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Lesser horseshoe bat

The lesser horseshoe bat (Rhinolophus hipposideros) is a small insectivorous in the family Rhinolophidae, distinguished by its complex, horseshoe-shaped nose leaf that aids in echolocation for navigating and hunting in the dark. Adults typically measure 35-45 mm in head-body length, with a of 35-43 mm, a of 200-255 mm, and a weight of 5-12 g, featuring greyish-brown fur on the back and paler underparts. Juveniles are greyer and reach adult size within a few weeks, while the hangs with its wings wrapped around its body at rest, a posture unique among bats. This bat inhabits a range spanning , , and southwestern Asia, with fragmented populations in the UK limited to , southwest , and western , where it favors temperate climates and diverse landscapes including woodlands, grasslands, and river valleys. It roosts in humid, sheltered sites such as attics, cellars, barns, caves, and mines; summer maternity colonies of 10-200 females form in warm buildings from May to July, while occurs solitarily or in small groups from October to April in cooler, stable underground sites at around 8-11°C. occurs low to the ground (rarely above 5 m) within 2-5 km of roosts, though individuals may travel up to 10 km in search of prey-rich areas near or dense . The diet consists primarily of small soft-bodied , including midges, moths, , lacewings, craneflies, and occasionally spiders or , captured using constant-frequency echolocation calls at 105-111 kHz to detect prey amid foliage. Bats often glean from surfaces or use temporary perches to consume larger items, emerging 20-30 minutes after sunset and remaining active throughout the night during the breeding season. Females give birth to a single pup in June or July after a of 45-60 days, with young able to fly within three weeks and becoming independent by two months; occurs in autumn, with sperm stored until . The species is non-migratory but may disperse up to 150 km between summer and winter roosts, exhibiting social behaviors like clustering in maternity groups for warmth. Globally classified as Least Concern by the IUCN but with a decreasing population trend due to habitat fragmentation, agricultural intensification, roost disturbance, and climate impacts, the lesser horseshoe bat is protected under the EU Habitats Directive and national laws in Europe, where targeted conservation has stabilized or increased some populations. In the UK and Ireland, it is a priority species, with efforts focusing on preserving traditional farm buildings and underground sites to support its recovery.

Taxonomy

Classification

The lesser horseshoe bat, Rhinolophus hipposideros, occupies the following position in the taxonomic hierarchy: kingdom Animalia; phylum Chordata; class Mammalia; order Chiroptera; family Rhinolophidae; genus Rhinolophus; species R. hipposideros (André, 1797). This classification places it among the Old World horseshoe bats, distinguished by their specialized nasal structures adapted for echolocation. The genus name Rhinolophus originates from the rhinos () and lophos (crest), reflecting the elaborate, crest-like noseleaf that covers the snout and aids in sound emission. The specific epithet hipposideros derives from Greek hippos (horse) and sideros (iron), evoking the shape of an iron horseshoe, which describes the distinctive U-shaped form of the anterior noseleaf. The was first validly described as Noctilio hipposideros by Édouard in 1797, later reassigned to Rhinolophus. Although traditionally attributed to Bechstein (1800) as Vespertilio hipposideros, a 2022 nomenclatural confirms André (1797) as the authority, resolving earlier ambiguities in dating and attribution. Within the Rhinolophidae, R. hipposideros is the recognized member of the hipposideros species group, defined by shared morphological traits such as the structure of the noseleaf and basioccipital bone. Recent taxonomic studies have scrutinized this group for potential cryptic diversity. The lesser horseshoe bat, Rhinolophus hipposideros, is currently recognized as comprising two subspecies. The nominate subspecies, R. h. hipposideros, occurs in the Maghreb region of North Africa and western Europe west of the Dnieper River, Bosporus, and Strait of Karpathos, with a diploid chromosome number of 2n = 54–56. The other subspecies, R. h. minimus, is distributed from eastern Europe (including Crimea and the Caucasus) through the Middle East to northeastern Africa (from Sudan to Djibouti), characterized by a diploid number of 2n = 58. These subspecies are distinguished primarily by genetic differences, with morphological and acoustic traits showing less clear separation. In a 2023 taxonomic revision, populations previously considered part of R. hipposideros in the were elevated to a distinct species, Rhinolophus midas, based on comprehensive analyses. This new species is restricted to areas around the and . The separation is supported by genetic evidence from (mtDNA) and nuclear DNA (nuDNA), showing significant divergence; morphological differences, including a smaller , distinct structure, and variations in dental traits; and acoustic distinctions in echolocation calls (echotypes). No have yet been defined for R. midas. The taxonomic status of easternmost populations of the R. hipposideros complex, such as those in West Turkestan, , and , remains unresolved and awaits further investigation with expanded sampling. Phylogenetically, R. hipposideros (sensu stricto) belongs to the monotypic hipposideros species group within the genus Rhinolophus, positioned as sister to other major clades in the Afro-Palaearctic lineage of the family Rhinolophidae.

Description

Physical characteristics

The lesser horseshoe bat (Rhinolophus hipposideros) is a small with a head-body length of 35–45 mm, a length of 37–44 mm, a of 192–254 mm, and a weight of 5–9 g. These measurements position it as the smallest species in , with a compact build suited to its agile flight. Its external appearance features soft, fluffy fur that covers the body, with the dorsal side exhibiting a smoky brown coloration and the ventral side paler grey. Juveniles display a darker grey fur overall, which lightens with age. The species is distinguished by a horseshoe-shaped noseleaf surrounding the nostrils, featuring a pointed lancet at the top, and ears that lack a tragus but include a broad antitragus for structural support. Additionally, it has small, strong feet adapted for clinging to surfaces. Sexual dimorphism is subtle, with females slightly larger than males, particularly in forearm length (averaging 38.5 mm in females versus 37.2 mm in males). The average lifespan in the wild is about 3 years, though individuals can reach up to 30 years in captivity.

Sensory adaptations

The lesser horseshoe bat (Rhinolophus hipposideros) exhibits specialized hearing adaptations that support its reliance on echolocation for navigation and foraging. Its pinnae are notably large, measuring 14–19 mm in length, which enhances directional sensitivity to incoming echoes by funneling and amplifying high-frequency sounds. The cochlea features a specialized acoustic fovea—a region with heightened neural representation and mechanical tuning—that processes constant-frequency components of echolocation signals with exceptional sharpness, allowing discrimination of subtle Doppler shifts from prey movement. This structure divides the cochlea into distinct functional zones, including an expanded basal region optimized for the bat's species-specific reference frequency around 110 kHz. Vision in the lesser horseshoe bat is functional but subordinate to auditory cues, suited primarily to dim-light environments. The eyes are relatively large compared to the bat's small body size (head-body length 35–45 mm), with a rod-dominated and large corneal surface area that improve sensitivity to low illumination levels during crepuscular activity. However, the is constrained by the prominent noseleaf, limiting its role in precise spatial tasks. Olfaction contributes to social interactions, such as and mate identification, supported by a retaining only about 10% non-functional genes—higher functionality than in many other echolocating bats. Tactile hairs distributed across the wing membranes serve as airflow sensors, providing feedback for flight stability and close-range prey detection by registering subtle air disturbances from . Adaptations for insectivory include a dental formula of I 1/1, C 1/1, P 2/3, M 3/3 (total 32 teeth), with robust molars featuring broad, rugose crushing surfaces and sharp cusps designed to fracture and process the chitinous exoskeletons of prey like and moths.

Habitat and distribution

Habitat preferences

The lesser horseshoe bat (Rhinolophus hipposideros) prefers warm, temperate environments such as woodlands, riparian zones, and landscapes, which provide suitable opportunities and . These bats are often found in sheltered valleys with dense scrub or extensive tree cover, favoring and areas up to 1,160 meters in during summer and extending to 2,000 meters in winter, particularly in southern parts of their range where higher altitudes are tolerated. They avoid open plains and arable lands, instead relying on linear features like hedgerows and treelines for in fragmented habitats. Roosting sites are critical and include natural caves, abandoned mines, and structures such as old churches, barns, and cellars, with maternity colonies typically forming in warm attics of historic during summer. Winter hibernacula demand high levels of 85–95% and stable temperatures between 8–12°C to minimize expenditure and prevent disturbance. These roosts are selected for their microclimatic stability, often in limestone-rich areas that maintain consistent conditions year-round. Microhabitat selection emphasizes proximity to water bodies and insect-abundant grounds, such as grazed grasslands, parklands, and edges within 2.5 km of roosts, where the bats hunt low-flying amid vegetation. As a thermophilic , the lesser horseshoe bat exhibits sensitivity to cold snaps, which can increase juvenile mortality and disrupt , underscoring their reliance on mild climatic conditions in core habitats.

Geographic distribution

The lesser horseshoe bat (Rhinolophus hipposideros) has a distribution centered in , extending from western eastward to , and southward into from the region (, , and ) to , as well as across Asia Minor including parts of the . Its range encompasses Mediterranean islands and reaches the northern limit in the , , and , though populations there are sparse; the eastern extent includes the southern and up to and . Recent observations indicate an expansion into , with the first confirmed winter hibernations recorded in northern underground sites during the 2024–2025 monitoring season, suggesting a northward shift possibly driven by climate warming. Two subspecies are recognized within this range: R. h. hipposideros predominates in the western portions, including most of , , and western Minor, while R. h. minimus occupies the eastern areas, from and the through the to northeastern (Sudan to ). These subspecies exhibit karyotypic differences, with R. h. minimus having a diploid number of 58 chromosomes, compared to 54–56 in R. h. hipposideros. Key regional populations highlight the species' patchy occurrence: in the , it is restricted to and southwest , with an estimated 50,400 individuals (as of 2018); in Ireland, approximately 15,000 bats (as of 2023) form an internationally significant population confined to western counties; the host notable colonies, particularly on where the species is relatively common in habitats; in , it inhabits coastal and lowland areas; and in , northern humid zones support smaller, rarer groups. Population trends reflect historical declines followed by regional recoveries. Across , lesser horseshoe bat numbers fell by approximately 50% after the due to habitat loss and roost disturbance, with dramatic reductions in western and central regions during the leading to local extinctions in some areas. However, recent monitoring shows increases, including a 77% rise in the UK from 1999 to 2012 and stable or growing populations since the early 2000s (with 2.36% annual growth to 2023), attributed to efforts and milder winters. In , long-term trends indicate a 192% increase from 1999 to 2023 based on surveys.

Behavior and ecology

Roosting and hibernation

The lesser horseshoe bat forms maternity colonies from April to September, primarily in warm, man-made structures such as attics, barns, and other buildings that provide stable, humid conditions conducive to reproduction and pup rearing. These colonies typically consist of 10-200 females, though larger aggregations up to 400 individuals have been documented in high-quality habitats. In maternity colonies, females cluster for and protection during pup rearing. Males generally roost solitarily or in small, separate groups away from maternity sites, often in similar warm locations but without the dense clustering of females. Juveniles and dispersing individuals utilize transition roosts, including night roosts—temporary resting sites during bouts—located within 1 to 2.5 km of the primary maternity colony to facilitate and digestion. In preparation for winter, lesser horseshoe bats migrate to sites from to , selecting cool, humid underground environments such as caves, mines, cellars, and souterrains that maintain stable microclimates. These sites are chosen for their ability to support , a state of metabolic suppression where body closely tracks ambient temperatures of 6 to 11°C, allowing the bats to minimize energy expenditure over the non-reproductive season. During , activity levels are low, with bats remaining largely immobile; however, in milder winters, they may periodically arouse to feed on available , preventing complete . Colony sizes in hibernacula vary but often include dozens to hundreds of individuals, with bats showing site fidelity by returning to the same locations annually. Within roosts, lesser horseshoe bats exhibit characteristic colony dynamics, hanging freely by their feet with noses pointed upward and wings wrapped around their bodies, particularly during or deep to conserve heat and reduce exposure. This posture is unique among many species and facilitates quick arousal if needed. The species is largely sedentary, with individuals typically moving 5-10 km between summer and winter roosts, though dispersals up to 150 km or more have been recorded in response to habitat availability or disturbance. These local movements underscore the bat's reliance on proximate, connected roosting networks for survival.

Foraging and diet

The lesser horseshoe bat (Rhinolophus hipposideros) is primarily insectivorous, with its consisting mainly of small flying captured during nocturnal foraging. Moths () form the largest component, comprising 31–53% of the by occurrence across various studies, followed by flies (Diptera) at 24–38%. Other prey includes neuropterans, hemipterans, trichopterans, and occasionally spiders or , though these rarely exceed 10–15% of the total volume. Foraging occurs primarily through aerial hawking close to the ground or and from foliage, targeting clutter edges in woodlands, orchards, and riparian areas as a specialist. Bats typically within 1–2.5 km of roosts, with 50% of activity concentrated within 600 m, though maximum ranges can extend to 4–5 km in resource-rich landscapes. Nightly consumption equates to 20–30% of body mass, reflecting the species' high metabolic demands for sustained flight and echolocation-based prey detection. Dietary composition varies seasonally, with moths dominating in summer due to abundance in woodlands and pastures, while winter feeding in mild conditions shifts toward crane flies () and fungus gnats (), comprising up to 24% and 13% respectively in British samples. Juveniles initially favor soft-bodied dipterans (52–53%) over harder lepidopterans (30–31%), transitioning to adult-like proportions by weaning at 6–7 weeks, which supports rapid growth amid elevated energy needs.

Reproduction

The lesser horseshoe bat exhibits a seasonal reproductive cycle adapted to its temperate . Mating primarily occurs during autumn swarming aggregations at entrances and other sites, where both sexes gather in large numbers for promiscuous copulations involving multiple partners. Females store viable in their reproductive tracts throughout , delaying fertilization until ovulation in spring, typically April. Following and fertilization in spring (typically ), lasts about 60 days, resulting in the birth of a single pup per female (twins are rare) from mid-June to early July. Newborn pups weigh around 1.8 g, are born hairless with closed eyes, and initially cling to their mothers' teats while being carried during foraging flights. Pups develop rapidly, opening their eyes at about 10 days, beginning to fly at 3 weeks, and becoming fully independent at 6–7 weeks when occurs. is typically reached by the end of the first year. Maternity roosts form in warm, stable sites such as buildings, mines, or caves starting in , comprising primarily pregnant and lactating females with up to 20% males and numbering 10–250 individuals. These colonies exhibit high site fidelity, and stable conditions within them contribute to elevated pup survival rates by providing consistent and protection.

Echolocation

The lesser horseshoe bat (Rhinolophus hipposideros) utilizes a high-duty-cycle echolocation system consisting of long constant-frequency (CF) pulses followed by brief frequency-modulated (FM) sweeps. The CF component typically centers at approximately 110 kHz, with frequencies ranging from 105 to 111 kHz and an average duration of 20–30 ms; the terminal FM sweep facilitates precise target ranging during the approach phase. Echolocation calls are generated in the and emitted through the nostrils, where the characteristic noseleaf directs and shapes the outgoing pulses for focused beam formation; this structure shows subtle , with males producing calls at lower frequencies (mean ~105 kHz) than females (mean ~109 kHz). The system relies on Doppler shift compensation to discern target velocity, as returning echoes from moving exhibit frequency shifts relative to the emitted CF pulse, enabling the bat to track prey dynamics; this allows detection of small (e.g., 9 mm targets) at distances of 2–5 m. Juveniles display a slight age-related frequency drop compared to adults, potentially reflecting developmental adjustments in . Unlike the (Rhinolophus ferrumequinum), which emits CF pulses at lower frequencies around 81 kHz for broader detection ranges, the lesser horseshoe bat's higher-frequency calls afford greater resolution suited to cluttered environments but limit maximum range.

Conservation

Status and threats

The lesser horseshoe bat (Rhinolophus hipposideros) is classified as Least Concern on the global , with a decreasing population trend. In , its status was updated to Least Concern in 2023, improving from Near Threatened in the 2016 assessment. The is protected under Annex II and IV of the EU , requiring the designation of Special Areas of Conservation and strict protection measures. Major threats to the species include habitat loss and fragmentation from agricultural intensification, development, and practices, which reduce available roosting and areas. Roost disturbance by human activities, such as in caves or renovations in buildings, can cause abandonment of maternity and sites. use in diminishes insect prey populations, impacting success, while poses risks through warmer winters that may shorten periods and increase energy demands. Populations have declined significantly across much of the range, with estimates indicating reductions exceeding 90% across much of since the due to habitat changes and pesticide exposure. In , ongoing declines are linked to habitat degradation from and . These trends highlight the vulnerability of peripheral populations at the range edges. Emerging issues include from artificial lighting, which disrupts foraging routes and emergence from roosts, potentially fragmenting habitats. The fungus () is present on European bats, including the lesser horseshoe bat, but does not currently cause mass mortality as in ; however, evolving pathogen dynamics could pose future risks.

Protection and

The lesser horseshoe bat (Rhinolophus hipposideros) is protected under international agreements, including Appendix II of the , which requires special protection for its habitats and populations across Europe. It is also covered by the Agreement on the Conservation of Populations of European Bats (EUROBATS), established under the Bonn Convention, which promotes coordinated conservation measures for all European bat species, including monitoring and habitat management. Nationally, in the , the species and its roosts are safeguarded by the (as amended), prohibiting intentional killing, injury, or disturbance without a license. Conservation initiatives include the Lesser Horseshoe Bat Species Action Plan for (2022–2026), which outlines targeted actions such as protection, habitat connectivity improvements, and public awareness to maintain the species' stronghold in western counties. At the European level, the Action Plan for the Conservation of All Bat Species in the (2018–2024), with complementary updates, supports multi-species strategies including legal enforcement, , and cross-border collaboration to address and threats like wind energy development. In the , the Bat Conservation Trust runs the National Bat Monitoring Programme, which tracks population trends through standardized surveys of and sites, informing adaptive management. As of the 2024 National Bat Monitoring Programme report, the population continues a long-term increase since 1999, though short-term trends (2019–2024) show stability or emerging declines in some regions. Recovery efforts have shown positive results, with the population increasing by approximately 75% in winter roost counts from 1999 to 2012, attributed to enhanced protections and habitat management. In , a national Species Action Plan drafted in 2013 has contributed to stabilizing populations through roost and landscape connectivity projects. Roost restoration initiatives, such as creating alternative structures during building renovations, have successfully supported colony persistence, with examples showing increased occupancy post-intervention. Future actions emphasize enhancing habitats via agri-environment schemes that promote diverse hedgerows and restoration to bolster prey availability. adaptation monitoring, including predictive modeling of range shifts due to warming temperatures, guides proactive adjustments. Ongoing surveys in , such as winter monitoring in 2025, aim to document potential northward expansion and inform regional protections.

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