NATURA SLOVENIAE 25(3): 181-200 Prejeto / Received: 21. 12. 2023 
NERETVA SCIENCE WEEK 2022 – SCIENTIFIC PAPER Sprejeto / Accepted: 30. 12. 2023 
 
DOI: 10.14720/ns.25.3.181-200 
 Biotehniška fakulteta Univerze v Ljubljani in Nacionalni inštitut za biologijo, Ljubljana, 2023 
 
 
 
Study of bats in the upper Neretva River valley (Bosnia 
and Herzegovina) 
 
 
Maja ZAGMAJSTER1, Vojo MILANOVIĆ2, Behare REXHEPI1, Ester PREMATE1, Špela BORKO1, 
Anton VORAUER3 
 
 
1University of Ljubljana, Biotechnical Faculty, Department of Biology, SubBioLab, Jamnikarjeva 101, SI-1000 Ljubljana, 
Slovenia; E-mails: maja.zagmajster@bf.uni-lj.si, behare.rexhepi@bf.uni-lj.si; ester.premate@bf.uni-lj.si; 
borko.spela@gmail.com 
2Centre for Karst and Speleology, Branilaca Sarajeva 30, 71 000 Sarajevo, Bosnia and Herzegovina;  
E-mail: milanovic.vojo9@gmail.com 
3Ecotone Environmental Office, Austria; E-mail: anton.vorauer@utanet.at 
 
 
Abstract. The upper Neretva River valley with river and tributaries, preserved forests, natural caves and 
buildings represents a very interesting area for bats. During two field expeditions in summer 2022, we collected 
data on bats using a variety of methods: inspecting the potential roosts, mist netting, and recording bat 
echolocation calls. We used manual and automatic ultrasound detectors most frequently at sites along and near 
the Neretva River. The morphological identification of some species was confirmed using DNA analyses. We 
detected at least 13 bat species in the study area extending from Krupac to Konjic. With one addition, known from 
the literature, 14 different bat species have been found in the upper Neretva River valley, presenting nearly a half 
of all bat species recorded in Bosnia and Herzegovina. We confirmed the presence of three species of the highest 
conservation concern in Europe, listed in the Annex II of the Habitats directive: lesser horseshoe bat (Rhinolophus 
hipposideros), greater horseshoe bat (Rhinolophus ferrumequinum), and greater mouse-eared bat (Myotis myotis). 
Our finding of the alcathoe bat (Myotis alcathoe) presents new confirmation of this species in Bosnia and 
Herzegovina. Also, observation of the parti-coloured bat (Vespertilio murinus) is one of the few records of the 
species in this country. The upper Neretva River valley can be considered as an area of high conservation 
importance for bats. Additional fieldwork, including during other seasons, would likely reveal more bat species and 
their sites in the area of the upper Neretva River valley. 
 
Key words: Bosnia and Herzegovina, Neretva River, bats, echolocation, mist netting, roosts, Rhinolophus 
ferrumequinum, Rhinolophus hipposideros, Myotis myotis, Myotis alcathoe 
 
Izvleček. Raziskava netopirjev v dolini zgornje reke Neretve (Bosna in Hercegovina) – Dolina 
zgornje reke Neretve je z reko in pritoki, ohranjenimi gozdovi, naravnimi jamami in zgradbami zelo zanimivo 
območje za netopirje. Na dveh terenskih odpravah poleti 2022 smo popisovali netopirje z različnimi metodami: 
pregledi potencialnih zatočišč, lovom z mrežami in snemanjem eholokacijskih klicev. Na več mestih ob reki Neretvi 
in v njeni bližini smo uporabili ročne in avtomatske ultrazvočne detektorje. Morfološko določitev nekaterih vrst smo 
potrdili z analizami DNK. Skupno smo v naši raziskavi na preučevanem območju od Krupca do Konjica zabeležili 
vsaj 13 vrst netopirjev. Ob upoštevanju še ene, iz literature znane vrste, je bilo na območju zgornje Neretve skupno 
zabeleženih 14 različnih vrst netopirjev, kar predstavlja skoraj polovico vseh zabeleženih v Bosni in Hercegovini. 
Potrdili smo prisotnost treh vrst, ki so v Evropi deležne največje varstvene pozornosti in so navedene v Prilogi II 
Direktive o habitatih: mali podkovnjak (Rhinolophus hipposideros), veliki podkovnjak (Rhinolophus ferrumequinum) 
in navadni netopir (Myotis myotis). Najdba nimfnega netopirja (Myotis alcathoe) je nova potrditev vrste v Bosni in 
Hercegovini. Tudi najdba dvobarvnega netopirja (Vespertilio murinus) je eno redkih opažanj te vrste v državi. 
Dolino zgornjega toka reke Neretve lahko štejemo za območje, ki je zelo pomembno za varstvo netopirjev. Dodatno 
terensko delo, ki bi vključevalo tudi druge letne čase, bi razkrilo še več vrst in njihovih nahajališč na tem območju. 
 
Ključne besede: Bosna in Hercegovina, reka Neretva, netopirji, eholokacija, lov z mrežami, zatočišča, Rhinolophus 
ferrumequinum, Rhinolophus hipposideros, Myotis myotis, Myotis alcathoe 
182 Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 
NATURA SLOVENIAE 25(3): 181-200 
Apstrakt. Studija šišmiša u dolini gornje rijeke Neretve (Bosna i Hercegovina) – Dolina gornje rijeke 
Neretve sa rijekom i pritokama, očuvanim šumama, prirodnim pećinama i građevinama predstavlja vrlo interesantno 
područje za šišmiše. U dvije terenske ekspedicije početkom i krajem ljeta 2022. godine prikupljali smo podatke o 
šišmišima različitim metodama: pregledom potencijalnih skloništa, mreženjem i snimanjem eholokacije šišmiša. 
Koristili smo ručne i automatske ultrazvučne detektore na više lokacija uz i u blizini rijeke Neretve. Morfološka 
identifikacija nekih vrsta potvrđena je analizom DNK. Na području istraživanja, koje se proteže od Krupca do 
Konjica, otkrili smo najmanje 13 vrsta šišmiša. Uz još jednu, poznatu iz literature, u dolini rijeke Neretve pronađeno 
je ukupno 14 različitih vrsta šišmiša, što predstavlja skoro polovinu svih vrsta zabilježenih u Bosni i Hercegovini. 
Potvrdili smo prisustvo tri vrste koje su od najveće važnosti za očuvanje u Evropi, navedene u Aneksu II Direktive 
o staništima: mali potkovasti šišmiš (Rhinolophus hipposideros), veliki potkovasti šišmiš (Rhinolophus 
ferrumequinum) i veliki mišouhi šišmiš (Myotis myotis). Naš nalaz patuljastog brkatog šišmiša (Myotis alcathoe) 
predstavlja novu potvrdu prisutnosti vrste u Bosni i Hercegovini. Također, posmatranje dvobojnog šišmiša 
(Vespertilio murinus) je jedan od rijetkih zapisa ove vrste u zemlji. Gornja dolina rijeke Neretve može se smatrati 
područjem od velikog značaja za očuvanje šišmiša. Dodatni terenski rad, uključujući druga godišnja doba, otkrio bi 
više vrsta šišmiša i njihovih lokacija u tom području. 
 
Ključne riječi: Bosna i Hercegovina, rijeka Neretva, šišmiši, eholokacija, mreže, skloništa, Rhinolophus 
ferrumequinum, Rhinolophus hipposideros, Myotis myotis, Myotis alcathoe 
 
 
 
Introduction 
 
 
In the past two decades, knowledge on bats in Bosnia and Herzegovina (BIH) has increased. 
Many of the studies refer to inventories of bats at hibernation sites, mostly from caves (e.g. 
Presetnik et al. 2016; Mulaomerović et al. 2021), but also inventories using other methods, like 
mist netting and ultrasound detectors (e.g. Babić et al. 2018). To date, 31 bat species were 
recorded in BIH (Zagmajster et al. 2010; Babić et al. 2018), yet data on the distribution and 
ecology of some species remain scarce. 
 
While a study of bat fauna was conducted in the lower Neretva River valley (Mulaomerović 
et al. 2015), the upper Neretva River valley in Eastern Bosnia and Herzegovina was amongst 
the regions with virtually no records of bats. The region is covered with well preserved pristine 
forests (that extend around the Neretva River’s main stream and its tributaries), caves in karstic 
hills above the valley, pastures, and a few human settlements and buildings. All these 
characteristics present a high potential for a rich bat community. Yet, there were only three 
records for three species of bats from or in close vicinity of the upper Neretva River valley area 
prior to our study. In a small cave close to the village of Pridvorica u Borču (Borač under Dumoš 
planina), nine individuals of Rhinolophus hipposideros were observed on 29.9.2019 (J. 
Mulaomerović, pers. comm.). Mirić & Paunović (1997) reported on finding Nyctalus leisleri on 
15.3.1968 at Boračko jezero NW from Glavatičevo, within 3 km distance from the Neretva River. 
The same locality and date is given for the record of Pipistrellus pipistrellus, which is deposited 
in the Natural History Museum in Belgrade (Serbia) (Zagmajster et al. 2010).  
 
To fill this knowledge gap, a study of bats was conducted over a few days in June/July and 
August 2022, the first period being during the »Neretva Science Week 2022«. This event was 
organised to improve the knowledge on biodiversity of the area, which is threatened by a series 
of hydropower plants that are planned to be built on the upper Neretva River and its tributaries. 
Bats were studied by combining different methods: visual checking of the potential roosts in 
Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 183 
NATURA SLOVENIAE 25(3): 181-200 
caves and buildings, recording the echolocation calls, and mist netting. Here, we present the 
results which revealed a rich bat community despite short time of research. 
 
 
 
Materials and methods 
 
Study area 
 
We conducted the study in the upper Neretva River valley, extending from Krupac to Konjic 
(Fig. 1). The surrounding hills above the river and its tributaries are overgrown by forests, with 
few villages and individual buildings (many abandoned) present. The bedrock in the valley is 
formed of Cretaceous flysch at the foothills and Cretaceous carbonate, dolomite, and marl higher 
up on the slopes. Few caves are known from karstic hills surrounding the river valley. 
 
 
Figure 1. Map of the the upper Neretva River valley, with marked localities where we searched for bats in June/July and 
August 2022. Different symbols mark different study methods (explained in the legend, with details in the text). Numbers 
refer to localities, listed in Tab. 2. The embedded map on the left shows the position of the study area within Bosnia 
and Herzegovina, while the map on the top right presents a close up to Ulog area (marked with black frame on the 
main map).  
Slika 1. Zemljevid doline zgornje Neretve z označenimi kraji, kjer smo netopirje iskali junija/julija in avgusta 2022. Z 
različnimi simboli so označene različne metode preučevanja (pojasnjene v legendi, podrobnosti pa v besedilu). Številke 
se nanašajo na lokalitete, ki so navedene v Tab. 2. Vstavljena karta na levi prikazuje položaj območja raziskave znotraj 
Bosne in Hercegovine, medtem ko karta desno zgoraj prikazuje približano območje okoli Uloga (označeno s črnim 
okvirjem na večji karti).  
Slika 1. Karta doline gornje Neretve, sa označenim lokalitetima na kojima smo tražili šišmiše u junu/julu i augustu 2022. 
Različiti simboli označavaju različite metode istraživanja (objašnjeno u legendi, s detaljima u tekstu). Brojevi se odnose 
na lokalitete, navedene u Tab. 2. Ugrađena mapa pokazuje položaj istraživanog područja unutar Bosne i Hercegovine, a 
mapa u gornjem desnem kutu područje Uloga približano (markirano s crnim okvirom). 
184 Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 
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Checking the potential roosts 
 
During the day, we checked two caves, »Velika Đeverđela« (the whole cave) and »Mala 
Đeverđela« (only the entrance part), and a few abandoned buildings in the village Ulog. We 
also checked one small abandoned concrete building in Ulog during the night (Tab. 2). We 
looked for bats by using strong headlamps or handheld lights.  
 
 
Mist netting and handling of individuals 
 
On nine nights we set polyester mist nets (mesh size 1.5 cm, and up to 3 m height) at places 
where we anticipated the bats' flight paths: at the bank or over the surface of the Neretva River, 
on the forest edge, and amongst buildings of the village Ulog (Fig. 1; Tab. 2). We set the mist 
nets at the time of sunset (around 20:30), and took them down at least three hours later (around 
midnight). Night bat inventories were carried out when there was no rain and air temperatures 
exceeded 10°C. 
 
We checked the mist nets every few minutes to remove bats immediately after capture and 
placed them in a cloth bag. We measured the bats (weight, forearm length, other species-
specific characters), determined their age class according to the presence of cartilage in the 
epiphysal part of the metacarpal bones (Dietz & von Helversen 2004) and determined their 
reproductive status (Dietz & von Helversen 2004; Dietz et al. 2009; Dietz & Kiefer 2016). In 
some species, we took a small patch of wing membrane tissue with a puncher (1.5 mm 
diameter) and stored it in 96% ethanol for subsequent molecular analyses (Wilmer & Barratt 
1996). Before release, we marked each bat by cutting a tuft of hair (scapular or abdominal 
region) or by colouring the ear with a marker to recognise it in the case of recapture within the 
same night. 
 
 
Molecular analyses 
 
To confirm morphological identification in some cryptic species, we sequenced the the 
cytochrome C oxidase subunit I (COI) gene. We extracted the DNA from the punched wing 
tissue, using MagMAX DNA Multi-Sample Kit (Thermo Fisher Scientific), and amplified a standard 
barcoding marker, COI, using the Folmers primer pair, LCO1490 and HCO2198 (Folmer et al. 
1994). After purification we sent the PCR products for sequencing to Macrogen Europe 
laboratory (Amsterdam, Netherlands). We handled and assembled the sequences in Geneious 
Prime and compared them to the available sequences of bat species in the public database 
GenBank, using the Basic Local Alignment Search Tool (BLAST, NCBI) as a computational 
method. We submitted the COI sequences from this study to GenBank and we provide them in 
Tab. 3. 
 
 
  
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Ultrasound detectors and sound analysis 
 
We conducted bat surveys using ultrasound detectors on the nights without rain and with 
temperatures over 10°C, using two approaches: by recording bat calls manually using a hand 
ultrasound detector and by installing automatic ultrasound detectors (Tab. 2).  
 
We used hand D240x ultrasound detectors (Pettersson Elektronik AB) at or in the vicinity of 
the mist netting sites (Tab. 2). While in heterodyne mode, we scanned the frequencies (from 
20 to 110 kHz; mostly around 40 kHz) to detect bat calls. Once detected, we recorded them in 
10 x time expansion mode and stored them on an external digital recorder Roland RH-09. 
 
We set the automatic ultrasound detector (Song Meter SM4BAT FS, Wildlife Acoustics) at 
different sites along the Neretva River, and once in the village of Ulog (Tab. 2). The microphone 
was set at least 1.5 m above the ground, directed towards the river surface and its bank 
vegetation. The ultrasound detector recorded bats through the whole night, starting half an 
hour before sunset, and finishing at sunrise. The device recorded the full spectrum of sound for 
eight seconds with two seconds breaks, with no limits put on the sensitivity triggering the 
recording. We transferred all recordings to a PC and ran them through the Kaleidoscope software 
(Kaleidoscope Lite, Wildlife Acoustics) to produce the first separation of files with call sequences 
from the files with recorded noise. Due to the high number of automatic recordings, we checked 
only a subset of recordings per night to identify bat taxa. 
 
We analysed the files with call sequences using the sound analysis program Batsound 4.0 
(Pettersson Elektronik AB). We identified species or species groups based on characteristics of 
the echolocation calls, measured from spectrogram and power spectrum (FFT, 2048 samples, 
Hanning window), and using different references on bat calls (e.g. Russ 1999; Russo & Jones 
2002; Barataud 2014; Middleton et al. 2014; Dietz & Kiefer 2016). We checked the shape of the 
calls, start and end frequency, frequency of maximum energy, and call duration for 
determinations (Russ 1999; Russo & Jones 2002). In some cases, recognition to species level 
was not possible due to similar calls between species and variation within species. In these 
cases, calls were attributed to species groups or genera. 
 
 
 
Results and discussion 
 
General overview 
 
During the summer of 2022, we recorded at least 13 different bat species within 17 different 
bat taxa (Tabs. 1, 2). Groups of two or more species are given for some cases, where they could 
not be further identified with the methods used – but are nevertheless still informative about 
the presence of the bats. We confirmed the presence of two species of Rhinolophidae and at 
least 11 species of Vespertilionidae (Tabs. 1, 2), belonging to one and six genera, respectively. 
 
  
186 Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 
NATURA SLOVENIAE 25(3): 181-200 
Our results represent a considerable increase in the knowledge on bats in the area, as 
previously only single records of three species existed. Of those, we reconfirmed the presence 
of two (R. hipposideros, P. pipistrellus), while the third, N. leisleri (Mirć & Paunović 1997), could 
not be confirmed unambiguously based on echolocation calls we checked. 
 
Table 1. Overview of bat taxa (species or species groups) recorded during summer 2022 in the upper Neretva River 
valley in BIH, together with abbreviations of species names and number of localities where they were observed. The list 
of localities is in Tab. 2. 
Tabela 1. Pregled taksonov netopirjev (vrst ali skupin vrst), zabeleženih poleti 2022 v dolini zgornje Neretve v BIH, 
skupaj s kraticami imen vrst in številom lokalitet, kjer so bili opaženi. Seznam lokalitet je v Tab. 2. 
Tabela 1. Pregled vrsta šišmiša (vrsta ili grupa vrsta) zabilježenih u ljetu 2022. godine u dolini gornje Neretve u BiH, 
zajedno sa skraćenicama naziva vrsta i brojem lokaliteta, na kojima su uočeni. Spisak lokaliteta nalazi se u Tab. 2. 
 
Most of the findings refer to observations with ultrasound detectors at the Neretva River 
bank which recorded the bats on their flight paths; either when they were flying by, or were 
present near the point of observation whilst hunting at the bank’s vegetation or above the river 
surface. In some cases, the bats were observed near the street lights in villages or near the 
houses, where many species hunt for insects attracted to light. This approach gathered a lot of 
information on bat presence, but in many cases exact determination to species level was not 
possible due to ther similarity of the echolocation calls (Barataud 2014; Middleton et al. 2014). 
We identified at least five different bat species with this approach  
(Tab. 2). 
 
Five out of nine mist netting evenings/nights were successful in catching at least one 
individual bat (Tab. 2). Even though this approach is time consuming, and limited to one site 
per night, it allows for the most exact species identifications, as individuals are held in the hand 
and directly observed and allow for samples for DNA analyses to be taken. We confirmed the 
presence of ten different bat species using this method (Tab. 2). 
 
No. Family 
Species/species group 
Abb. No. of 
localities 
 Rhinolophidae   
1 Rhinolophus hipposideros (Bechstein, 1800) Rhip 5 
2 Rhinolophus ferrumequinum (Schreber, 1774) Rfer 2 
 Vespertilionidae   
3 Myotis myotis (Borkhausen, 1797) Mmyo 1 
4 Myotis nattereri (Kuhl, 1817) Mnat 1 
5 Myotis mystacinus (Kuhl, 1817) Mmys 2 
6 Myotis alcathoe O. von Helversen & K.-G. Heller, 2001 Malc 1 
7 Myotis sp. Myosp 6 
8 Pipistrellus pipistrellus (Schreber, 1774) Ppip 4 
9 Pipistrellus pygmaeus (Leach, 1825) Ppyg 2 
10 Pipistrellus nathusii (Keyserling & Blasius, 1839) Pnat 1 
11 Pipistrellus kuhlii/nathusii Pkuh/nat 5 
12 Hypsugo savii (Bonaparte, 1837) Hsav 7 
13 Nyctalus noctula (Schreber, 1774) Nnoc 1 
14 Nyctalus noctula/lasiopterus Nnoc/las 2 
15 Vespertilio murinus Linnaeus, 1758 Vmur 1 
16 Vespertilio/Nyctalus/Eptesicus V/M/E 6 
17 Plecotus sp.  Plesp 1 
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Table 2. List of localities with bat taxa (species or species group) recorded in summer 2022 in upper Neretva River valley 
in BIH (number of individuals given in brackets in the column Taxa). Geographical coordinates are in decimal degrees 
(WGS84). Abbreviations: Method: OB – observation, RC – roost check, MN – mist netting, AUD – automatic ultrasound 
detector, HUD – hand ultrasound detector; Leg/Det. – legator/determinator:  AV – Anton Vorauer, BR – Behare 
Rexhepi, EP – Ester Premate, SA - Stefan Andjus, ŠB – Špela Borko, VM – Vojo Milanović, MZ – Maja Zagmajster. Taxa 
abbreviations are in the Tab. 1.  
Tabela 2. Seznam lokalitet s taksoni netopirjev (vrstami ali skupinami vrst), zabeleženimi poleti 2022 v dolini zgornje 
Neretve v BIH (število osebkov je podano v oklepaju  v stolpcu Taxa). Geografske koordinate so v decimalnih stopinjah 
(WGS84). Okrajšave: Metoda: OB – opazovanje, RC – preverjanje kotišč, MN – lovljenje z mrežo, AUD – avtomatski 
ultrazvočni detektor, HUD – ročni ultrazvočni detektor; Leg/Det – legator/določevalec: AV – Anton Vorauer,  
BR – Behare Rexhepi, EP – Ester Premate, SA – Stefan Andjus, ŠB – Špela Borko, VM – Vojo Milanović, MZ – Maja 
Zagmajster. Okrajšave taksonov so v Tab. 1. 
Tabela 2. Spisak lokaliteta vrtsa šišmiša (vrste ili grupe vrsta) evidentiranih tokom ljeta 2022. godine u dolini gornje 
Neretve u BiH (broj jedinki je u zagradama u koloni Taxa). Geografske koordinate su u decimalnim stepenima (WGS84). 
Skraćenice: Metoda: OB – posmatranje, RC – provjera skloništa, MN – mreženje, AUD – automatski ultrazvučni 
detektor, HUD – ručni ultrazvučni detektor; Leg/Det. – legator/određivać: AV – Anton Vorauer, BR – Behare Rexhepi, 
EP – Ester Premate, SA - Stefan Andjus, ŠB – Špela Borko, VM – Vojo Milanović, MZ – Maja Zagmajster. Skraćenice 
taksona nalaze se u Tab. 1. 
No Locality Coordinates Date Method Leg./ 
Det. 
Taxa Comments 
1 Ulog, near 
Mosque/Church 
43.41590, 
18.31067 
27.6. 
2022 
OB AV/ 
AV 
Plesp  
(1 flying) 
Observed in flight 
2 Ulog, downstream 
Camp 
43.42299, 
18.31037 
27.6. 
2022 
MN AV/ 
AV 
Nnoc (1), 
Vmur (1) 
Mist nets set at 
the river bank 
3 At the right bank of the 
Neretva, downstream 
from Swimming Beach, 
Ulog 
43.42485, 
18.30973 
28./29.6. 
2022 
AUD MZ/
MZ 
Rfer, 
Myosp, 
Ppip, 
Ppyg, 
Pkuh/nat, 
Hsav, 
Nnoc/las, 
V/N/E 
Microphone set 
toward the river  
4 At the right bank of the 
Neretva, upstream of 
the Cerova location 
43.37879, 
18.35610 
29./30.6. 
2022 
AUD MZ/
MZ 
Myosp, 
Pkuh/nat, 
Hsav, 
Nnoc/las, 
V/N/E 
Microphone set 
toward the river  
5 Near the bridge over 
Neretva, 600 m 
downstream of the 
Krupac confluence, 
Southeast from the 
farm 
43.33218, 
18.41821 
29.6. 
2022 
MN MZ, 
VM, 
(EP, 
SB, 
BR, 
SA)/
MZ, 
VM 
Mmyo (2), 
Mnat (1), 
Mmys (2), 
Malc (4), 
Pnat (1), 
Ppyg (1) 
Five mist nets set 
up, one (15 m) 
over the river, 
one near the 
forest edge (7 m), 
three over small 
dry channel (7 m, 
6 m, 10 m) 
6 Ulog »beach« 43.42420, 
18.30857 
29.6. 
2022 
MN AV/ 
AV 
Rfer (1)  
7 Ulog, between two 
houses, south from the 
bar-house, near the 
Cinema 
43.41648, 
18.31223 
30.6. 
2022 
MN AV, 
VM/ 
AV, 
VM 
Ppip (1) Two mist nets set 
up 
8 Velika Đeverđela cave, 
Ulog 
43.40587, 
18.28429 
30.6. 
2022 
BR MZ/
MZ 
Rhip (1 
humerus) 
In the chamber 
just after the 
vertical drop 
188 Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 
NATURA SLOVENIAE 25(3): 181-200 
No Locality Coordinates Date Method Leg./ 
Det. 
Taxa Comments 
9 Mala Đeverđela cave, 
Ulog 
43.40660, 
18.28416 
 
30.6. 
2022 
RC MZ/
MZ 
Rhip  
(1 flying) 
In the entrance 
chamber, before 
the vertical drop 
10 At the street light in 
Ulog, near the first road 
curve South from the 
Neretva 
43.41578, 
18.31244 
30.6./1.7.
2022 
AUD MZ/
MZ 
Pkuh/nat, 
Hsav, 
V/N/E 
 
11 At the right bank of 
Neretva River, 
Northeast from the bar 
house in Ulog 
43.41509, 
18.31475 
1.7. 
2022 
MN MZ, 
VM/- 
/ Two mist nets:  
6 m at the gravel 
road; 15 m above 
the river 
    HUD MZ/
MZ 
Myosp, 
Ppip, 
Hsav, 
Nnoc/las, 
V/N/E 
D240x 
Pettersoon, at the 
river bank, 
recordings 
12 Ulog, above the 
Mosque/Church old 
house 1 
43.41095, 
18.31061 
1.7.2022 RC AV/ 
AV 
Rhip (6) Pregnant females 
– visual 
observation 
13 Ulog, above the 
Mosque/Church old 
house 2 
43.41091, 
18.31061 
1.7.2022 RC AV/ 
AV 
Rhip (8) Pregnant females 
– visual 
observation 
14 Ulog, meadow east of 
the Camp 
43.41533, 
18.31282 
1.7.2022 MN AV/- /  
15 Abandoned small 
building, north from 
Neretva, Southeast 
from Ulog 
43.41507, 
18.31518 
1.7.2022 RC MZ/
MZ 
Rhip (1) Active, left the 
room when we 
checked on it (at 
about midnight) 
16 At the right bank of 
Neretva River, below 
the Grand River 
Adventure Ranch, 
Glavatičevo 
43.51044, 
18.09880 
2.7.2022 MN VM/- /  
    HUD VM, 
MZ/
MZ 
Myosp, 
Pkuh/nat, 
Hsav, 
V/N/E 
 
17 At the bank of Neretva 
River at the Brijestov 
bridge 
43.482282, 
18.226530 
4.8.2022 MN VM/- – Three mist nets:  
9 m, 3 m, 6 m 
above the river 
    AUD VM/
MZ 
Myosp, 
Hsav, 
V/N/E 
 
18 At the bank of Neretva 
River near Nedavic 
43.45797, 
18.32121 
5.8.2022 MN VM/ 
VM 
Mmys (3) Two mist nets:  
9 m and 6 m 
above the river 
    AUD VM/
MZ 
Hsav, 
Pkuh/nat, 
Myosp, 
V/N/E 
 
 
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There were few observations of bats in roosts, in one case this was only based on finding 
bony remnants which confirmed the species’ presence in the cave (Tab. 2). We recorded roosts 
of only one species, R. hipposideros, in a few buildings and caves, as this is the species that can 
be more easily observed due to hanging in exposed places (Dietz et al. 2009). In the short time 
for this research, only a few potential roosts could be checked, even though there were many 
more potentially suitable in the wider Neretva River valley (not only in buildings, but also in 
caves, rocky walls, and tree trunks).  
 
The species richest site during our survey was at the Neretva River near the Krupac 
confluence, where six different bat species were caught in mist nets (Tab. 2). Mist nets were 
set at the edge of the forest next to the Neretva, as well as over a small side river channel next 
to the main river bed. This confirms that forests around the river support a very diverse bat 
community, including species that are typical forest species (M. mystacinus, M. alcathoe) or  
species hunting near riparian vegetation (Pipistrellus spp.). Even though other mist netting 
events revealed fewer bat species, one to two, they still revealed the presence of rarely 
encountered species. These were mist nettings at the Neretva River, both in the wider 
surroundings of Ulog (Tab. 2). 
 
Measurements of bat individuals are within the typical ranges of all the species (Tab. 3). 
Most of the females had signs of lactation, indicating that they were rearing a juvenile. This 
further supports their presence in the valley and the importance of the area for reproduction of 
at least four bat species (Tab. 3). The tissues taken from some bats enabled exact identification 
to species level (see comments in the next section). 
 
Table 3. Measurements of the bats mist netted in summer 2022 in the upper Neretva River valley in BIH. For species 
abbreviations see Tab. 1, for localities and dates see Tab. 2. Abbreviations: FA - forearm length, W - weight, Repr.- 
reproduction state, M - male, F - female, Ad - adult, Sad - subadult, D1 - length of the first finger, D5 - length of the 
fifth finger, P3.2 - length of the second phalang of the third finger, P3.3 - length of the third phalang of the third finger, 
CM3 - length of the upper tooth row from canine to 3rd molar, P - premolar, in upper (superscripted number) or lower 
(subscripted number) tooth row. 
Tabela 3. Meritve netopirjev, ujetih v mreže poleti 2022 v zgornjem toku reke Neretve v BIH. Za kratice vrst glej Tab. 1, 
za nahajališča in datume glej Tab. 2. 2. Okrajšave: FA - dolžina podlakti, W - teža, Repr.- reprodukcijsko stanje, M - 
samec, F - samica, Ad - odrasel, Sad - subadult, D1 - dolžina prvega prsta, D5 - dolžina petega prsta, P3.2 - dolžina 
druge prstnice tretjega prsta, P3.3 - dolžina tretje prstnice tretjega prsta, CM3 - dolžina zgornje vrste zob od kanina do 
tretjega molarja, P - premolar, v zgornji (nadnapisana številka) ali spodnji (podnapisana številka) vrsti zob. 
Tabela 3. Mjerenja šišmiša koji su uhvaćeni u mreže tokom ljeta 2022. godine u dolini gornje Neretve u BiH. Za 
skraćenice vrsta vidi Tab. 1, za lokalitete i datume vidi Tab. 2. Skraćenice: FA - dužina podlaktice, W - težina, Repr.- 
stanje reprodukcije, M - mužijak, Ž - ženka, Ad - odrasla jedinka, Sad - pododrasla jedinka, D1 - dužina prvog prsta, D5 
- dužina petog prsta , P3.2 - dužina druge falange trećeg prsta, P3.3 - dužina treće falange trećeg prsta, CM3 - dužina 
gornjeg zubnog reda od očnjaka do 3. kutnjaka, P - premolar, u gornjem (nadpisani broj) ili donjem (potpisani broj) 
zubnom redu. 
Loc Time Species Sex Age FA 
[mm] 
W 
[g] 
Repr. Remarks, other measurements 
2 / Vmur M Ad 42.4 15 Inactive D5: 48.5 mm, D1: 6,1 mm 
2 / Nnoc M Ad 54.4 30 Inactive  
5 21:40 Mmyo M Ad 59.8 29.5 Active Enlarged testices, CM3: 10.5 mm, strong 
smell, black tip on tragus 
5 21:41 Mmyo M Ad 60.9 28.0 Active Enlarged testices, CM3: 10.2 mm, strong 
smell, black tip on tragus 
5 21:15 Mnat M Ad 38.1 7.0 Inactive Black epididymis; GenBank Acc.No. 
OR487194 
190 Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 
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Loc Time Species Sex Age FA 
[mm] 
W 
[g] 
Repr. Remarks, other measurements 
5 22:00 Mmys F Ad 34.4 7.0 Lactating D1: 6.0 mm, Tibia: 17.8 mm, dark ear, 
face brownish-pale 
5 22:26 Mmys F Sad 34.8 6.1 Nuliparous D1: 6.0 mm, Tibia: 17.6 mm, Foot:  
5.6 mm, dark ear, greyish, young 
5 21:15 Malc M Ad 33.1 4.5 Active Enlarged testices, D1: 5.3 mm, Tibia: 
15.6 mm, Foot: 4.4 mm, Upper teeth 
row: no cing obvious (Dietz et al. 2009). 
Species first identified as Mmys, but 
allocated to Malc according to DNA; 
GenBank Acc.No. OR487193  
5 21:43 Malc F Ad 32.8 5.5 Lactating D1: 4.3 mm, Tibia: 14.5 mm, Foot: 4.3 
mm, bright face; GenBank Acc. No. 
OR487195 
5 22:00 Malc F Ad 32.3 5.0 Lactating D1: 4.1 mm, Tibia: 14.9 mm, Foot: 4.4 
mm, light face, bright ears; GenBank 
Acc.No. OR487196 
5 22:00 Malc M Ad 31.6 4.0 Inactive D1: 4.9 mm, Tibia: 14.5 mm, Foot: 5.3 
mm, light face, black epididymis; 
GenBank Acc.No. OR487197 
5 22:30 Pnat M Ad 34.3 8.5 Active Enlarged testices, epidydimis slightly 
black, D5: 43.8 mm, upper teeth like Pnat 
(Dietz & von Helversen 2004); Ring: 
BIHA0140 
5 21:15 Ppyg F Ad 30.5 5.0 Lactating D5: 37.5 mm, P3.2: 7.7 mm, P3.3: 6.7 
mm, internal edge of the ear: 7.6 mm, 
orange glands (buccal and around 
vagina)  
6 / Rfer M Ad 54.7 24 Inactive  
7 23:00 Ppip F Ad 30.0 6.0 Lactating Wing membrane as in Dietz et al. (2004) 
for Ppip, echolocation upon release – 
around 45 kHz 
18 21:10 Mmys F Sad 36.0 6.0 Nuliparous No protocone, P3 < 2/3 P2; P3 < P2 ; dark 
ears, face; greyish fur, cartilage in joints; 
Ring: BIHA0114; GenBank Acc.No. 
OR487200 
18 / Mmys F Ad 34.9 5.0 Lactated No protocone, P3 ~ 2/3 P2; P3 < P2 ; dark 
ears, face; brownish fur; Ring: 
BIHA0115; GenBank Acc.No. OR487201 
18 21:20 
 
Mmys M Sad 35.9 5.5  No protocone, P3 < 2/3 P2; P3 < P2 ; dark 
ears, face; greyish fur, cartilage in joints; 
Ring: BIHA0116; DNA did not amplify 
 
  
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Comments to bat species 
 
Rhinolophidae, Rhinolophus spp. 
 
We recorded two species of the genus during our study, the smallest (R. hipposideros) and 
the largest (R. ferrumequinum) of this family in Europe (Dietz et al. 2009). Both are known to 
use caves and parts of buildings as shelters. In our study we only found roosts of  
R. hipposideros by recording individuals or small groups of individuals in some abandoned 
buildings and a cave (Tab. 2). The groups of individuals in some of the buildings in Ulog  
(Tab. 2) indicate that these were females forming nursery colonies. Individuals had fat bellies 
and looked to be close to gestation period (Fig. 2). 
 
We caught only one male of R. ferrumequinum in the mist net set near the Neretva River 
near Ulog (Tab. 2), but did not find any roosts of the species. Its presence in the region was 
further supported with the species echolocation calls detected at one site near the Neretva River 
near Krupac (Tab. 2). The species is known to feed in riparian habitats (Dietz et al. 2019), and 
considering the preserved habitats it could be expected to be more common in the Neretva 
River valley. There was an old record of the species from the town of Nevesinje (Felten et al. 
1977), but our observations confirm its current presence in the upper Neretva River valley.  
 
 
Figure 2. A group of Lesser horseshoe bats (Rhinolophus hipposideros) observed in an abandoned building in Ulog. Due 
to fat bellies of the individuals, and typical behaviour in forming summer groups, this is very likely a group of females 
forming a nursery colony (Photo: A. Vorauer). 
Slika 2. Skupina malih podkovnjakov (Rhinolophus hipposideros) opazovana v zapuščeni stavbi v Ulogu. Zaradi debelih 
trebuhov osebkov in značilnega vedenja pri oblikovanju poletnih skupin je zelo verjetno, da gre za skupino samic v 
porodniški koloniji (Foto: A. Vorauer). 
Slika 2. Grupa malih potkovastih šišmiša (Rhinolophus hipposideros) uočena u napuštenoj zgradi u Ulogu. Zbog debelih 
trbuha jedinki i tipičnog ponašanja pri formiranju ljetnih grupa, vrlo je vjerovatno da je riječ o grupi ženki koja formira 
prodničnu koloniju (Foto: A. Vorauer). 
192 Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 
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Vespertilionidae, Myotis spp. 
 
The biggest species of this genus, Myotis myotis, uses caves and abandoned buildings as 
roosts, and feeds mostly in the forest openings or along forest roads (Dietz et al. 2019). We 
caught one male (Fig. 3) at the Neretva River near Krupac, flying out of the forest at the river 
bank.  
 
 
Figure 3. The greater mouse-eared bat (Myotis myotis) caught in the mist net close to the Krupac confluence with 
Neretva (photo: Ester Premate).  
Slika 3. Navadni netopir (Myotis myotis), ujet v mrežo v bližini sotočja Krupca z Neretvo (foto: Ester Premate). 
Slika 3. Veliki mišouhi šišmiš (Myotis myotis) uhvaćen u mrežu u blizini ušća Krupca u Neretvu (foto: Ester Premate). 
 
Myotis nattereri is a medium sized species of the genus, being a typical forest species, 
dwelling in crevices in tree trunks, and feeding among the forest vegetation (Dietz et al. 2019). 
We caught one male at the Neretva River near Krupac, flying out of the forest at the river bank 
(Tab. 2). Due to a suitable habitat for roosting and feeding, it can be predicted that this species 
is resident here and common in the upper Neretva River area.  
 
There were two sites at which individuals of the smallest species of the genus were mist 
netted, belonging to the Myotis mystacinus species complex (M. mystacinus, brandtii, alcathoe, 
davidii, Dietz & Kiefer 2016). It is a group of morphologically similar species, where 
discrimination is not always unambiguous. All species of the complex are forest species, roosting 
in tree hollows, and feeding in the forest (Dietz et al. 2019). M. alcathoe’s description has been 
supported by using molecular differences (von Helversen et al. 2001; Dietz et al. 2009). In order 
to reliably identify species, we used DNA barcoding to help identify the species of this complex. 
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One of the animals first morphologically identified as M. mystacinus in the field was genetically 
identified as M. alcathoe, proving the difficulty in morphological discriminations of the species 
(Tab. 3). Also on the European level, distribution of this species is full of knowledge gaps, mainly 
due to under-sampling of the habitats of this bat species and misidentifications during fieldwork 
(Jan et al. 2010; Bashta et al. 2011). For example, the presence of M. alcathoe in Portugal was 
also confirmed using DNA barcoding, after an individual, morphologically determined as M. 
mystacinus, was analysed molecularly (Rebelo et al. 2020).  
 
Myotis alcathoe was confirmed only at Neretva near Krupac site, flying in the net set at the 
forest edge near Neretva. The first record of the species in the country comes from Sarajevo 
area, and was based on morphological characters only (Babić et al. 2018). Later, the species 
was confirmed for eastern Bosnia and Herzegovina also with molecular analyses (Verhees et al. 
2021). As Myotis alcathoe is a species bound to riparian and forest habitats, more findings of 
the species in the upper Neretva River could be expected. Proper management of the forest is 
crucial for preservation of the species (von Helversen et al. 2001; Coronado et al. 2017).  
 
The other bat species of the complex are here referred to as Myotis mystacinus (Tabs. 1,2). 
However, in a recent study, it has been suggested that Myotis davidii populations may have 
historically replaced the M. mystacinus s.str. in the Balkans, where even hybridising zones 
between the species may exist (Çoraman et al. 2020). Discrimination between two species is 
difficult both morphologically or based on mitochondrial DNA (Benda et al. 2016; Dietz & Kiefer 
2016; Çoraman et al. 2020). We report M. mystacinus s.l. here, but its exact species status, not 
only in the upper Neretva River valley but probably in the whole Bosnia and Herzegovina, 
remains to be resolved.  
 
The species of the genus Myotis sp. have characteristic frequency modulated shape 
echolocation calls, which are very similar to one another and species can only rarely be identified 
to species/species group level. The calls recorded near the Neretva River surface most likely 
belong to the species group M. daubentonii/capaccinii; species that typically hunt over water 
surfaces (Dietz et al. 2009). Due to this, their calls show typical interruptions due to the 
reflection from the water surface (Russ 1999) (Tab. 2). Further analyses of bat calls, as well as 
additional field work, including mist netting, may reveal further bat species of this genus in the 
area.   
 
Vespertilionidae, Pipistrellus spp. & Hypsugo savii 
 
Pipistrellus pipistrellus and P. pygmaeus are morphologically very similar, and were only 
recognised as separate species three decades ago (Jones & van Parijs 1993; Barlow & Jones 
1997). The two species can be distinguished based on echolocation calls (Russo & Jones 2000; 
Barataud 2014). In our study, we detected P. pygmaeus more often, whilst both species were 
caught in the mist nets. Pipistrellus pygmaues is a species that is more bound to riparian 
habitats, and indeed it was recorded at sites near the Neretva River (Tab. 1, 2).   
 
It was less than ten years ago that the species P. nathusii was confirmed to occur in BIH 
(Karapandža et al. 2014), with a few records in subsequent years (Husanović & Presetnik 2021). 
Our finding of the species at the Krupac site near the Neretva River is therefore one of the rare 
findings in the country. Part of the reason that the species distribution is poorly known could be 
the fact that its echolocation calls overlap with the echolocation calls of the species P. kuhlii, to 
194 Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 
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such an extent that discrimination is not possible (Kalko 1995; Russo & Jones 1999; Barataud 
2014; Dietz & Kiefer 2016). As the two species cannot be reliably discriminated based on 
echolocation calls, such observations are allocated to a species group P. kuhlii/nathusii, which 
was recorded in almost all sites where ultrasound detector observations were made (Tabs. 1, 
2). Social calls of the species are species-specific, and they were used to confirm the presence 
of P. nathusii during autumn in Mostar (Husanović & Presetnik 2021).  
 
Hypsugo savii was confirmed at all sites where bats were recorded with ultrasound detectors, 
showing the species is generally widespread in the region. The species typically roosts in rocky 
walls, but also in building walls, which are both present in the Neretva River valley. 
 
Vespertilionidae, other bat taxa recorded 
 
Species of the group Vespertilio murinus, Nyctalus spp., Eptesicus spp. (»V/N/E« in  
Tabs. 1, 2) use echolocation with a lot of overlap, therefore we conservatively used this call 
determination category for such cases. Even though this category includes the genus Eptesicus, 
we cannot confirm its presence based on our results, whilst we did detect species of the other 
two genera in our study (Tabs. 1, 2). 
 
In some cases we were able to identify species of the group Nyctalus noctula/Nyctalus 
lasiopterus, due to two characteristic types of echolocation calls, and low call frequency (Russo 
& Jones 2002; Barataud 2014). We decided to remain conservative in separating the two 
species, due to large frequency overlap, and kept the group category. However, some studies 
have shown that discrimination between the two species is possible in some cases (Estók & 
Siemers 2009). Considering that we mist netted the male of N. noctula, it is most probable that 
most of the echolocation calls belong to this species.  
 
Based on the amount of recording analyses for this study, we could not yet unambiguously 
confirm the presence of N. leisleri. The species is expected in the area as it is bound to large 
forested areas. The only reliable confirmation of species presence so far is the report from near 
Boračko jezero, close to Glavatičevo (Mirić & Paunović 1997).  
 
Reliable discrimination of V. murinus is possible only based on male display song calls 
(Zagmajster 2003), but they were not recorded during our study, as they are typically emitted 
during the mating season in autumn. The confirmation of the species presence in the Neretva 
river valley was therefore possible due to a successful mist netting event close to the Ulog 
region. The species typically feeds in riparian zones, finding many suitable habitats within the 
Neretva River valley (Dietz et al. 2009).   
 
The observation of the Plecotus bat is so far based on the visual observation of the flying 
individual made by AV. It is highly likely that the species of this genus are present in the area, 
yet the only data existing to date are from about 14 km from Ulog, close to the region of Zalomka 
– where echolocation calls of Plecotus auritus were recorded (Mulaomerović 2022).  
 
  
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Conservation notes 
 
We detected at least 13 bat species in the study area extending from Krupac to Konjic. With 
one additional species, known from the literature, 14 different bat species have been found in 
the upper Neretva river valley, presenting nearly a half of all bat species recorded in Bosnia and 
Herzegovina. The rich bat community in the Neretva River valley, despite the short time of 
research, proved that this area is potentially very rich in bat species and of high importance for 
conservation. All bat species are protected under the Bern convention, which is the basis for the 
designation of Emerald protection sites - since 2023, the upper Neretva River valley has been 
amongst such proposed sites (EEA 2023). Bats as migratory species are protected under the 
Bonn Convention (OJ L 1982), and a special agreement of this convention dedicated specifically 
to bat protection (Eurobats 1991), which was also officially signed by Bosnia and Herzegovina 
in 2021.  
 
In European Union (EU) member states, one of the most powerful international documents 
for the protection of bats is the EU Habitats Directive (OJ EC 1992). As accession to the EU is 
also a path being taken by Bosnia and Herzegovina, under this directive, one of the obligations 
shall be the list of proposed Sites of Community Importance. In this EU document, all bats are 
protected (Annex IV), but there are some species of even higher conservation concern, which 
need designation of areas for protection (Annex II). In our study, we recorded three bat species 
with the highest conservation concern in the Neretva River valley: R. ferrumequinum,  
R. hipposideros and M. myotis. According to the EU Habitats Directive, these species need the 
designation of special areas for their conservation. The upper Neretva River valley could be such 
an area for bats and should be included in such an European conservation network. 
 
Planned constructions in the Neretva River valley, including a set of hydropower plants and 
their corresponding reservoirs, accompanied by forest cuts due to the building processes, 
threaten to significantly change the pristine nature of the area. This will consequently change 
the habitats for bats, which have been shown to suffer from hydropower constructions and 
related changes of natural habitats (Zortéa et al. 2021). Considering the rich bat community in 
the area, hydropower plants on the Neretva River present a significant threat to bats in the 
area. It is also highly important to conduct further studies of bats in the area, including other 
seasons, to get a more complete overview of the bat diversity in the area. 
 
 
  
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Povzetek  
 
 
V BIH je bilo doslej zabeleženih 31 vrst netopirjev (Zagmajster et al. 2010; Babić et al. 2018), vendar je 
podatkov o razširjenosti in ekologiji nekaterih vrst malo. Za tri vrste netopirjev so bili z območja zgornje 
doline reke Neretve ali iz njene neposredne bližine zabeleženi le trije zapisi: Rhinolophus hipposideros je bil 
opažen 29.9.2019 (J. Mulaomerović, pers. comm.). Mirić & Paunović (1997) poročata o najdbi Nyctalus 
leisleri 15.3.1968 pri Boračko jezero SZ od Glavatičeva, v oddaljenosti 3 km od reke Neretve. Isto nahajališče 
in datum sta navedena tudi pri zapisu o Pipistrellus pipistrellus, ki je shranjen v Prirodoslovnem muzeju v 
Beogradu (Srbija) (Zagmajster et al. 2010). Da bi zapolnili to vrzel v znanju, smo v juniju/juliju in avgustu 
2022 opravili raziskavo netopirjev, pri čemer je bilo prvo obdobje v času »Neretvanskega tedna znanosti 
2022«. 
 
Raziskavo smo opravili v zgornjem delu doline reke Neretve, ki se razteza od Krupca do Konjica (Sl. 1). 
Čez dan smo pregledali dve jami, »Velika Đeverđela« (celotno jamo) in »Mala Đeverđela« (samo vhodni 
del), ter nekaj zapuščenih stavb v vasi Ulog. Devet noči smo postavili poliestrske mreže za meglo na mestih, 
kjer smo predvidevali preletne poti netopirjev (Sl. 1; Tab. 2). Za potrditev terenskih določitev pri nekaterih 
kriptičnih vrstah smo iz tkiva, odvzetega nekaterim netopirjem, sekvencirali gen COI. Preiskave netopirjev z 
ultrazvočnimi detektorji smo izvajali v nočeh brez dežja in ob temperaturah nad 10 °C, in sicer z ročnimi in 
avtomatskimi ultrazvočnimi detektorji (Tab. 2). 
 
Med 17 različnimi taksoni netopirjev smo zabeležili vsaj 13 različnih vrst netopirjev (Tab. 1, 2). Skupine 
dveh ali več vrst so navedene v nekaterih primerih, ko jih z uporabljeno metodo ni bilo mogoče natančneje 
določiti - vendar so še vedno informativne glede prisotnosti netopirjev. Potrdili smo prisotnost dveh vrst iz 
družine Rhinolophidae (Rhinolophus hipposideros, Rhinolophus ferrumequinum) in vsaj 11 vrst iz družine 
Vespertilionidae znotraj 15 različnih taksonov (Myotis myotis, Myotis nattereri, Myotis mystacinus s.l., Myotis 
alcathoe, Myotis sp., Pipistrellus pipistrellus, Pipistrellus pygmaeus, Pipistrellus nathusii, Pipistrellus 
kuhlii/nathusii, Hypsugo savii, Nyctalus noctule, Nyctalus noctula/lasiopterus, Vespertilio murinus, 
Vespertilio/Nyctalus/Eptesicus spp., Plecotus sp.) Zabeležili smo tudi nekaj redko opaženih vrst v BIH (Myotis 
alcathoe, Vespertilio murinus). 
 
Večina ugotovitev se nanaša na opazovanja z ultrazvočnimi detektorji. V petih od devet noči, ko smo 
lovili z mrežami, smo uspešno ujeli vsaj enega netopirja (Tab. 2). S to metodo smo potrdili prisotnost desetih 
različnih vrst netopirjev (Tab. 2). Le pri eni vrsti, R. hipposideros, smo zabeležili zatočišča v nekaj stavbah 
in jamah, od katerih so nekatere najverjetneje kotišča. Vrstno najbogatejša lokacija je bila na reki Neretvi 
pri sotočju s Krupcem, kjer smo v mreže ujeli šest različnih vrst netopirjev.  
 
Bogata združba netopirjev, ki smo jo odkrili v dolini reke Neretve kljub kratkemu času raziskave,  je 
pokazala, da je to območje potencialno zelo bogato z netopirji in zelo pomembno za njihovo ohranjanje. 
Vse vrste netopirjev so zavarovane. V skladu z EU Habitatno direktivo (OJ ES 1992) je treba za nekatere 
vrste netopirjev predlagati potencialno območje, pomembno za Skupnost (Priloga II). V naši raziskavi smo 
v dolini reke Neretve zabeležili tri take vrste netopirjev, ki so najpomembnejše za varstvo:  
R. ferrumequinum, R. hipposideros in M. myotis. 
 
Načrtovane gradnje v dolini reke Neretve, vključno z nizom hidroelektrarn in pripadajočih zadrževalnikov, 
ki jih spremlja krčenje gozdov, predstavljajo nevarnost, da se bistveno spremeni neokrnjena narava 
območja. To bo posledično spremenilo habitate za netopirje, na katere ima gradnja hidroelektrarn dokazano 
negativen vpliv (Zortéa et al. 2021). Glede na bogato vrstno pestrost lahko dolino zgornjega toka reke 
Neretve štejemo za območje velikega varstvenega pomena za netopirje. 
 
  
Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 197 
NATURA SLOVENIAE 25(3): 181-200 
Sažetak 
 
 
U BiH je do sada zabilježena 31 vrsta šišmiša (Zagmajster et al. 2010; Babić et al. 2018), ali podataka o 
rasprostranjenosti i ekologiji nekih vrsta je malo. Za tri vrste šišmiša iz ili u neposrednoj blizini gornje doline 
rijeke Neretve zabilježena su samo tri zapisa: Rhinolophus hipposideros uočeni su 29.9.2019  
(J. Mulaomerović, pers. comm.). Mirić & Paunović (1997) izvještavaju o pronalasku Nyctalus leisleri 
15.3.1968 na Boračkom jezeru SZ od Glavatičeva, na udaljenosti od 3 km od rijeke Neretve. Isti lokalitet i 
datum dat je i za zapis Pipistrellus pipistrellus, koji se nalazi u Prirodnjačkom muzeju u Beogradu (Srbija) 
(Zagmajster et al. 2010). Kako bi se popunila ova praznina u znanju, nekoliko dana u junu/julu i augustu 
2022. godine provedeno je istraživanje šišmiša, a prvi period je bio tokom »Neretvanske sedmice nauke 
2022.«. 
 
Istraživanje smo proveli u dolini gornje Neretve, koja se proteže od Krupca do Konjica (Sl. 1). Tokom 
dana smo proverili dve pećine, »Velika Đeverđela« (cela pećina) i »Mala Đeverđela« (samo ulazni deo), i 
nekoliko napuštenih objekata u selu Ulog. Devet noći smo postavljali poliesterske mreže na mjestima gdje 
smo predviđali puteve leta šišmiševa (Sl. 1; Tab. 2). Da bismo potvrdili terenska određivanja kod nekih 
kriptičnih vrsta, sekvencirali smo COI gen iz tkiva uzetog od nekih vrsta. Istraživanja vršili smo ručnim ili 
automatskim detektorima u noćima bez kiše i pri temperaturama preko 10°C (Tab. 2). 
 
Zabilježili smo najmanje 13 različitih vrsta šišmiša unutar 17 različitih taksona šišmiša (Tab. 1, 2). Za 
neke slučajeve date su grupe od dvije ili više vrsta, gdje se nisu mogle dalje identificirati s korištenom 
metodom - ali su i dalje informativne o prisutnosti šišmiša. Potvrđeno je prisustvo dvije vrste Rhinolophidae 
(Rhinolophus hipposideros, Rhinolophus ferrumequinum) i najmanje 11 vrsta iz porodice Vespertilionidae u 
okviru 15 različitih taksona (Myotis myotis, Myotis nattereri, Myotis mystacinus, Myotis alcathoe, Myotis sp., 
Pipistrellus pipistrellus, Pipistrellus pygmaeus, Pipistrellus nathusii, Pipistrellus kuhlii/nathusii, Hypsugo savii, 
Nyctalus noctula, Nyctalus noctula/lasiopterus, Vespertilio murinus, Vespertilio/Nyctalus/Eptesicus. 
Zabilježili smo i neke rijetko opažene vrste u BiH (Myotis alcathoe, Vespertilio murinus). 
 
Većina nalaza odnosi se na opažanja ultrazvučnim detektorima. Pet od devet pokušaja lova s mrežom 
bilo je uspješno u hvatanju najmanje jedne jedinke šišmiša (Tab. 2). Ovom metodom smo potvrdili prisustvo 
deset različitih vrsta (Tab. 2). Zabilježili smo skloništa samo jedne vrste, R. hipposideros, u nekoliko zgrada 
i pećina, od kojih su neke najvjerovatnije porodiljske kolonije. Vrstama najbogatije nalazište tokom našeg 
istraživanja zabilježeno je na rijeci Neretvi kod ušća Krupca, gdje je u mrežama uhvaćeno šest različitih vrsta 
šišmiša. 
 
Bogata zajednica šišmiša otkrivena u dolini rijeke Neretve i pored kratkog vremena istraživanja, dokazala 
je, da je ovo područje potencijalno vrlo bogato vrstama šišmiša i od velikog značaja za njihovo očuvanje. 
Sve vrste šišmiša su zaštićene različitim konvencijama. Prema Direktivi EU o staništima (OJ EC 1992), neke 
vrste šišmiša trebaju odrediti područja za zaštitu (Aneks II). U našem istraživanju zabilježili smo tri vrste 
šišmiša od najvećeg značaja za očuvanje u dolini rijeke Neretve: R. ferrumequinum, R. hipposideros i  
M. myotis. 
 
Planirana gradnja u dolini rijeke Neretve, uključujući niz hidroelektrana, njihovih odgovarajućih 
akumulacija, praćena sječom šuma, predstavlja prijetnju netaknuti prirodi područja. To će posljedično 
promijeniti staništa šišmiša, za koje se pokazao negativni utjecaj hidroenergetskih konstrukcija i sličnih 
promjena prirodnih staništa (Zortéa et al. 2021). S obzirom na bogatstvo vrsta, dolina gornje Neretve može 
se smatrati područjem od velikog značaja za očuvanje šišmiša. 
  
198 Maja ZAGMAJSTER et al.: Study of bats in the upper Neretva River valley ... / NSW 2022 – SCIENTIFIC PAPER 
NATURA SLOVENIAE 25(3): 181-200 
Acknowledgments 
 
 
We are grateful to the organising team of »Neretva Science Week 2022« for inviting us to the event and offering the 
opportunity to study bats. Stefan Andjus and Joshua Lim helped at two mist nettings, and Vera Knook collected the 
automatic detector from a site. We are grateful to Christian Dietz and Primož Presetnik for their opinion on V. murinus 
identification. The work was partly funded by Slovenian Research Agency through core funding P1-0184 and PhD grant to 
EP. EP was supported by the University Foundation of ing. Milan Lenarčič. 
 
 
 
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© 2023 Maja Zagmajster, Vojo Milanović, Behare Rexhepi, Ester Premate, Špela Borko and 
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