Volcano rabbit

The volcano rabbit (Romerolagus diazi), also known as the teporingo or zacatuche, is a small lagomorph endemic to the upper slopes of the Chichinautzin volcanic range in central Mexico, where it inhabits high-altitude bunchgrass prairies and sparse pine-alder woodlands between 2,800 and 4,250 meters elevation.^[1] Weighing 390–600 grams with short legs, rounded ears, a vestigial tail, and dense yellowish-brown fur, it represents one of the world's smallest rabbits and exhibits primitive traits resembling those of pikas, such as vocal communication and reliance on specific grasses for cover and diet.^[1] Primarily herbivorous, it depends heavily on zacatón bunchgrass (Muhlenbergia macroura) for foraging, burrowing, and thermal regulation in its cool, temperate habitat.^[1] Classified as Endangered on the IUCN Red List since its assessment in 2019, the species persists in a restricted area of roughly 280 km², with population estimates below 7,000 individuals amid ongoing declines driven by habitat loss.^[2]^[1] Major threats include agricultural conversion, urban expansion from nearby Mexico City, logging, and wildfires that destroy essential zacatón stands, exacerbating fragmentation in this narrow endemic range along the Trans-Mexican Volcanic Belt.^[2]^[1] Despite legal protections within national parks and CITES Appendix I listing prohibiting international trade, enforcement challenges and land-use pressures persist, underscoring the species' vulnerability as a relict lagomorph with limited dispersal capabilities.^[2]^[1] Semi-social in groups of 2–5 individuals, volcano rabbits display diurnal activity peaks at dawn and dusk, breeding year-round with females producing up to 4–5 litters annually of 1–4 young after a 38–40 day gestation, adaptations that support survival in their precarious montane refugia.^[1] Their vocalizations—including barks, squeaks, and foot-thumping—facilitate group cohesion and predator deterrence, highlighting behavioral specializations evolved in isolation from other leporids.^[1] Conservation efforts emphasize habitat restoration and monitoring, yet the species' dependence on undisturbed zacatón ecosystems renders it highly susceptible to anthropogenic and climatic perturbations.^[2]^[1]

Taxonomy

Etymology

The English common name "volcano rabbit" reflects the species' exclusive occurrence on the uppermost slopes of four inactive volcanoes—Popocatépetl, Iztaccíhuatl, El Pelado, and Nevado de Toluca—in the Trans-Mexican Volcanic Belt, at elevations between 3,000 and 4,300 meters.^[3]^[4] In Mexico, the species is referred to as zacatuche or teporingo, indigenous terms predating European contact. Zacatuche originates from Nahuatl, derived from zacatl (grass) combined with a root denoting rabbit, translating to "grass rabbit" in allusion to its dependence on dense bunchgrass (Zacaton) understory for cover and forage.^[5] The etymology of teporingo is uncertain and not well-documented in historical records.^[6] The scientific binomial Romerolagus diazi combines the monotypic genus Romerolagus, erected by American mammalogist C. Hart Merriam in 1896 to accommodate specimens from Mount Popocatépetl, with the specific epithet honoring Mexican engineer and geographer Agustín Díaz y Guerrero (1853–1927), who collected the type series in December 1892 on the northeastern slope of Volcán Iztaccíhuatl.^[5]^[7] Merriam's genus name derives from Matías Romero (1837–1898), Mexican Minister of the Treasury and a key diplomatic figure, compounded with the Greek lagos (hare).^[5] Ferrari-Pérez initially described it as Lepus diazi in 1893 within a report by the Mexican Geographical and Exploring Commission, based on Díaz's specimens from near San Martín Texmelucan, Puebla.^[8]^[6]

Phylogeny

The volcano rabbit (Romerolagus diazi) is classified within the family Leporidae of the order Lagomorpha, which comprises rabbits, hares, and pikas.^[9] Phylogenetic analyses based on molecular data, including mitochondrial genomes, position Romerolagus as basal within Leporidae, often as sister to the remaining genera excluding Pronolagus (African hares) and Nesolagus (striped rabbits).^[10]^[11] This topology aligns with maximum likelihood reconstructions using protein-coding genes from the complete mitochondrial genome of R. diazi, which shares the standard leporid gene order but underscores its early divergence.^[10] Within the broader Leporidae phylogeny, Romerolagus diverged from other North American lineages approximately 12.8 million years ago, stemming from a common leporid ancestor that radiated during the Miocene.^[12] Molecular supermatrix approaches incorporating nuclear and mitochondrial sequences reinforce this early split, distinguishing Romerolagus from more derived clades like Lepus (hares) and Sylvilagus (cottontails).^[13] Cytogenetic data, including G-banded chromosomes, suggest an ancestral karyotype shared with Lepus, supporting a common evolutionary origin for leporids rather than recent convergence.^[5] The phylogeographic history of R. diazi reflects adaptation to montane environments in the Trans-Mexican Volcanic Belt, shaped by interactions between climatic oscillations and volcanic activity since the Pleistocene.^[14] Molecular evidence from mitochondrial DNA indicates population expansions during interglacial periods and contractions linked to glacial advances and eruptions, such as those from Popocatépetl, fostering genetic structure across isolated subpopulations.^[14] This dynamic history, evidenced by haplotype diversity, highlights R. diazi as a mountain specialist with limited gene flow, contrasting with more vagile leporids.^[14]

Physical characteristics

Morphology

The volcano rabbit (Romerolagus diazi) is among the smallest rabbit species, with adults typically measuring 270–315 mm in total length, hind foot lengths of 42–55 mm, ear lengths of 40–45 mm, and body weights ranging from 380 to 600 grams.^[15] It possesses short hind legs and feet relative to its body size, small rounded ears, and a vestigial tail that is externally invisible.^[1]^[5] The fur is thick, soft, and dense, providing insulation in its high-altitude habitat; dorsal and lateral pelage is yellowish-brown, while the ventral surface is white or buff-colored.^[1] There is minimal sexual dimorphism in body size between males and females.^[16] The species exhibits several primitive morphological traits, including a reduced supraorbital process and unique dentition, distinguishing it from more derived leporids.^[5]^[16]

Adaptations

The volcano rabbit (Romerolagus diazi) exhibits morphological adaptations suited to its dense, high-altitude bunchgrass habitat, including short hind legs and feet that enable maneuvering through tall zacatón (Muhlenbergia spp.) tussocks rather than relying on the long leaps typical of open-country leporids.^[1] Small, rounded ears and a vestigial tail minimize snag risks in vegetation, while grizzled yellowish-brown dorsal fur provides camouflage against grassy substrates.^[1] Its compact body size (average adult mass 417–536 g) and large, laterally positioned eyes facilitate low-profile movement and wide-angle detection of threats in obstructed environments at elevations of 2,800–4,250 m.^[1] Behaviorally, the species relies on zacatón cover for predator evasion, lacking the speed of hares or larger rabbits, and constructs shallow burrows or nests within grass clumps for shelter and concealment of young.^[1] Unlike most leporids, R. diazi is vocal, producing high-pitched barks, growls, screams, and clicks—adaptations likely evolved for communication and alarm signaling in visually limited dense vegetation, where olfactory and auditory cues predominate over sight.^[1] Semi-social groups of 2–5 individuals use scent glands for territorial and reproductive signaling, enhancing coordination in patchy habitats.^[1] Physiologically, the volcano rabbit practices cecotrophy, reingesting soft feces to maximize nutrient and water extraction from fibrous forage.^[1] Its gut microbiota is enriched with fiber-degrading enzymes such as endo-1,4-β-xylanases (up to 363 sequences per sample), endoglucanases, and tannases (18 bacterial sequences identified), enabling efficient breakdown of phenolic-rich, lignified zacatón grasses that dominate its diet and provide both nutrition and structural habitat.^[17] These microbial adaptations support specialization on tough, low-quality vegetation unavailable or indigestible to generalist herbivores.^[17]

Distribution and habitat

Geographic range

The volcano rabbit (Romerolagus diazi) is endemic to central Mexico, with its geographic range restricted to the central-eastern portion of the Trans-Mexican Volcanic Belt. This area encompasses fragmented habitats primarily in the Sierra Nevada, including the slopes of Popocatépetl and Iztaccíhuatl volcanoes, and the Sierra Chichinautzin, spanning the states of México, Morelos, and Puebla.^[18]^[19] The species' distribution consists of three to four discontinuous core areas, with a total extent of approximately 386 km², all located within about 100 km south and east of Mexico City. Key sites include El Pelado volcano in the Sierra Chichinautzin and the volcanic highlands near Ajusco. Populations are absent from surrounding regions despite suitable elevations elsewhere in the volcanic belt, likely due to historical habitat fragmentation and specific ecological requirements.^[20]^[5]^[21] Historical records indicate a contraction of the range since the 1990s, with surveys confirming presence only in these limited patches rather than a continuous distribution across the belt. The proximity to urban expansion from Mexico City has intensified isolation of these remnants.^[3]

Habitat preferences

The volcano rabbit (Romerolagus diazi) is a habitat specialist primarily inhabiting subalpine bunchgrass grasslands at elevations ranging from 2,800 to 4,250 meters in the Trans-Mexican Volcanic Belt of central Mexico.^[1] These grasslands are dominated by dense zacatón bunchgrasses, such as Muhlenbergia macroura and Festuca tolucensis, which provide essential cover, foraging resources, and sites for burrows and runways.^[22] ^[23] Higher relative abundances occur in patches above 3,600 meters with rock cover below 15%, moderate slopes, and vegetation structures offering tussock density for predator evasion and thermoregulation.^[19] ^[22] The species avoids densely forested areas or those with sparse grass cover, though it tolerates open pine (Pinus spp.) woodlands where bunchgrasses persist in the understory, albeit at lower densities than in pure grasslands.^[24] Habitat suitability is further influenced by factors like soil moisture and altitude, with preferences for drier, higher sites supporting robust grass growth.^[23]

Behavior

Activity patterns

The volcano rabbit (Romerolagus diazi) displays primarily diurnal activity, remaining active throughout daylight hours but with pronounced peaks at dawn and dusk, during which foraging and movement are most intense.^[1]^[20] Midday periods typically involve quieter resting amid zacatón grass (Muhlenbergia macroura) clumps, minimizing exposure to heat and predators.^[5] Above-ground behaviors such as foraging, playing, chasing, and social interactions predominate during these active phases, often centered within grass tussocks that provide cover.^[5] Limited observations suggest potential nocturnal activity in captive settings, though wild populations show stronger diurnal tendencies tied to their high-altitude grassland habitat.^[5] These patterns align with crepuscular traits common in small lagomorphs, enhancing survival in predator-rich environments.^[1] Volcano rabbits exhibit semi-social behavior, typically living in small groups of 2 to 5 individuals that occupy shared burrow systems up to 5 m long and 40 cm deep.^[1] These groups are structured around a dominant breeding pair, with the female occupying the top position in the hierarchy; subordinate individuals, often juveniles or non-breeding adults, assist in territory maintenance but rarely reproduce.^[1] Social interactions include aggressive displays such as biting and chasing to establish dominance, particularly among males competing for breeding access, though larger captive groups of up to six have shown only one male and 1-2 females successfully breeding.^[20] Territories are defended collectively through vocalizations and thumping, with group cohesion aiding predator avoidance in their grassland habitat.^[15] Reproduction is aseasonal but intensifies during the summer rainy season, with females capable of producing 4-5 litters annually under optimal conditions.^[15] Gestation lasts 34-48 days, averaging approximately 39 days in controlled settings.^[25] Litters consist of 2-4 young on average, though captive records report variability up to 10 offspring across multiple litters from a single female; newborns weigh 25-32 g and are altricial, relying on maternal care in nested burrows lined with grass.^[25] ^[1] Weaning occurs at about 3 weeks, after which juveniles integrate into the group hierarchy, with sexual maturity reached by 6-8 months.^[25] In captivity, breeding success has been limited, with high juvenile mortality unless pairs are housed in enriched environments mimicking natural burrows.^[25]

Ecology

Diet and foraging

The volcano rabbit (Romerolagus diazi) maintains a herbivorous diet dominated by graminoids from zacatón bunchgrasses, which form dense clumps providing both sustenance and concealment in its high-altitude habitat. Primary food plants include Muhlenbergia macroura, Festuca amplissima, Festuca rosei, Stipa ichu, and Epicampes species, with these tussock grasses comprising the bulk of intake due to their abundance and nutritional value.^[8]^[26] Fecal pellet analysis from field studies in central Mexico indicates a diverse diet encompassing over 70 plant species, though graminoids consistently predominate year-round, supplemented by forbs, shrubs, and lichens. Grasses contribute up to 80% of the diet in wet seasons when fresh growth is plentiful, reflecting selective foraging for tender shoots and leaves within zacatón stands.^[27]^[26] In dry and cold periods, such as winter, consumption shifts toward woody browse including shrubs and tree bark to meet energy demands amid grass senescence, with digestibility trials confirming higher intake of M. macroura-based diets supporting body weight maintenance. Foraging behavior emphasizes cover-dependent grazing, where individuals clip and consume vegetation close to the ground within grass tussocks up to 2.5 meters tall, minimizing exposure to aerial predators while exploiting the structural refuge of these perennials.^[26]^[28]^[3]

Predators, parasites, and natural interactions

The volcano rabbit (Romerolagus diazi) is preyed upon by several native mammalian and avian carnivores in its high-altitude grassland habitat. Documented predators include long-tailed weasels (Mustela frenata), bobcats (Lynx rufus), coyotes (Canis latrans), and red-tailed hawks (Buteo jamaicensis), which exploit the rabbit's small size and ground-dwelling habits.^[1] ^[3] Analysis of coyote scat from multiple seasons has revealed bone fragments identifiable as volcano rabbit remains, indicating predation on adults occurs year-round.^[29] Bobcats similarly incorporate the species into their diet, though not as a primary food source, based on dietary studies in overlapping ranges.^[30] To counter predation risks, volcano rabbits employ cryptic behaviors, remaining concealed within dense zacaton grass (Muhlenbergia macroura) patches during daylight hours and using vocalizations—such as barks and squeaks—to alert group members of threats.^[1] ^[3] These adaptations reflect the species' reliance on vegetative cover for evasion, as open areas increase vulnerability to aerial and terrestrial hunters.^[1] The volcano rabbit hosts a range of endoparasites, primarily helminths adapted to lagomorph hosts. Nematodes such as Boreostrongylus romerolagi inhabit the duodenum, while other species including Trichostrongylus calcaratus and Longistriata dubia occur in the gastrointestinal tract.^[31] ^[1] Cestodes like Anoplocephaloides romerolagi have been identified as intestinal parasites in examined specimens.^[32] Ectoparasites include the tick Ixodes neotomae and the mite Cheyletiella mexicana, both recorded on wild individuals.^[5] These parasites likely impose physiological costs, though prevalence data remain limited due to the species' endangered status and challenging field conditions.^[33] Ecologically, volcano rabbits serve as a prey resource sustaining local predator populations, contributing to trophic dynamics in pine-bunchgrass ecosystems without evidence of mutualistic or competitive interactions with other herbivores beyond shared habitat use.^[30] Their burrowing and foraging may indirectly aerate soil and disperse grass seeds, but such roles await quantitative confirmation.^[1]

Conservation

Population status and trends

The volcano rabbit (Romerolagus diazi) is classified as Endangered on the IUCN Red List, with a current population trend assessed as decreasing due to ongoing habitat loss and fragmentation. Global population estimates indicate approximately 7,000 mature individuals across its restricted range of less than 400 km² in the Trans-Mexican Volcanic Belt, primarily in fragmented patches on volcanoes such as Iztaccíhuatl, Popocatépetl, and El Pelado.^[2]^[34] Local surveys provide varying figures; for instance, a study on El Pelado volcano estimated a mean population of 6,488 individuals (range: 2,478–12,120), recommending the lower bound for conservative planning.^[21] Population declines are attributed to habitat degradation from agricultural expansion, overgrazing by livestock, urbanization near Mexico City, and periodic wildfires, which reduce suitable bunchgrass cover essential for the species. No comprehensive range-wide surveys have been conducted since the early 2010s, but localized monitoring shows persistent fragmentation and absence from formerly occupied areas, with relative abundance indices dropping in burned habitats as of 2024.^[4]^[35] The species persists in fewer than 200 colonies, vulnerable to stochastic events, underscoring the need for updated demographic data to refine threat assessments.^[36]

Primary threats

The primary threats to the volcano rabbit (Romerolagus diazi) stem from anthropogenic habitat alteration and loss, which have reduced and fragmented its specialized grassland ecosystem on the slopes of Popocatépetl and Iztaccíhuatl volcanoes in central Mexico.^[19]^[1] Livestock grazing, particularly by cattle, degrades the zacatón bunchgrasses (Muhlenbergia macroura and M. quadridentata) essential for cover and foraging, as overgrazing compacts soil, reduces grass density, and favors invasive species, leading to local population declines especially in the Sierra Chichinautzin region.^[19] Agriculture, logging, and grass harvesting for construction and fuel further diminish habitat continuity, with estimates indicating a 15–20% loss over the past three generations due to these activities.^[37]^[1] Urban expansion and infrastructure development, including highway construction and residential growth around Mexico City, exacerbate fragmentation by isolating subpopulations and increasing edge effects that expose rabbits to predators and reduce genetic connectivity.^[38]^[1] Wildfires, often ignited for land clearance or grazing expansion, destroy dense grass cover critical for concealment, though natural fire regimes may historically play a role in maintaining habitat; human-induced burns now occur more frequently and intensely.^[1] Hunting, while documented historically, appears secondary to habitat pressures, with limited current evidence of widespread impact.^[4] Emerging climate pressures compound these issues, as projected temperature rises of 2–4°C by 2050 could shift suitable habitat upslope by up to 700 meters, compressing the species' range against volcanic summits and overlapping with intensified human land use.^[3] This elevational constraint, combined with reduced precipitation potentially stressing bunchgrasses, heightens vulnerability without adaptive migration corridors.^[39] Overall, these threats have contracted the species' distribution from an estimated original range of over 1,000 km² to less than 400 km², underscoring the need for targeted land-use restrictions.^[36]

Conservation efforts and outcomes

Conservation efforts for the volcano rabbit have primarily emphasized habitat management, research, and limited captive breeding initiatives. National and international policies in Mexico regulate human activities in the species' range, including restrictions on land use to mitigate fragmentation from agriculture and urbanization. These measures aim to preserve zacatón bunchgrass ecosystems on volcanic slopes, though enforcement remains inconsistent due to competing local economic pressures. Community-based programs, supported by organizations like the Zoological Society of London through its EDGE of Existence initiative, engage indigenous groups in monitoring and sustainable practices to reduce overgrazing by cattle, which studies identify as a key factor lowering rabbit occupancy. A 2020 survival blueprint outlines prioritized actions, including fire suppression protocols and restoration of degraded bunchgrass patches, recognizing the rabbit's dependence on unburned, ungrazed habitat for cover and foraging.^[40]^[19] Research and monitoring constitute a core component, with projects funded by the Rufford Foundation documenting distribution, population genetics, and responses to disturbances like wildfires. Field surveys from 2017 onward have mapped occupancy patterns, revealing that proximity to human settlements correlates with reduced abundance, informing targeted exclusion zones. Environmental education campaigns target local communities to curb incidental hunting during bird shoots and promote zacatón conservation, as the grass's overexploitation for thatching exacerbates habitat loss. Captive breeding trials at Chapultepec Zoo in Mexico City achieved initial successes in the early 2000s, producing offspring under semi-captive conditions, though scaling to viable reintroduction remains unproven due to logistical challenges and low genetic diversity risks.^[41]^[42]^[43] Outcomes have been modest, with no documented population recovery amid persistent threats. Post-wildfire assessments in the Ajusco-Chichinautzin corridor, burned in 2021, showed decreased rabbit occupancy and altered relative abundance indices in regenerating bunchgrass communities, indicating slow habitat recovery insufficient for population rebound. Legal protections against habitat conversion exist, but fragmentation continues, confining viable patches to less than 500 km² across disjointed volcanic sites. Genetic studies highlight inbreeding risks from isolation, underscoring the need for connectivity corridors that have not yet materialized. Overall, while research has refined threat models—prioritizing grazing exclusion over direct predator control—the species' endangered status persists, with efforts hampered by inadequate funding and enforcement in rural Mexican highlands.^[35]^[20]^[19]