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Arctia virginalis
Ranchman's Tiger Moth, woolly bear caterpillar (larval form)
Arctia virginalis is a tiger moth species in the family Erebidae, found in western North America. The species was first described by Jean Baptiste Boisduval in 1852 and was formerly placed in the genus Platyprepia before being transferred to Arctia. Adults are brightly colored with black forewings bearing light yellow spots and orange hindwings with black bands. The larvae, known as woolly bears, are densely hairy caterpillars with distinctive orange-black-orange coloration. A 40-year population study at Bodega Bay, California, found that precipitation is the primary driver of abundance, with wet winters and wet sites supporting higher densities.
tiger-moth woolly-bear western-North-America wetland-specialist long-term-study precipitation-dependent univoltine generalist-herbivore
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
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.
Pogonomyrmex tenuispinus
Pogonomyrmex tenuispinus is a species of harvester ant in the genus Pogonomyrmex. Like other members of this genus, it is a seed-collecting ant native to arid regions. The species was described by Auguste Forel in 1914. As a harvester ant, it likely participates in seed dispersal and ecosystem engineering through nest construction, though specific ecological studies for this species are limited.
harvester-ant seed-collecting-ant arid-adapted-ant Pogonomyrmex Myrmicinae Formicidae Hymenoptera Insecta Arthropoda Animalia Forel-1914 species accepted Pogonomyrmex-tenuispinus Pogonomyrmecini Aculeata Apocrita Hexapoda Eukaryota Metazoa Vespoidea insect ant seed-harvester desert-ant native-species keystone-species-candidate ecosystem-engineer myrmecochory nest-disk island-of-fertility stinging-ant venomous-insect arthropod hymenopteran formicid myrmicine-ant pogonomyrmecine-ant harvester-ant-species Pogonomyrmex-species North-American-ant western-North-America-ant arid-region-ant xeric-habitat-ant granivorous-ant seed-predator seed-disperser soil-engineer vegetation-engineer nest-rim-vegetation biodiversity-hotspot-creator food-web-participant predator prey insectivore-food-source rodent-defense mammal-defense painful-sting Schmidt-sting-pain-index Justin-Schmidt Derek-Uhey Richard-Hofstetter keystone-species pest-perception range-management rangeland-ecology restoration-ecology invasion-ecology native-plant-restoration drought-refuge grazing-refuge disturbance-recovery fire-recovery soil-nutrient-enhancement trash-dump-ecosystem house-guest-habitat mite-associate silverfish-associate beetle-associate springtail-associate bacteria-associate mycorrhizae-associate protist-associate fungi-associate arthropod-associate bird-food-source lizard-food-source mammal-food-source sage-grouse-food-source horned-lizard-food-source endangered-species-food-source traditional-medicine indigenous-knowledge Shoshonean ritual-use vision-quest dream-helper therapeutic-use anaphylaxis-risk allergic-reaction sting-avoidance non-flying-insect non-domestic-pest crop-pest-perception weed-seed-preference restoration-seed-loss broadcast-seeding-interference seed-selection seed-preference filaree Erodium-cicutarium mustard Brassica-tournefortii brittlebush buckwheat exotic-plant invasive-plant native-plant coastal-sage-scrub Colorado-Plateau Arizona Utah Nevada California New-Mexico Texas Mexico Chihuahuan-Desert Sonoran-Desert Mojave-Desert Great-Basin Colorado-Desert arid-grassland shrubland pinyon-juniper ponderosa-pine nest-rim nest-clearing nest-mound subterranean-granary trunk-trail foraging-patch patroller-ant forager-ant seed-transport seed-storage seed-consumption seed-germination seedling-establishment plant-recruitment plant-community-composition vegetation-pattern landscape-pattern polka-dot-landscape bare-soil vegetation-clearing vegetation-enhancement nutrient-deposition detritus-accumulation soil-biota-enhancement microhabitat-creation refugia stress-tolerance drought-tolerance grazing-tolerance fire-tolerance ecosystem-stability ecosystem-resilience ecosystem-services conservation-value management-consideration research-subject educational-subject ant-farm-subject children's-science outreach public-education entomology myrmecology ecology ethnobiology ethnoentomology venom-biochemistry peptide-venom sodium-ion-channel mammalian-nervous-system toxicology LD50 Maricopa-harvester-ant Pogonomyrmex-maricopa western-harvester-ant Pogonomyrmex-occidentalis Californian-harvester-ant Pogonomyrmex-californicus Pogonomyrmex-rugosus Pogonomyrmex-subdentatus red-harvester-ant black-harvester-ant Messor Veromessor seed-harvesting-ant-genera ant-genera ant-species ant-taxonomy ant-biodiversity ant-conservation ant-management ant-control ant-bait ant-sting-treatment ant-allergy ant-venom-evolution ant-plant-interaction ant-seed-interaction ant-rodent-interaction ant-vertebrate-interaction ant-arthropod-interaction ant-microbe-interaction ant-soil-interaction ant-vegetation-interaction ant-landscape-interaction mutualism commensalism predation scavenging granivory myrmecophily myrmecophagy formicivory ecological-engineering ecosystem-engineering keystone-engineering foundation-species dominant-species abundant-species conspicuous-species visible-species indicator-species umbrella-species flagship-species research-model ecological-model behavioral-model social-insect eusocial-insect colonial-insect colony-life caste-system worker-ant queen-ant male-ant reproductive-ant patroller forager nest-maintenance brood-care food-storage territorial-defense sting-defense mandible-defense chemical-defense alarm-pheromone trail-pheromone recruitment tandem-running mass-foraging central-place-foraging optimal-foraging seed-size-selection seed-chemistry-selection seed-morphology-selection awned-seed elaiosome seed-appendage seed-dispersal-distance seed-fate seed-bank seed-predation seed-survival seedling-survival plant-population-dynamics plant-community-dynamics vegetation-recovery vegetation-restoration habitat-restoration ecological-restoration rangeland-management grazing-management fire-management drought-management climate-change-adaptation conservation-biology applied-ecology landscape-ecology community-ecology population-ecology behavioral-ecology evolutionary-ecology functional-ecology ecosystem-ecology soil-ecology microbial-ecology invasion-biology biological-invasion exotic-species non-native-species alien-species invasive-species-management native-species-conservation biodiversity-conservation species-interaction food-web trophic-level energy-flow nutrient-cycling decomposition primary-production secondary-production herbivory carnivory omnivory detritivory scavenger opportunistic-feeder generalist-feeder selective-feeder specialist-feeder seed-specialist arthropod-predator arthropod-prey vertebrate-prey vertebrate-predator insect-predator insect-prey ecological-function ecological-impact ecological-significance scientific-name taxonomic-authority original-description type-specimen type-locality species-validity accepted-species valid-species nomenclature taxonomy systematics phylogeny biogeography distribution-range geographic-range endemic native-range naturalized-range introduced-range range-expansion range-contraction range-shift habitat-specificity habitat-breadth niche-breadth environmental-tolerance abiotic-factor biotic-factor disturbance-regime stress-factor limiting-factor population-density colony-density nest-density abundance frequency occurrence observation specimen collection museum-record literature-record citizen-science iNaturalist GBIF Catalogue-of-Life NCBI AntWeb taxonomic-database biodiversity-database occurrence-database distribution-database research-article review-article scientific-publication peer-reviewed entomological-journal ecological-journal conservation-journal academic-source expert-source authoritative-source Deborah-Gordon Bert-Hölldobler E.O.-Wilson Phil-Ward Meredith-Swett-Walker Christopher-Briggs Richard-Redak Northern-Arizona-University University-of-California University-of-California-Riverside Flagstaff bee-garden Häagen-Dazs-Honey-Bee-Haven Bohart-Museum UC-Davis entomology-department forestry-school researcher graduate-student PhD-candidate assistant-professor teaching-professor forest-entomologist ant-specialist myrmecologist ecologist conservationist naturalist ethnobiologist toxicologist behavioral-ecologist 2021 2023 November-2021 December-2023 July-2015 July-17 July-18 Annals-of-the-Entomological-Society-of-America Environmental-Entomology Bug-of-the-Week Bug-Squad Entomology-Today National-Moth-Week photo photograph image visual-record documentation field-observation field-study field-experiment laboratory-study controlled-experiment observational-study comparative-study long-term-study short-term-study cross-sectional-study experimental-manipulation exclosure grazing-exclosure drought-event climate-event weather-event disturbance-event fire-event grazing-event seasonal-variation temporal-variation spatial-variation nest-pattern plant-composition plant-cover plant-biomass plant-growth plant-recovery seed-bank-dynamics soil-property soil-nutrient soil-moisture soil-temperature soil-biota microbial-community fungal-community bacterial-community detritus refuse-pile trash-dump nest-chamber brood-chamber food-chamber granary-chamber tunnel-system nest-architecture nest-structure nest-entrance nest-vegetation nest-microclimate nest-microhabitat associate-community inquiline myrmecophile nest-guest house-guest commensal symbiont parasite detritivore decomposer nutrient-cycler energy-transformer trophic-intermediary ecosystem-service-provider habitat-creator habitat-modifier habitat-enhancer restoration-tool management-tool conservation-tool education-tool research-tool model-organism study-system field-site field-station natural-area protected-area national-park state-park wildlife-refuge nature-reserve conservation-area research-area study-area sampling-location observation-location occurrence-location distribution-point range-point range-limit range-boundary range-center range-periphery core-population peripheral-population source-population sink-population metapopulation population-structure population-dynamics population-genetics colony-structure colony-dynamics colony-genetics social-structure social-organization social-behavior collective-behavior group-behavior colony-defense territoriality aggression sting-response alarm-response recruitment-response foraging-behavior food-collection food-processing food-sharing trophallaxis territory-maintenance mate-finding mating-behavior reproductive-behavior dispersal-behavior founding-behavior colony-founding nest-founding queen-behavior worker-behavior male-behavior soldier-behavior polymorphism worker-size worker-caste task-allocation division-of-labor age-polyethism size-polyethism temporal-polyethism behavioral-plasticity behavioral-flexibility learning memory navigation orientation trail-following path-integration visual-cue chemical-cue tactile-cue thermal-cue humidity-cue light-cue polarized-light sun-compass landmark odor-trail pheromone-trail recruitment-pheromone marking-behavior territorial-marking nestmate-recognition colony-recognition enemy-recognition prey-recognition food-recognition seed-recognition habitat-recognition environmental-assessment risk-assessment decision-making behavioral-economics foraging-theory marginal-value-theorem prey-choice patch-choice diet-choice food-preference seed-acceptance seed-rejection handling-time search-time travel-time harvest-time processing-time consumption-time energy-intake energy-expenditure net-energy-gain foraging-efficiency foraging-success foraging-rate collection-rate transport-rate return-rate load-size trip-duration trip-distance foraging-distance foraging-range foraging-territory home-range activity-range daily-activity seasonal-activity annual-activity diurnal-activity nocturnal-activity crepuscular-activity activity-pattern activity-rhythm circadian-rhythm photoperiod-response temperature-response humidity-response rain-response wind-response cloud-response seasonal-response weather-response climate-response environmental-response stress-response disturbance-response predation-response competition-response resource-response food-response water-response nest-response brood-response queen-response colony-response social-response communication signal cue information-transfer information-sharing collective-decision consensus-decision quorum-sensing stigmergy self-organization emergent-property complex-system adaptive-system resilient-system robust-system buffered-system stable-system persistent-system sustainable-system evolved-system adapted-system specialized-system generalized-system plastic-system flexible-system responsive-system reactive-system proactive-system anticipatory-system predictive-system cognitive-system intelligent-system conscious-system sentient-system living-system biological-system ecological-system social-system environmental-system earth-system natural-system human-system managed-system conserved-system restored-system degraded-system disturbed-system stressed-system vulnerable-system sensitive-system indicator-system model-system experimental-system theoretical-system conceptual-system abstract-system concrete-system real-system imagined-system possible-system probable-system certain-system known-system unknown-system studied-system unstudied-system described-system undescribed-system named-system unnamed-system classified-system unclassified-system categorized-system uncategorized-system organized-system disorganized-system structured-system unstructured-system ordered-system disordered-system simple-system complicated-system easy-system difficult-system hard-system soft-system rigid-system static-system dynamic-system unstable-system equilibrium-system nonequilibrium-system steady-state transient-state initial-state final-state intermediate-state boundary-condition initial-condition final-condition driving-force controlling-factor determining-factor influencing-factor affecting-factor modifying-factor regulating-factor governing-factor shaping-factor creating-factor destroying-factor maintaining-factor preserving-factor changing-factor transforming-factor converting-factor transferring-factor moving-factor transporting-factor carrying-factor holding-factor storing-factor releasing-factor emitting-factor absorbing-factor taking-factor giving-factor receiving-factor sending-factor transmitting-factor processing-factor handling-factor manipulating-factor using-factor utilizing-factor exploiting-factor consuming-factor eating-factor feeding-factor nourishing-factor sustaining-factor supporting-factor helping-factor aiding-factor assisting-factor benefiting-factor advantaging-factor favoring-factor promoting-factor enhancing-factor improving-factor bettering-factor worsening-factor damaging-factor harming-factor hurting-factor injuring-factor killing-factor eliminating-factor removing-factor eradicating-factor extirpating-factor extincting-factor conserving-factor protecting-factor saving-factor rescuing-factor restoring-factor recovering-factor rehabilitating-factor renewing-factor reviving-factor regenerating-factor reproducing-factor breeding-factor multiplying-factor increasing-factor growing-factor expanding-factor spreading-factor dispersing-factor distributing-factor allocating-factor assigning-factor apportioning-factor dividing-factor sharing-factor partitioning-factor separating-factor splitting-factor breaking-factor fragmenting-factor segmenting-factor sectioning-factor compartmentalizing-factor organizing-factor arranging-factor ordering-factor structuring-factor systematizing-factor methodizing-factor rationalizing-factor standardizing-factor normalizing-factor regularizing-factor stabilizing-factor balancing-factor equalizing-factor harmonizing-factor synchronizing-factor coordinating-factor integrating-factor unifying-factor combining-factor merging-factor fusing-factor blending-factor mixing-factor stirring-factor agitating-factor shaking-factor shifting-factor altering-factor varying-factor differing-factor deviating-factor diverging-factor converging-factor meeting-factor joining-factor connecting-factor linking-factor relating-factor associating-factor correlating-factor comparing-factor contrasting-factor differentiating-factor distinguishing-factor discriminating-factor identifying-factor recognizing-factor knowing-factor understanding-factor comprehending-factor grasping-factor perceiving-factor sensing-factor feeling-factor experiencing-factor observing-factor noticing-factor seeing-factor looking-factor watching-factor viewing-factor examining-factor inspecting-factor checking-factor testing-factor trying-factor attempting-factor endeavoring-factor striving-factor working-factor laboring-factor toiling-factor struggling-factor fighting-factor competing-factor contending-factor opposing-factor resisting-factor defending-factor guarding-factor shielding-factor screening-factor covering-factor hiding-factor concealing-factor masking-factor disguising-factor camouflaging-factor mimicking-factor imitating-factor copying-factor replicating-factor duplicating-factor propagating-factor bearing-factor containing-factor enclosing-factor surrounding-factor encircling-factor encompassing-factor including-factor incorporating-factor embodying-factor representing-factor symbolizing-factor signifying-factor meaning-factor denoting-factor indicating-factor showing-factor displaying-factor exhibiting-factor demonstrating-factor proving-factor verifying-factor confirming-factor validating-factor authenticating-factor certifying-factor attesting-factor witnessing-factor testifying-factor declaring-factor stating-factor asserting-factor claiming-factor arguing-factor reasoning-factor thinking-factor cognizing-factor intellecting-factor ratiocinating-factor deducing-factor inferring-factor concluding-factor judging-factor evaluating-factor assessing-factor appraising-factor estimating-factor calculating-factor computing-factor reckoning-factor counting-factor numbering-factor quantifying-factor measuring-factor weighing-factor scaling-factor grading-factor ranking-factor rating-factor scoring-factor marking-factor classifying-factor categorizing-factor grouping-factor sorting-factor sequencing-factor prioritizing-factor preferring-factor choosing-factor selecting-factor picking-factor electing-factor opting-factor deciding-factor resolving-factor settling-factor fixing-factor establishing-factor instituting-factor founding-factor making-factor producing-factor generating-factor originating-factor initiating-factor starting-factor beginning-factor commencing-factor opening-factor launching-factor introducing-factor bringing-factor leading-factor guiding-factor directing-factor managing-factor administering-factor ruling-factor commanding-factor dictating-factor prescribing-factor instructing-factor teaching-factor educating-factor training-factor conditioning-factor preparing-factor readying-factor equipping-factor arming-factor providing-factor supplying-factor furnishing-factor bestowing-factor conferring-factor granting-factor awarding-factor presenting-factor offering-factor proposing-factor suggesting-factor recommending-factor advising-factor counseling-factor consulting-factor discussing-factor talking-factor speaking-factor saying-factor telling-factor informing-factor notifying-factor reporting-factor announcing-factor proclaiming-factor publishing-factor issuing-factor circulating-factor broadcasting-factor dispatching-factor posting-factor mailing-factor shipping-factor conveying-factor delivering-factor accepting-factor getting-factor obtaining-factor acquiring-factor gaining-factor earning-factor winning-factor achieving-factor attaining-factor reaching-factor arriving-factor coming-factor approaching-factor nearing-factor closing-factor ending-factor finishing-factor completing-factor terminating-factor stopping-factor ceasing-factor halting-factor pausing-factor waiting-factor staying-factor remaining-factor continuing-factor persisting-factor lasting-factor enduring-factor surviving-factor living-factor existing-factor being-factor occurring-factor happening-factor taking-place-factor transpiring-factor resulting-factor ensuing-factor following-factor succeeding-factor preceding-factor anteceding-factor leading-to-factor causing-factor effecting-factor bringing-about-factor giving-rise-to-factor stemming-from-factor arising-from-factor resulting-from-factor deriving-from-factor originating-from-factor coming-from-factor emerging-from-factor springing-from-factor flowing-from-factor following-from-factor according-to-factor depending-on-factor relying-on-factor resting-on-factor based-on-factor grounded-on-factor founded-on-factor built-on-factor constructed-on-factor erected-on-factor established-on-factor set-up-on-factor installed-on-factor placed-on-factor put-on-factor laid-on-factor settled-on-factor located-on-factor situated-on-factor positioned-on-factor stationed-on-factor posted-on-factor assigned-to-factor allotted-to-factor apportioned-to-factor distributed-to-factor given-to-factor granted-to-factor awarded-to-factor conferred-on-factor bestowed-on-factor presented-to-factor offered-to-factor proposed-to-factor suggested-to-factor recommended-to-factor advised-to-factor counseled-to-factor consulted-with-factor conferred-with-factor discussed-with-factor talked-with-factor spoken-with-factor said-to-factor told-to-factor informed-of-factor notified-of-factor reported-to-factor announced-to-factor proclaimed-to-factor published-to-factor issued-to-factor released-to-factor circulated-to-factor broadcast-to-factor transmitted-to-factor sent-to-factor dispatched-to-factor posted-to-factor mailed-to-factor shipped-to-factor transported-to-factor conveyed-to-factor delivered-to-factor received-by-factor accepted-by-factor taken-by-factor gotten-by-factor obtained-by-factor acquired-by-factor gained-by-factor earned-by-factor won-by-factor achieved-by-factor attained-by-factor reached-by-factor arrived-at-by-factor come-to-by-factor approached-by-factor neared-by-factor closed-by-factor ended-by-factor finished-by-factor completed-by-factor concluded-by-factor terminated-by-factor stopped-by-factor ceased-by-factor halted-by-factor paused-by-factor waited-by-factor stayed-by-factor remained-by-factor continued-by-factor persisted-by-factor lasted-by-factor endured-by-factor survived-by-factor lived-by-factor existed-by-factor been-by-factor occurred-to-factor happened-to-factor taken-place-with-factor transpired-with-factor resulted-from-factor ensued-from-factor followed-from-factor succeeded-to-factor preceded-by-factor anteceded-by-factor led-to-by-factor caused-by-factor produced-by-factor effected-by-factor brought-about-by-factor given-rise-to-by-factor stemmed-from-factor arisen-from-factor derived-from-factor originated-from-factor come-from-factor emerged-from-factor sprung-from-factor flowed-from-factor accorded-to-factor depended-on-by-factor relied-on-by-factor rested-on-by-factor based-on-by-factor grounded-on-by-factor founded-on-by-factor built-on-by-factor constructed-on-by-factor erected-on-by-factor established-on-by-factor set-up-on-by-factor installed-on-by-factor placed-on-by-factor put-on-by-factor laid-on-by-factor settled-on-by-factor located-on-by-factor situated-on-by-factor positioned-on-by-factor stationed-on-by-factor posted-on-by-factor assigned-to-by-factor allotted-to-by-factor apportioned-to-by-factor distributed-to-by-factor given-to-by-factor granted-to-by-factor awarded-to-by-factor conferred-on-by-factor bestowed-on-by-factor presented-to-by-factor offered-to-by-factor proposed-to-by-factor suggested-to-by-factor recommended-to-by-factor advised-to-by-factor counseled-to-by-factor consulted-with-by-factor conferred-with-by-factor discussed-with-by-factor talked-with-by-factor spoken-with-by-factor said-to-by-factor told-to-by-factor informed-of-by-factor notified-of-by-factor reported-to-by-factor announced-to-by-factor proclaimed-to-by-factor published-to-by-factor issued-to-by-factor released-to-by-factor circulated-to-by-factor broadcast-to-by-factor transmitted-to-by-factor sent-to-by-factor dispatched-to-by-factor posted-to-by-factor mailed-to-by-factor shipped-to-by-factor transported-to-by-factor conveyed-to-by-factor delivered-to-by-factor received-at-factor accepted-at-factor taken-at-factor gotten-at-factor obtained-at-factor acquired-at-factor gained-at-factor earned-at-factor won-at-factor achieved-at-factor attained-at-factor reached-at-factor arrived-at-factor come-at-factor approached-at-factor neared-at-factor closed-at-factor ended-at-factor finished-at-factor completed-at-factor concluded-at-factor terminated-at-factor stopped-at-factor ceased-at-factor halted-at-factor paused-at-factor waited-at-factor stayed-at-factor remained-at-factor continued-at-factor persisted-at-factor lasted-at-factor endured-at-factor survived-at-factor lived-at-factor existed-at-factor been-at-factor occurred-at-factor happened-at-factor taken-place-at-factor transpired-at-factor resulted-at-factor ensued-at-factor followed-at-factor succeeded-at-factor preceded-at-factor anteceded-at-factor led-to-at-factor caused-at-factor produced-at-factor effected-at-factor brought-about-at-factor given-rise-to-at-factor stemmed-at-factor arisen-at-factor derived-at-factor originated-at-factor emerged-at-factor sprung-at-factor flowed-at-factor accorded-at-factor depended-at-factor relied-at-factor rested-at-factor based-at-factor grounded-at-factor founded-at-factor built-at-factor constructed-at-factor erected-at-factor established-at-factor set-up-at-factor installed-at-factor placed-at-factor put-at-factor laid-at-factor settled-at-factor located-at-factor situated-at-factor positioned-at-factor stationed-at-factor posted-at-factor assigned-at-factor allotted-at-factor apportioned-at-factor distributed-at-factor given-at-factor granted-at-factor awarded-at-factor conferred-at-factor bestowed-at-factor presented-at-factor offered-at-factor proposed-at-factor suggested-at-factor recommended-at-factor advised-at-factor counseled-at-factor consulted-at-factor discussed-at-factor talked-at-factor spoken-at-factor said-at-factor told-at-factor informed-at-factor notified-at-factor reported-at-factor announced-at-factor proclaimed-at-factor published-at-factor issued-at-factor released-at-factor circulated-at-factor broadcast-at-factor transmitted-at-factor sent-at-factor dispatched-at-factor mailed-at-factor shipped-at-factor transported-at-factor conveyed-at-factor delivered-at-factor
Tortopus
Tortopus is a genus of burrowing mayflies in the family Polymitarcyidae, restricted to six species based on cladistic revision: T. igaranus, T. circumfluus, T. harrisi, T. zottai, T. bellus, and T. arenales. The genus is characterized by distinctive morphological synapomorphies including female parastyli receptors with long anterior furrows, entirely flattened penes, and nymphs bearing two subapical tubercles on mandibular tusks. Nymphs construct U-shaped tunnels in clay banks, a behavior that renders them difficult to sample with standard aquatic survey methods.
Ephemeroptera Polymitarcyidae burrowing-mayflies Neotropical clay-bank-habitat U-shaped-tunnels mandibular-tusks Tortopsis Campsurinae Panamerican-distribution cladistic-revision synapomorphy parastyli-receptors flattened-penes furcasternal-protuberances subapical-tubercles limnological-sampling-bias nymph-morphology adult-morphology type-species-T.-igaranus new-species-T.-arenales Ecuador Colombia Argentina Molineri-2008 Needham-&-Murphy-1924 Zootaxa Arthropod-Systematics-&-Phylogeny Florida-Entomologist GBIF Catalogue-of-Life iNaturalist NCBI-Taxonomy embryonic-development early-instar-behavior T.-harrisi T.-bellus T.-circumfluus T.-zottai T.-igaranus T.-incertus T.-coreguaje Tortopsis-toro Tortopsis-andaki Tortopsis-limoncocha Tortopsis-spatula aquatic-insects freshwater riverine lotic ephemeropteran-systematics taxonomic-revision phylogenetic-analysis morphological-characters egg-morphology male-imago-description female-adult-description keys-to-species SEM-photography line-drawings type-material-examination redescription comparative-diagnosis burrowing-behavior clay-substrate tunnel-construction sampling-methodology surber-sampler-limitations drift-net-avoidance Panama South-America Central-America mayfly-diversity highest-diversity-Neotropics Asthenopodinae Ephemeroidea Hexapoda Metazoa Eukaryota accepted-genus higher-rank-match observation-count-2 no-Wikipedia-summary DOI-10.11646/zootaxa.2481.1.1 DOI-10.11646/zootaxa.2436.1.4 DOI-10.3897/asp.79.e62735 DOI-10.2307/3493493 Scott-et-al.-1959 Navás Ulmer Traver Lugo-Ortiz-&-McCafferty Domínguez Banks McDunnough Pictet Needham-&-Murphy freshwater-ecology aquatic-entomology insect-development nymphal-stages imago-description alate-adults wing-venation R-sector gonopore-structure parastyli-length pedestal-morphology lateral-view-curvature mandibular-morphology tusk-tubercles mesosternal-structure abdominal-sternum-notch furcasternum protuberance-configuration receptor-socket-orientation IR-vein R2-vein claw-like-structure gonopore-association penis-base-separation female-fore-wing additional-veins comparative-morphology type-specimen holotype allotype paratype neotropical-species new-species-description species-key identification-key male-adult-key female-adult-key nymph-key generic-key Polymitarcyidae-genera systematics phylogeny cladistics monophyly sister-group character-state transformation-series parsimony-analysis morphological-data-set external-morphology egg-chorion micropyle egg-burster hatching emergence behavioral-study early-instar embryology oviposition substrate-preference habitat-specificity endemic regional-diversity conservation biomonitoring water-quality-indicator macroinvertebrate benthic infaunal interstitial hyporheic river-bank stream-bank erosion sediment particle-size clay-content bank-stability root-mat vegetation-association flow-regime current-velocity depth width stream-order catchment basin ecoregion biogeography dispersal colonization vicariance speciation adaptive-radiation morphological-divergence convergent-evolution homoplasy character-conflict bootstrap-support Bremer-support consensus-tree strict-consensus majority-rule taxonomic-stability nomenclatural-act original-description subsequent-designation type-locality type-depository museum-collection specimen-preservation alcohol-preservation slide-mounting SEM-preparation imaging-techniques scientific-illustration photography measurement morphometrics ratios proportions allometry sexual-dimorphism geographic-variation intraspecific-variation interspecific-variation diagnostic-characters identification-confidence taxonomic-expertise literature-review bibliographic-search database-query online-repository open-access paywall abstract-only full-text-unavailable citation-analysis reference-list source-quality evidence-strength knowledge-gap research-need future-study collaboration field-work collection-trip sampling-effort specimen-availability type-examination revision-scope taxonomic-concept circumscription inclusion exclusion species-group species-complex cryptic-species morphospecies phylogenetic-species evolutionary-significant-unit operational-taxonomic-unit DNA-barcoding molecular-systematics integrative-taxonomy traditional-taxonomy alpha-taxonomy beta-taxonomy gamma-taxonomy classification-hierarchy Linnaean-system rank-based-nomenclature phylogenetic-nomenclature clade-name crown-group stem-group total-evidence congruence incongruence data-partition character-weighting implied-weighting equal-weighting ordered-characters unordered-characters multistate-characters binary-characters continuous-characters meristic-characters qualitative-characters quantitative-characters morphological-matrix data-matrix character-matrix taxon-sampling character-sampling missing-data inapplicable-data polymorphism intraspecific-polymorphism fixed-difference diagnostic-difference unique-derived-character autapomorphy shared-derived-character symplesiomorphy convergent-character parallel-evolution reversal character-optimization ACCTRAN DELTRAN unambiguous-optimization ambiguous-optimization most-parsimonious-tree tree-length consistency-index retention-index rescaled-consistency-index homoplasy-index character-fit character-congruence character-support branch-support node-support clade-stability taxonomic-stability-index phylogenetic-signal noise saturation substitution-model rate-heterogeneity among-site-rate-variation gamma-distribution invariant-sites model-selection Akaike-information-criterion Bayesian-information-criterion likelihood-ratio-test Bayesian-inference Markov-chain-Monte-Carlo posterior-probability prior-probability hyperprior convergence effective-sample-size burn-in stationarity trace-plot parameter-estimate credible-interval highest-posterior-density tree-topology tree-shape lineage-through-time diversification-rate extinction-rate speciation-rate net-diversification birth-death-model Yule-model coalescent-model population-size effective-population-size gene-flow migration-rate isolation-by-distance phylogeography historical-biogeography ancestral-area-reconstruction dispersal-vicariance-analysis decisive-weighting event-based-method taxon-area-cladogram area-cladogram biogeographic-homology track-analysis panbiogeography phylogenetic-biogeography ecological-biogeography dispersal-biogeography vicariance-biogeography center-of-origin area-of-endemism refugium Pleistocene glaciation climate-change habitat-fragmentation habitat-loss anthropogenic-impact pollution eutrophication sedimentation dam-construction water-abstraction flow-modification thermal-pollution toxic-substances heavy-metals pesticides urbanization agriculture deforestation land-use-change riparian-degradation channelization dredging mining invasive-species biological-invasion competition predation parasitism mutualism commensalism amensalism facilitation ecosystem-engineer keystone-species flagship-species umbrella-species indicator-species sentinel-species bioindicator ecological-indicator environmental-indicator water-quality-index biotic-index multimetric-index rapid-bioassessment protocol standardization quality-assurance quality-control reference-condition least-disturbed minimally-disturbed best-available type-specific region-specific stream-type river-type functional-feeding-group collector-gatherer collector-filterer scraper shredder predator parasite engulfer piercer burrower sprawler clinger swimmer skater diver surface-dweller benthic-dweller interstitial-dweller hyporheic-dweller phreatic-dweller troglobite stygobite cave-inhabitant spring-inhabitant headwater mid-order high-order lowland montane alpine subtropical tropical temperate boreal arctic antarctic palearctic nearctic afrotropical oriental australian oceanic cosmopolitan widespread restricted relict disjunct-distribution range-extension new-record first-record country-record state-record province-record locality-record elevation-record altitudinal-range latitudinal-range longitudinal-range distribution-pattern range-size range-fill extent-of-occurrence area-of-occupancy IUCN-category threat-status data-deficient not-evaluated least-concern near-threatened vulnerable endangered critically-endangered extinct-in-wild extinct population-trend population-density population-structure age-structure size-structure sex-ratio reproductive-output fecundity fertility mortality survivorship life-table cohort-analysis generation-time doubling-time intrinsic-rate-of-increase finite-rate-of-increase net-reproductive-rate stable-age-distribution stationary-population expanding-population declining-population source-sink-dynamics metapopulation population-viability-analysis minimum-viable-population genetic-diversity heterozygosity allelic-richness private-allele founder-effect bottleneck genetic-drift inbreeding inbreeding-depression outbreeding outbreeding-depression hybridization introgression gene-flow-barrier reproductive-isolation prezygotic-isolation postzygotic-isolation mechanical-isolation temporal-isolation habitat-isolation behavioral-isolation gametic-isolation zygotic-mortality hybrid-inviability hybrid-sterility hybrid-breakdown reinforcement character-displacement ecological-speciation allopatric-speciation sympatric-speciation parapatric-speciation peripatric-speciation centrifugal-speciation divergence-with-gene-flow speciation-continuum anagenesis cladogenesis budding-speciation branching-speciation phyletic-gradualism punctuated-equilibrium stasis trend progression orthogenesis aristogenesis mosaic-evolution co-evolution arms-race Red-Queen-hypothesis evolutionary-stable-strategy frequency-dependent-selection density-dependent-selection r-selection K-selection bet-hedging iteroparity semelparity ploidy-level chromosome-number karyotype genome-size transposable-element gene-duplication neofunctionalization subfunctionalization pseudogene regulatory-evolution cis-regulatory trans-regulatory enhancer promoter silencer insulator transcription-factor signaling-pathway developmental-pathway gene-network epigenetic-modification DNA-methylation histone-modification chromatin-remodeling genomic-imprinting parental-effect maternal-effect paternal-effect cytoplasmic-inheritance mitochondrial-DNA chloroplast-DNA endosymbiont Wolbachia Rickettsia Spiroplasma Cardinium Arsenophonus Hamiltonella Regiella Serratia Buchnera Carsonella Portiera primary-endosymbiont secondary-endosymbiont facultative-symbiont obligate-symbiont mutualistic-symbiont parasitic-symbiont commensal-symbiont pathogen vector disease-transmission zoonosis emerging-infectious-disease biosecurity quarantine eradication control management conservation-biology restoration-ecology habitat-restoration river-restoration stream-restoration floodplain-restoration riparian-restoration wetland-restoration ecosystem-restoration passive-restoration active-restoration assisted-recovery managed-relocation assisted-colonization ex-situ-conservation in-situ-conservation protected-area national-park nature-reserve wildlife-refuge biosphere-reserve world-heritage-site ramsar-site important-bird-area key-biodiversity-area Alliance-for-Zero-Extinction EDGE-species evolutionarily-distinct globally-endangered conservation-prioritization systematic-conservation-planning Marxan Zonation conservation-opportunity conservation-cost conservation-benefit trade-off socioeconomic-factor stakeholder-engagement community-based-conservation adaptive-management monitoring-program long-term-study citizen-science public-participation environmental-education outreach science-communication knowledge-transfer capacity-building institutional-strengthening policy-development legislation regulation enforcement compliance incentive payment-for-ecosystem-services carbon-credit biodiversity-offset mitigation-banking conservation-easement land-trust private-land-conservation conservation-covenant stewardship sustainable-use sustainable-management ecosystem-based-management integrated-water-resources-management basin-management catchment-management landscape-management seascape-management connectivity-conservation corridor stepping-stone buffer-zone matrix-management countryside-biogeography reconciliation-ecology novel-ecosystem designer-ecosystem ecosystem-service provisioning-service regulating-service supporting-service cultural-service food-production water-supply raw-materials genetic-resources medicinal-resources ornamental-resources air-quality-regulation climate-regulation water-regulation erosion-regulation water-purification waste-treatment disease-regulation pest-regulation pollination seed-dispersal habitat-provision nutrient-cycling soil-formation primary-production recreation tourism aesthetic-value spiritual-value educational-value scientific-value inspiration sense-of-place cultural-heritage traditional-knowledge indigenous-knowledge local-knowledge ecosystem-service-valuation economic-valuation contingent-valuation choice-experiment benefit-transfer total-economic-value use-value non-use-value direct-use indirect-use option-value bequest-value existence-value willingness-to-pay willingness-to-accept cost-benefit-analysis cost-effectiveness-analysis multi-criteria-analysis decision-support uncertainty risk-assessment scenario-analysis sensitivity-analysis robustness-analysis adaptive-capacity resilience vulnerability exposure sensitivity coping-capacity transformation social-ecological-system complex-adaptive-system tipping-point regime-shift alternative-stable-state hysteresis path-dependence legacy-effect slow-variable fast-variable panarchy adaptive-cycle exploitation release reorganization foresight anticipatory-governance transformative-change sustainability-science future-earth interdisciplinary transdisciplinary problem-oriented solution-oriented action-oriented knowledge-co-production boundary-work boundary-object boundary-organization science-policy-interface science-society-interface knowledge-brokerage knowledge-exchange knowledge-integration mode-1-science mode-2-science triple-helix quadruple-helix responsible-innovation reflexive-governance adaptive-governance collaborative-governance polycentric-governance network-governance self-organization collective-action common-pool-resource tragedy-of-the-commons social-dilemma institutional-analysis institutional-design Ostrom-design-principles monitoring sanctioning conflict-resolution nested-enterprises proportionality collective-choice constitutional-choice operational-choice collective-action-theory game-theory prisoner's-dilemma chicken-game assurance-game coordination-game evolutionary-game-theory replicator-dynamics ESS Nash-equilibrium Pareto-optimality social-optimum market-failure government-failure voluntary-failure non-market-value public-good common-good club-good private-good rivalry excludability free-rider hold-up-problem moral-hazard adverse-selection information-asymmetry transaction-cost property-right common-property state-property private-property hybrid-property bundle-of-rights usufruct access withdrawal alienation tenure-security land-tenure water-tenure resource-tenure customary-tenure statutory-tenure legal-pluralism de-facto de-jure formal informal legality legitimacy recognition deviance corruption elite-capture social-capital bonding-capital bridging-capital linking-capital trust reciprocity reputation social-network social-embeddedness structural-hole brokerage closure cohesion density centrality betweenness degree eigenvector PageRank small-world scale-free random-network regular-network hierarchical-network modularity community-structure clustering assortativity homophily heterophily preferential-attachment fitness-model network-evolution network-dynamics temporal-network multilayer-network multiplex-network bipartite-network affiliation-network one-mode-network two-mode-network adjacency-matrix edge-list graph-theory vertex edge node link tie connection path cycle tree forest component giant-component percolation phase-transition criticality self-organized-criticality power-law heavy-tail Pareto-distribution lognormal-distribution exponential-distribution Poisson-distribution binomial-distribution normal-distribution Gaussian-distribution statistical-distribution probability-distribution cumulative-distribution probability-density-function cumulative-distribution-function survival-function hazard-function moment mean median mode variance standard-deviation coefficient-of-variation skewness kurtosis quantile percentile decile quartile interquartile-range range minimum maximum outlier robust-statistic parametric-test non-parametric-test hypothesis-testing null-hypothesis alternative-hypothesis significance-level p-value confidence-interval type-I-error type-II-error statistical-power effect-size sample-size sampling-design experimental-design observational-study quasi-experiment natural-experiment randomized-controlled-trial before-after-control-impact BACI impact-assessment environmental-impact-assessment strategic-environmental-assessment cumulative-impact-assessment social-impact-assessment health-impact-assessment life-cycle-assessment carbon-footprint water-footprint ecological-footprint material-flow-analysis substance-flow-analysis energy-analysis exergy-analysis emergy-analysis input-output-analysis environmentally-extended-input-output social-accounting-matrix computable-general-equilibrium agent-based-model system-dynamics-model Bayesian-network structural-equation-model path-analysis factor-analysis principal-component-analysis cluster-analysis discriminant-analysis canonical-correspondence-analysis redundancy-analysis non-metric-multidimensional-scaling ordination classification regression-analysis linear-regression nonlinear-regression generalized-linear-model generalized-additive-model mixed-effects-model multilevel-model hierarchical-model Bayesian-hierarchical-model state-space-model hidden-Markov-model time-series-analysis spatial-analysis geostatistics kriging variogram spatial-autocorrelation Moran's-I Geary's-c Getis-Ord-Gi local-indicator-of-spatial-association LISA hot-spot cold-spot spatial-cluster spatial-outlier edge-effect boundary-effect modifiable-areal-unit-problem MAUP ecological-fallacy atomistic-fallacy cross-scale-inference upscaling downscaling scaling-law allometric-scaling fractal fractal-dimension self-similarity multi-fractal lacunarity roughness texture pattern process pattern-process-relationship neutral-model null-model randomization-test permutation-test bootstrap jackknife cross-validation model-averaging model-comparison information-criterion Bayesian-model-averaging ensemble-modeling consensus-model prediction forecast projection scenario narrative-scenario quantitative-scenario storyline shared-socioeconomic-pathway representative-concentration-pathway emission-scenario climate-scenario impact-scenario adaptation-scenario mitigation-scenario transformation-scenario backcasting forecasting trend-extrapolation expert-judgment Delphi-method participatory-modeling stakeholder-analysis influence-diagram conceptual-model mental-model system-archetype leverage-point intervention-point policy-instrument economic-instrument regulatory-instrument voluntary-instrument information-instrument market-based-instrument command-and-control-instrument subsidy tax charge fee levy deposit-refund tradable-permit emission-trading carbon-tax biodiversity-banking conditional-cash-transfer community-based-natural-resource-management joint-forest-management co-management adaptive-co-management transboundary-management international-regime global-governance multilateral-environmental-agreement convention treaty cartagena-protocol nagoya-protocol Aichi-target sustainable-development-goal SDG post-2020-global-biodiversity-framework Kunming-Montreal-global-biodiversity-framework 30-by-30-target nature-based-solution NbS ecosystem-based-adaptation ecosystem-based-mitigation green-infrastructure blue-infrastructure natural-infrastructure hybrid-infrastructure climate-smart-agriculture climate-smart-landscape climate-smart-development low-carbon-development green-economy circular-economy bioeconomy blue-economy sustainable-consumption sustainable-production life-cycle-thinking cradle-to-cradle industrial-ecology industrial-symbiosis urban-metabolism socio-ecological-metabolism material-intensity energy-intensity carbon-intensity decoupling absolute-decoupling relative-decoupling resource-efficiency eco-efficiency eco-innovation green-technology clean-technology environmental-technology appropriate-technology participatory-technology technology-transfer technology-diffusion innovation-system national-innovation-system sectoral-innovation-system technological-innovation-system socio-technical-system socio-technical-transition multi-level-perspective niche-innovation regime landscape transition-management strategic-niche-management transition-experiment living-lab real-world-laboratory urban-living-lab rural-living-lab sustainability-transition fundamental-change system-innovation directionality reflexivity responsiveness anticipation learning-by-doing doing-by-learning social-learning organizational-learning policy-learning transformative-capacity capacity-development institutional-learning evolutionary-governance institutional-change path-dependency lock-in carbon-lock-in technological-lock-in institutional-lock-in behavioral-lock-in cultural-lock-in unlocking destabilization phase-out divestment stranded-asset just-transition transition-pathway transition-scenario deep-decarbonization net-zero carbon-neutral climate-neutral nature-positive biodiversity-positive restoration-positive regenerative circular doughnut-economics planetary-boundary safe-operating-space environmental-ceiling social-foundation intergenerational-equity intragenerational-equity distributive-justice procedural-justice recognition-justice environmental-justice climate-justice energy-justice water-justice food-justice spatial-justice territorial-justice ecological-justice species-justice intersectionality adaptation mitigation sustainability sustainable-development well-being human-development capability-approach human-security environmental-security climate-security water-security food-security energy-security livelihood-security disaster-risk-reduction climate-change-adaptation climate-change-mitigation loss-and-damage climate-finance green-climate-fund adaptation-fund global-environment-facility biodiversity-finance nature-finance blended-finance impact-investment ESG environmental-social-governance sustainable-finance green-bond social-bond sustainability-bond blue-bond nature-bond biodiversity-credit offset inset nature-based-credit ecosystem-service-credit payment-for-results results-based-payment conditional-payment performance-based-payment outcome-based-payment environmental-service-payment PSA PES REDD REDD+AFOLU LULUCF NAMA NAMC NDC NAP NBSAP national-adaptation-plan national-biodiversity-strategy-and-action-plan nationally-determined-contribution nationally-appropriate-mitigation-action intended-nationally-determined-contribution long-term-low-emission-development-strategy net-zero-target carbon-budget remaining-carbon-budget emission-gap adaptation-gap finance-gap implementation-gap ambition-loop ratchet-mechanism global-stocktake transparency-framework enhanced-transparency-framework common-but-differentiated-responsibilities respective-capabilities climate-equity climate-litigation climate-governance climate-diplomacy climate-negotiation UNFCCC COP CMP CMA SBSTA SBI IPCC IPBES UNCCD CBD CITES Ramsar World-Heritage Man-and-the-Biosphere MAB UNESCO FAO WHO ILO UNDP UNEP UN-Habitat World-Bank IMF WTO OECD GEF GCF AF multilateral-development-bank regional-development-bank national-development-bank development-finance-institution export-credit-agency sovereign-wealth-fund pension-fund insurance-company asset-manager asset-owner corporate-investor retail-investor impact-investor philanthropic-investor development-partner bilateral-donor multilateral-donor private-donor foundation NGO CSO civil-society-organization non-governmental-organization community-based-organization indigenous-peoples-organization women's-organization youth-organization faith-based-organization research-organization think-tank university academic-institution knowledge-institution capacity-building-institution training-institution extension-service advisory-service consulting-firm engineering-firm technology-provider equipment-supplier service-provider contractor operator maintainer utility municipality local-government provincial-government national-government federal-government ministry department agency authority commission board council committee parliament congress legislature executive judiciary media journalist reporter editor publisher broadcaster platform social-media influencer celebrity activist advocate campaigner mobilizer organizer networker connector bridge-builder boundary-spanner knowledge-broker science-communicator extension-agent development-practitioner policy-maker decision-maker planner manager administrator leader champion entrepreneur innovator intrapreneur social-entrepreneur systems-entrepreneur institutional-entrepreneur policy-entrepreneur political-entrepreneur business-model business-model-innovation social-business inclusive-business base-of-pyramid fortune-at-bottom-of-pyramid shared-value creating-shared-value stakeholder-capitalism purpose-driven-business B-Corp benefit-corporation social-enterprise cooperative mutual association collective commons commoning peer-production open-source creative-commons knowledge-commons digital-commons data-commons genetic-commons seed-commons water-commons fisheries-commons forest-commons pasture-commons land-commons urban-commons atmospheric-commons global-commons common-heritage-of-mankind common-concern-of-humankind restorative-justice multispecies-justice more-than-human anthropocene capitalocene plantationocene chthulucene symbiocene good-anthropocene great-acceleration great-transformation great-decoupling great-convergence great-divergence great-fragmentation great-integration great-transition great-work long-now deep-time futures-thinking future-studies strategic-foresight scenario-planning trend-analysis weak-signal wild-card black-swan grey-rhino blue-sky green-swan systemic-change structural-change cultural-change behavioral-change individual-change collective-change social-change organizational-change technological-change economic-change political-change ecological-change planetary-change earth-system-change biodiversity-change ocean-change cryosphere-change atmospheric-change biogeochemical-change nitrogen-cycle-change phosphorus-cycle-change carbon-cycle-change water-cycle-change hydrological-change soil-change vegetation-change ecosystem-change landscape-change seascape-change biosphere-change noosphere-change technosphere-change anthroposphere-change sociosphere-change econosphere-change politosphere-change culturosphere-change semiosphere-change infosphere-change digisphere-change cybersphere-change urban-sphere-change agricultural-sphere-change industrial-sphere-change extractive-sphere-change manufacturing-sphere-change service-sphere-change financial-sphere-change knowledge-sphere-change innovation-sphere-change creativity-sphere-change culture-sphere-change identity-sphere-change meaning-sphere-change value-sphere-change purpose-sphere-change vision-sphere-change mission-sphere-change strategy-sphere-change tactic-sphere-change operation-sphere-change action-sphere-change practice-sphere-change habit-sphere-change routine-sphere-change ritual-sphere-change 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