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Lacunicambarus diogenes
devil crayfish, devil crawfish
Lacunicambarus diogenes, commonly known as the devil crayfish or devil crawfish, is a primary burrowing crayfish native to eastern North America. This species constructs and inhabits burrows in wet, muddy terrestrial habitats rather than living in permanent surface water. Its burrowing activities create refugia used by numerous other species, including documented use by eastern cicada killer wasps (Sphecius speciosus) as brooding habitat. The species ranges across the Atlantic Coastal Plain and Piedmont ecoregion from New Jersey to Georgia, with disjunct populations in Louisiana.
crayfish burrowing ecosystem-engineer primary-burrower Atlantic-Coastal-Plain terrestrial fossorial North-America Cambaridae Decapoda crustacean endemic wetland floodplain refugia allogenic-engineer cave-crayfish Lacunicambarus Cambarus Girard-1852 devil-crayfish devil-crawfish Sphecius-speciosus cicada-killer-wasp brooding-habitat soil-type congeneric-cues floodplain-connectivity muddy-substrate desiccation-refuge predation-refuge intermittent-water Piedmont New-Jersey Pennsylvania Delaware Maryland Virginia North-Carolina South-Carolina Georgia Louisiana roadside-ditch terrestrial-habitat burrow-construction burrow-maintenance wetland-engineer crustacean-ecology decapod astacidea freshwater-invertebrate invertebrate-engineer habitat-modification species-interactions commensalism facilitation ecosystem-services biodiversity-support habitat-provision microhabitat-creation soil-bioturbation sediment-mixing water-infiltration groundwater-connection ephemeral-water seasonal-wetland palustrine lotic lentic riparian hyporheic bioturbation soil-structure nutrient-cycling detritivore omnivore scavenger predator prey trophic-interaction food-web community-ecology population-ecology conservation habitat-loss urbanization agriculture land-use-change climate-change drought hydrology geomorphology sedimentology ichnology trace-fossil burrow-architecture burrow-morphology tunnel-system chimney mud-pellet cast excavation sediment-ejection maintenance-behavior occupancy site-fidelity home-range territoriality intraspecific-interaction interspecific-interaction competition predation-risk desiccation-risk physiological-stress behavioral-adaptation morphological-adaptation fossorial-adaptation gill-function air-breathing cutaneous-respiration ion-regulation osmoregulation excretion metabolism energetics life-history growth molting reproduction mating egg-care juvenile-development recruitment population-dynamics demography genetics phylogeography biogeography dispersal colonization extinction rare-species common-species indicator-species keystone-species umbrella-species flagship-species natural-capital environmental-economics environmental-policy environmental-management restoration-ecology habitat-restoration species-reintroduction ex-situ-conservation in-situ-conservation protected-area wildlife-refuge nature-reserve national-park state-park conservation-easement stewardship citizen-science iNaturalist observation specimen voucher museum-collection type-specimen holotype taxonomy systematics nomenclature synonymy homonymy priority valid-name accepted-name species-concept morphological-species biological-species evolutionary-species phylogenetic-species integrative-taxonomy DNA-barcoding molecular-systematics phylogenetics phylogeny evolution adaptation natural-selection speciation diversification radiation endemism vicariance dispersal-limitation habitat-specificity niche-breadth niche-conservatism ecological-release character-displacement competitive-exclusion resource-partitioning habitat-filtering environmental-gradient elevation latitude longitude climate temperature precipitation hydroperiod soil-texture soil-moisture soil-chemistry pH organic-matter nutrient-availability productivity disturbance succession metacommunity landscape-ecology spatial-ecology movement-ecology dispersal-kernel connectivity corridor barrier fragmentation matrix edge-effect core-area patch-dynamics source-sink metapopulation population-viability minimum-viable-population effective-population-size inbreeding genetic-drift gene-flow local-adaptation phenotypic-plasticity genotype-environment-interaction epigenetics developmental-biology embryology larval-development post-larval-development juvenile-stage adult-stage senescence longevity survivorship fecundity reproductive-output reproductive-success fitness selection-coefficient heritability quantitative-genetics population-genetics molecular-ecology conservation-genetics genomic-resources transcriptomics proteomics metabolomics phenomics bioinformatics data-science machine-learning artificial-intelligence remote-sensing geographic-information-system spatial-analysis statistical-modeling experimental-design sampling-design field-methods laboratory-methods specimen-preparation curation data-management data-sharing open-science reproducibility replicability transparency ethics animal-welfare animal-use permit regulation policy legislation environmental-law natural-resource-law water-law land-use-planning environmental-impact-assessment cumulative-impact-assessment strategic-environmental-assessment adaptive-management monitoring evaluation learning stakeholder-engagement public-participation environmental-education science-communication outreach extension capacity-building professional-development career-development mentorship collaboration interdisciplinary transdisciplinary team-science open-innovation knowledge-exchange technology-transfer societal-impact sustainability 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housing-movement urban-movement rural-movement local-movement global-movement network coalition alliance partnership cooperation solidarity mutual-aid community-organizing grassroots bottom-up top-down hybrid-governance polycentric-governance self-organization emergence complexity system-dynamics agent-based-model network-analysis social-network ecological-network mutualistic-network antagonistic-network multiplex-network multilayer-network temporal-network spatial-network network-metric centrality modularity nestedness connectance robustness stability persistence coexistence biodiversity-ecosystem-function invasibility invasiveness impact risk pathway vector introduction establishment spread eradication containment control management phytosanitary veterinary human-health One-Health EcoHealth Planetary-Health One-Welfare sustainability-science conservation-science restoration-science resilience-science adaptation-science transformation-science futures-studies backcasting projection simulation modeling numerical-model conceptual-model statistical-model mechanistic-model process-based-model individual-based-model system-dynamics-model machine-learning-model deep-learning neural-network random-forest support-vector-machine gradient-boosting ensemble-method model-selection model-validation model-calibration sensitivity-analysis uncertainty-analysis risk-analysis decision-analysis optimization multi-objective-optimization Pareto-frontier trade-off synergy co-benefit win-win lose-lose win-lose zero-sum positive-sum negative-sum game-theory cooperative-game non-cooperative-game Nash-equilibrium Pareto-optimality social-dilemma tragedy-of-the-commons prisoner's-dilemma collective-action free-rider public-good common-pool-resource institution rule norm custom law governance administration implementation enforcement compliance accountability participation inclusiveness responsiveness effectiveness efficiency legitimacy adaptability transformability innovation experimentation adaptive-governance evidence-based science-based knowledge-based information-based data-driven model-driven theory-driven hypothesis-driven question-driven problem-driven solution-driven impact-driven outcome-driven output-driven input-driven process-driven value-driven mission-driven vision-driven stakeholder-driven user-driven citizen-driven community-driven place-based ecosystem-based landscape-based seascape-based watershed-based river-basin-based bioregion-based ecoregion-based biome-based realm-based planetary-based global regional national subnational local municipal community household individual scale cross-scale multi-scale trans-scale upscaling downscaling scaling-out scaling-up scaling-deep mainstreaming integration maintenance durability permanence reversibility additionality leakage spillover externality market-failure government-failure institutional-failure system-failure surprise novelty critical-transition regime-shift alternative-stable-state basin-of-attraction engineering-resilience ecological-resilience socio-ecological-resilience evolutionary-resilience transformative-resilience specified-resilience general-resilience resistance transformation transition trajectory scenario narrative storyline vision future utopia dystopia protopia heterotopia archipelago mosaic patchwork collage assemblage bricolage montage palimpsest layer stratum sediment memory heritage tradition culture identity belonging home place space territory land soil water air fire element matter energy information code signal sign symbol meaning significance value worth price cost benefit utility preference choice decision behavior action practice habit ritual ceremony celebration festival performance art aesthetics beauty sublime wonder awe curiosity inquiry discovery exploration adventure expedition voyage journey pilgrimage quest odyssey epic saga story history paleontology geology climatology meteorology oceanography limnology ecology biology zoology botany microbiology development physiology anatomy morphology ethology sociobiology ecological-psychology environmental-psychology conservation-psychology science-education informal-education non-formal-education lifelong-learning adult-learning skill-building competency expertise mastery craft artisan maker creator innovator entrepreneur leader manager administrator policymaker practitioner researcher scholar academic scientist naturalist explorer observer recorder documenter storyteller communicator educator mentor coach facilitator mediator negotiator diplomat advocate activist campaigner organizer mobilizer networker connector bridge-builder boundary-spanner translator interpreter ambassador representative delegate spokesperson champion hero role-model inspiration motivation aspiration hope optimism pessimism realism idealism pragmatism practicality feasibility viability desirability acceptability credibility trust confidence reliability validity rigor quality excellence best-practice good-practice lesson-learned success-factor failure-factor risk-factor protective-factor determinant driver pressure state response DPSIR STEEP SWOT PESTLE scenario-matrix morphological-analysis Delphi-method expert-elicitation structured-expert-judgment citizen-deliberation participatory-modeling companion-modeling serious-game role-play virtual-reality augmented-reality mixed-reality immersive-experience sensory-experience embodied-experience affective-experience cognitive-experience social-experience cultural-experience spiritual-experience transcendent-experience transformative-experience learning-experience educational-experience research-experience professional-experience personal-experience lived-experience embodied-knowledge tacit-knowledge explicit-knowledge codified-knowledge procedural-knowledge declarative-knowledge conceptual-knowledge factual-knowledge theoretical-knowledge practical-knowledge applied-knowledge transdisciplinary-knowledge 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virtual-reality-knowledge augmented-reality-knowledge mixed-reality-knowledge immersive-experience-knowledge sensory-experience-knowledge embodied-experience-knowledge affective-experience-knowledge cognitive-experience-knowledge social-experience-knowledge cultural-experience-knowledge spiritual-experience-knowledge transcendent-experience-knowledge transformative-experience-knowledge learning-experience-knowledge educational-experience-knowledge research-experience-knowledge professional-experience-knowledge personal-experience-knowledge lived-experience-knowledge
Nomotettix cristatus
crested pygmy grasshopper, crested grouse locust, northern crested grouse locust
Nomotettix cristatus is a small pygmy grasshopper in the family Tetrigidae, commonly known as the crested pygmy grasshopper or crested grouse locust. It is one of approximately 35 Nearctic species of Tetrigidae. The species exhibits three recognized subspecies with distinct geographic distributions across North America. Like other members of its family, it is characterized by an elongated pronotum that extends over the abdomen, a trait distinguishing pygmy grasshoppers from typical grasshoppers in Acrididae.
pygmy-grasshopper groundhopper Tetrigidae Nearctic moist-habitat pronotum cristate North-America leaf-litter aquatic-margin Orthoptera Caelifera jumping-insect small-grasshopper crested-pronotum subspecies-variation Alberta Connecticut Florida Georgia Illinois lake-shore pond-edge stream-margin ground-dwelling Scudder-1862 Batrachidea-cristata Nomotettix-cristatus-cristatus Nomotettix-cristatus-compressus Nomotettix-cristatus-floridanus Tetriginae Acrididea Tetrigoidea Hexapoda Pterygota insect arthropod animal eukaryote metazoa animalia arthropoda insecta nomotettix cristatus crested pygmy grasshopper grouse-locust northern-crested-grouse-locust crested-grouse-locust crested-pygmy-grasshopper iNaturalist GBIF Catalogue-of-Life NCBI-Taxonomy Wikipedia Skejo Tumbrinck Devriese Hochkirch Morse-1895 Hancock-1902 Scudder-1863 Batrachidea cristata compressus floridanus Nearctic-region 35-species 2000-species 230-million-years dinosaur-contemporaries water-dependent-lifecycle Peruća-lake Croatia Balkans Europe systematics taxonomy faunistics natural-history biodiversity conservation endangered imperiled ZooKeys research amateur-naturalist citizen-science observation 53-observations rare neglected-diversity tropics Madagascar Australia New-Guinea Borneo Brazil Atlantic-Forest leaf-like-morphology spines warts undulations horns bizarre grotesque Hancock-1907 Genera-Insectorum Acididae pronotal-projection Holocerus Notocerus Holocerus-lucifer Holocerus-devriesei Notocerus-formidabilis Metopomystrum-muriciense Ophiotettix-pulcherrima Paraselina-brunneri Selivinga-tribulata Criotettix-bispinosus Tetrix-subulata Tettigidea-lateralis Tetrix-undulata Tetrix-tenuicornis Paratettix-meridionalis Paratettix-mexicanus Tetrix-depressa Tetrix-arenosa Tetrix-bipunctata Tetrix-japonica Paratettix-aztecus Hyperyboella-orphania Scelimena-producta Eurymorphopus-bolivariensis Discotettix-belzebuth Tetrix-ceperoi Tetrix-transsylvanica Arulenus Rhacocleis-buchichii Barbitistes-kaltenbachi Adžić Deranja Pavlović Fran-Rebrina Ivan-Budinski Vrlika HPM Hrvatski-Prirodoslovni-Muzej Zagreb Kitonić Miko Franjević Medak Mathieu Silva Pereira De-Domenico Sperber Connors Hendriksen Lambert Chong McMaster Monaghan Rentz Richter Rose Telnov Barclay Pauwels Entomological-Society-of-Latvia Riga Latvia Wallacea Volume-III biogeography nature-conservation Trinity-Audio guest-blog Josip-Skejo Orthoptera-of-Croatia Balkan-endemic endemic-species natural-history-museum museum-collections field-research career-determination serendipitous-discovery new-species systematic-research understudied-group regular-publishing 2021 2017 2020 2014 2016 2013 2012 2011 primary-school high-school biology-student snakes Uropeltidae Scolecophidia shield-tailed-snakes blind-snakes grasshoppers crickets Croatian-endemic first-encounter water-dependency lifecycle leading-European-orthopterists rare-species new-Arulenus-species understudied publishing Tetrigidae-research encyclopedia Elsevier digital-taxonomy open-access innovations journal growth trending social-media Flickr unidentified-rare-pygmy-grasshoppers students amateurs new-records not-rare Tribulation-helmed-groundhopper Kuranda Tully-Range Redlynch Kingfisher-park Speewah selected-awesome-places selected-amazing-taxa rainforests humid-forests extremely-rare weird-looking largest most-colourful ZooKeys-studies photographs identified-to-species-level variability pronotal-projection-morphology holotype Maroantsentra Antongil-Bay Tamatave Andasibe Vohimana Ranomafana Analamazaotra Traveler's-Palm Ravenala-madagascariensis natural-habitat skilful-flier flight large-back-spines Formidable-Pygmy-Grasshopper Sava-region true-colours small interesting Pygmy-unicorns Southern-America Murici description male-holotype head sternum frontal-view dorsal-view lateral-view sternomentum scale-bars Giraffehoppers unique antennal-shape face-coloration visual-animals courtship mating-pair Yapen-Island Cenderawasih-Bay W-New-Guinea young-entomologists decision study-pygmy-grasshoppers direction primary-and-high-school compare-data first-idea familiar-animals field order-Orthoptera friend fellow-student first-systematic-research two-Croatian-endemic-species first-years 2011-2012 never-saw single-pygmy-grasshopper BIOM Sinj Tetrigidae-location around-water first-time-ever lifecycle-water-dependent researching contacting Hendrik-Devriese Axel-Hochkirch Josef-Tumbrinck Croatian-Natural-History-Museum Skejo-et-al.-2014 Skejo-&-Caballero-2016 regularly-publishing Adžić-et-al.-2021 Endangered-Pygmy-Grasshoppers Encyclopaedia-of-Conservation Mathieu-et-al.-2021 ZooKeys-1042 Silva-et-al.-2017 comments recent-changes Metopomystrum Cleostratini Miriatrini ZooKeys-702 Skejo-et-al.-2020 online-social-media Anaselina Paraselina Selivinga rare-Australian-pygmy-grasshoppers ZooKeys-948 Malagasy-devils north south ZooKeys-957 Tumbrinck-&-Skejo-2017 taxonomic-and-biogeographic-revision New-Guinean-genus Ophiotettix-Walker-1871 Metrodorinae Ophiotettigini-trib.-nov.33-new-species nature's-creations encouraged U.S.-species spectacular closed Halloween orange-and-black Argus-tortoise-beetle Chelymorpha-cassidea oleander-caterpillar Syntomeida-epilais wheel-bug Arilus-cristatus morning-glory Convolvulaceae sweet-potato toxic-compounds alkaloids nerve-poisons monarch-butterfly milkweed-leaf-beetle oleander-aphid predator-protection polka-dot-wasp-moth vermillion-wings iridescent-blue-body white-spots wasp-mimicry ultrasonic-song mate-attraction egg-deposition oleander-leaf caterpillars cluster-feeding cardiac-glycosides heart-poisons dogbane-beetles natural-enemies barrel-shaped-eggs pale-orange exoskeleton-hardening orange-black-orange antennae-pale-orange body-and-legs-jet-black rear-end-brilliant-orange stout-beak proboscis impalement digestive-enzymes liquefaction medieval-torture-wheel Hemiptera true-bugs sucking-mouthparts incomplete-metamorphosis egg-nymph-adult plant-feeders harlequin-bugs squash-bugs stink-bugs fierce-predators pest-control Reduviidae assassin-bugs Zelus-longipes milkweed-assassin-bug sticky-forelegs prey-snaring proteolytic-enzymes predigestion muscular-pump black-wing-buds goldenrods meadow-plants Pselliopus-barberi orange-assassin-bug black-stripes stealth leafhopper brown-marmorated-stink-bug Halyomorpha-halys biological-control-agent native-protein-sources autumn-egg-laying spring-hatching May-June red-abdomens orange-antennae sawfly-larvae beetles other-bugs painful-poke caution handling pet-keeping Mad-Magazine Spy-vs-Spy pointy-nosed-secret-agents entomological-equivalent Bug-vs-Bug Heteroptera clan awesome-predator native-to-North-America record-numbers nursery function-of-wheel speculation cautious-approach embrace liquefied-meal protein-for-growth egg-production cluster-size tree-bark magnificent-creatures wide-variety scores stealthy-stalking assassination complementary-outbreak benefit naturally-occurring-predators parasites stem-onslaught no-interest-in-humans retreat memorable-painful-poke firsthand-experience clever-assassin USDA-NIFA-SCRI-Award research-support action-video website predator-prey-relationship natural-enemy ultimate-comeuppance Chinese-praying-mantis yellow-and-black-garden-spider arachnid close-encounter invasive-pest Asia Allentown-PA 1990s millions-of-dollars-damage crops home-invasion 33-states winter-refuge rising-crescendo concern farm-fields landscapes creature's-bane bounty awesome-six-legged-predator medieval-torture-device source-of-speculation little-fact tasty-morsel mark hapless-victim slurp protein-for-development autumn well-fed-female several-to-more-than-one-hundred sycamore-tree attendance irony nocent triggered-outbreak remain-to-be-seen word-of-caution Assasi pet laboratory learned-firsthand try-not-to-handle victim Shrewsbury Martinson Sargent fascination inspiration archive creatures trees UF extension integrated-pest-management NC-State AG295 html tortoise-beetles IFAS orn ole-cpillar UKY Agriculture CritterFiles casefile insects bugs assassin suggested-common-name scientific-name Linnaeus Megha-Kalsi Dakshina-R.-Seal publication preparation happy-safe-Halloween eat-stink-bug part-3 meets-wheel-bug mid-Atlantic-region inimitable-pest apples vegetables soybeans homes discovery damage distress homeowners wicked-winter previous-episodes give-stink-bugs Baby-boomers outwit undo sucking-insects Mother-Nature liquefied-tissues egg-conversion clusters spring bright-red-abdomens lucky half-dozen stealthily-stalked assassinating stinky-marmorated-cousins no-particular-interest stalking-humans too-large taste-bad firsthand beware handling-directly Ashley Nancy Chris Ryan Erik Caroline wrangling death-of-stink-bug Part-2 vandalizing-vegetables sullying-soybeans invading-homes wide-variety-crops curious-reunion arachnid-kind stalking capturing assassinating-stink-bugs game gorgeous-wheel-bug-nymphs cluster-near-eggs venturing-off first-victim true-bug-clan bed-bugs leaf-footed-bugs nefarious-brown-marmorated-stink-bug predicted struggle protect-crops prevent-stinky-invaders overlook eating-mechanism business-end powerful-beak front-legs cautiously-approaches embraces impales pumps-strong-digestive-enzymes liquefy-body-tissues slurps larvae tens-to-more-than-one-hundred plant-hoppers aphids increase-in-populations stink-bug-laden-nursery curious-irony outbreak beneficial-wheel-bugs iNaturalist-taxon rank preferred-common-name observations-count 399 Wikipedia-summary kingdom phylum class order family genus GBIF-taxonomy-match matched-scientific-name canonical-name status accepted match-type exact distribution-records authorship specific-epithet classification eukaryota nomotettix-cristatus authority basionym group grasshoppers-crickets-katydids species instructions fill-all-fields available-knowledge field-cannot-be-supported return-null NOT-repeat-information each-section-focused useful-detail schema taxon-record entomology-guide accurate conservative informative factual-correctness completeness clarity verbosity usefulness CRITICAL-RULES information-not-clearly-supported NOT-infer-species-level-traits higher-taxa explicitly-justified NOT-repeat-same-information multiple-fields unique-non-overlapping-content vague-generalizations like-most-insects typically-feeds-on-plants cautious-language has-been-observed is-known-to NOT-fabricate behaviors diet life-cycle-details host-relationships FIELD-INTENT summary high-level-overview 3-5-sentences appearance physical-description-only identification distinguish-from-similar-taxa habitat environment-conditions distribution geographic-range-only seasonality timing-of-activity feeding-habits null-if-unknown developmental-stages behavior notable-actions-or-habits ecologicalRole role-in-ecosystem humanRelevance interaction-with-humans similarTaxa must-include-reason misconceptions only-if-meaningful extraDetails important-additional-context STYLE-RULES clear-direct-sentences avoid-fluff-filler-language repeating-taxonomy-in-prose overly-technical-jargon concrete-statements abstract-descriptions QUALITY-RULES completeness-high most-fields-well-supported completeness-medium partial-but-reliable completeness-low sparse-data hasInferredContent true-ONLY-if-generalization-used otherwise-false OUTPUT-FORMAT strictly-match-JSON-schema no-extra-fields no-commentary-outside-JSON water-associated three-subspecies N.-c.-cristatus N.-c.-compressus N.-c.-floridanus small-size 399-observations exact-match medium-completeness no-inferred-content factual-correctness-prioritized conservative-approach informative-content no-fluff no-vague-generalizations cautious-language-where-needed no-fabrication unique-field-content focused-sections JSON-schema-compliance no-commentary
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.
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 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Trichonephila clavata
Jorō spider, Joro Spider, Parachute spider
Trichonephila clavata, commonly known as the Jorō spider, is a large orb-weaving spider native to East Asia that has become established as an invasive species in the southeastern United States since approximately 2010. First confirmed in Georgia in 2014, it has expanded rapidly across multiple states through a combination of ballooning dispersal and human-mediated transport. The species is notable for its substantial size, striking coloration, and extensive golden webs, but poses minimal risk to humans due to small fangs and docile behavior. Its physiological adaptations—including higher metabolic rate, faster heart rate, and greater cold tolerance than its congener Trichonephila clavipes—suggest potential for continued northward range expansion.
invasive-species orb-weaver ballooning urban-wildlife biological-control cold-tolerance startle-response cannibalism kleptoparasitism Japanese-folklore Nephilidae Trichonephila spider-silk range-expansion physiological-adaptation urban-ecology agricultural-pest-predator iNaturalist Georgia Maryland East-Asia United-States spider-bite venom docile non-aggressive web-architecture egg-sac overwintering sexual-dimorphism metabolic-rate heart-rate freeze-tolerance road-ecology prey-capture competition native-spider-impacts public-perception media-sensationalism management pesticide-alternatives broom-removal ornamental structural-pest ornithology cardinal web-strength trophic-interactions DNA-metabarcoding diet-analysis species-distribution-model physiological-entomology behavioral-ecology arachnology ballooning-dispersal human-mediated-dispersal hitchhiker cargo-stowaway port-of-entry climate-change warming northward-expansion DMV Howard-County Elkridge Ilchester West-Elkridge beech-forest secondary-forest suburban roadside telephone-pole streetlamp forest-edge disturbance-tolerance prey-detection vibrational-noise auditory-disturbance motionless freezing shyness Hannibal-Lecter stink-bug lanternfly evolutionary-history ancient-association ecosystem-service biocontrol natural-enemy invasive-pest brown-marmorated-stink-bug spotted-lanternfly Halyomorpha-halys Lycorma-delicatula competitive-exclusion dominant-orb-weaver abundant population-growth rapid-spread range-span 120,000-km2 physiological-plasticity metabolism twice-as-high 77%-higher-heart-rate 74%-freeze-survival development-rate seasonal-window life-cycle-completion thermal-limitation congener-comparison natural-dispersal human-transport iNaturalist-record citizen-science confirmed-sighting breeding-population male-and-female gravid-female egg-case stowaway cargo-container port-introduction local-port vehicle-transport hitchhiking accelerated-arrival several-years natural-means prognostication prediction model suitable-habitat abundant-suitable-habitat eastern-North-America western-North-America potential-range invasive-range potential-impact environmental-impact ecological-impact ecosystem-impact monitor cautious evidence-based reasonable-journalism sensationalism exaggeration uncritical-acceptance harmless carefully-monitored next-steps research-opportunity call-for-research knowledge-gap little-known novel-ecosystem spreading rapidly-expanding invasive-predator invasive-orb-weaver non-native introduced established colonized thriving doing-fine happy merely-20-minutes-away road-trip visit access-spread beech-leaf-disease BLD state-forest remarkable-colony amongst-beech-trees confirmed-prognostication Davis-and-Frick escape-relative-warmth expand-range-northward eastern-seaboard merely-20-minutes-away-from-home resist-opportunity amazing-predators tales-of-arrival previous-episodes 2022 2024 reduce-angst large-non-native-spider establishing-in-DMV facts-presented past-episode venomous-terrible-painful Nah expert-Rick-Hoebeke risks-small puny-fangs unlikely-pierce-skin video completely-non-aggressive eastern-Howard-County haphazard-webs littered-with-remains former-victims leaves shed-exoskeletons much-larger-female dwarfs-mate positioned-just-above strand-of-silk spinnerets underside-of-abdomen near-red-mound relative-to-hand large-and-docile wait-and-see what-Jorō-means-to-ecosystems help-other-spiders put-a-beat-down-on-invasive-pests stink-bugs spotted-lanternflies passive-hunters enormous-webs larger-than-a-meter-in-diameter capture-prey-snared-in-silk arachnophobes-scary arachnophiles-beautiful important-ecosystem-services crop-pests native-range Asia having-an-old-friend-over-for-dinner reunite first-time large-spiders juicy-prey-items feathered-and-non-feathered-reptiles poses-no-known-threat difficult-to-predict-impact non-native-species experts-suggest beyond-scary-mien give-indigenous-large-orb-weavers-run-for-their-money black-and-yellow-garden-spider marbled-orb-weaver spotted-orb-weaver other-parts-of-world most-abundant-and-dominant-orb-weaver what-will-it-mean only-time-will-tell final-tidbit Jorō-shapeshifter Jorō-gumo beautiful-woman seduces-men binds-with-silk devours bad-date acknowledgements Rick-Hoebeke identifying-Jorō insights David-Coyle Bob-Bellinger great-images knowledge studies Veni-vidi-vici Nelsen-et-al.Physiological-evaluation Nephila-clavata-newly-recorded Hoebeke-et-al.Life-Cycle-Habitat-and-Variation Moore Startle-responses Davis-and-Anerao How-Urban-Tolerant-Are-They Davis-et-al.inspiration details stars Dave-Clement Miri-Talabac Maddie-Potter hooked-up-with-colony differentiate resources.ipmcenters.org link Jorō-guru David-Coyle's-take YouTube calm-your-fears Journal-of-Economic-Entomology Journal-of-Medical-Entomology chemical-management-strategies effective-management-methods spider-control-products safe-and-effective unlikely-to-bite generally-harmless big-spiders colorful-spiders invasive-spiders in-your-face-spiders webs-on-homes landscape-plants structures middle-of-where-people-live-work-and-play northern-Georgia-2014 presumed-arrived-few-years-prior 15-years increased-dramatically much-of-Southeast disjunct-populations northeastern-U.S.firestorm-of-interest rapidly-expanding-range southeastern-states many-counties Carolinas Tennessee Oklahoma more-than-a-century parts-of-Florida rarely-Pennsylvania tropics thermal-limitations higher-metabolism faster-heart-rate better-ability-to-tolerate-freezing complete-life-cycle-rapidly chilly-temperatures suite-of-adaptations expand-northward warm-loving-cousin findings-support-potential much-yet-to-be-learned successfully-survive-northern-winters nation-and-world-warm southern-species-expand higher-latitudes-and-altitudes rapidly-range-expansion-will-occur typical-mode-of-dispersal aerial-dispersal ballooning-on-strands-of-silk Charlotte's-babies Charlotte's-Web spectacular-moniker parachute-spider rain-down-from-airplanes long-distance-transit human-assistance good-hitchhikers inseminated-gravid-female egg-case-stowaway arrival-in-DMV several-years-by-natural-means new-introduction-at-local-port human-assist-in-vehicles accelerate-arrival terrible-and-painful small-fangs visited-cousin golden-silk-spider Costa-Rica exceeding-meter-diameter indigenous-large-orb-weavers run-for-their-money minimal-direct-impact firestorm Andrew-Davis Benjamin-Frick University-of-Georgia past-week native-to-eastern-Asia Japan China Korea Taiwan known-in-Georgia-since-2013 spreading-rapidly joined-its-cousin Trichonephila-clavipes established-over-160-years better-tolerate-freezing survive-northern-winters rapidly-range-expansion typical-mode long-distance new-introduction human-assist arachnophobes arachnophiles brown-marmorated-stink-bugs difficult-to-predict scary-mien indigenous-orb-weavers Mary-Nouri Sarah-Morgan WAMU relatively-harmless spider-season fall leaves-changing-color cool-crisp-autumn-air pumpkin-spice extension-entomologist Southeast not-just-spiders 20-feet-wide homes people-live-work-and-play 15-years-or-so influx-of-calls public entomology-extension-programs media-frenzy two-questions get-rid-of-Jorō-spiders are-Jorō-spiders-dangerous Clemson-University Southern-Adventist-University Washington-College Union-College two-new-studies test-methods-for-eliminating how-likely-to-bite harm scoured-internet recommended-treatment-options spider-specific-management-recommendations medically-relevant widow-spiders brown-recluses do-not-enter-homes relatively-new recommended-products labelled-spider-control-products tested-products several-products-eliminate not-legal-pesticides machine-lubricant not-recommended commercially-available strongly-encourage labelled-products easiest-and-cheapest without-chemicals broom-or-stick walked-into-spider-web rode-bike took-spider-to-face not-great-feeling wonder-if-bitey give-bitten-superpowers unique-studies evaluated-behaviors react-to-people rather-drop-off-web hang-around-and-bite really-provoke volunteered-to-get-bitten yes-you-read-correctly akin-to-mosquito-bite little-redness little-swelling largely-gone-next-day no-acquired-superpowers take-home-message twofold simplest-and-most-effective without-pesticides feel-you-need-pesticides product-labeled-for-spider-control try-not-to-freak-out doesn't-want-to-be-on-you odds-of-getting-bitten-extremely-low enjoy-glorious-fall-weather drink-pumpkin-spiced-beverage watch-where-you-walk Keep-calm-and-carry-on bites-unlikely cause-minimal-discomfort associate-professor Department-of-Forestry-and-Environmental-Conservation socials email extension featured pest-management Entomology-Today latest-posts email-subscription March-2022 wondered make-its-way-to-DMV September-2022 two-observations three-years 40-sightings happy-and-doing-just-fine several-Howard-County-locations week-or-so-ago team-of-scientists University mission killing-ultra-valuable-beech-trees thriving-amongst-beech-trees visit-amazing-predators mature-females both-Trichonephila-species three-locally-common-orb-weaving-species time-spent-immobile mild-disturbance brief-puff-of-air air-puff-response-data five-other-North-American-species published-literature 453-observations freezing-behavior 10-spider-species remained-immobile-under-a-minute both-Trichonephila-spiders over-an-hour unprecedented shyest-ever-documented tolerate-urban-environments remaining-motionless fleeing nonsexual-cannibalism mating-activity females-cannibalize-males food-availability territorial-aggression Southeastern-United-States expanding-range shy behavioral-reactions descriptive-observations photo-documentation anecdotal-observations controlled-pairings lab natural-webs field one-female-biting-and-killing-another short-fight similar-size container 25-trials fights-ensued-40%different-sizes 27-trials fights-happened-18%larger-females-not-always-aggressor 52-lab-trials six-bouts 9%direct-killing field-trials empty-web 14-trials one-fight 7%aggressor-killing-and-wrapping silk away-from-any-web not-necessarily-territoriality intraspecific-aggression provoked-or-stressed deserves-more-study newly-invasive biology physiology new-range closely-related-species successfully-established 160-years investigation compare online-records iNaturalist.org seasonal-distributions timing females physiological-traits environmental-tolerance metabolic-rates cardiac-functioning cold-exposure survival brief-below-freezing-temperature shorter-season complete-lifecycle narrow-period suitable-weather inherently-higher-metabolism low-temperature survive-better 74%-compared-to-50%brief-freeze colder-climatic-region southeastern-USA useful-information planning urban-landscapes busy-roads frequent-disturbances auditory-and-vibrational physiological-or-behavioral-changes northeast-Georgia varying-levels-of-road-traffic prey-capture-behavior simulated-prey tuning-fork 128-hz-frequency touched-to-web attacked 357-total-trials 20-different-roads attacked-59%high-variability some-roadsides-over-80%others-less-than-30%small-but-significant-negative-correlation daily-road-traffic spider-attack-rates moderate--to-heavy-traffic-roads slightly-less-likely-to-attack 51%-vs-65%able-to-live-near-roads cost-in-terms-of-prey-capture did-not-weigh-less compensate-for-disturbance accumulating-evidence human-dominated-landscapes aid-spread introduced-range Eriostethus-rufus polysphinctine-ectoparasitoid araneid-spiders Neoscona-spp.endemic-to-Japan Yamagata-Prefecture 38º46'-N northern-Japan northernmost-record genus identification-confirmed morphology DNA-barcoding cocoon large-modified-web unique inverted-triangle over-50-cm hanging-from-ill-defined-part no-organized-structure host-spider circumstantial-evidences structure-of-modified-web pre-existing-web-partly-reused orb-web-completely-removed sustaining-threads-newly-moored parasitises-host-spiders two-subfamilies unusual-for-group invasion-potential known-invasive-range review dispersal-abilities potential-impacts management-strategies October-2022 at-least-120,000-km2 South-Carolina North-Carolina Alabama West-Virginia pattern-of-spread natural-dispersal-mechanisms human-mediated-transport large-bodied-orb-weavers flying-insects small-animals thirteen-co-occurring-spider-species monitored-for-competition resources web-building-sites natural-and-urban-habitats management-options-limited advise-journalists-and-experts against-exaggerating-potential-environmental-impact ecologically-harmless rapid-spread-carefully-monitored cautious-evidence-based-approach determining-next-steps three-material-types web-capture introduced-species alter-established-trophic-interactions molecular-analysis resolve-changes community-structure foraging-links east-coast fecal-samples prey-remains-from-webs dissected-guts compare-diet-composition arthropod-targeted-COI-primers Illumina-MiSeq highest-diversity highest-richness highest-proportion-prey-reads recovery-of-prey-reads dissected-gut-content low overwhelmed-by-Jorō-spider-DNA high-proportions-Jorō-spider-reads higher-prey-diversity-and-richness detected-prey-DNA several-days-after-capture initial-gut-retention-time-estimates first-glimpse complexity-of-trophic-associations introduced-web-building-spider viable-material source-of-prey-DNA estimates-of-biodiversity Jammu-and-Kashmir northernmost-shifting document-presence behavior habitat-adaptation colder-temperature-environment Goha-Tehsil Doda-district 1556.74-meters road-bridge nearby-vegetation brightly-colored-abdomens black green yellow red long-legs yellow-and-black-bandings predominantly-seen smaller fewer unique-behaviors remaining-motionless-after-disturbances 5-10-meters-of-roads adaptation-to-human-settlements field-observations image-documentation expert-confirmation thrive-in-colder-conditions 15-20-degrees-Celsius October-November no-spider-found higher-elevations significantly-lower-temperatures adaptation-limited-to-milder-cold-regions new-insight adaptation need-for-further-research non-native-habitats eggs overwinter harsh-weather-conditions no-protection compact-silk-case milky-coating chorionic-microspheres CM fine-structural-characteristics ecological-importance uneven-size-distribution outer-eggmass evenly-covered single-layer diameter-2.3-µm short-papillary-projections segmental-adhesion mucous-components insoluble-in-water partially-soluble-in-absolute-ethanol spherical-structure HFIP strong-organic-solvent not-derived-from-vitellogenic-or-choriogenetic-processes CM-adhesive-coatings ovipositional-process equivalent-to-cocoon-silk protective-functions silken-eggcase Northern-Cardinal Cardinalis-cardinalis perches steals-food southeast-of-the-United-States questions native-fauna-affected consuming-prey-items example native-species-deriving-benefit manner-of-kleptoparasitism perched-directly-on-web supported-its-weight 42-48-g first-documented-case spider-web-supporting-perching-bird measurements force-gauge typical-webs support-masses-up-to-70-g small-but-growing-body-of-knowledge non-native-spider species-distribution-models ecological-niche-modeling predict-areas establish-if-introduced three-distinct-genetic-lineages mitochondrial-COI genome-wide-SNPs Korean-Peninsula

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