Rodents Of Europe

Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are native to all major land masses except for New Zealand, Antarctica, and several oceanic islands, though they have subsequently been introduced to most of these land masses by human activity.

Rodents are extremely diverse in their ecology and lifestyles and can be found in almost every terrestrial habitat, including human-made environments. Species can be arboreal, fossorial (burrowing), saltatorial/richochetal (leaping on their hind legs), or semiaquatic. However, all rodents share several morphological features, including having only a single upper and lower pair of ever-growing incisors. Well-known rodents include mice, rats, squirrels, prairie dogs, porcupines, beavers, guinea pigs, and hamsters. Rabbits, hares, and pikas, whose incisors also grow continually (but have two pairs of upper incisors instead of one), were once included with them, but are now considered to be in a separate order, the Lagomorpha. Nonetheless, Rodentia and Lagomorpha are sister groups, sharing a single common ancestor and forming the clade of Glires.

Most rodents are small animals with robust bodies, short limbs, and long tails. They use their sharp incisors to gnaw food, excavate burrows, and defend themselves. Most eat seeds or other plant material, but some have more varied diets. They tend to be social animals and many species live in societies with complex ways of communicating with each other. Mating among rodents can vary from monogamy, to polygyny, to promiscuity. Many have litters of underdeveloped, altricial young, while others are precocial (relatively well developed) at birth.

The rodent fossil record dates back to the Paleocene on the supercontinent of Laurasia. Rodents greatly diversified in the Eocene, as they spread across continents, sometimes even crossing oceans. Rodents reached both South America and Madagascar from Africa and, until the arrival of ''Homo sapiens'', were the only terrestrial placental mammals to reach and colonize Australia.

Rodents have been used as food, for clothing, as pets, and as laboratory animals in research. Some species, in particular, the brown rat, the black rat, and the house mouse, are serious pests, eating and spoiling food stored by humans and spreading diseases. Accidentally introduced species of rodents are often considered to be invasive and have caused the extinction of numerous species, such as island birds, the dodo being an example, previously isolated from land-based predators.

Characteristics

The distinguishing feature of the rodents is their pairs of continuously growing, razor-sharp, open-rooted incisors. These incisors have thick layers of enamel on the front and little enamel on the back. Because they do not stop growing, the animal must continue to wear them down so that they do not reach and pierce the skull. As the incisors grind against each other, the softer dentine on the rear of the teeth wears away, leaving the sharp enamel edge shaped like the blade of a chisel. Most species have up to 22 teeth with no canines or anterior premolars. A gap, or diastema, occurs between the incisors and the cheek teeth in most species. This allows rodents to suck in their cheeks or lips to shield their mouth and throat from wood shavings and other inedible material, discarding this waste from the sides of their mouths. Chinchillas and guinea pigs have a high-fiber diet; their molars have no roots and grow continuously like their incisors.

In many species, the molars are relatively large, intricately structured, and highly cusped or ridged. Rodent molars are well equipped to grind food into small particles. The jaw musculature is strong. The lower jaw is thrust forward while gnawing and is pulled backwards during chewing. Gnawing uses incisors and chewing uses molars, however, due to the cranial anatomy of rodents these feeding methods cannot be used at the same time and are considered to be mutually exclusive. Among rodents, the masseter muscles plays a key role in chewing, making up 60% - 80% of the total muscle mass among masticatory muscles and reflects rodents herbivorous diet. Rodent groups differ in the arrangement of the jaw muscles and associated skull structures, both from other mammals and amongst themselves. The Sciuromorpha, such as the eastern grey squirrel, have a large deep masseter, making them efficient at biting with the incisors. The Myomorpha, such as the brown rat, have enlarged temporalis and masseter muscles, making them able to chew powerfully with their molars. In rodents, masseter muscles insert behind the eyes and contribute to eye boggling that occurs during gnawing where the quick contraction and relaxation of the muscle causes the eyeballs to move up and down. The Hystricomorpha, such as the guinea pig, have larger superficial masseter muscles and smaller deep masseter muscles than rats or squirrels, possibly making them less efficient at biting with the incisors, but their enlarged internal pterygoid muscles may allow them to move the jaw further sideways when chewing. The cheek pouch is a specific morphological feature used for storing food and is evident in particular subgroups of rodents like kangaroo rats, hamsters, chipmunks and gophers which have two bags that may range from the mouth to the front of the shoulders. True mice and rats do not contain this structure but their cheeks are elastic due to a high degree of musculature and innervation in the region.

While the largest species, the capybara, can weigh as much as , most rodents weigh less than . Rodents have wide-ranging morphologies, but typically have squat bodies and short limbs. The fore limbs usually have five digits, including an opposable thumb, while the hind limbs have three to five digits. The elbow gives the forearms great flexibility. The majority of species are plantigrade, walking on both the palms and soles of their feet, and have claw-like nails. The nails of burrowing species tend to be long and strong, while arboreal rodents have shorter, sharper nails. Rodent species use a wide variety of methods of locomotion including quadrupedal walking, running, burrowing, climbing, bipedal hopping (kangaroo rats and hopping mice), swimming and even gliding.
Scaly-tailed squirrels and flying squirrels, although not closely related, can both glide from tree to tree using parachute-like membranes that stretch from the fore to the hind limbs. The agouti is fleet-footed and antelope-like, being digitigrade and having hoof-like nails. The majority of rodents have tails, which can be of many shapes and sizes. Some tails are prehensile, as in the Eurasian harvest mouse, and the fur on the tails can vary from bushy to completely bald. The tail is sometimes used for communication, as when beavers slap their tails on the water surface or house mice rattle their tails to indicate alarm. Some species have vestigial tails or no tails at all. In some species, the tail is capable of regeneration if a part is broken off.

Rodents generally have well-developed senses of smell, hearing, and vision. Nocturnal species often have enlarged eyes and some are sensitive to ultraviolet light. Many species have long, sensitive whiskers or vibrissae for touch or "whisking". Whisker action is mostly driven by the brain stem, which is itself provoked by the cortex. However Legg et al. 1989 find an alternate circuit between the cortex and whiskers through the cerebellar circuits, and Hemelt & Keller 2008 the superior colliculus. Some rodents have cheek pouches, which may be lined with fur. These can be turned inside out for cleaning. In many species, the tongue cannot reach past the incisors. Rodents have efficient digestive systems, absorbing nearly 80% of ingested energy. When eating cellulose, the food is softened in the stomach and passed to the cecum, where bacteria reduce it to its carbohydrate elements. The rodent then practices coprophagy, eating its own fecal pellets, so the nutrients can be absorbed by the gut. Rodents therefore often produce a hard and dry fecal pellet. Horn et al. 2013 makes the finding that rodents entirely lack the ability to vomit. In many species, the penis contains a bone, the baculum; the testes can be located either abdominally or at the groin.

Sexual dimorphism occurs in many rodent species. In some rodents, males are larger than females, while in others the reverse is true. Male-bias sexual dimorphism is typical for ground squirrels, kangaroo rats, solitary mole rats and pocket gophers; it likely developed due to sexual selection and greater male-male combat. Female-bias sexual dimorphism exists among chipmunks and jumping mice. It is not understood why this pattern occurs, but in the case of yellow-pine chipmunks, males may have selected larger females due to their greater reproductive success. In some species, such as voles, sexual dimorphism can vary from population to population. In bank voles, females are typically larger than males, but male-bias sexual dimorphism occurs in alpine populations, possibly because of the lack of predators and greater competition between males.

Distribution and habitat

One of the most widespread groups of mammals, rodents can be found on every continent except Antarctica. They are the only terrestrial placental mammals to have colonized Australia and New Guinea without human intervention. Humans have also allowed the animals to spread to many remote oceanic islands (e.g., the Polynesian rat). Rodents have adapted to almost every terrestrial habitat, from cold tundra (where they can live under snow) to hot deserts.

Some species such as tree squirrels and New World porcupines are arboreal, while some, such as gophers, tuco-tucos, and mole rats, live almost completely underground, where they build complex burrow systems. Others dwell on the surface of the ground, but may have a burrow into which they can retreat. Beavers and muskrats are known for being semiaquatic, but the rodent best-adapted for aquatic life is probably the earless water rat from New Guinea. Rodents have also thrived in human-created environments such as agricultural and urban areas.

Though some species are common pests for humans, rodents also play important ecological roles. Some rodents are considered keystone species and ecosystem engineers in their respective habitats. In the Great Plains of North America, the burrowing activities of prairie dogs play important roles in soil aeration and nutrient redistribution, raising the organic content of the soil and increasing the absorption of water. They maintain these grassland habitats, and some large herbivores such as bison and pronghorn prefer to graze near prairie dog colonies due to the increased nutritional quality of forage.

Extirpation of prairie dogs can also contribute to regional and local biodiversity loss, increased seed depredation, and the establishment and spread of invasive shrubs. Burrowing rodents may eat the fruiting bodies of fungi and spread spores through their feces, thereby allowing the fungi to disperse and form symbiotic relationships with the roots of plants (which usually cannot thrive without them). As such, these rodents may play a role in maintaining healthy forests.

In many temperate regions, beavers play an essential hydrological role. When building their dams and lodges, beavers alter the paths of streams and rivers and allow for the creation of extensive wetland habitats. One study found that engineering by beavers leads to a 33 percent increase in the number of herbaceous plant species in riparian areas. Another study found that beavers increase wild salmon populations. Meanwhile, some rodents are seen as pests, due to their wide range.

Behavior and life history

Feeding

Most rodents are herbivorous, feeding exclusively on plant material such as seeds, stems, leaves, flowers, and roots. Some are omnivorous and a few are predators. The field vole is a typical herbivorous rodent and feeds on grasses, herbs, root tubers, moss, and other vegetation, and gnaws on bark during the winter. It occasionally eats invertebrates such as insect larvae. The plains pocket gopher eats plant material found underground during tunneling, and also collects grasses, roots, and tubers in its cheek pouches and caches them in underground larder chambers.

The Texas pocket gopher avoids emerging onto the surface to feed by seizing the roots of plants with its jaws and pulling them downwards into its burrow. It also practices coprophagy. The African pouched rat forages on the surface, gathering anything that might be edible into its capacious cheek pouches until its face bulges out sideways. It then returns to its burrow to sort through the material it has gathered and eats the nutritious items.

Agouti species are one of the few animal groups that can break open the large capsules of the Brazil nut fruit. Too many seeds are inside to be consumed in one meal, so the agouti carries some off and caches them. This helps dispersal of the seeds as any that the agouti fails to retrieve are distant from the parent tree when they germinate. Other nut-bearing trees tend to bear a glut of fruits in the autumn. These are too numerous to be eaten in one meal and squirrels gather and store the surplus in crevices and hollow trees. In desert regions, seeds are often available only for short periods. The kangaroo rat collects all it can find and stores them in larder chambers in its burrow.

A strategy for dealing with seasonal plenty is to eat as much as possible and store the surplus nutrients as fat. Marmots do this, and may be 50% heavier in the autumn than in the spring. They rely on their fat reserves during their long winter hibernation. Beavers feed on the leaves, buds, and inner bark of growing trees, as well as aquatic plants. They store food for winter use by felling small trees and leafy branches in the autumn and immersing them in their pond, sticking the ends into the mud to anchor them. Here, they can access their food supply underwater even when their pond is frozen over.

Although rodents have been regarded traditionally as herbivores, most small rodents opportunistically include insects, worms, fungi, fish, or meat in their diets and a few have become specialized to rely on a diet of animal matter. A functional-morphological study of the rodent tooth system supports the idea that primitive rodents were omnivores rather than herbivores. Studies of the literature show that numerous members of the Sciuromorpha and Myomorpha, and a few members of the Hystricomorpha, have either included animal matter in their diets or been prepared to eat such food when offered it in captivity. Examination of the stomach contents of the North American white-footed mouse, normally considered to be herbivorous, showed 34% animal matter.

More specialized carnivores include the shrewlike rats of the Philippines, which feed on insects and soft-bodied invertebrates, and the Australian water rat, which devours aquatic insects, fish, crustaceans, mussels, snails, frogs, birds' eggs, and water birds. The grasshopper mouse from dry regions of North America feeds on insects, scorpions, and other small mice, and only a small part of its diet is plant material. It has a chunky body with short legs and tail, but is agile and can easily overpower prey as large as itself.

Social behavior

Rodents exhibit a wide range of types of social behavior ranging from the mammalian caste system of the naked mole-rat, the extensive "town" of the colonial prairie dog, through family groups to the independent, solitary life of the edible dormouse. Adult dormice may have overlapping feeding ranges, but they live in individual nests and feed separately, coming together briefly in the breeding season to mate. The pocket gopher is also a solitary animal outside the breeding season, each individual digging a complex tunnel system and maintaining a territory.

Larger rodents tend to live in family units where parents and their offspring live together until the young disperse. Beavers live in extended family units typically with a pair of adults, this year's kits, the previous year's offspring, and sometimes older young. Brown rats usually live in small colonies with up to six females sharing a burrow and one male defending a territory around the burrow. At high population densities, this system breaks down and males show a hierarchical system of dominance with overlapping ranges. Female offspring remain in the colony while male young disperse. The prairie vole is monogamous and forms a lifelong pair bond. Outside the breeding season, prairie voles live with others in small colonies. A male is not aggressive towards other males until he has mated, after which time he defends a territory, a female, and a nest against other males. The pair huddles together, grooms one another, and shares nesting and pup-raising responsibilities.

Among the most social of rodents are the ground squirrels, which typically form colonies based on female kinship, with males dispersing after weaning and becoming nomadic as adults. Cooperation in ground squirrels varies between species and typically includes making alarm calls, defending territories, sharing food, protecting nesting areas, and preventing infanticide. The black-tailed prairie dog forms large towns that may cover many hectares. The burrows do not interconnect, but are excavated and occupied by territorial family groups known as coteries. A coterie often consists of an adult male, three or four adult females, several nonbreeding yearlings, and the current year's offspring. Individuals within coteries are friendly with each other, but hostile towards outsiders.

Perhaps the most extreme examples of colonial behavior in rodents are the eusocial naked mole rat and Damaraland mole rat. The naked mole rat lives completely underground and can form colonies of up to 80 individuals. Only one female and up to three males in the colony reproduce, while the rest of the members are smaller and sterile, and function as workers. Some individuals are of intermediate size. They help with the rearing of the young and can take the place of a reproductive if one dies. The Damaraland mole rat is characterized by having a single reproductively active male and female in a colony where the remaining animals are not truly sterile, but become fertile only if they establish a colony of their own.

Communication

Olfactory

Rodents use scent marking in many social contexts including inter- and intra-species communication, the marking of trails and the establishment of territories. Their urine provides genetic information about individuals including the species, the sex and individual identity, and metabolic information on dominance, reproductive status and health. Compounds derived from the major histocompatibility complex (MHC) are bound to several urinary proteins. The odor of a predator depresses scent-marking behavior.

Rodents are able to recognize close relatives by smell and this allows them to show nepotism (preferential behavior toward their kin) and also avoid inbreeding. This kin recognition is by olfactory cues from urine, feces and glandular secretions. The main assessment may involve the MHC, where the degree of relatedness of two individuals is correlated to the MHC genes they have in common. In non-kin communication, where more permanent odor markers are required, as at territorial borders, then non-volatile major urinary proteins (MUPs), which function as pheromone transporters, may also be used. MUPs may also signal individual identity, with each male house mouse (''Mus musculus'') excreting urine containing about a dozen genetically encoded MUPs.

House mice deposit urine, which contains pheromones, for territorial marking, individual and group recognition, and social organization. Territorial beavers and red squirrels investigate and become familiar with the scents of their neighbors and respond less aggressively to intrusions by them than to those made by non-territorial "floaters" or strangers. This is known as the "dear enemy effect".

Auditory

Many rodent species, particularly those that are diurnal and social, have a wide range of alarm calls that are emitted when they perceive threats. There are both direct and indirect benefits of doing this. A potential predator may stop when it knows it has been detected, or an alarm call can allow conspecifics or related individuals to take evasive action. Several species, for example prairie dogs, have complex anti-predator alarm call systems. These species may have different calls for different predators (e.g. aerial predators or ground-based predators) and each call contains information about the nature of the precise threat. The urgency of the threat is also conveyed by the acoustic properties of the call.

Social rodents have a wider range of vocalizations than do solitary species. Fifteen different call-types have been recognized in adult Kataba mole rats and four in juveniles. Similarly, the common degu, another social, burrowing rodent, exhibits a wide array of communication methods and has an elaborate vocal repertoire comprising fifteen different categories of sound. Ultrasonic calls play a part in social communication between dormice and are used when the individuals are out of sight of each other.

House mice use both audible and ultrasonic calls in a variety of contexts. Audible vocalizations can often be heard during agonistic or aggressive encounters, whereas ultrasound is used in sexual communication and also by pups when they have fallen out of the nest.

Laboratory rats (which are brown rats, ''Rattus norvegicus'') emit short, high frequency, ultrasonic vocalizations during purportedly pleasurable experiences such as rough-and-tumble play, when anticipating routine doses of morphine, during mating, and when tickled. The vocalization, described as a distinct "chirping", has been likened to laughter, and is interpreted as an expectation of something rewarding. In clinical studies, the chirping is associated with positive emotional feelings, and social bonding occurs with the tickler, resulting in the rats becoming conditioned to seek the tickling. However, as the rats age, the tendency to chirp declines. Like most rat vocalizations, the chirping is at frequencies too high for humans to hear without special equipment, so bat detectors have been used for this purpose.

Visual

Rodents, like all placental mammals except primates, have just two types of light receptive cones in their retina, a short wavelength "blue-UV" type and a middle wavelength "green" type. They are therefore classified as dichromats; however, they are visually sensitive into the ultraviolet (UV) spectrum and therefore can see light that humans can not. The functions of this UV sensitivity are not always clear. In degus, for example, the belly reflects more UV light than the back. Therefore, when a degu stands up on its hind legs, which it does when alarmed, it exposes its belly to other degus and ultraviolet vision may serve a purpose in communicating the alarm. When it stands on all fours, its low UV-reflectance back could help make the degu less visible to predators. Ultraviolet light is abundant during the day but not at night. There is a large increase in the ratio of ultraviolet to visible light in the morning and evening twilight hours. Many rodents are active during twilight hours (crepuscular activity), and UV-sensitivity would be advantageous at these times. Ultraviolet reflectivity is of dubious value for nocturnal rodents.

The urine of many rodents (e.g. voles, degus, mice, rats) strongly reflects UV light and this may be used in communication by leaving visible as well as olfactory markings. However, the amount of UV that is reflected decreases with time, which in some circumstances can be disadvantageous; the common kestrel can distinguish between old and fresh rodent trails and has greater success hunting over more recently marked routes.

Tactile

Vibrations can provide cues to conspecifics about specific behaviors being performed, predator warning and avoidance, herd or group maintenance, and courtship. The Middle East blind mole rat was the first mammal for which seismic communication was documented. These fossorial rodents bang their head against the walls of their tunnels. This behavior was initially interpreted as part of their tunnel building behavior, but it was eventually realized that they generate temporally patterned seismic signals for long-distance communication with neighboring mole rats.

Footdrumming is used widely as a predator warning or defensive action. It is used primarily by fossorial or semi-fossorial rodents. The banner-tailed kangaroo rat produces several complex footdrumming patterns in a number of different contexts, one of which is when it encounters a snake. The footdrumming may alert nearby offspring but most likely conveys that the rat is too alert for a successful attack, thus preventing the snake's predatory pursuit. Several studies have indicated intentional use of ground vibrations as a means of intra-specific communication during courtship among the Cape mole rat. Footdrumming has been reported to be involved in male-male competition; the dominant male indicates its resource holding potential by drumming, thus minimizing physical contact with potential rivals.

Mating strategies

Some species of rodent are monogamous, with an adult male and female forming a lasting pair bond. Monogamy can come in two forms; obligate and facultative. In obligate monogamy, both parents care for the offspring and play an important part in their survival. This occurs in species such as California mice, oldfield mice, Malagasy giant rats and beavers. In these species, males usually mate only with their partners. In addition to increased care for young, obligate monogamy can also be beneficial to the adult male as it decreases the chances of never finding a mate or mating with an infertile female. In facultative monogamy, the males do not provide direct parental care and stay with one female because they cannot access others due to being spatially dispersed. Prairie voles appear to be an example of this form of monogamy, with males guarding and defending females within their vicinity.

In polygynous species, males will try to monopolize and mate with multiple females. As with monogamy, polygyny in rodents can come in two forms; defense and non-defense. Defense polygyny involves males controlling territories that contain resources that attract females. This occurs in ground squirrels like yellow-bellied marmots, California ground squirrels, Columbian ground squirrels and Richardson's ground squirrels. Males with territories are known as "resident" males and the females that live within the territories are known as "resident" females. In the case of marmots, resident males do not appear to ever lose their territories and always win encounters with invading males. Some species are also known to directly defend their resident females and the ensuing fights can lead to severe wounding. In species with non-defense polygyny, males are not territorial and wander widely in search of females to monopolize. These males establish dominance hierarchies, with the high-ranking males having access to the most females. This occurs in species like Belding's ground squirrels and some tree squirrel species.
thumbnail, A mating plug in a female Richardson's ground squirrel
Promiscuity, in which both males and females mate with multiple partners, also occurs in rodents. In species such as the white-footed mouse, females give birth to litters with multiple paternities. Promiscuity leads to increased sperm competition and males tend to have larger testicles. In the Cape ground squirrel, the male's testes can be 20 percent of its head-body length. Several rodent spe