By Allison Poor
Diversity
The family Spalacidae is a diverse old world group of fossorial and semi-fossorial rodents. This family consists of 36 species in 6 genera distributed among four subfamilies: the Myospalacinae, the Rhizomyinae, the Spalacinae, and the Tachyoryctinae. (Norris et al., 2004)
Geographic Range
Spalacids are old world rodents. They range from the Ukraine through the Balkans and the eastern Mediterranean, into Africa as far west as Libya and as far south as northern Tanzania, and in Asia from western China south to Sumatra and north to southern Siberia. (Corbert, 1984; Nowak, 1999; Vaughan, Ryan, and Czaplewski, 2000)
Habitat
Spalacids construct burrows in grasslands, scrublands, agricultural areas, and forests. They avoid deserts, preferring moist or semi-moist soils. Many inhabit mountainous regions, reaching elevations of up to 4,000 meters. (Corbert, 1984; Nowak, 1999)
These animals are found in the following types of habitat:
temperate
; tropical
; terrestrial
.
Terrestrial Biomes:
savanna or grassland
; chaparral
; forest
; scrub forest
; mountains
.
Other:
agricultural
; riparian
.
Systematic and Taxonomic History
Jansa and Weksler (2004) include a summary of the long, complicated taxonomic history of the family Spalacidae (as well as that of the rest of Muroidea). Alston (1876) designated Spalacidae as a family including the subfamilies Spalacinae and Bathyerginae, and classified the genus Myospalax seperately under the subfamily Siphneinae and the family Muridae. Thomas (1896) included two subfamilies under Spalacidae, Rhizomyinae and Spalacinae, and classified Myospalacinae as a subfamily of Muridae. Miller and Gidley (1918) placed the subfamilies Myospalacinae and Spalacinae in the family Spalacidae, separate from the Rhizomyidae, and Ellerman (1940, 1941) and Simpson (1945) proposed similar classifications, except that the former placed the Tachyoryctinae and the Myospalacinae within Muridae and the latter placed Myospalacini within Cricetidae. Chaline et al. (1977) did away with the family Spalacidae altogether, instead placing the subfamilies Spalacinae and Myospalacinae in the Cricetidae and the Tachyoryctoidinae and Rhizomyinae in the Rhizomyidae. Additionally, some authors have classified all the subfamilies of the current Spalacidae within the Muridae (Vaughan et al. 2000; Musser and Carleton 1993; Nowak 1991). However, recent molecular evidence has shown that the genera Rhizomys, Tachyoryctes, Myospalax, and Spalax form a monophyletic group sister to all other Muroidea, and hence these have been placed within a separate family, the Spalacidae (Jansa and Weksler 2004; Norris et al. 2004; Steppan et al. 2004). This classification actually follows that proposed by Tullberg (1899) over 100 years ago.
In recent years it has been suggested that the Myospalacinae are most closely related to the Cricetinae, based on a phylogenetic analysis using one Myospalax specimen (Michaux et al. 2001). However, other studies using multiple specimens have not supported this view, and instead place Myospalacinae basal to the other subfamilies within the Spalacidae (Jansa and Weksler 2004; Norris et al. 2004; Steppan et al. 2004). (Alston, 1876; Chaline, Mein, and Petter, 1977; Ellerman, 1940; Ellerman, 1941; Jansa and Weksler, 2004; Michaux, Reyes, and Catzeflis, 2001; Miller and Gidley, 1918; Musser and Carleton, 1993; Norris et al., 2004; Nowak, 1999; Simpson, 1945; Steppan, Adkins, and Anderson, 2004; Thomas, 1896; Tullberg, 1899; Vaughan, Ryan, and Czaplewski, 2000)
- loss or reduction of external eyes
- reduction of pinnae
- tail less than 50% of body length
- stocky body
- broad snout
- triangular braincase
- ovoid
infraorbital canal
- infraorbital canal not extending ventrally to roof of palate
- loss or reduction of
zygomatic plate
- infraorbital canal containing lacrimal canal
- small to medium sized
incisive foramina
- well-developed neck musculature with prominant attachment points on occipitum
- small reduction of third
molar relative to first and second molars - equal number of cusps on second and third molars
- distinct orientation of
manubrium of malleus
- Nucleotide characters in a suite of genes including: Lechitin cholesterol acetyltransferase (LCAT), von Willebrand’s factor (vWF), interphotoreceptor retinoid binding protein (IRBP), growth hormone receptor (GHR), breast cancer 1 (BRCA1), recombination activating gene 1 (RAG1), and the c-myc oncogene.
Physical Description
Spalacids are adapted for a fossorial or semifossorial lifestyle. They have stout, rounded, molelike bodies, reduced eyes and external ears, short, dense fur, and short limbs. Their heads are broad and they have powerful neck muscles. The wide
incisors project forward in front of the lips in all except for the myospalacines (which dig with their forearms instead of with their incisors) (Norris et al. 2004). Body sizes for this family range from the diminutive Spalax leucodon, measuring 130 mm in body length and weighing just 100 grams, to the hefty Rhizomys sumatrensis, measuring 480 mm and weighing up to to 4 kg. In some species, such as Tachyoryctes splendens, males are larger, and in others, there is no discernible sexual dimorphism. (Corbert, 1984; Flynn, 1990; Norris et al., 2004; Nowak, 1999; Vaughan, Ryan, and Czaplewski, 2000)
Some key physical features:
endothermic
; homoiothermic; bilateral symmetry
.
Sexual dimorphism:
sexes alike, male larger; sexes alike; male larger.
Reproduction
The only mating systems that have been reported for spalacids are polygyny and polygynandry. Males and females of most spalacid species only associate for a short time during courtship and mating. (Nowak, 1999)
Mating systems:
polygynous
; polygynandrous (promiscuous)
.
Spalacids usually have either one or two litters per year. Females of some species have a postpartum estrus, becoming pregnant again as soon as they give birth. Other females only have a single litter in their lifetime. The time of breeding varies between and within species, and depends on location. Gestation lasts between four and seven weeks, and anywhere from one to five young are born per litter. (Corbert, 1984; Nowak, 1999)
Key reproductive features:
semelparous
; iteroparous
; seasonal breeding
; year-round breeding
; gonochoric/gonochoristic/dioecious (sexes separate); sexual
; fertilization
(internal
); internal
; viviparous
; post-partum estrous.
Female spalacids constuct underground nests in which they give birth to altricial young. Males do not help care for their offspring. Females of most species nurse their young for four to six weeks, and the young leave the nest at two to three months. (Corbert, 1984; Nowak, 1999)
Parental investment:
altricial
; pre-fertilization (provisioning, protecting: female); provisioning; protecting (female); female; pre-hatching/birth (provisioning: female, protecting: female); provisioning (female); female; protecting (female); female; pre-weaning/fledging (provisioning: female, protecting: female); provisioning (female); female; protecting (female); female; pre-independence (protecting: female); protecting (female); female.
Lifespan/Longevity
Maximum longevity for species in this family is 4.5 years, but most probably do not live more than a year in the wild. (Nowak, 1999)
Behavior
Spalacids spend most of their lives underground, although some may come to the surface occasionally to forage. They are active year round, with timing of daily activity varying by species. These rodents construct elaborate tunnel systems, digging with their incisors (spalacines, rhizomyines and tachyoryctines) or foreclaws (rhizomyines, tachyoryctines and myospalacines) and using their hind feet or snout to push soil out behind them. Tunnel systems include well-defined areas for sleeping, foraging, and defecating. Spalacids are solitary, with each animal inhabiting its own burrow system and defending a territory that varies in size depending on the age, sex, and size of the individual. Generally, spalacids are sedentary, but a few species migrate seasonally when food becomes scarce. (Corbert, 1984; Flynn, 1990; Nevo, 1999; Norris et al., 2004; Nowak, 1999; Savic and Nevo, 1990; Vaughan, Ryan, and Czaplewski, 2000)
Key behaviors:
terricolous; fossorial
; diurnal
; nocturnal
; crepuscular
; motile
; migratory
; sedentary
; solitary
; territorial
.
Communication and Perception
Because they spend most of their time underground, spalacids do not have much use for vision, and therefore, their eyes are much reduced, although they remain functional and light sensitive in some. Their sense of touch is well-developed, and many have tactile hairs on the sides of their head. They also have good hearing and sense of smell. Pheromones and scent-marking are important means of communication in this group. Some species are known to communicate with each other by drumming their heads against the walls of their tunnels and sensing the vibrations created by others. Many are known to make grunting or hissing noises when threatened. (Nevo, 1999; Nowak, 1999)
Other communication keywords:
pheromones
; scent marks
; vibrations
.
Perception channels:
visual
; tactile
; acoustic
; vibrations
; chemical
.
Food Habits
The spalacid diet consists largely of roots, bulbs, rhizomes, and other underground plant parts. Shoots, leaves, seeds, fruit, insects, and other arthropods are eaten occasionally by some species. Many store large quantities of food in their underground burrow systems. (Corbert, 1984; Nowak, 1999; Vaughan, Ryan, and Czaplewski, 2000)
Primary Diet:
carnivore
(insectivore
, eats non-insect arthropods); insectivore
; eats non-insect arthropods; herbivore
(folivore
, frugivore
, granivore
); folivore
; frugivore
; granivore
; omnivore
.
Behaviors:
stores or caches food
.
Predation
- owls (Strigidae)
- snakes (Serpentes)
- eagles (Accipitridae)
- mammalian carnivores (Carnivora)
Spalacids sometimes fall prey to nocturnal hunters, such as owls, when they emerge above ground to forage. Other predators include snakes, eagles, and small mammalian carnivores. When confronted by a predator, spalacids may fight fearlessly, rushing the enemy and biting viciously with their formidable incisors. Their fossorial lifestyle may be their primary defense against predators. (Flynn, 1990; Nowak, 1999)
Ecosystem Roles
Spalacids are primary consumers, and they are a food source for a number of predators. Because of their extensive digging activity, spalacids affect the distribution of nutrients, air, and water in the soil, and therefore impact plant diversity. Also, other small animals sometimes shelter in their burrows. Finally, spalacids are parasitized by nematodes, ixodid ticks, gamasid mites, and fleas. (Flynn, 1990; Ganzorig et al., 1999; Litvinov and Sapegina, 2003; Zhang, Liu, and Du, 2004; Zhang, Zhang, and Liu, 2003)
Key ways these animals impact their ecosystem:
creates habitat; soil aeration
.
- nematodes Nematoda
- ixodid ticks Ixodidae
- gamasid mites Gamasida
- fleas Siphonaptera
Economic Importance for Humans: Negative
When present in agricultural areas, spalacids may feed on the roots of crops and cause considerable damage. (Nowak, 1999)
Ways that these animals might be a problem for humans:
crop pest.
Economic Importance for Humans: Positive
Many species of spalacids are eaten by native tribes, and the skin and bones of some are used as charms and for medicinal purposes, respectively. Also, some species of spalacids are important for medical research. (Chariyalertsak et al., 1997; Nowak, 1999; Zhang, Zhang, and Liu, 2003; Zhou, Zhou, and Zhang, 2004)
Ways that people benefit from these animals:
food
; body parts are source of valuable material; source of medicine or drug
; research and education.
Conservation
Of the 36 species in this family, 7 are listed as vulnerable (Chinese zokors, Eospalax fontanierii, sandy blind mole rats, Spalax arenarius, giant blind mole rats, Spalax giganteus, Balkan blind mole rats, Spalax graecus, greater blind mole rats, Spalax microphthalmus, lesser blind mole rats, Spalax leucodon, and big-headed mole rats, Tachyoryctes macrocephalus), 3 are listed as lower risk (three Myospalax species), and 2 are listed as data-deficient (two Tachyoryctes species) by the IUCN. (IUCN, 2004)
Contributors
Allison Poor (author), University of Michigan.
Tanya Dewey (editor), Animal Diversity Web, University of Michigan Museum of Zoology.

