Soricomorpha

Diversity

Order Soricomorpha includes three extant families, Soricidae (true shrews), Talpidae (moles, shrew-moles, and desmans), Solenodontidae (solenodons), and one extinct family, Nesophontidae (West Indies shrews). The name Soricomorpha was designated by Gregory in 1910 and means 'shrew-formed'. Members of this order were previously included in the now defunct order, Insectivora . The most abundant family in order Soricomorpha is Soricidae , which encompasses about 300 species, followed by family Talpidae , which includes about 42 species, and family Solenodontidae , with two extant and two extinct species. The smallest members of this order are pygmy white-toothed shrews , which weigh about 3 grams and are about 35 mm long, from head to body. Cuban solenodons are the largest species in this order, and can be up to 600 mm long. Shrews are very abundant and thrive in high latitudes. Due to their small body size and surface area to mass ratio, most soricomorphs must consume large amounts of food and do not hibernate. Families Solenodontidae and Talpidae include fossorial species, many of which are significantly adapted to living underground. Solenodons are very rare; only 37 Cuban solenodons have ever been caught.

Geographic Range

Members of order Soricomorpha are found in the Nearctic, Palearctic, Neotropical, Ethiopian, and Oriental ranges, they are only excluded from landmasses in the Australian and Antarctic regions. Family Solenodontidae (solenodons) includes two extant species, Cuban and Haitian solenodons , which reside on Cuba and Hispaniola, respectively. Family Soricidae (shrews) includes three subfamilies, Soricinae , Myosoricinae and Crocidurinae . Subfamily Soricinae inhabits North and Central America, Europe and most of Asia. Subfamily Myosoricinae is mostly found in central and southern Africa, they are especially concentrated from Cameroon to Tanzania. Subfamily Crocidurinae inhabits Africa and most of Southern Asia. Family Talpidae (moles and desmans) are found mostly in Europe and Asia. They are also found in Southern Canada and most of the United States. Desmans specifically are found in Europe and Russia.

• Biogeographic Regions
• nearctic nearctic
  • introduced introduced
  • native native
• palearctic palearctic
  • introduced introduced
  • native native
• oriental oriental
  • introduced introduced
  • native native
• ethiopian ethiopian
  • native native
• neotropical neotropical
  • introduced introduced

• Other Geographic Terms
• cosmopolitan cosmopolitan

Habitat

Members of families Solenodontidae and Talpidae are fossorial; this is reflected in their fusiform body and short powerful limbs. Their tunnel networks vary in depth; some moles are able to dig as deep as 150 cm. Moles' tunnels are usually 4 to 5 cm wide. Moles live under pastures, woodlands, and gardens and do not do well in acidic soil. Some aquatic species of moles are found near fresh water, or brackish and somewhat salty water. Members of family Soricidae prefer areas that are not too dry, although some species are capable of surviving in the desert. These animals occupy spaces under damp leaf litter. Shrews must remain near wet areas due to their small body size and the resulting high rate of desiccation. Desmans of family Talpidae are semi-aquatic, found in a wide range of aquatic habitats such as swamps, streams, and rivers. Russian desmans prefer ponds and marshes, while Pyrenean desmans prefer fast moving water. Their feet are webbed and fimbriated for swimming.

• Habitat Regions
• terrestrial terrestrial
• freshwater freshwater

• Terrestrial Biomes
• taiga taiga
• savanna or grassland savanna or grassland
• forest forest

• Aquatic Biomes
• lakes and ponds
• rivers and streams

• Wetlands
• marsh marsh
• swamp swamp
• bog bog

• Other Habitat Features
• suburban suburban
• agricultural agricultural
• riparian riparian

Systematic and Taxonomic History

Currently, 4 families are included in order Soricomorpha , Solenodontidae , Soricidae , Talpidae and the extinct family, Nesophontidae . Previously, these families, along with several others, were included in the now defunct order, Insectivora . Recent members of order Insectivora are now separated into 3 different orders, including Afrosoricida , Erinaceomorpha , and Soricomorpha , however, historically, orders Macroscelidea , Scandentia , and Dermoptera were also included in this large order of insectivores. In 1945, Simpson declared that order Insectivora was little more than “a scrap basket for small mammals”, this sentiment was largely shared by other mammalogists. Prior to its dissolution, order Insectivora included solenodons, tenrecs, golden moles, hedgehogs, gymnures, shrews, moles, desmans, flying lemurs, tree shrews, and elephant shrews. Currently, species included in order Soricomorpha include solenodons, shrews, moles, and desmans. However, the current grouping is also under a great deal of debate and may be subject to change. For instance, order Soricomorpha closely groups shrews and moles, although recent mitochondrial genomic studies have suggested a very close relationship between shrews and hedgehogs, but not a similarly close relationship between hedgehogs and moles. Likewise, other studies have suggested solenodons are more closely related to tenrecs or myomorph rodents, than other soricomorphs. The only officially recognized synapomorphy within this grouping is an elongated alisphenoid canal. Soricomorpha was originally used to separate groups of species that did not fit with other members of order Insectivora . While they were still included in order Insectivora , their grouping was placed in a separate suborder, Lipotyphla . Occasionally, orders Soricomorpha and Erinaceomorpha are still grouped together into Lipotyphla . The relationship between species described herein is largely based on Hutterer’s definitions as recorded in Wilson and Reeder 2005. There are, however, potential discrepancies between this source and other respected sources.

Members of order Soricomorpha are often the subject of evolutionary research due to their primitive characteristics, which are likely similar to some of the earliest mammals. Most members of family Nesophontidae became extinct about 500 years ago, although evidence suggests a few species survived until the early 18th century. Based on fossil evidence, family Nesophontidae included 8 species, within a single genus, Nesophontes . The grouping of nesophontids within order Soricomorpha is questionable, as their relationship to other members of the order is difficult to trace. Likewise, family Solenodontidae is also a questionable addition to the order. This family includes one genus, Solenodon , and 4 species, two of which are extinct, however, Cuban solenodons and Marcano's solenodons have previously been grouped into their own genera, Atopogale and Antillogale , respectively. Family Soricidae includes 3 subfamilies and 26 genera. The oldest specimens of this family date back to the Eocene epoch in North America. Genetically, there is evidence that family Soricidae may have a close relationship with bats . Divisions within family Talpidae are still being debated, but currently include 3 subfamilies and 17 genera.

Physical Description

Shrews have a long, slim rostrum, small eyes, and short ear pinnae. Their five-toed feet are unspecialized, except for enlarged claws in semi-fossorial species and fringes of stiff hair on semi-aquatic species. Shrews have dilambdodont molars, with W-shaped ectolophs and unicuspid premolars. Their coat is typically gray or brown. Some species of shrews are venomous. In these species, their first lower incisors have a crude channel that carries the toxin from the venom-producing sub-maxillary glands, which open near the base of their lower incisors. There are three subfamilies within family Soricidae : Soricinae , Crocidurinae , and Myosoricinae . The vast majority of shrews are no bigger than 12.7 cm long, excluding the length of their tail, and weigh no more than 0.04 kg.

Family Talpidae includes moles and desmans. Most of these animals have modified heads and forelimbs for fossorial life. Their eyes are small and are often hidden beneath the skin. To make up for their lack of vision, these animals have a long, slender, nearly naked snout, with an extraordinary sensory system. Their ears lack pinnae and their fur is characteristically velvety and smooth. Their upper molars have W-shaped ectolophs, giving them dilambdodont dentition, which effectively pierces through an insect's exoskeleton. Their forelimbs are rotated such that their digits are pointed to the side, their palms face backward, and their elbows point upward. Their phalanges are short and their claws are long. Their peculiar forelimb orientation is a modification for fossorial life, which helps them efficiently “swim” through soil. Moles have tens of thousands of touch receptors located on their snout in the Eimer’s organ, which helps them forage along their tunnel systems. Moles adapted to aquatic environments, including desmans, have webbed feet and a fringe of stiff hair that helps them swims. Overall, fully grown moles range from about 6.1 to 43.2 cm in body length and weigh 0.2 to 3.5 kg.

Members of family Solenodontidae , commonly known as solenodons, have five-toed feet and a relatively long, nearly hairless tail. Their snout is long, slender, and highly flexible and extends beyond their lower jaw. Their eyes are small, and they have prominent pinnae. No auditory bullae are present and their zygomatic arch is incomplete. Their dental formula is 3/3, 1/1, 3/3, 3/3. Their second lower incisors have a deep groove that carries toxic saliva produced by glands under their lower incisors. Their upper molars have a V-shaped ectoloph, giving them zalambdodont dentition. They are generally brown, but occasionally they are black. Their undersides are lighter in comparison. Adult solenodons can reach body lengths of 60 cm; their tail may add another 15 to 20 cm. Their body weight ranges from 0.6 to 1 kg.

Most of the information regarding members of family Nesophontidae comes from skulls and skeletal remains, because only a few species survived into the 1900s, and all species are now thought to be extinct. The skulls of these animals lack zygomatic arches, jugal bones, or auditory bullae. These animals were characteristically similar to solenodons, with long, flexible snouts and small eyes.

• Other Physical Features
• endothermic endothermic
• homoiothermic
• bilateral symmetry bilateral symmetry
• venomous venomous

• Sexual Dimorphism
• male larger

Reproduction

Shrews breed during March and April. Males travel short to long distances to find areas with a higher density of females. To attract mates, shrews give off musky odors produced by their lateral glands, or make characteristic movements, such as tail-wagging. Species of shrews may have different mating systems. For instance, Asian lesser white-tooted shrews , common white toothed shrews , and Eurasian water shrews are all thought to be promiscuous, while, greater white-toothed shrews and bicolored shrews are thought to be monogamous. The division between monogamy and promiscuity in shrew species is thought to be related to their climate, shrews in temperate regions are often promiscuous, whereas highly seasonal shrew species are often monogamous. Moles mate from late February to early March. They produce one litter a year. Little is known about the mating systems of mole species; however, eastern moles and European moles are thought to maintain a polygynous mating system. Solenodons may be polygynous or promiscuous. They are capable of having 2 litters within a year; their litters are generally composed of 1 to 3 young. Males breed year round, whereas females go in and out of estrous, without a recognizable pattern. Thus, their breeding period is undefined. Among solenodons, courting behaviors include scent-marking and mutual sniffing. Since most families within this order are solitary, there is little effect on their social structure due to mating behaviors.

• Mating System
• monogamous monogamous
• polygynous polygynous
• polygynandrous (promiscuous) polygynandrous (promiscuous)

Shrews from northern temperate areas breed two or more times a year during April until September, although they are most reproductively active during the summer. Tropical species breed throughout the year, although they breed less frequently during the dry season. Females usually build nests made of leaves and grass in inconspicuous places, such as inside tunnels or under rocks. Their gestation period commonly lasts three to four weeks. Litter sizes among shrew species varies between three to seven young. Their offspring grow rapidly and leave their mother in three to four weeks. Moles have litters of about three to five offspring in early to mid-summer. Females build nests of leaves, grasses, and other plant fibers in well-drained soils. Their nests are usually built under objects, such as stumps or rocks to provide protection. Moles generally have about a 40-day gestation period. Their young are helpless and naked at birth, but mature quickly. Young leave the nest in four to six weeks and are sexually mature at ten months. Solenodons generally have two litters every year. They do not have a specific breeding season. They have about a 50-day gestation period and give birth to one to three offspring. Females nurse their young with their two mammae, which are located more caudally than is common among mammals. Young nurse for less than three months, but stay with their mother until her next litter. Soricomorphs can arrest blastocyst development when an extreme environmental condition is encountered. Each blastocyst floats freely in suspended animation in the reproductive tract until environmental conditions become favorable for implantation.

• Key Reproductive Features
• iteroparous iteroparous
• seasonal breeding seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual sexual
• viviparous viviparous
• delayed implantation delayed implantation

All soricomorphs provide very high fat, nutritious milk for their young through their mammae. Soricomorph males offer little, to no parental care. Nursing time is relatively short in this order, lasting about 3 months at most. During reproduction and lactation, females increase their food intake drastically. To produce high-calorie milk, females eat up to 3-times their normal intake when weaning young. When conditions are harsh, females draw on their own energy stores, which may lead to a loss in body mass. Due to their highly nutritious milk, postnatal growth is rapid. Some shrew species from subfamily Crocidurinae perform an unusual maternal behavior. When a female takes her offspring into the forest to explore and learn foraging skills, the mother leads the offspring in a row, with each young shrew using their teeth to grasp the hair on the rump of the shrew in front of them, this is known as a 'caravan'.

• Parental Investment
• altricial altricial
• female parental care female parental care
• pre-fertilization
  • protecting
    • female
• pre-hatching/birth
  • provisioning
    • female
  • protecting
    • female
• pre-weaning/fledging
  • provisioning
    • female
  • protecting
    • female
• pre-independence
  • provisioning
    • female
  • protecting
    • female

Lifespan/Longevity

Shrews usually live up to a year in the wild. There are reports of moles living to seven years in captivity. However, on average, moles live to about two years. Small mammals like shrews and moles have a rather short lifespan, which is attributed to their small body mass, high metabolic rates, rapid sexual maturity, relatively large litter size, and short gestation periods. On the other hand, solenodons live to an average of 6 years in captivity.

Behavior

Most shrews are nocturnal, however a few species forage during the day. Shrews are mainly insectivorous, but many species are functional omnivores. With their diminutive body mass and high surface to volume ratio they must forage often to avoid exhausting their energy stores. As a result, shrews are unable to hibernate. Due to their high respiratory water loss and small body size, shrews inhabit moist habitats, often near water. Unlike most mammals, some species of shrews produce venom, delivered through the groove in their lower incisors. Shrews are aggressive towards conspecifics and members of other species. Upon encountering another shrew, they often attack continuously, until one eventually dies. They typically avoid one another by marking small territories with distinct scents and thus avoid fights. Many species of shrews use high-frequency sounds for inter-specific communication, prey detection, and orientation.

Moles are largely fossorial, burrowing underground, creating large tunnels for living. Some species are mainly active at night, but others are active both during the day and night. Burrowing is generally done in moist soils. The tunnels they create are often visible above ground, and are known as 'mole runs'. These shallow tunnels are four to five cm in diameter and are primarily created when moles search for invertebrate prey. A 'molehill' is a rounded circular mound of dirt that is created when moles bring dirt up as they dig deep tunnels into the earth. Within these deep tunnels are complex branching networks of defended tunnels, which extend to 150 cm deep and include nest chambers. Deep chambers are also used to escape the cold. Since moles are mostly underground, where there is essentially no light, they are not visually oriented. Rather, they sense their environment through electroreception and chemoreception; their receptors are usually located in their snout area. Although some species of moles are social, most are solitary.

Like shrews and moles, solenodons are nocturnal. They are fossorial animals that shelter in caves, crevices, and under logs. Solenodons construct extensive networks of tunnels that reach depths greater than 20 cm below ground. These animals are omnivorous, but prefer animal matter, primarily preying on invertebrates. Due to the rocky, bushy, or forested areas they inhabit, solenodons move slowly in a zigzag path with a waddling gait. Solenodons have a highly developed sense of touch, smell, and hearing. Their vocalizations consist of grunts, shrieks, and clicks. Their clicks may be used as a form of echolocation to help them find prey. They may use echolocation not only to find food, but to help them visualize their environment. Solenodons produce toxic saliva and use this poison to immobilize their prey for consumption. Generally, these animals are solitary.

• Key Behaviors
• terricolous
• fossorial fossorial
• nocturnal nocturnal
• motile motile
• sedentary sedentary
• solitary solitary
• territorial territorial

Communication and Perception

Members of order Soricomorpha are solitary and only interact to mate or establish a home range. This solitary behavior makes their social communication insignificant. Moles have reduced eyes and use smell, sound, and mostly touch to perceive their environment. Eimer's organ, are very sensitive to touch, and are located on the rhinarium of some mole species. Their reduced eyes are the result of their underground lifestyle; they live in the dark and rarely surface. They use their naked snout to direct themselves. Some mole species utilize chemoreception through their nose to produce electrical impulses, which helps them understand their surroundings. Moles' fur does not lie flat on their bodies, because they are often backing up through tunnels, flat fur would be a hindrance. Shrews live above ground and have more acute vision, they also use their sense of smell and sound, and some species even use echolocation to perceive their environment. Similar to bats, they use a high frequency sound, but unlike bats, they do not use it to find prey. Shrews are also solitary, but do communicate when necessary, using tail wagging and foot drumming. Solenodons use vocal communication, such as shrieks, grunts, and clicks. These animals may also use a rudimentary form of echolocation. Both shrews and solenodons have whiskers on their snouts, which helps their depth perception, since their sight is poor. Whiskers and long bristly hairs are common among animals that don’t rely on sight. They are especially common around the face, but can appear other places based on the animal’s environment and need for extra sensing.

• Communication Channels
• visual visual
• tactile tactile
• chemical chemical

• Other Communication Modes
• scent marks scent marks
• vibrations vibrations

• Perception Channels
• visual visual
• tactile tactile
• acoustic acoustic
• echolocation echolocation
• vibrations vibrations
• chemical chemical

Food Habits

All soricomorphs are considered insectivorous. Moles primarily feed on larval insects, earthworms, snails, and other invertebrates, although they will eat almost anything that falls into their tunnels. Many mole species eat roots and other plant parts, and some aquatic species also eat fish and frogs. Moles have a much slower metabolic rate than shrews and eat about one-half of their body weight daily. Since shrews are very small, have short fur, and do not store fat well, they eat massive amounts of food to survive. Each day, they eat about 80 to 90% of their body weight; some species are able to eat up to four times that amount. They are generally omnivorous, but their diets mainly include insects, small vertebrates, fruits, and seeds. When food is scarce, shrews will eat lizards, frogs, small mammals, and even other shrews, especially during the winter months. During the night time, solenodons forage above ground, feeding on spiders, earthworms, insects, and other invertebrates. They obtain their food by rooting in the ground with their snouts and tearing through rotten logs. Their diet can also include some carrion, as well as small reptiles and amphibians.

• Primary Diet
• carnivore carnivore
  • eats terrestrial vertebrates
  • piscivore piscivore
  • insectivore insectivore
  • eats non-insect arthropods
  • molluscivore molluscivore
  • scavenger scavenger
• herbivore herbivore
  • frugivore frugivore
  • granivore granivore
• omnivore omnivore

Predation

Members of order Soricomorpha have many predators; however, their largest population stressors are habitat related. Solenodons are threatened by habitat loss, as well as predation by feral cats and dogs . They are also preyed upon by Asian mongooses , which were introduced to their habitat to control snake and rat populations. Shrews and moles have a variety of aerial predators, such as owls, including barn owls , tawny owls , barred owls , great horned owls , screech owls , and long-eared owls , bats such as greater false vampire bats , and raptors such as kestrels , red-tailed hawks , red-shouldered hawks , and broad-winged hawks . Shrews and moles are also occasionally preyed on by terrestrial mammals, however, their strong odor likely acts as a deterrent for many species. Domestic and feral cats for instance often kill shrews, but do not consume them, likely due to their odor. Shrews and moles may be eaten by ermines , European pine martens , European otters and other mustelids , as well as skunks and foxes . Snakes, such as reticulated pythons have also been known to feed on shrews. Anti-predatory adaptations such as mimicking, aposematic coloration, or cryptic behavior is not seen in order Soricomorpha , instead, moles hide in their burrows and shrews and solenodons are capable of sprinting at high speeds. Likewise, their small body size and dull coloration, helps shrews and solenodons blend into the forest floor, making them difficult to spot.

Ecosystem Roles

Moles dig extensive underground tunnels, which helps aerate the soil and helps plants grow. They also eat the larvae of some parasitic invertebrates that feed on crops and other garden plants. Their tunnels are also inhabited by other animals, such as shrews. Soricomorphs host fleas, which in turn can cause diseases that are harmful to other animals and even humans. Soricomorphs are not a major carrier of rabies or other harmful diseases. They are among the most successful mammalian orders and have become so independently of other creatures.

• Ecosystem Impact
• soil aeration soil aeration

Economic Importance for Humans: Positive

By constructing tunnels, moles help aerate the soil, which keeps the soil fertile and promotes plant growth. They also eat whatever drops into their tunnels, which often include larvae of harmful crop and garden pests. Moles do not eat plant roots, so they do not directly harm plants. Shrews eat a large amount of insects and may even prey on mice; however, they are not an effective pest control method because shrews themselves can become pests. Likewise, solenodons also consume a large quantity of invertebrate pests. Russian desmans were at one time hunted for their scent, which was used as a base for perfumes. Historically, mole fur has also been treasured for its use in fashion.

• Positive Impacts
• body parts are source of valuable material
• produces fertilizer
• controls pest population

Economic Importance for Humans: Negative

Moles do not eat plant roots, but their tunnels can be harmful to plants. They may dig up plant roots, which is especially bad for young plants. Their tunnels can also give other animals access to the roots. Their tunnels are considered unsightly and a disturbance to pastures, golf courses, and lawns. This can cause a serious problem for pastures, because mole tunnels reduce the amount of usable land for grazing and they encourage weed growth, which is not nutritious for grazers. These tunnels can also damage drainage systems and underground power lines and machinery. Shrews can become household pests by contaminating stored food and causing an unpleasant smell. They tend to use a single area for urinating and defecating, which makes the odor more intense. Shrews produce venom and some reports have suggested that their bites can be extremely painful, causing swelling and a burning sensation at the bite location for up to a week. Other reports have suggested that their bites do not impact humans. Otherwise, shrews are generally harmless. Solenodons are not a major pest, but they can damage gardens while digging for food. Solenodons are venomous, although they do not often bite, when provoked, their bites can cause discomfort. Their venom is not deadly and is usually only used to subdue prey.

• Negative Impacts
• injures humans
  • bites or stings
  • venomous venomous
• crop pest
• household pest

Conservation Status

According to the International Union for Conservation of Nature (IUCN), 28 species of shrews are critically endangered; 28 species are endangered; 55 species are vulnerable; and four are near threatened. Species that are critically endangered, endangered, or vulnerable are at a high risk of extinction. In other words, more than one-third of all shrew species are at some risk. The U.S. Fish and Wildlife Service has included Buena Vista Lake ornate shrews on the endangered list. Many at risk shrew species live in limited areas with very small populations. This puts them at a higher danger of extinction because a single act of human disruption or a natural disaster could annihilate the entire community. Among moles, the IUCN names two critically endangered species; five endangered species; and three vulnerable species, all of which are facing high risks of extinction in the wild. The U.S. Fish and Wildlife Service has not included any moles on the endangered list. Overall, one quarter of all mole species are at the risk of extinction. Like shrews, at risk moles live in small populations and areas that are disappearing due to human disturbance. In addition, introduced species are outcompeting native species in many areas. The two extant species of solenodons are endangered according to both the IUCN and the U.S. Fish and Wildlife Service. They too are at a very high risk of extinction in the wild. Scientists presume the cause of their decline includes habitat destruction and hunting by domestic dogs and cats.

Additional Links

Encyclopedia of Life

Contributors

Vincent Adam Patsy (author), University of Michigan-Ann Arbor, Jane Yaewon Song (author), University of Michigan-Ann Arbor, Robin Elizabeth Weber (author), University of Michigan-Ann Arbor, Leila Siciliano Martina (author, editor), Texas State University, Phil Myers (editor), University of Michigan-Ann Arbor.

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