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Amegilla

Amegilla is a genus of robust, solitary bees in the family Apidae, subfamily Apinae, and tribe Anthophorini, comprising approximately 260 species worldwide.^[1] These bees are characterized by their long tongues, protuberant clypeus, short antennae, and dark metasoma with pale tergal hairbands, with many species featuring colorful, iridescent abdominal bands, such as the blue bands in species like A. cingulata.^[1]^[2] Native to the Old World, Amegilla species are distributed across Africa, Europe, Asia, Australia, and the Pacific islands, thriving in diverse habitats from arid regions and shrublands to farmlands and forests.^[1]^[3] They are ground-nesting bees that dig burrows in soil or soft substrates, exhibiting solitary behavior with males often emerging before females and mating occurring on flowers.^[1]^[3] Amegilla bees play a crucial ecological role as polylectic pollinators, visiting a wide range of plants from families such as Lamiaceae, Fabaceae, and Boraginaceae, and are particularly noted for their ability to perform buzz pollination, where they vibrate flowers to release pollen from poricidal anthers.^[1]^[3] This specialized technique makes them essential for pollinating crops like tomatoes, eggplants, and chilies, as well as wild plants such as bottlebrush and grass trees, contributing to biodiversity and agriculture in their native ranges.^[3] Fast-flying and active in warm summer conditions, these bees hover while foraging and are generally non-aggressive, though capable of stinging.^[1]^[4] The genus includes several subgenera, such as Asaropoda, Micramegilla, Notomegilla, and Zonamegilla, reflecting its taxonomic diversity and evolutionary adaptations across regions.^[2]

Taxonomy

Genus classification

Amegilla is a genus of bees belonging to the family Apidae, subfamily Apinae, and tribe Anthophorini.^[5] The genus was established by Heinrich Friese in 1897.^[6] Subsequent taxonomic revisions have divided Amegilla into multiple subgenera, reflecting its morphological and geographical diversity across the Old World.^[2] Bees in the genus Amegilla are identified by several key diagnostic traits, including a robust body form, an elongated proboscis adapted for accessing deep floral nectaries, a prominently protuberant clypeus, relatively short antennae, the absence of arolia between the tarsal claws, and a shortened marginal cell in the forewings.^[1] These features distinguish Amegilla from closely related genera such as Anthophora, which typically exhibit larger body sizes, the presence of arolia, and less distinctly banded metasomal pubescence.^[7]^[8]

Subgenera and phylogeny

The genus Amegilla is subdivided into approximately 11 subgenera, primarily recognized through morphological cladistic analyses that emphasize characters such as metasomal tergal hair structure, male sternal morphology, and gonostylar features.^[9] Key subgenera include Asaropoda (characterized by turret-building nesting behaviors and fluffy pubescence), Micramegilla (with compact body forms and metallic sheen), Notomegilla (featuring broad metasomal bands), and Zonamegilla (noted for extensive species diversity exceeding 50 taxa and iridescent coloration).^[9]^[2] Other recognized subgenera encompass Glossamegilla (adapted with elongated proboscides for deep-corolla flowers), Zebramegilla, Dizonamegilla, Megamegilla, Aframegilla, Ackmonopsis, and Atrugilla, though some classifications, such as Michener's 2007 revision, questioned their monophyly and opted against formal subdivision.^[9]^[10] Phylogenetic relationships within Amegilla are derived from combined morphological and molecular studies, positioning the genus as a monophyletic clade within the tribe Anthophorini, closely allied to Anthophora based on shared apomorphic traits like wing venation and ground-nesting habits.^[11] Cladistic analyses using 51 morphological characters across 26 anthophorine taxa reveal two primary Amegilla lineages: one with narrow, triangular male eighth sternites (encompassing six subgenera) and another with broad, quadrate sternites (five subgenera), reflecting gradual evolutionary transitions via intermediate species groups.^[9] The genus exhibits Old World origins, likely in northern Southeast Asia during the Oligocene, with multiple dispersal events facilitating its diversification across the Palearctic, Afrotropical, Indomalayan, and Australasian realms.^[11] Molecular phylogenies incorporating mitochondrial cytochrome oxidase I sequences further support subgeneric boundaries, particularly in regional revisions.^[2] Amegilla comprises over 250 species worldwide, with recent taxonomic revisions refining counts in specific regions; for instance, a 2017 study delineated 43 species (31 in Notomegilla and 12 in Zonamegilla) across the Australian subgenera Notomegilla and Zonamegilla, while approximately 36 species are recognized in the West Palearctic.^[1]^[2]^[4] Evolutionary adaptations within the genus include the development of buzz pollination (floral sonication), which has arisen independently multiple times in Anthophorini and is linked to specialization on poricidal flowers, enhancing pollen collection efficiency and driving diversification in bee-flower interactions.^[12] This behavior, facilitated by thoracic vibrations, correlates phylogenetically with xerophilic habits and polylectic foraging, underscoring Amegilla's role in the tribe's adaptive radiation.^[9]^[11]

Physical characteristics

Morphology

Amegilla bees are medium-sized insects, typically measuring 10-12 mm in length, with a robust, stout build that supports their ground-nesting and foraging lifestyles.^[13]^[1] The head is golden brown and features short antennae—12 segments in females and 13 in males—along with a protuberant clypeus that aids in soil excavation for nesting.^[3]^[1] Thoracic adaptations include a densely pubescent mesonotum for insulation, while the legs are notably hairy, particularly the hind legs with a scopa formed by branched hairs that facilitate pollen collection.^[13] The proboscis is long, enabling efficient nectar extraction from deep flowers.^[1] The abdomen, or metasoma, is dark and robust, featuring transverse stripes with light hairbands that contribute to its overall structure.^[3] Wing venation, characterized by three submarginal cells and a straight basal vein, supports the bees' fast-flying and hovering capabilities essential for their agile foraging.^[13] These iridescent bands on the abdomen enhance visibility during flight but are secondary to the structural adaptations described.^[3]

Coloration and sexual dimorphism

Amegilla bees are renowned for their striking abdominal coloration, characterized by metallic blue or green bands created by specialized iridescent pubescence on the metasomal terga. These bands, which result from light-diffracting structures in the hairs, provide a signature visual trait for many species in the genus, particularly in subgenera like Zonamegilla and Notomegilla. For instance, in Amegilla cingulata, the abdomen displays bold metallic blue bands against a dark background, enhancing the bee's iridescent appearance under sunlight.^[14]^[15] Sexual dimorphism in coloration is evident primarily through the number of complete abdominal bands, with females typically exhibiting four and males five, as seen in species such as Amegilla cingulata and Amegilla zonata. This difference aids in rapid field identification, alongside other traits like males' smaller body size, longer antennae (with 13 segments versus 12 in females), and lack of a sting, while females are larger and possess stronger tibial spurs adapted for digging nests.^[3]^[15]^[16] Coloration varies across subgenera and species, with some lacking prominent bands altogether. In Amegilla chlorocyanea, bands may appear pale blue or cream rather than intensely metallic, and fading or orange tinges can occur, complicating identification but underscoring the role of these iridescent hairs in taxonomic keys and field guides. Such variations highlight how abdominal banding serves as a key diagnostic feature for differentiating Amegilla species.^[14]^[17]

Distribution and habitat

Geographic range

The genus Amegilla is distributed across the Old World, primarily in tropical and subtropical regions of Africa, Asia, Australia, and parts of Europe, with no recorded presence in the Americas.^[18] Species are native to southern and eastern Africa, including Madagascar, where they inhabit diverse landscapes from coastal lowlands to highlands.^[3] In Asia, the genus extends from India through Southeast Asia to southern China, with notable concentrations in the Indo-Australian Archipelago, including Borneo and New Guinea.^[19] The highest species diversity occurs in Australia and Southeast Asia, where multiple subgenera such as Asaropoda, Notomegilla, and Zonamegilla are represented.^[17] In Australia, Amegilla species exhibit broad distributions, often centered in arid and semi-arid zones, with examples like Amegilla cingulata widespread across most of the continent including the Northern Territory, except Tasmania.^[20]^[21] Other species, such as A. chlorocyanea, are prevalent in southern regions, while tropical and subtropical east-coast populations include A. bombiformis and A. rhodoscymna.^[22] The subgenus Asaropoda is particularly extensive, spanning Australia, New Guinea, and the Bismarck Archipelago.^[17] In the West Palearctic region, 36 species are documented, primarily in Mediterranean and southern areas including 11 in Europe.^[4]^[1] While most Amegilla species remain within their native ranges and exhibit limited long-distance migration due to their largely sedentary nature, some have expanded through human-mediated introductions. For instance, A. pulchra, native to eastern Australia, has become widespread in the Pacific islands, including Fiji, potentially altering local pollination dynamics.^[23] Fast flight capabilities facilitate local dispersal within suitable habitats.^[22]

Habitat preferences

Amegilla bees primarily inhabit arid and semi-arid regions, where they thrive in environments such as scrublands, grasslands, rangelands, and coastal dunes.^[24]^[3] These bees are particularly diversified in areas featuring Mediterranean vegetation, extending their presence to subtropical and tropical zones with low rainfall.^[1] While the genus is most abundant in Australia and parts of Africa and Asia, species like Amegilla dawsoni are specialized for sparsely vegetated, open landscapes in arid interiors.^[25]^[26] For nesting, Amegilla species favor loose, sandy, loam, or clay soils that allow for burrowing, often selecting soft sandstone banks, clay washouts, or even mud bricks in human-modified areas.^[24] These sites are typically chosen on sloping ground protected from flooding, with nests constructed in close proximity to abundant flowering plants to facilitate foraging.^[14] Such preferences ensure access to nectar and pollen resources while minimizing exposure to environmental stressors. Amegilla bees exhibit strong adaptations to hot, dry climates, with many species active during warmer months and capable of multiple generations per year in subtropical conditions.^[14] Some, like those in Mediterranean-influenced habitats, tolerate seasonal variations but generally avoid regions with extreme temperature fluctuations or excessive humidity.^[1] Regarding habitat fragmentation, these bees show a clear preference for large, undisturbed soil patches, as urbanization and impervious surfaces disrupt nesting aggregations and foraging routes.^[14]

Ecology

Nesting and life cycle

Amegilla bees are solitary digger bees, with each female independently excavating and maintaining her own burrow in the ground, lacking any cooperative or eusocial interactions with conspecifics. Although nests are often solitary, females may aggregate in suitable sites, forming loose congregations where burrows are spaced as closely as 0.5 cm apart, facilitating shared microhabitats without division of labor.^[3]^[27] Nest structure typically involves a vertical main tunnel extending to a cluster of brood cells at the terminus; entrances may feature a tumulus or mud turret 1–2 cm high formed from excavated soil. For example, in A. zonata and A. dawsoni, tunnels are 4–30 cm deep with 2–21 brood cells that are cylindrical or urn-shaped, measuring 1.3–2 cm in length and 0.4–0.7 cm in diameter, with walls lined by a thin layer of waterproof wax secreted by the female's Dufour's gland to prevent desiccation. Provisioning follows a mass-provisioning strategy, where females deposit a compact mass of pollen mixed with nectar (e.g., averaging 0.07 g per cell in A. zonata) before oviposition, after which the cell is capped with a multilayered plug of soil and secretions. Cells are oriented vertically and branched off the main tunnel, with progressive construction allowing for multiple offspring per nest. Nests are preferentially built in soft, bare soils like clay, sand, or disturbed earth, aligning with broader habitat preferences for well-drained substrates.^[27]^[28]^[29] The life cycle varies by species and region, ranging from univoltine (one generation per year, as in A. dawsoni, with activity from July to September) to multivoltine (multiple generations in warmer climates, up to three per year in A. zonata). Eggs are creamy white, cylindrical, and laid singly atop the provision mass; larvae progress through four instars, initially feeding on the pollen-nectar mixture and later consuming the wax lining, before defecating and entering a motionless prepupal stage. For instance, in A. zonata, pupation occurs within the cell without a silken cocoon, lasting about 14 days, with emerging adults cleaning themselves before exiting in warm seasons to synchronize with floral resources. Overwintering may involve diapause in the prepupal stage, lasting several months to a year or more in arid environments, with some individuals surviving longer under laboratory conditions.^[28]^[27] Parental care is minimal and maternal, confined to nest construction, provisioning, and capping, after which females abandon the site with no further investment in offspring development. Females may briefly guard the nest entrance by remaining in the shaft during vulnerable periods, deterring parasites such as mutillid wasps and dipterans through presence or swarming behavior, but no prolonged vigilance or progressive feeding occurs, consistent with the solitary nature of the genus.^[28]^[30]^[2]

Pollination and foraging

Amegilla bees are effective pollinators, particularly through buzz pollination, where females vibrate their thoracic muscles to dislodge pollen from poricidal anthers in flowers such as those of tomatoes (Solanum lycopersicum) and blueberries (Vaccinium spp.). For example, A. murrayensis vibrates at frequencies around 350 Hz. This sonication behavior involves grasping the flower and tapping the anthers with the head, enabling efficient pollen release and transfer to the stigma, which enhances fruit set and seed production in buzz-dependent crops like tomatoes and eggplants (Solanum melongena).^[31]^[32]. Unlike some bumblebees that bite anthers, Amegilla species rely on rapid head knocks, allowing quicker visits of about 1 second per flower and potentially higher pollen extraction rates compared to other sonicators (e.g., in A. murrayensis).^[31]. As polylectic foragers, Amegilla bees visit a wide range of plant families, including Solanaceae, Fabaceae, Lamiaceae, Boraginaceae, and Cucurbitaceae, collecting both nectar and pollen to provision larvae, though they do not produce honey for storage.^[1]^[33]. Their long tongues enable access to deep corollas, and they exhibit fast, hovering flight during foraging, often without landing on shallow flowers. In some species, such as A. quadrifasciata, visitation rates exceed 18 flowers per minute.^[1]^[34]. For example, in greenhouse settings with A. chlorocyanea, females typically make around 9 pollen-collecting flights per day, buzzing approximately 9.3 flowers per minute during each, resulting in over 1,100 vibrations delivered daily, contributing to superior pollen transfer efficiency for larval provisioning and crop pollination.^[35]. Foraging activity in Amegilla is diurnal, peaking midday when temperatures and sunshine are highest, aligning with their adaptations to warm, arid, and Mediterranean habitats where they interact with specialized flora such as drought-tolerant species in Fabaceae and Solanaceae.^[1]^[36]. This timing optimizes nectar and pollen collection from sun-exposed blooms, though females also forage from dawn to dusk on diverse crops and weeds, preferring yellow and blue flowers for efficient resource gathering.^[33].

Behavior

Behaviors in Amegilla vary by species, but common patterns include the following. Amegilla bees are primarily solitary, lacking the eusocial structures of honeybees or bumblebees, but they exhibit aggregative nesting behaviors where females construct burrows in close proximity, often forming loose clusters that can include thousands of nests in a single site.^[29] This aggregative pattern enhances resource efficiency by concentrating nests near optimal foraging areas and soil conditions suitable for burrowing.^[37] Communication among Amegilla individuals is limited and relies on pheromonal cues rather than complex visual or auditory signals like dances. Cuticular hydrocarbons serve as pheromones that facilitate recognition and repulsion between individuals.^[38] Unlike social bees, Amegilla do not form hives or engage in elaborate communal signaling. Interspecific interactions include kleptoparasitism, where cuckoo bees of the genus Thyreus invade Amegilla nests to lay eggs, exploiting the host's provisions for their offspring.^[39] Amegilla also face competition for floral resources from introduced honeybees (Apis mellifera), which can deplete nectar and pollen sources, potentially reducing foraging success for native species like Amegilla in shared habitats.^[40] Territoriality is pronounced in females, who aggressively defend nest entrances by remaining stationed within the burrow shaft to deter intruders, occasionally deploying stings as a last resort despite their generally non-aggressive nature.^[37]^[3]

Reproduction and mating

Amegilla bees display a promiscuous mating system in which males employ alternative tactics to secure multiple mates, while females typically mate only once. Territorial males defend emergence sites adjacent to nests, aggressively chasing rivals and waiting for virgin females to emerge from pupation before attempting copulation, which lasts 1.5 to 18 minutes on average. Smaller males, in contrast, patrol flower patches, pouncing on and mating with receptive females encountered during foraging, often without establishing territory. This dual strategy allows males to maximize encounters, with larger individuals dominating nest sites through combat and smaller ones relying on mobility at flowers.^[41]^[37] Courtship relies primarily on chemical cues, with cuticular hydrocarbons (CHCs) on the exoskeleton of virgin females serving as attractants to guide patrolling or territorial males. Virgin females produce a distinct CHC profile that elicits male pursuit and mounting, whereas mated or nesting females alter their CHCs, rendering them unattractive and reducing male harassment. Males may also hover near nest entrances to detect emerging females via these scents, though no elaborate visual displays have been documented beyond general hovering and darting behaviors.^[38]^[37] Following mating, females engage in oviposition by constructing individual brood cells in the nest, provisioning them with pollen and nectar, and laying a single egg on the food mass before sealing the cell with wax and soil. In species such as A. dawsoni, offspring size and sex are determined early in this process, with females adjusting provisioning effort—through more or fewer foraging trips—to produce larger daughters or major sons versus smaller minor sons, resulting in a seasonally variable sex ratio that balances parental investment overall.^[42]^[43] During the reproductive period, particularly nesting, females show heightened aggression to defend brood cells, blocking nest entrances and swarming potential intruders such as parasites or predators. They possess a mild stinging capability, delivering a painful but non-lethal jab comparable to a pinprick when directly threatened or handled roughly, without the autotomy seen in honey bees; males exhibit less defensiveness overall.^[37]^[3]

Conservation and threats

Population status

The genus Amegilla includes approximately 260 species worldwide, with the majority not considered globally threatened, though assessments remain limited for most.^[1] In Europe, 11 species have been evaluated under the IUCN Red List criteria as of 2014, with several classified as Least Concern (e.g., A. albigena, A. canifrons, A. garrula, A. quadrifasciata) and others as Data Deficient due to insufficient information on population sizes and trends.^[44]^[1] In Australia, a key diversity hotspot with over 60 species across subgenera such as Asaropoda (21 species), Zonamegilla (12+ species), and Notomegilla (2 species), populations appear stable in native arid and semi-arid zones, where species like A. dawsoni emerge in aggregations of thousands during seasonal breeding.^[22]^[45] However, some Australian endemics exhibit regional declines and vulnerability, such as A. dawsoni, which faces risks from habitat disturbance in its restricted claypan nesting sites in Western Australia.^[46]^[29] Europe represents a lower diversity area by comparison, with no endemic Amegilla species listed as threatened.^[44] Few Amegilla species have formal IUCN global assessments, reflecting broader gaps in bee taxonomy and monitoring; for instance, A. dawsoni lacks an official IUCN category but is noted for localized conservation concerns.^[47] Population trends are largely unknown or stable where monitored, with no widespread evidence of global declines but indications of regional pressures in arid habitats.^[44] Ongoing monitoring occurs through citizen science platforms like iNaturalist, which has documented thousands of observations across species ranges, and targeted regional surveys in Australia that track emergence events and habitat occupancy.^[48]^[49]

Human impacts

Human activities pose significant threats to Amegilla populations, primarily through habitat alteration and chemical exposure. Agriculture and urbanization have led to extensive land clearing in arid and semi-arid regions where these ground-nesting bees thrive, fragmenting nesting sites and reducing access to suitable soils for burrowing. For instance, development for farming and urban expansion in Australia removes bare ground and native vegetation essential for Amegilla species like the blue-banded bee (A. cingulata), disrupting their solitary nesting habits and limiting population connectivity.^[26]^[50] Pesticide use further exacerbates these pressures, as foraging Amegilla bees encounter residues on flowers during pollination activities, leading to direct toxicity and sublethal effects on navigation and reproduction. Insecticides applied in agricultural fields have been linked to elevated mortality rates among wild bees, including native Australian species, with exposure reducing foraging efficiency and overall colony viability in treated areas. Herbicides compound the issue by eliminating wildflowers, depriving bees of nectar and pollen sources critical for their buzz-pollination role in crops like tomatoes.^[50]^[26] Climate change intensifies these anthropogenic impacts by altering phenological synchrony between Amegilla bees and their floral resources, as shifting flowering times due to warmer temperatures mismatch peak foraging periods. Increased droughts in arid habitats dry out soils, making it harder for these bees to excavate nests and provision brood, potentially leading to higher larval mortality and reduced reproductive success. In Australia, such changes threaten the persistence of species adapted to stable semi-arid conditions, with projections indicating further habitat unsuitability under continued warming.^[50]^[26] Mitigation efforts focus on promoting Amegilla as key native pollinators and restoring degraded habitats to bolster populations. In Australia, initiatives like the Queensland Government's Biodiversity Conservation Strategy emphasize planting native flora such as grevilleas and banksias to provide year-round forage, while bee hotels offer artificial nesting sites for solitary species in urban areas. Public awareness campaigns highlight their role in crop pollination, encouraging reduced pesticide use and habitat preservation projects that reconnect fragmented arid landscapes. These actions, including weed control and feral species management by organizations like the Australian Wildlife Conservancy, aim to enhance ecosystem resilience for Amegilla bees.^[50]^[26]

References

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Apidae – euglossine, euglossines, honey bees, stingless bees. Subfamily, Apinae. Tribe, Anthophorini. Direct Children: Genus, Amegilla Friese, 1897.
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Amegilla cingulata is one of the most visually stunning bees, with a goldy thorax, and black with metallic blue bands on the abdomen.Amegilla - Zonamegilla And... · Taxonomy · Threats<|control11|><|separator|>
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an introduced buzz‐pollinating bee Amegilla pulchra (Hymenoptera ...
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### Threats to Australian Native Bees (Including Amegilla/Blue-Banded Bees)
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[33]
Foraging Behavior and Pollen Transport by Flower Visitors of the ...
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[34]
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Aug 6, 2025 · ... These bees help in the pollination of various crops and enhance the productivity of the crops. The foraging activity was observed to ...
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Dawson's Burrowing Bee (Amegilla dawsoni)
Like many other 'solitary' bees, Dawson's Bee exhibits gregarious tendencies and females usually nest in close proximity to one another. An active nesting ...
[36]
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[38]
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May 29, 2025 · An example of the Thyreus genus, which are kleptoparasites of Amegilla – fantastic pollinators, especially of plants that require sonication. As ...
[39]
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[42]
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[44]
[PDF] European Red List of Bees - IUCN Portals
Amegilla, Ancyla, Anthophora, Habropoda, Melecta and Thyreus species ... conservation status of the species in the wild. Research is also required to ...
[45]
Amegilla Asarapoda - www.anba.org.au
This subgenus contains 21 species and is widely distributed across Australia. Like the blue banded bees, they vigorously vibrate certain flowers to extract ...
[46]
Dawson's burrowing bee at risk from tourists and careless drivers ...
Aug 28, 2022 · There are calls for the bees to be protected from careless drivers; The bees are coming to the end of their life cycles, and will become active ...
[47]
Dawson's Burrowing Bee (Amegilla dawsoni) - iNaturalist
Amegilla dawsoni, sometimes called the Dawson's burrowing bee, is a species of bee that nests by the thousands in arid claypans in Western Australia.
[48]
Blue-banded and Allied Digger Bees (Genus Amegilla) - iNaturalist
Amegilla is a large genus of bees in the tribe Anthophorini. Several species have blue metallic bands on the abdomen, and are referred to as "blue-banded bees".
[49]
https://www.bushheritage.org.au/news/burrowing-bees-emerge-in-their-thousands
[50]
What's the buzz? Your guide to Australian native bees
May 20, 2025 · Habitat loss. One of the biggest threats to native bees is simply losing their homes. Land clearing for development, agriculture, and mining ...