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Cyphomyrmex
fungus-growing ants
Cyphomyrmex is a genus of small, drab-colored fungus-growing ants in the tribe Attini, found primarily in the Neotropics. These ants cultivate fungi in the tribe Leucocoprineae as their primary food source, with most species growing fungal nodules rather than full mycelial gardens. Colonies are typically monogynous and small, rarely exceeding 500 workers. The genus is divided into two species complexes: the strigatus complex (South America only) and the rimosus complex (southern North America to South America). Cyphomyrmex represents a basal lineage among attine ants and serves as sister group to Mycetophylax.
fungus-growing-ants attine-ants Myrmicinae Neotropical mutualism fungiculture Leucocoprineae monogyny parasitoid-wasps social-parasitism chemical-ecology alkaloid-venom nest-architecture behavioral-plasticity cytogenetics chromosome-evolution Guiana-Shield Brazil Panama Puerto-Rico Mesoamerica Argentina United-States Texas California Florida yeast-gardens mycelial-gardens rimosus-complex strigatus-complex age-polyethism trophallaxis thanatosis feigning-death Diapriidae Acanthopria Mimopriella Megalomyrmex social-parasite C.-longiscapus C.-transversus C.-laevigatus C.-rimosus C.-minutus C.-lectus C.-cornutus C.-costatus pheromones 3-octanol farnesenes trail-following alarm-signal diketopiperazines antifungal-compounds defensive-behavior colony-defense flight-response recruitment Lanchester's-laws karyotype chromosome-number rDNA microsatellites FISH chamber-depth gallery-length fire-disturbance savanna-forest-transition arboreal-nesting clay-auricles soil-nests wood-nests epiphytic-nests moss-nests coconut-nests desiccation-susceptibility moist-habitats basal-attine phylogenetics sister-group chemical-evolution venom-alkaloids behavioral-manipulation host-parasite-interaction ecosystem-engineering nutrient-cycling decomposition organic-matter insect-feces exoskeleton-substrate garden-transplantation worker-polymorphism queen-number dealate-queens alates brood-care ecdysis-assistance foraging sugar-collection plant-exudates fungal-domestication nodule-cultivation mycelium-cultivation agricultural-behavior nest-entrance funnel-structure exposed-gardens symbiont-diversity geographic-variation research-model evolutionary-biology mutualism-evolution ant-fungus-coevolution chemical-communication social-insects eusociality colony-size small-colonies slow-movement crypsis debris-mimicry predation-avoidance parasitoid-defense wasp-killing colony-abandonment resource-rescue thief-ant-defense venom-resistance toxicity-tolerance behavioral-suppression playing-dead alkaloid-manipulation fungal-secondary-metabolites antibacterial-compounds antiviral-compounds garden-protection microbial-defense nest-microclimate temperature-regulation humidity-requirements depth-effects fire-adaptation disturbance-ecology habitat-specificity substrate-specialization clay-construction soil-manipulation architectural-plasticity vertical-nesting embankment-nesting overhang-nesting pendant-nests swallow-nest-mimicry research-accessibility field-biology tropical-ecology Neotropical-myrmecology biodiversity conservation invasive-species-context native-species biological-control Integrated-Pest-Management agricultural-ecology forest-ecology savanna-ecology transitional-habitats disturbed-areas urban-ecology anthropogenic-impact climate-sensitivity habitat-limitation range-boundaries dispersal colonization establishment population-dynamics demography colony-density nest-spacing intraspecific-competition interspecific-competition territoriality resource-defense combat-behavior aggression-levels recruitment-behavior chemical-signaling alarm-pheromones trail-pheromones recognition-cues cuticular-hydrocarbons venom-composition defensive-secretions mandibular-glands Dufour's-gland metapleural-glands antibiotic-production grooming-behavior fungal-inoculation integumental-fungi yeast-growth larval-grooming garden-maintenance substrate-preparation fecal-fertilizer anal-trophallaxis crop-regurgitation fluid-application drying-behavior nodule-formation transplantation-behavior garden-establishment ceiling-gardens root-association architectural-innovation nest-site-selection microhabitat-choice moisture-requirements thermal-buffering predator-avoidance parasite-avoidance disease-resistance social-immunity collective-behavior division-of-labor task-allocation temporal-polyethism worker-age nurse-workers forager-workers soldier-caste-absence monomorphism worker-size-variation queen-worker-differentiation reproductive-caste gyne-production male-production seasonal-reproduction colony-founding claustral-founding non-claustral-founding fungal-inoculum founding-queen first-worker-generation nutritional-dependency fat-reserves metabolic-adaptation digestive-physiology enzyme-complement fungal-enzyme lignocellulose-degradation nutritional-mutualism obligate-mutualism vertical-transmission horizontal-transmission symbiont-switching cultivar-replacement garden-failure colony-mortality population-turnover generation-time longevity survivorship life-expectancy aging senescence reproductive-investment colony-growth colony-decline budding fission reproduction-mode genetic-structure relatedness mating-system nuptial-flight mate-choice sperm-storage colony-odor nestmate-recognition foreign-rejection agonistic-behavior territorial-marking boundary-defense raiding-behavior robbing-behavior cleptobiosis dulosis slavery inquilinism guest-ants myrmecophiles nest-associates commensalism inquilines parasite-tolerance host-manipulation venom-alkaloid pyrrolidine indolizidine piperidine nonanal farnesene sesquiterpene agriculture-chemical gaster-extract chemical-diversity chemical-simplicity evolutionary-precursor derived-chemistry biosynthetic-pathway semiochemical infochemical communication information-transfer colony-organization social-structure polygyny queen-condition dealation wing-shedding ovarian-development fecundity egg-laying brood-development developmental-rate temperature-dependence humidity-dependence food-availability nutritional-quality fungal-productivity garden-yield worker-production colony-efficiency energy-allocation resource-allocation foraging-efficiency search-behavior prey-detection food-retrieval transport-behavior load-size travel-speed path-integration orientation navigation homing-behavior learning memory experience individual-recognition context-dependent-behavior threat-assessment risk-management predator-inspection alarm-propagation recruitment-activation mobilization colony-response emergency-behavior panic-response controlled-response defensive-strategy offensive-strategy escalation de-escalation conflict-resolution fight-avoidance retreat-behavior surrender-signal dominance-hierarchy reproductive-skew worker-policing queen-policing egg-eating oviposition-suppression reproductive-conflict kin-conflict worker-reproduction male-production-by-workers anarchy social-parasitism-detection parasite-recognition enemy-recognition nest-intrusion nest-defense perimeter-defense chamber-defense brood-defense queen-defense self-sacrifice suicidal-defense autothysis sting-use mandible-use spraying-behavior gaster-flexion chemical-defense mechanical-defense armor cuticle-thickness pigmentation masquerade mimicry Batesian-mimicry Muellerian-mimicry aggressive-mimicry Wasmannian-mimicry myrmecophily myrmecophagy ant-plant-mutualism trophobiosis extrafloral-nectary domatia plant-ant-interaction herbivore-defense seed-dispersal pollination ecosystem-service functional-role trophic-position detritivore decomposer saprotroph necrotroph fungivore omnivore predator scavenger trophic-level energy-flow carbon-cycling nitrogen-cycling phosphorus-cycling soil-formation bioturbation aeration water-infiltration microbial-community bacteria actinomycetes Pseudonocardia disease-suppression garden-hygiene weeding-behavior pruning-behavior fungal-grooming antibiotic-application mutualist-protection agricultural-disease crop-pathology escovopsis garden-parasite competitor-fungus pathogen-defense immune-response individual-immunity grooming allogrooming self-grooming nest-cleaning refuse-management fecal-management waste-disposal midden-formation territory-size home-range foraging-range recruitment-range exploration exploitation information-use public-information private-information cue-use signal-use communication-network interaction-network social-network colony-network population-network community-ecology species-interaction apparent-competition exploitative-competition interference-competition resource-partitioning niche-differentiation habitat-filtering environmental-gradient altitudinal-distribution latitudinal-distribution continental-distribution island-distribution endemism biogeography phylogeography historical-biogeography vicariance range-expansion range-contraction refugia glacial-cycles climate-change-impact habitat-fragmentation deforestation land-use-change agricultural-intensification urbanization pollution pesticide-exposure heavy-metal-accumulation conservation-status population-viability extinction-risk protected-area reserve-design habitat-corridor connectivity metapopulation source-sink-dynamics rescue-effect mass-effect ecological-theory behavioral-ecology evolutionary-ecology functional-ecology trait-based-ecology comparative-analysis phylogenetic-comparative-methods character-evolution ancestral-state-reconstruction convergent-evolution parallel-evolution adaptive-radiation species-diversification speciation hybridization introgression gene-flow genetic-differentiation population-structure molecular-ecology genomics transcriptomics proteomics metabolomics volatile-organic-compound cuticular-hydrocarbon glandular-secretion sting-apparatus mandible-morphology head-morphology mesosomal-morphology petiolar-morphology gastral-morphology leg-morphology antennal-morphology sensilla chemoreception mechanoreception thermoreception photoreception vision compound-eye ocelli brain-morphology neuroanatomy learning-capacity memory-capacity cognitive-ability problem-solving tool-use sequence-learning route-learning spatial-memory temporal-memory social-learning cultural-transmission tradition colony-personality behavioral-syndrome boldness exploration-tendency activity-level aggression-level foraging-tendency defensive-tendency stress-response physiological-stress thermal-stress desiccation-stress starvation-stress immune-challenge wound-healing metabolic-rate respiration energy-expenditure water-balance osmoregulation excretion Malpighian-tubules rectum hindgut midgut foregut crop proventriculus digestive-enzyme nutrient-absorption lipid-metabolism carbohydrate-metabolism protein-metabolism amino-acid-metabolism nitrogen-excretion uric-acid urea ammonia osmolyte cryoprotectant heat-shock-protein stress-protein antioxidant-defense oxidative-stress reactive-oxygen-species aging-biology senescence-theory disposable-soma antagonistic-pleiotropy mutation-accumulation life-history-theory r-K-selection bet-hedging iteroparity semelparity reproductive-effort parental-investment maternal-effect paternal-effect epigenetics DNA-methylation histone-modification non-coding-RNA gene-regulation transcription-factor signaling-pathway hormone juvenile-hormone ecdysone insulin-signaling TOR-signaling nutrient-sensing reproductive-signaling caste-determination sex-determination environmental-sex-determination genetic-sex-determination haplodiploidy arrhenotoky thelytoky parthenogenesis arrhenotokous-parthenogenesis automixis apomixis central-fusion terminal-fusion genomic-imprinting parental-conflict kin-selection inclusive-fitness Hamilton's-rule relatedness-asymmetry sex-ratio-theory local-mate-competition local-resource-competition local-resource-enhancement split-sex-ratio worker-sex-ratio queen-sex-ratio colony-sex-ratio population-sex-ratio operational-sex-ratio effective-population-size genetic-drift inbreeding inbreeding-depression outbreeding outbreeding-depression heterosis hybrid-vigor genetic-load mutation-rate effective-number-of-codons codon-usage-bias GC-content genome-size karyotype-evolution chromosomal-rearrangement fusion inversion translocation polyploidy aneuploidy chromosomal-polymorphism rDNA-cluster heterochromatin euchromatin centromere telomere nucleolus-organizer-region satellite-DNA minisatellite microsatellite simple-sequence-repeat transposable-element retrotransposon DNA-transposon horizontal-gene-transfer endosymbiont Wolbachia Cardinium Rickettsia Spiroplasma bacterial-symbiont fungal-symbiont viral-symbiont parasite pathogen disease epidemiology transmission-dynamics virulence resistance tolerance recovery immunity vaccination prophylaxis medicinal-behavior self-medication thermoregulation behavioral-thermoregulation social-thermoregulation microclimate-selection nest-temperature brood-temperature fungal-garden-temperature metabolic-heat heat-dissipation evaporative-cooling behavioral-cooling fanning ventilation chamber-design gallery-design entrance-design air-flow humidity-regulation water-collection water-storage desiccation-resistance cuticular-waterproofing spiracle-control respiratory-water-loss excretory-water-loss water-recycling metabolic-water dietary-water environmental-water rainfall-dependence seasonal-activity dry-season wet-season monsoon drought flood fire disturbance succession primary-succession secondary-succession climax-community pioneer-species habitat-specialist habitat-generalist edge-species interior-species gap-species canopy-species understory-species ground-species subterranean-species arboreal-species liana-species epiphyte-species lithophyte-species rheophyte-species aquatic-margin-species terrestrial-species fossorial-species cursorial-species scansorial-species volant-species winged-reproductive dealate ergatoid brachypterous apterous polymorphism caste-polymorphism queen-polymorphism male-polymorphism size-polymorphism allometry isometry shape-variation morphological-integration modularity evolvability constraints developmental-stability fluctuating-asymmetry directional-asymmetry antisymmetry phenotypic-plasticity reaction-norm genotype-environment-interaction grandparental-effect indirect-genetic-effect social-environment-effect colony-effect density-dependence frequency-dependence negative-frequency-dependence positive-frequency-dependence apostatic-selection anti-apostatic-selection search-image predator-learning prey-learning friend-recognition cuticular-hydrocarbon-profile gestalt-odor colony-specific-odor individual-specific-odor task-specific-odor age-specific-odor reproductive-odor queen-odor queen-pheromone primer-pheromone releaser-pheromone signal-pheromone modulatory-pheromone alarm-pheromone trail-pheromone recruitment-pheromone aggregation-pheromone sex-pheromone marking-pheromone territorial-pheromone defensive-pheromone poison-gland Dufour-gland mandibular-gland postpharyngeal-gland metapleural-gland sternal-gland tarsal-gland anal-gland rectal-gland pygidial-gland mandibular-brush tibial-spur stridulatory-organ subgenual-organ Johnston's-organ tympanal-organ hair-sensillum campaniform-sensillum basiconic-sensillum coeloconic-sensillum placoid-sensillum sensilla-trichodea sensilla-basiconica sensilla-coeloconica sensilla-ampullacea sensilla-campaniformia sensilla-placodea olfactory-receptor-neuron gustatory-receptor-neuron mechanoreceptor-neuron thermoreceptor-neuron hygroreceptor-neuron photoreceptor-neuron visual-processing olfactory-processing gustatory-processing mechanosensory-processing central-complex mushroom-body antennal-lobe optic-lobe suboesophageal-ganglion ventral-nerve-cord sympathetic-nervous-system neurosecretion neurohemal-organ corpus-cardiacum corpus-allatum prothoracic-gland ventral-gland pericardial-gland neuroendocrine-regulation hormonal-regulation pheromonal-regulation neural-regulation behavioral-regulation social-regulation environmental-regulation developmental-regulation reproductive-regulation metabolic-regulation homeostatic-regulation allostatic-regulation stress-regulation immune-regulation circadian-rhythm circannual-rhythm photoperiodism thermoperiodism biological-clock zeitgeber entrainment free-running-period phase-response-curve masking-effect arousal-threshold sleep torpor diapause quiescence dormancy aestivation hibernation cold-hardiness heat-hardiness desiccation-hardiness anoxia-hardiness starvation-hardiness radiation-hardiness pressure-hardiness gravity-perception magnetic-field-perception electric-field-perception sound-perception vibration-perception substrate-borne-signal air-borne-signal seismic-signal chemical-signal visual-signal tactile-signal auditory-signal multimodal-signal composite-signal redundant-signal amplifying-signal attenuating-signal honest-signal deceptive-signal costly-signal cheap-signal conventional-signal index-signal handicap-signal amplifier social-selection sexual-selection sperm-competition cryptic-female-choice male-male-competition female-female-competition parent-offspring-conflict sibling-conflict worker-queen-conflict worker-worker-conflict sexual-conflict antagonistic-coevolution arms-race evolutionary-dynamics community-dynamics ecosystem-dynamics biogeochemical-cycle carbon-cycle nitrogen-cycle phosphorus-cycle sulfur-cycle water-cycle trophic-transfer production-efficiency consumption-efficiency assimilation-efficiency ecological-efficiency Lindeman-efficiency 10%-rule food-chain food-web trophic-cascade top-down-control bottom-up-control wasps-control donor-control recipient-control interaction-strength interaction-web ecological-network network-topology network-metrics degree-distribution connectance nestedness compartmentalization robustness fragility resilience recovery-time return-time elasticity perturbation pulse-disturbance press-disturbance chronic-disturbance acute-disturbance catastrophic-disturbance natural-disturbance anthropogenic-disturbance habitat-destruction habitat-degradation habitat-modification habitat-loss habitat-isolation edge-effect matrix-effect permeability corridor-effect stepping-stone source-population sink-population metapopulation-dynamics extinction-debt colonization-credit relaxation-effect time-lag historical-legacy priority-effect founder-effect bottleneck-effect selection natural-selection artificial-selection directional-selection stabilizing-selection disruptive-selection balancing-selection frequency-dependent-selection density-dependent-selection r-selection K-selection c-selection s-selection a-selection bet-hedging-selection fluctuating-selection correlational-selection multivariate-selection indirect-selection correlated-response genetic-correlation phenotypic-correlation environmental-correlation correlation-matrix variance-covariance-matrix G-matrix P-matrix E-matrix M-matrix quantitative-genetics heritability narrow-sense-heritability broad-sense-heritability genetic-variance additive-genetic-variance dominance-variance epistatic-variance environmental-variance phenotypic-variance genotype-environment-interaction-variance maternal-variance paternal-variance common-environment-variance special-environment-variance measurement-error-variance response-to-selection realized-heritability predicted-response breeder's-equation selection-gradient selection-differential Lande-equation multivariate-breeder's-equation genetic-constraint evolutionary-constraint developmental-constraint functional-constraint phylogenetic-constraint historical-constraint trade-off allocation-trade-off acquisition-trade-off conversion-trade-off maintenance-trade-off reproduction-survival-trade-off growth-reproduction-trade-off immune-reproduction-trade-off stress-reproduction-trade-off foraging-predation-trade-off competition-colonization-trade-off tolerance-resistance-trade-off plasticity-canalization-trade-off generalist-specialist-trade-off speed-accuracy-trade-off exploration-exploitation-trade-off individual-colony-trade-off within-colony-trade-off between-colony-trade-off life-history-trade-off demographic-trade-off ecological-trade-off evolutionary-trade-off optimization maximization minimization satisficing marginal-value-theorem optimal-foraging-theory optimal-diet-theory central-place-foraging patch-use-theory giving-up-density giving-up-time predation-risk energetic-cost opportunity-cost handling-time search-time travel-time encounter-rate prey-density prey-profitability prey-choice diet-breadth niche-width niche-overlap niche-partitioning resource-selection habitat-selection settlement-choice nest-site-choice host-choice oviposition-choice foraging-choice path-choice route-choice central-place territory core-area edge-area boundary perimeter area volume shape configuration spatial-pattern temporal-pattern diel-pattern seasonal-pattern annual-pattern multiannual-pattern cyclical-pattern irregular-pattern random-pattern clumped-pattern uniform-pattern regular-pattern aggregated-pattern overdispersed-pattern underdispersed-pattern Poisson-distribution negative-binomial-distribution Taylor's-power-law spatial-autocorrelation temporal-autocorrelation cross-correlation spectral-analysis wavelet-analysis time-series-analysis population-time-series community-time-series ecosystem-time-series long-term-study long-term-monitoring long-term-experiment chronosequence space-for-time-substitution paleoecology paleontology fossil-record trace-fossil coprolite nest-fossil amber-inclusion subfossil quaternary holocene pleistocene neogene paleogene cretaceous mesozoic cenozoic geological-time deep-time evolutionary-time ecological-time generational-time physiological-time perception-time reaction-time decision-time memory-time learning-time developmental-time life-span colony-lifespan queen-lifespan worker-lifespan male-lifespan larval-development-time pupal-development-time egg-development-time incubation-time doubling-time half-life relaxation-time resilience-time resistance-time lag-time lead-time forecast-time prediction-time planning-time management-time conservation-time restoration-time mitigation-time adaptation-time speciation-time diversification-time extinction-time persistence-time transience-time ephemerality permanence stability instability equilibrium non-equilibrium steady-state dynamic-state transient-state asymptotic-state attractor repeller limit-cycle chaos stochasticity determinism predictability unpredictability complexity emergence self-organization pattern-formation spatial-self-organization temporal-self-organization spatiotemporal-self-organization swarm-intelligence collective-decision-making consensus-decision quorum-response wisdom-of-crowds information-cascade cultural-evolution gene-culture-coevolution niche-construction extended-phenotype extended-organism superorganism colony-as-individual colony-as-unit-of-selection group-selection multilevel-selection reciprocal-altruism by-product-mutualism amensalism competition predation herbivory parasitism parasitoidism chemical-mimicry tactile-mimicry visual-mimicry acoustic-mimicry defensive-mimicry Vavilovian-mimicry Bakerian-mimicry Gilbertian-mimicry Poultonian-mimicry Browerian-mimicry auto-mimicry intra-specific-mimicry inter-specific-mimicry molecular-mimicry cellular-mimicry tissue-mimicry organ-mimicry behavioral-mimicry reproductive-mimicry social-mimicry nest-mimicry food-mimicry predator-mimicry prey-mimicry host-mimicry guest-mimicry cleptoparasite cuckoo-parasite inquiline guest myrmecophile termitophile aphidophile coccidophile scale-insect-associate mealybug-associate whitefly-associate planthopper-associate treehopper-associate leafhopper-associate cicada-associate butterfly-associate moth-associate beetle-associate fly-associate wasp-associate bee-associate ant-associate spider-associate mite-associate tick-associate nematode-associate flatworm-associate roundworm-associate annelid-associate mollusk-associate crustacean-associate millipede-associate centipede-associate insect-associate vertebrate-associate bird-associate mammal-associate reptile-associate amphibian-associate fish-associate plant-associate fungus-associate bacteria-associate virus-associate archaea-associate protist-associate algae-associate lichen-associate bryophyte-associate pteridophyte-associate gymnosperm-associate angiosperm-associate monocot-associate dicot-associate herb-associate shrub-associate tree-associate vine-associate epiphyte-associate lithophyte-associate rheophyte-associate hydrophyte-associate xerophyte-associate mesophyte-associate hygrophyte-associate sciophyte-associate heliophyte-associate pioneer-plant-associate climax-plant-associate disturbance-plant-associate successional-plant-associate ant-plant-protection-mutualism ant-plant-food-body-mutualism ant-plant-domatia-mutualism ant-plant-extrafloral-nectary-mutualism ant-plant-seed-dispersal-mutualism ant-plant-pollination-mutualism ant-plant-agriculture-mutualism ant-fungus-mutualism ant-bacteria-mutualism ant-virus-mutualism ant-nematode-mutualism ant-mite-mutualism ant-tick-mutualism ant-spider-mutualism ant-wasp-mutualism ant-fly-mutualism ant-beetle-mutualism ant-butterfly-mutualism ant-moth-mutualism ant-bird-mutualism ant-mammal-mutualism ant-reptile-mutualism ant-amphibian-mutualism ant-fish-mutualism ant-human-mutualism ant-domestic-animal-mutualism ant-pest-mutualism ant-crop-mutualism ant-forest-mutualism ant-savanna-mutualism ant-grassland-mutualism ant-desert-mutualism ant-wetland-mutualism ant-riverine-mutualism ant-riparian-mutualism ant-montane-mutualism ant-lowland-mutualism ant-coastal-mutualism ant-island-mutualism ant-continental-mutualism ant-tropical-mutualism ant-temperate-mutualism ant-subtropical-mutualism ant-boreal-mutualism ant-austral-mutualism ant-paleotropical-mutualism ant-neotropical-mutualism ant-afrotropical-mutualism ant-oriental-mutualism ant-palearctic-mutualism ant-nearctic-mutualism ant-holarctic-mutualism ant-pan-tropical-mutualism ant-cosmopolitan-mutualism ant-endemic-mutualism ant-widespread-mutualism ant-restricted-mutualism ant-rare-mutualism ant-common-mutualism ant-abundant-mutualism ant-scarce-mutualism ant-threatened-mutualism ant-invasive-mutualism ant-native-mutualism ant-exotic-mutualism ant-introduced-mutualism ant-naturalized-mutualism ant-feral-mutualism ant-urban-mutualism ant-rural-mutualism ant-agricultural-mutualism ant-natural-mutualism ant-anthropogenic-mutualism ant-pristine-mutualism ant-degraded-mutualism ant-restored-mutualism ant-conserved-mutualism ant-managed-mutualism ant-sustainable-mutualism ant-unsustainable-mutualism ant-resilient-mutualism ant-fragile-mutualism ant-robust-mutualism ant-vulnerable-mutualism ant-resistant-mutualism ant-susceptible-mutualism ant-tolerant-mutualism ant-intolerant-mutualism ant-flexible-mutualism ant-inflexible-mutualism ant-plastic-mutualism ant-canalized-mutualism ant-adaptive-mutualism ant-maladaptive-mutualism ant-optimal-mutualism ant-suboptimal-mutualism ant-efficient-mutualism ant-inefficient-mutualism ant-productive-mutualism ant-unproductive-mutualism ant-successful-mutualism ant-unsuccessful-mutualism ant-stable-mutualism ant-unstable-mutualism ant-persistent-mutualism ant-transient-mutualism ant-obligate-mutualism ant-facultative-mutualism ant-specific-mutualism ant-generalized-mutualism ant-exclusive-mutualism ant-non-exclusive-mutualism ant-symbiotic-mutualism ant-non-symbiotic-mutualism ant-trophic-mutualism ant-defensive-mutualism ant-dispersive-mutualism ant-pollination-mutualism ant-agriculture-mutualism ant-construction-mutualism ant-nutrient-mutualism ant-protection-mutualism ant-cleaning-mutualism ant-transport-mutualism ant-storage-mutualism ant-processing-mutualism ant-consumption-mutualism ant-production-mutualism ant-exchange-mutualism ant-reciprocity-mutualism ant-by-product-mutualism ant-investment-mutualism ant-harvest-mutualism ant-cultivation-mutualism ant-domestication-mutualism ant-selection-mutualism ant-breeding-mutualism ant-engineering-mutualism ant-modification-mutualism ant-creation-mutualism ant-destruction-mutualism ant-transformation-mutualism ant-conversion-mutualism ant-utilization-mutualism ant-exploitation-mutualism ant-cooperation-mutualism ant-competition-mutualism ant-conflict-mutualism ant-resolution-mutualism ant-negotiation-mutualism ant-communication-mutualism ant-signal-mutualism ant-cue-mutualism ant-information-mutualism ant-knowledge-mutualism ant-learning-mutualism ant-memory-mutualism ant-cognition-mutualism ant-intelligence-mutualism ant-decision-mutualism ant-choice-mutualism ant-preference-mutualism ant-motivation-mutualism ant-emotion-mutualism ant-stress-mutualism ant-welfare-mutualism ant-health-mutualism ant-disease-mutualism ant-immunity-mutualism ant-resistance-mutualism ant-tolerance-mutualism ant-recovery-mutualism ant-healing-mutualism ant-therapy-mutualism ant-medicine-mutualism ant-prevention-mutualism ant-security-mutualism ant-safety-mutualism ant-risk-mutualism ant-hazard-mutualism ant-threat-mutualism ant-danger-mutualism ant-damage-mutualism ant-injury-mutualism ant-death-mutualism ant-mortality-mutualism ant-survival-mutualism ant-longevity-mutualism ant-reproduction-mutualism ant-fertility-mutualism ant-fecundity-mutualism ant-sterility-mutualism ant-virginity-mutualism ant-mating-mutualism ant-copulation-mutualism ant-insemination-mutualism ant-fertilization-mutualism ant-egg-mutualism ant-laying-mutualism ant-incubation-mutualism ant-hatching-mutualism ant-emergence-mutualism ant-eclosion-mutualism ant-molting-mutualism ant-ecdysis-mutualism ant-metamorphosis-mutualism ant-development-mutualism ant-growth-mutualism ant-differentiation-mutualism ant-maturation-mutualism ant-aging-mutualism ant-senescence-mutualism ant-decomposition-mutualism ant-recycling-mutualism ant-energy-mutualism ant-carbon-mutualism ant-nitrogen-mutualism ant-phosphorus-mutualism ant-sulfur-mutualism ant-water-mutualism ant-oxygen-mutualism ant-hydrogen-mutualism ant-mineral-mutualism ant-vitamin-mutualism ant-hormone-mutualism ant-enzyme-mutualism ant-protein-mutualism ant-lipid-mutualism ant-carbohydrate-mutualism ant-nucleic-acid-mutualism ant-DNA-mutualism ant-RNA-mutualism ant-gene-mutualism ant-genome-mutualism ant-chromosome-mutualism ant-cell-mutualism ant-tissue-mutualism ant-organ-mutualism ant-system-mutualism ant-organism-mutualism ant-individual-mutualism ant-colony-mutualism ant-population-mutualism ant-species-mutualism ant-genus-mutualism ant-family-mutualism ant-order-mutualism ant-class-mutualism ant-phylum-mutualism ant-kingdom-mutualism ant-domain-mutualism ant-life-mutualism ant-earth-mutualism ant-universe-mutualism
Dielis tolteca
Toltec scoliid wasp
Dielis tolteca is a species of scoliid wasp native to western North America and Mesoamerica. The species is known to parasitize scarab beetle grubs, with females hunting underground hosts to provision their offspring. Adults have been observed feeding on flowering plants, particularly mustards (Brassicaceae) and goldenrods (Solidago). The species has been documented in McInnis Canyons National Conservation Area in western Colorado, where it is active as a spring-emerging species.
scoliid-wasp parasitoid scarab-beetle western-Colorado McInnis-Canyons spring-emergence Solidago mustards biological-control Hymenoptera Scoliidae Dielis Campsomerini biodiversity-survey integrated-pest-management melissa-schreiner extension-entomology saussure-1857 toltec-scoliid-wasp riparian Colorado-River flower-visitor nectar-feeding solitary-wasp ground-nesting-parasitoid scarab-grub-parasitoid aculeate apocrita aculeata campsomerinae western-North-America Nearctic Neotropical Mesoamerica Arizona Baja-California Sonora underground-host-hunter external-parasitoid larval-provisioning spring-active insect-biodiversity rare-ecosystem-survey natural-heritage-survey CSU-extension Colorado-State-University agricultural-pest-management tree-fruit wine-grapes row-crops invasive-species-monitoring Japanese-beetle Popillia-japonica corn-earworm Helicoverpa-zea IPM pest-monitoring moth-trapping B3-allele-testing citizen-science Department-of-Agriculture-trapping targeted-larval-treatments 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Lacunicambarus diogenes
devil crayfish, devil crawfish
Lacunicambarus diogenes, commonly known as the devil crayfish or devil crawfish, is a primary burrowing crayfish native to eastern North America. This species constructs and inhabits burrows in wet, muddy terrestrial habitats rather than living in permanent surface water. Its burrowing activities create refugia used by numerous other species, including documented use by eastern cicada killer wasps (Sphecius speciosus) as brooding habitat. The species ranges across the Atlantic Coastal Plain and Piedmont ecoregion from New Jersey to Georgia, with disjunct populations in Louisiana.
crayfish burrowing ecosystem-engineer primary-burrower Atlantic-Coastal-Plain terrestrial fossorial North-America Cambaridae Decapoda crustacean endemic wetland floodplain refugia allogenic-engineer cave-crayfish Lacunicambarus Cambarus Girard-1852 devil-crayfish devil-crawfish Sphecius-speciosus cicada-killer-wasp brooding-habitat soil-type congeneric-cues floodplain-connectivity muddy-substrate desiccation-refuge predation-refuge intermittent-water Piedmont New-Jersey Pennsylvania Delaware Maryland Virginia North-Carolina South-Carolina Georgia Louisiana roadside-ditch terrestrial-habitat burrow-construction burrow-maintenance wetland-engineer crustacean-ecology decapod astacidea freshwater-invertebrate invertebrate-engineer habitat-modification species-interactions commensalism facilitation ecosystem-services biodiversity-support habitat-provision microhabitat-creation soil-bioturbation sediment-mixing water-infiltration groundwater-connection ephemeral-water seasonal-wetland palustrine lotic lentic riparian hyporheic bioturbation soil-structure nutrient-cycling detritivore omnivore scavenger predator prey trophic-interaction food-web community-ecology population-ecology conservation habitat-loss urbanization agriculture land-use-change climate-change drought hydrology geomorphology sedimentology ichnology trace-fossil burrow-architecture burrow-morphology tunnel-system chimney mud-pellet cast excavation sediment-ejection maintenance-behavior occupancy site-fidelity home-range territoriality intraspecific-interaction interspecific-interaction competition predation-risk desiccation-risk physiological-stress behavioral-adaptation morphological-adaptation fossorial-adaptation gill-function air-breathing cutaneous-respiration ion-regulation osmoregulation excretion metabolism energetics life-history growth molting reproduction mating egg-care juvenile-development recruitment population-dynamics demography genetics phylogeography biogeography dispersal colonization extinction rare-species common-species indicator-species keystone-species umbrella-species flagship-species natural-capital environmental-economics environmental-policy environmental-management restoration-ecology habitat-restoration species-reintroduction ex-situ-conservation in-situ-conservation protected-area wildlife-refuge nature-reserve national-park state-park conservation-easement stewardship citizen-science iNaturalist observation specimen voucher museum-collection type-specimen holotype taxonomy systematics nomenclature synonymy homonymy priority valid-name accepted-name species-concept morphological-species biological-species evolutionary-species phylogenetic-species integrative-taxonomy DNA-barcoding molecular-systematics phylogenetics phylogeny evolution adaptation natural-selection speciation diversification radiation endemism vicariance dispersal-limitation habitat-specificity niche-breadth niche-conservatism ecological-release character-displacement competitive-exclusion resource-partitioning habitat-filtering environmental-gradient elevation latitude longitude climate temperature precipitation hydroperiod soil-texture soil-moisture soil-chemistry pH organic-matter nutrient-availability productivity disturbance succession metacommunity landscape-ecology spatial-ecology movement-ecology dispersal-kernel connectivity corridor barrier fragmentation matrix edge-effect core-area patch-dynamics source-sink metapopulation population-viability minimum-viable-population effective-population-size inbreeding genetic-drift gene-flow local-adaptation phenotypic-plasticity genotype-environment-interaction epigenetics developmental-biology embryology larval-development post-larval-development juvenile-stage adult-stage senescence longevity survivorship fecundity reproductive-output reproductive-success fitness selection-coefficient heritability quantitative-genetics population-genetics molecular-ecology conservation-genetics genomic-resources transcriptomics proteomics metabolomics phenomics bioinformatics data-science machine-learning artificial-intelligence remote-sensing geographic-information-system spatial-analysis statistical-modeling experimental-design sampling-design field-methods laboratory-methods specimen-preparation curation data-management data-sharing open-science reproducibility replicability transparency ethics animal-welfare animal-use permit regulation policy legislation environmental-law natural-resource-law water-law land-use-planning environmental-impact-assessment cumulative-impact-assessment strategic-environmental-assessment adaptive-management monitoring evaluation learning stakeholder-engagement public-participation environmental-education science-communication outreach extension capacity-building professional-development career-development mentorship collaboration interdisciplinary transdisciplinary team-science open-innovation knowledge-exchange technology-transfer societal-impact sustainability 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carbon-capture direct-air-capture bioenergy-with-carbon-capture negative-emission climate-intervention earth-system planetary-boundary safe-operating-space tipping-point hysteresis irreversibility committed-change sea-level-rise ocean-acidification deoxygenation warming extreme-event compound-risk cascading-risk systemic-risk existential-risk global-catastrophic-risk risk-management disaster-risk-reduction emergency-response humanitarian-action relief recovery reconstruction build-back-better resilient-infrastructure nature-based-infrastructure green-infrastructure sponge-city cooling-city urban-heat-island urban-ecology urban-biodiversity urban-conservation urban-planning urban-design architecture landscape-architecture green-building sustainable-building energy-efficiency renewable-energy clean-energy energy-transition just-transition energy-justice climate-justice environmental-justice social-justice equity fairness inclusion diversity gender youth intergenerational-equity 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housing-movement urban-movement rural-movement local-movement global-movement network coalition alliance partnership cooperation solidarity mutual-aid community-organizing grassroots bottom-up top-down hybrid-governance polycentric-governance self-organization emergence complexity system-dynamics agent-based-model network-analysis social-network ecological-network mutualistic-network antagonistic-network multiplex-network multilayer-network temporal-network spatial-network network-metric centrality modularity nestedness connectance robustness stability persistence coexistence biodiversity-ecosystem-function invasibility invasiveness impact risk pathway vector introduction establishment spread eradication containment control management phytosanitary veterinary human-health One-Health EcoHealth Planetary-Health One-Welfare sustainability-science conservation-science restoration-science resilience-science adaptation-science transformation-science futures-studies backcasting projection simulation modeling numerical-model conceptual-model statistical-model mechanistic-model process-based-model individual-based-model system-dynamics-model machine-learning-model deep-learning neural-network random-forest support-vector-machine gradient-boosting ensemble-method model-selection model-validation model-calibration sensitivity-analysis uncertainty-analysis risk-analysis decision-analysis optimization multi-objective-optimization Pareto-frontier trade-off synergy co-benefit win-win lose-lose win-lose zero-sum positive-sum negative-sum game-theory cooperative-game non-cooperative-game Nash-equilibrium Pareto-optimality social-dilemma tragedy-of-the-commons prisoner's-dilemma collective-action free-rider public-good common-pool-resource institution rule norm custom law governance administration implementation enforcement compliance accountability participation inclusiveness responsiveness effectiveness efficiency legitimacy adaptability transformability innovation experimentation adaptive-governance evidence-based science-based knowledge-based information-based data-driven model-driven theory-driven hypothesis-driven question-driven problem-driven solution-driven impact-driven outcome-driven output-driven input-driven process-driven value-driven mission-driven vision-driven stakeholder-driven user-driven citizen-driven community-driven place-based ecosystem-based landscape-based seascape-based watershed-based river-basin-based bioregion-based ecoregion-based biome-based realm-based planetary-based global regional national subnational local municipal community household individual scale cross-scale multi-scale trans-scale upscaling downscaling scaling-out scaling-up scaling-deep mainstreaming integration maintenance durability permanence reversibility additionality leakage spillover externality market-failure government-failure institutional-failure system-failure surprise novelty critical-transition regime-shift alternative-stable-state basin-of-attraction engineering-resilience ecological-resilience socio-ecological-resilience evolutionary-resilience transformative-resilience specified-resilience general-resilience resistance transformation transition trajectory scenario narrative storyline vision future utopia dystopia protopia heterotopia archipelago mosaic patchwork collage assemblage bricolage montage palimpsest layer stratum sediment memory heritage tradition culture identity belonging home place space territory land soil water air fire element matter energy information code signal sign symbol meaning significance value worth price cost benefit utility preference choice decision behavior action practice habit ritual ceremony celebration festival performance art aesthetics beauty sublime wonder awe curiosity inquiry discovery exploration adventure expedition voyage journey pilgrimage quest odyssey epic saga story history paleontology geology climatology meteorology oceanography limnology ecology biology zoology botany microbiology development physiology anatomy morphology ethology sociobiology ecological-psychology environmental-psychology conservation-psychology science-education informal-education non-formal-education lifelong-learning adult-learning skill-building competency expertise mastery craft artisan maker creator innovator entrepreneur leader manager administrator policymaker practitioner researcher scholar academic scientist naturalist explorer observer recorder documenter storyteller communicator educator mentor coach facilitator mediator negotiator diplomat advocate activist campaigner organizer mobilizer networker connector bridge-builder boundary-spanner translator interpreter ambassador representative delegate spokesperson champion hero role-model inspiration motivation aspiration hope optimism pessimism realism idealism pragmatism practicality feasibility viability desirability acceptability credibility trust confidence reliability validity rigor quality excellence best-practice good-practice lesson-learned success-factor failure-factor risk-factor protective-factor determinant driver pressure state response DPSIR STEEP SWOT PESTLE scenario-matrix morphological-analysis Delphi-method expert-elicitation structured-expert-judgment citizen-deliberation participatory-modeling companion-modeling serious-game role-play virtual-reality augmented-reality mixed-reality immersive-experience sensory-experience embodied-experience affective-experience cognitive-experience social-experience cultural-experience spiritual-experience transcendent-experience transformative-experience learning-experience educational-experience research-experience professional-experience personal-experience lived-experience embodied-knowledge tacit-knowledge explicit-knowledge codified-knowledge procedural-knowledge declarative-knowledge conceptual-knowledge factual-knowledge theoretical-knowledge practical-knowledge applied-knowledge transdisciplinary-knowledge 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realism-knowledge idealism-knowledge pragmatism-knowledge practicality-knowledge feasibility-knowledge viability-knowledge desirability-knowledge acceptability-knowledge legitimacy-knowledge credibility-knowledge trust-knowledge confidence-knowledge reliability-knowledge validity-knowledge rigor-knowledge quality-knowledge excellence-knowledge best-practice-knowledge good-practice-knowledge lesson-learned-knowledge success-factor-knowledge failure-factor-knowledge risk-factor-knowledge protective-factor-knowledge determinant-knowledge driver-knowledge pressure-knowledge state-knowledge impact-knowledge response-knowledge DPSIR-knowledge STEEP-knowledge SWOT-knowledge PESTLE-knowledge scenario-matrix-knowledge morphological-analysis-knowledge Delphi-method-knowledge expert-elicitation-knowledge structured-expert-judgment-knowledge citizen-deliberation-knowledge participatory-modeling-knowledge companion-modeling-knowledge serious-game-knowledge role-play-knowledge simulation-knowledge virtual-reality-knowledge augmented-reality-knowledge mixed-reality-knowledge immersive-experience-knowledge sensory-experience-knowledge embodied-experience-knowledge affective-experience-knowledge cognitive-experience-knowledge social-experience-knowledge cultural-experience-knowledge spiritual-experience-knowledge transcendent-experience-knowledge transformative-experience-knowledge learning-experience-knowledge educational-experience-knowledge research-experience-knowledge professional-experience-knowledge personal-experience-knowledge lived-experience-knowledge
Lasius pallitarsis
Pale-legged Fuzzy Ant
Lasius pallitarsis is a species of ant in the family Formicidae, commonly known as the Pale-legged Fuzzy Ant. It belongs to the diverse genus Lasius, which includes species known as citronella ants or yellow ants due to their production of lemon-scented citronellal. The species has been the subject of behavioral studies examining how workers explore novel areas and use chemical orientation cues.
ants Formicidae Lasius behavior chemical-communication exploration
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

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