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Blarina

Blarina is a of relatively large in the family Soricidae, subfamily Soricinae, and tribe Blarinini, comprising five extant species endemic to : the (B. brevicauda), (B. carolinensis), Elliot's short-tailed shrew (B. hylophaga), short-tailed shrew (B. peninsulae), and Sherman's short-tailed shrew (B. shermani). These are distinguished by their robust, cylindrical bodies measuring 7–13 cm in length (including a short of 1–3 cm), velvety fur typically grayish-brown above and paler below, small eyes and concealed ears, and a pointed snout adapted for and terrestrial foraging. Native to habitats across the eastern and and southern , they prefer moist, vegetated areas such as and mixed , grasslands, brushy fields, and forest edges with ample leaf litter or ground cover for burrowing and concealment. A defining feature of the genus Blarina is its members' production of a potent neurotoxic saliva, unique among North American mammals, secreted from enlarged submaxillary glands and delivered via grooved lower incisors to subdue and small prey. This , composed of proteins like blarina toxin, can immobilize animals several times their size, such as mice or salamanders, facilitating predation on , , centipedes, and occasionally carrion or seeds. With high metabolic rates requiring near-constant feeding (up to their body weight daily), Blarina exhibit aggressive, territorial behavior, constructing elaborate underground systems for nesting and storing food. Their adaptations, including strong forelimbs and 32 red-tipped teeth for grinding tough exoskeletons, enable them to thrive in diverse ecological roles as both predators and occasional prey for , snakes, and mammals.

Taxonomy

Etymology

The genus Blarina was established by the British zoologist in 1838 as a within the genus Sorex (previously classified under Corsira), in his systematic revision published in the Proceedings of the Zoological Society of . introduced the name to distinguish a group of North American short-tailed shrews characterized by their robust build and reduced tail length compared to other Sorex species. Gray designated the type species of Blarina as Corsira (Blarina) talpoides Gray, 1838, which was based on Sorex talpoides Gapper, 1830—a junior synonym of Sorex brevicauda Say, , now recognized as Blarina brevicauda, the . This designation anchored the subgenus to specimens from eastern , reflecting Gray's effort to reorganize the diverse and often confused of soricids based on morphological traits like and pelage. In 1842, French naturalist René Primevère Lesson elevated Blarina to full generic status in his Nouveau tableau du Royaume Animal, separating it definitively from Sorex due to differences in cranial structure and tail proportions. Subsequent reclassifications, particularly in the late 19th and early 20th centuries, refined the genus's boundaries; for instance, Merriam's 1895 revision of American shrews confirmed Blarina as distinct from related genera like Notiosorex and incorporated additional based on geographic variation. Today, Blarina remains a well-defined in the Soricidae, encompassing five extant primarily distributed across , though the specific status of B. peninsulae (as a full species or of B. carolinensis) remains debated based on karyotypic and genetic evidence as of 2025.

Classification

Blarina is classified within the kingdom Animalia, phylum Chordata, class Mammalia, order Eulipotyphla, family Soricidae, subfamily Soricinae, tribe Blarinini, and genus Blarina. The genus belongs to the red-toothed shrews, characterized by reddish pigmentation on their teeth due to iron deposits, a trait shared across the subfamily Soricinae. Phylogenetically, Blarina forms a monophyletic group within Soricidae, with molecular analyses confirming its close relationship to other North American soricines; specifically, the genus is positioned as to Cryptotis, the small-eared , based on mitochondrial sequences and broader soricid phylogenies that place both genera in a distinct from Old World soricines like Sorex. This positioning reflects a shared evolutionary history in the Nearctic region, with Blarina diverging from Cryptotis-like ancestors during the to transition, though some studies suggest potential affinities to Notiosorex in broader tribal rearrangements. The fossil record of Blarina extends from the late to the Recent, with ancestral forms emerging from the blarinine stem in the middle to late , as evidenced by early records like Blarina gidleyi from deposits. Key Pleistocene fossils include those attributable to Blarina carolinensis, known from over 25 sites across nine U.S. states, indicating a historical range expansion northward during periods before post-Pleistocene retraction. The genus's paleobiogeography shows persistence in both warm-moist and cool-moist faunas through the Blancan to Rancholabrean land-mammal ages, underscoring its adaptability amid climate shifts.

Description

Morphology

Members of the genus Blarina are robust, short-legged characterized by a cylindrical body, long pointed , and short , distinguishing them from other North American soricids. Across the five species, total length ranges from 70 to 140 mm, with head and body lengths typically 66–110 mm and lengths of 12–30 mm; weights vary from 5 to 30 g, with B. brevicauda being the largest and B. peninsulae the smallest. These dimensions reflect adaptations for a lifestyle, with larger body sizes in northern species correlating to colder climates. The pelage is dense and velvety, providing and ; dorsal fur is typically grayish-brown to slate-gray, often with a silvery tinge, while ventral fur is paler gray. Seasonal variation occurs, with winter pelage darker and thicker than summer coats. Cranially, Blarina shrews possess a robust with a broad braincase and short rostrum, adapted for powerful action; eyes and ears are small and concealed in the fur, emphasizing reliance on other senses. The dental formula is I 3/1, C 1/1, P 3/1, M 3/3 = 32 teeth, featuring red pigmentation on the tips of the upper incisors, which may aid in identification and is unique among soricines. The is short, comprising less than 50% of head and body length, and covered in fine , aiding during burrowing. Limbs are short with thick forefeet equipped with strong claws for tunnels and turnover, while hind feet are narrower for .

Physiology

Blarina species possess exceptionally high metabolic rates, among the highest recorded for mammals, which demand continuous energy intake to prevent . Individuals must consume food equivalent to up to three times their body weight daily and feed every 2-3 hours to maintain vital functions, as their small size results in rapid heat loss and depletion. This elevated supports their active lifestyle but limits survival without food to mere hours, with full body turnover occurring in 9-24 hours depending on and . To against short-term , Blarina can store reserves, particularly during summer, enhancing overwinter survival. A distinctive physiological is the system, centered in the enlarged submaxillary salivary glands, which secrete blarina (BLTX), a 4.6 kDa exhibiting kallikrein-like activity. BLTX enhances catalytic efficiency compared to non-toxic homologs, leading to prey immobilization through mechanisms such as a precipitous drop in () and induced , resulting in without direct neurotoxic effects. This capability, rare among mammals, facilitates efficient subduing of larger or more mobile prey. Sensory physiology in Blarina emphasizes olfaction and tactile perception over . The is acute, aiding in prey detection and environmental navigation, while vibrissae on the provide highly sensitive touch cues for in low-light or cluttered habitats. plays a minor role, with small, degenerated eyes limiting resolution to basic light detection. For thermoregulation, Blarina maintain euthermy through a high augmented by the heat increment of feeding, where postprandial oxygen consumption rises by approximately 18% for up to 2 hours, elevating body temperature by 0.6°C to offset cold exposure. During food scarcity, they can enter reversible or shallow , reducing metabolic demands to conserve energy, though this is less pronounced than in smaller soricines.

Distribution and habitat

Geographic range

The genus Blarina is endemic to and occupies a broad continental range across eastern and central regions, extending from the Canadian provinces of and in the east to southern in the west, and southward through the central and to eastern and south-central . This distribution is primarily confined to areas east of the , with the genus reaching its western limits in the . The latitudinal extent of Blarina spans roughly 26°N to 50°N, encompassing diverse physiographic provinces from the to the valley, though some northern populations exhibit disjunct occurrences separated by unsuitable habitats. Species-specific ranges within the vary, with B. brevicauda dominating the northern portions and B. carolinensis and B. shermani confined more southerly, as detailed in subsequent sections. Fossil records provide evidence of post-Pleistocene range expansions for Blarina, with specimens from sites in and indicating persistence in southern refugia during glacial maxima, followed by northward recolonization as ice sheets retreated approximately 10,000–15,000 years ago. Phylogeographic analyses further support this pattern, revealing genetic signatures of historical fragmentation and subsequent demographic expansions across formerly glaciated landscapes in eastern .

Habitat preferences

Blarina shrews exhibit a semifossorial , primarily inhabiting moist, vegetated environments that provide suitable conditions for burrowing and cover. They favor forests, grasslands, shrublands, and the edges of wetlands, where dense vegetation and leaf litter layers offer protection and microhabitat stability. These habitats are typically mesic, with consistent moisture supporting the that form a key part of their , though the shrews avoid extremes of or . Soil composition is critical for Blarina species, as their semifossorial habits require loose, friable substrates that allow efficient and . They thrive in damp but well-drained soils rich in , such as those under or grassy , and actively avoid dense clay soils that impede digging or periodically flooded areas that could collapse burrows. This preference for workable earth enables them to create extensive subterranean networks, often just below the surface or deeper in stable ground. Within their North American distribution, Blarina shrews occur from lowlands to higher elevations, reaching up to approximately 1,300 m in mountainous regions like the Appalachians, where they continue to select sites with abundant cover and leaf litter for concealment. This altitudinal tolerance reflects their adaptability to varied terrains, provided the core requirements of moisture and soil are met.

Behavior

Activity patterns

Blarina species exhibit a polyphasic activity characterized by short bursts of activity interspersed with periods of rest, driven by their exceptionally high metabolic rates that necessitate frequent to meet energetic demands. Observations indicate bimodal peaks in activity, typically occurring shortly after sunset and , though individuals remain active to varying degrees throughout the 24-hour cycle, with overall activity comprising only about 16% of the day. This pattern aligns with their strongly nocturnal tendencies under entrained light-dark cycles, but free-running rhythms in constant conditions reveal persistent low-level activity without strict ultradian feeding bouts. As semifossorial mammals, Blarina navigate both surface runways—often those created by larger mammals like voles—and subsurface tunnels within the top 10–50 cm of , facilitating efficient movement in leaf litter, , or under . Their involves quick, jerky runs with sudden stops, enabling bursts of speed reaching up to 4.25 m/s in juveniles during or exploration, though average travel is more measured to conserve . They also employ ultrasonic vocalizations for echolocation to navigate in low-visibility conditions. These movements support territorial defense, with individuals patrolling home ranges averaging about 2.5 (0.1–1.8 depending on and sex) during active periods. Seasonally, activity levels decline in winter, where Blarina enter near-torpid states during inactive periods, reducing metabolic rate, body temperature, and to cope with and limited food availability, often relying on subnivean under snow cover. In contrast, activity intensifies during the breeding season (typically to ), with increased and movement to support , though overall patterns remain influenced by temperature, showing greater daytime consistency in colder conditions.

Social organization

Blarina shrews exhibit a predominantly solitary , with adults maintaining exclusive territories that are vigorously defended against intruders. These territories are roughly comparable to their home range sizes and allow residents to monopolize resources while minimizing energy expenditure on defense. Territories are primarily defended through scent marking along boundaries using specialized dermal glands, which deters nomadic individuals from entering, and through vocalizations such as high-pitched chirps and trills that serve as warning signals during encounters. Populations of Blarina consist of a mix of resident individuals, who occupy stable territories, and nomadic that wander between areas, particularly during periods of low prey availability. densities vary with quality, typically ranging from 5 to 30 individuals per in optimal old-field or forested environments, though they can fluctuate dramatically up to over 100 per in peak conditions. is a key aspect of social interactions, with frequent fights occurring between residents and intruders, often involving chases, threats, and venomous bites delivered via the shrew's specialized salivary glands. These bites, containing a potent , can incapacitate rivals and reinforce territorial boundaries, though such confrontations are energetically costly. Rare instances of communal nesting have been noted among females, particularly during , where tolerance of conspecifics may temporarily override solitary tendencies to share systems for . Territorial patrols are influenced by the shrews' polyphasic activity cycles, with increased vigilance during crepuscular periods to monitor boundaries.

Ecology

Diet and foraging

Blarina species exhibit a primarily carnivorous and insectivorous diet, consisting mainly of such as (annelids), snails (gastropods), lepidopteran larvae, centipedes (chilopods), arachnids, and other , supplemented occasionally by small vertebrates including salamanders, small , and reptiles, as well as minor amounts of plant material and seeds. This dietary composition reflects their opportunistic predation on soil-dwelling and surface abundant in their forested habitats. Foraging in Blarina occurs predominantly through ambush predation within subterranean tunnels and leaf litter, where individuals remain stationary or move slowly to detect prey or scents before striking. They utilize ultrasonic clicks as an echolocation-like to navigate dark tunnels and locate prey, emitting pulses in the 30-60 kHz range to interpret echoes for spatial orientation and . Upon capturing prey, even those larger than themselves, Blarina deliver a venomous bite via grooved lower incisors to immobilize it rapidly, facilitating handling and transport. Due to their elevated metabolic rate, Blarina individuals must consume approximately their body weight in food daily, often approaching or equaling 1-2 times body mass under normal conditions to sustain energy demands. Excess prey is frequently cached in burrows rather than consumed immediately, with studies showing that over 85% of captured items are stored for later retrieval, enhancing during periods of . This frequent feeding pattern is essential to support their high physiological requirements for and activity.

Reproduction

Blarina shrews exhibit a breeding season that typically spans from March to October, enabling females to produce 2 to 4 litters annually. Litter sizes generally range from 4 to 8 young, though extremes of 2 to 10 have been recorded across species. Gestation periods last 21 to 30 days, resulting in the birth of altricial young that are blind, hairless, and weigh approximately 1 gram. These offspring are weaned at 20 to 25 days and reach within 2 to 3 months, allowing many individuals to breed in their first year. Parental care is provided solely by females, who construct and maintain nests from grass and leaves, nurse the young, and aggressively defend them from predators; males play no role in rearing. In the wild, Blarina individuals typically have a lifespan of 1 to 2 years, with reproduction ceasing after this period.

Species

Diversity

The genus Blarina comprises five currently recognized of short-tailed , all endemic to : B. brevicauda, B. carolinensis, B. hylophaga, B. peninsulae, and B. shermani . Taxonomic debates persist regarding the distinctiveness of certain taxa, such as the status of B. shermani relative to B. carolinensis in southern regions and B. peninsulae as a potential of B. carolinensis, with some studies proposing synonymy based on morphological and genetic overlap, though molecular and karyotypic evidence supports separation . The of Blarina occurred primarily during the Pleistocene epoch, when ancestral forms diverged in response to glacial cycles and the development of diverse biomes across eastern , including forests, grasslands, and transitional habitats . Fossil records indicate that B. brevicauda or a close precursor served as the basal lineage, with speciation driven by isolation in refugia south of the , leading to adaptations for varied ecological niches . Genetic variation among Blarina species is marked by distinct s and limited interspecific hybridization, reflecting strong . Most species exhibit a diploid number (2n) of 48 and a fundamental number (FN) of 48, though B. carolinensis shows variability with 2n ranging from 37 to 46 and FN of 44–45, while B. peninsulae has a distinct (2n = 46); contact zones between species reveal minimal , with hybridization rates below 5% in overlapping ranges .

Key species accounts

The (Blarina brevicauda) is the largest species in the , reaching weights of up to 30 g and total lengths exceeding 110 mm, with a robust build, short , and dense slate-gray fur. It is widespread across the and southern , primarily inhabiting mixed and deciduous forests, as well as edges of fields and wetlands. The (B. carolinensis) is notably smaller than its northern counterpart, typically weighing 15–20 g and measuring 7–10 cm in total length, featuring soft, uniformly grayish-brown fur and a proportionally short, furred . Its range spans the , from southward through the to eastern , favoring woodlands, brushy fields, and moist deciduous areas. Elliot's short-tailed shrew (B. hylophaga), an intermediate-sized weighing 13–17 g with total lengths of 92–121 mm, exhibits a stocky form with concealed ears and a pointed adapted to open habitats. It occurs across prairies and grasslands, extending through the from to and eastward to and . The short-tailed shrew (B. peninsulae) is a medium-sized endemic, similar in build to B. carolinensis but with subtle morphological distinctions such as a distinct , and it inhabits coastal marshes and wetlands of southern . Its restricted range and vulnerability to habitat loss from and sea-level rise have led to concerns for its conservation, with ongoing efforts focused on preservation. The Sherman's short-tailed shrew (B. shermani) is a medium-sized , typically weighing 16–22 g and measuring 80–105 mm in total length, with soft grayish-brown fur and a short . It is endemic to peninsular , from central regions southward, inhabiting dense herbaceous areas, edges of marshes and basin wetlands, mesic flatwoods, and moist mixed forests.

Ecoepidemiology

Disease reservoirs

Short-tailed of the Blarina, particularly B. brevicauda, serve as significant hosts for several zoonotic pathogens, contributing to the maintenance and transmission of diseases in North American ecosystems. They are competent reservoirs for , the of , with rates detected via xenodiagnosis reaching up to 78.6% in tested populations from endemic areas in during the early 1990s. Similarly, Blarina species host Babesia microti, the protozoan parasite responsible for human babesiosis, exhibiting competence levels exceeding 17% in wildlife surveys from the , where patent parasitemias have been observed in over 60% of sampled individuals at certain sites. These high prevalences underscore Blarina's role in amplifying pathogen circulation among vertebrate hosts, though their overall contribution to rates is moderated by lower infestation levels on immatures compared to . Beyond Lyme disease agents, Blarina shrews harbor other tick-borne viruses, including Powassan virus (a flavivirus transmitted by Ixodes scapularis ticks), for which they demonstrate reservoir competence through direct viral detection in tissues and positive associations between shrew bloodmeals in ticks and site-level infection prevalence. Hantaviruses, such as Camp Ripley virus, have also been identified in B. brevicauda, with prevalence up to 25% in lung tissues from Minnesota populations, indicating shrews as potential reservoirs for these rodent- and soricomorph-associated pathogens despite their evolutionary divergence from classic rodent-borne strains. Transmission of these pathogens primarily occurs via questing Ixodes scapularis ticks, which acquire infections during bloodmeals on infected shrews and subsequently pass them to other hosts, including humans. Studies since the 1990s, including xenodiagnostic experiments and field surveillance, have consistently confirmed Blarina's reservoir competence for both bacterial and viral agents. Ecological traits of Blarina species enhance their role as disease reservoirs. Their high local population densities, which can fluctuate 3- to 4-fold annually in fragmented habitats, combined with burrowing behaviors, increase encounters with ground-dwelling vectors like I. scapularis larvae and nymphs in leaf litter and interfaces. This contact facilitates spillover, as support a substantial proportion of larval feedings—up to 35% overall and 55% of those on infected ticks in northeastern sites—thereby sustaining enzootic cycles despite their short lifespans. Ongoing research highlights how these factors position Blarina as inconspicuous yet impactful contributors to dynamics in wildlife.

Interactions with humans

Short-tailed shrews of the genus Blarina, particularly B. brevicauda, occasionally interact with humans through defensive bites when handled, delivering venomous that causes localized , swelling, and burning sensations akin to a . These effects can persist for hours to days, with symptoms including numbness, throbbing, and in rare cases, systemic reactions such as drowsiness or cramps, though no fatalities have been reported. The venom contains paralytic peptides like BPP2, which activate T-type calcium channels (hCav3.2) involved in , leading to and severe . Such envenomations are uncommon, typically occurring during handling or accidental encounters in residential areas. In ecoepidemiological contexts, Blarina species serve as reservoirs for zoonotic pathogens, amplifying transmission risks to humans primarily through ecological overlap in urban-rural interfaces. For instance, B. brevicauda hosts Borrelia burgdorferi, the causative agent of Lyme disease, feeding approximately 55% of infected Ixodes scapularis tick larvae and contributing to over 20% of infected nymphs that quest for human hosts. This role elevates Lyme disease incidence in North American forests and suburbs, where shrews' high reservoir competence (42%) sustains tick infection rates. Blarina shrews also harbor orthohantaviruses, such as Camp Ripley virus, which has the potential to cause hantaviral disease in humans similar to hemorrhagic fever with renal syndrome via aerosolized excreta exposure, though no direct shrew-human transmission or confirmed cases have been documented. Additionally, they carry Powassan virus, a flavivirus transmitted by ticks that leads to encephalitis in humans, with shrews acting as amplifying hosts in endemic areas. A novel henipavirus, Camp Hill virus, was detected in B. brevicauda tissues in Alabama, phylogenetically related to human-pathogenic viruses like Langya henipavirus, posing potential spillover risks through contact with infected shrews or their secretions in shared habitats. No human cases of Camp Hill virus infection have been confirmed, but the shrews' proximity to human populations heightens surveillance needs for paramyxoviral emergences. As of November 2025, no human infections with Camp Hill virus or Camp Ripley virus have been reported, but surveillance continues due to proximity to human habitats. Urban adaptation by Blarina species further increases exposure opportunities, such as via agricultural activities or vector-mediated pathways.

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