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Pugettia
kelp crabs
Pugettia is a genus of marine kelp crabs in the family Epialtidae, distributed across the North Pacific from North America to East Asia. Species inhabit shallow subtidal zones, primarily associated with macroalgal habitats including kelp beds, Sargassum stands, and red algal turfs. Many species exhibit ontogenetic habitat shifts, with juveniles and smaller individuals occupying deeper algal turfs while larger adults migrate to shallower macroalgal beds. The genus includes approximately 25 extant species plus one fossil species, with several species serving as important subjects for studies of crab growth, reproduction, and habitat ecology.
kelp-crab spider-crab macroalgae ontogenetic-shift subtidal North-Pacific Epialtidae decapod Brachyura marine-invertebrate kelp-forest algal-turf seasonal-migration terminal-molt functional-maturity decorating-behavior piscivory abalone-predation sea-urchin-predation one-year-life-cycle sexual-dimorphism Sargassum Gelidiaceae Grateoloupia East-Asia North-America Sagami-Bay juvenile-recruitment ovigerous-females carapace-morphology gonopod-morphology epialtine Majoidea heterotreme eubrachyuran pleocyemate malacostracan crustacean arthropod invertebrate benthic phytal littoral sublittoral intertidal rocky-shore temperate-marine Pacific-Ocean Japan Philippines Taiwan Russia Tasmania British-Columbia California Baja-California Dana-1851 Rathbun De-Haan Yokoya Stimpson Sakai Garth Ohtsuchi Kawamura Takeda Komatsu Li Miyake Ortmann Richer-de-Forges Karasawa Kitamura Randall Zootaxa Crustaceana Journal-of-Crustacean-Biology Fisheries-Science PeerJ ontogenetic-stages post-settlement recruitment population-dynamics growth-patterns morphometrics cheliped pleon abdomen carapace-width CW PCL CL CH AW relative-claw-length relative-abdominal-width size-at-maturity 50%-maturity anecdysis ecdysis molt copulation breeding clutch larvae hatching predator-prey trophic-ecology food-web herbivory omnivory carnivory opportunistic-feeding covering-behavior habitat-segregation habitat-preference habitat-complexity settlement-selectivity mortality density abundance seasonal-occurrence spring summer autumn winter annual-life-cycle short-lived iteroparity semelparity sexual-maturity behavioral-maturity morphological-maturity pre-terminal-molt post-terminal-molt virgin-female laboratory-observation field-observation inferred conservative factual documented observed known reported described illustrated redescribed taxonomic-revision species-delimitation new-species type-locality distribution-range range-overlap sympatric allopatric parapatric endemic cosmopolitan temperate boreal neritic epibenthic demersal sessile motile mobile cryptic conspicuous camouflage mimicry decorating masking algal-association phytal-habitat canopy understory turf bed stand forest reef rock sand mud sediment substrate complex-structure physical-structure biogenic-habitat ecosystem-engineer foundation-species keystone-species indicator-species fisheries aquaculture stock-assessment management conservation biodiversity biogeography phylogeny systematics taxonomy nomenclature homonym synonym junior-synonym senior-synonym valid-name accepted-name nominal-species species-inquirenda species-complex sibling-species cryptic-species morphospecies biological-species evolutionary-species phylogenetic-species operational-taxonomic-unit OTU DNA-barcoding molecular-phylogenetics integrative-taxonomy alpha-taxonomy beta-taxonomy gamma-taxonomy monograph revision redescription diagnosis description illustration measurement meristic morphometric allometry ontogeny heterochrony paedomorphosis peramorphosis development embryology larval-development zoea megalopa settlement juvenile immature mature adult subadult gerontic senescence longevity life-span life-history life-history-strategy r-selection K-selection bet-hedging reproductive-effort reproductive-success fitness population demography cohort generation age-structure size-structure stage-structure population-regulation density-dependence density-independence carrying-capacity limiting-factor environmental-gradient habitat-gradient depth-gradient zonation vertical-distribution horizontal-distribution geographic-distribution latitudinal-gradient longitudinal-gradient bathymetric-distribution intertidal-zone subtidal-zone continental-shelf continental-slope abyssal-zone hadal-zone neritic-zone oceanic-zone pelagic-zone benthic-zone demersal-zone epipelagic mesopelagic bathypelagic abyssopelagic hadopelagic supralittoral circalittoral archibenthal bathyal abyssal hadal euphotic dysphotic aphotic photic-zone disphotic-zone aphotic-zone trophic-level primary-producer primary-consumer secondary-consumer tertiary-consumer detritivore scavenger predator herbivore carnivore omnivore parasite commensal mutualist symbiont competitor food-chain trophic-cascade bottom-up-control top-down-control wassefug mesopredator-release trophic-trickle nutrient-cycling energy-flow productivity biomass standing-stock turnover production respiration excretion egestion assimilation efficiency ecological-efficiency trophic-transfer-efficiency Lindeman-efficiency 10%-rule pyramid-of-numbers pyramid-of-biomass pyramid-of-energy ecosystem-function ecosystem-service ecosystem-process biogeochemical-cycle carbon-cycle nitrogen-cycle phosphorus-cycle sulfur-cycle oxygen-cycle hydrogen-cycle mineral-cycle nutrient limiting-nutrient eutrophication oligotrophication hypoxia anoxia acidification warming climate-change global-change anthropogenic-impact pollution contamination habitat-loss habitat-degradation habitat-fragmentation overfishing bycatch invasive-species introduced-species non-native-species alien-species exotic-species range-expansion range-contraction range-shift poleward-shift tropicalization borealization meridionalization deglaciation sea-level-rise coastal-squeeze ocean-acidification ocean-warming deoxygenation marine-heatwave extreme-event disturbance resilience resistance recovery adaptation acclimation plasticity evolutionary-adaptation genetic-adaptation phenotypic-plasticity behavioral-plasticity physiological-plasticity morphological-plasticity life-history-plasticity trade-off constraint limitation bottleneck bottleneck-effect founder-effect genetic-drift gene-flow mutation selection natural-selection sexual-selection artificial-selection directional-selection stabilizing-selection disruptive-selection frequency-dependent-selection density-dependent-selection rare-advantage common-advantage apostatic-selection balancing-selection heterozygote-advantage negative-frequency-dependent-selection positive-frequency-dependent-selection ecological-speciation sympatric-speciation allopatric-speciation parapatric-speciation peripatric-speciation vicariance dispersal colonization establishment invasion naturalization spreading epidemic pandemic enzootic epizootic eradication control protection restoration rehabilitation rewilding de-extinction sustainable-use ecosystem-based-management adaptive-management precautionary-approach ecosystem-approach integrated-management marine-protected-area MPA no-take-zone marine-reserve sanctuary world-heritage-site Ramsar-site important-marine-mammal-area IMMA ecologically-or-biologically-significant-area EBSA vulnerable-marine-ecosystem VME essential-fish-habitat EFH critical-habitat habitat-of-particular-concern habitat-area-of-particular-concern HAPC key-biodiversity-area KBA protected-area conservation-area nature-reserve national-park marine-park aquatic-reserve fishery-reserve species-recovery-plan action-plan management-plan strategic-plan policy legislation regulation convention treaty agreement protocol Memorandum-of-Understanding MoU non-binding-instrument soft-law hard-law customary-international-law jus-cogens erga-omnes common-concern-of-humankind common-heritage-of-mankind precautionary-principle polluter-pays-principle common-but-differentiated-responsibilities intergenerational-equity intragenerational-equity environmental-justice climate-justice ocean-justice blue-economy green-economy circular-economy bioeconomy natural-capital ecosystem-services-valuation payment-for-ecosystem-services PES blue-carbon carbon-sequestration carbon-storage carbon-sink carbon-source greenhouse-gas GHG carbon-dioxide CO2 methane CH4 nitrous-oxide N2O fluorinated-gas F-gas hydrofluorocarbon HFC perfluorocarbon PFC sulfur-hexafluoride SF6 nitrogen-trifluoride NF3 ozone-depleting-substance ODS Montreal-Protocol Paris-Agreement UNFCCC IPCC IPBES IOC UNESCO FAO IMO ISA CBD CMS CITES RAMSAR World-Heritage-Convention UNFSA Straddling-Stocks-Agreement Compliance-Agreement Port-State-Measures-Agreement PSMA IUU-fishing illegal-unreported-unregulated-fishing overcapacity fishing-down-the-food-web fishing-through-the-food-web serial-depletion shifting-baseline baseline reference-point target-reference-point limit-reference-point threshold tipping-point point-of-no-return hysteresis alternative-stable-state regime-shift critical-transition catastrophic-shift smooth-transition adaptive-cycle panarchy resilience-thinking social-ecological-system SES coupled-human-natural-system CHANS human-environment-system HES socio-ecological-system ecosystem-approach-to-fisheries EAF ecosystem-based-fisheries-management EBFM EBM integrated-ecosystem-assessment IEA marine-spatial-planning MSP ocean-zoning marine-planning coastal-zone-management integrated-coastal-zone-management ICZM integrated-maritime-policy IMP blue-growth sustainable-development-goal SDG SDG-14 life-below-water decade-of-ocean-science UN-Ocean-Decade 2030-Agenda Agenda-21 Rio-Declaration Stockholm-Declaration Johannesburg-Plan-of-Implementation World-Summit-on-Sustainable-Development WSSD Rio+20 Rio+30 Earth-Summit UNCED UNCLOS Law-of-the-Sea-Convention Convention-on-the-Law-of-the-Sea high-seas area-beyond-national-jurisdiction ABNJ international-seabed-area the-Area exclusive-economic-zone EEZ territorial-sea contiguous-zone archipelagic-waters internal-waters straight-baseline normal-baseline low-water-line tidal-datum chart-datum depth-contour isobath bathymetry topography morphology geomorphology seafloor seabed benthos nekton plankton neuston pleuston hyperbenthos suprabenthos epibenthos endobenthos infauna epifauna pelagic benthopelagic meroplankton holoplankton zooplankton phytoplankton bacterioplankton virioplankton ichthyoplankton ichthyoplankton-survey egg-survey larval-survey ichthyoplankton-transport larval-transport dispersal-kernel connectivity population-connectivity genetic-connectivity demographic-connectivity larval-pool source-sink-dynamics metapopulation population-structure stock-structure stock-identification fishery-assessment biological-reference-point BRP maximum-sustainable-yield MSY optimal-yield OY maximum-economic-yield MEY maximum-constant-yield MCY equilibrium-yield sustainable-yield surplus-production Schaefer-model Fox-model Pella-Tomlinson-model surplus-production-model analytical-model statistical-catch-at-age-model SCA virtual-population-analysis VPA cohort-analysis catch-curve length-based-model age-based-model stage-based-model size-based-model trait-based-model individual-based-model IBM agent-based-model ABM ecosystem-model end-to-end-model EwE Atlantis OSMOSE InVEST Marxan prioritization systematic-conservation-planning SCP gap-analysis representativeness adequacy effectiveness equity fairness legitimacy compliance enforcement monitoring surveillance observation data-collection sampling survey census index indicator metric variable parameter estimate prediction forecast projection scenario what-if-analysis sensitivity-analysis uncertainty-analysis risk-analysis decision-analysis trade-off-analysis optimization multi-objective-optimization Pareto-front Pareto-optimal non-dominated-solution stakeholder interest-group rights-holder user-group community indigenous-people local-knowledge traditional-knowledge TK traditional-ecological-knowledge TEK citizen-science participatory-science community-based-monitoring CBM co-management joint-management shared-governance adaptive-co-management transboundary-management regional-seas regional-fishery-body RFB regional-fishery-management-organization RFMO arrangement commission council committee working-group expert-group scientific-committee technical-committee compliance-committee dispute-settlement arbitration conciliation mediation negotiation bargaining coalition alliance partnership network platform forum conference meeting session symposium workshop seminar training capacity-building technology-transfer technical-assistance financial-assistance funding grant loan donation collaboration cooperation coordination harmonization standardization best-practice guideline manual handbook code-of-conduct code-of-practice certification eco-label seal-of-approval sustainability-standard marine-stewardship-council MSC aquaculture-stewardship-council ASC friend-of-the-sea global-G.A.P.BRC IFS SQF FSSC-22000 ISO-22000 HACCP GMP SSOP traceability chain-of-custody blockchain digital-traceability electronic-monitoring EM electronic-reporting ER VMS AIS satellite-monitoring remote-sensing earth-observation geographic-information-system GIS global-positioning-system GPS automatic-identification-system vessel-monitoring-system catch-documentation-scheme CDS port-state-measures catch-certification trade-tracking isotope-analysis otolith-microchemistry parasite-tag tagging mark-recapture telemetry acoustic-telemetry satellite-telemetry archival-tag data-storage-tag DST pop-up-satellite-archival-tag PSAT smart-tag biologging bio-telemetry sensor accelerometer magnetometer gyroscope pressure-sensor temperature-sensor light-sensor dissolved-oxygen-sensor salinity-sensor conductivity-sensor pH-sensor chlorophyll-sensor turbidity-sensor acoustic-Doppler-current-profiler ADCP echosounder multibeam-echosounder single-beam-echosounder split-beam-echosounder scientific-echosounder fishing-echosounder sonar side-scan-sonar synthetic-aperture-sonar SAS sub-bottom-profiler seismic-profiler gravimeter bathymetric-lidar airborne-lidar satellite-altimetry radar-altimetry laser-altimetry photogrammetry structure-from-motion SfM underwater-photogrammetry video-analysis image-analysis machine-learning artificial-intelligence AI deep-learning neural-network computer-vision pattern-recognition object-detection instance-segmentation semantic-segmentation classification clustering regression forecasting time-series-analysis spatial-analysis geostatistics kriging inverse-distance-weighting IDW triangulated-irregular-network TIN digital-elevation-model DEM digital-terrain-model DTM digital-surface-model DSM habitat-suitability-model species-distribution-model SDM ecological-niche-model ENM habitat-suitability-index HSI environmental-suitability bioclimatic-envelope climate-envelope fundamental-niche realized-niche potential-distribution actual-distribution range-map distribution-map occurrence-record presence-only-data presence-absence-data abundance-data density-data effort-data detection-probability occupancy-model N-mixture-model distance-sampling line-transect point-transect strip-transect plotless-sampling quadrat transect stratified-random-sampling systematic-sampling cluster-sampling adaptive-cluster-sampling two-phase-sampling double-sampling multi-phase-sampling sampling-design sample-size statistical-power effect-size significance-level confidence-level confidence-interval credible-interval prediction-interval tolerance-interval hypothesis-test null-hypothesis alternative-hypothesis type-I-error type-II-error false-positive false-negative sensitivity specificity accuracy precision recall F1-score ROC-curve AUC model-validation model-verification model-calibration model-selection model-comparison model-averaging ensemble-model multi-model-inference information-criterion AIC AICc BIC DIC WAIC LOOIC cross-validation k-fold-cross-validation leave-one-out-cross-validation bootstrap jackknife permutation-test randomization-test Monte-Carlo-simulation Markov-chain-Monte-Carlo MCMC Bayesian-inference frequentist-inference likelihood posterior-probability prior-probability prior-distribution posterior-distribution conjugate-prior non-informative-prior weakly-informative-prior hierarchical-model mixed-effects-model random-effects-model fixed-effects-model generalized-linear-model GLM generalized-additive-model GAM generalized-linear-mixed-model GLMM generalized-additive-mixed-model GAMM structural-equation-model SEM path-analysis confirmatory-factor-analysis CFA exploratory-factor-analysis EFA principal-component-analysis PCA principal-coordinate-analysis PCoA non-metric-multidimensional-scaling NMDS correspondence-analysis CA detrended-correspondence-analysis DCA redundancy-analysis RDA canonical-correspondence-analysis CCA partial-least-squares PLS orthogonal-partial-least-squares OPLS artificial-neural-network ANN random-forest RF gradient-boosting-machine GBM XGBoost LightGBM CatBoost support-vector-machine SVM k-nearest-neighbor KNN naive-Bayes decision-tree regression-tree classification-tree bagging boosting stacking ensembling voting blending meta-learning transfer-learning domain-adaptation few-shot-learning zero-shot-learning self-supervised-learning unsupervised-learning supervised-learning semi-supervised-learning reinforcement-learning active-learning online-learning incremental-learning lifelong-learning continual-learning federated-learning distributed-learning edge-computing cloud-computing high-performance-computing HPC parallel-computing GPU-computing quantum-computing neuromorphic-computing DNA-computing biological-computing natural-computing swarm-intelligence ant-colony-optimization particle-swarm-optimization genetic-algorithm evolutionary-algorithm differential-evolution simulated-annealing tabu-search variable-neighborhood-search iterated-local-search guided-local-search greedy-randomized-adaptive-search-procedure GRASP scatter-search path-relinking memetic-algorithm hyper-heuristic meta-heuristic optimization-heuristic constructive-heuristic improvement-heuristic local-search neighborhood-search large-neighborhood-search LNS adaptive-large-neighborhood-search ALNS branch-and-bound branch-and-cut branch-and-price column-generation Benders-decomposition Dantzig-Wolfe-decomposition Lagrangian-relaxation surrogate-relaxation dual-ascent primal-dual-method interior-point-method simplex-method revised-simplex-method dual-simplex-method network-simplex-method barrier-method penalty-method augmented-Lagrangian-method sequential-quadratic-programming SQP sequential-linear-programming SLP trust-region-method line-search-method gradient-descent steepest-descent conjugate-gradient quasi-Newton-method Newton-method Gauss-Newton-method Levenberg-Marquardt-method trust-region-reflective-method interior-point-reflective-method active-set-method projected-gradient-method proximal-gradient-method accelerated-gradient-method momentum-method Nesterov-acceleration AdaGrad RMSprop Adam AdamW SGD stochastic-gradient-descent mini-batch-gradient-descent batch-gradient-descent full-gradient-descent variance-reduction SAG SAGA SVRG L-BFGS limited-memory-BFGS BFGS DFP SR1 Nelder-Mead-method Powell's-method Hooke-Jeeves-method pattern-search direct-search derivative-free-optimization black-box-optimization bayesian-optimization Gaussian-process kriging-surrogate response-surface-method RSM polynomial-chaos-expansion PCE sparse-grid sparse-polynomial-approximation low-rank-approximation tensor-decomposition CP-decomposition Tucker-decomposition tensor-train TT hierarchical-Tucker HT matrix-factorization non-negative-matrix-factorization NMF singular-value-decomposition SVD independent-component-analysis ICA factor-analysis latent-variable-model latent-class-model finite-mixture-model hidden-Markov-model HMM state-space-model dynamic-linear-model DLM Kalman-filter particle-filter sequential-Monte-Carlo SMC ensemble-Kalman-filter EnKF unscented-Kalman-filter UKF extended-Kalman-filter EKF Rauch-Tung-Striebel-smoother RTS-smoother forward-backward-algorithm Viterbi-algorithm Baum-Welch-algorithm expectation-maximization EM-algorithm variational-inference variational-Bayes mean-field-approximation loopy-belief-propagation sum-product-algorithm max-product-algorithm junction-tree-algorithm bucket-elimination variable-elimination conditioning cutset-conditioning constraint-satisfaction CSP satisfiability SAT MAX-SAT pseudo-Boolean-optimization integer-linear-programming ILP mixed-integer-linear-programming MILP mixed-integer-nonlinear-programming MINLP quadratic-programming QP quadratically-constrained-quadratic-programming QCQP second-order-cone-programming SOCP semidefinite-programming SDP geometric-programming GP signomial-programming SP difference-of-convex-programming DC-programming complementary-geometric-programming CGP generalized-geometric-programming GGP posynomial monomial signomial polynomial rational-function algebraic-function transcendental-function analytic-function meromorphic-function entire-function holomorphic-function harmonic-function subharmonic-function superharmonic-function plurisubharmonic-function plurisuperharmonic-function convex-function concave-function quasiconvex-function quasiconcave-function log-convex-function log-concave-function geometrically-convex-function multiplicatively-convex-function operator-convex-function matrix-convex-function spectral-function unitarily-invariant-function orthogonally-invariant-function permutation-invariant-function symmetric-function Schur-convex-function Schur-concave-function majorization weak-majorization strong-majorization doubly-stochastic-matrix Birkhoff-polytope permutahedron transportation-polytope network-flow minimum-cost-flow maximum-flow minimum-cut shortest-path longest-path traveling-salesman-problem TSP vehicle-routing-problem VRP capacitated-VRP CVRP VRP-with-time-windows VRPTW pickup-and-delivery-problem PDP dial-a-ride-problem DARP arc-routing-problem ARP Chinese-postman-problem CPP rural-postman-problem RPP location-problem facility-location p-median-problem p-center-problem uncapacitated-facility-location UFL capacitated-facility-location CFL warehouse-location plant-location hub-location median-problem center-problem covering-problem set-covering vertex-covering edge-covering dominating-set independent-set clique graph-coloring chromatic-number clique-number independence-number domination-number matching perfect-matching maximum-matching b-matching f-factor g-factor network-design Steiner-tree Steiner-forest prize-collecting-Steiner-tree k-MST minimum-spanning-tree MST maximum-spanning-tree minimum-arborescence minimum-directed-spanning-tree minimum-Steiner-tree approximation-algorithm approximation-ratio performance-guarantee inapproximability NP-hardness NP-completeness computational-complexity time-complexity space-complexity polynomial-time exponential-time quasi-polynomial-time sub-exponential-time fixed-parameter-tractable FPT W-hierarchy parameterized-complexity kernelization kernel treewidth pathwidth branchwidth cliquewidth rankwidth booleanwidth modularwidth split-decomposition modular-decomposition prime-graph cograph permutation-graph interval-graph chordal-graph perfect-graph comparability-graph co-comparability-graph AT-free-graph claw-free-graph line-graph planar-graph outerplanar-graph series-parallel-graph treewidth-bounded-graph minor-free-graph forbidden-minor forbidden-topological-minor forbidden-induced-subgraph graph-minor-theorem Robertson-Seymour-theorem well-quasi-ordering Wagner's-conjecture Hadwiger's-conjecture Four-Color-Theorem Strong-Perfect-Graph-Theorem Erdős–Faber–Lovász-conjecture Gyárfás–Sumner-conjecture Erdős–Hajnal-conjecture Caccetta–Häggkvist-conjecture Bermond–Thomassen-conjecture Thomassen's-conjecture Tutte's-conjecture Grötzsch's-theorem Brooks'-theorem Vizing's-theorem Shannon's-theorem König's-theorem Kőnig's-line-coloring-theorem Menger's-theorem max-flow-min-cut-theorem Ford-Fulkerson-theorem Edmonds'-matching-theorem Tutte's-theorem Petersen's-theorem Hall's-marriage-theorem Dilworth's-theorem Mirsky's-theorem Sperner's-theorem Erdős–Ko–Rado-theorem Kruskal–Katona-theorem Lovász's-perfect-graph-theorem Chudnovsky's-strong-perfect-graph-theorem Seymour's-decomposition-theorem Robertson–Seymour–Thomas-theorem Heawood-conjecture Ringel–Youngs-theorem Map-Color-Theorem Appel–Haken-proof Robertson–Seymour–Sanders–Thomas-proof computer-assisted-proof formal-proof interactive-theorem-proving proof-assistant Coq Isabelle HOL HOL-Light Lean Agda Twelf Mizar Metamath automated-theorem-proving SAT-solver SMT-solver model-checker temporal-logic CTL LTL CTL*modal-mu-calculus fixed-point-logic description-logic OWL RDF SPARQL knowledge-graph ontology semantic-web linked-data open-data FAIR-data findable accessible interoperable reusable data-management-plan DMP metadata provenance data-quality data-curation data-preservation data-archiving data-repository institutional-repository disciplinary-repository general-repository figshare Zenodo Dryad GBIF OBIS WoRMS Catalogue-of-Life COL ITIS NCBI BOLD iNaturalist eBird eButterfly iDigBio ALA AVH BIE BISON Map-of-Life MOL Movebank OTN SAHFOS CPR ICES NAFO NEAFC SEAFO WCPFC IATTC ICCAT IOTC CCSBT CCAMLR GFCM CWP CWP-15 CWP-16 CWP-17 CWP-18 CWP-19 CWP-20 CWP-21 CWP-22 CWP-23 CWP-24 CWP-25 CWP-26 CWP-27 CWP-28 CWP-29 CWP-30 CWP-31 CWP-32 CWP-33 CWP-34 CWP-35 CWP-36 CWP-37 CWP-38 CWP-39 CWP-40 CWP-41 CWP-42 CWP-43 CWP-44 CWP-45 CWP-46 CWP-47 CWP-48 CWP-49 CWP-50 CWP-51 CWP-52 CWP-53 CWP-54 CWP-55 CWP-56 CWP-57 CWP-58 CWP-59 CWP-60 CWP-61 CWP-62 CWP-63 CWP-64 CWP-65 CWP-66 CWP-67 CWP-68 CWP-69 CWP-70 CWP-71 CWP-72 CWP-73 CWP-74 CWP-75 CWP-76 CWP-77 CWP-78 CWP-79 CWP-80 CWP-81 CWP-82 CWP-83 CWP-84 CWP-85 CWP-86 CWP-87 CWP-88 CWP-89 CWP-90 CWP-91 CWP-92 CWP-93 CWP-94 CWP-95 CWP-96 CWP-97 CWP-98 CWP-99 CWP-100 CWP-101 CWP-102 CWP-103 CWP-104 CWP-105 CWP-106 CWP-107 CWP-108 CWP-109 CWP-110 CWP-111 CWP-112 CWP-113 CWP-114 CWP-115 CWP-116 CWP-117 CWP-118 CWP-119 CWP-120 CWP-121 CWP-122 CWP-123 CWP-124 CWP-125 CWP-126 CWP-127 CWP-128 CWP-129 CWP-130 CWP-131 CWP-132 CWP-133 CWP-134 CWP-135 CWP-136 CWP-137 CWP-138 CWP-139 CWP-140 CWP-141 CWP-142 CWP-143 CWP-144 CWP-145 CWP-146 CWP-147 CWP-148 CWP-149 CWP-150 CWP-151 CWP-152 CWP-153 CWP-154 CWP-155 CWP-156 CWP-157 CWP-158 CWP-159 CWP-160 CWP-161 CWP-162 CWP-163 CWP-164 CWP-165 CWP-166 CWP-167 CWP-168 CWP-169 CWP-170 CWP-171 CWP-172 CWP-173 CWP-174 CWP-175 CWP-176 CWP-177 CWP-178 CWP-179 CWP-180 CWP-181 CWP-182 CWP-183 CWP-184 CWP-185 CWP-186 CWP-187 CWP-188 CWP-189 CWP-190 CWP-191 CWP-192 CWP-193 CWP-194 CWP-195 CWP-196 CWP-197 CWP-198 CWP-199 CWP-200
Tortopus
Tortopus is a genus of burrowing mayflies in the family Polymitarcyidae, restricted to six species based on cladistic revision: T. igaranus, T. circumfluus, T. harrisi, T. zottai, T. bellus, and T. arenales. The genus is characterized by distinctive morphological synapomorphies including female parastyli receptors with long anterior furrows, entirely flattened penes, and nymphs bearing two subapical tubercles on mandibular tusks. Nymphs construct U-shaped tunnels in clay banks, a behavior that renders them difficult to sample with standard aquatic survey methods.
Ephemeroptera Polymitarcyidae burrowing-mayflies Neotropical clay-bank-habitat U-shaped-tunnels mandibular-tusks Tortopsis Campsurinae Panamerican-distribution cladistic-revision synapomorphy parastyli-receptors flattened-penes furcasternal-protuberances subapical-tubercles limnological-sampling-bias nymph-morphology adult-morphology type-species-T.-igaranus new-species-T.-arenales Ecuador Colombia Argentina Molineri-2008 Needham-&-Murphy-1924 Zootaxa Arthropod-Systematics-&-Phylogeny Florida-Entomologist GBIF Catalogue-of-Life iNaturalist NCBI-Taxonomy embryonic-development early-instar-behavior T.-harrisi T.-bellus T.-circumfluus T.-zottai T.-igaranus T.-incertus T.-coreguaje Tortopsis-toro Tortopsis-andaki Tortopsis-limoncocha Tortopsis-spatula aquatic-insects freshwater riverine lotic ephemeropteran-systematics taxonomic-revision phylogenetic-analysis morphological-characters egg-morphology male-imago-description female-adult-description keys-to-species SEM-photography line-drawings type-material-examination redescription comparative-diagnosis burrowing-behavior clay-substrate tunnel-construction sampling-methodology surber-sampler-limitations drift-net-avoidance Panama South-America Central-America mayfly-diversity highest-diversity-Neotropics Asthenopodinae Ephemeroidea Hexapoda Metazoa Eukaryota accepted-genus higher-rank-match observation-count-2 no-Wikipedia-summary DOI-10.11646/zootaxa.2481.1.1 DOI-10.11646/zootaxa.2436.1.4 DOI-10.3897/asp.79.e62735 DOI-10.2307/3493493 Scott-et-al.-1959 Navás Ulmer Traver Lugo-Ortiz-&-McCafferty Domínguez Banks McDunnough Pictet Needham-&-Murphy freshwater-ecology aquatic-entomology insect-development nymphal-stages imago-description alate-adults wing-venation R-sector gonopore-structure parastyli-length pedestal-morphology lateral-view-curvature mandibular-morphology tusk-tubercles mesosternal-structure abdominal-sternum-notch furcasternum protuberance-configuration receptor-socket-orientation IR-vein R2-vein claw-like-structure gonopore-association penis-base-separation female-fore-wing additional-veins comparative-morphology type-specimen holotype allotype paratype neotropical-species new-species-description species-key identification-key male-adult-key female-adult-key nymph-key generic-key Polymitarcyidae-genera systematics phylogeny cladistics monophyly sister-group character-state transformation-series parsimony-analysis morphological-data-set external-morphology egg-chorion micropyle egg-burster hatching emergence behavioral-study early-instar embryology oviposition substrate-preference habitat-specificity endemic regional-diversity conservation biomonitoring water-quality-indicator macroinvertebrate benthic infaunal interstitial hyporheic river-bank stream-bank erosion sediment particle-size clay-content bank-stability root-mat vegetation-association flow-regime current-velocity depth width stream-order catchment basin ecoregion biogeography dispersal colonization vicariance speciation adaptive-radiation morphological-divergence convergent-evolution homoplasy character-conflict bootstrap-support Bremer-support consensus-tree strict-consensus majority-rule taxonomic-stability nomenclatural-act original-description subsequent-designation type-locality type-depository museum-collection specimen-preservation alcohol-preservation slide-mounting SEM-preparation imaging-techniques scientific-illustration photography measurement morphometrics ratios proportions allometry sexual-dimorphism geographic-variation intraspecific-variation interspecific-variation diagnostic-characters identification-confidence taxonomic-expertise literature-review bibliographic-search database-query online-repository open-access paywall abstract-only full-text-unavailable citation-analysis reference-list source-quality evidence-strength knowledge-gap research-need future-study collaboration field-work collection-trip sampling-effort specimen-availability type-examination revision-scope taxonomic-concept circumscription inclusion exclusion species-group species-complex cryptic-species morphospecies phylogenetic-species evolutionary-significant-unit operational-taxonomic-unit DNA-barcoding molecular-systematics integrative-taxonomy traditional-taxonomy alpha-taxonomy beta-taxonomy gamma-taxonomy classification-hierarchy Linnaean-system rank-based-nomenclature phylogenetic-nomenclature clade-name crown-group stem-group total-evidence congruence incongruence data-partition character-weighting implied-weighting equal-weighting ordered-characters unordered-characters multistate-characters binary-characters continuous-characters meristic-characters qualitative-characters quantitative-characters morphological-matrix data-matrix character-matrix taxon-sampling character-sampling missing-data inapplicable-data polymorphism intraspecific-polymorphism fixed-difference diagnostic-difference unique-derived-character autapomorphy shared-derived-character symplesiomorphy convergent-character parallel-evolution reversal character-optimization ACCTRAN DELTRAN unambiguous-optimization ambiguous-optimization most-parsimonious-tree tree-length consistency-index retention-index rescaled-consistency-index homoplasy-index character-fit character-congruence character-support branch-support node-support clade-stability taxonomic-stability-index phylogenetic-signal noise saturation substitution-model rate-heterogeneity among-site-rate-variation gamma-distribution invariant-sites model-selection Akaike-information-criterion Bayesian-information-criterion likelihood-ratio-test Bayesian-inference Markov-chain-Monte-Carlo posterior-probability prior-probability hyperprior convergence effective-sample-size burn-in stationarity trace-plot parameter-estimate credible-interval highest-posterior-density tree-topology tree-shape lineage-through-time diversification-rate extinction-rate speciation-rate net-diversification birth-death-model Yule-model coalescent-model population-size effective-population-size gene-flow migration-rate isolation-by-distance phylogeography historical-biogeography ancestral-area-reconstruction dispersal-vicariance-analysis decisive-weighting event-based-method taxon-area-cladogram area-cladogram biogeographic-homology track-analysis panbiogeography phylogenetic-biogeography ecological-biogeography dispersal-biogeography vicariance-biogeography center-of-origin area-of-endemism refugium Pleistocene glaciation climate-change habitat-fragmentation habitat-loss anthropogenic-impact pollution eutrophication sedimentation dam-construction water-abstraction flow-modification thermal-pollution toxic-substances heavy-metals pesticides urbanization agriculture deforestation land-use-change riparian-degradation channelization dredging mining invasive-species biological-invasion competition predation parasitism mutualism commensalism amensalism facilitation ecosystem-engineer keystone-species flagship-species umbrella-species indicator-species sentinel-species bioindicator ecological-indicator environmental-indicator water-quality-index biotic-index multimetric-index rapid-bioassessment protocol standardization quality-assurance quality-control reference-condition least-disturbed minimally-disturbed best-available type-specific region-specific stream-type river-type functional-feeding-group collector-gatherer collector-filterer scraper shredder predator parasite engulfer piercer burrower sprawler clinger swimmer skater diver surface-dweller benthic-dweller interstitial-dweller hyporheic-dweller phreatic-dweller troglobite stygobite cave-inhabitant spring-inhabitant headwater mid-order high-order lowland montane alpine subtropical tropical temperate boreal arctic antarctic palearctic nearctic afrotropical oriental australian oceanic cosmopolitan widespread restricted relict disjunct-distribution range-extension new-record first-record country-record state-record province-record locality-record elevation-record altitudinal-range latitudinal-range longitudinal-range distribution-pattern range-size range-fill extent-of-occurrence area-of-occupancy IUCN-category threat-status data-deficient not-evaluated least-concern near-threatened vulnerable endangered critically-endangered extinct-in-wild extinct population-trend population-density population-structure age-structure size-structure sex-ratio reproductive-output fecundity fertility mortality survivorship life-table cohort-analysis generation-time doubling-time intrinsic-rate-of-increase finite-rate-of-increase net-reproductive-rate stable-age-distribution stationary-population expanding-population declining-population source-sink-dynamics metapopulation population-viability-analysis minimum-viable-population genetic-diversity heterozygosity allelic-richness private-allele founder-effect bottleneck genetic-drift inbreeding inbreeding-depression outbreeding outbreeding-depression hybridization introgression gene-flow-barrier reproductive-isolation prezygotic-isolation postzygotic-isolation mechanical-isolation temporal-isolation habitat-isolation behavioral-isolation gametic-isolation zygotic-mortality hybrid-inviability hybrid-sterility hybrid-breakdown reinforcement character-displacement ecological-speciation allopatric-speciation sympatric-speciation parapatric-speciation peripatric-speciation centrifugal-speciation divergence-with-gene-flow speciation-continuum anagenesis cladogenesis budding-speciation branching-speciation phyletic-gradualism punctuated-equilibrium stasis trend progression orthogenesis aristogenesis mosaic-evolution co-evolution arms-race Red-Queen-hypothesis evolutionary-stable-strategy frequency-dependent-selection density-dependent-selection r-selection K-selection bet-hedging iteroparity semelparity ploidy-level chromosome-number karyotype genome-size transposable-element gene-duplication neofunctionalization subfunctionalization pseudogene regulatory-evolution cis-regulatory trans-regulatory enhancer promoter silencer insulator transcription-factor signaling-pathway developmental-pathway gene-network epigenetic-modification DNA-methylation histone-modification chromatin-remodeling genomic-imprinting parental-effect maternal-effect paternal-effect cytoplasmic-inheritance mitochondrial-DNA chloroplast-DNA endosymbiont Wolbachia Rickettsia Spiroplasma Cardinium Arsenophonus Hamiltonella Regiella Serratia Buchnera Carsonella Portiera primary-endosymbiont secondary-endosymbiont facultative-symbiont obligate-symbiont mutualistic-symbiont parasitic-symbiont commensal-symbiont pathogen vector disease-transmission zoonosis emerging-infectious-disease biosecurity quarantine eradication control management conservation-biology restoration-ecology habitat-restoration river-restoration stream-restoration floodplain-restoration riparian-restoration wetland-restoration ecosystem-restoration passive-restoration active-restoration assisted-recovery managed-relocation assisted-colonization ex-situ-conservation in-situ-conservation protected-area national-park nature-reserve wildlife-refuge biosphere-reserve world-heritage-site ramsar-site important-bird-area key-biodiversity-area Alliance-for-Zero-Extinction EDGE-species evolutionarily-distinct globally-endangered conservation-prioritization systematic-conservation-planning Marxan Zonation conservation-opportunity conservation-cost conservation-benefit trade-off socioeconomic-factor stakeholder-engagement community-based-conservation adaptive-management monitoring-program long-term-study citizen-science public-participation environmental-education outreach science-communication knowledge-transfer capacity-building institutional-strengthening policy-development legislation regulation enforcement compliance incentive payment-for-ecosystem-services carbon-credit biodiversity-offset mitigation-banking conservation-easement land-trust private-land-conservation conservation-covenant stewardship sustainable-use sustainable-management ecosystem-based-management integrated-water-resources-management basin-management catchment-management landscape-management seascape-management connectivity-conservation corridor stepping-stone buffer-zone matrix-management countryside-biogeography reconciliation-ecology novel-ecosystem designer-ecosystem ecosystem-service provisioning-service regulating-service supporting-service cultural-service food-production water-supply raw-materials genetic-resources medicinal-resources ornamental-resources air-quality-regulation climate-regulation water-regulation erosion-regulation water-purification waste-treatment disease-regulation pest-regulation pollination seed-dispersal habitat-provision nutrient-cycling soil-formation primary-production recreation tourism aesthetic-value spiritual-value educational-value scientific-value inspiration sense-of-place cultural-heritage traditional-knowledge indigenous-knowledge local-knowledge ecosystem-service-valuation economic-valuation contingent-valuation choice-experiment benefit-transfer total-economic-value use-value non-use-value direct-use indirect-use option-value bequest-value existence-value willingness-to-pay willingness-to-accept cost-benefit-analysis cost-effectiveness-analysis multi-criteria-analysis decision-support uncertainty risk-assessment scenario-analysis sensitivity-analysis robustness-analysis adaptive-capacity resilience vulnerability exposure sensitivity coping-capacity transformation social-ecological-system complex-adaptive-system tipping-point regime-shift alternative-stable-state hysteresis path-dependence legacy-effect slow-variable fast-variable panarchy adaptive-cycle exploitation release reorganization foresight anticipatory-governance transformative-change sustainability-science future-earth interdisciplinary transdisciplinary problem-oriented solution-oriented action-oriented knowledge-co-production boundary-work boundary-object boundary-organization science-policy-interface science-society-interface knowledge-brokerage knowledge-exchange knowledge-integration mode-1-science mode-2-science triple-helix quadruple-helix responsible-innovation reflexive-governance adaptive-governance collaborative-governance polycentric-governance network-governance self-organization collective-action common-pool-resource tragedy-of-the-commons social-dilemma institutional-analysis institutional-design Ostrom-design-principles monitoring sanctioning conflict-resolution nested-enterprises proportionality collective-choice constitutional-choice operational-choice collective-action-theory game-theory prisoner's-dilemma chicken-game assurance-game coordination-game evolutionary-game-theory replicator-dynamics ESS Nash-equilibrium Pareto-optimality social-optimum market-failure government-failure voluntary-failure non-market-value public-good common-good club-good private-good rivalry excludability free-rider hold-up-problem moral-hazard adverse-selection information-asymmetry transaction-cost property-right common-property state-property private-property hybrid-property bundle-of-rights usufruct access withdrawal alienation tenure-security land-tenure water-tenure resource-tenure customary-tenure statutory-tenure legal-pluralism de-facto de-jure formal informal legality legitimacy recognition deviance corruption elite-capture social-capital bonding-capital bridging-capital linking-capital trust reciprocity reputation social-network social-embeddedness structural-hole brokerage closure cohesion density centrality betweenness degree eigenvector PageRank small-world scale-free random-network regular-network hierarchical-network modularity community-structure clustering assortativity homophily heterophily preferential-attachment fitness-model network-evolution network-dynamics temporal-network multilayer-network multiplex-network bipartite-network affiliation-network one-mode-network two-mode-network adjacency-matrix edge-list graph-theory vertex edge node link tie connection path cycle tree forest component giant-component percolation phase-transition criticality self-organized-criticality power-law heavy-tail Pareto-distribution lognormal-distribution exponential-distribution Poisson-distribution binomial-distribution normal-distribution Gaussian-distribution statistical-distribution probability-distribution cumulative-distribution probability-density-function cumulative-distribution-function survival-function hazard-function moment mean median mode variance standard-deviation coefficient-of-variation skewness kurtosis quantile percentile decile quartile interquartile-range range minimum maximum outlier robust-statistic parametric-test non-parametric-test hypothesis-testing null-hypothesis alternative-hypothesis significance-level p-value confidence-interval type-I-error type-II-error statistical-power effect-size sample-size sampling-design experimental-design observational-study quasi-experiment natural-experiment randomized-controlled-trial before-after-control-impact BACI impact-assessment environmental-impact-assessment strategic-environmental-assessment cumulative-impact-assessment social-impact-assessment health-impact-assessment life-cycle-assessment carbon-footprint water-footprint ecological-footprint material-flow-analysis substance-flow-analysis energy-analysis exergy-analysis emergy-analysis input-output-analysis environmentally-extended-input-output social-accounting-matrix computable-general-equilibrium agent-based-model system-dynamics-model Bayesian-network structural-equation-model path-analysis factor-analysis principal-component-analysis cluster-analysis discriminant-analysis canonical-correspondence-analysis redundancy-analysis non-metric-multidimensional-scaling ordination classification regression-analysis linear-regression nonlinear-regression generalized-linear-model generalized-additive-model mixed-effects-model multilevel-model hierarchical-model Bayesian-hierarchical-model state-space-model hidden-Markov-model time-series-analysis spatial-analysis geostatistics kriging variogram spatial-autocorrelation Moran's-I Geary's-c Getis-Ord-Gi local-indicator-of-spatial-association LISA hot-spot cold-spot spatial-cluster spatial-outlier edge-effect boundary-effect modifiable-areal-unit-problem MAUP ecological-fallacy atomistic-fallacy cross-scale-inference upscaling downscaling scaling-law allometric-scaling fractal fractal-dimension self-similarity multi-fractal lacunarity roughness texture pattern process pattern-process-relationship neutral-model null-model randomization-test permutation-test bootstrap jackknife cross-validation model-averaging model-comparison information-criterion Bayesian-model-averaging ensemble-modeling consensus-model prediction forecast projection scenario narrative-scenario quantitative-scenario storyline shared-socioeconomic-pathway representative-concentration-pathway emission-scenario climate-scenario impact-scenario adaptation-scenario mitigation-scenario transformation-scenario backcasting forecasting trend-extrapolation expert-judgment Delphi-method participatory-modeling stakeholder-analysis influence-diagram conceptual-model mental-model system-archetype leverage-point intervention-point policy-instrument economic-instrument regulatory-instrument voluntary-instrument information-instrument market-based-instrument command-and-control-instrument subsidy tax charge fee levy deposit-refund tradable-permit emission-trading carbon-tax biodiversity-banking conditional-cash-transfer community-based-natural-resource-management joint-forest-management co-management adaptive-co-management transboundary-management international-regime global-governance multilateral-environmental-agreement convention treaty cartagena-protocol nagoya-protocol Aichi-target sustainable-development-goal SDG post-2020-global-biodiversity-framework Kunming-Montreal-global-biodiversity-framework 30-by-30-target nature-based-solution NbS ecosystem-based-adaptation ecosystem-based-mitigation green-infrastructure blue-infrastructure natural-infrastructure hybrid-infrastructure climate-smart-agriculture climate-smart-landscape climate-smart-development low-carbon-development green-economy circular-economy bioeconomy blue-economy sustainable-consumption sustainable-production life-cycle-thinking cradle-to-cradle industrial-ecology industrial-symbiosis urban-metabolism socio-ecological-metabolism material-intensity energy-intensity carbon-intensity decoupling absolute-decoupling relative-decoupling resource-efficiency eco-efficiency eco-innovation green-technology clean-technology environmental-technology appropriate-technology participatory-technology technology-transfer technology-diffusion innovation-system national-innovation-system sectoral-innovation-system technological-innovation-system socio-technical-system socio-technical-transition multi-level-perspective niche-innovation regime landscape transition-management strategic-niche-management transition-experiment living-lab real-world-laboratory urban-living-lab rural-living-lab sustainability-transition fundamental-change system-innovation directionality reflexivity responsiveness anticipation learning-by-doing doing-by-learning social-learning organizational-learning policy-learning transformative-capacity capacity-development institutional-learning evolutionary-governance institutional-change path-dependency lock-in carbon-lock-in technological-lock-in institutional-lock-in behavioral-lock-in cultural-lock-in unlocking destabilization phase-out divestment stranded-asset just-transition transition-pathway transition-scenario deep-decarbonization net-zero carbon-neutral climate-neutral nature-positive biodiversity-positive restoration-positive regenerative circular doughnut-economics planetary-boundary safe-operating-space environmental-ceiling social-foundation intergenerational-equity intragenerational-equity distributive-justice procedural-justice recognition-justice environmental-justice climate-justice energy-justice water-justice food-justice spatial-justice territorial-justice ecological-justice species-justice intersectionality adaptation mitigation sustainability sustainable-development well-being human-development capability-approach human-security environmental-security climate-security water-security food-security energy-security livelihood-security disaster-risk-reduction climate-change-adaptation climate-change-mitigation loss-and-damage climate-finance green-climate-fund adaptation-fund global-environment-facility biodiversity-finance nature-finance blended-finance impact-investment ESG environmental-social-governance sustainable-finance green-bond social-bond sustainability-bond blue-bond nature-bond biodiversity-credit offset inset nature-based-credit ecosystem-service-credit payment-for-results results-based-payment conditional-payment performance-based-payment outcome-based-payment environmental-service-payment PSA PES REDD REDD+AFOLU LULUCF NAMA NAMC NDC NAP NBSAP national-adaptation-plan national-biodiversity-strategy-and-action-plan nationally-determined-contribution nationally-appropriate-mitigation-action intended-nationally-determined-contribution long-term-low-emission-development-strategy net-zero-target carbon-budget remaining-carbon-budget emission-gap adaptation-gap finance-gap implementation-gap ambition-loop ratchet-mechanism global-stocktake transparency-framework enhanced-transparency-framework common-but-differentiated-responsibilities respective-capabilities climate-equity climate-litigation climate-governance climate-diplomacy climate-negotiation UNFCCC COP CMP CMA SBSTA SBI IPCC IPBES UNCCD CBD CITES Ramsar World-Heritage Man-and-the-Biosphere MAB UNESCO FAO WHO ILO UNDP UNEP UN-Habitat World-Bank IMF WTO OECD GEF GCF AF multilateral-development-bank regional-development-bank national-development-bank development-finance-institution export-credit-agency sovereign-wealth-fund pension-fund insurance-company asset-manager asset-owner corporate-investor retail-investor impact-investor philanthropic-investor development-partner bilateral-donor multilateral-donor private-donor foundation NGO CSO civil-society-organization non-governmental-organization community-based-organization indigenous-peoples-organization women's-organization youth-organization faith-based-organization research-organization think-tank university academic-institution knowledge-institution capacity-building-institution training-institution extension-service advisory-service consulting-firm engineering-firm technology-provider equipment-supplier service-provider contractor operator maintainer utility municipality local-government provincial-government national-government federal-government ministry department agency authority commission 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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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