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Tortopus
Tortopus is a genus of burrowing mayflies in the family Polymitarcyidae, restricted to six species based on cladistic revision: T. igaranus, T. circumfluus, T. harrisi, T. zottai, T. bellus, and T. arenales. The genus is characterized by distinctive morphological synapomorphies including female parastyli receptors with long anterior furrows, entirely flattened penes, and nymphs bearing two subapical tubercles on mandibular tusks. Nymphs construct U-shaped tunnels in clay banks, a behavior that renders them difficult to sample with standard aquatic survey methods.
Ephemeroptera Polymitarcyidae burrowing-mayflies Neotropical clay-bank-habitat U-shaped-tunnels mandibular-tusks Tortopsis Campsurinae Panamerican-distribution cladistic-revision synapomorphy parastyli-receptors flattened-penes furcasternal-protuberances subapical-tubercles limnological-sampling-bias nymph-morphology adult-morphology type-species-T.-igaranus new-species-T.-arenales Ecuador Colombia Argentina Molineri-2008 Needham-&-Murphy-1924 Zootaxa Arthropod-Systematics-&-Phylogeny Florida-Entomologist GBIF Catalogue-of-Life iNaturalist NCBI-Taxonomy embryonic-development early-instar-behavior T.-harrisi T.-bellus T.-circumfluus T.-zottai T.-igaranus T.-incertus T.-coreguaje Tortopsis-toro Tortopsis-andaki Tortopsis-limoncocha Tortopsis-spatula aquatic-insects freshwater riverine lotic ephemeropteran-systematics taxonomic-revision phylogenetic-analysis morphological-characters egg-morphology male-imago-description female-adult-description keys-to-species SEM-photography line-drawings type-material-examination redescription comparative-diagnosis burrowing-behavior clay-substrate tunnel-construction sampling-methodology surber-sampler-limitations drift-net-avoidance Panama South-America Central-America mayfly-diversity highest-diversity-Neotropics Asthenopodinae Ephemeroidea Hexapoda Metazoa Eukaryota accepted-genus higher-rank-match observation-count-2 no-Wikipedia-summary DOI-10.11646/zootaxa.2481.1.1 DOI-10.11646/zootaxa.2436.1.4 DOI-10.3897/asp.79.e62735 DOI-10.2307/3493493 Scott-et-al.-1959 Navás Ulmer Traver Lugo-Ortiz-&-McCafferty Domínguez Banks McDunnough Pictet Needham-&-Murphy freshwater-ecology aquatic-entomology insect-development nymphal-stages imago-description alate-adults wing-venation R-sector gonopore-structure parastyli-length pedestal-morphology lateral-view-curvature mandibular-morphology tusk-tubercles mesosternal-structure abdominal-sternum-notch furcasternum protuberance-configuration receptor-socket-orientation IR-vein R2-vein claw-like-structure gonopore-association penis-base-separation female-fore-wing additional-veins comparative-morphology type-specimen holotype allotype paratype neotropical-species new-species-description species-key identification-key male-adult-key female-adult-key nymph-key generic-key Polymitarcyidae-genera systematics phylogeny cladistics monophyly sister-group character-state transformation-series parsimony-analysis morphological-data-set external-morphology egg-chorion micropyle egg-burster hatching emergence behavioral-study early-instar embryology oviposition substrate-preference habitat-specificity endemic regional-diversity conservation biomonitoring water-quality-indicator macroinvertebrate benthic infaunal interstitial hyporheic river-bank stream-bank erosion sediment particle-size clay-content bank-stability root-mat vegetation-association flow-regime current-velocity depth width stream-order catchment basin ecoregion biogeography dispersal colonization vicariance speciation adaptive-radiation morphological-divergence convergent-evolution homoplasy character-conflict bootstrap-support Bremer-support consensus-tree strict-consensus majority-rule taxonomic-stability nomenclatural-act original-description subsequent-designation type-locality type-depository museum-collection specimen-preservation alcohol-preservation slide-mounting SEM-preparation imaging-techniques scientific-illustration photography measurement morphometrics ratios proportions allometry sexual-dimorphism geographic-variation intraspecific-variation interspecific-variation diagnostic-characters identification-confidence taxonomic-expertise literature-review bibliographic-search database-query online-repository open-access paywall abstract-only full-text-unavailable citation-analysis reference-list source-quality evidence-strength knowledge-gap research-need future-study collaboration field-work collection-trip sampling-effort specimen-availability type-examination revision-scope taxonomic-concept circumscription inclusion exclusion species-group species-complex cryptic-species morphospecies phylogenetic-species evolutionary-significant-unit operational-taxonomic-unit DNA-barcoding molecular-systematics integrative-taxonomy traditional-taxonomy alpha-taxonomy beta-taxonomy gamma-taxonomy classification-hierarchy Linnaean-system rank-based-nomenclature phylogenetic-nomenclature clade-name crown-group stem-group total-evidence congruence incongruence data-partition character-weighting implied-weighting equal-weighting ordered-characters unordered-characters multistate-characters binary-characters continuous-characters meristic-characters qualitative-characters quantitative-characters morphological-matrix data-matrix character-matrix taxon-sampling character-sampling missing-data inapplicable-data polymorphism intraspecific-polymorphism fixed-difference diagnostic-difference unique-derived-character autapomorphy shared-derived-character symplesiomorphy convergent-character parallel-evolution reversal character-optimization ACCTRAN DELTRAN unambiguous-optimization ambiguous-optimization most-parsimonious-tree tree-length consistency-index retention-index rescaled-consistency-index homoplasy-index character-fit character-congruence character-support branch-support node-support clade-stability taxonomic-stability-index phylogenetic-signal noise saturation substitution-model rate-heterogeneity among-site-rate-variation gamma-distribution invariant-sites model-selection Akaike-information-criterion Bayesian-information-criterion likelihood-ratio-test Bayesian-inference Markov-chain-Monte-Carlo posterior-probability prior-probability hyperprior convergence effective-sample-size burn-in stationarity trace-plot parameter-estimate credible-interval highest-posterior-density tree-topology tree-shape lineage-through-time diversification-rate extinction-rate speciation-rate net-diversification birth-death-model Yule-model coalescent-model population-size effective-population-size gene-flow migration-rate isolation-by-distance phylogeography historical-biogeography ancestral-area-reconstruction dispersal-vicariance-analysis decisive-weighting event-based-method taxon-area-cladogram area-cladogram biogeographic-homology track-analysis panbiogeography phylogenetic-biogeography ecological-biogeography dispersal-biogeography vicariance-biogeography center-of-origin area-of-endemism refugium Pleistocene glaciation climate-change habitat-fragmentation habitat-loss anthropogenic-impact pollution eutrophication sedimentation dam-construction water-abstraction flow-modification thermal-pollution toxic-substances heavy-metals pesticides urbanization agriculture deforestation land-use-change riparian-degradation channelization dredging mining invasive-species biological-invasion competition predation parasitism mutualism commensalism amensalism facilitation ecosystem-engineer keystone-species flagship-species umbrella-species indicator-species sentinel-species bioindicator ecological-indicator environmental-indicator water-quality-index biotic-index multimetric-index rapid-bioassessment protocol standardization quality-assurance quality-control reference-condition least-disturbed minimally-disturbed best-available type-specific region-specific stream-type river-type functional-feeding-group collector-gatherer collector-filterer scraper shredder predator parasite engulfer piercer burrower sprawler clinger swimmer skater diver surface-dweller benthic-dweller interstitial-dweller hyporheic-dweller phreatic-dweller troglobite stygobite cave-inhabitant spring-inhabitant headwater mid-order high-order lowland montane alpine subtropical tropical temperate boreal arctic antarctic palearctic nearctic afrotropical oriental australian oceanic cosmopolitan widespread restricted relict disjunct-distribution range-extension new-record first-record country-record state-record province-record locality-record elevation-record altitudinal-range latitudinal-range longitudinal-range distribution-pattern range-size range-fill extent-of-occurrence area-of-occupancy IUCN-category threat-status data-deficient not-evaluated least-concern near-threatened vulnerable endangered critically-endangered extinct-in-wild extinct population-trend population-density population-structure age-structure size-structure sex-ratio reproductive-output fecundity fertility mortality survivorship life-table cohort-analysis generation-time doubling-time intrinsic-rate-of-increase finite-rate-of-increase net-reproductive-rate stable-age-distribution stationary-population expanding-population declining-population source-sink-dynamics metapopulation population-viability-analysis minimum-viable-population genetic-diversity heterozygosity allelic-richness private-allele founder-effect bottleneck genetic-drift inbreeding inbreeding-depression outbreeding outbreeding-depression hybridization introgression gene-flow-barrier reproductive-isolation prezygotic-isolation postzygotic-isolation mechanical-isolation temporal-isolation habitat-isolation behavioral-isolation gametic-isolation zygotic-mortality hybrid-inviability hybrid-sterility hybrid-breakdown reinforcement character-displacement ecological-speciation allopatric-speciation sympatric-speciation parapatric-speciation peripatric-speciation centrifugal-speciation divergence-with-gene-flow speciation-continuum anagenesis cladogenesis budding-speciation branching-speciation phyletic-gradualism punctuated-equilibrium stasis trend progression orthogenesis aristogenesis mosaic-evolution co-evolution arms-race Red-Queen-hypothesis evolutionary-stable-strategy frequency-dependent-selection density-dependent-selection r-selection K-selection bet-hedging iteroparity semelparity ploidy-level chromosome-number karyotype genome-size transposable-element gene-duplication neofunctionalization subfunctionalization pseudogene regulatory-evolution cis-regulatory trans-regulatory enhancer promoter silencer insulator transcription-factor signaling-pathway developmental-pathway gene-network epigenetic-modification DNA-methylation histone-modification chromatin-remodeling genomic-imprinting parental-effect maternal-effect paternal-effect cytoplasmic-inheritance mitochondrial-DNA chloroplast-DNA endosymbiont Wolbachia Rickettsia Spiroplasma Cardinium Arsenophonus Hamiltonella Regiella Serratia Buchnera Carsonella Portiera primary-endosymbiont secondary-endosymbiont facultative-symbiont obligate-symbiont mutualistic-symbiont parasitic-symbiont commensal-symbiont pathogen vector disease-transmission zoonosis emerging-infectious-disease biosecurity quarantine eradication control management conservation-biology restoration-ecology habitat-restoration river-restoration stream-restoration floodplain-restoration riparian-restoration wetland-restoration ecosystem-restoration passive-restoration active-restoration assisted-recovery managed-relocation assisted-colonization ex-situ-conservation in-situ-conservation protected-area national-park nature-reserve wildlife-refuge biosphere-reserve world-heritage-site ramsar-site important-bird-area key-biodiversity-area Alliance-for-Zero-Extinction EDGE-species evolutionarily-distinct globally-endangered conservation-prioritization systematic-conservation-planning Marxan Zonation conservation-opportunity conservation-cost conservation-benefit trade-off socioeconomic-factor stakeholder-engagement community-based-conservation adaptive-management monitoring-program long-term-study citizen-science public-participation environmental-education outreach science-communication knowledge-transfer capacity-building institutional-strengthening policy-development legislation regulation enforcement compliance incentive payment-for-ecosystem-services carbon-credit biodiversity-offset mitigation-banking conservation-easement land-trust private-land-conservation conservation-covenant stewardship sustainable-use sustainable-management ecosystem-based-management integrated-water-resources-management basin-management catchment-management landscape-management seascape-management connectivity-conservation corridor stepping-stone buffer-zone matrix-management countryside-biogeography reconciliation-ecology novel-ecosystem designer-ecosystem ecosystem-service provisioning-service regulating-service supporting-service cultural-service food-production water-supply raw-materials genetic-resources medicinal-resources ornamental-resources air-quality-regulation climate-regulation water-regulation erosion-regulation water-purification waste-treatment disease-regulation pest-regulation pollination seed-dispersal habitat-provision nutrient-cycling soil-formation primary-production recreation tourism aesthetic-value spiritual-value educational-value scientific-value inspiration sense-of-place cultural-heritage traditional-knowledge indigenous-knowledge local-knowledge ecosystem-service-valuation economic-valuation contingent-valuation choice-experiment benefit-transfer total-economic-value use-value non-use-value direct-use indirect-use option-value bequest-value existence-value willingness-to-pay willingness-to-accept cost-benefit-analysis cost-effectiveness-analysis multi-criteria-analysis decision-support uncertainty risk-assessment scenario-analysis sensitivity-analysis robustness-analysis adaptive-capacity resilience vulnerability exposure sensitivity coping-capacity transformation social-ecological-system complex-adaptive-system tipping-point regime-shift alternative-stable-state hysteresis path-dependence legacy-effect slow-variable fast-variable panarchy adaptive-cycle exploitation release reorganization foresight anticipatory-governance transformative-change sustainability-science future-earth interdisciplinary transdisciplinary problem-oriented solution-oriented action-oriented knowledge-co-production boundary-work boundary-object boundary-organization science-policy-interface science-society-interface knowledge-brokerage knowledge-exchange knowledge-integration mode-1-science mode-2-science triple-helix quadruple-helix responsible-innovation reflexive-governance adaptive-governance collaborative-governance polycentric-governance network-governance self-organization collective-action common-pool-resource tragedy-of-the-commons social-dilemma institutional-analysis institutional-design Ostrom-design-principles monitoring sanctioning conflict-resolution nested-enterprises proportionality collective-choice constitutional-choice operational-choice collective-action-theory game-theory prisoner's-dilemma chicken-game assurance-game coordination-game evolutionary-game-theory replicator-dynamics ESS Nash-equilibrium Pareto-optimality social-optimum market-failure government-failure voluntary-failure non-market-value public-good common-good club-good private-good rivalry excludability free-rider hold-up-problem moral-hazard adverse-selection information-asymmetry transaction-cost property-right common-property state-property private-property hybrid-property bundle-of-rights usufruct access withdrawal alienation tenure-security land-tenure water-tenure resource-tenure customary-tenure statutory-tenure legal-pluralism de-facto de-jure formal informal legality legitimacy recognition deviance corruption elite-capture social-capital bonding-capital bridging-capital linking-capital trust reciprocity reputation social-network social-embeddedness structural-hole brokerage closure cohesion density centrality betweenness degree eigenvector PageRank small-world scale-free random-network regular-network hierarchical-network modularity community-structure clustering assortativity homophily heterophily preferential-attachment fitness-model network-evolution network-dynamics temporal-network multilayer-network multiplex-network bipartite-network affiliation-network one-mode-network two-mode-network adjacency-matrix edge-list graph-theory vertex edge node link tie connection path cycle tree forest component giant-component percolation phase-transition criticality self-organized-criticality power-law heavy-tail Pareto-distribution lognormal-distribution exponential-distribution Poisson-distribution binomial-distribution normal-distribution Gaussian-distribution statistical-distribution probability-distribution cumulative-distribution probability-density-function cumulative-distribution-function survival-function hazard-function moment mean median mode variance standard-deviation coefficient-of-variation skewness kurtosis quantile percentile decile quartile interquartile-range range minimum maximum outlier robust-statistic parametric-test non-parametric-test hypothesis-testing null-hypothesis alternative-hypothesis significance-level p-value confidence-interval type-I-error type-II-error statistical-power effect-size sample-size sampling-design experimental-design observational-study quasi-experiment natural-experiment randomized-controlled-trial before-after-control-impact BACI impact-assessment environmental-impact-assessment strategic-environmental-assessment cumulative-impact-assessment social-impact-assessment health-impact-assessment life-cycle-assessment carbon-footprint water-footprint ecological-footprint material-flow-analysis substance-flow-analysis energy-analysis exergy-analysis emergy-analysis input-output-analysis environmentally-extended-input-output social-accounting-matrix computable-general-equilibrium agent-based-model system-dynamics-model Bayesian-network structural-equation-model path-analysis factor-analysis principal-component-analysis cluster-analysis discriminant-analysis canonical-correspondence-analysis redundancy-analysis non-metric-multidimensional-scaling ordination classification regression-analysis linear-regression nonlinear-regression generalized-linear-model generalized-additive-model mixed-effects-model multilevel-model hierarchical-model Bayesian-hierarchical-model state-space-model hidden-Markov-model time-series-analysis spatial-analysis geostatistics kriging variogram spatial-autocorrelation Moran's-I Geary's-c Getis-Ord-Gi local-indicator-of-spatial-association LISA hot-spot cold-spot spatial-cluster spatial-outlier edge-effect boundary-effect modifiable-areal-unit-problem MAUP ecological-fallacy atomistic-fallacy cross-scale-inference upscaling downscaling scaling-law allometric-scaling fractal fractal-dimension self-similarity multi-fractal lacunarity roughness texture pattern process pattern-process-relationship neutral-model null-model randomization-test permutation-test bootstrap jackknife cross-validation model-averaging model-comparison information-criterion Bayesian-model-averaging ensemble-modeling consensus-model prediction forecast projection scenario narrative-scenario quantitative-scenario storyline shared-socioeconomic-pathway representative-concentration-pathway emission-scenario climate-scenario impact-scenario adaptation-scenario mitigation-scenario transformation-scenario backcasting forecasting trend-extrapolation expert-judgment Delphi-method participatory-modeling stakeholder-analysis influence-diagram conceptual-model mental-model system-archetype leverage-point intervention-point policy-instrument economic-instrument regulatory-instrument voluntary-instrument information-instrument market-based-instrument command-and-control-instrument subsidy tax charge fee levy deposit-refund tradable-permit emission-trading carbon-tax biodiversity-banking conditional-cash-transfer community-based-natural-resource-management joint-forest-management co-management adaptive-co-management transboundary-management international-regime global-governance multilateral-environmental-agreement convention treaty cartagena-protocol nagoya-protocol Aichi-target sustainable-development-goal SDG post-2020-global-biodiversity-framework Kunming-Montreal-global-biodiversity-framework 30-by-30-target nature-based-solution NbS ecosystem-based-adaptation ecosystem-based-mitigation green-infrastructure blue-infrastructure natural-infrastructure hybrid-infrastructure climate-smart-agriculture climate-smart-landscape climate-smart-development low-carbon-development green-economy circular-economy bioeconomy blue-economy sustainable-consumption sustainable-production life-cycle-thinking cradle-to-cradle industrial-ecology industrial-symbiosis urban-metabolism socio-ecological-metabolism material-intensity energy-intensity carbon-intensity decoupling absolute-decoupling relative-decoupling resource-efficiency eco-efficiency eco-innovation green-technology clean-technology environmental-technology appropriate-technology participatory-technology technology-transfer technology-diffusion innovation-system national-innovation-system sectoral-innovation-system technological-innovation-system socio-technical-system socio-technical-transition multi-level-perspective niche-innovation regime landscape transition-management strategic-niche-management transition-experiment living-lab real-world-laboratory urban-living-lab rural-living-lab sustainability-transition fundamental-change system-innovation directionality reflexivity responsiveness anticipation learning-by-doing doing-by-learning social-learning organizational-learning policy-learning transformative-capacity capacity-development institutional-learning evolutionary-governance institutional-change path-dependency lock-in carbon-lock-in technological-lock-in institutional-lock-in behavioral-lock-in cultural-lock-in unlocking destabilization phase-out divestment stranded-asset just-transition transition-pathway transition-scenario deep-decarbonization net-zero carbon-neutral climate-neutral nature-positive biodiversity-positive restoration-positive regenerative circular doughnut-economics planetary-boundary safe-operating-space environmental-ceiling social-foundation intergenerational-equity intragenerational-equity distributive-justice procedural-justice recognition-justice environmental-justice climate-justice energy-justice water-justice food-justice spatial-justice territorial-justice ecological-justice species-justice intersectionality adaptation mitigation sustainability sustainable-development well-being human-development capability-approach human-security environmental-security climate-security water-security food-security energy-security livelihood-security disaster-risk-reduction climate-change-adaptation climate-change-mitigation loss-and-damage climate-finance green-climate-fund adaptation-fund global-environment-facility biodiversity-finance nature-finance blended-finance impact-investment ESG environmental-social-governance sustainable-finance green-bond social-bond sustainability-bond blue-bond nature-bond biodiversity-credit offset inset nature-based-credit ecosystem-service-credit payment-for-results results-based-payment conditional-payment performance-based-payment outcome-based-payment environmental-service-payment PSA PES REDD REDD+AFOLU LULUCF NAMA NAMC NDC NAP NBSAP national-adaptation-plan national-biodiversity-strategy-and-action-plan nationally-determined-contribution nationally-appropriate-mitigation-action intended-nationally-determined-contribution long-term-low-emission-development-strategy net-zero-target carbon-budget remaining-carbon-budget emission-gap adaptation-gap finance-gap implementation-gap ambition-loop ratchet-mechanism global-stocktake transparency-framework enhanced-transparency-framework common-but-differentiated-responsibilities respective-capabilities climate-equity climate-litigation climate-governance climate-diplomacy climate-negotiation UNFCCC COP CMP CMA SBSTA SBI IPCC IPBES UNCCD CBD CITES Ramsar World-Heritage Man-and-the-Biosphere MAB UNESCO FAO WHO ILO UNDP UNEP UN-Habitat World-Bank IMF WTO OECD GEF GCF AF 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seed-commons water-commons fisheries-commons forest-commons pasture-commons land-commons urban-commons atmospheric-commons global-commons common-heritage-of-mankind common-concern-of-humankind restorative-justice multispecies-justice more-than-human anthropocene capitalocene plantationocene chthulucene symbiocene good-anthropocene great-acceleration great-transformation great-decoupling great-convergence great-divergence great-fragmentation great-integration great-transition great-work long-now deep-time futures-thinking future-studies strategic-foresight scenario-planning trend-analysis weak-signal wild-card black-swan grey-rhino blue-sky green-swan systemic-change structural-change cultural-change behavioral-change individual-change collective-change social-change organizational-change technological-change economic-change political-change ecological-change planetary-change earth-system-change biodiversity-change ocean-change cryosphere-change atmospheric-change biogeochemical-change 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