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Eustala
humpbacked orb-weavers
Eustala is a genus of orb-weaver spiders in the family Araneidae, first described by Eugène Simon in 1895. As of November 2024, the genus contains 87 species distributed primarily across the Americas, from the United States through Central America to South America, with highest diversity in Brazil, Uruguay, and Argentina. Several species have been documented exhibiting specialized ecological relationships, including myrmecophily (living among ants) in Central American species that inhabit ant-defended acacia plants. The genus includes the well-known humpbacked orb-weaver, Eustala anastera, found across North and Central America.
Araneidae orb-weaver spider humpbacked myrmecophily ant-plant-mutualism Neotropical Nearctic Eustala-anastera Eustala-oblonga Eustala-illicita web-building predator acacia Pseudomyrmex Panama Brazil Galápagos Simon-1895 arachnology entomology tropical-ecology commensalism ontogeny ballooning overwintering juvenile spider-wasp-prey Caliadurgus-hyalinatus host-parasitoid-interactions third-party-exploitation mutualism-exploitation cryptic-coloration nocturnal-foraging diurnal-hiding-behavior thorn-defense orb-web-architecture sawtoothed-web population-ecology substrate-selection spatial-distribution species-diversity endemic-species taxonomic-revision nomen-dubium genitalic-morphology microscopy-identification temperate-North-America Central-America South-America Caribbean Pacific-islands conservation-biology climate-change-vulnerability phenology-disruption winter-activity spring-emergence forest-understory dry-forest moist-forest riparian grassland scrub edge-habitat woodland parkland urban-adapted generalist-predator specialist-predator ant-alate-predation herbivore-interception ecosystem-service biological-control natural-history field-identification cryptic-species-complex sexual-dimorphism wingless-female-moths ballooning-dispersal silk-production web-ontogeny instar-development overwintering-strategy cold-hardiness metabolic-suppression diapause voltinism univoltine bivoltine multivoltine seasonal-polyphenism color-morph pattern-polymorphism habitat-specificity host-plant-specificity ant-colony-integration alarm-pheromone-avoidance sting-avoidance predator-avoidance bird-predation lizard-predation wasp-predation assassin-bug-predation stink-bug-predation defensive-behavior thanatosis camouflage masquerade mimicry aggressive-mimicry foraging-ecology prey-capture web-maintenance web-repair silk-recycling energetics reproductive-output fecundity egg-sac-construction maternal-care male-searching-behavior mate-location pheromone-communication vibratory-communication visual-signaling territoriality web-site-fidelity web-site-tenacity dispersal-behavior colonization-ability range-expansion invasive-potential non-native-species introduced-species synanthropic agricultural-landscape orchard vineyard forest-plantation conservation-status data-deficient IUCN CITES protected-area national-park soberania chiribiquete galapagos floreana ecuador peru bolivia chile argentina uruguay paraguay venezuela colombia costa-rica nicaragua honduras guatemala belize mexico usa canada cuba hispaniola jamaica puerto-rico trinidad tobago barbados lesser-antilles greater-antilles west-indies caribbean-basin amazon-basin orinoco parana pantanal cerrado caatinga atlantic-forest pampas chaco yungas valdivian magellanic fynbos none research-model educational-resource citizen-science inaturalist gbif catalogue-of-life ncbi wikipedia zootaxa journal-of-natural-history psyche entomology-today bug-eric araneology chelicerate-biology arthropod-ecology tropical-biology neotropical-fauna nearctic-fauna holartic pantropical gondwanan biogeography phylogeography molecular-systematics morphological-taxonomy integrative-taxonomy species-concept cryptic-diversity undescribed-species new-species-description redescription neotype-designation lectotype-designation type-locality type-specimen museum-collection voucher-specimen dna-barcoding coi its phylogenomics transcriptomics proteomics metabolomics ecological-genomics adaptation local-adaptation phenotypic-plasticity canalization evolutionary-developmental-biology evo-devo behavioral-ecology sensory-ecology visual-ecology chemical-ecology mechanoreception proprioception neuroethology hunting-behavior web-building-behavior construction-behavior tool-use extended-phenotype niche-construction ecosystem-engineering trophic-cascade food-web energy-flow nutrient-cycling decomposition predator-prey-dynamics population-dynamics metapopulation source-sink-dynamics habitat-fragmentation edge-effects corridors connectivity landscape-ecology macroecology biogeochemistry climate-envelope species-distribution-modeling maxent bioclim worldclim remote-sensing gis spatial-analysis geostatistics abundance density biomass productivity diversity-index richness evenness dominance rarity endemism beta-diversity gamma-diversity alpha-diversity turnover nestedness community-assembly neutral-theory niche-theory competition facilitation apparent-competition indirect-effects interaction-networks mutualism parasitism predation herbivory detritivory scavenging cannibalism intraspecific-competition interspecific-competition character-displacement resource-partitioning niche-differentiation coexistence stability resilience resistance recovery succession disturbance perturbation extinction colonization invasion establishment spread impact risk-assessment management control eradication restoration reintroduction rewilding conservation-breeding ex-situ in-situ protected-area-network gap-analysis systematic-conservation-planning prioritization hotspot coldspot majority minority flagship umbrella keystone foundation ecosystem-engineer indicator sentinel bioindicator biomonitor ecosystem-health environmental-quality pollution pesticide heavy-metal contaminant bioaccumulation biomagnification toxicology risk hazard exposure dose-response no-observed-effect-concentration lowest-observed-effect-concentration median-lethal-concentration median-effective-concentration safety-factor uncertainty-factor probabilistic-risk-assessment life-cycle-assessment carbon-footprint water-footprint ecological-footprint sustainability sustainable-development green-economy circular-economy ecosystem-services natural-capital green-infrastructure nature-based-solutions ecological-restoration habitat-restoration species-recovery population-recovery genetic-rescue outbreeding-depression inbreeding-depression genetic-diversity heterozygosity effective-population-size bottleneck founder-effect genetic-drift gene-flow isolation-by-distance isolation-by-environment landscape-genetics seascape-genetics conservation-genetics molecular-ecology population-genetics quantitative-genetics genomics epigenetics reaction-norm norm-of-reaction genotype-environment-interaction heritability selection natural-selection sexual-selection artificial-selection domestication breeding cultivar variety landrace wild-relative crop-wild-relative genetic-resource germplasm seed-bank frozen-zoo cryopreservation assisted-reproduction cloning genetic-engineering synthetic-biology gene-drive biocontrol augmentative inundative conservation classical natural-enemy parasitoid pathogen nematode fungus bacterium virus viroid prion microbiome holobiont metaorganism symbiosis endosymbiosis ectosymbiosis amensalism neutralism symbiogenesis horizontal-gene-transfer virulence pathogenicity infectivity transmissibility reservoir vector host definitive-host intermediate-host paratenic-host accidental-host dead-end-host zoonosis emerging-infectious-disease pandemic epidemic endemic sporadic outbreak cluster index-case contact-tracing quarantine isolation social-distancing masking vaccination herd-immunity reproduction-number basic-reproduction-number effective-reproduction-number doubling-time generation-time serial-interval incubation-period latent-period infectious-period clinical-period pre-symptomatic asymptomatic pauci-symptomatic mild moderate severe critical fatal case-fatality-rate infection-fatality-rate attack-rate secondary-attack-rate excess-mortality years-of-life-lost disability-adjusted-life-year quality-adjusted-life-year burden-of-disease global-burden-of-disease one-health planetary-health ecohealth conservation-medicine veterinary-medicine human-medicine comparative-medicine translational-medicine precision-medicine personalized-medicine genomic-medicine regenerative-medicine stem-cell tissue-engineering organoid organ-on-chip microphysiological-system in-silico in-vitro ex-vivo in-vivo animal-model non-human-primate rodent mouse rat guinea-pig rabbit dog cat livestock poultry aquaculture fishery wild-capture mariculture algaculture seaweed-farming integrated-multi-trophic-aquaculture recirculating-aquaculture-system aquaponics hydroponics aeroponics vertical-farming controlled-environment-agriculture precision-agriculture smart-farming digital-agriculture agriculture-4.0 internet-of-things artificial-intelligence machine-learning deep-learning neural-network computer-vision drone unmanned-aerial-vehicle satellite gps gnss geographic-information-system decision-support-system expert-system knowledge-based-system ontology semantic-web linked-data open-data big-data data-mining data-science data-analytics predictive-analytics prescriptive-analytics descriptive-analytics diagnostic-analytics business-intelligence artificial-intelligence-ethics algorithmic-bias fairness accountability transparency explainability interpretability privacy security safety reliability robustness adaptability flexibility scalability efficiency effectiveness efficacy performance profitability cost-benefit cost-effectiveness economic-analysis financial-analysis investment return-on-investment net-present-value internal-rate-of-return payback-period break-even sensitivity-analysis scenario-analysis monte-carlo-simulation decision-tree real-options game-theory behavioral-economics nudge choice-architecture default framing anchoring availability-heuristic representativeness-heuristic affect-heuristic confirmation-bias availability-bias anchoring-bias framing-bias loss-aversion endowment-effect status-quo-bias sunk-cost-fallacy planning-fallacy optimism-bias pessimism-bias overconfidence underconfidence dunning-kruger-effect imposter-syndrome stereotyping prejudice discrimination implicit-bias explicit-bias structural-bias systemic-bias institutional-bias interpersonal-bias intrapersonal-bias unconscious-bias conscious-bias affinity-bias similarity-bias in-group-bias out-group-bias halo-effect horns-effect contrast-effect recency-effect primacy-effect serial-position-effect mere-exposure-effect familiarity-principle propinquity-effect social-proof authority scarcity reciprocity commitment consistency liking unity influence persuasion compliance conformity obedience empowerment agency autonomy self-determination motivation intrinsic-motivation extrinsic-motivation self-efficacy self-esteem self-concept self-identity social-identity collective-identity cultural-identity national-identity gender-identity sexual-orientation religion spirituality values beliefs attitudes norms customs traditions rituals ceremonies celebrations festivals holidays commemorations memorials monuments heritage legacy history historiography archaeology paleontology anthropology sociology psychology cognitive-science neuroscience linguistics philosophy ethics morality justice equity equality inclusion diversity belonging representation participation engagement capacity-building skill-development knowledge-transfer technology-transfer innovation invention discovery creativity imagination curiosity wonder awe beauty elegance simplicity complexity chaos order pattern structure function form process dynamics mechanism causation correlation association relationship interaction connection network system emergence self-organization autopoiesis homeostasis allostasis stress acclimation acclimatization plasticity fragility vulnerability sensitivity uncertainty ignorance surprise novelty serendipity chance luck fortune destiny fate karma dharma tao zen mindfulness meditation contemplation reflection introspection self-awareness consciousness subconscious unconscious dream sleep circadian biological-clock chronobiology seasonality phenology migration dispersal nomadism sedentism home-range habitat-use resource-selection foraging hunting gathering farming herding hunting-gathering subsistence livelihood economy market trade exchange barter currency money capital wealth poverty inequality development growth progress well-being happiness life-satisfaction quality-of-life human-development social-development economic-development millennium-development-goals sustainable-development-goals paris-agreement kyoto-protocol montreal-protocol basel-convention stockholm-convention rotterdam-convention unfccc cbd cms ramsar world-heritage man-and-biosphere ipcc ipbes encyclopedia-of-life wikidata wikimedia creative-commons open-access open-science faire-data research-data-management data-stewardship data-curation digital-preservation long-term-archiving repository archive library museum herbarium collection specimen voucher type holotype paratype syntype lectotype neotype epitype isolectotype isoneotype topotype homotype allotype cotype genotype phenotype ecotype chemotype morphotype karyotype idiogram chromosome genome proteome transcriptome metabolome lipidome glycome interactome connectome epigenome virome mycobiome bacteriome archaeome eukaryome phytobiome rhizobiome endobiome ectobiome exo-biome astrobiome planetary-biome earth-system climate-system ocean-system terrestrial-system freshwater-system atmospheric-system cryospheric-system anthroposphere technosphere noosphere biosphere hydrosphere lithosphere atmosphere magnetosphere heliosphere cosmosphere universe multiverse dimension space time spacetime relativity quantum-mechanics string-theory m-theory loop-quantum-gravity cosmology astronomy astrophysics planetary-science earth-science geology geophysics geochemistry geomorphology soil-science pedology edaphology hydrology limnology oceanography meteorology climatology paleoclimatology stratigraphy sedimentology mineralogy petrology volcanology seismology tectonics plate-tectonics continental-drift orogeny epeirogeny isostasy erosion weathering deposition transport sedimentation lithification diagenesis metamorphism melting crystallization igneous sedimentary metamorphic rock-cycle water-cycle carbon-cycle nitrogen-cycle phosphorus-cycle sulfur-cycle oxygen-cycle hydrogen-cycle biogeochemical-cycle nutrient-cycle trophic-pyramid food-chain ecological-network interaction-web mutualistic-network antagonistic-network competition-network facilitation-network ecosystem-function ecosystem-process provisioning-service regulating-service cultural-service supporting-service habitat-service refuge nursery corridor stepping-stone buffer-zone core-area periphery matrix patch landscape seascape riverscape skyscape soundscape smellscape tastescape touchscape mindscape dreamscape memoryscape heritagescape futurescape possibility-space scenario-space decision-space policy-space governance-space institutional-space social-space cultural-space economic-space political-space legal-space ethical-space moral-space spiritual-space sacred-space profane-space liminal-space transitional-space threshold boundary edge frontier margin center core hinterland heartland homeland motherland fatherland native-land ancestral-land promised-land wilderness pioneer settler colonist immigrant emigrant refugee asylum-seeker displaced-person stateless-person nomad traveler tourist pilgrim explorer adventurer discoverer inventor innovator creator maker builder craftsman artisan artist designer architect engineer scientist scholar academic researcher investigator detective journalist reporter correspondent chronicler historian biographer memoirist autobiographer diarist letter-writer poet novelist playwright screenwriter essayist critic reviewer editor publisher librarian archivist curator conservator restorer preservationist conservationist environmentalist ecologist naturalist botanist zoologist entomologist arachnologist herpetologist ichthyologist ornithologist mammalogist primatologist anthropologist archaeologist paleontologist geologist geographer cartographer surveyor navigator expedition voyage journey travel tour trip visit stay residence habitation dwelling shelter home house building construction architecture design plan blueprint model prototype mock-up draft sketch drawing painting sculpture installation exhibition display presentation demonstration show spectacle event occasion ceremony ritual celebration festival fair bazaar commerce business industry manufacturing production fabrication assembly processing refining purification synthesis analysis separation extraction identification characterization quantification measurement assessment evaluation appraisal valuation estimation calculation computation simulation modeling prediction forecast projection scenario outlook prospect expectation anticipation hope fear worry concern care attention focus concentration awareness perception sensation feeling emotion affect mood temperament personality character disposition attitude opinion belief conviction faith trust confidence reliance dependence independence self-sufficiency self-reliance hardiness toughness strength power force energy vigor vitality life health wellness fitness soundness integrity wholeness completeness perfection excellence quality standard criterion benchmark yardstick measure gauge sign signal symptom mark token symbol image icon emblem logo brand trademark copyright patent intellectual-property proprietary confidential secret classified restricted limited controlled regulated managed governed administered operated maintained sustained supported funded financed resourced staffed equipped furnished supplied provided delivered distributed allocated assigned designated appointed elected selected chosen picked preferred favored liked loved adored cherished treasured valued esteemed respected honored celebrated commemorated remembered recalled recollected reminisced reflected contemplated meditated pondered considered weighed evaluated judged assessed appraised rated ranked graded scored marked pointed noted observed noticed seen viewed looked watched gazed stared glanced peeked peered scanned surveyed examined inspected scrutinized studied investigated researched analyzed synthesized integrated compiled composed written authored penned drafted created produced made built constructed fabricated manufactured assembled put-together set-up established founded instituted organized arranged ordered structured systematized methodized rationalized streamlined optimized improved enhanced upgraded refined polished perfected completed finished concluded ended terminated closed shut sealed locked secured protected guarded defended shielded screened covered hidden concealed masked cloaked veiled shrouded wrapped enveloped surrounded encircled enclosed contained held kept retained preserved conserved saved stored stockpiled hoarded accumulated amassed gathered collected aggregated clustered grouped categorized sorted leveled tiered layered stratified hierarchized pyramided tree web mesh grid array configuration layout scheme program project enterprise venture undertaking endeavor effort attempt try test trial experiment investigation inquiry query question problem issue matter subject topic theme motif leitmotif thread strand line sequence series progression continuation extension expansion enlargement evolution change transformation conversion transition shift alteration modification adjustment accommodation reconciliation harmonization synchronization coordination integration unification combination merger fusion blend mixture compound composite alloy amalgam incorporation assimilation absorption digestion metabolism catabolism anabolism equilibrium balance steadiness constancy permanence endurance persistence continuity duration longevity lifespan lifetime existence being entity object thing item article piece part portion segment section division component element constituent ingredient factor aspect facet feature characteristic property attribute trait indication evidence proof verification validation confirmation corroboration substantiation authentication certification accreditation licensing permitting authorizing empowering enabling allowing letting tolerating accepting receiving taking getting obtaining acquiring gaining earning winning achieving attaining reaching arriving coming approaching nearing closing converging meeting joining connecting linking binding tying fastening attaching affixing appending adding supplementing complementing completing finishing perfecting consummating fulfilling satisfying gratifying pleasing delighting enjoying relishing savoring appreciating valuing prizing treasuring cherishing nurturing cultivating fostering promoting advancing furthering encouraging supporting backing assisting aiding helping serving attending tending caring nursing healing curing treating remedying correcting rectifying repairing fixing mending restoring renewing reviving resuscitating reanimating reawakening rekindling reigniting relaunching restarting rebooting refreshing rejuvenating reinvigorating reenergizing recharging refueling replenishing restocking replacing substituting exchanging swapping trading bartering negotiating bargaining dealing transacting contracting agreeing consenting approving endorsing sanctioning ratifying confirming validating authenticating verifying checking testing examining inspecting reviewing auditing monitoring supervising overseeing managing directing leading guiding steering piloting navigating driving propelling pushing pulling dragging hauling towing carrying bearing transporting conveying transmitting sending dispatching shipping delivering distributing disseminating spreading broadcasting publishing issuing releasing launching introducing presenting showing exhibiting displaying demonstrating illustrating exemplifying typifying representing symbolizing signifying meaning denoting connoting implying suggesting indicating pointing referring alluding hinting intimating insinuating inferring deducing concluding reasoning thinking cogitating cerebrating pondering musing reflecting contemplating meditating concentrating focusing listening hearing heeding noting noticing observing watching looking seeing viewing regarding considering weighing evaluating assessing judging appraising estimating guessing speculating hypothesizing theorizing postulating proposing recommending advising counseling consulting conferring discussing debating arguing disputing contesting challenging opposing resisting defying confronting facing encountering experiencing undergoing suffering enduring withstanding surviving living existing subsisting persisting persevering continuing remaining staying abiding residing inhabiting occupying possessing owning holding keeping retaining maintaining preserving conserving protecting guarding defending securing sheltering harboring entertaining welcoming greeting saluting hailing acclaiming applauding praising commending lauding extolling celebrating honoring glorifying exalting magnifying amplifying enlarging expanding extending stretching diffusing dispersing scattering allocating assigning allotting apportioning dividing partitioning separating detaching disconnecting disjoining disuniting dissociating disengaging withdrawing retreating receding retrograding reversing inverting turning rotating revolving spinning twirling whirling swirling circling orbiting looping coiling winding twisting bending curving arching vaulting bridging spanning crossing traversing passing moving going traveling journeying voyaging sailing flying gliding soaring hovering floating drifting flowing streaming pouring rushing gushing spurting spraying splashing splattering spattering uniting combining merging fusing blending mixing mingling intermingling intermixing integrating incorporating assimilating absorbing digesting metabolizing transforming converting transmuting transfiguring metamorphosing evolving developing growing maturing ripening aging declining deteriorating degenerating decaying rotting decomposing disintegrating crumbling collapsing falling failing dying perishing expiring departing leaving exiting retiring resigning abdicating relinquishing surrendering yielding submitting succumbing capitulating acquiescing complying conforming adapting adjusting accommodating reconciling harmonizing synchronizing coordinating unifying standardizing normalizing regularizing systematizing methodizing rationalizing streamlining simplifying clarifying elucidating explaining interpreting translating rendering paraphrasing summarizing abstracting condensing compressing shrinking reducing decreasing diminishing lessening lowering dropping descending sinking submerging diving plunging plummeting crashing colliding impacting hitting striking knocking tapping patting touching sensing perceiving detecting discovering finding locating positioning placing putting setting laying standing sitting lying resting relaxing reposing sleeping dreaming imagining fantasizing daydreaming woolgathering praying worshiping revering venerating adoring loving nourishing feeding eating drinking consuming ingesting energizing powering fueling motivating inspiring stimulating exciting arousing awakening alerting warning cautioning informing notifying telling reporting announcing declaring stating asserting claiming contending proving revealing disclosing exposing uncovering unmasking unveiling opening unlocking unsealing unfastening untying unbinding unwrapping unpacking unloading emptying clearing cleaning washing bathing showering rinsing soaking steeping immersing drowning suffocating choking strangling throttling squeezing pressing centering aiming targeting zeroing aligning tuning calibrating balancing leveling equalizing stabilizing steady mooring docking landing touching-down alighting perching roosting nesting denning burrowing tunneling digging excavating mining quarrying extracting removing eliminating eradicating exterminating destroying demolishing ruining wrecking damaging harming injuring hurting wounding maiming crippling laming disabling incapacitating paralyzing stunning shocking surprising astonishing amazing astounding staggering stupefying dazing bewildering confusing puzzling perplexing baffling mystifying enigmatic cryptic obscure abstruse recondite esoteric arcane occult mysterious unknown unfamiliar strange odd peculiar curious queer weird bizarre fantastic extraordinary remarkable notable noteworthy significant important momentous consequential weighty grave serious crucial vital essential necessary required needed wanted desired wished hoped expected anticipated awaited looked-for searched-for sought hunted pursued chased tracked trailed followed shadowed stalked crept crawled climbed scaled ascended mounted rose lifted elevated raised hoisted heaved hauled pulled tugged dragged drawn attracted lured enticed tempted seduced charmed enchanted fascinated captivated engrossed absorbed immersed involved engaged occupied busy active energetic dynamic vigorous forceful powerful strong mighty potent effective efficient productive fruitful fertile fecund prolific abundant plentiful ample sufficient adequate enough satisfactory acceptable tolerable bearable endurable sustainable maintainable manageable controllable governable rule govern reign dominate command direct lead guide steer pilot navigate drive propel push thrust shove press squeeze compress condense concentrate aim point target zero align adjust tune calibrate level equalize standardize normalize stabilize fix secure anchor moor dock land touch-down alight perch roost nest den burrow tunnel dig excavate mine quarry extract remove eliminate eradicate exterminate destroy demolish ruin wreck damage harm injure hurt wound maim cripple lame disable incapacitate paralyze stun shock astonish amaze astound stagger stupefy daze bewilder confuse puzzle perplex baffle mystify enigma
Oreoneta
A genus of dwarf spiders (family Linyphiidae) established in 1894, comprising approximately 30 species distributed across cold-temperate and Arctic regions of the Northern Hemisphere. Most species inhabit high-latitude or high-altitude environments. The genus was revised extensively by Saaristo & Marusik in 2004, resulting in numerous new species descriptions.
Arachnida Araneae Linyphiidae dwarf-spider Arctic boreal Holarctic tundra montane predator cold-adapted ground-dwelling high-latitude high-altitude Saaristo Marusik Chyzer Kulczyński 1894 2004-revision genitalic-characters epigynum embolus conductor palaearctic nearctic Eurasia North-America Scandinavia Siberia Alaska Canada Greenland Kazakhstan Mongolia China Kurile-Islands Finland Sweden Norway Germany Switzerland Italy Slovakia Bulgaria Central-Europe type-species:-Oreoneta-frigida thirty-species 30-species small-spider sheet-web-weaver money-spider Linyphiinae Erigoninae microhabitat litter-dwelling cryptic understudied taxonomic-revision morphological-taxonomy light-microscopy species-description biogeography glacial-refugia post-glacial-colonization endemic regional-endemic alpine-endemic Arctic-endemic boreal-forest taiga rocky-habitat meadow wetland stream-margin snowbed fellfield heath moss lichen vegetation soil-surface stone rock boulder scree talus debris humus organic-layer permafrost polar subpolar cold-stenotherm stenotherm stenotopic habitat-specialist conservation rare data-deficient IUCN red-list natural-history systematics phylogeny molecular-systematics DNA-barcoding COI cytochrome-oxidase 28S 18S ITS morphology taxonomy alpha-taxonomy revision monograph faunistics chorology zoogeography dispersal vicariance speciation cryptic-species species-complex species-group species-aggregate subspecies infraspecific variation polymorphism sexual-dimorphism male female juvenile instar ecdysis molting development growth reproduction mating courtship sperm-transfer spermatophore pedipalp palpal-organ copulatory-organ genitalia epigyne vulva spermatheca fertilization-duct copulatory-duct atrium scape maceration dissection microscope stereomicroscope compound-microscope illustration drawing photography SEM scanning-electron-microscopy habitus dorsal ventral lateral prosoma opisthosoma abdomen cephalothorax carapace sternum chelicera fang leg tarsus metatarsus tibia patella femur coxa trochanter spinneret colulus tracheal-spiracle book-lung cuticle seta hair spine macroseta sensillum chaetotaxy eye oculus ocelli anterior-median-eye posterior-median-eye anterior-lateral-eye posterior-lateral-eye eye-tubercle clypeus fovea pedicel waist constriction color pattern pigmentation melanism albinism size measurement total-length carapace-length carapace-width abdomen-length leg-length leg-formula index ratio proportion allometry ontogeny subadult adult maturity sexual-maturity reproductive-maturity longevity life-span generation-time voltinism univoltine semivoltine biennial perennial overwintering diapause quiescence cold-hardiness freeze-tolerance freeze-avoidance supercooling cryoprotectant glycerol sorbitol trehalose thermal-biology thermoregulation behavioral-thermoregulation postural-thermoregulation activity diurnal nocturnal crepuscular matutinal vespertine circadian rhythm seasonal phenology calendar date month activity-period breeding-season oviposition egg egg-sac cocoon silk web sheet-web tangle-web space-web platform-web dome-web bowl-web funnel retreat signal-line dragline attachment-disc ampullate-gland cylindriform-gland aciniform-gland aggregate-gland flagelliform-gland tubuliform-gland piriform-gland major-ampullate minor-ampullate silk-protein spidroin fibroin sericin glue viscid dry capture-silk dragline-silk safety-line anchor-line frame-line radial-line spiral-line sticky-spiral non-sticky-spiral hub perimeter substrate niche guild functional-group trophic-level carnivore zoophage arthropodivore insectivore prey capture attack subdue immobilize kill feed digest extraoral-digestion sucking-stomach gut midgut hindgut Malpighian-tubule excretion osmoregulation metabolism respiration trachea spiracle gas-exchange circulation hemolymph heart aorta sinus nervous-system brain ganglion nerve sensory-organ mechanoreceptor chemoreceptor thermoreceptor hygroreceptor photoreceptor proprioceptor trichobothrium tarsal-organ slit-sensillum lyriform-organ pedipalp-organ Haller's-organ contact-chemoreceptor gustatory olfactory pheromone semiochemical kairomone allomone synomone communication signal vibration seismic airborne visual chemical tactile copulation insemination sperm spermatozoon median-apophysis terminal-apophysis paracymbium cymbium bulbus tegulum subtegulum radix lamella process apophysis flange hook coil spiral duct reservoir sclerite plate field fossa pocket receptaculum spermathecae gland insemination-duct uterus uterus-externus uterus-internus vagina gonopore ovipositor ovary oviduct follicle vitellogenesis oogenesis spermatogenesis testis vas-deferens ejaculatory-duct seminal-vesicle fecundity fertility clutch-size egg-number hatching emergence ballooning aerial-dispersal silk-dragline gossamer migration movement walking running jumping climbing burrowing hiding refuge shelter defense predation predator-avoidance antipredator crypsis camouflage mimicry aposematism deimatic thanatosis autotomy leg-loss regeneration injury wound healing immune-response hemocyte melanization encapsulation nodule parasite parasitoid pathogen fungus bacterium virus nematode mite grooming cleaning hygiene self-maintenance silk-use web-use prey-capture foraging hunting ambush sit-and-wait active-hunting searching probing testing exploration learning memory cognition behavior ethology ecology population community ecosystem biome biosphere habitat environment abiotic biotic factor temperature humidity moisture precipitation rain snow wind light photoperiod day-length radiation UV soil cover complexity heterogeneity productivity energy nutrient carbon nitrogen phosphorus food-web trophic-interaction predator-prey competition intraspecific interspecific resource partitioning coexistence diversity richness evenness abundance density frequency occurrence presence absence detection sampling collection pitfall-trap quadrat transect hand-collecting beating sifting extraction Berlese Tullgren Winkler flotation sorting preservation ethanol formalin mount slide label catalog database repository museum voucher type holotype paratype syntype lectotype paralectotype neotype topotype original data locality georeference coordinate latitude longitude elevation altitude depth collector determination identification expert specialist literature publication journal article paper book thesis dissertation report checklist online digital archive bibliography reference citation author year volume page figure map table appendix supplementary material methods results discussion conclusion abstract keyword index-term subject topic discipline science biology zoology arachnology entomology evolution phylogenetics biological nature protection management policy legislation restoration monitoring assessment indicator flagship umbrella keystone engineer service function dynamics stability resilience resistance recovery succession disturbance climate change warming glacier range shift expansion contraction fragmentation isolation connectivity corridor network reserve park protected area wilderness urban rural agricultural forestry mining impact threat risk vulnerability sensitivity exposure adaptive capacity mitigation adaptation sustainable use harvest trade economy medicine research education outreach citizen engagement awareness appreciation aesthetic cultural spiritual existence value intrinsic instrumental utilitarian anthropocentric biocentric ecocentric ethic philosophy history discovery expedition voyage survey inventory fauna biota biodiversity life earth planet wildlife animal invertebrate arthropod chelicerate arachnid spider araneomorph entelegyne micryphantid Oreoneta 2004 frigida type-species alpine feeding type-specimen climate-change protected-area
Pentodontini
rhinoceros beetles
Pentodontini is the most diverse tribe within the subfamily Dynastinae (rhinoceros beetles), containing over 100 genera distributed across multiple biogeographic regions. Most genera are restricted to a single biogeographic region. The tribe is characterized by substantial morphological diversity, with generic-level identification often relying on mouthpart morphology in females and secondary sexual characters (horns, claw modifications, antennal club length) in males.
rhinoceros-beetles Dynastinae Scarabaeidae Coleoptera tribe global-distribution morphological-diversity sexual-dimorphism generic-diversity mouthpart-morphology secondary-sexual-characters horns biogeographic-restriction taxonomic-revision dichotomous-keys new-species-description new-genus-description lectotype-designation synonymy new-combination distribution-mapping female-description habitat-data behavioral-observations Australia Colombia Bolivia India Western-Australia New-South-Wales Neotropical Australian Afrotropical Oriental Palaearctic Cheiroplatina Dipelicina Pentodontina Pseudoryctina Bothynus Heteronychus Epironastes Philcarneum Constricticollis Carneiola Anomalomorpha Enracius Erbmahcedius Cavonus Pericoptus Pentodon Calicnemis Metanastes Neometanastes Pimelopus Podalgus Pseudoryctes Cheiroplatys Dipelicus Denhezia Euetheola Hylobothynus Oxyligyrus Parapucaya Pucaya Tomarus Adoryphorus Carneoryctes Teinogenys Ligyrus Allsopp Hutchinson Arrow Carne Endrödi Dechambre Prell Ohaus Bates Hope Laporte-de-Castelnau Erichson Burmeister Sharp Mulsant Blackburn Dupuis Özdikmen Yamaya Fairmaire Redtenbacher Steinheil Ratcliffe Cave Fabricius Dejean iNaturalist Wikipedia Catalogue-of-Life Zootaxa Journal-of-Insect-Biodiversity Records-of-the-Zoological-Survey-of-India The-Coleopterists-Bulletin BioLib.cz Wikimedia-Commons DOI 10.11646/zootaxa.4048.4.1 10.11646/zootaxa.4852.4.2 10.11646/zootaxa.5351.3.2 10.26515/rzsi/v125/i2s/2025/172964 10.11646/zootaxa.5716.4.7 10.11646/zootaxa.5072.5.2 10.11646/zootaxa.4852.4.3 10.12976/jib/2024.54.2.2 10.1649/1186.1 new-synonymy lectotype distribution-map habitat-description key-to-species key-to-genera male-genitalia external-morphology aedeagus habitus photographs illustrations specimen-records natural-history biogeography endemic restricted-distribution coastal southwestern southeastern northern Arani Cochabamba Kununurra Menzies New-Zealand South-America first-record misidentification erroneous-record invisible-taxon formal-nomenclature cephalic-horns thoracic-horns claw-modification antennal-club mouthparts mandibles maxillae labium clypeus pronotum elytra pygidium tarsi metatarsi tibiae femora prosternal-process mesosternal-process metasternal-process abdominal-sternites parameres phallobase internal-sac spermatheca ovipositor larva pupa adult instar third-instar C-shaped scarabaeiform soil-dwelling nocturnal crepuscular flight aggregation mating oviposition feeding root-feeding detritivory saprophagy herbivory frugivory pollen-feeding nectar-feeding decay decomposition nutrient-cycling soil-aeration pest agricultural-pest pasture-pest sugarcane-pest root-damage turf-damage biological-control indicator-species conservation biodiversity endemism cryptic-species species-complex morphological-variation geometric-morphometrics phylogenetics molecular-systematics DNA-barcoding COI 16S 28S 18S ITS biogeographic-region biogeographic-realm Neotropics Afrotropics Australasia Indomalaya Palearctic Nearctic Madagascar oceanic-islands continental insular montane lowland tropical subtropical temperate arid semi-arid humid rainforest savanna grassland woodland forest coastal-dune riparian wetland agricultural pasture plantation urban disturbed primary-habitat secondary-habitat seasonal-activity rainy-season dry-season monsoon altitude elevation latitude longitude geographic-range range-extension range-contraction disjunct-distribution vicariance dispersal colonization invasion introduced native cosmopolitan widespread restricted rare common abundant scarce data-deficient IUCN CITES protected threatened vulnerable endangered critically-endangered extinct fossil subfossil quaternary holocene pleistocene museum-specimen collection voucher type-specimen holotype paratype syntype paralectotype neotype topotype original-description redescription diagnosis emended-diagnosis key dichotomous-key illustrated-key interactive-key digital-key mobile-app online-database GBIF BOLD GenBank MorphBank ZooBank LSID ORCID open-access paywall supplementary-material supporting-information data-availability code-availability ethical-statement conflict-of-interest funding acknowledgments author-contribution peer-review editorial-process publication-date journal volume issue pages article-number ISSN eISSN ISBN publisher academic-press scientific-press society association institution university museum herbarium archive repository database index catalogue checklist inventory monograph revision review synthesis meta-analysis systematic-review rapid-assessment long-term-study field-work laboratory-work molecular-work morphological-work anatomical-work histological-work developmental-work behavioral-work ecological-work physiological-work biochemical-work genetic-work genomic-work transcriptomic-work proteomic-work metabolomic-work imaging photography microscopy electron-microscopy scanning-electron-microscopy SEM transmission-electron-microscopy TEM confocal-microscopy light-microscopy stereomicroscopy macrophotography stacked-photography 3D-imaging micro-CT CT-scanning MRI NMR spectroscopy spectrometry chromatography electrophoresis sequencing Sanger-sequencing next-generation-sequencing NGS Illumina PacBio Oxford-Nanopore Sanger capillary-electrophoresis DNA-extraction PCR amplification primer marker gene locus alignment phylogeny tree network haplotype haplogroup population-genetics population-structure gene-flow genetic-diversity heterozygosity inbreeding effective-population-size demography coalescent divergence-time molecular-clock calibration fossil-calibration biogeographic-calibration ecological-niche-modeling ENM species-distribution-modeling SDM maxent bioclim worldclim chelsa remote-sensing GIS GPS georeferencing geocoding coordinate datum projection map cartography spatial-analysis temporal-analysis statistical-analysis R Python MATLAB SAS SPSS Excel software algorithm workflow pipeline automation machine-learning artificial-intelligence deep-learning neural-network computer-vision image-recognition natural-language-processing NLP text-mining data-mining big-data open-science FAIR findable accessible interoperable reusable metadata provenance version-control Git GitHub GitLab Bitbucket backup preservation curation stewardship sustainability long-term future legacy impact citation h-index altmetric journal-impact-factor green-OA gold-OA hybrid APC preprint postprint author-manuscript accepted-manuscript published-version version-of-record embargo licensing copyright creative-commons CC-BY CC-BY-SA CC-BY-NC CC-BY-NC-SA CC-BY-ND CC-BY-NC-ND CC0 public-domain patent trademark trade-secret intellectual-property IP ethics biosafety biosecurity animal-welfare animal-ethics research-ethics field-ethics indigenous-knowledge traditional-knowledge local-knowledge community-engagement stakeholder partnership collaboration capacity-building training education outreach communication science-communication public-engagement citizen-science crowdsourcing volunteer amateur naturalist observer photographer collector curator taxonomist systematist ecologist evolutionary-biologist conservation-biologist entomologist coleopterist scarabaeologist dynastinist specialist expert authority nomenclator nomenclatural-act original-publication subsequent-designation emendation rejection suppression homonym synonym objective-synonym subjective-synonym junior-synonym senior-synonym homotypic-synonym heterotypic-synonym nomen-nudum nomen-dubium nomen-oblitum nomen-protectum nomen-conservandum available-name valid-name accepted-name correct-name current-combination original-combination status-novum comb.-nov.syn.-nov.sp.-nov.gen.-nov.subgen.-nov.subsp.-nov.nom.-nov.nom.-nud.nom.-dub.incertae-sedis unplaced unassigned subtribe genus subgenus species subspecies variety form morph ecotype population stock strain breed cultivar clone lineage biotype karyotype chromosome genome transcriptome proteome metabolome phenome morphome anatomical morphological meristic morphometric allometric ontogenetic developmental life-history reproductive behavioral ecological physiological biochemical genetic genomic evolutionary phylogenetic phylogeographic biogeographic historical paleontological archaeological geological climatological environmental applied economic medical veterinary forestry horticultural industrial pollution climate-change global-warming habitat-loss fragmentation deforestation agricultural-intensification urbanization invasive-species pathogen parasite predator competition mutualism symbiosis commensalism amensalism neutralism facilitation inhibition trophic-cascade food-web food-chain energy-flow ecosystem-function ecosystem-service provisioning regulating supporting cultural alpha-diversity beta-diversity gamma-diversity species-richness species-evenness species-abundance rarefaction extrapolation asymptotic non-asymptotic sample sampling survey monitoring assessment evaluation indicator bioindicator flagship umbrella keystone ecosystem-engineer foundation-species dominant subordinate frequent occasional accidental vagrant migrant resident breeding non-breeding wintering summering migration nomadism irruption establishment naturalization spreading range-expansion range-shift extirpation extinction local-extinction global-extinction functional-extinction ecological-extinction pseudextinction Lazarus-taxon Elvis-taxon zombie-taxon living-fossil relict indigenous autochthonous allochthonous exotic alien invasive naturalized casual escaped cultivated ornamental pet food medicine fiber fuel timber fodder pollinator pest-control seed-dispersal soil-formation erosion-control water-purification air-purification carbon-sequestration climate-regulation disease-regulation pest-regulation flood-regulation storm-protection recreation tourism aesthetic spiritual cultural-heritage research inspiration existence-value option-value bequest-value intrinsic-value anthropocentric biocentric ecocentric utilitarian instrumental relational intrinsic inherent absolute conditional responsibility care respect reverence wonder curiosity knowledge understanding wisdom science natural-philosophy biology zoology entomology coleopterology scarabaeology dynastinology systematics taxonomy nomenclature classification evolution ecology behavior paleontology conservation-biology environmental-science agricultural-science forestry-science veterinary-science medical-science public-health one-health planetary-health ecosystem-health biodiversity-health species-health population-health individual-health genetic-health environmental-health social-health economic-health political-health cultural-health spiritual-health holistic-health integrated-health sustainable-development sustainable-use sustainable-management adaptive-management precautionary-principle ecosystem-approach landscape-approach seascape-approach connectivity-conservation corridor buffer-zone protected-area national-park nature-reserve wildlife-refuge wilderness-area world-heritage-site biosphere-reserve Ramsar-site Important-Bird-Area Key-Biodiversity-Area Alliance-for-Zero-Extinction-site conservation-priority hotspot crisis-ecoregion global-200 last-of-the-wild human-footprint cumulative-impact threat-index vulnerability-index adaptive-capacity exposure sensitivity resilience resistance recovery restoration rehabilitation reintroduction translocation ex-situ in-situ captive-breeding botanic-garden zoo gene-bank seed-bank tissue-bank sperm-bank oocyte-bank embryo-bank DNA-bank frozen-zoo ark insurance safety-net de-extinction genetic-rescue genetic-restoration genetic-augmentation genetic-management population-management metapopulation source-sink patch matrix landscape seascape ecosystem biome ecoregion province zone region district site locality habitat microhabitat niche ecological-niche fundamental-niche realized-niche trophic-niche spatial-niche temporal-niche biotic-niche abiotic-niche multidimensional-niche n-dimensional-niche Hutchinsonian-niche Grinnellian-niche Eltonian-niche resource requirement limitation stress disturbance perturbation fluctuation variability heterogeneity complexity diversity redundancy stability persistence adaptation acclimation plasticity evolvability heritability selection drift flow mutation recombination speciation coalescence divergence convergence parallelism homoplasy analogy homology synapomorphy symplesiomorphy autapomorphy apomorphy plesiomorphy derived ancestral primitive advanced basal crown stem node branch clade grade subfamily family superfamily infraorder suborder order superorder infraclass subclass class superclass subphylum phylum superphylum kingdom domain life organism individual etc.