IntroductionTop

Studies about the biology and ecology of chiropterans have increased in recent decades. Among them, many have focused on the metazoan endoparasites and ectoparasites associated with bats and their high specificity with these hosts (Dick, 2007; Dick & Patterson, 2007; Walldorf & Mehlhorn, 2014; Tlapaya-Romero et al., 2019; Minaya et al., 2020).

Bats are hosts to a large number of endoparasites and ectoparasites. The most common ectoparasites that live on bats belong to the groups of mites, ticks and insects, especially bugs (Hemiptera), fleas (Siphonaptera) and bat flies (Diptera) (Frank et al., 2014). This last group of ectoparasites is the most specific and diverse on bats (Reeves & Lloyd, 2019).

Desmodus rotundus (Geoffroy, 1810) is distributed from northern Mexico throughout Central America and part of South America to southern Chile (Barquez et al., 2015). It is a hematophagous bat, known as a typical vampire, and feeds on the blood of birds and mammals (Quintana & Pacheco, 2007). Reports of ectoparasites on vampire bats have been studied extensively, and more than 33 species of ectoparasites including bat flies, mites and soft ticks have been recorded on this bat species (Frank et al., 2014). In Peru, ectoparasites of D. rotundus have been reported by Guerrero (1996a), Mendoza-Uribe & Chavez-Chorocco (2003), Autino et al. (2011) and Venzal et al. (2012).

The objective of this study is to report the ectoparasitic fauna of the vampire bat D. rotundus and additionally to compile a checklist that summarizes and includes all the records of bat ectoparasites in Peru.

Material and methodsTop

Individuals of D. rotundus were captured in an abandoned railway cave in San Bartolomé (11°54’11.5”S, 76°30’12.5” W; 1710 m.a.s.l.) in September 2016, near La Esperanza bridge, located 3 km east of the San Bartolomé district , Huarochirí Province, Lima, Peru. This town belongs to the Yungas natural region or Warm valley.

The ectoparasites were extracted with entomological forceps and brushes soaked in alcohol (Tlapaya-Romero et al., 2015). All arthropods were preserved in 70% ethanol. Chewing lice were rinsed in 20% KOH for 24 h, cleared in clove oil, and mounted in Canada balsam (Price et al., 2003). Diptera were placed in a solution of 70% ethanol, 5% glycerin, and 25% distilled water (Autino et al., 2011). Additionally, some tick specimens were analyzed with scanning electron microscopy (SEM).

For the identification of lice species, the taxonomic key of Price et al. (2003) was used. For Diptera, the keys of Wenzel (1976), Guerrero (1993, 1995a, 1996a), and the pictorial keys of Graciolli & Carvalho (2001) were used. Ticks were identified using the taxonomic keys of Barros-Battesti et al. (2013) with the support of the descriptions of Venzal et al. (2012). The values of prevalence (P%) and mean intensity (IM) were calculated according to Bush et al. (1997). The ectoparasitic specimens were deposited in the collection of Helminths Parasites and Related Invertebrates – HPIA – of the zoological collection of the Museum of Natural History of the Universidad Nacional Federico Villarreal – MUFV –, Lima, Peru. Accession numbers are listed in Table 1.

Table 1.—Ectoparasitic arthropods in 27 individuals of *Desmodus rotundus* from Lima, Peru, found in this study. Voucher numbers are also indicated (MUFV: Museum of Natural History of the Universidad Nacional Federico Villarreal).

Tabla 1.—Artrópodos ectoparásitos de 27 individuos de *Desmodus rotundus* de Lima, Perú, base de este estudio. Se indican también los códigos de los ejemplares (MUFV: Museum of Natural History of the Universidad Nacional Federico Villarreal).
Species	Stage	N	A	P%	MI	Voucher MUFV
DIPTERA
Strebla wiedemanni	Adult	6	7	22.22	1.17	HPIA: 183
Trichobius parasiticus	Adult	3	4	11.11	1.33	HPIA: 184
PHTHIRAPTERA
Campanulotes compar	Adult	2	4	7.41	2.00	HPIA: 185
Columbicola columbae	Adult	2	3	7.41	1.50	HPIA: 186
Colpocephalum sp.	Adult	1	1	3.70	1.00	HPIA: 187
IXODIDA
Ornithodoros peruvianus	nymph, Larvae	14	183	51.85	13.07	HPIA: 188

A= abundance, MI= mean intensity, N= number of infected host individuals, P= prevalence.

The checklist was compiled from data obtained from searches of the physical and virtual literature in the country’s university libraries, scientific collections, museums, and databases published until March 2020.

This review includes a list of ectoparasites, indicating their hosts, location, host specificity (HS), and the respective reference. Host Specificity was classified as Monoxenous (Mon), Stenoxenous (Est), Oligoxenous (Oli) and Polyxenous (Pol), according to Marshall (1981), and Gettinger (2018).

ResultsTop

ECTOPARASITES IN THE COMMON VAMPIRE BAT

Twenty-seven specimens of the common vampire D. rotundus were captured from the district of San Bartolomé, Huarochirí Province, Lima, Peru. Of this population, 19 individuals (70.4%) were parasitized by at least one ectoparasitic species. The taxa and ectoparasitic species registered on the D. rotundus sample are shown in Table 1.

The community of parasites on D. rotundus consisted of adult specimens of two species of bat flies in the family Streblidae, nymphs and larvae of a tick species belonging to the family Argasidae and three species of lice (considered accidental) of the families Menoponidae and Philopteridae (Figure 1). Among the collected species, Ornithodoros peruvianus Kohls, Clifford & Jones 1969 was the ectoparasite with the highest prevalence (P% = 51.85), Mean intensity (IM = 13.7) and abundance (s = 183).

Fig. 1.—Ectoparasites of the common vampire Desmodus rotundus from San Bartolomé district, Huarochirí Province, Lima, Peru. A. Colpocephalum sp. B. Campanulotes compar. C. Columbicola columbae. D. Trichobius parasiticus. E. Strebla wiedemanni. F. Ornithodoros peruvianus.

Fig. 1.—Ectoparásitos del vampiro común Desmodus rotundus del distrito de San Bartolomé, provincia de Huarochirí, Lima, Perú. A. Colpocephalum sp. B. Campanulotes compar. C. Columbicola columbae. D. Trichobius parasiticus. E. Strebla wiedemanni. F. Ornithodoros peruvianus.

CHECKLIST OF THE ECTOPARASITES OF BATS IN PERU

In our review, we found 104 species of ectoparasites in bats from Peru in two major groups: Hexapoda (Diptera, Hemiptera, Siphonaptera) and Arachnida (Mesostigmata and Ixodida). Of these 104 species, 99 were identified at the species level, and the rest at genus level only. The groups with the highest number of species were the families Streblidae (Diptera) (s = 59) and Spinturnicidae (Mesostigmata) (s = 20) (Table 2). Among the species with the higher number of hosts, Aspidoptera falcata Wenzel, 1976 and Megistopoda aranea (Coquillett, 1899) were recorded on eight host species each (Table 2).

Table 2.—Checklist of reported ectoparasitic arthropods in bats from Peru.

Table 2.—Lista de los registros de artrópodos ectoparásitos en murciélagos del Perú.
Ectoparasite	Host	Locality	HS	Reference
Class: Arachnida
Order: Mesostigmata
Family: Macronyssidae
Parichoronyssus alexanderfaini Morales-Malacara & Guerrero (2020)	Rhinophylla pumilio	Madre de Dios	Mon	Morales-Malacara & Guerrero (2020)
Parichoronyssus bakeri Morales-Malacara & Guerrero (2007)	Phyllostomus elongatus, P. hastatus	Cusco, Madre de Dios	Ste	Morales-Malacara & Guerrero (2007)
Parichoronyssus euthysternum Radovsky, 1967	Lophostoma silvicola, Sturnira erythromos, S. lilium, S. magna, S. oporaphilum	Cusco, Madre de Dios	Oli	Morales-Malacara & Guerrero (2020)
Parichoronyssus lopezi Morales-Malacara, 1996	Artibeus glaucus, Chiroderma salvini, Platyrrhinus brachycephalus	Cusco, Madre de Dios	Oli	Morales-Malacara & Guerrero (2020)
Parichoronyssus radovskyi Morales-Malacara, 1992	Lophostoma silvicola	Cusco	Mon	Morales-Malacara & Guerrero (2020)
Parichoronyssus sclerus Radovsky, 1966	Anoura geoffroyi	Cusco	Mon	Morales-Malacara & Guerrero (2020)
Family: Spelaeorhynchidae
Spelaeorhynchus praecursor Neumann, 1 902	Agoura cultrate, A. geoffroyi	Cusco, Madre de Dios	Ste	Gettinger et al. (2020)
Spelaeorhynchus soaresi Peracchi, 1990	Carollia brevicauda, C. perspicillata	Cusco, Madre de Dios	Ste	Gettinger et al. (2020)
Family: Spinturnicidae
Periglischrus acutisternus Machado-Allison, 1964	Phyllostomus elongatus, P. hastatus	Cusco, Madre de Dios	Ste	Gettinger (2018)
Periglischrus caligus Kolenati, 1857	Glossophaga soricina	lca	Mon	Mendoza-Uribe & Chavez-Chorocco (2003)
Periglischrus gameroi Machado-Allison & Antequera, 1971	Lonchorhina spp.	Cusco, Madre de Dios	Ste	Gettinger (2018)
Periglischrus grandisoma Herrin & Tipton, 1975	unknown	Cusco, Madre de Dios	--	Gettinger (2018)
Periglischrus herrerai Machado-Allison, 1965	Desmodus rotundus	Cusco, lca, Madre de Dios	Mon	Mendoza-Uribe & Chavez-Chorocco (2003); Gettinger (2018)
Periglischrus hopkinsi Machado-Allison, 1965	Lionycteris spurrelli	Cusco, Madre de Dios	Mon	Gettinger (2018)
Periglischrus iheringi Oudemans, 1902	Artibeus fraterculus	lca	Mon	Mendoza-Uribe & Chavez-Chorocco (2003)
Periglischrus micronycteridis Furman, 1966	Micronycteris megalotis, M. minuta	Cusco, Madre de Dios	Ste	Gettinger (2018)
Periglischus ojasti Machado-Allison, 1964	Sturnira erythromos, S. lilium, S. magna, S. oporaphilum, S. tildae	Cusco, Madre de Dios	Ste	Gettinger (2018)
Periglischrus paracutisternus Machado-Allison & Antequera, 1971	Trachops cirrhosus	Cusco, Madre de Dios	Mon	Gettinger (2018)
Periglischrus paravargasi Herrin & Tipton 1975	Anoura caudifer	Cusco, Madre de Dios	Mon	Gettinger (2018)
Periglischrus ramirezi Machado-Allison and Antequera, 1971	Rhinophylla pumilio	Cusco, Madre de Dios	Mon	Gettinger (2018)
Periglischrus tonatii Herrin & Tipton, 1975	Lophostoma carrikeri, L. silvicolum	Cusco, Madre de Dios	Ste	Gettinger (2018)
Periglischrus torrealbai Machado-Allison, 1965	Phyllostomus elongatus, P. hastatus	Cusco, Madre de Dios	Ste	Gettinger (2018)
Periglischrus vargasi Hoffmann, 1944	Anoura geoffroyi	Cusco, Madre de Dios	Mon	Gettinger (2018)
Order: Ixodida
Superfamilia: Ixodoidea
Family: Argasidae
Antricola sp.	Pteronotus parnellii rubiginosus	Huánuco	Mon	Need et al. (1991)
Ornithodoros hasei (Schulze, 1935)	Artibeus sp., Myotis albescens, Noctilio albiventris, Rhinophylla pumilio	Lima, Loreto	Pol	Need et al. (1991); Díaz et al. (2007)
Ornithodoros peruvianus Kohls, Clifford & Jones, 1969	Desmodus rotundus, Glossophaga sp., Molossus currentium	Lima, Loreto, Piura, Ucayali	Pol	Kohls et al. (1969); Present study
Ornithodoros sp.	unknown	Amazonas	Mon	Need et al. (1991)
Class: Hexapoda
Order: Siphonaptera
Family: Ischnopsyllidae
Myodopsylla wolffsohni wolffsohni (Rothschild, 1903)	Myotis albescens	Loreto	Mon	Autino et al. (2011)
Sternopsylla distincta (Rothschild, 1903)	Tadarida brasiliensis	Cusco	Mon	Autino et al. (1999)
Family: Pulicidae
Craneopsylla minerva wolffhuegeli (Rothschild, 1909)	Myotis nigricans**	unknown	Mon	Autino & Claps (2000)
Rhynchopsyllus pulex (Haller, 1880)	Molossus currentium, Histiotus sp.	Huánuco, unknown	Pol	Autino et al. (1999), Tipton & Machado-Allison (1972)
Order: Phthiraptera
Family: Menoponidae
Colpocephalum sp.	Desmodus rotundus**	Lima	Mon	Present study
Family: Philopteridae
Columbicola columbae (Linnaeus, 1758)	Desmodus rotundus**	Lima	Mon	Present study
Campanulotes compar (Burmeister, 1838)	Desmodus rotundus**	Lima	Mon	Present study
Order: Hemiptera
Family: Polyctenidae
Hesperoctenes angustatus Ferris & Usinger, 1939	Molossus molossus	Unknown, Piura	Mon	Autino et al. (1999); Bonifaz et al. (2020)
Hesperoctenes fumarius (Westwood, 1874)	Molossus rufus, Phyllostomus hastatus	Cusco	Pol	Maa (1961)
Order: Díptera
Family: Nycteribiidae
Basilia anceps Guimarães & D'Andretta, 1956	Myotis nigricans	Huánuco	Mon	Guimarães & D'Andretta (1956); Guimarães (1972)
Basilia constricta Guimarães & D'Andretta, 1956	Myotis nigricans, Uroderma bilobatum	Huánuco	Pol	Guimarães (1972)
Basilia dubia Guimarães & D'Andretta, 1956	Myotis albescens	Cusco	Mon	Guimarães & D'Andretta (1956); Guimarães (1972)
Basilia ferrisi Schuurmans-Stekhoven, Jr., 1931	unknown	Loreto, Ucayali	--	Guimarães & D'Andretta (1956); Guimarães (1972)
Basilia manu Guerrero, 1996	Myotis albescens, M. nigricans, M. riparius, M. simus	Madre de Dios	Ste	Guerrero (1996c)
Basilia mimoni Theodor & Peterson, 1964	Gardnerycteris crenulatum	Loreto	Mon	Theodor & Peterson (1964)
Basilia peruvia Guimarães & D'Andretta, 1956	Myotis nigricans, Tadarida brasiliensis	Cusco, Huánuco, Ucayali	Pol	Guimarães & D'Andretta (1956); Graciolli et al. (2007)
Basilia rugosa Schuurmans-Stekhoven, Jr., 1942	unknown	Ayacucho	--	Graciolli et al. (2007)
Hershkovitzia inaequalis Theodor, 1967	Thyroptera discifera	Loreto	Mon	Theodor (1967); Graciolli et al. (2007)
Family: Streblidae
Anastrebla modestini Wenzel, 1966	Anoura geoffroyi, A. peruana	Lima	Ste	Claps et al. (2005); Bonifaz et al. (2020)
Anatrichobius scorzai Wenzel, 1966	Myotis oxyotus	Cusco	Mon	Wenzel et al. (1966)
Aspidoptera falcata Wenzel, 1976	Carollia perspicillata, Desmodus rotundus, Rhinophylla pumilio, Sturnira lilium, S. magna, S. tildae, Sturnira sp. unknown	Cusco, Lima, Loreto, Madre de Dios, Pasco	Oli	Guerrero (1995b); Claps et al. (2005); Autino et al. (2011)
Aspidoptera phyllostomatis (Perty, 1833)	Artibeus fraterculus, A. jamaicensis, A. lituratus, A. obscurus, A. planirostris	Loreto, Madre de Dios, Pasco, Piura	Ste	Guerrero (1995b); Autino et al. (2011), Bonifaz et al. (2020)
Exastinion clovisi (Pessóa & Guimarães, 1936)	Anoura geoffroyi	Lima	Mon	Claps et al. (2005)
Mastoptera minuta (Costa-Lima, 1921)	Carollia brevicauda, Lophostoma silvicolum, Phyllostomus elongatus, P. hastatus	Loreto, Madre de Dios	Oli	Guerrero (1995b); Autino et al. (2011)
Megistopoda aranea (Coquillett, 1899)	Artibeus fraterculus, A. jamaicensis, A. lituratus, A. planirostris, Desmodus rotundus, Platyrrhinus brachycephalus, Uroderma bilobatum, unknown	Lambayeque, Lima, Loreto, Madre de Dios, Pasco, Piura	Oli	Wenzel (1970); Guerrero (1994b); Claps et al. (2005); Autino et al. (2011); Bonifaz et al. (2020)
Megistopoda proxima (Séguy, 1926)	Sturnira lilium, S. tildae, Sturnira sp.	Loreto, Madre de Dios, Pasco	Ste	Guerrero (1994b); Autino et al. (2011)
Metelasmus pseudopterus Coquillett, 1907	Artibeus fraterculus, A. jamaicensis, A. lituratus, A. planirostris, Artibeus sp., Dermanura anderseni	Loreto, Madre de Dios, Pasco, Piura.	Oli	Guerrero (1996a); Autino et al. (2011), Bonifaz et al. (2020)
Neotrichobius bisetosus Wenzel, 1976	Artibeus jamaicensis, A. obscurus	Madre de Dios	Ste	Guerrero (1994b)
Neotrichobius delicatus (Machado-Allison, 1966)	Dermanura anderseni, D. gnoma, Noctilio albiventris, Molossus rufus, Rhinophylla fischerae, R. pumilio	Loreto, Pasco	Pol	Guerrero (1994b); Autino et al. (2011)
Neotrichobius ectophyllae Wenzel, 1976	Mesophylla macconnelli	Loreto	Mon	Autino et al. (2011)
Noctiliostrebla aitkeni Wenzel, 1966	Noctilio leporinus	Loreto	Mon	Autino et al. (2011)
Noctiliostrebla dubia (Rudow, 1871)	Artibeus lituratus	Pasco	Mon	Guerrero (1995b)
Noctiliostrebla traubi Wenzel, 1966	Noctilio leporinus	Tumbes	Mon	Wenzel et al. (1966)
Paradyschiria fusca Speiser, 1900	Noctilio albiventris, Noctilio leporinus, unknown	Loreto, Pasco	Ste	Guerrero (1995b); Autino et al. (2011)
Paradyschiria lineata Kessel, 1925	Noctilio albiventris, Rhinophylla pumilio	Pasco	Pol	Guerrero (1995b)
Paradyschiria parvula Falcoz, 1931	Myotis simus, Noctilio albiventris, Noctilio sp.	Loreto, Madre de Dios	Pol	Guerrero (1995b), Autino et al. (2011)
Parastrebla handleyi Wenzel, 1966	Micronycteris megalotis	Madre de Dios	Mon	Guerrero (1994b)
Paratrichobius dunni (Curran, 1935)	Platyrrhinus brachycephalus, Uroderma bilobatum, Vampyriscus bidens	Loreto, Madre de Dios	Oli	Guerrero (1994b); Autino et al. (2011)
Paratrichobius longicrus (Miranda-Ribeiro, 1907)	Artibeus lituratus, Carollia perspicillata	Loreto, Madre de Dios	Olí	Guerrero (1994b); Autino et al. (2011)
Phalconomus sp.	Piatalina genovensium	Arequipa	Mon	Velazco et al. (2014)
Pseudostrebla ribeiroi Costa-Lima, 1921	Lophostoma silvicolum	Loreto, Madre de Dios	Mon	Guerrero (1994b); Autino et al. (2011)
Speiseria ambigua Kessel, 1925	Carollia brevicauda, C. perspicillata, Myotis sp., Piatalina genovensium, Trachops cirrhosus	Lima, Loreto, Madre de Dios, Pasco, Piura	Pol	Guerrero (1994b, 1996b); Claps et al. (2005); Velazco et al. (2014)
Speiseria magnioculus Wenzel, 1976	Diphylla ecaudata	Madre de Dios	Mon	Guerrero (1994b)
Speiseria peytonae Wenzel, 1976	Carollia brevicauda	Madre de Dios	Mon	Guerrero (1994b)
Strebla alvarezi Wenzel, 1966	Lonchophylla thomasi	Madre de Dios	Mon	Guerrero (1996a)
Strebla consocia Wenzel, 1966	Lophostoma brasiliense, L silvicolum, Phyllostomus elongatus, P. hastatus, Phyllostomus sp., Platyrrhinus brachycephalus	Loreto, Madre de Dios	Oli	Wenzel et al. (1966); Guerrero (1996a); Autino et al. (2011)
Strebla curvata Wenzel, 1976	Carollia brevicauda, Glossophaga commissarisi	Loreto, Madre de Dios	Oli	Guerrero (1996a); Autino et al. (2011)
Strebla diaemi Wenzel, 1966	Diaemus youngi	Loreto	Mon	Wenzel et al. (1966)
Strebla diphyllae Wenzel, 1966	Diphylla ecaudata	Madre de Dios	Mon	Guerrero (1996a)
Strebla galindoi Wenzel, 1966	Tonatia saurophila	Loreto	Mon	Autino et al. (2011)
Strebla guajiro (Garcia & Casal, 1965)	Carollia brevicauda, C. perspicillata, Carollia sp., Glossophaga soricina	Loreto, Pasco, unknown	Oli	Guerrero (1996a); Autino et al. (2011)
Strebla hertigi Wenzel, 1966	Carollia brevicauda, Phyllostomus discolor, P. elongatus	Loreto, Piura	Oli	Wenzel et al. (1966); Autino et al. (2011)
Strebla kohlsi Wenzel, 1966	Glossophaga soricina, Lophostoma silvicolum, Phylloderma stenops	Loreto, Madre de Dios	Oli	Guerrero (1996a); Autino et al. (2011)
Strebla kohlsi-Complex mirabilis	Phyllostomus hastatus, Trachops cirrhosus	Loreto	Oli	Autino et al. (2011)
Strebla mirabilis (Waterhouse, 1879)	Phyllostomus elongatus, P. hastatus, Phyllostomus sp., Trachops cirrhosus, unknown	Junin, Loreto, Madre de Dios	Oli	Wenzel et al. (1966); Guerrero (1996a)
Strebla machadoi Wenzel, 1966	Gardnerycteris crenulatum, Micronycteris minuta	Loreto, Madre de Dios	Oli	Guerrero (1996a); Autino et al. (2011)
Strebla obtusa Wenzel, 1976	Trinycteris nicefori	Loreto	Mon	Autino et al. (2011)
Strebla paramirabilis Wenzel, 1976	Lophostoma silvicolum	Loreto	Mon	Autino et al. (2011)
Strebla tonatie (Kessel, 1924)	Micronycteris megalotis, Lophostoma silvicolum	Cusco, Loreto	Oli	Wenzel et al. (1966); Autino et al. (2011)
Strebla wiedemanni Kolenati, 1856	Desmodus rotundus, Sturnira tildae	Apurímac, Cusco, Junín, Lima, Loreto, Piura, Puno, San Martín	Oli	Wenzel (1970); Guerrero (1996a); Claps et al. (2005); Autino et al. (2011); Present study
Trichobius caecus Edwards, 1918	Carollia perspicillata, Phyllostomus elongatus	Amazonas, Madre de Dios	Oli	Cáceres et al. (1992); Ibáñez & Jara (2008)
Trichobius costalimai Guimarães, 1938	Phyllostomus discolor	unknown	Mon	Wenzel et al. (1966)
Trichobius diaemi Wenzel, 1976	Diaemus youngi	Huánuco	Mon	Guerrero (1995a)
Trichobius diphyllae Wenzel, 1966	Diphylla ecaudata	Madre de Dios	Mon	Guerrero (1995a)
Trichobius dugesii Townsend, 1891	Glossophaga soricina	Junín, Madre de Dios	Mon	Wenzel et al. (1966); Guerrero (1995a)
Trichobius dugesioides Wenzel, 1966	Carollia perspicillata, Trachops cirrhosus	Huánuco, Madre de Dios	Oli	Guerrero (1995a); Guerrero (1996b)
Trichobius dybasi Wenzel, 1966	Lophostoma silvicolum	Piura	Mon	Wenzel (1970)
Trichobius furmani Wenzel, 1966	Carollia perspicillata, Desmodus rotundus, Glossophaga soricina, Lophostoma silvicolum, Molossus molossus, Phyllostomus hastatus	Junín, Huánuco, Puno	Pol	Wenzel et al. (1966); Guerrero (1995a)
Trichobius handleyi Wenzel, 1976	Micronycteris megalotis	Madre de Dios	Mon	Guerrero (1995a)
Trichobius joblingi Wenzel, 1966	Carollia brevicauda, C. castanea, C. perspicillata, Glossophaga soricina, Molossus molossus, Phyllostomus elongatus, Saccopteryx bilineata	Amazonas, Huánuco, Junín, Lima, Loreto, Madre de Dios, Pasco, Piura, Puno	Pol	Wenzel (1970), Cáceres et al. (1992); Guerrero (1995a); Ibáñez & Jara (2008), Bonifaz et al. (2020)
Trichobius lionycteridis Wenzel, 1966	Lionycteris spurrelli	Cusco	Mon	Wenzel et al. (1966)
Trichobius lonchophyllae Wenzel, 1966	Lonchophylla thomasi	Madre de Dios	Mon	Guerrero (1994a); Guerrero (1996b)
Trichobius longipes (Rudow, 1871)	Phyllostomus elongatus, P. hastatus	Amazonas, Madre de Dios, Pasco	Ste	Cáceres et al. (1992); Guerrero (1994a, 1996b); Ibáñez & Jara (2008)
Trichobius parasiticus Gervais, 1844	Carollia castanea, Desmodus rotundus, Rhinophylla pumilio, Sturnira lilium, S. tildae	Cusco, Huánuco, Huaraz, lca, Lima, Loreto, Madre de Dios, Pasco, Piura	Oli	Wenzel (1970); Elliot et al. (1985); Cáceres et al. (1992); Guerrero (1995a), Present study
Trichobius uniformis Curran, 1935	Glossophaga commissarisi, G. soricina	Madre de Dios, unknown	Ste	Wenzel et al. (1966); Guerrero (1996b)
Trichobius sp.	Platyrrhinus albericoi, P. infuscus, P. masu, P. nigellus	Cusco	Ste	Graciolli & Carvalho (2012)
Xenotrichobius sp.	Noctilio albiventris	Madre de Dios	Mon	Guerrero (1996b)

HS= Host Specificity: Mon= Monoxenous, St= Stenoxenous, OIi= Oligoxenous, Pol= Polyxenous

HS= Especificidad de hospedador: Mon= Monoxeno, St= Estenoxeno, OIi= Oligoxeno, Pol= Polixeno

** Accidental host

We found at least one record of some species of ectoparasite in seventy-five species of bats in Peru. These bat species are distributed in seven families, with Phyllostomidae being the one with the most reports of parasitized bats (s = 58). The bat species with the highest number of reported ectoparasitic species were Lophostoma silvicolum d’Orbigny, 1836, Phyllostomus elongatus (Geoffroy, 1810) and Phyllostomus hastatus (Pallas, 1767) with ten species each, followed by Carollia perspicillata (Linnaeus, 1758) and Lophostoma silvicolum d’Orbigny, 1836. Accidental ectoparasites were not considered in this count.

According to the level of specificity of the ectoparasites for their hosts, species with monoxenic associations (s = 47) were the most common, followed by oligoxenic ones (s = 21). Species of Diptera were observed to present the four types of association (Table 3).

Table 3.—Types of association according to host-ectoparasite specificity.

Tabla 3.—Tipos de asociación de acuerdo a su especificidad hospedador-ectoparásito.
Order Ectoparasites	Host specificity
Order Ectoparasites	Dip	Hem	Sip	Mes	Ixo	Total
Monoxenous	30	1	3	11	2	47
Stenoxenous	9	0	0	9	0	18
Oligoxenous	19	0	0	2	0	21
Polyxenous	8	1	1	0	2	12

Ixo=Ixodida, Dip= Diptera, Hem= Hemiptera, Mes=Mesostigmata, Sip= Siphonaptera.

In regards to the geographical distribution of ectoparasites and their hosts in Peru, 19 of the 24 departments in the Peruvian territory present at least some reports of an association of ectoparasites and bats. The departments of Madre de Dios and Loreto have the highest number of records, 40 and 37 species of ectoparasites, respectively, and 40 species of hosts in Loreto, followed by Madre de Dios with 30 species of hosts. Cajamarca, Huancavelica, La Libertad, Moquegua, and Tacna are the only five departments with no reports of ectoparasites on bats (Figure 2).

Fig. 2.—Distribution of reports of ectoparasites and their hosts in the regions of Peru.

Fig. 2.—Distribución de los registros de ectoparásitos y sus hospedadores en los departamentos del Perú.

DiscussionTop

ECTOPARASITES IN THE COMMON VAMPIRE BAT

It has been reported that the typical vampire bat D. rotundus hosts seven species of ectoparasites in Peru: Aspidoptera falcata (Claps et al., 2005), Megistopoda aranea (Claps et al., 2005), Ornithodorus peruvianus (Kohls et al., 1969), Periglischrus herrerai Machado-Allison, 1965 (Mendoza-Uribe & Chavez-Chorocco, 2003; Gettinger, 2018), Strebla wiedemanni Kolenati, 1856 (Wenzel, 1970; Claps et al., 2005; Autino et al., 2011), Trichobius furmani Wenzel, 1966 (Wenzel et al., 1966; Guerrero, 1995a), and Trichobius parasiticus Gervais, 1844 (Wenzel, 1970; Elliot et al., 1985; Guerrero, 1995a). Of these species, O. peruvianus, S. wiedemanni, and T. parasiticus were also found in our study.

The most studied families of ectoparasites on bats are the Streblidae and Nycteribiidae due to their specificity to their hosts (Aguiar & Antonini, 2016; Reeves & Lloyd, 2019). The family Streblidae has been reported typically associated with bats of the family Phyllostomidae (Aguiar & Antonini, 2016); S. wiedemanni and T. parasiticus have been reported in the present study with relatively low P% and MI (P% = 22.2, MI = 1.17 and P% = 11, MI = 1.33, respectively) on D. rotundus. Different studies have reported a broad range of values of these variables for these two ectoparasites on D. rotundus. Aguiar & Antonini (2011), in studies on the same host in Brazil, found intermediate values in S. wiedemanni (P% = 43.6, IM = 3.57), and in T. parasiticus (P% = 29.5, IM = 2.09); Rojas et al. (2008), in Costa Rica, observed low values in S. wiedemanni (P% = 11.9, IM = 2.12) and high values in T. parasiticus (P% = 91.4, IM = 5.65); González-Ávalos et al. (2014), in Mexico, found low values for S. wiedemanni (P% = 3.2) and high for T. parasiticus (P% = 94.3); Aguiar & Antonini (2016), in Brazil, found intermediate values for S. wiedemanni (P% = 44, IM = 3.6) and T. parasiticus (P% = 30, IM = 2.1); Guerrero (2019), in Venezuela, found intermediate values for S. wiedemanni (P% = 53.04, IM = 4.27) and low values for T. parasiticus (P% = 16.4, IM = 30.2). These differences may indicate that the two species present a fluctuating behavior, acting as the nucleus, secondary or satellite species of the parasitic community of D. rotundus, which may be influenced by different factors: environmental (specific to each ecosystem), or degree of disturbance (Lafferty & Kuris, 2005).

Early reports noted that one of the groups that are not usually found in bats is chewing lice (Phthiraptera). However, there are few reports of this significant association being considered incidental. In one of the first reports of mallophagans on bats, Gerberg & Goble (1941) noted two cases: a species of the genus Physconelloides Ewing, 1927, probably Physconelloides galapagensis (Kellogg & Kuwana, 1902), on Carollia perspicillata (reported as Hemiderma perspicilliatum) and another species, Geomydoecus (Geomydoecus) geomydis (Osborn, 1891), collected from Leptonycteris nivalis (Saussure, 1860) in Nuevo León, México. In this study, only presence was reported, but no plausible explanation for accidental parasitism was provided; still, Gerberg & Goble (1941) speculated that mallophagans can occur naturally in bats.

Another finding was by Cerny & Scholz (1983), who reported Brueelia straminea (Denny, 1842) (Philopteridae) in a single specimen of Pipistrellus kuhlii (Kuhl, 1817) in the town of Beyugdash, near Gobustan, Republic of Azerbaijan. Cerny & Scholz (1983) hypothesized that the transmission of B. straminea to P. kuhlii must have occurred in a place where the accidental host and the specific host, Dendrocopos major (Linnaeus, 1758), shared the same refuge for a particular time.

In our study, three species of lice, Columbicola columbae (Linnaeus, 1758), Campanulotes compar (Burmeister, 1838), and Colpocephalum sp. were collected from the coat of D. rotundus. These ectoparasite species are considered typical of columbiform birds, specifically the common pigeon Columba livia Gmelin, 1789 (González & Roldán, 2010; Castro et al., 2017; Abdullah et al., 2018). It is also known that these species often parasitize domestic birds such as the common chicken Gallus gallus domesticus (Linnaeus, 1758) (de Chirinos et al., 2001; Ferreira et al., 2013; Jassim & Hadi, 2019). Thus, we can consider two possibilities to explain the infestation of these lice in the vampire bat; the first is that D. rotundus has become parasitized when coming into contact with the common pigeon, perhaps when feeding on it; and the second possibility, when feeding on the domestic chicken. Of the two, we consider the latter most likely, based on reports of the selectivity and preference of D. rotundus for the blood of the domestic hen over other domestic and wild animals, probably because they are more predictable and easily available for D. rotundus (Bobrowiec et al., 2015).

Based on the above and the results presented in this study, we consider that lice parasitosis in bats is an accidental interaction, and that it is rare, with very few reports. We do not share the idea that it is a natural interaction, as implied by Gerberg & Goble (1941).

Ornithodoros peruvianus was described by Kohls et al. (1969) from three hosts, with D. rotundus as a typical host collected from the department of Lima. Venzal et al. (2012) studied the same ectoparasite on the same host raised in Chile and provided new morphological data and a tentative diagnosis of O. peruvianus; however, the material considered by the latter authors showed severe damage to the hypostome, an organ of taxonomic importance. Finally, Muñoz-Leal et al. (2020) contributed with a complete redescription of the larva, nymph, and adult stages of female and male O. peruvianus collected from the same host in northern Chile, which helped identify the specimens in this study.

CHECKLIST FOR ECTOPARASITES ON BATS IN PERU

In Peru, the diversity of mammals has been estimated at 559 species (Pacheco et al., 2018), of which 189 are species that belong to the order Chiroptera (Velazco, 2020). In this study, we present a summary of the reports of ectoparasites of bats in Peru, in which we found information for 75 species of bats, that is, there is no such information for 60.31% (s = 114) of bat species in Peru, which calls for further studies.

Previous checklists have summarized the fauna of endoparasites (Santos & Gibson, 2015; Minaya et al., 2020) and ectoparasites (Guerrero, 1997; Frank et al., 2014) on bats from South America including Peru. Frank et al. (2014) listed 38 species of ectoparasites, including 36 bat fly species (Streblidae 30, Nycteribiidae 6) and two Siphonaptera species. However, compared to our checklist, 59 species are registered in the Streblidae family (96.67% more) and four in Siphonaptera (100% more).

According to the specificity of the ectoparasites for their bat host, a greater variety of types of association was seen in the flies, presenting the four types of established associations. Even so, the monoxenous species were the most numerous. This high host specificity was also observed in other studies (Dick, 2007; Dick & Patterson, 2007). According to these authors, the behavior of the hosts, which live in large groups in close physical contact with other species of bats is not sufficient to increase the occurrence of oligoxenous or polyxenous species. The higher prevalence of monoxenous species could be explained by: (1) a mutual adaptation between host and its parasite that results in an immunological reaction that reduces the physical defense of the host (Dick & Patterson, 2007; Frank et al., 2014); and (2) the co-evolution of both groups playing a vital role in the specificity of the host, which is supported by the correlation between the phylogenies of the Streblidae and Nycteribiidae with their respective hosts (Patterson et al., 1998; Dick & Patterson, 2007). However, the latter is questioned by Graciolli & Carvalho (2012), who used phylogenic comparisons to show that some species, like Trichobius phyllostomae group (Streblidae), do not seem to have co-speciated with their bat hosts (Subfamily Stenodermatinae), with colonization of a new host providing a more likely explanation for this association.

The other group with higher specificity for their hosts is Acarina Mesostigmata, where only monoxenous, stenoxenous and oligoxenous species have been found. Within this group, the Spinturnicidae presented the highest number of species. According to Gettinger (2018), the family Spinturnicidae is one of the most specific in bats of the family Phyllostomidae compared to other host families, and polygenic species are not known.

Regarding the reported geographic distribution of ectoparasites and their hosts, some regions like Loreto and Madre de Dios have the highest number records because they have been comparatively more studied, probably because of the high diversity of bats that can be found in these regions, where the ecoregions of high-altitude rain forests (high jungle) and Amazonian tropical forest (low jungle) predominate. Most reports of bat ectoparasites in Peru result from the efforts of Autino et al. (2011) in Loreto, and Guerrero (1994b; 1995b) in Madre de Dios, which added significantly to this checklist.

It is worth mentioning that the species Artibeus jamaicensis Leach, 1821, Pteronotus parnellii rubiginosus Gray, 1843, and Molossus currentium Thomas, 1901 are not distributed in Peru according to Miller et al. (2016), Solari (2016) and Barquez & Diaz (2016), respectively. However, they have been considered in this checklist because they were cited in the works of Guerrero (1994b, 1995b, 1996a), Need et al. (1991), Kohls et al. (1969), and Autino et al. (1999). A review of the material deposited in the respective collections mentioned in these works is necessary to confirm species assignments.

AcknowledgmentsTop

The authors thank Gustavo Graciolli (Universidade Federal de Mato Grosso do Sul, Campo Grande, Brazil) for the kindness in answering our questions, for his comments, and for providing bibliographic material for the development of this work, and Omar Amin (Institute of Parasitic Diseases and Parasitology Center -Parasitology, Arizona, USA) and Brenton Ladd (Universidad Científica del Sur, Lima, Peru) for the suggestions and comments.

ReferencesTop


○	Abdullah, S.H., Mohammed, A.A. & Saeid, N.M., 2018. Study of ecto and haemo parasites in domestic pigeons (Columba livia domestica) in Sulaimani province, Kurdistan region/Iraq. Journal of Zankoy Sulaimani, 20(1): 37-44. https://doi.org/10.17656/jzs.10640
○	Aguiar, L.M.D.S. & Antonini, Y., 2011. Descriptive ecology of bat flies (Diptera: Hippoboscoidea) associated with vampire bats (Chiroptera: Phyllostomidae) in the cerrado of Central Brazil. Memórias do Instituto Oswaldo Cruz, 106(2): 170-176. https://doi.org/10.1590/S0074-02762011000200009
○	Aguiar, L.M.D.S. & Antonini, Y., 2016. Prevalence and intensity of Streblidae in bats from a Neotropical savanna region in Brazil. Folia Parasitologica, 63: 024 024. https://doi.org/10.14411/fp.2016.024
○	Autino, A.G. & Claps, G.L., 2000. Catalogue of the ectoparasitic insects of the bats of Argentina. Insecta Mundi, 14(4): 193-209.
○	Autino, A.G., Claps, G.L. & Barquez, R.M., 1999. Insectos ectoparásitos de murciélagos de las Yungas de la Argentina. Acta Zoológica Mexicana (nueva serie), 78: 119-169.
○	Autino, A.G., Claps, G.L., Barquez, R.M. & Díaz, M.M., 2011. Ectoparasitic insects (Diptera: Streblidae and Siphonaptera: Ischnopsyllidae) of bats from Iquitos and surrounding areas (Loreto, Peru). Memórias do Instituto Oswaldo Cruz, 106(8): 917-925. https://doi.org/10.1590/S0074-02762011000800004
○	Barquez, R. & Diaz, M., 2016. Molossus currentium. The IUCN Red List of Threatened Species 2016: e.T88087340A22107231. /https://doi.org/10.2305/IUCN.UK.2016-3.RLTS.T88087340A22107231.en
○	Barquez, R., Perez, S., Miller, B. & Diaz, M., 2015. Desmodus rotundus. The IUCN Red List of Threatened Species 2015: e.T6510A21979045. https://doi.org/10.2305/IUCN.UK.2015-4.RLTS.T6510A21979045.en
○	Barros-Battesti, D.M., Ramirez, D.G., Landulfo, G.A., Faccini, J. L.H., Dantas-Torres, F., Labruna, M.B., Venzal, J.L. & Onofrio, V.C., 2013. Immature argasid ticks: diagnosis and keys for Neotropical region. Revista Brasileira de Parasitologia Veterinária, 22(4): 443-456. https://doi.org/10.1590/S1984-29612013000400002
○	Bobrowiec, P.E.D., Lemes, M.R. & Gribel, R., 2015. Prey preference of the common vampire bat (Desmodus rotundus, Chiroptera) using molecular analysis. Journal of Mammalogy, 96(1): 54-63. https://doi.org/10.1093/jmammal/gyu002
○	Bonifaz, E., Mena, J.L. & Oporto, R., 2020. Moscas de murciélagos en algunas localidades de la costa peruana. Revista Peruana de Biología, 27(2): 241-250. https://doi.org/10.15381/rpb.v27i2.17881
○	Bush, A.O., Lafferty, K.D., Lotz, J.M. & Shostak, A.W., 1997. Parasitology meets ecology on its own terms: Margolis et al. revisited. The Journal of Parasitology, 83(4): 575-583. https://doi.org/10.2307/3284227
○	Cáceres, I., Elliot, A., Orozco, R. & Campomanes, L., 1992. Identificación de espécies de Trichobius (Gervais, 1984) (Diptera, Streblidae) em murciélagos de la Reserva de Tambopata, Madre de Dios, Peru. Revista Peruana de Medicina Tropical UNMSM, 6: 99-102.
○	Castro, J., Naupay, A., Orozco, K., Rodríguez, S., Díaz, Y., Navarro, J. & Purca, N., 2017. Ectoparásitos de Columba livia Linnaeus, 1758 (Aves: Columbiformes) del Distrito de Carmen de La Legua, Callao, Perú. The Biologist (Lima), 15(2): 425-435.
○	Cerny, V. & Scholz, T., 1983. Finding of Mallophaga on Pipistrellus kuhli (Chiroptera). Folia Parasitologica, 30(3): 268 268.
○	Claps, G.L., Autino, A.G. & Barquez, R.M., 2005. Streblidae de murciélagos de Lima: dos citas nuevas para Perú. Revista de la Sociedad Entomológica Argentina, 64(1-2): 95-98.
○	de Chirinos, N.I.M., Chirinos, Á.R., Hinestroza, Y., Inicarte, M.F., Manco, M. & Meléndez, A., 2001. Prevalencia de ectoparásitos en gallinas de corral (Gallus-gallus domesticus) del municipio San Francisco, Estado Zulia, Venezuela. Revista Científica de la Facultad de Ciencias Veterinarias, 11(4): 348-355.
○	Díaz, M.M., Nava, S., Venzal, J.M., Sánchez, N. & Guglielmone, A.A., 2007. Tick collections from the Peruvian Amazon, with new host records for species of Ixodes Latreille, 1795 (Acari: Ixodidae) and Ornithodoros Koch, 1844 (Acari: Argasidae). Systematic and Applied Acarology, 12(2): 127-133. https://doi.org/10.11158/saa.12.2.6
○	Dick, C.W., 2007. High host specificity of obligate ectoparasites. Ecological Entomology, 32 (5): 446–450. https://doi.org/10.1111/j.1365-2311.2007.00836.x
○	Dick, C.W. & Patterson, B.D., 2007. Against all odds: explaining high host specificity in dispersal-prone parasites. International Journal for Parasitology, 37(8-9): 871-876. https://doi.org/10.1016/j.ijpara.2007.02.004
○	Elliot, A., Cáceres, I. & Macedo, S., 1985. Ectoparásitos de Desmodus rotundus I. Trichobius parasiticus, Gervais 1844 (Diptera, Streblidae). Boletín del Instituto de Medicina Tropical “Daniel Carrión”, (UNMSM)., 4: 18-21.
○	Ferreira, C.G.T., Mafra, C., Bezerra, A.C.D.S., de Carvalho, O.V., Júnior, A.S. & de Almeida, M.R., 2013. Campanulotes compar (Burmeister, 1838) (Phthiraptera: Ischnocera) in chickens (Gallus gallus domesticus) from Rio Grande do Norte State, Brazil. The reemergence of an ectoparasite?. Veterinary Parasitology, 195(1-2): 203-204. https://doi.org/10.1016/j.vetpar.2012.12.047
○	Frank, R., Münster, J., Schulze, A., Liston, A. & Klimpel, S., 2014. Macroparasites of Microchiroptera: bat ectoparasites of Central and South America. In: Klimpel, S. & Mehlhorn, H. (eds.), Bats (Chiroptera) as vectors of diseases and parasites. Springer-Verlag. Berlin & Heidelberg: 87–130.
○	Gerberg, E.J. & Goble, F.C., 1941. Two unusual records of Mallophaga from bats. Journal of Mammalogy, 22(4): 454-454. https://doi.org/10.1093/jmammal/22.4.454-a
○	Gettinger, D., 2018. Checklist of bloodfeeding mites (Acari: Spinturnicidae) from the wings of Bats (Mammalia: Chiroptera) in the Manú Biosphere Reserve, Peru. MANTER: Journal of Parasite Biodiversity, (Occasional Papers)., 10: 1-9. https://doi.org/10.13014/K2DJ5CVZ
○	Gettinger, D., Epperson, M., Hermasillo, C. & Gardner, S., 2020. The Enigmatic Spelaeorhynchidae Oudemans, 1902 (Acari: Mesostigmata) blood-feeding ectoparasites infesting Neotropical bats, with catalog and notes on a collection from the Manú Biosphere Reserve in Peru. MANTER: Journal of Parasite Biodiversity, (Occasional Papers)., 12: 1-6. https://doi.org/10.32873/unl.dc.manter12
○	González, E. & Roldán, J., 2010. Mallophagos en aves y mamíferos domésticos circunstancialmente presentes en el Mercado Mayorista de Trujillo y zonas aledañas. Revista Peruana de Parasitología, 18: 21-22.
○	González-Ávalos, C., Macías-Díaz, D.A., Macip-Castañeda, A., Mendez-Andrade, A., Ortiz-García, K.P. & Rivera-García, K.D., 2014. Ectoparásitos (Diptera: Streblidae) asociados a Desmodus rotundus en el sur del Estado de Puebla, México. Entomología Mexicana, 1: 973–977.
○	Graciolli, G., Autino, A.G. & Claps, G.L., 2007. Catalogue of American Nycteribiidae (Diptera, Hippoboscoidea). Revista Brasileira de Entomologia, 51(2): 142-159. https://doi.org/10.1590/S0085-56262007000200004
○	Graciolli, G. & Carvalho, C.J.D., 2001. Moscas ectoparasitas (Diptera, Hippoboscoidea) de morcegos (Mammalia, Chiroptera) do estado do Paraná. II. Streblidae: chave pictórica para gêneros e espécies. Revista Brasileira de Zoologia, 18(3): 907-960. https://doi.org/10.1590/S0101-81752001000300026
○	Graciolli, G. & Carvalho, C.J., 2012. Do fly parasites of bats and their hosts coevolve? Speciation in Trichobius phyllostomae group (Diptera, Streblidae) and their hosts (Chiroptera, Phyllostomidae) suggests that they do not. Revista Brasileira de Entomologia, 56 (4): 436-450. https://doi.org/10.1590/S0085-56262012000400007
○	Guerrero, R., 1993. Catálogo de los Streblidae (Diptera: Pupipara) parásitos de murciélagos (Mammalia: Chiroptera) del Nuevo Mundo. I. Clave para los géneros y Nycterophiliinae. Acta Biológica Venezuelica, 14(4): 61–75.
○	Guerrero, R., 1994 a. Catálogo de los Streblidae (Diptera: Pupipara) parásitos de murciélagos (Mammalia: Chiroptera) del Nuevo Mundo. II. Los grupos: pallidus, caecus, major, uniformis y longipes del género Trichobius Gervais, 1844. Acta Biológica Venezuelica, 15(1): 1 –18.
○	Guerrero, R., 1994 b. Catálogo de los Streblidae (Diptera: Pupipara) parásitos de murciélagos (Mammalia: Chiroptera) del Nuevo Mundo. IV. Trichobiinae con alas desarroladas. Boletín de Entomología Venezolana, 9(2): 161 –192.
○	Guerrero, R., 1995 a. Catálogo de los Streblidae (Diptera: Pupipara) parásitos de murciélagos (Mammalia: Chiroptera) del Nuevo Mundo. III. Los grupos: dugesii, dunni y phyllostomae del género Trichobius Gervais, 1844. Acta Biológica Venezuelica, 15(3-4): 1 –27.
○	Guerrero, R., 1995 b. Catálogo de los Streblidae (Diptera: Pupipara) parásitos de murciélagos (Mammalia: Chiroptera) del Nuevo Mundo. V. Trichobiinae con alas reducidas o ausentes y misceláneos. Boletín de Entomología Venezolana, 10(2): 135–160.
○	Guerrero, R., 1996 a. Catálogo de los Streblidae (Diptera: Pupipara) parásitos de murciélagos (Mammalia: Chiroptera) del Nuevo Mundo. VI. Streblinae. Acta Biológica Venezuelica, 16(2): 1–25.
○	Guerrero, R., 1996 b. Streblidae (Diptera: Pupipara) parásitos de los murciélagos de Pakitza, Parque Nacional Manu (Perú). In: Wilson, D.E. & Sandoval, A. (eds.), Manu: The biodiversity of southeastern Peru. Smithsonian Institution. Washington, DC: 627-641.
○	Guerrero, R., 1996 c. The Basilia juquiensis species-group (Diptera: Nycteribiidae) with description of a new species from Pakitza, Perú. In: Wilson, D.E. & Sandoval, A. (eds.), Manu: The biodiversity of southeastern Peru. Smithsonian Institution. Washington, DC: 665-674.
○	Guerrero, R., 1997. Catálogo de los Streblidae (Diptera: Pupipara) parásitos de murciélagos (Mammalia: Chiroptera) del Nuevo Mundo. VII. Lista de especies, hospedadores y países. Acta Biológica Venezuelica, 17(1): 9 –24.
○	Guerrero, R., 2019. Streblidae (Diptera: Pupipara) de Venezuela. Sistemática, Ecología y Evolución. Editorial Académica Española. Wroclaw. 590 pp.
○	Guimarães, L.R., 1972. Venezuelan nycteribiid batflies (Diptera: Nycteribiidae). Brigham Young University Science Bulletin, (Biological Series)., 17(1): 1-10.
○	Guimarães, L.R. & D’Andretta, M.A.V., 1956. Sinopse dos Nycteribiidae (Diptera) do Novo Mundo. Arquivos de Zoologia do Estado de São Paulo, 10: 1-184.
○	Ibáñez, N.H. & Jara, C.A., 2008. Registro de Trichobius (Diptera: Streblidae) en murciélagos de Amazonas, Perú. Rebiol, 28(2): 87-90.
○	Jassim, S.Y. & Hadi, A.M., 2019. Isolation and identification of lice on some species of Columbidae family. Advances in Animal and Veterinary Sciences, 7(9): 806-809. https://doi.org/10.17582/journal.aavs/2019/7.9.806.809
○	Kohls, G.M., Clifford, C.M. & Jones, E.K., 1969. The systematics of the subfamily Ornithodorinae (Acarina: Argasidae). IV. Eight new species of Ornithodoros from the Western Hemisphere. Annals of the Entomological Society of America, 62 (5): 1035 –1043. https://doi.org/10.1093/aesa/62.5.1035
○	Lafferty, K.D. & Kuris, A.M., 2005. Parasitism and environmental disturbances. In: Thomas, R.F., Thomas, F., Renaud, R.F. & Guegan, J.F. (Eds.). Parasitism and ecosystems. Oxford University Press. Oxford: 113-123.
○	Maa, T.C., 1961. Records and descriptions of Polyctenidae (Hemiptera). Pacific Insects, 3(1): 1-10.
○	Marshall, A.G., 1981. The sex ratio in ectoparasitic insects. Ecological Entomology, 6(2): 155-174. https://doi.org/10.1111/j.1365-2311.1981.tb00602.x
○	Mendoza-Uribe, L. & Chavez-Chorocco, J., 2003. New records of mites (Gamasida: Spinturnicidae) on Peruvian bats (Quiroptera: Phyllostomidae). Revista de Biología Tropical, 51(1): 276-277.
○	Miller, B., Reid, F., Arroyo-Cabrales, J., Cuarón, A.D. & de Grammont, P.C., 2016. Artibeus jamaicensis. The IUCN Red List of Threatened Species 2016: e.T88109731A21995883. https://doi.org/10.2305/IUCN.UK.2016-3.RLTS.T88109731A21995883.en
○	Minaya, D.A., Saez, F.G., Chero, J.C., Cruces, C.C. & Iannacone, J., 2020. Helminth parasites of bats (Chiroptera, Phyllostomidae) in the Department of Junin, Peru and Check list of records made in Peru. Revista del Museo Argentino de Ciencias Naturales, 20(1): 57-73.
○	Morales-Malacara, J.B. & Guerrero, R., 2007. A new species of Parichoronyssus (Acari: Dermanyssoidea: Macronyssidae) from bats of the genus Phyllostomus (Chiroptera: Phyllostomidae) in Peru and Venezuela, with keys to the species of Parichoronyssus. Journal of medical Entomology, 44(1): 8-13. https://doi.org/10.1093/jmedent/41.5.08
○	Morales-Malacara, J. B. & Guerrero, R., 2020. Two new species and new records of mites of the Genus Parichoronyssus (Acari: Macronyssidae) From South American bats (Chiroptera), with a key to the known species of the genus. Journal of Medical Entomology, 57(2): 404-417. https://doi.org/10.1093/jme/tjz196
○	Muñoz-Leal, S., González-Acuña, D., Labruna, M.B. & Venzal, J.M., 2020. Redescription of the larva, and description of the nymphal and adult stages of Ornithodoros peruvianus Kohls, Clifford & Jones, 1969 (Acari: Argasidae). Systematic Parasitology, 97: 201-215. https://doi.org/10.1007/s11230-020-09908-6
○	Need, J.T., Dale, W.E., Keirans, J.E. & Dasch, G.A., 1991. Annotated list of ticks (Acari: Ixodidae, Argasidae) reported in Peru: distribution, hosts, and bibliography. Journal of Medical Entomology, 28 (5): 590-597. https://doi.org/10.1093/jmedent/28.5.590
○	Pacheco, V., Inche, B. & Wust, W., 2018. Mamíferos del Perú. Grupo La República Publicaciones S.A., Lima. 120 p.
○	Patterson, B.D., Ballard, J.W. Wenzel, R.L., 1998. Distributional evidence for cospeciation between neotropical bats and their bat fly ectoparasites. Studies on Neotropical Fauna and Environment, 33 (2):76 –84. https://doi.org/10.1076/snfe.33.2.76.2152
○	Price, R.D., Hellenthal, R.A., Palma, R.L., Johnson, K.P. & Clayton, D.H. (Eds.), 2003. The chewing lice: world checklist and biological overview. Illinois Natural History Survey, Illinois.
○	Quintana, H. & Pacheco, V., 2007. Identificación y distribución de los murciélagos vampiros del Perú. Revista Peruana de Medicina Experimental y Salud Publica, 24(1): 81-88.
○	Reeves, W.K. & Lloyd, J.E., 2019. Louse Flies, Keds, and Bat Flies (Hippoboscoidea). In: Mullen, G.R. & Durden, L.A. (eds.) Medical and Veterinary Entomology. Mullen. Academic Press. New York: 421-438.
○	Rojas, A., Jiménez, A., Vargas, M., Zumbado, M. & Herrero, M.V., 2008. Ectoparasites of the common vampire bat (Desmodus rotundus) in Costa Rica: parasitism rates and biogeographic trends. Mastozoología Neotropical, 15(2): 181-187.
○	Santos, C.P. & Gibson, D.I., 2015. Checklist of the helminth parasites of South American bats. Zootaxa, 3937(3): 471-499. https://doi.org/10.11646/zootaxa.3937.3.3
○	Solari, S., 2016. Pteronotus parnellii. The IUCN Red List of Threatened Species 2016: e.T88017638A22077695. https://doi.org/10.2305/IUCN.UK.2016-3.RLTS.T88017638A22077695.en
○	Theodor, O., 1967. An Illustrated catalogue of the Rothschild collection of Nycteribiidae (Diptera) in the British Museum (Natural History). Publication, 655. Trustees of the British Museum (Natural History). London. 506 pp.
○	Theodor, O., & Peterson, B.V., 1964. On some new species of Nycteribiidae (Diptera: Pupipara). The Great Basin Naturalist, 24: 107-115. https://doi.org/10.5962/bhl.part.22782
○	Tlapaya-Romero, L., Horváth, A., Gallina-Tessaro, S., Naranjo, E.J. & Gómez, B., 2015. Prevalencia y abundancia de moscas parásitas asociadas a una comunidad de murciélagos cavernícolas en La Trinitaria, Chiapas, México. Revista Mexicana de Biodiversidad, 86(2): 377-385. https://doi.org/10.1016/j.rmb.2015.04.012
○	Tlapaya-Romero, L., Ibáñez-Bernal, S. & Santos-Moreno, A., 2019. New records of bat flies (Diptera: Streblidae) in Oaxaca, Mexico. Revista Mexicana de Biodiversidad, 90: e902894. https://doi.org/10.22201/ib.20078706e.2019.90.2894
○	Tipton, V.J. & Machado-Allison, C.E., 1972. Fleas of Venezuela. Brigham Young University Science Bulletin, (Biological Series)., 17: 1-122.
○	Velazco, P.M., 2020. Murciélagos del Perú. Available at: http://www.paulvelazco.com/murcielagos_peru.html [accessed: 04/04/2020].
○	Velazco, P.M., Autino, A.G. & Claps, G.L., 2014. New record of the ectoparasite insect Speiseria ambigua Kessel, 1925 (Diptera: Streblidae) of Platalina genovensium Thomas, 1928 (Chiroptera: Phyllostomidae) from Peru. Check List, 10(6): 1525-1527. https://doi.org/10.15560/10.6.1525
○	Venzal, J., González-Acuña, D., Mangold, A. & Guglielmone, A., 2012. Ornithodoros peruvianus Kohls, Clifford & Jones (Ixodoidea: Argasidae) in Chile: a tentative diagnosis. Neotropical Entomology, 41 (1): 74-78. https://doi.org/10.1007/s13744-011-0011-y
○	Walldorf, V. & Mehlhorn, H., 2014. Bats: A glimpse on their astonishing morphology and lifestyle. In: Klimpel, S. & Mehlhorn, H. (eds.). Bats (Chiroptera) as vectors of diseases and parasites. Springer-Verlag. Berlin & Heidelberg: 7-24.
○	Wenzel, R.L., 1970. Family Streblidae. In: Papavero, N. (ed.). A catalogue of the Diptera of the Americas south of the United States, Museu de Zoologia. Universidade de São Paulo, São Paulo: 1-25.
○	Wenzel, R.L., 1976. The streblid batflies of Venezuela (Diptera: Streblidae). Brigham Young University Science Bulletin, (Biological Series)., 20: 1-177.
○	Wenzel, R.L., Tipton, V.J. & Kiewlicz, A., 1966. The Streblid batflies of Panamá (Diptera Calypterae: Streblidae). In: Wenzel, R.L. & Tipton, V.J. (Eds.). Ectoparasites of Panama. Publications of Field Museum of Natural History Chicago: 405-675.

ECTOPARASITIC FAUNA ON THE COMMON VAMPIRE BAT DESMODUS ROTUNDUS (GEOFFROY, 1810) (CHIROPTERA: PHYLLOSTOMIDAE) FROM HUAROCHIRI, LIMA, AND A CHECKLIST OF ECTOPARASITES IN BATS OF PERU