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Muroidea

Muroidea is a large superfamily of rodents within the suborder Myomorpha of the order Rodentia, encompassing over 1,700 species distributed across six families: Cricetidae (including hamsters, voles, and New World rats and mice), Muridae (Old World rats, mice, and gerbils), Nesomyidae (African and Malagasy rodents), Spalacidae (mole rats and bamboo rats), Calomyscidae (mouse-like hamsters), and Platacanthomyidae (spiny dormice).^[1]^[2] These rodents are defined by distinctive morphological traits, such as a keyhole-shaped infraorbital foramen in the skull, a broad zygomatic plate, and a sciurognathous (squirrel-like) lower jaw articulation, which support their monophyly as confirmed by paleontological, morphological, and molecular evidence.^[1]^[3] Muroidea represents more than 25% of all mammal species and occupies diverse habitats—from forests and deserts to urban environments—on every continent except Antarctica, often introduced by humans to new regions.^[1] The superfamily's evolutionary origins trace back to the middle Eocene in Asia, with fossils appearing in Europe by the early Oligocene and in Africa by the early Miocene, reflecting adaptive radiations that have led to its exceptional diversity.^[3] Muroids exhibit a wide range of body sizes, from the tiny African pygmy mouse (Mus minutoides) at about 3-5 grams to over 2 kg in species like the Flores giant rat (Papagomys armandvillei).^[1]^[4] Their dentition features ever-growing incisors for gnawing and high-crowned molars adapted for grinding varied diets, including seeds, insects, and vegetation, enabling ecological versatility.^[5] Behaviorally, muroids display polygynandrous mating systems, high reproductive rates (with some species producing multiple litters per year), and flexible activity patterns, from nocturnal foraging to diurnal burrowing, contributing to their success as both commensal species and ecosystem engineers.^[1] Muroidea holds significant ecological, economic, and scientific importance; many species, such as the house mouse (Mus musculus) and Norway rat (Rattus norvegicus), are common pests that impact agriculture and human health by vectoring diseases, while others serve as key prey for predators and indicators of environmental health.^[5] In research, muroids like laboratory mice and rats have been instrumental in biomedical studies due to their genetic tractability and physiological similarities to humans, with Muridae comprising the majority of rodent species used in experiments.^[5] Many species face conservation challenges, with a significant proportion listed as threatened on the IUCN Red List, driven by habitat loss and invasive introductions, underscoring the need for targeted protection of this diverse clade.^[6]

Characteristics

Morphology and physiology

Muroid rodents exhibit a range of shared morphological traits that reflect their adaptability across diverse habitats, though with considerable variation within the superfamily. Most species are small to medium in size, with head-body lengths typically ranging from 5 to 30 cm and weights from a few grams to over 2 kg, as seen in extremes like the pygmy mouse (Baiomys) and the large cloud rat (Phloeomys).^[7] They generally possess elongated snouts that house enlarged nasal cavities, supporting a dominant olfactory system essential for navigation and communication.^[8] Prominent, ever-growing incisors—yellowish due to iron deposits for strength—enable efficient gnawing of tough vegetation and materials, a hallmark of rodent dentition.^[9] Tails are often furred for balance and thermoregulation but can be scaly in some arid-adapted forms, and while clavicles are present, they are reduced relative to other mammals, providing shoulder girdle flexibility for burrowing and maneuvering in confined spaces.^[10] Physiologically, muroids are characterized by high basal metabolic rates, often exceeding those of similar-sized mammals, which support their predominantly nocturnal lifestyles and facilitate rapid reproductive cycles with short generation times.^[11] This elevated metabolism correlates with efficient energy processing, including adaptations in the digestive system such as a semiglandular stomach divided into a nonglandular forestomach for microbial fermentation and a glandular hindstomach for acid secretion, aiding the breakdown of fibrous diets.^[12] Some taxa, notably hamsters in the Cricetinae, possess internal, fur-lined cheek pouches that extend to the shoulders, allowing temporary storage and transport of seeds and food without impeding locomotion.^[13] The dental formula is characteristically 1/1:0/0:0/0–3:0–3:3/3, featuring a single pair of upper and lower incisors per jaw, absent canines and premolars, and up to three molars per quadrant; these molars are typically hypsodont (ever-growing) with abrasive crowns suited to gritty, plant-based diets.^[14] Morphological variations across muroid clades highlight evolutionary divergences tied to locomotor modes. Arboreal species, such as certain oryzomyine rats (e.g., in the genus Mindomys), often feature long, semi-prehensile tails covered in tactile hairs that aid in grasping branches and balance during climbing.^[15] In contrast, fossorial forms like blind mole-rats (Spalacinae) display specialized burrowing adaptations, including reduced eyes and external ears, shortened limbs with robust foreclaws, and a cylindrical body for efficient soil displacement.^[16] Sensory systems in muroids emphasize olfaction and hearing over vision in many cases, reflecting their crepuscular or subterranean tendencies. A large olfactory bulb and extensive nasal turbinates provide acute smell for detecting pheromones and food, dominating sensory input.^[8] Hearing is highly developed, with sensitivity to ultrasonic frequencies for echolocation-like communication in some species, while vision varies: diurnal or open-habitat forms have forward-facing eyes for binocular depth perception, but subterranean taxa exhibit degenerated visual systems with minimal eye development.^[17]

Reproduction and development

Muroid rodents are characterized by high reproductive output, which supports their rapid population dynamics and adaptability to variable environments. Most species are polyestrous, capable of producing multiple litters annually, often 3 to 5 or more under optimal conditions, allowing for continuous breeding when resources are available.^[18] Gestation periods are generally short, ranging from 18 to 30 days across the superfamily; for example, house mice (Mus musculus) have gestations of 19–21 days, while voles (Microtus spp.) average 21 days and hamsters (Phodopus spp.) around 16–20 days.^[19]^[20]^[21] Litter sizes are typically large to maximize survival chances, averaging 4–12 young per litter, though this varies by subfamily and social structure—solitary species like some mice may produce up to 10–12 pups, while more social voles average 3–6.^[18]^[19]^[20] The young are predominantly altricial, born blind, hairless, and helpless, requiring intensive maternal care, though some species in subfamilies like Gerbillinae exhibit slightly more precocial traits with fur at birth.^[1] Mating systems within Muroidea display considerable diversity, reflecting adaptations to ecological pressures. Promiscuity or polygynandry predominates in many taxa, such as the house mouse (Mus musculus), where both males and females mate with multiple partners to increase genetic diversity and reproductive success.^[22] In contrast, monogamy with long-term pair bonds occurs in certain voles, notably the prairie vole (Microtus ochrogaster), where males form stable partnerships that enhance offspring survival through biparental care.^[23] Ovulation mechanisms also vary; while most muroids are spontaneous ovulators, some, like golden hamsters (Mesocricetus auratus), exhibit induced ovulation triggered by copulation, synchronizing reproduction with mating opportunities.^[24] This high fecundity is physiologically supported by efficient metabolic rates that allocate energy toward gamete production and lactation, enabling quick recovery between litters.^[18] Developmental stages in muroids are accelerated to facilitate early independence and further reproduction. Pups grow rapidly, with eyes opening at 10–14 days and weaning occurring around 21 days in species like mice and voles; sexual maturity is typically reached within 1–3 months, allowing females to breed in their first year.^[19]^[20] Parental care is primarily maternal, involving nest building, nursing, and grooming, but varies by species—males provide substantial assistance in some hamsters, such as the Djungarian hamster (Phodopus campbelli), where fathers help clean pups, retrieve strays, and even assist during birth.^[21] In high-density populations, behaviors like infanticide emerge as a density-dependent regulation mechanism; for instance, in crowded house mouse colonies, unrelated adults may kill pups to redirect resources toward their own offspring, reducing competition.^[25] These traits collectively contribute to the superfamily's remarkable reproductive flexibility and population resilience.

Distribution and ecology

Geographic distribution

Muroidea originated in Eurasia during the Eocene epoch, with the earliest fossil records appearing in Asia during the late Eocene to early Oligocene around 35-30 million years ago, marking the initial diversification of the superfamily within this continent.^[2] Subsequent radiations expanded their presence to Africa via early Miocene migrations across Afro-Arabian land connections, while further dispersals within Asia occurred through continental corridors. To the Americas, ancestral lineages, particularly within Cricetidae, migrated overland from Asia via Beringian land bridges between approximately 10 and 5 million years ago, facilitating the establishment of diverse cricetid groups in North America before further southward expansion.^[26]^[27] Today, muroids exhibit a near-global distribution, inhabiting all continents except Antarctica and avoiding extreme polar regions, with the highest abundances in temperate and tropical zones across Eurasia, Africa, and the Americas. They dominate rodent faunas in these areas, comprising about 26% of global mammalian diversity through adaptive radiations in varied ecosystems.^[28] Invasive species, such as the black rat (Rattus rattus), have further extended ranges to Oceania and remote oceanic islands, where they were introduced by human activities, often disrupting native biotas.^[29]^[30]^[31] Dispersal patterns of muroids reflect a combination of natural and anthropogenic processes, including overland migrations that enabled continental colonizations, such as the Cricetidae's Beringian crossing, and island colonizations like the Nesomyidae's arrival in Madagascar from eastern Africa during the early Miocene. Human-mediated introductions, particularly of Rattus rattus following European seafaring expansions after the 14th century, propelled the black rat to a worldwide distribution, including ports, islands, and even isolated archipelagos in the Pacific. These patterns underscore muroids' remarkable dispersal capabilities, driven by ecological opportunism and human vectors.^[32]^[33]^[34] Biogeographic hotspots for muroid diversity include Southeast Asia, where Muridae exhibit high endemism due to insular radiations in the Philippines, Sulawesi, and surrounding islands, and Madagascar, home to endemic Nesomyidae representing ancient vicariant divergences. These regions harbor significant portions of muroid species in localized assemblages, highlighting centers of evolutionary innovation tied to tectonic isolation and climatic stability.^[35]^[36]

Habitat and behavior

Muroid rodents exhibit remarkable habitat versatility, inhabiting a wide array of environments from arid deserts and grasslands to temperate forests and urban settings. Fossorial species like gerbils (Gerbillinae) thrive in desert ecosystems, where they construct extensive burrow systems to escape daytime heat and predators, often preferring sandy or loose soils that facilitate digging. In contrast, grassland dwellers such as voles (Arvicolinae) favor open fields with dense vegetation cover for foraging and nesting, while some arboreal members of the Murinae, such as certain tree rats, adapt to forested canopies using climbing adaptations to access fruits and insects. This ecological flexibility allows muroids to exploit diverse niches, including semiaquatic habitats near rivers and lakes for species like water voles.^[1]^[37]^[38] Behaviorally, muroids display varied diets ranging from herbivorous (seeds and vegetation in voles and hamsters) to omnivorous or insectivorous patterns, with many species engaging in food hoarding to buffer against scarcity; for instance, hamsters (Cricetinae) scatter-hoard seeds in cheek pouches for later retrieval, a strategy enhanced under food deprivation. Activity patterns are predominantly nocturnal or crepuscular to minimize predation risk, though some diurnal exceptions exist in safer environments. Social structures vary widely, from solitary lifestyles in rats (Rattus spp.), which defend territories via aggression, to highly colonial groups in lemmings and voles, where communal nesting and cooperative defense promote group cohesion in open habitats. Communication relies on scent marking for territory delineation and ultrasonic vocalizations for alarm signals or mate attraction, supplemented by tactile and vibrational cues in social interactions.^[1]^[39]^[38] Foraging behaviors emphasize efficiency and risk avoidance, with muroids using agility, camouflage, or rapid burrowing to evade predators like owls and snakes; hoarding not only secures resources but also aids in predator deterrence by reducing surface exposure time. Defense mechanisms include tail autotomy in some species and aposematic coloration in others, while group living in colonial forms enhances vigilance through collective alarm calls. Ecologically, muroids serve as key prey for numerous predators, contributing to food web dynamics, and play vital roles in seed dispersal—such as roof rats (Rattus rattus) transporting nonnative seeds via frugivory—and soil aeration through burrowing, which improves nutrient cycling. However, invasive muroids like house mice (Mus musculus) and Norway rats (Rattus norvegicus) cause significant agricultural damage by consuming crops such as grains and sugarcane, leading to economic losses and altered native ecosystems.^[1]^[40]

Evolution

Phylogenetic relationships

Muroidea forms the sister group to Dipodoidea within the suborder Myomorpha, a relationship strongly supported by molecular phylogenies based on nuclear and mitochondrial genes.^[41] The superfamily exhibits a basal divergence from the family Platacanthomyidae, which represents the earliest split among extant muroid lineages and is estimated to have occurred around 45 million years ago in the Eocene.^[29] Subsequent to this basal split, Muroidea diversified into major clades including Spalacidae, Calomyscidae, Nesomyidae, Cricetidae, and Muridae (which encompasses subfamilies such as Deomyinae, Gerbillinae, and Murinae).^[29] These relationships were first robustly resolved using sequences from the interphotoreceptor retinoid-binding protein (IRBP) gene, highlighting the monophyly of these groups despite some early polytomies in shallower branches.^[42] Uncertainties persist in the exact relationships among Nesomyidae, Cricetidae, and Muridae due to rapid early radiations. The evolutionary history of Muroidea is marked by a rapid radiation during the late Eocene to Oligocene, approximately 40–30 million years ago, coinciding with global cooling and habitat fragmentation that facilitated adaptive diversification.^[43] This period saw the emergence of key lineages, with diversification rates accelerating further in the Miocene through episodic bursts driven by ecological opportunities in expanding grasslands and forests.^[44] For instance, a comprehensive phylogeny encompassing 900 species revealed increasing net diversification rates across muroid clades, particularly within Murinae and Sigmodontinae during the early to middle Miocene around 20 million years ago, underscoring a pattern of density-dependent slowdown following initial radiations.^[29] Recent advances have refined these timelines using integrated datasets. A Bayesian tip-dated analysis incorporating fossil calibrations estimated the crown age of Muroidea at approximately 40 million years ago, with major biogeographic events like the African radiation of Nesomyinae occurring in the Oligocene.^[2] Complementing this, mitogenomic studies from 2025 have clarified relationships within Murinae, resolving previously ambiguous tribal affiliations and highlighting rapid diversification in African lineages such as Deomyinae, supported by complete mitochondrial genomes that reveal distinct geographic clades.^[45] These phylogenies rely on concatenated nuclear (e.g., BRCA1, RAG1) and mitochondrial DNA sequences, calibrated with fossil priors, though some unresolved polytomies persist in early branches due to rapid successive speciations.

Fossil history

The fossil record of Muroidea traces back to the Middle Eocene in Asia, where early stem representatives exhibit primitive myomorphous jaw mechanics indicative of the group's divergence from other rodents. Fossils such as Pappocricetodon antiquus from middle Eocene deposits in China, dating to approximately 45–40 million years ago (mya), represent some of the oldest known members, characterized by cricetid-like dental patterns and zygomasseteric structures transitional to fully myomorphous forms. By the late Eocene, further Asian specimens, including those from Kazakhstan and China, show a clearer shift toward the defining myomorph jaw configuration, marking an important evolutionary step in masticatory adaptation.^[46]^[47]^[48] Major extinct groups related to Muroidea include the Eomyidae, considered basal to crown-group muroids, which flourished from the Oligocene to the Miocene across Eurasia and North America. Eomyids, such as genera Eomys and Ligerimys, are documented in key European sites like Sansan in France (middle Miocene, ~14–12 mya), where diverse assemblages reveal early cricetid-like forms alongside other small mammals, and in North American localities such as the John Day Formation in Oregon (late Oligocene to early Miocene, ~30–18 mya), yielding abundant remains that highlight faunal turnover and dispersals. These families exhibit specialized dental morphologies for herbivory, with eomyids often dominating Oligo-Miocene rodent faunas before declining.^[49]^[50]^[51] Evolutionary milestones in the muroid fossil record include significant diversifications during the Miocene, with cricetid ancestors emerging in Asia around 20 mya, as seen in early forms like Democricetodon from Siwalik deposits in Pakistan and related sites. These developments coincide with global climatic cooling and habitat shifts, fostering radiations into arid and forested environments. In the Pleistocene, adaptations to Ice Age conditions are evident in fossils of woolly voles and lemmings (e.g., Dicrostonyx spp.), whose remains from Eurasian permafrost sites (~2.6–0.01 mya) show furred paws and dense pelage suited to snowy tundras, reflecting rapid morphological responses to glacial cycles.^[52]^[53] The muroid fossil record reveals minor extinction events, particularly in the Pliocene (~5.3–2.6 mya), where some Eurasian and North American lineages, including late-surviving eomyids and primitive cricetids, experienced localized losses linked to habitat fragmentation and cooling climates. Gaps persist in the record from Africa and South America due to poor preservation and limited exposures; the earliest African muroids appear in the late Oligocene (~25 mya) but remain sparse until the Miocene, while South American arrivals are documented only from the late Miocene (~7–5 mya), with pre-Pliocene evidence virtually absent.^[54]^[2]^[55]

Classification

Taxonomic hierarchy

Muroidea is a superfamily of rodents within the suborder Myomorpha of the order Rodentia, encompassing approximately 1,890 species distributed across six families: Platacanthomyidae, Spalacidae, Calomyscidae, Nesomyidae, Cricetidae, and Muridae.^[56] These families are further divided into 19 subfamilies and approximately 310 genera, reflecting the group's extensive diversification.^[56]^[1] The superfamily's type genus is Mus, with historical synonyms including Myoidea (proposed by Gill in 1872).^[3]^[43] The taxonomic framework of Muroidea originated with Illiger's establishment of the superfamily in 1811, but its initial broad delineation as a cohesive group was advanced by Simpson in 1945, who included diverse taxa like arvicolines and cricetines under a unified Muridae and Cricetidae, treating them as polyphyletic assemblages based on morphological traits.^[43] Subsequent refinements in the late 20th century incorporated molecular data, leading to the elevation of Nesomyidae to family status in the 1990s after cytochrome b analyses revealed paraphyly in the traditional Nesomyinae subfamily.^[57] Ongoing debates have centered on the placement of Calomyscidae, initially subsumed within Muridae due to morphological similarities but recognized as a distinct family in modern classifications following phylogenetic evidence of its basal position relative to other eumuroids.^[29]^[58] Current consensus on Muroidea's hierarchy is primarily drawn from the Mammal Diversity Database (version 2024c), which updates Wilson and Reeder's Mammal Species of the World (3rd edition, 2005) through genomic approaches and taxonomic revisions.^[59]^[56] Recent mitogenomic studies, including those from 2025, have prompted adjustments such as refined tribal boundaries and potential subfamily splits within Muridae, based on analyses of mitochondrial genomes that resolve previously ambiguous relationships among Old World murines.^[60]^[45]

Diversity and species counts

The superfamily Muroidea encompasses approximately 1,890 species, representing about 27% of global mammalian diversity.^[61] This estimate reflects recent phylogenetic analyses that account for undescribed forms and cryptic diversity, updating earlier reconstructions such as a 2017 study incorporating 900 species.^[29] Within Muroidea, the family Muridae dominates with around 1,043 species, followed by Cricetidae with approximately 803 species; Nesomyidae includes about 67 species, the majority of which are endemic to Madagascar.^[61] Patterns of diversity in Muroidea highlight elevated speciation rates in tropical regions, particularly Asia, where roughly 40% of species occur due to historical biogeographic opportunities in Southeast Asian archipelagos and karst habitats. Molecular delimitation methods have revealed extensive cryptic diversity, such as in the genus Rattus, where phylogeographic studies identify multiple independent lineages exceeding 100 putative forms across invasive populations.^[62] Additionally, around 10% of muroid species, primarily from genera like Rattus and Mus, function as global pests through invasive spread, impacting ecosystems on islands and continents.^[63] Conservation assessments indicate that approximately 26% of Muroidea species are threatened, according to IUCN evaluations, with 32 critically endangered and 70 endangered, driven by habitat fragmentation and invasive congeners.^[1] Endemism hotspots like Madagascar amplify risks, where about 50% of Nesomyidae species face extinction threats from deforestation and climate shifts, underscoring the superfamily's sensitivity to anthropogenic pressures.^[64]

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Feb 15, 2023 · The narrow-headed vole, collared lemming and common vole were the most abundant small mammal species across the Eurasian Late Pleistocene steppe ...
[53]
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Summary of each segment:
[54]
Pliocene colonization and adaptive radiations in Australia and New ...
Aug 7, 2025 · ... (Muroidea: Murinae) | The old endemic rodents of Australia and New Guinea (Sahul) represent one or more large adaptive radiations including ...
[55]
Mammal Species of the World. A taxonomic and geographic ...
A taxonomic and geographic reference. 2005. Don E. Wilson & DeeAnn M. Reeder (Eds.). Ed. 3, 2 vols., 2142 pp. Johns Hopkins University Press, Baltimore.
[56]
[PDF] Molecular Phylogeny and Biogeography of the Native Rodents of ...
cates: (1) rodents invaded Madagascar only once, (2) of Madagascar. Specifically, this study examines whether they came from Asia not from Africa as is ...Missing: colonization | Show results with:colonization
[57]
Phylogenetic Position of the Rodent Genus Typhlomys and the ...
We conclude that both Myodonta and Muroidea originated in Eurasia and that the 3 earliest divergences within Muroidea were restricted to the Eurasian ...Missing: review | Show results with:review
[58]
New insights into the mitogenomic phylogeny and evolutionary ...
Mar 27, 2025 · The current classification of Murinae includes 15 tribes ( Wilson and Mittermeier 2018 ). Previous studies have obtained different phylogenetic ...
[59]
Phylogeography of multiple lineages of black rats Rattus rattus ...
Mar 4, 2025 · With approximately 1,620 species (30% of mammal diversity) of the Muroidea ... [2024]) and 37 specimens of other Rattus species, such as ...
[60]
How many species of mammals are there? | Journal of Mammalogy
Feb 1, 2018 · The largest mammalian families are in the order Rodentia—Muridae (834 species versus 730 in MSW3) and Cricetidae (792 species versus 681 in MSW ...
[61]
Nesomyidae (African and Malagasy endemic rats and mice)
Nesomyidae is a diverse group of African rodents that has recently proven to be even more diverse than originally thought.Missing: colonization | Show results with:colonization
[62]
Invasive rats on tropical islands: Their population biology and ...
The three most invasive rat species, black or ship rat Rattus rattus, brown or Norway rats, R. norvegicus and Pacific rat, R. exulans have been incrementally ...
[63]
The macroevolutionary impact of recent and imminent mammal ...
Jan 10, 2023 · One hundred four of the extant lemurs (95%) are threatened with extinction, and of these 33 are classified as Critically Endangered (30% of all ...<|separator|>