Cold-tolerance

Guides

Cucujus
flat bark beetles
Cucujus is a genus of flat bark beetles in the family Cucujidae, containing approximately 25 recognized species distributed across the Holarctic region. Species are dorso-ventrally compressed, brightly colored in red or red-and-black patterns, and range from 6–25 mm in length. The genus is most diverse in Asia, with indigenous species in North America, Europe, and Asia. Several species, notably C. cinnaberinus and C. clavipes, have been subjects of ecological and physiological research due to their conservation status and extreme cold tolerance adaptations.
saproxylic predator Holarctic cold-tolerance vitrification conservation Natura-2000 dead-wood subcortical flagellum
Cucujus puniceus
Red Flat Bark Beetle
Cucujus puniceus is a species of flat bark beetle in the family Cucujidae, distributed across northern North America. The species has been extensively studied for its exceptional cold tolerance mechanisms. Larvae survive winter temperatures through deep supercooling and vitrification, with documented survival to at least –100°C. The species is treated as a synonym of Cucujus clavipes by some authorities, or as a distinct species or subspecies by others.
cold-tolerance vitrification supercooling cryoprotectants overwintering boreal subarctic
Entomobrya nivalis
Cosmopolitan Springtail
Entomobrya nivalis, commonly known as the cosmopolitan springtail, is a globally distributed species of slender springtail in the family Entomobryidae. Adults measure approximately 2 mm in length and display distinctive coloration with dark transverse bands on a yellow or white background. The species is notable for its cold tolerance, facilitated by antifreeze compounds in its hemolymph, enabling survival in temperate and polar regions. Juveniles inhabit leaf litter while adults migrate to lichen-covered tree bark.
springtail Collembola cosmopolitan-distribution cold-tolerance forest-decomposer
Hyles euphorbiae
spurge hawk-moth, Leafy Spurge Hawkmoth
Hyles euphorbiae, the spurge hawk-moth, is a European hawkmoth in the family Sphingidae. First described by Linnaeus in 1758, this species is notable for its use in biological pest control against leafy spurge (Euphorbia esula). The species comprises a complex with multiple ecotypes showing variation in cold tolerance and larval pattern morphology across its range. Adults are strong fliers with distinctive wing patterns, while larvae feed on spurge plants and display conspicuous black, white, and red coloration.
Sphingidae hawkmoth biological-control Euphorbia diapause cold-tolerance species-complex
Ilybius oblitus
Ilybius oblitus is a predaceous diving beetle in the family Dytiscidae, first described by Sharp in 1882. It occurs in North America, with confirmed records from Ontario, Canada. The species has been documented to tolerate low temperatures, though it remains susceptible to freezing. Observations of this species are sparse, with limited biological data available.
Dytiscidae predaceous-diving-beetle North-America cold-tolerance Ontario
Megabruchidius dorsalis
honey locust seed beetle, Asian seed beetle
Megabruchidius dorsalis is an invasive seed beetle (Chrysomelidae: Bruchinae) native to Asia that has established populations across Europe and North America. The species is oligophagous on Caesalpinioideae seeds, primarily developing within pods of Gleditsia species. Adults exhibit freeze-avoidant cold tolerance strategy with supercooling points around -10°C. The beetle has demonstrated host range expansion in its introduced range, colonizing North American Gymnocladus dioica in addition to its native Asian Gleditsia hosts.
invasive-species seed-beetle Bruchinae Gleditsia-pest cold-tolerance host-range-expansion urban-entomology
Myllocerus
oriental broad-nosed weevils, ash weevils, grey weevils, Sri Lankan weevils
Myllocerus is a large genus of oriental broad-nosed weevils in the family Curculionidae, comprising at least 330 described species. Species within this genus are predominantly distributed across the Indian subcontinent and surrounding regions, with several species recognized as significant agricultural pests. Notable pest species include Myllocerus undecimpustulatus undatus (Sri Lankan weevil), M. maculosus (cotton grey weevil), and M. viridanus (ash weevil), which attack diverse crop plants including cereals, legumes, vegetables, and fruit trees. The genus exhibits typical weevil morphology with broad rostrums and root-feeding larval stages combined with foliage-feeding adults.
Curculionidae Entiminae agricultural-pest root-feeding-larvae foliage-feeding-adults polyphagous sexual-dimorphism invasive-species India Sri-Lanka Florida integrated-pest-management biopesticides cold-tolerance mating-behavior soil-dwelling-larvae
Myllocerus undecimpustulatus
Sri Lanka weevil, yellow-headed ravenous weevil
Myllocerus undecimpustulatus is an oriental broad-nosed weevil (Curculionidae: Entiminae) native to Sri Lanka and the Indian subcontinent. It is a polyphagous pest with a wide host range that has established invasive populations in Florida, USA, since at least 2000. The species exhibits distinctive sexual behavior including female abdominal rocking as a mate-strength test and prolonged copulatory guarding by males. It shows seasonal variation in cold tolerance and feeding activity, with winter-collected adults consuming more leaf material after cold exposure than summer-collected individuals.
invasive pest polyphagous sexual-dimorphism cold-tolerance Sri-Lanka Florida biopesticide-susceptibility mating-behavior copulatory-guarding
Myllocerus undecimpustulatus undatus
Sri Lankan weevil
Myllocerus undecimpustulatus undatus, the Sri Lankan weevil, is a polyphagous curculionid pest native to Sri Lanka that has spread to India, Pakistan, and the United States. It was first identified in Florida in 2000 on Citrus species and has since established populations in the state. The species exhibits sexual dimorphism with females typically larger than males. Research has documented its cold tolerance limitations, mating behavior involving female abdominal rocking and prolonged copulatory guarding, and susceptibility to certain biopesticides.
invasive-species agricultural-pest polyphagous sexual-dimorphism curculionidae sri-lankan-weevil florida cold-tolerance mating-behavior biopesticide
Periplaneta japonica
Japanese cockroach, Yamato cockroach
Periplaneta japonica is a cold-tolerant cockroach native to Japan, adapted to cooler northern climates. It possesses a flexible life cycle with facultative nymphal diapause, allowing nymphs to overwinter once or twice before reaching maturity. The species produces a unique viscous proteinaceous secretion in nymphs that enables active defense against ant predators. First documented in the United States in 2012 in New York City, it has been observed to survive outdoors in freezing temperatures, distinguishing it from most urban cockroach pests.
cockroach invasive-species cold-tolerance diapause urban-pest Japan New-York defensive-secretion DNA-barcoding-identification Periplaneta Blattidae Blattodea
Phragmatobia fuliginosa
Ruby Tiger, Ruby Tiger Moth
Phragmatobia fuliginosa, the Ruby Tiger, is a moth in the subfamily Arctiinae of family Erebidae. It has a wingspan of 35–45 mm and is distinguished by dark reddish-brown forewings with a blackish comma-shaped spot and bright carmine to rose-red hindwings. The species is widely distributed across the Palearctic and into northern North America, with multiple recognized subspecies including the North American P. f. rubricosa. It exhibits geographic variation in generation time: univoltine in northern regions with adults flying in June, and bivoltine in southern Britain with adults appearing April–June and again August–September.
moth Arctiinae tiger-moth ruby-tiger polyphagous cold-tolerance genome-sequenced pyrrolizidine-alkaloids bivoltine univoltine Palearctic Nearctic
Polistes annularis
Ringed Paper Wasp, Jack Spaniard Wasp, Red Paper Wasp
Polistes annularis is a primitively eusocial paper wasp native to the eastern United States, distinguished by its large size and distinctive red-and-black coloration with a prominent yellow ring on the abdomen. The species exhibits complex social behavior including cooperative nest founding by multiple foundresses, dominance hierarchies, and unique overwintering strategies involving honey storage. Nests are constructed in aggregations under overhangs near water bodies, with colonies typically containing 500 or more cells. Unlike many temperate Polistes species, P. annularis shows relatively robust cold tolerance and has been extensively studied as a model organism for social insect behavior and kin recognition.
social-wasp paper-wasp primitively-eusocial nest-aggregation overwintering honey-storage Müllerian-mimicry foundress-association dominance-hierarchy kin-recognition cold-tolerance model-organism caterpillar-predator riparian-habitat annual-colony-cycle queen-succession cuticular-hydrocarbons subgenus-Aphanilopterus Polistinae Vespidae Hymenoptera eastern-United-States red-and-black-coloration ringed-abdomen orange-antennae cliff-nesting hibernacula usurpation altruism reproductive-competition juvenile-hormone ovarian-development microsatellite-markers maternity-assignment behavioral-ecology social-insect nest-architecture petiole-construction foundress-eviction worker-emergence sexual-dimorphism structural-sex-determination trehalose cold-survival ant-predation parasitoid Elasmus-polistis Chalcoela-iphitalis biological-control pollinator sting-allergen antigen-5 Linnaeus-1763 Fabricius-1804 etymology-Greek-polistes Latin-annularis-ringed
Protophormia terraenovae
northern blowfly, blue-bottle fly, blue-assed fly, blackbottle
Protophormia terraenovae is a large, metallic blue-green blowfly with a Holarctic distribution. It is the most cold-tolerant calliphorid species, occurring from the Arctic to temperate regions. The species is economically significant as a cause of myiasis in livestock and valuable in maggot debridement therapy due to its selective consumption of necrotic tissue and antibiotic secretions. Its temperature-dependent development makes it a key forensic indicator for post-mortem interval estimation.
forensic-entomology maggot-therapy myiasis carrion-ecology cold-tolerance Holarctic calliphorid blowfly necrophagous post-mortem-interval wound-strike livestock-pest antibiotic-secretion photoperiodism circadian-rhythm diapause
Pytho americanus
dead log beetle
Pytho americanus is a species of dead log beetle in the family Pythidae. It is native to North America, with records from Canada and the northern United States. The species exhibits notable cold tolerance adaptations, including the ability to overwinter in both larval and adult stages and the synthesis of glycerol during cold acclimation. These physiological traits allow it to persist in climates with severe winter conditions.
cold-tolerance cryoprotectant overwintering dead-wood North-America beetle
Trichonephila clavata
Jorō spider, Joro Spider, Parachute spider
Trichonephila clavata, commonly known as the Jorō spider, is a large orb-weaving spider native to East Asia that has become established as an invasive species in the southeastern United States since approximately 2010. First confirmed in Georgia in 2014, it has expanded rapidly across multiple states through a combination of ballooning dispersal and human-mediated transport. The species is notable for its substantial size, striking coloration, and extensive golden webs, but poses minimal risk to humans due to small fangs and docile behavior. Its physiological adaptations—including higher metabolic rate, faster heart rate, and greater cold tolerance than its congener Trichonephila clavipes—suggest potential for continued northward range expansion.
invasive-species orb-weaver ballooning urban-wildlife biological-control cold-tolerance startle-response cannibalism kleptoparasitism Japanese-folklore Nephilidae Trichonephila spider-silk range-expansion physiological-adaptation urban-ecology agricultural-pest-predator iNaturalist Georgia Maryland East-Asia United-States spider-bite venom docile non-aggressive web-architecture egg-sac overwintering sexual-dimorphism metabolic-rate heart-rate freeze-tolerance road-ecology prey-capture competition native-spider-impacts public-perception media-sensationalism management pesticide-alternatives broom-removal ornamental structural-pest ornithology cardinal web-strength trophic-interactions DNA-metabarcoding diet-analysis species-distribution-model physiological-entomology behavioral-ecology arachnology ballooning-dispersal human-mediated-dispersal hitchhiker cargo-stowaway port-of-entry climate-change warming northward-expansion DMV Howard-County Elkridge Ilchester West-Elkridge beech-forest secondary-forest suburban roadside telephone-pole streetlamp forest-edge disturbance-tolerance prey-detection vibrational-noise auditory-disturbance motionless freezing shyness Hannibal-Lecter stink-bug lanternfly evolutionary-history ancient-association ecosystem-service biocontrol natural-enemy invasive-pest brown-marmorated-stink-bug spotted-lanternfly Halyomorpha-halys Lycorma-delicatula competitive-exclusion dominant-orb-weaver abundant population-growth rapid-spread range-span 120,000-km2 physiological-plasticity metabolism twice-as-high 77%-higher-heart-rate 74%-freeze-survival development-rate seasonal-window life-cycle-completion thermal-limitation congener-comparison natural-dispersal human-transport iNaturalist-record citizen-science confirmed-sighting breeding-population male-and-female gravid-female egg-case stowaway cargo-container port-introduction local-port vehicle-transport hitchhiking accelerated-arrival several-years natural-means prognostication prediction model suitable-habitat abundant-suitable-habitat eastern-North-America western-North-America potential-range invasive-range potential-impact environmental-impact ecological-impact ecosystem-impact monitor cautious evidence-based reasonable-journalism sensationalism exaggeration uncritical-acceptance harmless carefully-monitored next-steps research-opportunity call-for-research knowledge-gap little-known novel-ecosystem spreading rapidly-expanding invasive-predator invasive-orb-weaver non-native introduced established colonized thriving doing-fine happy merely-20-minutes-away road-trip visit access-spread beech-leaf-disease BLD state-forest remarkable-colony amongst-beech-trees confirmed-prognostication Davis-and-Frick escape-relative-warmth expand-range-northward eastern-seaboard merely-20-minutes-away-from-home resist-opportunity amazing-predators tales-of-arrival previous-episodes 2022 2024 reduce-angst large-non-native-spider establishing-in-DMV facts-presented past-episode venomous-terrible-painful Nah expert-Rick-Hoebeke risks-small puny-fangs unlikely-pierce-skin video completely-non-aggressive eastern-Howard-County haphazard-webs littered-with-remains former-victims leaves shed-exoskeletons much-larger-female dwarfs-mate positioned-just-above strand-of-silk spinnerets underside-of-abdomen near-red-mound relative-to-hand large-and-docile wait-and-see what-Jorō-means-to-ecosystems help-other-spiders put-a-beat-down-on-invasive-pests stink-bugs spotted-lanternflies passive-hunters enormous-webs larger-than-a-meter-in-diameter capture-prey-snared-in-silk arachnophobes-scary arachnophiles-beautiful important-ecosystem-services crop-pests native-range Asia having-an-old-friend-over-for-dinner reunite first-time large-spiders juicy-prey-items feathered-and-non-feathered-reptiles poses-no-known-threat difficult-to-predict-impact non-native-species experts-suggest beyond-scary-mien give-indigenous-large-orb-weavers-run-for-their-money black-and-yellow-garden-spider marbled-orb-weaver spotted-orb-weaver other-parts-of-world most-abundant-and-dominant-orb-weaver what-will-it-mean only-time-will-tell final-tidbit Jorō-shapeshifter Jorō-gumo beautiful-woman seduces-men binds-with-silk devours bad-date acknowledgements Rick-Hoebeke identifying-Jorō insights David-Coyle Bob-Bellinger great-images knowledge studies Veni-vidi-vici Nelsen-et-al.Physiological-evaluation Nephila-clavata-newly-recorded Hoebeke-et-al.Life-Cycle-Habitat-and-Variation Moore Startle-responses Davis-and-Anerao How-Urban-Tolerant-Are-They Davis-et-al.inspiration details stars Dave-Clement Miri-Talabac Maddie-Potter hooked-up-with-colony differentiate resources.ipmcenters.org link Jorō-guru David-Coyle's-take YouTube calm-your-fears Journal-of-Economic-Entomology Journal-of-Medical-Entomology chemical-management-strategies effective-management-methods spider-control-products safe-and-effective unlikely-to-bite generally-harmless big-spiders colorful-spiders invasive-spiders in-your-face-spiders webs-on-homes landscape-plants structures middle-of-where-people-live-work-and-play northern-Georgia-2014 presumed-arrived-few-years-prior 15-years increased-dramatically much-of-Southeast disjunct-populations northeastern-U.S.firestorm-of-interest rapidly-expanding-range southeastern-states many-counties Carolinas Tennessee Oklahoma more-than-a-century parts-of-Florida rarely-Pennsylvania tropics thermal-limitations higher-metabolism faster-heart-rate better-ability-to-tolerate-freezing complete-life-cycle-rapidly chilly-temperatures suite-of-adaptations expand-northward warm-loving-cousin findings-support-potential much-yet-to-be-learned successfully-survive-northern-winters nation-and-world-warm southern-species-expand higher-latitudes-and-altitudes rapidly-range-expansion-will-occur typical-mode-of-dispersal aerial-dispersal ballooning-on-strands-of-silk Charlotte's-babies Charlotte's-Web spectacular-moniker parachute-spider rain-down-from-airplanes long-distance-transit human-assistance good-hitchhikers inseminated-gravid-female egg-case-stowaway arrival-in-DMV several-years-by-natural-means new-introduction-at-local-port human-assist-in-vehicles accelerate-arrival terrible-and-painful small-fangs visited-cousin golden-silk-spider Costa-Rica exceeding-meter-diameter indigenous-large-orb-weavers run-for-their-money minimal-direct-impact firestorm Andrew-Davis Benjamin-Frick University-of-Georgia past-week native-to-eastern-Asia Japan China Korea Taiwan known-in-Georgia-since-2013 spreading-rapidly joined-its-cousin Trichonephila-clavipes established-over-160-years better-tolerate-freezing survive-northern-winters rapidly-range-expansion typical-mode long-distance new-introduction human-assist arachnophobes arachnophiles brown-marmorated-stink-bugs difficult-to-predict scary-mien indigenous-orb-weavers Mary-Nouri Sarah-Morgan WAMU relatively-harmless spider-season fall leaves-changing-color cool-crisp-autumn-air pumpkin-spice extension-entomologist Southeast not-just-spiders 20-feet-wide homes people-live-work-and-play 15-years-or-so influx-of-calls public entomology-extension-programs media-frenzy two-questions get-rid-of-Jorō-spiders are-Jorō-spiders-dangerous Clemson-University Southern-Adventist-University Washington-College Union-College two-new-studies test-methods-for-eliminating how-likely-to-bite harm scoured-internet recommended-treatment-options spider-specific-management-recommendations medically-relevant widow-spiders brown-recluses do-not-enter-homes relatively-new recommended-products labelled-spider-control-products tested-products several-products-eliminate not-legal-pesticides machine-lubricant not-recommended commercially-available strongly-encourage labelled-products easiest-and-cheapest without-chemicals broom-or-stick walked-into-spider-web rode-bike took-spider-to-face not-great-feeling wonder-if-bitey give-bitten-superpowers unique-studies evaluated-behaviors react-to-people rather-drop-off-web hang-around-and-bite really-provoke volunteered-to-get-bitten yes-you-read-correctly akin-to-mosquito-bite little-redness little-swelling largely-gone-next-day no-acquired-superpowers take-home-message twofold simplest-and-most-effective without-pesticides feel-you-need-pesticides product-labeled-for-spider-control try-not-to-freak-out doesn't-want-to-be-on-you odds-of-getting-bitten-extremely-low enjoy-glorious-fall-weather drink-pumpkin-spiced-beverage watch-where-you-walk Keep-calm-and-carry-on bites-unlikely cause-minimal-discomfort associate-professor Department-of-Forestry-and-Environmental-Conservation socials email extension featured pest-management Entomology-Today latest-posts email-subscription March-2022 wondered make-its-way-to-DMV September-2022 two-observations three-years 40-sightings happy-and-doing-just-fine several-Howard-County-locations week-or-so-ago team-of-scientists University mission killing-ultra-valuable-beech-trees thriving-amongst-beech-trees visit-amazing-predators mature-females both-Trichonephila-species three-locally-common-orb-weaving-species time-spent-immobile mild-disturbance brief-puff-of-air air-puff-response-data five-other-North-American-species published-literature 453-observations freezing-behavior 10-spider-species remained-immobile-under-a-minute both-Trichonephila-spiders over-an-hour unprecedented shyest-ever-documented tolerate-urban-environments remaining-motionless fleeing nonsexual-cannibalism mating-activity females-cannibalize-males food-availability territorial-aggression Southeastern-United-States expanding-range shy behavioral-reactions descriptive-observations photo-documentation anecdotal-observations controlled-pairings lab natural-webs field one-female-biting-and-killing-another short-fight similar-size container 25-trials fights-ensued-40%different-sizes 27-trials fights-happened-18%larger-females-not-always-aggressor 52-lab-trials six-bouts 9%direct-killing field-trials empty-web 14-trials one-fight 7%aggressor-killing-and-wrapping silk away-from-any-web not-necessarily-territoriality intraspecific-aggression provoked-or-stressed deserves-more-study newly-invasive biology physiology new-range closely-related-species successfully-established 160-years investigation compare online-records iNaturalist.org seasonal-distributions timing females physiological-traits environmental-tolerance metabolic-rates cardiac-functioning cold-exposure survival brief-below-freezing-temperature shorter-season complete-lifecycle narrow-period suitable-weather inherently-higher-metabolism low-temperature survive-better 74%-compared-to-50%brief-freeze colder-climatic-region southeastern-USA useful-information planning urban-landscapes busy-roads frequent-disturbances auditory-and-vibrational physiological-or-behavioral-changes northeast-Georgia varying-levels-of-road-traffic prey-capture-behavior simulated-prey tuning-fork 128-hz-frequency touched-to-web attacked 357-total-trials 20-different-roads attacked-59%high-variability some-roadsides-over-80%others-less-than-30%small-but-significant-negative-correlation daily-road-traffic spider-attack-rates moderate--to-heavy-traffic-roads slightly-less-likely-to-attack 51%-vs-65%able-to-live-near-roads cost-in-terms-of-prey-capture did-not-weigh-less compensate-for-disturbance accumulating-evidence human-dominated-landscapes aid-spread introduced-range Eriostethus-rufus polysphinctine-ectoparasitoid araneid-spiders Neoscona-spp.endemic-to-Japan Yamagata-Prefecture 38º46'-N northern-Japan northernmost-record genus identification-confirmed morphology DNA-barcoding cocoon large-modified-web unique inverted-triangle over-50-cm hanging-from-ill-defined-part no-organized-structure host-spider circumstantial-evidences structure-of-modified-web pre-existing-web-partly-reused orb-web-completely-removed sustaining-threads-newly-moored parasitises-host-spiders two-subfamilies unusual-for-group invasion-potential known-invasive-range review dispersal-abilities potential-impacts management-strategies October-2022 at-least-120,000-km2 South-Carolina North-Carolina Alabama West-Virginia pattern-of-spread natural-dispersal-mechanisms human-mediated-transport large-bodied-orb-weavers flying-insects small-animals thirteen-co-occurring-spider-species monitored-for-competition resources web-building-sites natural-and-urban-habitats management-options-limited advise-journalists-and-experts against-exaggerating-potential-environmental-impact ecologically-harmless rapid-spread-carefully-monitored cautious-evidence-based-approach determining-next-steps three-material-types web-capture introduced-species alter-established-trophic-interactions molecular-analysis resolve-changes community-structure foraging-links east-coast fecal-samples prey-remains-from-webs dissected-guts compare-diet-composition arthropod-targeted-COI-primers Illumina-MiSeq highest-diversity highest-richness highest-proportion-prey-reads recovery-of-prey-reads dissected-gut-content low overwhelmed-by-Jorō-spider-DNA high-proportions-Jorō-spider-reads higher-prey-diversity-and-richness detected-prey-DNA several-days-after-capture initial-gut-retention-time-estimates first-glimpse complexity-of-trophic-associations introduced-web-building-spider viable-material source-of-prey-DNA estimates-of-biodiversity Jammu-and-Kashmir northernmost-shifting document-presence behavior habitat-adaptation colder-temperature-environment Goha-Tehsil Doda-district 1556.74-meters road-bridge nearby-vegetation brightly-colored-abdomens black green yellow red long-legs yellow-and-black-bandings predominantly-seen smaller fewer unique-behaviors remaining-motionless-after-disturbances 5-10-meters-of-roads adaptation-to-human-settlements field-observations image-documentation expert-confirmation thrive-in-colder-conditions 15-20-degrees-Celsius October-November no-spider-found higher-elevations significantly-lower-temperatures adaptation-limited-to-milder-cold-regions new-insight adaptation need-for-further-research non-native-habitats eggs overwinter harsh-weather-conditions no-protection compact-silk-case milky-coating chorionic-microspheres CM fine-structural-characteristics ecological-importance uneven-size-distribution outer-eggmass evenly-covered single-layer diameter-2.3-µm short-papillary-projections segmental-adhesion mucous-components insoluble-in-water partially-soluble-in-absolute-ethanol spherical-structure HFIP strong-organic-solvent not-derived-from-vitellogenic-or-choriogenetic-processes CM-adhesive-coatings ovipositional-process equivalent-to-cocoon-silk protective-functions silken-eggcase Northern-Cardinal Cardinalis-cardinalis perches steals-food southeast-of-the-United-States questions native-fauna-affected consuming-prey-items example native-species-deriving-benefit manner-of-kleptoparasitism perched-directly-on-web supported-its-weight 42-48-g first-documented-case spider-web-supporting-perching-bird measurements force-gauge typical-webs support-masses-up-to-70-g small-but-growing-body-of-knowledge non-native-spider species-distribution-models ecological-niche-modeling predict-areas establish-if-introduced three-distinct-genetic-lineages mitochondrial-COI genome-wide-SNPs Korean-Peninsula