BIOTROPIA Vol. 30 No. 3, 2023: 365 - 373 DOI: 10.11598/btb.2023.30.3.1963 365 DIVERSITY OF ECTOPARASITES ON BATS IN DRAMAGA, BOGOR, INDONESIA VIRGILIUS MARTIN KELAKE KEDANG1, YANRI RIZKY NATANAEL SIMANGUNSONG1, SUSI SOVIANA2, UPIK KESUMAWATI HADI2 AND SUPRIYONO2* 1School of Veterinary Medicine and Biomedical Sciences, IPB University, Bogor 16680, Indonesia 2Division of Parasitology and Medical Entomology, School of Veterinary Medicine and Biomedical Sciences, IPB University, Bogor 16680, Indonesia Received 26 March 2023 / Revised 1 July 2023 / Accepted 2 July 2023 ABSTRACT Ectoparasites infestation is one of the major health problems affecting animals, including bats, which are known as reservoir hosts for various pathogens. Several reports have shown that a comprehensive understanding of ectoparasites on these animals is crucial from a public health perspective. Therefore, this study aims to identify the diversity of ectoparasites on bats in Dramaga, Bogor, Indonesia. The samples were captured at night with a mist net and then identified using the species identification key. Ectoparasites were collected from the body of the samples and identified using a microscope. A total of 56 bats from 9 species, namely Cynopterus brachyotis, Cynopterus sphinx, Cynopterus titthaecheilus, Macroglossus sobrinus, Rousettus leschenaultii, Myotis muricola, Nycteris javanica, Pipistrellus javanicus, and Scotophilus kuhlii were obtained in this study. The results of ectoparasites identification showed the presence of Basilia spp., Eucampsipoda sp., Leptocyclopodia ferrarii (Nycteribiidae), Raymondia sp. (Streblidae), Meristaspis spp., Spinturnix spp. (Spinturnicidae), and ticks (Ixodidae). The total prevalence of infested bats was 51.7%, with females tending to have a higher intensity compared to males. Bats species with the highest prevalence of infestation were Rousettus leschenaultii and Myotis muricola. Meanwhile, this study found no ectoparasites on Macroglossus sobrinus and Scotophilus kuhlii. Keywords: bats, bats flies, Bogor, ectoparasite, Indonesia INTRODUCTION Bats are known to play an essential role in maintaining the ecological balance of ecosystems, serving as seed transmitters, pollinators, and natural insect control (Suyanto 2001). Despite their significant ecological functions, these animals also serve as the natural reservoir of various pathogens. Previous studies showed that bats had unique immune systems, thereby enabling them to carry various zoonotic viruses per host species compared to rodents (Irving et al. 2021; Luis et al. 2013). Several emerging infectious diseases have also been linked to these animals bats as a source of zoonotic pathogens, including Ebola virus, SARS coronavirus, Nipah virus, and Hendra virus (Brook & Dobson 2015; Calisher et al. 2006). Furthermore, studies conducted in Indonesia showed their association with potential zoonotic diseases (Diptyanusa et al. 2021; Tsang et al. 2021). This indicates that the study of bats and their potential to transmit diseases to humans or other animals demands thorough investigation. One of the health issues affecting bats is ectoparasites infestation, which warrants further investigation to comprehend the ecology and health of these creatures. The commonly associated ectoparasites belong to the class Insecta (bats flies and bats flea) and Arachnida (ticks and mites). The species within the class Insecta are known to belong to the family Nycteribiidae, Streblidae, Hippoboscidae, and Ischnopsyllidae. Meanwhile, those from class Arachnida are often members of the family Argasidae, Ixodidae, and Spinturnicidae (Fajri et al. 2018; Azhar et al. 2015). Previous studies showed that these ectoparasites in bats are vectors of pathogens, such as Bartonella spp., Rickettsia spp., and certain viruses with the ability *Corresponding author, email: supriyono84@apps.ipb.ac.id BIOTROPIA Vol. 30 No. 3, 2023 366 to cause diseases in humans or animals (Szentiványi et al. 2019; Reeves et al. 2016). Several studies have been carried out on bat ectoparasites in several Southeast Asia countries, including Malaysia (Azhar et al. 2015), Singapore (Lim et al. 2020), and the Philippines (Alvarez et al. 2015). Based on previous findings, studies related to these ectoparasites in Indonesia remains significantly limited, despite the presence of over 230 distinct bat species in this region (Maryanto et al. 2020). Existing reports in Indonesia primarily focused on recording ectoparasite infestation in different locations and species, especially megabats (Nangoy et al. 2021; Sauqi et al. 2021; Fajri et al. 2018). Based on these findings, there is a need to conduct further studies to understand the characteristics of ectoparasites, including their specific host and potential role in transmitting diseases. Dramaga is a subdistrict located in Bogor, Indonesia, which has been extensively explored due to bats species diversity in Dramaga. Several studies have also been carried out in and around the IPB Dramaga Campus area and its surrounding villages. According to Mustari et al. (2014) and Mustari (2020), at least 10 species of bats were found in IPB Dramaga Campus. Another report showed the presence of 11 species in villages around the area (Sumirto 2013). Due to the role of these animals in maintaining ecosystem balance, there is a need to carry out further investigations on their ecology and health problems, such as ectoparasites infestation. Therefore, this study aims to identify the diversity of ectoparasites on bats in Dramaga, Bogor, Indonesia. MATERIALS AND METHODS Study Sites and Sampling Techniques Bats used in this study were obtained from the seven locations in Dramaga, Bogor, Indonesia from November 2021 to February 2022. The habitats selected as sampling locations included arboretums, agricultural land areas, and rural areas. Furthermore, reconnaissance surveys were conducted in the area before the study began to identify the location where bats were likely to congregate. The survey targeted the regions where bats were expected to fly, such as fruiting trees or congregation sites of insects, to determine the appropriate sampling locations. The procedures for sample capturing in this study referred to the Guidelines for Bats Reservoir Data Collection published by the National Institute of Health Research and Development, Ministry of Health, Indonesia. The samples were captured at night using mist-net techniques. The mist nets were installed in determined locations and placed approximately 5 meters above the ground. The installation was carried out for four hours from 18.00 to 22.00 and monitored every hour. Bats were collected as detected trapped in the mist net, placed into a cotton bag, and transported to a nearby station for further identification (Balitbangkes 2015). Bats Identification and Ectoparasites Collection Captured bats were identified based on morphological features and morphometries using the species identification key from the Field Guide of Bats in Indonesia (Suyanto 2001). The identified samples were then observed to determine the presence of ectoparasites infestation. The collection procedure for ectoparasites was carried out using small tweezers. Subsequently, the parasites obtained were placed in a labeled vial containing 70% alcohol (Balitbangkes 2015), and bats were released back to the initial location. Ectoparasites Identification The initial grouping of ectoparasites was carried out based on their morphological characteristics. Bats flies were distinguished by their large size resembling a fly with or without wings, while mites and ticks were distinguished by their relatively small size compared to bat flies. Subsequently, the collected ectoparasites were identified using a microscope. Bats flies were identified using the publications of Maa (1971, 1968, 1962) and Theodor (1967, 1959), while mites were identified using the publication of Baker & Delfinado (1964) and Delfinado & Baker (1963). Ticks ectoparasites were identified using the publication of Hoogstraal (1955). Data Analysis Data on captured bats were analyzed descriptively by making a table that included the species, the number of captured bats, the number Diversity of Ectoparasite on Bats in Dramaga, Bogor, Indonesia – Kedang et al. 367 of bats infested with ectoparasites, the prevalence of ectoparasites infestation, and the intensity of ectoparasites. Data on the collected ectoparasites were analyzed descriptively by making a table containing the types of ectoparasites, hosts, sex, and the number of collected samples per host. RESULTS AND DISCUSSION Bats Diversity A total of 56 bats were obtained in this study, belonging to the suborders Megachiroptera and Microchiroptera. The samples represented nine species, namely Cynopterus brachyotis, Cynopterus sphinx, Cynopterus titthaecheilus, Macroglossus sobrinus, Rousettus leschenaultii, Myotis muricola, Nycteris javanica, Pipistrellus javanicus, and Scotophilus kuhlii. The results showed that the most captured species was Cynopterus brachyotis, with a total of 20 samples (35.7%). The least caught species was Scotophilus kuhlii, with only one sample (1.8%), as shown in Table 1. All bats species found in this study had a conservation status of least concern, except Nycteris javanica, which was considered vulnerable. Nycteris javanica was an endemic bats and was only found in Java and Timor (Waldien & Wiantoro 2021). This study found a total of 29 samples that were infested with ectoparasites, accounting for 51.7% of the total population. The species with the highest prevalence (80%) of infestation were Rousettus leschenaultia and Myotis muricola. Meanwhile, there was no infestation on Macroglossus sobrinus and Scotophilus kuhlii, as shown in Table 1. The prevalence and intensity of ectoparasites in each bat sex are presented in Table 2. The results showed that the number and intensity of ectoparasites varied widely among the samples’ sex and species. Female Rousettus lechenaultii species had the highest intensity of infestation. Female bats tended to have a higher intensity of ectoparasites compared to males, but their population was smaller. Table 1 Bats species and prevalence of ectoparasites infestation in bats Bats Species Number of Captured Bats Number of Infested Bats Prevalence (%) Megachiroptera (Megabat) Cynopterus brachyotis 20 11 55 Cynopterus sphinx 5 2 40 Cynopterus titthaecheilus 11 6 54.5 Macroglossus sobrinus 4 0 0 Rousettus leschenaultii 5 4 80 Subtotal 45 23 51.1 Microchiroptera (Microbat) Myotis muricola 5 4 80 Nycteris javanica 2 1 50 Pipistrellus javanicus 3 1 33.3 Scotophilus kuhlii 1 0 0 Subtotal 11 6 54.5 Total 56 29 51.7 Table 2. Intensity of ectoparasites based on bats species and sex Bats Species Infested Bats/Captured Bats Total Ectoparasites Intensity of Ectoparasites Male Female Male Female Male Female Cynopterus brachyotis 10/19 1/1 20 2 2.0 2.0 Cynopterus sphinx 1/3 1/2 3 1 3.0 1.0 Cynopterus titthaecheilus 4/6 2/5 5 2 1.25 1.0 Rousettus leschenaultii 2/3 2/2 8 24 4.0 12.0 Myotis muricola 4/5 0/0 27 0 6.75 0 Nycteris javanica 1/2 0/0 1 0 1.0 0 Pipistrellus javanicus 1/3 0/0 2 0 2.0 0 Macroglossus sobrinus 0/4 0/0 0 0 0 0 Scotophilus kuhlii 0/1 0/0 0 0 0 0 Total 23/46 6/10 66 29 2.87 4.83 BIOTROPIA Vol. 30 No. 3, 2023 368 Figure 1 Bats species captured in Dramaga, Bogor: (A) Cynopterus titthaecheilus with ectoparasites (white circle), (B) Macroglossus sobrinus, (C) Rousettus leschenaultii, (D) Nycteris javanica This was the first study to record the diversity of ectoparasites on bats in Dramaga, Bogor. The results could be used to monitor the status of bats species in Dramaga, Bogor, in line with conservation efforts. This study also presented the first record of ectoparasites infestation on vulnerable endemic bats species, namely Nycteris javanica. The species was endemic to Java and Timor Islands and not commonly found in nature due to its declining population and conservation status being vulnerable (Waldien & Wiantoro 2021). Due to the absence of ectoparasites on some species, further studies must be carried out to determine the possibility of infestation on bats. This study found no infestation in Macroglossus sobrinus and Scotophilus kuhlii, possibly due to the lack of representative samples. Therefore, future reports must explore ectoparasites that could affect these species. Further studies were also needed due to the inability of this current study to cover all bat species reported by Mustari (2020), such as Kerivoula hardwickii, Rhinolophus affinis, and Hipposideros diadema. Ectoparasites Diversity Ectoparasites found in this study were from Insecta and Acarina classes, as shown in Table 3. A total of 95 samples were collected representing three groups, namely bats flies (family Nycteribiidae and Streblidae), mites (family Spinturnicidae), and ticks (family Ixodidae). Nycteribiidae bats flies were found on five microbats and megabats, namely Myotis muricola, Cynopterus brachyotis, Cynopterus sphinx, Cynopterus titthaecheilus, and Rousettus leschenaultii. Streblidae species was only discovered on Nycteris javanica. Mites were discovered on Rousettus leschenaultii, Myotis muricola, and Pipistrellus javanicus, while ticks were only found on Myotis. Furthermore, Rousettus leschenaultia had the highest ectoparasites infestation compared to others in Dramaga. The differences in the number of infestation were due to the variations in number of captured individual bats in each species. Diversity of Ectoparasite on Bats in Dramaga, Bogor, Indonesia – Kedang et al. 369 Table 3 Collected ectoparasites found on bats in Dramaga, Bogor No Ectoparasites Species Number of Ectoparasites Total C.b C.s C.t M.sb R.l M.m N.j P.j S.k Bats flies 1 Basilia spp. 0 0 0 0 0 10 0 0 0 10 2 Eucampsipoda sp. 0 0 0 0 21 0 0 0 0 21 3 Leptocyclopodia ferrarii 22 4 7 0 0 0 0 0 0 33 4 Raymondia sp. 0 0 0 0 0 0 1 0 0 1 Mites 1 Meristaspis spp. 0 0 0 0 11 0 0 0 0 11 2 Spinturnix spp. 0 0 0 0 0 16 0 2 0 18 Ticks 1 Ixodidae 0 0 0 0 0 1 0 0 0 1 Total 22 4 7 0 32 27 1 2 0 95 Notes: C.b: Cynopterus brachyotis, C.s: Cynopterus sphinx, C.t: Cynopterus titthaecheilus, M.sb: Macroglossus sobrinus, R.l: Rousettus leschenaultii, M.m: Myotis muricola, N.j: Nycteris javanica, P.j: Pipistrellus javanicus, S.k: Scotophilus kuhlii. Ectoparasites from the family Nycteribiidae were the most collected ectoparasites in this study. Furthermore, Nycteribiid bats flies were ectoparasites commonly found in megabats from the genus Cynopterus and Rousettus (Nangoy et al. 2021). Species from the genus Cynopterus were reported to be the primary hosts of Leptocyclopodia flies (Maa 1975). Several studies collected Leptocyclopodia ferrarii from Cynopterus brachyotis in Indonesia (Nangoy et al. 2021; Sauqi et al. 2021). Eucampsipoda bats flies were common on megabats, and they had infested Eonycteris spelaea and Rousettus leschenaultii in Southeast Asia countries (Lim et al. 2020; Fajri et al. 2018; Azhar et al. 2015). Basilia ectoparasites were commonly obtained in megabats and microbats, and Poerwanto et al. (2020) reported their presence on Miniopterus schreibersii in Yogyakarta, Indonesia. Ectoparasites from the genus Raymondia included Streblidae bats flies found in microbats (Azhar et al. 2015; Maa 1962). These species could be distinguished from Nycteribiidae samples by the presence of the wings (Azhar et al. 2015). This study only found Raymondia sp. on Nycteris javanica. This result was the first record of ectoparasites infestation by the genus Raymondia on Nycteris javanica. Figure 2 Ectoparasites collected from bats in Dramaga, Bogor: (A) Basilia sp. collected from Myotis muricola, (B) Eucampsipoda sp. collected from Rousettus leschenaultii, (C) Leptocyclopodia ferrarii collected from Cynopterus spp., (D) Raymondia sp. collected from Nycteris javanica, (E) Mite Meristaspis sp. ♀ and (F) Meristaspis sp. ♂ collected from Rousettus leschenaultii, (G) Spinturnix sp. collected from Myotis muricola and Pipistrellus javanicus, and (H) Ixodid tick (Ixodidae) collected from Myotis muricola BIOTROPIA Vol. 30 No. 3, 2023 370 Mites and ticks were found in the genus Rousettus, Myotis, and Pipistrellus. Mite ectoparasites infesting bats in Indonesia were from the Spinturnicidae family, while the ticks were from the Ixodidae and Argasidae families (Fajri et al. 2018). Several studies reported ticks and mites ectoparasites infestation on bats in Indonesia (Poerwanto et al. 2020; Fajri et al. 2018). The results of Meristaspis sp. and Spinturnix sp. mites from the family Spinturnicidae and ticks larvae from the family Ixodidae were consistent with previous studies on arachnid infestation on bats. Bat-Ectoparasites Interaction Ectoparasites in bats spent almost their entire life cycle on the host through blood consumption (Hiller et al. 2019; Bordes et al. 2008). The intensity and prevalence of ectoparasites were affected by several factors, including host- specificity, habitat, nesting, gender, diet, and social behavior (Nangoy et al. 2021; Hiller et al. 2019; Ramanantsalama et al. 2018). Therefore, their distribution and abundance were associated with the distribution and abundance of the host (Putra, 2014; Ter Hofstede & Fenton, 2005). The colony size of each bat species could also influence social behavior, leading to differences in the prevalence of ectoparasites infestation. Species that shared the same roosting site in larger groups increased the possibility of ectoparasites transmission between the bats (Putra 2014). Bats in genera Cynopterus, Rousettus, and Pipistrellus were generally known to roost from medium to large colonies (Garg et al. 2015; Kumar et al. 2015; Gay et al. 2014). Different roosting behavior was found in Macroglossus species, which tended to roost alone or in small colonies occupying different sites (Putra 2014; Gould 1978). The absence of ectoparasites infestation on Macroglossus bats in this study was allegedly related to the behavior of the species. Several reports showed that ectoparasites infestation in bats had an association with sex. In this study, females tended to have a higher intensity of ectoparasites compared to males, but they had few infested individuals due to the small population. The results were consistent with Nangoy et al. (2021) on Pteropodid bats in Sulawesi, where females had a higher intensity due to their high susceptibility caused by several factors. Fluctuations in the reproductive cycle, such as pregnancy and lactation, could suppress immunity and increase susceptibility to parasites. During this period, the female species tended to spend more time in roosting sites, increasing contact with ectoparasites or the other infested bats (Tai et al. 2022; Nangoy et al. 2021; Webber et al. 2015). A previous study reported that extended stay in roosting sites also increased ectoparasite exposure (Lim et al. 2020). Compared to the results of this study, Lim et al. (2020) found that male Cynopterus brachyotis and Eonycteris spealea had a higher level of infestation. This was because they spent more time in the roost site due to defense. Grooming activities commonly carried out to expel parasites also affected infiltration among sexes since males spent more time in grooming activities related to ectoparasites consumption compared to females (Ramanantsalama et al. 2018). Meanwhile, Godinho et al. (2013) found no association between grooming activity and the number of parasites. Due to the variation in literature, further studies must be carried out to investigate ectoparasites infestation in relation to sex. Ectoparasites in bats generally had certain specific hosts, and could only be found in some species. Bats flies were obligate and specialized parasitic organisms found on the fur and wing membranes. Furthermore, this study found a genus-specific host pattern in ectoparasites. Leptocyclopodia ferarri bats flies obtained had specific hosts from the Cynopterus spp. bats. These findings were consistent with several studies, where species from the genus Cynopterus were primary hosts for Leptocyclopodia spp. (Lim et al. 2020; Azhar et al. 2015). Based on the results, Meristaspis mites tended to infest megabats, while Spinturnix mites infested microbats. The results were in line with previous studies, which recorded the infestation of Meristaspis spp. mites on megabats from the genus Macroglossus and Rousettus (Fajri & Armiani 2021; Fajri et al. 2018). Meanwhile, the study by Zania et al. (2022) reported the Spinturnix mite infestation on Rousettus bats in Banyuwangi. This variation in findings suggested that the mites did not have a genus-specific pattern. Diversity of Ectoparasite on Bats in Dramaga, Bogor, Indonesia – Kedang et al. 371 The presence of ectoparasites in bats contributed to the spread of pathogens between individual members in the colony. Some of the parasites had various hosts and could be vectors for certain pathogens. Several studies found Bartonella spp. bacteria in Leptocyclopodia bats flies collected from the megabats in Malaysia and the Philippines (Low et al. 2022; Morse et al. 2012). Bartonella spp. was a bacteria that caused bartonellosis and was considered zoonosis (Chomel & Kasten 2010). Bacterium Bartonella spp. and Rickettsia spp. were also found in mites from the genus Spinturnix collected from Myotis myotis bats in Poland (Szubert-Kruszyńska et al. 2019). Feng et al. (2017) found the Khaeng Koi virus in Eucampsipoda sundaica from Rousettus leschenaultia bats in China. The results of bats ectoparasites in Dramaga proved that ectoparasites infestation was a major health problem in bats. The ability of ectoparasites to act as vectors of various pathogens demands further studies on their distribution in each bats species, as well as their public health importance. CONCLUSION In conclusion, ectoparasites were one of the causes of health-related problems in bats with health importance due to their ability to act as vectors of various pathogens. Ectoparasites affecting bats in Dramaga were from the family Nycteribiidae, Streblidae, Spinturnicidae, and Ixodidae. The results showed that the total prevalence of infested samples in the study location was 51.7%. Furthermore, the female samples tended to have a higher intensity of ectoparasites compared to males. The species with the highest prevalence of infestation were Rousettus leschenaultii and Myotis muricola. This study found no ectoparasites infestation on Macroglossus sobrinus and Scotophilus kuhlii. Based on these findings, further studies were needed as some species of bats required large sample sizes. REFERENCES Azhar I, Khan FAA, Ismail N, Abdullah MT. 2015. Checklist of bat flies (Diptera: Nycteribiidae and Streblidae) and their associated bat hosts in Malaysia. Check List 11(5): 1777. Alvarez JD, Lit Jr IL, Alviola PA. 2015. Bat flies (Diptera: Nycteribiidae) from Mount Makiling, Luzon Island: New host and distribution records, with a checklist of species found in the Philippines. Check List 11(1): 1509. Baker EW, Delfinado MD. 1964. Spinturnicidae of South East Asia and The Pacific Region. Pacific Insects 6(4): 571-91. Balitbangkes. 2015. Pedoman Pengumpulan Data Reservoir (Kelelawar) di Lapangan. [Guidelines for Reservoar (Bat) Data Collection in the Field]. Jakarta (ID): Badan Penelitian dan Pengembangan Kesehatan. Bordes F, Morand S, Ricardo G. 2008. Bat fly species richness in Neotropical bats: correlations with host ecology and host brain. Oecologia 158(1): 109-16. Brook CE, Dobson AP. 2015. Bats as ‘special’ reservoirs for emerging zoonotic pathogens. Trends in Microbiology 23(3): 172-80. Calisher HC, Childs JE, Field HE, Holmes KV, Schountz T. 2006. Bats: Important Reservoir Hosts of Emerging Viruses. Clinical Microbiology Reviews 19(3): 531-45. Chomel BB, Kasten RW. 2010. Bartonellosis, an increasingly recognized zoonosis. Journal of Applied Microbiology 109(3): 743-50. Delfinado MD, Baker EW. 1963. Mites of The Family Spinturnicidae From The Philippines (Acarina). Pacific Insects 5(4): 905-20. Diptyanusa A, Herini ES, Indarjulianto S, Satoto TBT. 2021. The detection of Japanese encephalitis virus in Megachiropteran bats in West Kalimantan, Indonesia: A potential enzootic transmission pattern in the absence of pig holdings. International Journal for Parasitology: Parasites and Wildlife 14: 280-86. Fajri SR, Armiani S. 2021. A Prevalence, Intensity, and Associated of Ectoparasitic Fauna among Cave- Dwelling Bats from Lombok Island West Nusa Tenggara. Prisma Sains: Jurnal Pengkajian Ilmu dan Pembelajaran Matematika dan IPA IKIP Mataram 9(1): 141-51. Fajri SR, Primawati SN, Hadi I, Tresnani G. 2018. Ectoparasites diversity of bats species collected from Southern Lombok, West Nusa Tenggara, Indonesia. Journal of Biological Series 1(2): 31-41. Feng Y, Li Y, Fu S, Li X, Song J, Zhang H, Yang W, Zhang Y, Pan H, Liang G. 2017. Isolation of Kaeng Khoi virus (KKV) from Eucampsipoda sundaica bat flies in China. Virus Research 238: 94-100. Garg KM, Chattopadhyay B, Doss DPS, Kumar AKV, Kandula S, Ramakrishnan U. 2015. Males and Females Gain Differentially from Sociality in a Promiscuous Fruit Bat Cynopterus sphinx. PLoS ONE 10(3): e0122180. BIOTROPIA Vol. 30 No. 3, 2023 372 Gay N, Olival KJ, Bumrungsri S, Siriaroonrat B, Bourgarel M, Morand S. 2014. Parasite and viral species richness of Southeast Asian bats: Fragmentation of area distribution matters. International Journal for Parasitology: Parasites and Wildlife. 3: 161-70. Godinho LN, Cripps JK, Coulson G, Lumsden LF. 2013. The effect of ectoparasites on the grooming behaviour of Gould's wattled bat (Chalinolobus gouldii): An experimental study. Acta Chiropterologica. 15(2): 463-72. Gould E. 1978. Foraging Behavior of Malaysian Nectar- Feeding Bats. Biotropica 10(3): 184-93. Hiller T, Brändel SD, Honner B, Page RA, Tschapka M. 2019. Parasitization of bats by bat flies (Streblidae) in fragmented habitats. Biotropica 52(3): 488-501. Hoogstraal H. 1955. Bat Ticks of The Genus Argas (Ixodoidea, Argasidae) I The Subgenus Chiropterargas. Fieldiana: Zoology 37: 579-600. Irving AT, Ahn M, Goh G, Anderson DE, Wang LF. 2021. Lessons from the host defences of bats, a unique viral reservoir. Nature 589: 363-70. Kumar M, Priya YS, Mathur V, Elangovan V. 2015. Roost Selection and Roosting Ecology of Fulvous Fruitbat, Rousettus leschenaulti (Pteropodidae). International Journal of Advanced Biological Research 5(1): 62-68. Lim ZX, Hitch AT, Lee BPYH, Low DHW, Neves ES, Borthwick SA, Smith GJD, Mendenhall IH. 2020. Ecology of bat flies in Singapore: A study on the diversity, infestation bias and host specificity (Diptera: Nycteribiidae). International Journal of Parasitology: Parasites and Wildlife 12: 29-33. Low VL, Tan TK, Tohiran KA, Lim YAL, AbuBakar S, Nasir DM. 2022. A novel clade of bat-associated Bartonella detected in the bat fly Leptocyclopodia ferrari (Diptera: Nycteribiidae) parasitizing Cynopterus brachyotis (Chiroptera: Pteropodidae). Veterinary Microbiology 264: 109284. Luis AD, Hayman DTS, O'Shea TJ, Cryan PM, Gilbert AT, Pulliam JRC, Mills JN, Timonin ME, Willis CKR, Cunningham AA, Fooks AR, Rupprecht CE, Wood JLN, Webb CT. 2013. A comparison of bats and rodents as reservoirs of zoonotic viruses: are bats special?. Proceedings of the Royal Society B: Biological Sciences 280: 20122753. Maa TC. 1962. Records and Descriptions of Nycteribiidae and Streblidae (Diptera). Pacific Insects 4 (2): 417-436. Maa TC. 1968. Additions to The Cyclopodiinae Part I (Diptera: Nycteribiidae). Pacific Insects 10 (1): 1-23. Maa TC. 1971. Review of The Streblidae (Diptera) Parasitic on Megachiropteran Bats. Pacific Insects Monograph 28: 213-243. Maa TC. 1975. On new Diptera Pupipara from The Oriental Region. Pacific Insect 16(4): 465-486. Maryanto I, Maharadatunkamsi, Achmadi AS, Wiantoro S, Sulistyadi E, Yoneda M, Suyanto A, Sugardjito J. 2020. Checklist of the mammals of Indonesia. Jakarta (ID): Research Center for Biology Indonesian Institute of Sciences (LIPI). Morse SF, Olival KJ, Kosoy M, Billeter S, Patterson BD, Dick CW, Dittmar K. 2012. Global distribution and genetic diversity of Bartonella in bat flies (Hippoboscoidea, Streblidae, Nycteribiidae). Infection, Genetics and Evolution 12: 1717-23. Mustari AH, Zulkarnain I, Rinaldi D. 2014. Keanakaragaman Jenis dan Penyebaran Mamalia di Kampus IPB Dramaga Bogor. [Species Diversity and Distribution of Mammals in IPB Dramaga Campus, Bogor]. Media Konservasi 19(2): 117-25. Mustari AH. 2020. Biodiversitas di Kampus IPB University: Mammalia, Burung, Amfibi, Reptil, Kupu-Kupu, dan Tumbuhan. [Biodiversity in IPB University Campus: Mammals, Birds. Amphibians, Reptiles, Butterflies, and Plants]. Bogor (ID): IPB Press. Nangoy M, Ransaleleh T, Lengkong H, Koneri R, Latinne A, Kyes RC. 2021. Diversity of fruit bats (Pteropodidae) and their ectoparasites in Batuputih Nature Tourism Park, Sulawesi, Indonesia. Biodiversitas 22(6): 3075-82. Putra MIH. 2014. Hubungan Inang-Ektoparasit pada Kelelawar Pemakan Buah di Kampus Universitas Indonesia Depok. [Host-Ectoparasite Relationship on Fruit Bats in Universitas Indonesia, Depok]. [Undergraduate Thesis]. Retrieved from Universitas Indonesia Repository. Poerwanto SH, Ridhwan LR, Giyantolin G, Ginawati D, Paramitha DPR. 2020. Keanekaragaman Ektoparasit pada Kelelawar Subordo Microchiroptera di Goa Jepang Bukit Plawangan, Sleman, Yogyakarta. [Ectoparasites Diversity of Microchiroptera Bats Suborder in Jepang Cave, Plawangan Hill, Sleman, Yogyakarta]. Jurnal Veteriner 21(4): 629-36. Ramanantsalama RV, Andrianarimisa A, Raselimanana AP, Goodman SM. 2018. Rates of hematophagous ectoparasite consumption during grooming by an endemic Madagascar fruit bat. Parasites Vectors 11: 330. Reeves WK, Beck J, Orlova MV, Daly JL, Pippin K, Revan F, Loftis AD. 2016. Ecology of bats, their ectoparasites, and associated pathogens on Saint Kitts Island. J Med Entomol 53(5): 1218-25. Sauqi MS, Restiadi TI, Koesdarto S, Hastutiek P, Setiawan B, Wijaya A. 2021. Identification of ectoparasites and endoparasites on fruit bats (Cynopterus brachyotis) in Ketapang Timur Village, Ketapang Sub-District, Sampang District. Journal of Parasite Science 5(2): 35-40. Sumirto A. 2013. Keanekaragaman Jenis Kelelawar di Desa Cikarawang Kecamatan Dramaga Kabupaten Diversity of Ectoparasite on Bats in Dramaga, Bogor, Indonesia – Kedang et al. 373 Bogor Provinsi Jawa Barat. [Bats Diversity in Cikarawang Village Dramaga District Bogor Regency West Java Province]. [Undergraduate Thesis]. Retrieved from IPB University Repository. Suyanto A. 2001. Panduan Lapangan Kelelawar di Indonesia. [Field Guide of Bats in Indonesia]. Bogor (ID): Puslitbang Biologi LIPI. Szentiványi T, Christe P, Glaizot O. 2019. Bat Flies and Their Microparasites: Current Knowledge and Distribution. Frontiers in Veterinary Science 6: 115. Szubert-Kruszyńska A, Stańczak J, Cieniuch S, Podsiadły E, Postawa T, Michalik J. 2019. Bartonella and Rickettsia infections in haematophagous Spinturnix myoti mites (Acari: Mesostigmata) and their bat host, Myotis myotis (Yangochiroptera: Vespertilionidae), from Poland. Microbial Ecology 77(3): 759-768. Tai YL, Lee YF, Kuo YM, Kuo YJ. 2022. Effects of host state and body condition on parasite infestation of bent-wing bats. Frontiers in Zoology 19(1): 1-13. Ter Hofstede HM, Fenton MB. 2005. Relationship between roost preferences, ectoparasite density, and grooming behaviour of neotropical bats. Journal of Zoology. 266: 333-340. Theodor O. 1959. A Revision of The Genus Cyclopodia (Nycteribiidae, Diptera). Parasitology 49(1-2): 242-308. Theodor O. 1967. An Illustrated Catalogue of The Rothschild Collection of Nycteribiidae (Diptera) in The British Museum (Natural History). London (UK): British Museum (Natural History). Tsang SM, Low DHW, Wiantoro S, Smith I, Jayakumar J, Simmons NB, Vijaykrishna D, Lohman DJ, Mendenhall IH. 2021. Detection of Tioman Virus in Pteropus vampyrus Near Flores, Indonesia. Viruses 13: 563. Waldien DL, Wiantoro S. 2021. Nycteris javanica. The IUCN Red List of Threatened Species 2021: e.T14932A22013241. Webber QMR, McGuire LP, Steven B. 2015. Host behaviour, age and sex correlate with ectoparasite prevalence and intensity in a colonial mammal, the little brown bat. Behaviour 152(1): 83-105. Zania FO, Geraldine AP, Putri CK, Edila R, Yudhana A. 2022. Laporan Kasus Infestasi Spinturnix spp. pada Kelelawar Pemakan Buah (Rousettus spp.) di Banyuwangi. [Case report of Spinturnix spp. infestation on Fruit Bats (Rousettus spp.) in Banyuwangi]. Media Kedokteran Hewan 33(3): 233-43.