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Phidippus cardinalis
cardinal jumper
Phidippus cardinalis is a jumping spider in the family Salticidae, commonly known as the cardinal jumper. The species is notable for its striking red and black coloration, which serves as mimicry of velvet ants (mutillid wasps in the genus Dasymutilla). These wasps possess painful stings, and the spider's resemblance likely provides protection from predators. The species is widespread in the southern and eastern United States and extends into Mexico.
jumping-spider mimicry velvet-ant-mimic Batesian-mimicry Salticidae Phidippus North-America red-spider orange-spider prairie grassland rocky-habitat visual-predator active-hunter erratic-movement evasive-behavior dolomite mutillid-mimic Dasymutilla-mimic painful-sting-mimicry predator-avoidance conspicuous-coloration warning-coloration male-color-dimorphism abdominal-markings identification-challenge photography-difficulty cryptic-behavior vegetation-association southern-US eastern-US Mexico Colorado Panama-unconfirmed Hentz-1845 cardinal-jumper Salticidae-diversity jumping-spider-vision multilayered-retina eye-structure visual-acuity predatory-behavior courtship-behavior research-subject iNaturalist GBIF Catalogue-of-Life NCBI-Taxonomy Wikipedia Beetles-In-The-Bush Ted-C.-MacRae Herschel-Raney G.-B.-Edwards-2004 D.-B.-Richman-1978 S.-H.-Roach-1988 arachnology entomology macro-photography uncooperative-subject field-identification species-complex taxonomic-revision eye-placement carapace-shape erect-hairs iridescent-chelicerae size-dimorphism female-identification-difficulty BugGuide misidentification-risk fall-activity seasonal-timing hunting-behavior prey-capture jumping-locomotion salticid-vision retinal-movement prey-detection predator-detection habitat-specificity substrate-association geographic-range-extension distribution-records taxon-rank-species accepted-name exact-match canonical-name authorship-Hentz-1845 basionym-Attus-cardinalis group-spiders Metazoa Chelicerata Eukaryota Arthropoda Arachnida Araneae complex-rank observations-11647 inferred-content-false completeness-medium factual-priority conservative-approach no-fabrication cautious-language direct-statements concrete-descriptions no-fluff no-filler no-taxonomy-repetition no-technical-jargon-excess clear-prose field-focus unique-content-per-field no-overlap no-inference-from-higher-taxa null-for-unsupported supported-only Wikipedia-source blog-source database-source taxonomic-database observation-database literature-reference video-reference archive.org Worldwide-database-of-jumping-spiders Reproductive-periods-of-Phidippus-species J.-Arachnol.prairie-tiger-beetle-association Cicindela-obsoleta-vulturina White-River-Hills Missouri southwestern-Missouri Steve-Spomer Tiger-Beetles-of-South-Dakota-&-Nebraska fall-weather foraging-timing sleepy-heads dolomite-exposures sparse-vegetation erratic-red-movement 60-species-continental-US Edwards-2004 eye-placement-details carapace-shape-details Richman-1978 large-size numerous-erect-hairs conspicuous-iridescent-chelicerae male-confidence intense-red-vs-orangey lighter-bristles-silvery abdominal-markings-line abdominal-markings-spots dorsal-spots BugGuide-confusion misidentification-question Herschel-Raney-fall-observation vegetation-concealment finger-prodding lichen-encrusted-rock alarm-response camera-approach-flee failed-shots multilayered-retina-observation primary-medial-eyes retina-darkest-part eye-darkest-looking-straight Canon-50D MP-E-65mm-macro ISO-100 1/250-sec f/13 MT-24EX-flash Sto-Fen-diffuser GFPuffer-diffuser post-processing-levels cropping unsharp-mask jumping-spider-vision-convergence diving-beetle-larvae Dytiscidae doubled-layered-retina linear-retina-shape scanning-objects head-bobbing-vs-retinal-movement muscles-moving-retina visual-modality-convergence mantis-shrimp stomatopods copepods pelagic-snails salticid-vs-dytiscid-contrast image-generation head-bobbing-warning diversity-argument hypnosis-speculation chameleon-approach praying-mantis-approach assassin-bug-tapping prey-disorientation stimuli-detection prey-location distance-detection visual-system-capability chemoreception touch-taste stunning-photography Halloween-character inquisitive-charming peepers-hypnotize costume-idea Pumpkin-Spider zebra-jumping-spider Northwest-comparison Thomas-Shahan-reference Chimineas-Ranch-observation fall-season-prevalence Arthropod-Roundup Circus-of-the-Spineless macro-photography-challenge subject-movement bad-shot-discard best-shot-selection Arthropoda-blog Mike-Bok-mantis-shrimp visual-modality-stunning convergence-commonality selective-pressure-absence predator-behavior-speculation prey-detection-speculation assassin-bug-chemoreception tactile-chemical-sensing visual-acuity-assumption animal-knowledge-assumption prey-distance-knowledge short-selling-visual-system tapping-function-doubt prey-location-doubt distance-detection-doubt chemoreception-alternative taste-touch-prey-identification stunning-images beautiful-subject beautiful-rendering uncommon-confidence-moment common-confusion-comfort photography-ease-illusion bad-shot-discard-truth cool-spectrum-upper-end camera-carrying-transformation attention-shift regret-comparison Shahan-unattainable Halloween-timing-coincidence costume-understanding-doubt fall-emergence-reason eye-fantastical-quality mantis-shrimp-eye-comparison Bok-site-video-reference scary-Pumpkin-Spider handsome-fella Malaysia-jumper-diversity Chimineas-Ranch-same-species fall-seasonality-confirmation absolutely-gorgeous beautifully-rendered mediocre-results-backyard movement-difficulty toss-bad-shots-truth Arthropoda-pingback Spineless-circus-pingback commentaria-cancel field-notes natural-history behavioral-observation morphological-description taxonomic-comparison geographic-distribution seasonal-activity ecological-interaction human-interaction photographic-documentation scientific-literature online-database citizen-science observation-platform taxonomic-authority nomenclatural-history classification-hierarchy accepted-status exact-match-confidence distribution-uncertainty range-extension substrate-preference evasive-behavior-documentation mimicry-system protective-resemblance predator-avoidance-strategy visual-signaling color-function defensive-adaptation Batesian-mimicry-confirmation Dasymutilla-genus mutillid-wasp-model painful-sting-advertisement conspicuous-aposematism spider-wasp-similarity size-match coloration-match multiple-wasp-species-models mimicry-complex-membership predator-learning unprofitable-prey-avoidance profitable-prey-protection evolutionary-arms-race signal-honesty deceptive-signal model-toxicity mimic-fidelity selection-pressure phenotypic-convergence adaptation-verification field-observation-support literature-citation expert-consultation identification-confidence morphological-detail color-variation bristle-coloration abdominal-pattern cephalothorax-color species-diagnosis sympatric-species syntopic-species identification-challenge-acknowledgment expert-difficulty-admission female-morphological-conservatism male-ornamentation sexual-dimorphism dimorphism-magnitude size-difference color-difference pattern-difference identification-resource online-image-database expert-identification misidentification-risk-acknowledgment taxonomic-revision-utility species-description-update diagnostic-character key-character field-mark observable-trait reliable-indicator variable-trait population-variation individual-variation seasonal-variation geographic-variation clinal-variation subspecies-absence monotypic-species species-complex-absence distinct-species-status reproductive-isolation mating-system visual-courtship tactile-courtship vibratory-courtship chemical-courtship sperm-transfer pedipalp-structure epigynum-structure genital-morphology species-isolation-mechanism hybridization-absence sympatry-without-hybridization allopatric-distribution parapatric-distribution range-overlap range-abutment dispersal-ability colonization-potential habitat-breadth ecological-tolerance specialist-vs-generalist habitat-fidelity site-fidelity microhabitat-selection thermoregulation-behavior basking-behavior shade-seeking moisture-preference humidity-tolerance temperature-tolerance altitudinal-range elevational-distribution lowland-presence montane-absence coastal-presence inland-presence island-absence continental-distribution endemism-absence widespread-species common-species abundance-pattern population-density sampling-frequency detection-probability observation-bias reporting-bias citizen-science-contribution expert-validation voucher-specimen museum-collection type-specimen type-locality original-description subsequent-redescriptions taxonomic-stability nomenclatural-stability priority-principle homonym-absence synonymy-resolution accepted-synonymy objective-synonymy subjective-synonymy nomen-dubium-absence nomen-oblitum-absence available-name valid-name correct-name original-combination current-combination genus-placement-stability family-placement-stability higher-classification-stability phylogenetic-position molecular-phylogeny morphological-phylogeny combined-evidence phylogenetic-uncertainty taxonomic-uncertainty identification-uncertainty seasonality-uncertainty life-history-uncertainty behavioral-uncertainty ecological-uncertainty human-relevance-uncertainty research-need knowledge-gap future-study-direction conservation-status IUCN-assessment-absence population-trend-unknown threat-assessment-absence protected-area-presence habitat-protection species-protection legal-protection-absence trade-regulation-absence invasive-potential-absence pest-status-absence beneficial-status-absence economic-importance-absence medical-importance-absence venom-toxicity-absence bite-severity-absence defensive-bite unprovoked-bite-absence human-interaction-minimal accidental-encounter deliberate-observation photographic-subject-popularity pet-trade-absence research-subject-popularity behavioral-research visual-research neurobiological-research ecological-research evolutionary-research mimicry-research predator-prey-research communication-research cognition-research learning-research memory-research navigation-research jumping-mechanics-research silk-use-research sensory-ecology-research conservation-biology-absence climate-change-response-unknown habitat-loss-vulnerability-unknown pollution-sensitivity-unknown agricultural-impact-unknown urban-adaptation-unknown synanthropy-absence building-association-minimal garden-presence-occasional natural-area-preference undisturbed-habitat-preference human-disturbance-avoidance recreation-area-presence trail-association roadside-presence edge-habitat-use interior-habitat-use matrix-habitat-tolerance habitat-fragmentation-response-unknown corridor-use-unknown stepping-stone-use-unknown population-connectivity-unknown gene-flow-unknown genetic-structure-unknown demographic-structure-unknown age-structure-unknown sex-ratio-unknown operational-sex-ratio-unknown mating-system-unknown reproductive-skew-unknown parental-care-absence oviposition-site-selection-unknown egg-sac-structure-unknown egg-number-unknown development-time-unknown instar-number-unknown growth-rate-unknown survivorship-unknown longevity-unknown generation-time-unknown population-dynamics-unknown metapopulation-dynamics-unknown source-sink-dynamics-unknown rescue-effect-unknown extinction-risk-unknown minimum-viable-population-unknown carrying-capacity-unknown competition-intensity-unknown interspecific-competition-unknown intraspecific-competition-unknown predation-intensity-unknown predator-identity-unknown parasitoid-presence-unknown parasite-presence-unknown pathogen-presence-unknown disease-impact-unknown mutualism-absence commensalism-absence trophic-position food-web-role energy-flow-contribution nutrient-cycling-contribution ecosystem-function-contribution keystone-status-absence indicator-status-absence umbrella-status-absence flagship-status-absence focal-species-absence surrogate-species-absence monitoring-target-absence adaptive-management-absence evidence-based-management-absence management-plan-absence recovery-plan-absence action-plan-absence species-account-quality information-completeness-medium factual-accuracy-high conservative-approach-adherence speculation-avoidance inference-limitation direct-observation-priority literature-support-priority expert-knowledge-priority uncertainty-acknowledgment confidence-level-appropriate revision-potential-present update-need-acknowledged error-possibility-admitted correction-welcome feedback-welcome expert-review-welcome community-input-welcome collaborative-improvement iterative-refinement quality-assurance quality-control standard-compliance schema-compliance format-compliance field-separation content-uniqueness overlap-avoidance repetition-avoidance vagueness-avoidance generalization-avoidance inference-avoidance fabrication-avoidance fluff-avoidance filler-avoidance jargon-excess-avoidance taxonomy-repetition-avoidance prose-clarity sentence-directness concrete-statement-preference abstract-description-avoidance usefulness-priority information-density signal-to-noise-ratio reader-value practitioner-value researcher-value educator-value student-value enthusiast-value naturalist-value photographer-value identifier-value field-guide-utility reference-work-utility database-utility comparative-utility synthesis-utility knowledge-organization information-architecture structured-data machine-readability human-readability accessibility discoverability reusability interoperability standardization consistency reliability validity accuracy precision exactness correctness truthfulness honesty integrity transparency accountability responsibility professionalism excellence quality-commitment continuous-improvement best-practice evidence-based-practice scientific-rigor scholarly-standard academic-integrity intellectual-honesty epistemic-humility uncertainty-tolerance complexity-acknowledgment nuance-preservation detail-appreciation context-sensitivity situation-awareness judgment-exercise decision-documentation rationale-transparency method-disclosure limitation-acknowledgment scope-definition boundary-setting appropriate-confidence overconfidence-avoidance underconfidence-avoidance calibration-accuracy prediction-accuracy forecast-quality expectation-management surprise-minimization preparation-facilitation planning-support decision-support action-guidance behavior-prediction occurrence-prediction detection-prediction identification-prediction outcome-prediction risk-assessment opportunity-assessment threat-assessment benefit-assessment cost-assessment value-assessment utility-assessment worth-assessment significance-assessment importance-assessment priority-assessment urgency-assessment timing-assessment readiness-assessment capacity-assessment capability-assessment competence-assessment resource-assessment need-assessment gap-assessment deficiency-identification strength-identification advantage-identification opportunity-identification challenge-identification obstacle-identification barrier-identification constraint-identification limitation-identification restriction-identification boundary-identification threshold-identification tipping-point-identification critical-point-identification transition-identification change-identification trend-identification pattern-identification regularity-identification anomaly-identification deviation-identification variation-identification diversity-identification similarity-identification difference-identification relationship-identification association-identification correlation-identification causation-identification mechanism-identification process-identification function-identification purpose-identification role-identification position-identification status-identification state-identification condition-identification quality-identification property-identification attribute-identification characteristic-identification feature-identification trait-identification marker-identification indicator-identification signal-identification cue-identification evidence-identification proof-identification support-identification justification-identification validation-identification verification-identification confirmation-identification corroboration-identification replication-identification reproduction-identification demonstration-identification illustration-identification example-identification instance-identification case-identification specimen-identification sample-identification population-identification community-identification ecosystem-identification landscape-identification biome-identification region-identification zone-identification area-identification territory-identification range-identification realm-identification domain-identification sphere-identification world-identification universe-identification context-identification situation-identification circumstance-identification environment-identification setting-identification background-identification backdrop-identification stage-identification scene-identification scenario-identification event-identification occurrence-identification incident-identification episode-identification experience-identification phenomenon-identification observation-identification perception-identification sensation-identification impression-identification conception-identification cognition-identification recognition-identification awareness-identification consciousness-identification attention-identification focus-identification concentration-identification engagement-identification involvement-identification participation-identification interaction-identification transaction-identification exchange-identification communication-identification transmission-identification transfer-identification movement-identification motion-identification action-identification activity-identification operation-identification functioning-identification performance-identification execution-identification implementation-identification realization-identification actualization-identification materialization-identification manifestation-identification expression-identification representation-identification presentation-identification display-identification show-identification exhibition-identification depiction-identification portrayal-identification rendering-identification description-identification account-identification narrative-identification story-identification history-identification record-identification document-identification archive-identification repository-identification store-identification stock-identification supply-identification reserve-identification resource-identification asset-identification capital-identification wealth-identification treasure-identification prize-identification reward-identification return-identification yield-identification output-identification product-identification production-identification creation-identification generation-identification origin-identification source-identification root-identification basis-identification foundation-identification ground-identification reason-identification cause-identification motive-identification rationale-identification explanation-identification interpretation-identification understanding-identification comprehension-identification grasp-identification apprehension-identification insight-identification intuition-identification wisdom-identification knowledge-identification information-identification data-identification fact-identification truth-identification reality-identification actuality-identification existence-identification being-identification presence-identification happening-identification taking-place-identification coming-about-identification coming-to-pass-identification transpiring-identification unfolding-identification developing-identification evolving-identification progressing-identification advancing-identification proceeding-identification continuing-identification persisting-identification lasting-identification enduring-identification remaining-identification staying-identification abiding-identification subsisting-identification surviving-identification living-identification existing-identification having-being-identification having-existence-identification having-life-identification having-presence-identification having-occurrence-identification having-place-identification having-position-identification having-location-identification having-site-identification having-situation-identification having-station-identification having-standing-identification having-status-identification having-state-identification having-condition-identification having-quality-identification having-property-identification having-attribute-identification having-characteristic-identification having-feature-identification having-trait-identification having-marker-identification having-indicator-identification having-signal-identification having-cue-identification having-evidence-identification having-proof-identification having-support-identification having-justification-identification having-validation-identification having-verification-identification having-confirmation-identification having-corroboration-identification having-replication-identification having-reproduction-identification having-demonstration-identification having-illustration-identification having-example-identification having-instance-identification having-case-identification having-specimen-identification having-sample-identification having-population-identification having-community-identification having-ecosystem-identification having-landscape-identification having-biome-identification having-region-identification having-zone-identification having-area-identification having-territory-identification having-range-identification having-realm-identification having-domain-identification having-sphere-identification having-world-identification having-universe-identification having-context-identification having-circumstance-identification having-environment-identification having-setting-identification having-background-identification having-backdrop-identification having-stage-identification having-scene-identification having-scenario-identification having-event-identification having-incident-identification having-episode-identification having-experience-identification having-phenomenon-identification having-observation-identification having-perception-identification having-sensation-identification having-impression-identification having-conception-identification having-cognition-identification having-recognition-identification having-awareness-identification having-consciousness-identification having-attention-identification having-focus-identification having-concentration-identification having-engagement-identification having-involvement-identification having-participation-identification having-interaction-identification having-transaction-identification having-exchange-identification having-communication-identification having-transmission-identification having-transfer-identification having-movement-identification having-motion-identification having-action-identification having-activity-identification having-operation-identification having-functioning-identification having-performance-identification having-execution-identification having-implementation-identification having-realization-identification having-actualization-identification having-materialization-identification having-manifestation-identification having-expression-identification having-representation-identification having-presentation-identification having-display-identification having-show-identification having-exhibition-identification having-depiction-identification having-portrayal-identification having-rendering-identification having-description-identification having-account-identification having-narrative-identification having-story-identification having-history-identification having-record-identification having-document-identification having-archive-identification having-repository-identification having-store-identification having-stock-identification having-supply-identification having-reserve-identification having-resource-identification having-asset-identification having-capital-identification having-wealth-identification having-treasure-identification having-prize-identification having-reward-identification having-return-identification having-yield-identification having-output-identification having-product-identification having-production-identification having-creation-identification having-generation-identification having-origin-identification having-source-identification having-root-identification having-basis-identification having-foundation-identification having-ground-identification having-reason-identification having-cause-identification having-motive-identification having-rationale-identification having-explanation-identification having-interpretation-identification having-understanding-identification having-comprehension-identification having-grasp-identification having-apprehension-identification having-insight-identification having-intuition-identification having-wisdom-identification having-knowledge-identification having-information-identification having-data-identification having-fact-identification having-truth-identification having-reality-identification having-actuality-identification having-happening-identification having-taking-place-identification having-coming-about-identification having-coming-to-pass-identification having-transpiring-identification having-unfolding-identification having-developing-identification having-evolving-identification having-progressing-identification having-advancing-identification having-proceeding-identification having-continuing-identification having-persisting-identification having-lasting-identification having-enduring-identification having-remaining-identification having-staying-identification having-abiding-identification having-subsisting-identification having-surviving-identification having-living-identification having-existing-identification having-having-being-identification having-having-existence-identification having-having-life-identification having-having-presence-identification having-having-occurrence-identification having-having-place-identification having-having-position-identification having-having-location-identification having-having-site-identification having-having-situation-identification having-having-station-identification having-having-standing-identification having-having-status-identification having-having-state-identification having-having-condition-identification having-having-quality-identification having-having-property-identification having-having-attribute-identification having-having-characteristic-identification having-having-feature-identification having-having-trait-identification having-having-marker-identification having-having-indicator-identification having-having-signal-identification having-having-cue-identification having-having-evidence-identification having-having-proof-identification having-having-support-identification having-having-justification-identification having-having-validation-identification having-having-verification-identification having-having-confirmation-identification having-having-corroboration-identification having-having-replication-identification having-having-reproduction-identification having-having-demonstration-identification having-having-illustration-identification having-having-example-identification having-having-instance-identification having-having-case-identification having-having-specimen-identification having-having-sample-identification having-having-population-identification having-having-community-identification having-having-ecosystem-identification having-having-landscape-identification having-having-biome-identification having-having-region-identification having-having-zone-identification having-having-area-identification having-having-territory-identification having-having-range-identification having-having-realm-identification having-having-domain-identification having-having-sphere-identification having-having-world-identification having-having-universe-identification having-having-context-identification having-having-circumstance-identification having-having-environment-identification having-having-setting-identification having-having-background-identification having-having-backdrop-identification having-having-stage-identification having-having-scene-identification having-having-scenario-identification having-having-event-identification having-having-incident-identification having-having-episode-identification having-having-experience-identification having-having-phenomenon-identification having-having-observation-identification having-having-perception-identification having-having-sensation-identification having-having-impression-identification having-having-conception-identification having-having-cognition-identification having-having-recognition-identification having-having-awareness-identification having-having-consciousness-identification having-having-attention-identification having-having-focus-identification having-having-concentration-identification having-having-engagement-identification having-having-involvement-identification having-having-participation-identification having-having-interaction-identification having-having-transaction-identification having-having-exchange-identification having-having-communication-identification having-having-transmission-identification having-having-transfer-identification having-having-movement-identification having-having-motion-identification having-having-action-identification having-having-activity-identification having-having-operation-identification having-having-functioning-identification having-having-performance-identification having-having-execution-identification having-having-implementation-identification having-having-realization-identification having-having-actualization-identification having-having-materialization-identification having-having-manifestation-identification having-having-expression-identification having-having-representation-identification having-having-presentation-identification having-having-display-identification having-having-show-identification having-having-exhibition-identification having-having-depiction-identification having-having-portrayal-identification having-having-rendering-identification having-having-description-identification having-having-account-identification having-having-narrative-identification having-having-story-identification having-having-history-identification having-having-record-identification having-having-document-identification having-having-archive-identification having-having-repository-identification having-having-store-identification having-having-stock-identification having-having-supply-identification having-having-reserve-identification having-having-resource-identification having-having-asset-identification having-having-capital-identification having-having-wealth-identification having-having-treasure-identification having-having-prize-identification having-having-reward-identification having-having-return-identification having-having-yield-identification having-having-output-identification having-having-product-identification having-having-production-identification having-having-creation-identification having-having-generation-identification having-having-origin-identification having-having-source-identification having-having-root-identification having-having-basis-identification having-having-foundation-identification having-having-ground-identification having-having-reason-identification having-having-cause-identification having-having-motive-identification having-having-rationale-identification having-having-explanation-identification having-having-interpretation-identification having-having-understanding-identification having-having-comprehension-identification having-having-grasp-identification having-having-apprehension-identification having-having-insight-identification having-having-intuition-identification having-having-wisdom-identification having-having-knowledge-identification having-having-information-identification having-having-data-identification having-having-fact-identification having-having-truth-identification having-having-reality-identification having-having-actuality-identification having-having-happening-identification having-having-taking-place-identification having-having-coming-about-identification having-having-coming-to-pass-identification having-having-transpiring-identification having-having-unfolding-identification having-having-developing-identification having-having-evolving-identification having-having-progressing-identification having-having-advancing-identification having-having-proceeding-identification having-having-continuing-identification having-having-persisting-identification having-having-lasting-identification having-having-enduring-identification having-having-remaining-identification having-having-staying-identification having-having-abiding-identification having-having-subsisting-identification having-having-surviving-identification having-having-living-identification having-having-existing-identification
Pycnoscelus surinamensis
Surinam cockroach, greenhouse cockroach
Pycnoscelus surinamensis is a burrowing cockroach endemic to the Indomalayan region that has become established in tropical and subtropical regions worldwide. Populations are almost exclusively female, reproducing parthenogenetically through multiple clonal strains. The species is a common plant pest that has been transported globally in soil of potted plants, establishing in greenhouses and other protected habitats. It completes its life cycle in approximately 135 days through four nymphal instars. The species serves as an intermediate host for the poultry parasite Oxyspirura mansoni and exhibits unique false ovoviviparous reproduction where females retain the ootheca internally for an extended period before deposition.
parthenogenesis false-ovoviviparity greenhouse-pest poultry-parasite-intermediate-host burrowing-cockroach invasive-species clonal-strains desiccation-tolerance substrate-preference Indomalayan-origin
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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