irish dataset. The results of this study are dis- cussed in relation to the difficulty of reporting a bat species presence based on echolocation calls alone and the suitability of Ireland for both species.
Key words: distribution, detection, Chiroptera, species presence, Ireland
RIASSUNTO - Verifica della presenza di Barbastella barbastellus e Nyctalus noctula e descrizione delle ecolocalizzazioni di N. leisleri in Irlanda. La presenza di Nyctalus noctu- la and Barbastella barbastellus in Irlanda è stata segnalata nel 1997, sulla base dell‟analisi di ecolocalizzazioni. Da allora, essa non è stata ulteriormente confermata. Per accertare la presenza di B. barbastellus nell‟area di Lough Derg, sono stati effettuati transetti con bat
111 Buckley et al. detector (Pettersson D1000X) e registrazioni da punti fissi per un periodo di tre notti (SD1 Anabat e Pettersson D1000X). Su un totale di 1011 registrazioni, nessuna è risultata attribu- ibile a B. barbastellus. Per N. noctula, 98 Nyctalus sp. ecolocalizzazioni registrate in 5 qua- drati (30 km2) distribuiti sulla costa orientale dell‟isola sono state analizzate (frequenza massima e durata) e confrontate con i dati disponibili in letteratura sulle ecolocalizzazioni delle due specie e con 220 ecolocalizzazioni di N. leisleri (Dartry e Phoenix Park, Contea di Dublino). I valori dei diversi parametri di tutte le ecolocalizzazioni concordano con quelli riferiti a N. leisleri, benché in parte si sovrappongano a quelli noti per N. noctula. Nel com- plesso i risultati suggeriscono che quest‟ultima specie non sia presente in Irlanda e vengono discussi in relazione alla difficoltà di accertare la presenza di una specie in base alle sole ecolocalizzazioni e alla vocazionalità dell‟isola per entrambe le nottole.
Parole chiave: distribuzione, accertamento della presenza, Chiroptera, Irlanda
DOI: 10.4404/Hystrix-22.1.4472
INTRODUCTION and I. Ahlén, respectively, in the atlas of European Mammals (Mitchell-Jones To date, nine species of bat have been et al. 1999). Based on Mitchell-Jones et confirmed as resident in Ireland, na- al. (1999), subsequent publications mely Myotis nattereri, M. daubentonii, have considered the two species to be M. mystacinus, Pipistrellus pipistrellus, present in Ireland (e.g. Dietz et al. P. pygmaeus, P. nathusii, Nyctalus lei- 2009; C. Dietz personal communication sleri, Plecotus auritus and Rhinolophus 2010). However, to date, these two re- hipposideros (Marnell et al. 2009). cords have not been confirmed, as the These nine species have been con- species identification was only based firmed by capture of individuals and on echolocation calls and no individu- the presence of maternity roosts. Re- als have been captured nor have breed- cently, Boston et al. (2010) reported on ing sites been found for either species. the search for M. brandtii, but con- This could potentially be due to the cluded that there is currently insuffi- roosting behaviour of both species, cient data to confirm that this species is since they generally roost in trees as resident in Ireland. The other two spe- opposed to buildings, so they may have cies that have been reported in Ireland been overlooked (Boonman 2000; are Nyctalus noctula and Barbastella Russo et al. 2004). However, the erec- barbastellus, each reported based on tion and monitoring of 60 Schwegler echolocation calls from a single local- bat boxes in Portumna forest park for ity, Phoenix Park, County (Co.) Dublin the last 11 years has failed to find any on the east coast for the former and B. barbastellus in the woodland despite Portumna forest, Co. Galway in the annual surveys (Teesdale 2006; K. Mc west for the latter. These two records Aney personal communication 2010). were obtained in 1997 (K. Mc Aney Previous studies carried out in Ireland personal communication 1999) and have not been able to confirm the pres- first mentioned as personal communi- ence of either species despite numerous cations from I. Ahlén and H.J. Baagøe surveys totalling hundreds of recording
112 Buckley et al. Status of N. noctula and B. barbastellus in Ireland detector (Pettersson D1000X) e registrazioni da punti fissi per un periodo di tre notti (SD1 hours (e.g. Daubenton‟s bat waterway the same location and species show Anabat e Pettersson D1000X). Su un totale di 1011 registrazioni, nessuna è risultata attribu- monitoring scheme; Aughney et al. echolocation call variation across their ibile a B. barbastellus. Per N. noctula, 98 Nyctalus sp. ecolocalizzazioni registrate in 5 qua- range (e.g. O'Farrell et al. 2000). 2 2009; Batlas project). Since 2003, Bat drati (30 km ) distribuiti sulla costa orientale dell‟isola sono state analizzate (frequenza Conservation Ireland has been monitor- Therefore, the description of a species‟ massima e durata) e confrontate con i dati disponibili in letteratura sulle ecolocalizzazioni ing bats using a car-based transect echolocation calls as well as their delle due specie e con 220 ecolocalizzazioni di N. leisleri (Dartry e Phoenix Park, Contea di scheme (Roche et al. 2009). Currently, variation across space is required for Dublino). I valori dei diversi parametri di tutte le ecolocalizzazioni concordano con quelli 2 riferiti a N. leisleri, benché in parte si sovrappongano a quelli noti per N. noctula. Nel com- 28 survey squares (30 km ) are moni- accurate species identification. When plesso i risultati suggeriscono che quest‟ultima specie non sia presente in Irlanda e vengono tored annually and N. leisleri is the foraging, N. noctula and N. leisleri discussi in relazione alla difficoltà di accertare la presenza di una specie in base alle sole third most frequently encountered bat switch between two distinguishable call ecolocalizzazioni e alla vocazionalità dell‟isola per entrambe le nottole. with 2715 calls recorded between 2003 types (Waters et al. 1995; Parsons and and 2008 (Roche et al. 2009). A num- Jones 2000), the first signal consists of Parole chiave: distribuzione, accertamento della presenza, Chiroptera, Irlanda ber of the survey squares are located on a low bandwidth, long duration shallow the east coast of Ireland close to the frequency modulated (FM) call and the DOI: 10.4404/Hystrix-22.1.4472 location of the original 1997 N. noctula second has a higher bandwidth call record. A similar car-based scheme has with shorter duration (more broadband INTRODUCTION and I. Ahlén, respectively, in the atlas been piloted in Britain where N. noc- FM). N. noctula is reported to alternate of European Mammals (Mitchell-Jones tula and B. barbastellus have been suc- between the two call types more fre- To date, nine species of bat have been et al. 1999). Based on Mitchell-Jones et cessfully recorded using this methodol- quently than N. leisleri, which is rarely confirmed as resident in Ireland, na- al. (1999), subsequent publications ogy (Russ et al. 2008). mentioned as doing so (Zingg 1988; mely Myotis nattereri, M. daubentonii, have considered the two species to be Since Fenton and Bell (1981) and Zbinden 1989; Waters et al. 1995; Par- M. mystacinus, Pipistrellus pipistrellus, present in Ireland (e.g. Dietz et al. Ahlén (1981), echolocation calls have sons and Jones 2000; Russo and Jones P. pygmaeus, P. nathusii, Nyctalus lei- 2009; C. Dietz personal communication been widely and successfully used to 2002; Obrist et al. 2004; Papadatou et sleri, Plecotus auritus and Rhinolophus 2010). However, to date, these two re- identify bat species using different al. 2008). Acoustically, these two spe- hipposideros (Marnell et al. 2009). cords have not been confirmed, as the classification methods such as dis- cies are frequently separated in the These nine species have been con- species identification was only based criminant function analyses, neural field by differences in the frequency of firmed by capture of individuals and on echolocation calls and no individu- networks, synergetic algorithms, etc. maximum energy (Fpeak) of their echo- the presence of maternity roosts. Re- als have been captured nor have breed- (Vaughan et al. 1997; Parsons and location calls. As in many other bat cently, Boston et al. (2010) reported on ing sites been found for either species. Jones 2000; Parsons 2001; Russo and species (e.g. Parsons 1997; Barclay et the search for M. brandtii, but con- This could potentially be due to the Jones, 2002; Obrist et al. 2004; Preato- al. 1999; Gillam and McCracken 2007; cluded that there is currently insuffi- roosting behaviour of both species, ni et al. 2005; Papadatou et al. 2008). Russo et al. 2007; Soisook et al. 2008), cient data to confirm that this species is since they generally roost in trees as Similarly to direct identification by there is geographic variability in echo- resident in Ireland. The other two spe- opposed to buildings, so they may have trained listeners, these classification location call parameters within N. cies that have been reported in Ireland been overlooked (Boonman 2000; methods „compare‟ a suite of meas- leisleri across Europe. Average Fpeak of are Nyctalus noctula and Barbastella Russo et al. 2004). However, the erec- urements (e.g. time and frequency pa- N. leisleri has been reported to vary barbastellus, each reported based on tion and monitoring of 60 Schwegler rameters) from calls of individuals, from 25.5 kHz in England to 31.7 kHz echolocation calls from a single local- bat boxes in Portumna forest park for whose species identity is unknown, to in Greece with intermediate values be- ity, Phoenix Park, County (Co.) Dublin the last 11 years has failed to find any reference calls from identified indi- ing reported from Italy, Switzerland on the east coast for the former and B. barbastellus in the woodland despite viduals, and then, based on similarity, and England (Zingg 1988; Waters et al. Portumna forest, Co. Galway in the annual surveys (Teesdale 2006; K. Mc assign the unknown calls to a species or 1995; Vaughan et al. 1997; Parsons and west for the latter. These two records Aney personal communication 2010). species group. Species misidentifica- Jones 2000; Russo and Jones 2002; were obtained in 1997 (K. Mc Aney Previous studies carried out in Ireland tion can happen when two or more spe- Obrist et al. 2004; Papadatou et al. personal communication 1999) and have not been able to confirm the pres- cies have very similar calls (e.g. Myotis 2008). Although this variation could first mentioned as personal communi- ence of either species despite numerous species) or when reference and un- partly be explained by different re- cations from I. Ahlén and H.J. Baagøe surveys totalling hundreds of recording known echolocation calls are not from cording situations (e.g. emergence vs.
113 Buckley et al. foraging) of bats flying in different 2003; Pētersons et al. 2010; C. Corben habitats (e.g. open vs. semi-cluttered), personal communication 2009; Puech- recording of bats in similar situations maille unpublished data). and habitats in Greece, France and Ire- If these species are indeed resident in land confirmed the existence of large Ireland, then there is a legal obligation variations due to the bats themselves to monitor and protect them. N. noctula (Puechmaille unpublished data). Simi- and B. barbastellus are currently listed larly, characteristics of echolocation as Annex IV of the EU Habitats Direc- calls of N. noctula vary geographically tive, while B. barbastellus is also listed across Europe (Zbinden 1989; Vaughan as Annex II, thus requiring the designa- et al. 1997; Parsons and Jones 2000; tion of Special Areas of Conservation Russo and Jones 2002; Obrist et al. (92/43/EEC). 2004; Papadatou et al. 2008), but, on The aim of this study was to follow up average, its Fpeak is lower than in N. on the 1997 records of N. noctula and leisleri. This important geographic B. barbastellus and try to determine the variation in both species‟ echolocation presence of these two bat species. The calls renders uncertain species identifi- status of N. noctula was assessed cations without local reference calls. through the analysis of calls recorded Therefore, given that the only record of in the last five years from eastern Ire- N. noctula in Ireland is from echoloca- land and the status of B. barbastellus tion calls, knowledge on echolocation was assessed through both active and calls of N. leisleri from Ireland is of passive detector surveys carried out in paramount importance, although rarely June 2010 in Portumna forest and two reported. other woodland sites on the shores of B. barbastellus also emits two alternat- Lough (lake) Derg. ing call types. Type 1 calls are broad- band sweeping from 36 kHz to 28 kHz MATERIALS AND METHODS with a mean Fpeak of 32.8 kHz while type 2 calls have a narrow band FM 1. B. barbastellus component with starting frequency of The presence of the species was investi- 45 kHz followed by a broadband FM gated in three woodland sites (Portumna component ending on average at about forest, Rosturra forest and Raheen forest) 32 kHz with a mean Fpeak of 40 kHz in the vicinity of Lough Derg (Fig. 1) be- (Denzinger et al. 2001). In contrast to tween the 15th and the 17th of June 2010. Nyctalus species, echolocation calls of The first site was selected as it was the B. barbastellus show little variation source of the 1997 B. barbastellus record across Europe (Parsons and Jones and the two other sites were chosen as they 2000; Russo and Jones 2002; Obrist et represented suitable woodlands within al. 2004) and are quite distinctive from proximity of Portumna forest. echolocation calls of any other Euro- 1.1 Site descriptions pean bat species (Ahlén and Baagøe 1999), so that reliable species identifi- Portumna forest is located in Co. Galway cation can be done from good quality on the northern shore of Lough Derg, adja- echolocation call recordings (Ahlén cent to the town of Portumna (Fig. 1). It is
114 Buckley et al. Status of N. noctula and B. barbastellus in Ireland foraging) of bats flying in different 2003; Pētersons et al. 2010; C. Corben habitats (e.g. open vs. semi-cluttered), personal communication 2009; Puech- recording of bats in similar situations maille unpublished data). and habitats in Greece, France and Ire- If these species are indeed resident in land confirmed the existence of large Ireland, then there is a legal obligation variations due to the bats themselves to monitor and protect them. N. noctula (Puechmaille unpublished data). Simi- and B. barbastellus are currently listed larly, characteristics of echolocation as Annex IV of the EU Habitats Direc- calls of N. noctula vary geographically tive, while B. barbastellus is also listed across Europe (Zbinden 1989; Vaughan as Annex II, thus requiring the designa- et al. 1997; Parsons and Jones 2000; tion of Special Areas of Conservation Russo and Jones 2002; Obrist et al. (92/43/EEC). 2004; Papadatou et al. 2008), but, on The aim of this study was to follow up average, its Fpeak is lower than in N. on the 1997 records of N. noctula and leisleri. This important geographic B. barbastellus and try to determine the variation in both species‟ echolocation presence of these two bat species. The calls renders uncertain species identifi- status of N. noctula was assessed cations without local reference calls. through the analysis of calls recorded Therefore, given that the only record of in the last five years from eastern Ire- N. noctula in Ireland is from echoloca- land and the status of B. barbastellus tion calls, knowledge on echolocation was assessed through both active and calls of N. leisleri from Ireland is of passive detector surveys carried out in June 2010 in Portumna forest and two paramount importance, although rarely reported. other woodland sites on the shores of Figure 1 - Location of the study sites in Ireland with details of the Lough Derg region. B. barbastellus also emits two alternat- Lough (lake) Derg. ing call types. Type 1 calls are broad- approximately 600 ha in size. The main woodland consists of mature tree stands band sweeping from 36 kHz to 28 kHz MATERIALS AND METHODS canopy tree species are conifers, including dominated by Q. petraea and cleared areas Pinus sylvestris, Larix sp. and Picea abies. planted with a mixture of native tree and with a mean Fpeak of 32.8 kHz while type 2 calls have a narrow band FM 1. B. barbastellus However, there are also some broadleaf shrub species. stands with mature Quercus sp., and Fagus component with starting frequency of The presence of the species was investi- sylvatica. Native woodland (which in Ire- 1.2 Survey methodology 45 kHz followed by a broadband FM gated in three woodland sites (Portumna land includes mainly broadleaves such as component ending on average at about forest, Rosturra forest and Raheen forest) Q. robur, Q. petraea, Fraxinus excelsior, Portumna and Raheen forests were sur- 32 kHz with a mean Fpeak of 40 kHz in the vicinity of Lough Derg (Fig. 1) be- Alnus glutinosa and various Salix spp., veyed using walked transects and passive (Denzinger et al. 2001). In contrast to tween the 15th and the 17th of June 2010. among others) mainly occurs on the lake monitoring, while only passive monitoring Nyctalus species, echolocation calls of The first site was selected as it was the shore and northern edge of the forest. was carried out in Rosturra forest. Three B. barbastellus show little variation source of the 1997 B. barbastellus record Raheen forest is located in Co. Clare, 2 km transect routes were walked in Portumna across Europe (Parsons and Jones and the two other sites were chosen as they south of Scarriff village (Fig. 1). It is ap- (4, 3.4 and 4.3 km) over two nights, while 2000; Russo and Jones 2002; Obrist et represented suitable woodlands within proximately 128 ha in size. This forest con- one transect route was walked in Raheen al. 2004) and are quite distinctive from proximity of Portumna forest. sists of stands of very mature Quercus spp. forest (3.9 km) in one night. Each route echolocation calls of any other Euro- and mixed broadleaves (native and non- was mapped using an Etrex GPS (Garmin). 1.1 Site descriptions native) and conifer stands of varying age. Transects were walked at a constant pace, pean bat species (Ahlén and Baagøe Finally, Rosturra forest is located in Co. from 40 minutes after sunset until 00:30 1999), so that reliable species identifi- Portumna forest is located in Co. Galway Galway, 4.2 km northeast of Woodford am. All bat activity was manually recorded cation can be done from good quality on the northern shore of Lough Derg, adja- village (Fig. 1). This 39 ha semi-natural in real time on hearing using D1000X de- echolocation call recordings (Ahlén cent to the town of Portumna (Fig. 1). It is
115 Buckley et al. tectors (Pettersson Elktronik AB) set with a Generally, individuals with Fpeak <50 kHz pre-recording time of 2 seconds so as not to were classified as P. pipistrellus while miss any bat pass. Headphones were used those with Fpeak >52 kHz were classified as with heterodyne mode in one channel and P. pygmaeus and intermediate ones (50 frequency division in the other. The detec- kHz
116 Buckley et al. Status of N. noctula and B. barbastellus in Ireland tectors (Pettersson Elktronik AB) set with a Generally, individuals with Fpeak <50 kHz lowing a clear identification of the start and the following one. All calculations and pre-recording time of 2 seconds so as not to were classified as P. pipistrellus while end of the call. The five squares are located graphs were done in R 2.10.1 (Ihaka and miss any bat pass. Headphones were used those with Fpeak >52 kHz were classified as in the east and north east of the island and Gentleman 1996; R Development Core with heterodyne mode in one channel and P. pygmaeus and intermediate ones (50 square O04 includes locations within 5 km Team 2009). frequency division in the other. The detec- kHz
All calls recorded from the D1000X were visually observed while recorded, which (Pettersson Elktronik AB) to measure Fpeak analysed using the software BatSound, ver- corroborated the identification of the spe- using the power spectrum window. We Two hundred and twenty N. leisleri sion 4 (Pettersson Elktronik AB). When- cies recorded. Recordings were typically determined the call duration by measuring echolocation calls from Phoenix Park ever possible, calls were identified to spe- 10-20 seconds long and captured the range the time between the start and the end of and Dartry Park were studied (Tab. 2). cies, except for Myotis species which were of calls emitted by individuals while forag- each call from the amplitude shown in the These calls included search phase echo- just identified to genus. All calls recorded ing in the open. A total of 220 calls were oscillogram window. For calls recorded in location as well as approach phase calls from the Anabat were analysed using the analysed from six recordings, which repre- Phoenix Park and Dartry Park, call Fpeak, and feeding buzzes (Simmons et al. software AnalookW version 3.7o (written sented the range of calls emitted by the bats start and end time were measured using the 1979). In many species, feeding buzzes by Chris Corben). Pipistrellus calls were on these three evenings. function „Pulse Characteristics Analysis‟ can be divided into “buzz I” and “buzz only identified to genus from Anabat re- Ninety-eight good quality calls from 2005 available in BatSound. The „mark position- cordings while real time recordings ob- to 2009 were selected from five (J06, N77, ing‟ was enabled and measurements were II”, the start of the latter being charac- tained with the D1000X were identified to O04, S78, T05) of the 28 squares where the visually checked and corrected if neces- terized by an abrupt drop in frequency species level whenever possible. Real-time car-based monitoring scheme is operated sary. Intervals between pulses (IBP) were (e.g. Surlykke et al. 1993). Fpeak of recordings of Pipistrellus species were (Fig. 5). Good quality calls were defined as calculated as the absolute time difference search phase calls was typically in the identified to species level based on Fpeak. calls with a good signal to noise ratio, al- between the start of one call and the start of range of 20.5-25.5 kHz and often showed
117 Buckley et al.
Table 1 - Summary of the number of times each species or group of species were recorded during the surveys at Portumna, Raheen and Rosturra. Sounds of Pipistrellus spp. recorded with the Anabat system were not identified to species level and are, therefore, all included in the Pipistrellus spp. field, hence data were not available (N/A) for P. pygmaeus and P. pipistrellus separately (An: Anabat, T: Transect, D: D1000X, Tot: Total).
Portumna Raheen Rosturra
An T1 T2 T3 D1 Tot An. T1 D1 D2 Tot An Tot
Pipistrellus spp. 231 53 12 6 0 302 34 0 29 0 63 33 398 P. pygmaeus N/A 58 26 11 0 95 N/A 7 105 3 115 N/A 210 P. pipistrellus N/A 50 21 10 0 81 N/A 0 87 0 87 N/A 168 Myotis spp. 97 12 11 5 0 125 2 2 15 17 36 34 195 Nyctalus leisleri 0 14 15 4 0 33 0 1 0 0 1 0 34 Plecotus sp. 0 0 0 0 1 1 1 1 0 0 2 3 6 Tot 328 187 85 36 1 637 37 11 236 20 304 70 1011
Figure 2 - Typical pattern of search phase echolocation calls of N. leisleri in the open. a pattern of frequency alternation (see Feeding buzz I reached the highest Fpeak Fig. 2 and Tab. 2). Intervals between we recorded in the species (Tab. 2 and pulses averaged 227.4 ms with a Fig. 4). Fpeak of feeding buzz II was maximum of 376 ms. Approach phase approximately 7 kHz lower than in calls were characterized by a higher buzz I (Tab. 2). All four types of calls Fpeak, typically between 25.5 to 30.3 can be found on one sequence pro- kHz, a shorter duration and a higher duced by the same individual. Some of repetition rate (Fig. 3 and Tab. 2). the recordings from which calls were
118 Buckley et al. Status of N. noctula and B. barbastellus in Ireland
Table 1 - Summary of the number of times each species or group of species were recorded Table 2 - Characteristics of echolocation calls of N. leisleri in Ireland recorded from a sta- during the surveys at Portumna, Raheen and Rosturra. Sounds of Pipistrellus spp. recorded tionary position (columns 1 to 4) and comparison with the car transect data (last column) in with the Anabat system were not identified to species level and are, therefore, all included Ireland. Average, standard deviation (SD), minimum (min.) and maximum (max.) values in the Pipistrellus spp. field, hence data were not available (N/A) for P. pygmaeus and P. are presented for each parameter (1frequency with maximum energy; 2call duration; pipistrellus separately (An: Anabat, T: Transect, D: D1000X, Tot: Total). 3interval between pulses). IBP was not measured for the Car Transect data (N/A) as most recordings (300 ms long) only contained a single call. Portumna Raheen Rosturra Search phase Approach phase Buzz I Buzz II Car transect An T1 T2 T3 D1 Tot An. T1 D1 D2 Tot An Tot
Pipistrellus spp. 231 53 12 6 0 302 34 0 29 0 63 33 398 Sample size 123 33 56 8 98 1 P. pygmaeus N/A 58 26 11 0 95 N/A 7 105 3 115 N/A 210 Fpeak (SD) 23.0 (1.21) 27.1 (1.09) 31.7 (1.66) 24.5 (1.46) 25.6 (2.41) P. pipistrellus N/A 50 21 10 0 81 N/A 0 87 0 87 N/A 168 Min-max 20.50-25.50 25.6-30.3 28.7-34.7 22.5-26.4 21.6-31.7 Myotis spp. 97 12 11 5 0 125 2 2 15 17 36 34 195 Duration2 (SD) 15.3 (2.28) 8.9 (2.77) 1.8 (1.02) 0.45 (0.15) 9.2 (3.13) Nyctalus leisleri 0 14 15 4 0 33 0 1 0 0 1 0 34 Min-max 8.3-20.3 3.9-15.0 0.13-4.22 0.25-0.72 3.56-17 Plecotus sp. 0 0 0 0 1 1 1 1 0 0 2 3 6 IBP3 (SD) 227.4 (48.4) 127.4 (61.4) 14.4 (12.9) 5.4 (0.16) N/A Tot 328 187 85 36 1 637 37 11 236 20 304 70 1011 Min-max 87-376 23-255 0.48-70.26 5.07-5.49 N/A
Figure 2 - Typical pattern of search phase echolocation calls of N. leisleri in the open. Figure 3 - Typical pattern of N. leisleri search phase echolocation calls followed by an „ap- proach like phase‟ sequence (middle) and returning to search phase towards the end. Circles represent peak frequency and triangles call duration. a pattern of frequency alternation (see Feeding buzz I reached the highest Fpeak Fig. 2 and Tab. 2). Intervals between we recorded in the species (Tab. 2 and measured also contained N. leisleri so- car-based bat monitoring squares (J06: pulses averaged 227.4 ms with a Fig. 4). F of feeding buzz II was peak cial calls similar to those described in 18, N77: 32, O04: 14, S78: 16, T05: maximum of 376 ms. Approach phase approximately 7 kHz lower than in Pfalzer (2002) (type B and M) and fig- 18). A plot of F vs. call duration for calls were characterized by a higher buzz I (Tab. 2). All four types of calls peak ure 5.12 in Russ (1999). the car transect calls and the reference F , typically between 25.5 to 30.3 can be found on one sequence pro- peak Ninety eight high quality calls were calls described above showed a near kHz, a shorter duration and a higher duced by the same individual. Some of selected for the final analysis from the complete overlap between the two data- repetition rate (Fig. 3 and Tab. 2). the recordings from which calls were
119 Buckley et al.
Figure 4 - Sequence of N. leisleri echolocation calls showing the approach phase calls and feeding buzz I and buzz II. Circles represent peak frequency and triangles call duration.
Figure 5 - Echolocation peak frequency versus call duration for search phase and approach phase calls (●), feeding buzz I (♦) and feeding buzz II (■) for N. leisleri recorded in Ireland. Triangles (▲) represent echolocation calls recorded during the car transects from eastern Ireland (squares on the map of Ireland).
120 Buckley et al. Status of N. noctula and B. barbastellus in Ireland
Figure 4 - Sequence of N. leisleri echolocation calls showing the approach phase calls and feeding buzz I and buzz II. Circles represent peak frequency and triangles call duration.
Figure 6 - Echolocation peak frequency vs. call duration for N. leisleri recorded from sta- tionary position (+) and car transect data (¤). The average values for the two data sets are presented by their respective symbols enlarged. Published averages (large squares, triangles and circles), one standard deviation (solid line) and range (dashed line) of published N. noc- tula (open) and N. leisleri (filled) calls from England and Wales are also presented. For published data, different types of calls were presented whenever available in the original publication (e.g. type 1 and type 2 calls; Parsons and Jones, 2000).
sets‟ parameters (Fig. 5). Fig. 6 pre- quency range previously reported for sents published data on N. leisleri and N. leisleri in England while a few calls N. noctula from England and shows slightly exceeded, by a few millisec- that although some calls from Ireland onds, the previous maximum duration overlap with the published upper calls reported (Fig. 6). Waters et al. (1995) of British N. noctula (Vaughan et al. and Parsons et al. (2000) did not in- 1997; Parsons and Jones, 2000), none clude minimum and maximum values of the calls recorded in Ireland fall for the parameters measured but only within the lower frequency range of included their standard deviation from Figure 5 - Echolocation peak frequency versus call duration for search phase and approach British N. noctula calls (16.8-20 kHz, which we could estimate the 99% con- phase calls (●), feeding buzz I (♦) and feeding buzz II (■) for N. leisleri recorded in Ireland. Vaughan et al., 1997; see also Fig. 6). fidence intervals assuming a normal Triangles (▲) represent echolocation calls recorded during the car transects from eastern Fpeak of all 318 calls fell within the fre- distribution of the data. By doing so, Ireland (squares on the map of Ireland).
121 Buckley et al. our data entirely falls within the range land with large, mature trees and a well for N. leisleri calls reported by those developed understory (Sierro 1999; authors (data not shown). The maxi- Eriksson et al. 2004; Russo et al. 2004, mum call duration reported in our data 2005, 2010; Hillen et al. 2009; Rebelo set (20.3 ms) was less than 2/3rd of the and Jones 2010). Its diet is dominated maximum reported for N. noctula in by Lepidoptera (Sierro and Arlettaz England (33.5 ms, Vaughan et al. 1997; 1997), indicating a very specialised see also Fig. 6). niche. The question then arises, is Ire- land suitable from a climatic and habi- DISCUSSION tat perspective for this species? Recent predictive modelling for the distribu- Barbastella barbastellus tion of B. barbastellus in Europe shows suitable climatic conditions in the east This study could not confirm the pres- coast and midlands (including the ence of this species in the Lough Derg Lough Derg region) of Ireland (Rebelo region. However, it is important to note 2009). Nevertheless, this model does that all known Irish species were re- not consider habitat variables (e.g. for- corded during survey work except Rhi- est cover), which would be an impor- nolophus hipposideros, whose range tant factor for the presence of the spe- does not extend that far east (Kelleher cies (H. Rebelo personal communica- 2004) and P. nathusii, which has been tion 2010). Currently Ireland has a very only recently confirmed to be a resident reduced forest cover (Gallagher et al. species in Ireland (Russ and Montgom- 2001), although there is currently suit- ery 1998), breeding only in the north able forest habitat on the east coast, east (Russ et al. 2001). Plecotus auri- particularly in Co. Wicklow (Perrin et tus, which emits “quiet” echolocation al. 2006). Ireland‟s forest cover was calls, similar to B. barbastellus (Russ reduced to less than 1% by the nine- 1999) was also recorded at each site teenth century (McCracken 1971). surveyed, indicating the thoroughness Therefore, it is possible that this spe- of this acoustic survey. Myotis dauben- cies occurred here in the past but be- tonii can emit a social call that overlaps came extinct due to habitat loss. Al- in call structure with the type 2 call of though B. barbastellus is not known for B. barbastellus (Denzinger et al. 2001; long distance flights (Hutterer et al. call of type A in Pfalzer 2002). There- 2005), other slow flying species such as fore, it cannot be excluded that the re- M. mystacinus and P. auritus managed ported recording from 1997 may have to colonise Ireland after the last glacial been from M. daubentonii social calls; maximum. An investigation of the sub- however, we were unable to get access fossil bat bones of Irish caves would be to the unpublished 1997 collected data required to test if B. barbastellus also to investigate this possibility. reached Ireland in the same period. B. barbastellus is vulnerable through- out Europe (Temple and Terry 2007) 2. Nyctalus noctula and is highly specialised in its habitat requirements, generally occurring in The analyses of our data set suggest mature broadleaved and mixed wood- that N. noctula is absent from Ireland
122 Buckley et al. Status of N. noctula and B. barbastellus in Ireland our data entirely falls within the range land with large, mature trees and a well and show that the lower echolocation to the Irish N. leisleri echolocation for N. leisleri calls reported by those developed understory (Sierro 1999; calls of Irish N. leisleri compared to calls. authors (data not shown). The maxi- Eriksson et al. 2004; Russo et al. 2004, British conspecifics could complicate If N. noctula is absent from Ireland it is mum call duration reported in our data 2005, 2010; Hillen et al. 2009; Rebelo correct species identification without unlikely to be due to Ireland‟s island set (20.3 ms) was less than 2/3rd of the and Jones 2010). Its diet is dominated local reference calls. When the call status, given that this species is a long maximum reported for N. noctula in by Lepidoptera (Sierro and Arlettaz dataset from Dartry/Phoenix Park and distant migrant (Hutterer et al. 2005). England (33.5 ms, Vaughan et al. 1997; 1997), indicating a very specialised the car transect squares were compared Therefore, the Irish Sea, particularly see also Fig. 6). niche. The question then arises, is Ire- to the published call parameters for between south western Scotland and land suitable from a climatic and habi- peak frequency and call duration for N. north eastern Ireland (approx. 20 km) DISCUSSION tat perspective for this species? Recent leisleri in England (Waters et al. 1995; should not have been a barrier for colo- predictive modelling for the distribu- Vaughan et al. 1997; Parsons and Jones nisation. The potential distribution of Barbastella barbastellus tion of B. barbastellus in Europe shows 2000), they were found to fall within N. noctula in Europe has not been suitable climatic conditions in the east the known call range for this species modelled, but, similarly to B. bar- This study could not confirm the pres- coast and midlands (including the (see also Russ 1999). However, our bastellus, this species is reliant on ence of this species in the Lough Derg Lough Derg region) of Ireland (Rebelo study reveals that N. leisleri from Ire- woodland for foraging (Mackie and region. However, it is important to note 2009). Nevertheless, this model does land use on average a lower Fpeak than Racey 2007) and roosting, with a heavy that all known Irish species were re- not consider habitat variables (e.g. for- N. leisleri from Britain and we hy- reliance on woodpecker holes (Boon- corded during survey work except Rhi- est cover), which would be an impor- pothesise that this lower F could be man 2000). Woodpeckers (Dendroco- tant factor for the presence of the spe- peak nolophus hipposideros, whose range explained by the absence of N. noctula pos major) became extinct in Ireland in cies (H. Rebelo personal communica- does not extend that far east (Kelleher in Ireland. This would give N. leisleri historical times due to deforestation 2004) and P. nathusii, which has been tion 2010). Currently Ireland has a very the possibility to exploit the niche oc- (D'Arcy 1999) so N. noctula might only recently confirmed to be a resident reduced forest cover (Gallagher et al. cupied by N. noctula in the continent have occurred in Ireland in the past, species in Ireland (Russ and Montgom- 2001), although there is currently suit- and Britain and would potentially lead and became extinct along with wood- ery 1998), breeding only in the north able forest habitat on the east coast, to more prey being available, which peckers due to loss of habitat. How- east (Russ et al. 2001). Plecotus auri- particularly in Co. Wicklow (Perrin et tus, which emits “quiet” echolocation al. 2006). Ireland‟s forest cover was could explain why N. leisleri is com- ever, the “re-colonization” of the east- calls, similar to B. barbastellus (Russ reduced to less than 1% by the nine- mon in Ireland (O'Sullivan 1994). Fur- ern coast of Ireland by D. major in the 1999) was also recorded at each site teenth century (McCracken 1971). ther work should however be carried last few years (O'Halloran 2009) might surveyed, indicating the thoroughness Therefore, it is possible that this spe- out to test this hypothesis. Irish N. create favourable roosting conditions of this acoustic survey. Myotis dauben- cies occurred here in the past but be- leisleri calls do partly overlap with the for N. noctula and favour their coloni- tonii can emit a social call that overlaps came extinct due to habitat loss. Al- published call parameters for N. noc- sation from Britain. Furthermore, cur- in call structure with the type 2 call of though B. barbastellus is not known for tula but a large space parameter (Fpeak < rent policies promoting re-forestation B. barbastellus (Denzinger et al. 2001; long distance flights (Hutterer et al. 20 kHz and call duration > 21 ms, see and afforestation with broadleaf trees call of type A in Pfalzer 2002). There- 2005), other slow flying species such as Fig. 6) was missing from the Irish data might also help to create habitats suit- fore, it cannot be excluded that the re- M. mystacinus and P. auritus managed set when compared to the published able for both species. ported recording from 1997 may have to colonise Ireland after the last glacial range of N. noctula echolocation calls Although it is impossible to prove the been from M. daubentonii social calls; maximum. An investigation of the sub- in Britain (Vaughan et al. 1997; Par- absence of a species (e.g. Puechmaille however, we were unable to get access fossil bat bones of Irish caves would be sons and Jones 2000). We hypothesize et al. 2009), there is currently insuffi- to the unpublished 1997 collected data required to test if B. barbastellus also that if N. noctula were present in Ire- cient evidence to state that N. noctula to investigate this possibility. reached Ireland in the same period. land, then the Irish Nyctalus sp. call and B. barbastellus occur in Ireland. B. barbastellus is vulnerable through- dataset would contain calls of longer Further surveys should be carried out in out Europe (Temple and Terry 2007) 2. Nyctalus noctula duration and lower Fpeak. We were un- Irish woodlands to determine the status and is highly specialised in its habitat able to get access to the 1997 unpub- of both species, which, for the moment, requirements, generally occurring in The analyses of our data set suggest lished original data set identified as N. should not be considered as present or mature broadleaved and mixed wood- that N. noctula is absent from Ireland noctula to investigate how it compares resident in the island. We would also
123 Buckley et al. recommend that species should only be Boonman M. 2000. Roost selection by noc- considered as present in a country or tules (Nyctalus noctula) and Dauben- landmass if there are undeniable data to ton's bats (Myotis daubentonii). J. Zool., support such a statement. 251: 385-389. Boston E.S.M., Buckley D.J., Bekaert M., Gager Y., Lundy M.G., Scott D.D., ACKNOWLEDGMENTS Prodöhl P.A., Montgomery W.I., Mar- nell F., Teeling E.C. 2010. The status of The authors would like to thank Kate Mc the cryptic bat species, Myotis Aney, Tina Aughney, Serena Dool and mystacinus and Myotis brandtii in Ire- Liam Kavanagh for their assistance with land. Acta Chiropterol. 12(5): 457-461. this study. Thanks to Danilo Russo and an D'Arcy G. 1999. Ireland's lost birds, Four anonymous reviewer whose comments Courts Press, Dublin. enhanced the quality of this paper. This Denzinger A., Siemers B.M., Schaub A., project was funded by the National Parks Schnitzler H.-U. 2001. Echolocation by and Wildlife Service, the Heritage Council the Barbastelle bat, Barbastella bar- and the Irish Research Council for Science bastellus. J. Comp. Physiol., 187: 521- Engineering and Technology (IRCSET). 528. Dietz C., Von Helversen O., Dietmar N. REFERENCES 2009. Bats of Britain, Europe & North- west Africa, A & C Black Publishers Ahlén I. 1981. Identification of Scandina- Ltd., London. vian bats by their sounds. The Swedish Eriksson A., de Jong J., Ahlén I. 2004. University of Agricultural Sciences, Habitat selection in a colony of Bar- Department of Wildlife Ecology, Upp- bastella barbastellus in south Sweden. sala. Institutionen för naturvårdsbiologi, Ahlén I. 2003. Inventerengen av barbastell Uppsala. (Barbastella barbastellus) 1999-2003 i Fenton M.B., Bell G.P. 1981. Recognition Sverige. Swedish Environmental Pro- of species of insectivorous bats by their tection Agency. echolocation calls. J. Mammal. 62(2): Ahlén I., Baagøe H.J. 1999. Use of ultra- 233-243. sound detectors for bat studies in Gallagher G., Dunne S., Jordan P., Stanley Europe: experiences from field identifi- B. 2001. Ireland's forest inventory and cation, surveys, and monitoring. Acta planning system. Department of the Chiropterol. 1(2): 137-150. Marine and Natural Resources, Wex- Aughney T., Langton S., Roche N. 2009. ford. All Ireland Daubenton's bat waterway Gillam E.H., McCracken G.F. 2007. Vari- monitoring scheme 2006-2008. Irish ability in the echolocation of Tadarida Wildlife Manuals, No. 42. National brasiliensis: effects of geography and Park and Wildlife Service, Department local acoustic environment. Anim. Be- of the Environment, Heritage and Local hav. 74: 277-286. Government, Dublin, Ireland. Hillen J., Kiefer A., Veith M. 2009. Forag- Barclay R.M.R., Fullard J.H., Jacobs D.S. ing site fidelity shapes the spatial or- 1999. Variation in the echolocation ganisation of a population of female calls of the hoary bat (Lasiurus ciner- western barbastelle bats. Biol. Conserv. eus): influence of body size, habitat 142: 817-823. structure, and geographic location. J. Hutterer R., Ivanova T., Meyer-Cords C., Zool. 77: 530-534. Rodrigues L. 2005. Bat migrations in
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The car-based bat monitoring scheme S., Pidinchedda E., Jones G. 2007. Di- for Ireland: synthesis report 2003-2008. vergent echolocation call frequencies in
Irish Wildlife Manuals, No. 39. Na- insular rhinolophids (Chrioptera): a tional Park and Wildlife Service, De- case of character displacement? J. Bio- geogr. 34: 2129-2138. partment of the Environment, Heritage Sierro A. 1999. Habitat selection by bar- and Local Government, Dublin, Ireland, bastelle bats (Barbastella barbastellus) Russ J. 1999. The Bats of Britain & Ire- in the Swiss Alps (Valais). J. Zool. land: echolocation calls, sound analysis, 248(4): 429-432. and species identification, Alana Ecol- Sierro A., Arlettaz R. 1997. Barbastelle ogy Ltd, London. bats (Barbastella spp.) specialize in the Russ J., Briggs P., Wembridge D. 2008. predation of moths: implications for The Bats & Roadside Mammals Survey foraging tactics and conservation. Acta 2008: Final Report on Fourth Year of Oecol. 18(2): 91-106. Study. Bat Conservation Trust, London. Simmons J.A., Fenton M.B., O'Farrell M.J. Russ J., Montgomery W.I. 1998. Nathusius' 1979. Echolocation and pursuit of prey pipistrelle bats (Pipistrellus nathusii, by bats. Science 203: 16-21. Keyserling & Blasius 1839) breeding in Soisook P., Bumrungsri S., Satasook C., Ireland. J. Zool. 245: 345-349. Thong V.D., Bu S.S.H., Harrison D.L., Russ J.M., Hutson A.M., Montgomery Bates P.J.J. 2008. A taxonomic review W.I., Racey P.A., Speakman J.R. 2001. of Rhinolophus stheno and R. ma- The status of Nathusius' pipistrelle layanus (Chiroptera: Rhinolophidae) (Pipistrellus nathusii Keyserling & from continental Southeast Asia: an Blasius, 1939) in the Bristish Isles. J. evaluation of echolocation call fre- Zool. 254: 91-100. quency in discriminating between cryp- Russo D., Cistrone L., Garonna A.P., Jones tic species. Acta Chiropterol. 10(2): G. 2005. Spatial and temporal patterns 221-242. of roost use by tree-dwelling barbastelle Surlykke A., Miller L.A., Møhl B., Ander- bats Barbastella barbastellus. Ecogra- sen B.B., Christensen-Dalsgaard J., Jør- phy 28: 769-776. gensen M.B. 1993. Echolocation in two
127