Posted on Authorea 26 Jul 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162729215.55251866/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. al costetocnet.Ti okdmntae,frtefis ie h rsneo ihnpopulation within of presence the time, first the for repea- demonstrates, were undisturbed two, work (i) for This except contexts: contexts. measured, different two traits two the behavioural in All across territory. twice, of table deteriorated measured understanding partially were (ii) damselfly our and behaviours improve the territory The to in repeatability. needed traits, their is behavioural studies culated nine tested instance, More For studied species literature. developed. We and the splendens, be generalities. measured in to and traits underrepresented need patterns of highly and trait-correlation variety over currently studied wider research are poorly a invertebrates ecological still including on are within focused areas acknowledgement some studies and However, field interest decades. two increasing past received the has personality Animal EnvironmentalAbstract and 0000-0003-1086-7567. Fish ORCID Wildlife, Ume˚a, [email protected]; of 83 Sweden; Department 901 Sciences, Agricultural Studies, of University Swedish Brodin; Tomas author Corresponding Ume˚a, Sweden 2 1 Golab – J. wild Maria highly the under in wild the personality the in in insect damselfly personality Adult adult testing an for in species differences model personality promising a population This as within contexts. conditions. splendens of two environmental Calopteryx the presence controlled propose across the further repeatable (ii) time, and We were and first boldness, two, territory wild. for (iii) the undisturbed except (i) and for measured, contexts: aggressiveness demonstrates, traits different (ii) work two behavioural courtship, in All twice, (i) the territory. measured in deteriorated experiments: were traits, partially behaviours behavioural three The nine of are studied tested repeatability. array We invertebrates species their an on and generalities. calculated measured from and focused traits patterns studies splendens, of trait-correlation field variety Calopteryx of instance, wider damselfly understanding For a our including improve developed. studies to be More decades. needed to two is literature. need past the and the in studied underrepresented over highly research poorly currently ecological still within are acknowledgement areas and some interest However, increasing received has personality Animal Abstract 2021 26, July 2 1 Go a Maria as splendens studies Calopteryx field – for wild model the in personality insect Adult wds nvriyo giutrlSine maCampus Umea - Sciences Agricultural of University PAS Swedish Conservation Nature of Institute nttt fNtr osrain oihAaeyo cecs Krak´ow, Poland Sciences, of Academy Polish Conservation, Nature of Institute wds nvriyo giutrlSine,Dprmn fWllf,Fs n niomna Studies, Environmental and Fish Wildlife, of Department Sciences, Agricultural of University Swedish rma ra ftreeprmns i orsi,(i grsieesad(i)blns,adcal- and boldness, (iii) and aggressiveness (ii) courtship, (i) experiments: three of array an from l a˛b 1 zmnSniegula Szymon , 1 .Sniegula S. , 1 .Anto A. , 1 nre Anto Andrzej , aotrxsplendens Calopteryx l 1 .Brodin T. , 1 l 1 n oa Brodin Tomas and , 2 samdlfrfil studies field for model a as 2 Calopteryx Posted on Authorea 26 Jul 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162729215.55251866/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. net’sxa n oilbhvor ersn ra ait fpeoeata r aeo betin absent because or is rare This are that 2018). phenomena al., of et variety (Keiser great research a insects behavioural represent years animal behaviours recent in vertebrates in social on However, role and al., studies important et to 2013). sexual compared (Larsen an Logue, insects’ when Earth and play scarce on Mather still to animals 2014; are started of taxa Schuett, group Kralj-Fiˇser this and numerous on 2001; studies most (Gosling, personality the 2018), represent Stork, invertebrates 2017; that of fact consequences the evolutionary Osborn Despite 2010; and Braithwaite, and ecological (Archard mice the field 2017). populations natural natural Briffa, striped understanding in in and or for 2012) measured above Weissing, applicability 2012) and organism The (Wolf high field. Quinn, model personality and a animal of and laboratory in be (Cole the personality may in tits of both assessment conditions great consistently (2017) well-designed (Niemel¨a on behaved that Briffa itself individuals studies denote assessments. translocation that and examples hand, the showed personality by Osborn 2016) other influence biased al., the anemones, be et might (Yuen On therefore sea environment can 2017). on experiments laboratory Dingemanse, transplant example to and of recent the kind field between this a from correlation from In transition no Results was 2015). the there al., that but conditions, et showed zebra on field (McCowan Study natural and 2015b). For situations and laboratory al., environments. two laboratory in et two (Fisher in both only the both personality conditions showed activity in artificial finches between and in estimates repeatable correlations exploration personality was of any shyness lab-based similar However, repeatability find conditions. showed compare provide to crickets to others fail on designed whereas some studies Studies example, results, behaviour studies. validate different field field to show and important corresponding personality is lab with of it lab assessments lab the the field-based in in and patterns findings not behavioural of irrelevant ways relevance ecologically the patterns in for potential Archard behavioural behave this irrelevant may by of ecologically Because conditions (reviewed showing laboratory mates hence in potential and 2014). individuals environment of affected Dingemanse, (Niemel¨a consequence, natural are and pool laboratory a the often the reduced for As experiments of representative and lab-based homogeneity 2010). learning, since selection trapping, Braithwaite, unfortunate relaxed selective and is stress, and skew captivity This artificial e.g. environment, constraints: 2020). Fisher of 2013; al., number Maestripieri, and et a studies Carere by Hertel few 2010; Braithwaite, animals, 2015b; and captive-bred organism (Archard al., and study wild captive et the given on in a experiments personality for on laboratory focused of applicable have number 2013). be large al., to the et order metrics with (Carter Finally, Compared in phenomena information caution 2014). relevant the with (Koski, ecologically of chosen traits represent be understanding measured to sociability, should in-depth rarely and aggressiveness, traits and bring personality (boldness, commonly may the between personality that of associations drasti- animal traits 2014). possible is of Schuett, other the Kralj-Fiˇser personality and Five” ignoring and (e.g. lab-based invertebrate “Big activity), traits to on the behavioural and compared studies and exploration concerns studies of species studies of amount field currently most number the of research Further, the personality underrepresentation Also, to animal a 2010). disproportionate Repea- of Braithwaite, is cally individuality, 2004). aspects and there of several al., (Archard instance, consistency However, et For of tests 2016). Sih estimate criticism. al., 2017; standardized intense et a 2017, face (Roche provides al., and that personality et interactions metric is, Lichtenstein social informative that 2010; rates, highly Weiss, a speciation and is change, distribu- environmental tability Briffa geographic to (e.g. species response consequences important use, invasive, have space fitness studies be as: Personality to 2016). aspects tendencies al., such et tions, determining Roche implications (e.g. has evolutionary decades context and two and past ecological time the across in behaviour interest in growing differences received consistent inter-individual as defined personality, Animal propose conditions. Introduction environmental further controlled We highly under wild. wild words: the the Key in in personality testing damselfly for adult species model an promising in differences personality esnlt,fil xeiet,suisi h id aotrxslnes repeatability splendens, Calopteryx wild, the in studies experiments, field personality, 2 aotrxsplendens Calopteryx sa as Posted on Authorea 26 Jul 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162729215.55251866/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. nbd oorto,wt lerflcigdr igsosi h ideo h ig fmls hc satrait a is which males, of wings the of middle the in spots wing dark reflecting rese- blue with ecological 2006). colouration, Dijkstra, and 1988; body (Askew, in evolutionary Europe is in rivers for Odonata lowland within inhabiting systems ly families model studied intensively prime most (C´ordoba-Aguilar, as the 2008) of considered One (C´ordoba-Aguilar are 2008). arch damselflies and Dragonflies species Study different two territory. in a assessed of Methods was deterioration repeatability partial The after nature. in and 2013). syndromes territories) al., the that (males’ et in indicate study patches results (Carter first contexts Our original the system undisturbed and is study on time this this contexts: Since over to boldness. repeatability applicable (iii) their and are aggressiveness for (ii) tested behaviour, on courtship traits (i) behavioural axes: of behavioural lab-populations set existing a replace damselfly or report re-stock we 2013). as Ankeny, the Here, well and expand as both Leonelli to 2009; organisms, need al., model we 2020). how et traits of al., was behavioural (Behringer et of breadth (Vossen including zebrafish taxonomic unaffected study studies wild was of and recent zebrafish relevance of number lab-reared ecological a behaviour the the while the increase example, anxiolytic, to that For the Hence, the showed to when 2010). exposure in behaviour conditions the Braithwaite, novel zebrafish resulted by to changed and affect likely reaction can (Archard behavioural than pharmaceuticals its populations more anxiolytic in homogeneous, natural change have in a generations to This and many environments compared laboratories. for organisms, stable reared to model biology been adaptation significant molecular has species most 2001), of the Sokolowski, of environments 2006; one Roberts, non-natural, instance, 2012; For al., research. et 2013), be of Ankeny, (Kain organisms not ( may lines and of and some fruitfly Leonelli limitations for group their the problems, have models larger decades, scientific as several specific a over useful lab-conditions very on representing under research studied species and been integrative bred has non-human and intensively as comparative (i.e., 2013), of for al., organisms tests et used multiple model (Carter Also, trait usually established given 2004). which a Already al., passerine, describing is et a in which validity ( (Whittingham for areas, higher guppies habitats adequate open of on closed be in be done more could not boldness) in trait, may of species personality upon, 2003), proxy different one preyed two a Daniel, are in (as and traits and predator (Blumstein different al., inhabits, a flies kangaroos et measure to fruit e.g. (Sih conspecifics’ actually behaviours response in for may a mating testing sources used test/metric and of instance, 2019) same food presence For al., the at in et 2014). that aggregation (Brodin (Koski, hiding possible mosquitofish 2011), include: received in is (Koski, These shoal It have chimpanzees to behaviours. 2014). tendency in is traits of 2018), ‘sociability‘ grooming range al., behavioural term 2008), et whole these The (Scott al., a instance, but et for For (Cote proxy evolution, 2014). smell (Koski, a and studies as ecology personality used animal being species mating in the Naguib, to in attention related and became little behaviours Oers role example, relatively Van which For extraordinary 2007; aspects. sociability) al., personality an R´eale and other et 2013; of have activity Logue, understanding limited and exploration, have (Mather we aggressiveness, studies 2013), (boldness, personality animal five’ for big blueprint 2015b). 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All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162729215.55251866/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. mnil)o h eietml prahstepr fteopnn’ oyta s‘utbefrbiting’. be for simply may ‘suitable direc- head the body is of the the that pro- depending of when chance, parts trait body by some situation this arise opponent’s Also, may that movements. a the intruder damselflies suggest the of requires chasing biting we the part 1990) behaviour of studies, Waage, the tion/intensity/frequency and biting earlier approaches (Marden The and contest male aerial data measures. resident is In our he the personality on males of for of based (mandible) unsuitable type intruder, three is an the biting bably situation. of to the which (Koskim¨aki et to evaluate regard intensity adequately to varying With react have of and males conflict interaction Resident a every approach 2013). initiate territory, for Opaev, given and facing a and approach pass Panov or either non-territorial 2009; may immediately Also, al., males retreat non-territorial and 1976). activity resident Parker, this and During the and (Briffa mates. 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We repeatable In was repeatable. hits” significantly of “number were Disscussion et only boldness (Chaput-Bardy aggression and species to courtship the connected to for traits density the related intermediate traits ca. an 1996). behavioural was is Stettmer, All which procedure, 2010; river, al., mark-release the et daily of Kuitunen standardised section 2010; a m al., 1 on per based individual assessed 1 density, population bootstrap estimated the coefficient. set The repeatability we for models intervals rpt confidence Re- In replication estimate one 1. properly 2017). and Results in to 0 al., replication between order residual value et between in and takes level Stoffel 1000 correlates coefficient group The to trait 2016; of number individual). Team, particular sum case the how Core our over Development (in about variance unit level information (R nine group the of the v.4.0.2. gives ratio for variance as R coefficient calculated in (R) repeatability ‘rptR coefficient quantifying package peatability by the personality) using (i.e., traits consistency behavioural assessed We analyses 2004). Statistical al., et Sih 2005; Reale, and ae hwacaatrsi adrn eaiu,ta s arllre etoso ie looking river a of sections larger patrol is, that behaviour, wandering characteristic a show males 5 .splendens C. .splendens C. esrdin measured C. is Posted on Authorea 26 Jul 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162729215.55251866/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. 99 iaJtye l,19,19;Sv-oh n opr 95 n rdto GlbadSigl,pers. Sniegula, and (Golab Hilfert-R¨uppell, predation 1997; and Ruppell, 1995) and Hooper, and Hilfert Siva-Jothy 2011; 1995; Del-Claro, 1995, (Gib- macrophytes al., and environmental of temperature, et Guillermo-Ferreira Siva-Jothy structure air experiments 1992; and 1999; penetration, our composition Pain, (sunlight velocity), In and hetero- microclimate temperature 2009). environmental for: bons (current, al., quality standardized controlling water were et is speed), many Quinn field wind and R´eale, 2013; the natural and in were Dingemanse personality factors 2004; animal (Bell, studying geneity when difficulty – common (Golab Braithwaite, One observations and the (Archard procedure during the damselflies reduced to Hilfert-R¨uppell, adjusting/habituating they of 1999). from lab- 2010; since behaviour damselflies behaviour to prevented natural/free damselfly and compared influence obs.) adult methods pers. relevance, studying not the for specifically, did ecological promising More and trauma, in particularly damselflies. (Archard handling are calopterygid superior, study stressors adult are this (Niemel¨a including unnatural in studies animals factors presented additional most field environmental impose on by that experiments can modified conclude based conditions there significantly we since artificial be 2010), However, and experiments. can Braithwaite, lab-based 2014) expression predicts with Dingemanse, gene always field compare that and not crickets not evidence captivity on did (Niemel¨a growing study we in and estimates another study is repeatability personality with our bias line that may In experiments in 2017). and translocation is Dingemanse, in 2015a) This procedure 2015b). al., handling al., that et et showing consistency (Fisher (Fisher behavioural nature individual lifetimes In in that crickets. found adult personality They on laboratory. over researchers conducted the steady in of been and group has is field a the wild (https://www.wildcrickets.org/) 2015). in the Crickets” both (Strong, in “Wild personality season research studied on project mating personality large-scale based advanced of and most is interesting day the an group a of hand, and mating one as ratio other enter/visit insects behaviours sex to the Among mating-related group propensity on in the whose personality as But reindeers, no such profile. consistency showing subdominant factors behavioural studies than proximate on are rather beneficial research there plasticity example a Hence, behavioural for of 2010). offspring favour 2004). al., partner the adaptive would et be environments al., and sexual (Dingemanse would unpredictable 2007) et a traits al., or (Dall choose behavioural R´eale variable et what mates and 2013; highly assess al., offspring potential can et her for partner) (Korsten (choosing predicts for costs heritable that female cognitive theory are predictable, male with reduces is traits accordance in environment personality it in instance that all since for is given shown This behaviours two Also, 2007). been the mating good Magurran, also addition, a and in reacting has In potentially Magellan consistency such, displays when 2016; 1999). as al., mating but (Corbet, and, et away to repeatable, consistent (Biro chase engagement also guppies not were than Consistent was engagement personality. rather which and for attract 2). display intruder, (Tab. metric to dive an consistency wants courtship: to individual to male showed related reaction resident traits female the the approaching female to an 2013). a similar to al., to appear react the et to may identify (Carter needed to trait species male crucial The given resident be (Sih a a might trait in axis time personality trait behavioural The a targeted given escalated estimating the a above During for for for The away. test test proxies observ.). chased right multiple suitable pers. is developing the most Golab and intruder choosing 2005; 2008) an of al., in (R¨uppell Bell, et which importance occur and aggression the after can of illustrates flights hitting results or proxy pursuit collision discussed brief better some are fights a disputes aerial be longer the to of most seemed for but hits useful not of are number hence Generally, and the all contrast, them In of opponents combination in a their bites calopterygids. or compute conclude of in factors, we to number studies physical above, and able or personality the trajectory opponent environmental be Summarizing flight the social, would another. to compute either one odonates reaction on can bite that for precisely and time evidence to 2012) one the order no have al., that in is dragonflies et fighting that there during (Bybee fact movements 2012) the insects body (Olberg, despite among Additionally, prey systems area. their wing visual of a advanced of most centre the the of instance fo Calopteryx p.mlscmeefrrsucsdrn eilcnet Mre n ag,1990), Waage, and (Marden contests aerial during resources for compete males spp. 6 .splendens C. r lsi ris depending traits, plastic are .splendens C. rlu campestris Gryllus ntewild. the in Posted on Authorea 26 Jul 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162729215.55251866/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. el .. aksn .. akwk,KL,20.Terpaaiiyo eaiu:amt-nlss Animal stickleback meta-analysis. a behaviour: of of repeatability populations The https://doi.org/10.1016/j.anbehav.2008.12.022 2009. two 771–783. K.L., 77, https://doi.org/10.1111/j.1420- Laskowski, Behaviour and S.J., 464–473. Hankison, A.M., individuals 18, Bell, Biology between Evolutionary differences of 9101.2004.00817.x Journal Behavioural aculeatus). 2004. Manual. (Gasterosteus 871–878. Laboratory A.M., A 68, wall: Organisms: Sociobiol Bell, the Model Ecol Emerging Press. on 2009. Laboratory Behav (Eds.), mirror Harbor fish. R. Spring Mirror, Krumlauf, in Cold A., 2014. aggression Johnson, R., J.G., measuring Behringer, Frommen, for tests F., mirror Koch, of S., https://doi.org/10.1007/s00265-014-1698-7 value Wanner, predictive M., Colchester.the Taborsky, Books, V., Harley Europe. England. Balzarini, of Essex, dragonflies Books, The Harley 1988. ed. R.R., 2nd tempera- Askew, Europe, animal of of Dragonflies studies 2004. in R.R., https://doi.org/10.1111/j.1469-7998.2010.00714.x populations Askew, 149–160. wild 281, of Zoology importance of The Journal 2010. ment. V.A., Braithwaite, by G.A., supported Archard, was TB Sciences. of Academy were References Polish AA Umea. Conservation, and Sciences, Nature SS Agricultural MJG, of of 2014/15/N/NZ8/00338). University Institute no. Swedish by (grant part Centre in Science National TB, supported by SS, supported MJG, was writing MJG Manuscript AA. SS, MJG, Acknowledgements analysis: and collection Data SS. AA. MJG, design: Project interest. of conflict Contribution no Author have they that declare authors The Dryad. in archived be Interests will Competing data be the not manuscript, hence the aiming accepting should when Upon selection and trait consistent proper behaviour. not of individual accessibility were in need Data differences aggressiveness the behaviours of emphasises to studied implications This environmental related in ecological experiments. they key exists understand traits or addition, of to that tests the In variability manipulations personality of eye. natural relevant for Two naked the and useful represents the species. controlled study only this enables our using which of that of swiftly fidelity suggest repertoire in measured site We elaborate and damselfly strong parameters. an observed adult a has easily have an species be also The in can conditions. repeatability natural adult that behavioural relevant behaviours that ecologically demonstrate 2005). under suggest that al., personality results R¨uppell first et insect 2004; Our the (Corbet, wild. is behaviours work courtship the of our repertoire (Golab summarize, elaborated These males To non-territorial 2014). very of (Koski, a gathering studies and 2017), personality al., 2013) future et al., Golab for 2004; et useful (Corbet, potentially patrolling be territory include: could traits that 2018), al., personality et animal Five”(Keiser that in lacking emphasise be we can often Here that conditions. results environmental our varying to to robustness a due adds field-studies This methods). in details obs., Calopteryx p.epesohreooial motn eaius eodte“Big the beyond behaviours, important ecologically other express spp. .splendens C. 7 savr rmsn oe raimfrstudying for organism model promising very a is Posted on Authorea 26 Jul 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162729215.55251866/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. aito? n nmlProaiis eair hsooy n vlto.Uiest fCiaoPes pp. Press, Chicago of University Evolution. personality and animal Physiology, maintaining Behavior, selection 201–220. Personalities: natural for Animal evidence in: the variation?, is https://doi.org/10.1016/j.tree.2009.07.013 What persona- 81–89. 2013. R´eale, animal D., 25, N.J., norms: Dingemanse, Evol. reaction Ecol. Behavioural Trends 2010. plasticity. J., Publishing, individual Wright, Wildlife D., meets British Reale, lity A.J.N., Europe. Kazem, and N.J., Britain Dingemanse, of Dragonflies the to Guide Gillingham. Field 2006. K.-D.B., Dijkstra, https://doi.org/10.1111/j.1461- indi- 734–739. 2851–2858. consistent 7, 275, personality: Letters of Sci Ecology ecology perspective. Biol behavioural 0248.2004.00618.x adaptive The Proc an 2004. from fitness. 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Nida a Posted on Authorea 26 Jul 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162729215.55251866/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. odesDsac oratBnma itnevle er- of Near traits value: behavioural distance of Binomial (R) Repeatability 2. Table. react to Distance resident focal until [s] Time intruder to Reaction Boldness Aggressiveness Description resident focal until [s] Time female to Reaction Traits Courtship Experiment iet eunTm s asdutlthe until passed [s] Time accuracy an with [m; Distance resident a times many How return to Time resident a times many How distance Escape Hits Bites male a describing value nominal A a of alighting of Number Engagement display Dive .splendens C. 13 nBia in l iariver. Nida a oeta .maway 0.5m was than decoy more bird the male when – Far escaped 0.5m than decoy closer bird was the when escaped male male intruder presented the toward moved female a toward moved oebc a 8 s 180 was to back male come the for waited the observer time maximum The attack. predatory the after territory his to returned resident attack simulation predatory the male after resident flew focal the 0.5m] of intruder the hit male intruder the bit male trial experimental the during perching aside, flights no chasing, patrolling, no 25%: perches; aside, flies intruders, away chases territory, perch patrols not male does 50%: aside, fly not does intruders, away chases a territory, patrols male 75%: experiment); the during not flying (does stop perch not does period), experimental the his during leave territory not (does aside not fly does intruders, territory, away a chases patrols water, tandem, on a dives form to attempts male 100%: courtship: to devotion in display courtship common (a water surface the on male resident Calopteryx sp.) Posted on Authorea 26 Jul 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.162729215.55251866/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. iue2 xeieti h ed – A (B). field. ballast the a in using Experiment reduction area 2. patch Figure experimental Sequential (A). females Bia by substrate in site ( Study vegetation 1. Figure legends Figure aegudt sigline. fishing a to glued male oaoeo natans Potamogeton ie .6 .0 ,0.347 0, 0.364 0.109 0.282 CI 95% 0.160 0.338 0.396 return to Time 0.332 0, P distance Escape react to Distance 0.492 0.055 0.338 R Hits 0.167 Bites intruder to Reaction 0.396 Engagement display Dive female to Reaction Trait l iarvricuigslce eeainpths i ace ffloating of patches Six patches. vegetation selected including river Nida a aiuae oeulsz,ue strioisb ae n oviposition and males by territories as used size, equal to manipulated ) .splendens C. 14 eaegudt sigln;B–Flying – B line; fishing a to glued female 0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 .9,0.440 0.098, 0.491 0.161, 0.553 0.174, 0.541 0.113, 0.624 0.339, 0.465 0.184, 0.553 0.174, .splendens C.