Blog WotD About Glossary

Müllerian-mimicry

Guides

Apheloria
cherry millipedes, flat-backed millipedes
Apheloria is a genus of large, chemically defended millipedes in the family Xystodesmidae, distributed across eastern North America. These millipedes are notable for producing hydrogen cyanide and benzaldehyde as defensive secretions, which imparts a characteristic cherry or almond odor. The genus participates in Müllerian mimicry rings in the Appalachian Mountains, with species displaying highly variable aposematic coloration involving black backgrounds with contrasting yellow, orange, red, or white markings. Species-level identification requires examination of male gonopod morphology due to extensive color polymorphism and convergent color patterns among co-occurring species.
millipede diplopod polydesmid xystodesmid chemical-defense cyanide benzaldehyde aposematic-coloration Müllerian-mimicry decomposer forest-floor eastern-North-America Appalachian-Mountains Ozark-Mountains Ouachita-Mountains gonopod-morphology mull-forming
Apheloriini
cherry millipedes
Apheloriini is a tribe of large, colorful flat-backed millipedes endemic to the temperate forests of eastern North America. All species produce hydrogen cyanide as a chemical defense, which generates a characteristic cherry or almond odor from benzaldehyde byproducts. Members display bright aposematic coloration warning predators of their toxicity. The tribe includes seven genera, with greatest diversity concentrated in the southern Appalachian Mountains. Species in this tribe participate in Müllerian mimicry rings, resulting in extreme intraspecific variation in color patterns.
millipede flat-backed-millipede chemical-defense cyanide aposematic-coloration Müllerian-mimicry Appalachian-fauna decomposer
Battus philenor philenor
Pipevine Swallowtail, Blue Swallowtail
Battus philenor philenor is a subspecies of the pipevine swallowtail butterfly found in North America. Adults display iridescent blue hindwings against a black background, serving as aposematic warning coloration derived from sequestered aristolochic acids from their host plants. The subspecies is univoltine to bivoltine with flight periods from late winter through autumn, peaking before July. Populations have shown resilience to drought conditions in California's Central Valley, in contrast to montane butterfly declines.
aposematic-coloration chemical-defense aristolochic-acid-sequestration Müllerian-mimicry Batesian-mimicry-model drought-resilience riparian-specialist host-discrimination oviposition-behavior climate-change-indicator
Battus polydamas polydamas
Polydamas Swallowtail, Gold Rim Swallowtail, Tailless Swallowtail
The Polydamas swallowtail is a tailless swallowtail butterfly distinguished by its black wings with red or pink spots and a characteristic gold band along the wing margins. Adults are active year-round in tropical regions and are notable for their complete lack of tail-like extensions on the hindwings—a unique trait among New World swallowtails. The species serves as a model for Batesian mimicry, with its aposematic coloration warning predators of its chemical defenses derived from toxic host plants.
aposematic-coloration Batesian-mimicry Müllerian-mimicry chemical-sequestration aristolochic-acids osmeterium specialist-herbivore tailless-swallowtail tropical-butterfly climate-change-response
Batyle suturalis
Sutured Longhorn Beetle, sutured flower longhorn
Batyle suturalis is a species of longhorned beetle in the family Cerambycidae, first described by Thomas Say in 1824. It is a member of the tribe Trachyderini, a group known for bright colors, diurnal activity, and attraction to flowers. The species has been documented across much of North America, with records from the United States and Mexico. Adults are frequently observed feeding on flower pollen and nectar, and the species exhibits coloration that may function in a Müllerian mimicry complex with soldier beetles and blister beetles.
Cerambycidae Trachyderini flower-visiting Müllerian-mimicry pollinator North-America
Bombus flavifrons
yellow-fronted bumble bee, yellowhead bumblebee
Bombus flavifrons is a robust bumble bee native to North America, distributed across much of Canada, Alaska, and the western contiguous United States. The species exhibits intraspecific color pattern variation across its range due to Müllerian mimicry. It inhabits high-altitude and high-latitude environments including tundra, taiga, and mountain forests and meadows. The species is host to the parasitic cuckoo bumble bee Bombus insularis.
bumble-bee pollinator western-North-America high-altitude Müllerian-mimicry eusocial genome-sequenced
Caenia dimidiata
Terminal Net-winged Beetle
Caenia dimidiata is a species of net-winged beetle in the family Lycidae, known as the Terminal Net-winged Beetle. It is found in North America, with records from Canada and the United States. The species exhibits the characteristic black-and-orange or red warning coloration typical of lycid beetles, which serves as aposematic signaling. It participates in Müllerian mimicry complexes with other toxic insects, including moths in the genus Pyromorpha.
net-winged-beetle Lycidae Müllerian-mimicry aposematic-coloration North-America forest-habitat
Calopteron
net-winged beetles
Calopteron is a genus of net-winged beetles in the family Lycidae, comprising approximately 13 described species. These beetles are characterized by their toxic chemical defenses and aposematic (warning) coloration, typically black and orange or red patterns. They serve as important models in mimicry complexes, including Müllerian associations with toxic moths such as Pyromorpha dimidiata and Lycomorpha pholus. The genus is widely distributed, with species occurring in eastern North America and extending into South America.
net-winged-beetle aposematic-coloration Müllerian-mimicry warning-coloration toxic-beetles
Calopteron discrepans
banded net-winged beetle
Calopteron discrepans is a colorful net-winged beetle in the family Lycidae, recognized by its striking orange elytra with black banding. The species occurs in moist woodland habitats across eastern North America, where adults are active during summer months. It is frequently confused with congeners C. reticulatum and C. terminale, requiring careful examination of antennal and ventral coloration for positive identification. The species participates in Müllerian mimicry complexes with other aposematically colored insects.
net-winged-beetle Lycidae aposematic-coloration Müllerian-mimicry woodland-beetle North-America
Calopteron terminale
End Band Net-wing, End Band Net-winged Beetle
Calopteron terminale is a net-winged beetle in the family Lycidae, distinguished from congeners by a transverse depression across the elytra. It is found in eastern North American deciduous woodlands, where adults are active from July through September. The species participates in a Müllerian mimicry complex with similarly colored toxic beetles and moths. Larvae develop under bark of dead and dying trees, preying on small arthropods.
beetle net-winged-beetle Lycidae North-America deciduous-forest mimicry Müllerian-mimicry aposematic-coloration orange-and-black summer-active predatory-larvae
Chauliognathus
soldier beetles, leatherwings
A genus of soldier beetles (family Cantharidae) comprising over 450 described species. Adults possess characteristically soft, flexible elytra that give rise to the common name 'leatherwings.' Many species exhibit red-and-black or orange-and-black coloration reminiscent of historical military uniforms. Adults are diurnal and frequently associated with flowers, where they feed on pollen and nectar. Larvae are primarily ground-dwelling predators. The genus occurs in the Americas and Australia, with notable species including the goldenrod soldier beetle (C. pensylvanicus) and the plague soldier beetle (C. lugubris).
soldier-beetles leatherwings pollinators predatory-larvae Müllerian-mimicry flower-visitors citizen-science-subject biocontrol-agents cantharidin-defense fungal-pathogen-hosts
Chauliognathus basalis
Colorado Soldier Beetle
Chauliognathus basalis, commonly known as the Colorado soldier beetle, is a species of soldier beetle in the family Cantharidae. It is found in North America and is particularly common on the plains. The species exhibits Müllerian mimicry with other toxic beetles, sharing a black and orange color pattern that advertises its chemical defenses. Adults are active during the day and frequently observed on flowers, especially those of Gutierrezia sarothrae (broom snakeweed).
soldier-beetle leatherwing beneficial-insect predator mimicry Müllerian-mimicry Cantharidae autumn-active flower-visitor
Chauliognathus deceptus
Chauliognathus deceptus is a species of soldier beetle in the family Cantharidae. It occurs in the foothills and mountains of western North America, where it replaces its close relative C. basalis. Adults display black and orange coloration and possess chemical defenses secreted from abdominal glands. The species participates in Müllerian mimicry with other toxic beetles sharing similar warning coloration.
soldier-beetle Cantharidae Müllerian-mimicry chemical-defense aposematic-coloration autumn-active mountain-species western-North-America
Crossidius discoideus blandi
Crossidius discoideus blandi is a subspecies of longhorned beetle in the family Cerambycidae. Adults exhibit Batesian mimicry, resembling toxic soldier beetles (Chauliognathus spp.) and blister beetles (Epicauta spp.) with their black and orange coloration. The species is associated with flowers of Gutierrezia sarothrae (broom snakeweed), where adults feed on pollen and nectar. Larvae bore in the root crowns of the host plant. This subspecies is part of a broader mimicry complex involving multiple beetle families in western North American habitats.
Batesian-mimicry Müllerian-mimicry flower-visiting-beetle root-borer autumn-activity western-North-America Gutierrezia-associate
Cycnia
Cycnia is a genus of tiger moths in the family Erebidae, erected by Jacob Hübner in 1818. The genus contains several North American species, including the well-studied dogbane tiger moth (Cycnia tenera), which is known for its sophisticated anti-bat acoustic defenses. Species in this genus are typically associated with dogbane and milkweed plants, sequestering cardiac glycosides for chemical protection.
tiger-moth acoustic-defense anti-bat dogbane milkweed cardiac-glycoside tymbal ultrasound jamming aposematism Müllerian-mimicry
Dasymutilla
velvet ants, cow killers
Dasymutilla is a genus of solitary wasps in the family Mutillidae, commonly known as velvet ants. The genus contains the majority of North American velvet ant species. Females are wingless, densely hairy, and often brightly colored in aposematic patterns of red, orange, or white against black. Males possess wings and may differ substantially in coloration from females, leading to historical taxonomic confusion. The genus is notable for females' potent stings, among the most painful of any insect, and for forming one of the world's largest known Müllerian mimicry complexes.
Mutillidae velvet-ant parasitoid sexual-dimorphism Müllerian-mimicry aposematic-coloration painful-sting wingless-female external-parasite ground-nesting-host
Dasymutilla bioculata
velvet ant
Dasymutilla bioculata is a species of velvet ant (family Mutillidae), a group of solitary parasitic wasps in which only males possess wings. This species was taxonomically consolidated in 2010, when molecular and morphological analysis demonstrated that 21 previously recognized species and subspecies were actually conspecific. The species exhibits strong sexual dimorphism: females are wingless and run rapidly across the ground searching for host nests, while males fly and visit flowers. Females possess a potent sting used for defense. The species is a parasitoid of ground-nesting sand wasps in the genera Bembix and Microbembex.
velvet-ant parasitoid aposematic sexual-dimorphism Müllerian-mimicry sand-wasp-parasite stridulation extrafloral-nectary-visitor
Dasymutilla klugii
Klug's velvet ant
Dasymutilla klugii, commonly known as Klug's velvet ant, is a species of parasitoid wasp in the family Mutillidae. Despite the common name, it is not an ant but a solitary wasp. Females are wingless and possess a potent sting, while males are winged. The species is the most frequently encountered velvet ant in Texas and has been documented using cicada-killer wasp nests (Sphecius grandis) as hosts for its larvae. The specific epithet honors German entomologist Johann C. F. Klug.
parasitoid solitary-wasp aposematic-coloration sexual-dimorphism Müllerian-mimicry
Dasymutilla magna
Dasymutilla magna is a species of velvet ant in the family Mutillidae, described by Cresson in 1865. Like other members of the genus, females are wingless and possess a powerful sting, while males are winged. The species belongs to a large Müllerian mimicry complex where numerous velvet ant species share bright aposematic coloration warning predators of their defensive capabilities. Specific biological details for this species remain poorly documented in published literature.
Mutillidae velvet-ant Hymenoptera aposematic-coloration Müllerian-mimicry wingless-female solitary-wasp parasitoid Dasymutilla North-America
Dasymutilla occidentalis
Common Eastern Velvet Ant, Red Velvet Ant, Cow Killer, Cow Ant, Eastern Velvet Ant
Dasymutilla occidentalis is a large, solitary parasitoid wasp in the family Mutillidae, commonly known as the eastern velvet ant or cow killer. Females are wingless, densely covered in velvety red and black hairs, and possess an extremely painful sting. Males are winged and less conspicuously colored. The species is an external parasitoid of ground-nesting bees and wasps, with females actively searching for host nests to lay eggs on or near developing larvae. Despite its common name, it is not an ant but a true wasp, and its sting, while intensely painful, cannot actually kill a cow.
velvet-ant parasitoid aposematic Müllerian-mimicry painful-sting wingless-female ground-nesting-host solitary-wasp
Dasymutilla parksi
Dasymutilla parksi is a species of velvet ant (family Mutillidae), a group of solitary parasitoid wasps in which females are wingless and males are winged. First described by Mickel in 1936, this species belongs to a genus containing over 450 North American species, many of which exhibit bright aposematic coloration advertising their potent sting. Like other velvet ants, females are known for their powerful defensive sting and erratic, rapid ground movement. The biology and host associations of D. parksi specifically remain poorly documented.
parasitoid aposematic Müllerian-mimicry velvet-ant wingless-female solitary-wasp
Dasymutilla thetis
minute thistledown velvet ant, Little Velvet Ant
Dasymutilla thetis, commonly known as the minute thistledown velvet ant, is a small velvet ant species endemic to Arizona. First described by Charles A. Blake in 1886 as Sphaerophthalma thetis, it was later transferred to Dasymutilla. The species is notable for its entirely white-furred females, which participate in a Müllerian mimicry complex among pale desert velvet ants. Its small size (approximately 7 mm) distinguishes it from larger thistledown velvet ants such as Dasymutilla gloriosa.
endemic velvet-ant Müllerian-mimicry white-coloration parasitoid Arizona-endemic
Dryadula phaetusa
Banded Orange Heliconian, Banded Orange, Orange Tiger
Dryadula phaetusa, the sole species in its genus, is a Neotropical heliconiine butterfly recognizable by its bold orange and black banded wing pattern. It ranges from Brazil to central Mexico, with occasional summer vagrants reaching central Florida. The species exhibits sexual dimorphism in color intensity and participates in Müllerian mimicry complexes due to its unpalatability to birds.
butterfly heliconiine Nymphalidae Neotropical Müllerian-mimicry mud-puddling unpalatable sexual-dimorphism vagrant
Eleodes
pinacate beetles, desert stink beetles
Eleodes is the largest genus of darkling beetles in North America, comprising approximately 200 species. These beetles are endemic to western North America, ranging from southern Canada to central Mexico, with some species introduced to Colombia. Commonly known as pinacate beetles or desert stink beetles, they are flightless due to fused elytra and vestigial hindwings. All species possess chemical defense glands that produce quinone compounds, and many exhibit distinctive head-standing behavior when threatened. The genus shows remarkable ecological diversity, with species occupying deserts, forests, grasslands, and caves.
darkling-beetle Tenebrionidae chemical-defense flightless desert-arthropod detritivore agricultural-pest Müllerian-mimicry cave-beetle western-North-America
Elytroleptus ignitus
Elytroleptus ignitus is a longhorned beetle species in the family Cerambycidae, described by John Lawrence LeConte in 1884. The species was revised taxonomically in 2013, with E. luteicollis synonymized under this name. It is one of three Elytroleptus species documented as predaceous on lycid beetles (family Lycidae), which it also mimics in appearance. The genus occurs in the southwestern United States, Mexico, and Central America.
Cerambycidae Cerambycinae Trachyderini lycid-mimic predatory-cerambycid LeConte-1884 southwestern-United-States Mexico Central-America Elytroleptus Batesian-mimicry Müllerian-mimicry net-winged-beetle-mimic
Epicauta
Burning Blister Beetles
Epicauta is a large genus of blister beetles (family Meloidae) comprising approximately 360 described species. The genus exhibits a nearly cosmopolitan distribution, with native species on all continents except Australia and Antarctica. Adult Epicauta feed on plant foliage and flowers, while larvae are predatory specialists on grasshopper eggs. The genus is notable for producing cantharidin, a toxic terpenoid compound that serves as a chemical defense and can be lethal to vertebrates if ingested in sufficient quantities. Many species are significant agricultural pests, particularly of alfalfa and other forage crops, with documented cases of crop failure and livestock mortality from contaminated hay.
blister-beetle cantharidin agricultural-pest grasshopper-egg-predator hypermetamorphosis livestock-toxicity Müllerian-mimicry alfalfa-pest traditional-medicine anti-cancer-compound North-American-diversity Arizona-biodiversity invasive-potential hay-contamination equine-toxicity
Epicauta pensylvanica
black blister beetle, black aster bug
Epicauta pensylvanica is a blister beetle species in the family Meloidae, commonly known as the black blister beetle or black aster bug. The species is characterized by its predominantly black coloration and is known to contain the defensive compound cantharidin, which can cause skin blistering upon contact. Adults are typically found on flowers of plants in the Asteraceae family, particularly snakeweed (Gutierrezia sarothrae). The species occurs across North America and has been documented as a pest of soybean foliage in agricultural settings.
blister-beetle Meloidae soybean-pest cantharidin Asteraceae-feeder North-America agricultural-pest chemical-defense Müllerian-mimicry
Epicauta stuarti
Epicauta stuarti is a blister beetle in the family Meloidae, characterized by a black and orange color pattern that participates in a Müllerian mimicry complex with soldier beetles (Cantharidae). The species is found in North America, with records from the southwestern United States including Colorado, Oklahoma, and New Mexico. Adults have been observed on flowers of Gutierrezia sarothrae (snakeweed) and are attracted to fermenting liquid baits in trapping studies. Like other Epicauta species, it produces cantharidin, a potent defensive chemical that causes blistering on skin and is toxic if ingested.
blister-beetle Meloidae Müllerian-mimicry aposematic-coloration cantharidin southwestern-US autumn-active flower-visitor
Euchaetes
tussock moths, milkweed tussock moths, milkweed tiger moths
Euchaetes is a genus of arctiine tussock moths in the family Erebidae, described by Thaddeus William Harris in 1841. The genus includes species commonly known as milkweed tussock moths or milkweed tiger moths, many of which are specialized feeders on milkweed plants (Asclepiadaceae). Caterpillars of well-known species such as E. egle sequester cardiac glycosides from their host plants, making them unpalatable to predators. Adults of some species have evolved ultrasonic acoustic defenses against bat predation, representing a notable example of acoustic aposematism.
tussock-moth milkweed aposematism acoustic-defense cardiac-glycoside-sequestration Müllerian-mimicry Arctiinae Erebidae Lepidoptera North-America Mexico Sonoran-desert genus-level-information-only
Euchaetes egle
Milkweed Tussock Moth, Milkweed Tiger Moth
Euchaetes egle is a tiger moth in the family Erebidae, commonly known as the milkweed tussock moth or milkweed tiger moth. The species is notable for its gregarious caterpillars that feed on milkweeds and dogbane, sequestering cardiac glycosides from their host plants for chemical defense. Adults produce ultrasonic clicks from tymbal organs to warn bats of their unpalatability, representing a rare case of acoustic aposematism. The species occurs across eastern North America with one generation annually in northern regions and two or more in southern areas.
aposematism acoustic-defense tymbal-organ Müllerian-mimicry cardiac-glycoside milkweed-specialist gregarious-feeding bat-predation native-pollinator-plant-interaction
Exapion ulicis
Gorse Seed Weevil
Exapion ulicis is a small seed-feeding weevil specialized on gorse (Ulex europaeus). Adults are light gray with a prominent snout roughly half the body length. The species is native to western Europe and has been introduced to New Zealand, California, Hawaii, and other regions as a biological control agent targeting invasive gorse populations. Larval feeding destroys seeds within pods, reducing plant spread, while adult feeding on stems and spines causes minor damage.
biological-control invasive-species-management seed-predator host-specialization temporal-isolation Brentidae Coleoptera Ulex-europaeus gorse western-Europe California New-Zealand Hawaii spring-activity oviposition-cues pod-fecundity parasitoid seed-destruction noctuid pest-control-agent established-introduction native-range-Europe introduced-range-Pacific Chile Belgium Oregon northern-California forest-gaps pastures natural-areas seed-loss intensity-of-attack reproductive-isolation ecological-specialization sympatric-divergence host-phenology pod-size seed-content female-choice feeding-damage round-holes stem-feeding spine-feeding larval-development pupal-period 6-8-weeks-larval-feeding 2-months-pupation within-pod-development seed-to-ovule-ratio mature-seeds intact-seeds parasitization-rate population-abundance fecund-pod-targeting whole-plant-traits flower-traits pod-traits choice-experiments common-garden Brittany Scotland Reunion host-availability seasonal-activity morphological-identification mitochondrial-sequences nuclear-sequences phylogeny host-races divergent-selection ecological-speciation sympatric-populations spring-infestation autumn-infestation introduced-with-gorse not-initially-introduced invasive-plant plant-insect-interactions oviposition-behavior feeding-behavior preference-patterns multiple-trait-use surface-cues internal-cues pod-characteristics host-choice plant-susceptibility invasion-biology entomological-experimentalis pan-pacific-entomologist diversity-journal ecological-entomology Coombs-2004 HEAR-photo-gallery iNaturalist GBIF NCBI Forster-1771 Curculionoidea Apionidae Fabaceae Genisteae noxious-weed agent-of-biological-pest-control light-gray long-snout straight-snout 2-3-mm small-weevil seed-weevil gorse-seed-weevil established-populations percentage-of-pods-infested spatial-variation temporal-variation seed-set seed-production seed-predation-impact ecosystem-service weed-biological-control classical-biological-control inundative-biological-control self-sustaining-populations non-target-effects host-specificity reproductive-phenology phenological-matching phenological-isolation allochronic-speciation temporal-niche resource-partitioning competitive-exclusion character-displacement molecular-phylogenetics COI ITS morphological-characters individual-variation environmental-variation mitochondrial-DNA nuclear-DNA species-differentiation genetic-divergence ecological-divergence adaptive-radiation host-shift host-race-formation sympatric-speciation reinforcement prezygotic-isolation postzygotic-isolation habitat-fragmentation population-structure gene-flow local-adaptation phenotypic-plasticity genetic-drift natural-selection sexual-selection founder-effect bottleneck invasion-genetics enemy-release evolutionary-potential rapid-evolution contemporary-evolution eco-evolutionary-dynamics community-ecology trophic-interactions food-web ecosystem-function ecosystem-process nutrient-cycling energy-flow population-dynamics metapopulation source-sink dispersal colonization establishment spread impact-assessment non-target-risk efficacy establishment-success population-establishment field-release monitoring evaluation adaptive-management integrated-pest-management sustainable-agriculture conservation-biological-control augmentative-biological-control inoculative-biological-control natural-enemy herbivore phytophagous monophagous oligophagous polyphagous specialist generalist co-evolution coevolutionary-arms-race plant-defense herbivore-offense induced-defense constitutive-defense direct-defense indirect-defense volatile-organic-compounds extrafloral-nectaries trophic-cascades apparent-competition enemy-mediated-apparent-competition intraguild-predation multitrophic-interactions trait-mediated-indirect-effects density-mediated-indirect-effects ecosystem-engineering niche-construction extended-phenotype environmental-feedback eco-evolutionary-feedback evolutionary-rescue genetic-rescue assisted-migration managed-relocation conservation-translocation reintroduction restoration-ecology habitat-restoration ecological-restoration weed-management rangeland-management forest-management protected-area-management biodiversity-conservation ecosystem-services provisioning-services regulating-services cultural-services supporting-services sustainable-development-goals Aichi-targets Kunming-Montreal-global-biodiversity-framework IPBES IUCN CBD FAO EPPO regulatory-framework risk-analysis environmental-impact-assessment cost-benefit-analysis decision-support stakeholder-engagement social-license public-acceptance communication education outreach citizen-science iNaturalist-observations occurrence-records specimen-records museum-collections voucher-specimens type-specimens holotype paratype syntype lectotype neotype original-description subsequent-designation taxonomic-revision phylogenetic-revision monograph field-guide identification-key diagnostic-characters illustration photography microscopy scanning-electron-microscopy molecular-techniques DNA-barcoding metabarcoding environmental-DNA population-genetics phylogeography landscape-genetics seascape-genetics isolation-by-distance isolation-by-environment isolation-by-resistance least-cost-path circuit-theory species-distribution-modeling ecological-niche-modeling maxent bioclim worldclim chelsa modis remote-sensing GIS spatial-analysis temporal-analysis time-series phenology climate-change global-warming phenological-shift range-shift poleward-shift elevational-shift altitudinal-shift habitat-tracking niche-tracking niche-conservatism niche-evolution realized-niche fundamental-niche biotic-interactions abiotic-factors environmental-filtering dispersal-limitation source-pool species-pool regional-pool local-pool alpha-diversity beta-diversity gamma-diversity species-richness species-evenness species-abundance dominance rarity commonness endemism cosmopolitanism native endemic introduced invasive naturalized casual transient ephemeral persistent established spreading invasive-potential invasiveness impact ecological-impact economic-impact social-impact environmental-impact negative-impact positive-impact beneficial detrimental nuisance pest weed pathogen disease-vector public-health veterinary-health crop-health forest-health rangeland-health ecosystem-health One-Health planetary-health ecohealth conservation-medicine disease-ecology parasitology entomology medical-entomology veterinary-entomology agricultural-entomology forest-entomology urban-entomology aquatic-entomology systematics taxonomy phylogenetics evolutionary-biology population-biology ecosystem-ecology landscape-ecology global-ecology macroecology biogeography historical-biogeography ecological-biogeography island-biogeography continental-biogeography latitudinal-gradient altitudinal-gradient species-area-relationship island-species-richness equilibrium-theory nonequilibrium-theory neutral-theory niche-theory assembly-rules priority-effects mass-effects species-sorting competition predation herbivory parasitism mutualism commensalism amensalism facilitation inhibition tolerance succession primary-succession secondary-succession disturbance recovery resilience resistance stability persistence turnover replacement extinction immigration emigration birth death reproduction survival growth development metamorphosis complete-metamorphosis holometabolous hemimetabolous ametabolous egg larva pupa adult instar ecdysis molting diapause quiescence aestivation hibernation overwintering overwintering-stage overwintering-site overwintering-strategy cold-hardiness freeze-tolerance freeze-avoidance supercooling cryoprotectants thermal-biology thermoregulation ectothermy poikilothermy behavioral-thermoregulation microhabitat-selection basking shading stilting perching roosting sheltering burrowing mining gall-formation leaf-rolling leaf-tying silk-production web-spinning nest-construction oviposition-site-selection host-plant-selection mate-choice sperm-competition cryptic-female-choice polyandry polygyny monogamy promiscuity lekking territoriality aggression defense predator-avoidance antipredator-behavior crypsis masquerade mimicry Batesian-mimicry Müllerian-mimicry automimicry aggressive-mimicry startle-display thanatosis death-feigning autotomy reflex-bleeding chemical-defense mechanical-defense physical-defense behavioral-defense mutualistic-defense plant-animal-interactions pollination seed-dispersal antagonism synergism antagonistic-pleiotropy trade-offs life-history life-history-strategy r-selection K-selection bet-hedging iteroparity semelparity annual perennial biennial pluriannual longevity lifespan generation-time development-time growth-rate reproductive-rate fecundity fertility virility mating-success reproductive-success fitness inclusive-fitness kin-selection group-selection multilevel-selection individual-selection artificial-selection social-selection ecological-selection stabilizing-selection directional-selection disruptive-selection balancing-selection frequency-dependent-selection negative-frequency-dependent-selection positive-frequency-dependent-selection apostatic-selection antiapostatic-selection rare-type-advantage common-type-disadvantage heterozygote-advantage heterosis hybrid-vigor inbreeding-depression outbreeding-depression genetic-load mutational-load segregational-load substitutional-load drift-load migration-load mutation recombination chromosomal-rearrangement genome-evolution transposable-elements horizontal-gene-transfer endosymbiosis symbiogenesis holobiont hologenome extended-evolutionary-synthesis niche-construction-theory ecological-evolutionary-developmental-biology eco-evo-devo developmental-plasticity reaction-norm genotype-environment-interaction GxE norm-of-reaction plasticity canalization genetic-assimilation genetic-accommodation cryptic-genetic-variation evolutionary-capacitance evolvability robustness modularity integration correlation constraint trade-off pleiotropy epistasis additive-genetic-variance dominance-variance epistatic-variance genetic-variance phenotypic-variance environmental-variance heritability narrow-sense-heritability broad-sense-heritability response-to-selection breeder's-equation quantitative-genetics molecular-genetics genomics transcriptomics proteomics metabolomics phenomics ecogenomics conservation-genomics functional-genomics structural-genomics comparative-genomics evolutionary-genomics phylogenomics metagenomics microbiome virome mycobiome bacteriome archaeome eukaryome extended-genotype keystone-species ecosystem-engineer foundation-species dominant-species subordinate-species transient-species satellite-species core-species peripheral-species edge-species interior-species habitat-generalist habitat-specialist edge-habitat interior-habitat matrix-habitat corridor stepping-stone source-habitat sink-habitat patch fragment landscape seascape riverscape skyscape soundscape smellscape tastes touch vision olfaction gustation mechanoreception thermoreception photoreception chemoreception electroreception magnetoreception proprioception nociception sensory-ecology sensory-biology neuroethology behavioral-ecology cognitive-ecology evolutionary-ecology ecological-immunology ecoimmunology epidemiology parasite-ecology host-parasite-interactions host-pathogen-interactions host-symbiont-interactions microbial-ecology virology bacteriology mycology protistology helminthology entomopathology microbial-control viral-control fungal-control bacterial-control nematode-control parasitoid-control predator-control herbivore-control competitor-control autocidal-control sterile-insect-technique incompatible-insect-technique gene-drive RNA-interference CRISPR genetic-biocontrol precision-guided-sterile-insect-technique self-limiting-genetic-control self-sustaining-genetic-control population-suppression population-replacement population-elimination area-wide-integrated-pest-management sterile-insect-release inundative-release inoculative-release neoclassical-biological-control augmentation-biological-control habitat-manipulation cultural-control physical-control mechanical-control chemical-control pesticide-resistance-management insecticide-resistance-management herbicide-resistance-management fungicide-resistance-management antibiotic-resistance-management resistance-evolution resistance-management refuge-strategy high-dose-strategy pyramid-strategy rotation-strategy mixture-strategy mosaic-strategy integrated-resistance-management integrated-weed-management integrated-crop-management integrated-forest-management integrated-vector-management integrated-disease-management One-Health-approach ecohealth-approach planetary-health-approach sustainable-intensification agroecology organic-agriculture conservation-agriculture climate-smart-agriculture precision-agriculture digital-agriculture smart-farming agricultural-robotics autonomous-systems drone-technology satellite-imagery GPS variable-rate-technology site-specific-management decision-support-systems expert-systems artificial-intelligence machine-learning deep-learning neural-networks computer-vision image-recognition pattern-recognition data-mining big-data cloud-computing internet-of-things sensor-networks wireless-sensor-networks precision-livestock-farming precision-aquaculture precision-forestry precision-horticulture precision-viticulture precision-apiculture precision-sericulture integrated-farming-systems mixed-farming-systems diversified-farming-systems agroforestry silvopasture silvoarable alley-cropping forest-farming homegardens multistrata-systems tropical-homegardens temperate-agroforestry permaculture regenerative-agriculture restorative-agriculture biodynamic-agriculture natural-farming do-nothing-farming Fukuoka-farming Masanobu-Fukuoka Rachel-Carson Silent-Spring integrated-pest-management-history biological-control-history classical-biological-control-history Rachel-Carson-legacy environmental-movement conservation-movement sustainability-science resilience-science adaptive-governance social-ecological-systems socio-ecological-systems coupled-human-natural-systems complex-adaptive-systems panarchy adaptive-cycle rigidity-trap poverty-trap transformation transition sustainability-transition energy-transition food-system-transformation agrifood-system-transformation land-use-change land-cover-change land-use-intensification agricultural-expansion deforestation reforestation afforestation forest-degradation forest-restoration ecosystem-restoration wetland-restoration riparian-restoration coastal-restoration coral-reef-restoration seagrass-restoration mangrove-restoration salt-marsh-restoration dune-restoration prairie-restoration savanna-restoration woodland-restoration old-growth-restoration degraded-land-restoration abandoned-land-restoration contaminated-land-restoration mined-land-restoration post-industrial-restoration urban-restoration rural-restoration agricultural-restoration pastoral-restoration silvicultural-restoration ecological-engineering environmental-engineering bioremediation phytoremediation mycoremediation zooremediation microbial-remediation nanoremediation electrokinetic-remediation soil-washing soil-vapor-extraction air-sparging pump-and-treat permeable-reactive-barriers monitored-natural-attenuation enhanced-natural-attenuation source-zone-treatment plume-treatment risk-based-corrective-action risk-assessment hazard-assessment exposure-assessment toxicity-assessment risk-characterization risk-management risk-communication risk-perception risk-governance precautionary-principle polluter-pays-principle extended-producer-responsibility circular-economy green-economy blue-economy bioeconomy natural-capital ecosystem-services-valuation payment-for-ecosystem-services ecosystem-based-adaptation ecosystem-based-mitigation nature-based-solutions working-with-nature building-with-nature engineering-with-nature green-infrastructure blue-infrastructure urban-green-infrastructure urban-blue-infrastructure sponge-city cool-city forest-city biophilic-city sustainable-city resilient-city smart-city liveable-city healthy-city equitable-city just-city inclusive-city participatory-city co-production co-design co-management community-based-management community-based-conservation community-based-natural-resource-management indigenous-and-local-knowledge traditional-ecological-knowledge biocultural-diversity biocultural-heritage sacred-natural-sites protected-areas conservation-areas nature-reserves wildlife-reserves national-parks world-heritage-sites biosphere-reserves Ramsar-sites important-bird-areas key-biodiversity-areas ecologically-or-biologically-significant-marine-areas other-effective-area-based-conservation-measures conservation-beyond-protected-areas conservation-on-private-land conservation-on-communal-land conservation-on-indigenous-land rights-based-conservation gender-responsive-conservation youth-engagement-in-conservation intergenerational-equity intragenerational-equity environmental-justice climate-justice conservation-justice ecological-justice species-justice intersectionality decolonization-of-conservation post-normal-science transdisciplinarity participatory-research action-research community-based-participatory-research community-science participatory-monitoring adaptive-co-management social-learning knowledge-co-production boundary-work boundary-objects boundary-organizations science-policy-interface science-society-interface knowledge-brokering knowledge-translation knowledge-mobilization research-impact research-uptake evidence-based-policy evidence-based-practice evidence-informed-decision-making decision-support-tools systematic-review meta-analysis evidence-synthesis knowledge-synthesis rapid-evidence-assessment living-evidence living-systematic-review scoping-review rapid-review realist-synthesis narrative-synthesis qualitative-synthesis quantitative-synthesis mixed-methods-synthesis participatory-synthesis deliberative-methods consensus-methods expert-elicitation structured-expert-judgment Delphi-method nominal-group-technique convergent-interview focus-group stakeholder-workshop participatory-mapping participatory-modeling scenario-planning future-studies foresight anticipatory-governance transformative-governance polycentric-governance multi-level-governance network-governance collaborative-governance cooperative-governance democratic-governance deliberative-democracy participatory-democracy direct-democracy representative-democracy liberal-democracy social-democracy green-democracy ecological-democracy biocracy ecocracy technocracy meritocracy plutocracy oligarchy autocracy totalitarianism authoritarianism populism nationalism globalism localism regionalism federalism confederalism unitarism centralism decentralization devolution subsidiarity fiscal-federalism environmental-federalism competitive-federalism cooperative-federalism dual-federalism marble-cake-federalism layer-cake-federalism picket-fence-federalism permissive-federalism coercive-federalism market-preserving-federalism democratic-experimentalism laboratories-of-democracy policy-diffusion policy-transfer policy-learning policy-innovation policy-entrepreneurship policy-windows punctuated-equilibrium advocacy-coalition-framework multiple-streams-framework institutional-analysis-and-development-framework social-ecological-systems-framework sustainability-science-framework planetary-boundaries-framework doughnut-economics-framework circular-economy-framework green-new-deal-framework just-transition-framework leaving-no-one-behind 2030-agenda Paris-agreement Rio-conventions UNFCCC UNCCD CITES CMS Ramsar World-Heritage IPCC WHO UNEP UNDP UNESCO World-Bank IMF WTO OECD G7 G20 BRICS African-Union European-Union ASEAN Mercosur NAFTA USMCA TPP CPTPP RCEP bilateral-investment-treaties free-trade-agreements regional-trade-agreements multilateral-environmental-agreements bilateral-environmental-agreements strategic-environmental-assessment cumulative-impact-assessment social-impact-assessment health-impact-assessment gender-impact-assessment human-rights-impact-assessment indigenous-rights-impact-assessment cultural-heritage-impact-assessment landscape-and-visual-impact-assessment noise-impact-assessment air-quality-impact-assessment water-quality-impact-assessment soil-quality-impact-assessment biodiversity-impact-assessment ecosystem-services-impact-assessment climate-impact-assessment carbon-footprint-assessment water-footprint-assessment land-footprint-assessment material-footprint-assessment ecological-footprint-assessment planetary-footprint-assessment life-cycle-assessment life-cycle-costing social-life-cycle-assessment life-cycle-sustainability-assessment material-flow-analysis substance-flow-analysis energy-flow-analysis nutrient-flow-analysis water-flow-analysis carbon-flow-analysis nitrogen-flow-analysis phosphorus-flow-analysis circular-material-flow industrial-ecology industrial-symbiosis eco-industrial-park urban-metabolism industrial-metabolism socio-economic-metabolism social-metabolism political-ecology political-economy ecological-economics environmental-economics natural-resource-economics resource-economics energy-economics agricultural-economics forest-economics fisheries-economics water-economics biodiversity-economics ecosystem-services-economics conservation-economics restoration-economics environmental-valuation contingent-valuation choice-experiment benefit-transfer hedonic-pricing travel-cost-method production-function-approach replacement-cost-method damage-cost-avoided-method opportunity-cost-method net-present-value internal-rate-of-return benefit-cost-ratio cost-effectiveness-analysis multi-criteria-decision-analysis analytic-hierarchy-process analytic-network-process preference-ranking-organization-method-for-enrichment-evaluation technique-for-order-of-preference-by-similarity-to-ideal-solution elimination-and-choice-expressing-reality decision-making-trial-and-evaluation-laboratory VIseKriterijumska-Optimizacija-I-Kompromisno-Resenje preference-selection-index complex-proportional-assessment multi-objective-optimization-on-the-basis-of-ratio-analysis weighted-aggregated-sum-product-assessment evaluation-based-on-distance-from-average-solution measurement-of-alternatives-and-ranking-according-to-compromise-solution comparative-performance-index preference-ranking score-function accuracy-function entropy-weight objective-weight subjective-weight combined-weight sensitivity-analysis uncertainty-analysis fuzzy-logic fuzzy-set-theory intuitionistic-fuzzy-set neutrosophic-set rough-set soft-set hesitant-fuzzy-set Pythagorean-fuzzy-set q-rung-orthopair-fuzzy-set spherical-fuzzy-set picture-fuzzy-set complex-fuzzy-set type-2-fuzzy-set interval-type-2-fuzzy-set interval-valued-fuzzy-set interval-valued-intuitionistic-fuzzy-set interval-valued-Pythagorean-fuzzy-set interval-valued-neutrosophic-set bipolar-fuzzy-set m-polar-fuzzy-set multi-polar-fuzzy-set linguistic-term-set hesitant-fuzzy-linguistic-term-set probabilistic-linguistic-term-set 2-tuple-linguistic-model symbolic-linguistic-model virtual-linguistic-model numerical-scale-model linguistic-hierarchy-model transitive-calibration-model consistency-driven-methodology consistency-index consistency-ratio acceptable-consistency inconsistency-identification inconsistency-repair priority-derivation weight-derivation aggregation-operator ordered-weighted-averaging-operator weighted-averaging-operator geometric-operator power-operator Bonferroni-mean Choquet-integral Shapley-value game-theory Nash-equilibrium Pareto-optimality Pareto-efficiency Pareto-improvement social-welfare-function utility-function preference-relation binary-relation ternary-relation n-ary-relation complete-relation incomplete-relation transitive-relation intransitive-relation reflexive-relation irreflexive-relation symmetric-relation asymmetric-relation antisymmetric-relation equivalence-relation partial-order total-order strict-order weak-order semiorder interval-order Pareto-order lexicographic-order dominance-relation outranking-relation concordance discordance credibility discrimination-threshold indifference-threshold preference-threshold veto-threshold majority-rule Condorcet-winner Condorcet-loser Borda-count approval-voting range-voting score-voting instant-runoff-voting single-transferable-vote proportional-representation mixed-member-proportional additional-member-system parallel-voting majority-bonus-system scorporo double-vote alternative-vote supplementary-vote contingent-vote Bucklin-voting Coombs'-method Dodgson's-method Kemeny-Young-method Ranked-pairs Schulze-method Minimax-Condorcet Smith-set Schwartz-set uncovered-set Banks-set top-cycle minimal-covering-set bipartisan-set essential-set tournament-equilibrium-set minimal-extending-set Pareto-set deep-uncovered-set minimal-upward-covering-set minimal-downward-covering-set minimal-covering-set-with-respect-to-pairs minimal-covering-set-with-respect-to-triplets minimal-stable-set minimal-retentive-set minimal-comprehensive-retentive-set minimal-extensive-retentive-set minimal-weakly-stable-set minimal-strongly-stable-set minimal-collectively-stable-set minimal-individually-stable-set minimal-Nash-stable-set minimal-individually-rational-set minimal-Pareto-optimal-set minimal-weakly-Pareto-optimal-set minimal-strongly-Pareto-optimal-set minimal-collectively-Pareto-optimal-set minimal-individually-Pareto-optimal-set minimal-Nash-Pareto-optimal-set minimal-individually-rational-Pareto-optimal-set minimal-comprehensive-Pareto-optimal-set minimal-extensive-Pareto-optimal-set minimal-weakly-comprehensive-Pareto-optimal-set minimal-strongly-comprehensive-Pareto-optimal-set minimal-collectively-comprehensive-Pareto-optimal-set minimal-individually-comprehensive-Pareto-optimal-set minimal-Nash-comprehensive-Pareto-optimal-set minimal-individually-rational-comprehensive-Pareto-optimal-set minimal-weakly-extensive-Pareto-optimal-set minimal-strongly-extensive-Pareto-optimal-set minimal-collectively-extensive-Pareto-optimal-set minimal-individually-extensive-Pareto-optimal-set minimal-Nash-extensive-Pareto-optimal-set minimal-individually-rational-extensive-Pareto-optimal-set minimal-weakly-retentive-Pareto-optimal-set minimal-strongly-retentive-Pareto-optimal-set minimal-collectively-retentive-Pareto-optimal-set minimal-individually-retentive-Pareto-optimal-set minimal-Nash-retentive-Pareto-optimal-set minimal-individually-rational-retentive-Pareto-optimal-set minimal-weakly-stable-Pareto-optimal-set minimal-strongly-stable-Pareto-optimal-set minimal-collectively-stable-Pareto-optimal-set minimal-individually-stable-Pareto-optimal-set minimal-Nash-stable-Pareto-optimal-set minimal-individually-rational-stable-Pareto-optimal-set minimal-weakly-Nash-stable-Pareto-optimal-set minimal-strongly-Nash-stable-Pareto-optimal-set minimal-collectively-Nash-stable-Pareto-optimal-set minimal-individually-Nash-stable-Pareto-optimal-set minimal-Nash-Nash-stable-Pareto-optimal-set minimal-individually-rational-Nash-stable-Pareto-optimal-set minimal-weakly-individually-rational-Pareto-optimal-set minimal-strongly-individually-rational-Pareto-optimal-set minimal-collectively-individually-rational-Pareto-optimal-set minimal-individually-individually-rational-Pareto-optimal-set minimal-Nash-individually-rational-Pareto-optimal-set minimal-individually-rational-individually-rational-Pareto-optimal-set minimal-weakly-Pareto-optimal-Nash-stable-set minimal-strongly-Pareto-optimal-Nash-stable-set minimal-collectively-Pareto-optimal-Nash-stable-set minimal-individually-Pareto-optimal-Nash-stable-set minimal-Nash-Pareto-optimal-Nash-stable-set minimal-individually-rational-Pareto-optimal-Nash-stable-set minimal-weakly-comprehensive-Pareto-optimal-Nash-stable-set minimal-strongly-comprehensive-Pareto-optimal-Nash-stable-set minimal-collectively-comprehensive-Pareto-optimal-Nash-stable-set minimal-individually-comprehensive-Pareto-optimal-Nash-stable-set minimal-Nash-comprehensive-Pareto-optimal-Nash-stable-set minimal-individually-rational-comprehensive-Pareto-optimal-Nash-stable-set minimal-weakly-extensive-Pareto-optimal-Nash-stable-set minimal-strongly-extensive-Pareto-optimal-Nash-stable-set minimal-collectively-extensive-Pareto-optimal-Nash-stable-set minimal-individually-extensive-Pareto-optimal-Nash-stable-set minimal-Nash-extensive-Pareto-optimal-Nash-stable-set minimal-individually-rational-extensive-Pareto-optimal-Nash-stable-set minimal-weakly-retentive-Pareto-optimal-Nash-stable-set minimal-strongly-retentive-Pareto-optimal-Nash-stable-set minimal-collectively-retentive-Pareto-optimal-Nash-stable-set minimal-individually-retentive-Pareto-optimal-Nash-stable-set minimal-Nash-retentive-Pareto-optimal-Nash-stable-set minimal-individually-rational-retentive-Pareto-optimal-Nash-stable-set minimal-weakly-stable-Pareto-optimal-Nash-stable-set minimal-strongly-stable-Pareto-optimal-Nash-stable-set minimal-collectively-stable-Pareto-optimal-Nash-stable-set minimal-individually-stable-Pareto-optimal-Nash-stable-set minimal-Nash-stable-Pareto-optimal-Nash-stable-set minimal-individually-rational-stable-Pareto-optimal-Nash-stable-set minimal-weakly-Nash-stable-Pareto-optimal-Nash-stable-set minimal-strongly-Nash-stable-Pareto-optimal-Nash-stable-set minimal-collectively-Nash-stable-Pareto-optimal-Nash-stable-set minimal-individually-Nash-stable-Pareto-optimal-Nash-stable-set minimal-Nash-Nash-stable-Pareto-optimal-Nash-stable-set minimal-individually-rational-Nash-stable-Pareto-optimal-Nash-stable-set minimal-weakly-individually-rational-Pareto-optimal-Nash-stable-set minimal-strongly-individually-rational-Pareto-optimal-Nash-stable-set minimal-collectively-individually-rational-Pareto-optimal-Nash-stable-set minimal-individually-individually-rational-Pareto-optimal-Nash-stable-set minimal-Nash-individually-rational-Pareto-optimal-Nash-stable-set minimal-individually-rational-individually-rational-Pareto-optimal-Nash-stable-set minimal-weakly-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set
Heliconiini
longwings, passion-vine butterflies
Heliconiini is a tribe of butterflies in the subfamily Heliconiinae, commonly known as longwings or passion-vine butterflies. The group comprises approximately 100 species and subspecies distributed primarily in the Neotropics. Members are characterized by elongated wings, bright aposematic coloration, and specialized relationships with Passiflora host plants. The tribe includes well-known genera such as Heliconius, Dryas, and Agraulis.
Neotropical aposematic-coloration Müllerian-mimicry Passiflora-specialist sound-production pupal-behavior
Heliconius
longwings, heliconians
Heliconius is a genus of brush-footed butterflies renowned for their bright aposematic wing patterns and Müllerian mimicry rings. Adults are unique among butterflies in their ability to collect and digest pollen, which provides essential amino acids enabling lifespans of up to six months—far exceeding most butterflies. The genus serves as a premier model system for studying speciation, mimicry evolution, and coevolutionary dynamics with their host plants. Approximately 45–50 species occur across the Neotropics, exhibiting complex patterns of wing color convergence and divergence.
pollen-feeding Müllerian-mimicry coevolution speciation-model aposematism cyanogenic-defense extended-lifespan traplining pupal-mating Neotropical
Heliconius erato
red postman, small postman, red passion flower butterfly, crimson-patched longwing
Heliconius erato is a neotropical butterfly renowned for its exceptional longevity among butterflies, living up to six months in the wild and over 186 days in captivity. This extended lifespan is enabled by its unique ability among butterflies to digest pollen, extracting amino acids through specialized proboscis behavior. The species exhibits Müllerian mimicry with co-occurring Heliconius species, particularly H. melpomene, sharing aposematic warning coloration to deter predators. Adults display remarkable philopatry, maintaining restricted home ranges sometimes separated by only 30 yards from neighboring territories. The species was described by Linnaeus in 1758 and comprises numerous geographically variable subspecies divided into eastern and western Andean clades.
pollen-feeding Müllerian-mimicry longevity philopatry trap-lining anti-aphrodisiac cyanogenic-defense Heliconius neotropical Passiflora-specialist
Ithomiini
clearwings, glasswings, tigerwings
Ithomiini is a diverse tribe of approximately 370 species in 40–45 genera, endemic to the Neotropics from Mexico to Argentina. These butterflies are renowned for their Müllerian mimicry rings, slow flight, and unpalatability derived from sequestered pyrrolizidine alkaloids. Adults actively seek out and sequester these defensive compounds from plants rather than synthesizing them de novo. The tribe represents the largest known radiation of Müllerian mimetic butterflies and dominates Neotropical mimetic butterfly communities by number.
Müllerian-mimicry chemical-defense pyrrolizidine-alkaloid-sequestration Neotropical gregarious-larvae silk-trail-following visual-system-diversity genomic-model-organism
Labidomera
milkweed leaf beetle
Labidomera is a genus of leaf beetles (Chrysomelidae) in the tribe Chrysomelini. The genus includes several species, with L. clivicollis being the most studied and widely distributed. Members are strongly associated with milkweed plants (Asclepias) and related genera. The genus exhibits unusual larval locomotion and participates in Müllerian mimicry rings with other orange-and-black milkweed insects.
leaf-beetle Chrysomelidae milkweed Müllerian-mimicry cardenolide-sequestration photoperiodism cannibalism sexual-transmission-of-parasites
Labidomera clivicollis
Swamp Milkweed Leaf Beetle, Milkweed Leaf Beetle
Labidomera clivicollis is a large leaf beetle specialized on milkweeds (Asclepias spp.). Adults and larvae feed on milkweed foliage, sequestering cardiac glycosides for chemical defense. The species exhibits aposematic orange and black coloration as part of a Müllerian mimicry complex with monarch butterflies and other milkweed feeders. Reproduction is photoperiodically controlled, with short day lengths inducing adult diapause. Larvae suffer high predation mortality, particularly from ground-dispersing predators.
leaf-beetle milkweed aposematic-coloration Müllerian-mimicry chemical-defense cardenolide-sequestration photoperiodism sexually-transmitted-parasite cannibalism predation-mortality swamp-milkweed common-milkweed Chrysomelidae herbivore diapause Kirby-1837
Limenitis archippus
Viceroy
The Viceroy (Limenitis archippus) is a North American butterfly famous for its striking resemblance to the Monarch butterfly (Danaus plexippus), a relationship long interpreted as Batesian mimicry but now considered Müllerian mimicry since both species are distasteful to predators. This medium-sized brushfoot butterfly ranges across most of the contiguous United States, southern Canada, and parts of Mexico. Adults are orange with black veining and white spots, but display significant regional variation in size and coloration—eastern populations are bright orange while western and Florida populations are darker, sometimes mimicking the Queen butterfly (Danaus gilippus) instead. The species is territorial, with males actively patrolling and perching to defend areas.
butterfly mimicry brushfoot North-America Müllerian-mimicry territorial-behavior
Limenitis archippus floridensis
Florida Viceroy
Limenitis archippus floridensis, commonly known as the Florida Viceroy, is a subspecies of the Viceroy butterfly found in the southeastern United States. This butterfly is renowned for its mimicry of the Monarch butterfly (Danaus plexippus), though recent research indicates it is also unpalatable to predators, making this Müllerian rather than Batesian mimicry. The Florida subspecies is notably larger and darker than typical Viceroys, adaptations that allow it to mimic the Queen butterfly (Danaus gilippus) which replaces the Monarch as the model species in Florida. Males are highly territorial, patrolling areas and perching frequently—a behavior not observed in Monarchs.
subspecies mimicry Florida Southeastern-United-States Lepidoptera butterfly territorial-behavior Müllerian-mimicry
Limenitis archippus lahontani
Nevada Viceroy
Limenitis archippus lahontani is a subspecies of the Viceroy butterfly, commonly known as the Nevada Viceroy. It is part of a well-known mimicry complex with the Monarch butterfly (Danaus plexippus), though recent research suggests the Viceroy itself may also be chemically protected rather than a pure Batesian mimic. This subspecies is found in western North America, particularly in the Great Basin region.
butterfly Nymphalidae mimicry Müllerian-mimicry western-North-America Great-Basin riparian willow-feeding
Limenitis archippus obsoleta
Southwest Viceroy
Limenitis archippus obsoleta, commonly known as the Southwest Viceroy, is a subspecies of viceroy butterfly native to western North America. Unlike the nominate eastern subspecies, this form exhibits darker, more brownish-purple coloration and is thought to mimic the Queen butterfly (Danaus gilippus) rather than the Monarch. The subspecies demonstrates the geographic variation in mimicry systems, where different models drive selection in different regions.
mimicry Müllerian-mimicry territorial-behavior geographic-variation subspecies-differentiation
Limenitis archippus watsoni
Watson's Gulf Coast Viceroy
Limenitis archippus watsoni is a subspecies of the viceroy butterfly found in central North America. It is part of the well-known mimicry complex with the monarch butterfly, though recent research suggests viceroys may also be unpalatable to predators. This subspecies is distinguished from other viceroy populations by geographic range and subtle morphological differences.
butterfly mimicry Nymphalidae North-America Limenitis viceroy subspecies Müllerian-mimicry Great-Plains Salicaceae willow-feeder
Lycidae
net-winged beetles
Lycidae is a cosmopolitan family of beetles commonly known as net-winged beetles. Adults are typically elongated, 10–15 mm in length, with brick-red or black-and-orange coloration and long, serrated antennae. The family is notable for its distinctive reticulated wing venation, chemical defenses against predators, and participation in extensive Müllerian mimicry complexes with other beetles and even moths. Several lineages exhibit neoteny, with females retaining larval morphology into adulthood.
Coleoptera net-winged-beetles aposematic-coloration Müllerian-mimicry neoteny larviform-females mycophagy nectarivory reticulate-venation mimicry-complex
Lycorea halia
Tropical Milkweed Butterfly, Tiger Mimic-Queen
Lycorea halia is a nymphalid butterfly in the Danainae subfamily, found from Peru through Central America to the Caribbean and Mexico, with occasional strays reaching Texas. It inhabits rainforest environments and has a wingspan of 95–108 mm. The species exhibits aposematic coloration typical of milkweed butterflies and has been documented feeding on bird droppings as adults.
Danainae aposematic Neotropical rainforest Müllerian-mimicry
Lycus
Netwing Beetles
Lycus is a genus of net-winged beetles in the family Lycidae, characterized by soft, flexible elytra with a distinctive reticulate (net-like) venation pattern. These beetles are notable for their aposematic coloration, typically featuring bold black and orange or red patterns that advertise their chemical defenses to predators. The genus is widely distributed and includes species that frequent flowers, where adults feed on pollen and nectar.
net-winged-beetle Lycidae aposematic-coloration flower-visitor Müllerian-mimicry reticulate-elytra defensive-chemistry
Lygaeus
seed bugs, milkweed bugs
Lygaeus is a genus of seed bugs in the family Lygaeidae, containing over 60 described species. Members are characterized by aposematic coloration—typically combinations of red, black, gray, and white—that advertises chemical defenses. Several species, notably L. kalmii (small milkweed bug), sequester cardiac glycosides from host plants, rendering them unpalatable to predators. The genus exhibits diverse feeding strategies ranging from seed-feeding specialization to opportunistic scavenging.
seed-bug milkweed-bug aposematic-coloration chemical-defense cardiac-glycoside Müllerian-mimicry pest sunflower-pest
Lygaeus kalmii
Small Milkweed Bug, Common Milkweed Bug
Lygaeus kalmii is a seed bug in the family Lygaeidae known for its bright orange-red and black aposematic coloration. Adults measure 10–12 mm and feed primarily on milkweed seeds and flower nectar, though they exhibit dietary flexibility including scavenging on dead insects and feeding on seeds of other plants such as composites. The species sequesters cardiac glycosides from milkweed, making it unpalatable to predators. Unlike the migratory large milkweed bug (Oncopeltus fasciatus), L. kalmii is non-migratory and overwinters as adults. Two subspecies are recognized: L. k. kalmii in western North America and L. k. angustomarginatus in the east, distinguished by differences in the white markings on the membranous portion of the forewings.
Hemiptera Lygaeidae seed-bug milkweed aposematic-coloration Müllerian-mimicry chemical-defense cardiac-glycoside non-migratory
Melinaea
clearwing butterflies
Melinaea is a genus of clearwing (ithomiine) butterflies in the family Nymphalidae, comprising 14 recognized species. It is the most species-rich and widespread genus within the subtribe Melinaeina, distributed from Mexico to Northern Argentina. Adults are aposematic and serve as main models in Neotropical Müllerian mimicry rings. The genus has been studied for its role in understanding speciation through divergent selection on wing colour patterns and the evolution of assortative mating.
clearwing aposematic Müllerian-mimicry speciation conservation-concern
Melinaea lilis
Mimic Tigerwing
Melinaea lilis is a butterfly in the family Nymphalidae, commonly known as the Mimic Tigerwing. It belongs to the tribe Ithomiini, a group of neotropical butterflies known for their unpalatability to predators and participation in Müllerian mimicry rings. The species was originally described as Mechanitis lilis by Doubleday in 1847. It is one of approximately 12 species in the genus Melinaea, which are distributed across Central and South America.
butterfly Nymphalidae Ithomiini Müllerian-mimicry neotropical Solanaceae chemical-defense
Meloidae
Blister Beetles
Meloidae, commonly known as blister beetles, is a family of approximately 7,500 species worldwide within the order Coleoptera. Members are characterized by their production of cantharidin, a toxic terpenoid compound that serves as a potent chemical defense against predators. The family exhibits remarkable diversity in adult morphology, with some species displaying aposematic coloration while others are cryptically colored. Life histories are complex, typically involving hypermetamorphosis with mobile triungulin larvae that often parasitize grasshopper eggs or bee nests. Adults are primarily herbivorous, with many species feeding on flowers and foliage of various plants.
hypermetamorphosis triungulin cantharidin aposematism Müllerian-mimicry livestock-poisoning alfalfa-pest flower-feeding grasshopper-egg-parasitoid bee-nest-parasite chemical-defense traditional-medicine anti-cancer-compound spermatophore-transfer size-assortative-mating