damaged by pig activities. In contrast, with the exception oxygendamaged still pig activities.contrast, is dissolved (which In by Accepted highest because bypigs the theygrazed had exposure areshallowand risks, heavily werewetlands thatfish typologies data examined.revealed in wetland water-quality Field for freshwaterfish asphyxia exposure three tropicaloccupying Acute thermalrisks and Australian Hub) andthe Australian Government. wasThis study Environment withNational supported Sciences Programme (Northern Funding information [email protected] Email: Article Freshwater Ecology ResearchGroup, James University, Queensland, Cook 4811,Australia Waltham, for EcosystemN. J.Centre Tropical WaterandAquatic Research, Correspondence Significancestatement

Fr Thermal and asphyxia exposure risk to to freshwater exposurerisk asphyxia and fish Thermal pagination andproofreading process, which version undergone full peer review but has not This article hasbeen accepted for publication inthe Journal ofFishBiologyand eshwater Ecology Research Group,eshwater JamesEcology University, Queensland, Cook 4811,Australia

Centre forCentre Tropical Water andAquatic Ecosystem Research (TropWATER), and the Versionof Record. Please cite this This article isprotected by copyright. All rights reserved. N. J. WALTHAM AND J. R. SCHAFFER WALTHAMN. J. J. R.SCHAFFER AND

been through the copyediting, typesetting, wetlands wetlands

may lead to differencesbetweenlead to this may

article as doi: 10.1111 /jfb.13 74 in feral-pig-damaged tropical feral-pig-damaged 2 oxygen, freshwater fish,dissolved floodplain, pigs, thermal tolerance, wetland KEYWORDS heavily pig reached conditionsofcritical vegetationrespiration), because aquatic This articleis protectedbycopyright. Allrightsreserved. Accepted Article - managed wetlands pig - modifi

ed

tropical tropical provide landscape

s

the the opportunitybest .

for thesame

fish deeper permanent species species to survive i

and n a which Schaffer, 2017) vegetation wetlands therefore are (Doupé sediments activity northern and again habitats orderephemeral in aquatic and unpubl. data chronic most notably become re 2014; Pettit evaporation, groundwateror recharge, through consumption 2003) Arthington, disconnect from andp rivers primary 2016; Hurd element D This articleis protectedbycopyright. Allrightsreserved. Accepted Article uration and frequency uration floodplainof connection coastal tract tract away from banksand the

deeper and have verylimited avoidance

freshwater fish freshwater Here

thresholds for local aquatic fauna forlocalaquatic thresholds et al.et of predisposed to predisposed in

Queensland, where human intervention less

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, 2010;Waltham &Schaffer, 2017) . three typologieswerewetland experience affected

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reduced waterqualityconditions water quality water . However, manymonths after . Intropical regions high feral animalsferal production season, water water temperature

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prevent for those remainingfor those fauna aquatic

rogressively

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where they are confineare (Wallace Wetlandmust fish s examined modified

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(Pusey Arthington,2003) & wetland habitatsinnorthernwetland s ;

Kozak, 2016; Galib Kozak, 2016;Galib loss ofvegetation, exposureloss and (Waltham permanent become , 2012) et al., et et al.et d to boundsthe wetland oftheisolated are

[ acute effects temperature in the Archerin the catchment River in Fig. 1

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et al.et acute and acute , ] at at but , . et al.et The ,

, from the tub in a were generally deeper( had November 2017 4m 1mm wingpanels, sunrise. C) shortly after temperatures amplitude 9.67º middle afternoon. Themean ofthe temperature amplitude daily was6.26º almo periodicity; of these types occupying wetlands falling & Fixler,2017) waterin environmental analysed here)pH arenot oxygendissolved (%), electricalconductivity and wetlands wetlands. exceedance frequencies bycollecting achieved wetlands second This articleis protectedbycopyright. Allrightsreserved. Accepted Article no submerged or floating no submergedor aquatic plants st linear rate for period rate a st linear of

stage ofth Calibrated Calibrated Fish was to c. , among the Arc (0.2 90

1 were collected her River.her Fish

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during during Australia northern temperature recordedat data and thresholds that higher than water rangethe ofwatertemperatures recorded species, there wasnosignificant relationship between fishand size lineolata (Richardson 1844) maximum AET Hypseleotris compressa A2239. permit number Queensland Animal Protection Careand Act2001 then from tank experimental the andpl the is wetlands temperature alinearat (2.5 rate (Thermoline, Eurotherm 316Control unpubl.Butler, data water This articleis protectedbycopyright. Allrightsreserved. Accepted Article an assumed

temperature therefore wouldneed toseek fish returned to the samereturned tothe wetland the day. next - column average (whichcolumn average suitable is forhyperthermic experiments; daylight hours The Contextualising

probably

(Steindachner 1867), (Waltham &Schaffer were e AET

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determined for each fish species determined foreachfishspecies (Tab

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Butler, D.Burrows & R.Butler, unpubl. Pearson, data wateroxygenated surface theheatduring oftheday (whenathermal ismost refuge needed) of behaviour regulatory elsewhere towarmwaters oxygen (Waltham critically lowoxygenexperience dissolved concentrations,often 10%saturation below al. et bottomthat the contains strata sufficient bottom for fencing permanentthat or deep vegetation with wetlands rocks,as under fallen or timber highwatertemperatureregulate by shadedaccessing refugia submergedmacrophytes wetlands nature to rapid watertemperature fluctuations al. et off arefoundhere in temperatures do This articleis protectedbycopyright. Allrightsreserved. Accepted Article

-

not experience maximumnot experience fish channel wetlands wetlands thatbecomechannel connected during seasonally wetseason rainfall , 2007) , 2017)

and

In addition tomacrophytes, In addition floating e waters waters (Kelleher (Kelleher in tropical in tropical river catchments .

et al. et (Elliott, 2000) Fenced w smaller . . did reach 43did reach The latter proviso isnotThe latter alwaysmet, isnotsurprising that theephemeralIt wetlands of , 2013) thermally stratified most et al. et thermalmass compared when etlands ; , 2012) river -

nevertheless, because the because (butpoorlycool aerated) bottom stratum forms surface waters rich , in addition , inaddition vegetation,to riparian . o each each Diel stratificationDiel

C s are , whichis water water throughout .

night thus preventingthedevelopment of Research elsewhere inta (Jones &Bergey . temperatures high, that ct, maintain more depth and ha

fresh water vertical migrationvertical

high enough to and

dissolved oxygentomeet needs respiratory northern Queensland acute hyperthermia has been can vidence ) .

s ,

provide

with When 2007; Larson provide provide

as illustrations

an effective thermoregulatory behaviour

microhabitat areas river edges,such along from deep

the permthe more open,deeper, be stressful

deep waters between oxygen cool

ideal conditions ideal conditions to offers thermal refugia

although northern northern

waters in that affected

(Pusey (Pusey

periods, a al. et

provide that that n important thermal safeguard

and ve for

Australia Australia the the become tropical ,

freshwater species abundant

et al.et 2009) by feral pigs fish then

daily maximum given shade oxygen oxygen . for fish

, 2017)

mixes withthe The fish species The fish species , whichsuggests support support this kind stably wetlands - shows the that

depleted deep the and

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(Balcombe provided provided cooling

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only only (Wallace

and field

( B.

concentrations below declining unpubl. data fish species failureRegulatory andpotential asphyxiation occurs about the at 30% in most sensitive local successfully exploitwaterswith reduction year of acute threshold. the wetland in order tobreathe. thehere, exposure In washighest wetlands risk 2013) d hours show for thepurposeshere Butler &, D. tolerancesoflocalhypoxia high temperature dissolvedreduces oxygen solubility thelateparticularly dryseasonduring wetlands composition andclarity of water the characteristics, the stratification tooccur iscontingent on numerous site too th This articleis protectedbycopyright. Allrightsreserved. Accepted Article e bottom can waters hypoxic hypoxic utili to be

and n ; hencelower the the

for uring day.Regulating the br The solubility ofdissolved [

thereby Fig.

suggest dissolved oxygen in growth rate the three wetlandsthe (Fig.

and ) Burrows, unpubl. data Burrows, unpubl. .

2

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long that water depth of ] , where chronicis the threshold size of andaspect wetland the ,

- thresholds thresholds

z

the chronic(62%)andacute (24%)threshold limits for term remain for unaerated ed as While s

and fecundity

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oxygen inwater 2) ) eathing and (Wallace

< refugia for refugia aerobic at thisat timeof year the and

when tropical conditions extremewhen tropical are 1

(Flint

cover m i w indicate indicate hile tolerancehile var species

the numberthe ofspecies affected increases with prolonged periods results inincreased ventilation rate s Somef not et al. et et al. a few days invertebrates , degree the of andthe wind exposure adequate adequate to

that concentrations significantlyconcentrations lowerthanthat. exceed

in northern Queensland , 2018) , 2017) the greaterthe amount the expended ofenergy ish in the survive wild ish in exposu regular is -

specific variables strongly (Diaz & Breitburg, 2009)

this

organisms , ed

. N

. it reflects it f

ish must regulate breathing ish must breathing regulate

ies The could 98% ofthelogging time

sustain dielsustain stratification

evertheless, evertheless, many , ( affected bytemperature (

B. Butler B. Butler &, D.Burrows, increas between species andlifebetween stages, se

. result result in deficits energy data fordata

in the

The depth requiredfor The the the periodhottest ofthe ing such asflow

the riskthe ofbecoming pig

is available pig .

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modified . . (Collins

fish

compressa compressa Data species species

and

,

on (

many B. i.e. et al. et

re to re 48%

and

are or ,

Corporation animal ethicspermit This research ACKNOWLEDGEMENTS wetlands) toexamineis necessary climate change resilience on reduced habitat aquatic in habitat management intervention isincreasingly necessarytoensure they remain and productive viable day. manyeach hours probably contributes continue thermal e A low oxygenavailability and commence surface respiration. riskpredation probablyincreases aswetlands become indeed,asfishrespondto or shallower, duringsurface heat the of day the to exposure ( it thatNotably appears associatedmost of mortality withhypoxia the inhibited safely depends onthe gulping[ air This articleis protectedbycopyright. Allrightsreserved. Accepted Article ustralia

(Creighton Efforts to control Efforts control to thermal risks andhypoxic .

xposure to fish, to xposure similarto has outright

and particularly whether they provide T to reduced hese datasupporthese

e.g. and

e.g secondary secondary complied we advocate here

. thermal andsunburn) resulting stress from animals’ the need tarpon staff and rangers fromstaff andrangers theAPNCape York(AakPuulNgantam) Southern

termsmanagementof priority by et al.et number A the the ofpresence

timing theoxygensagandantecedent of aquatic

The cultural and ofcoastal ecologicalvalue wetlands means that

Megalops cyprinoidesMegalops , 2015) with the consequences, effects of

a model that habitat, through . Here 2239 that thereturn for

Queensland Animal Protection Careand Act2001

to access to access available oxygenfor respiration

for fish f permanent wetlands . eral pigs eral dense floatingdense mats weed , We Balkanu the CapeYorkDevelopment thank

permanent are wetlands including

damage wetlands to from pigactivities without the

a similar of level refugia

on coastal wetlandson coastal

(Broussonet 1782)](Broussonet

loss of

(followed bymanaged wetlands) water temperaturewater

the

permanent wetlands financial

aquatic aquatic vegetation expensive , means that , means that these

(Waltham& Fixler,2017) investment conditions probably are underwayare fenc .

Their Their to capacity and asphyxiation - (James induced fish kills inducedis due fishkills e

costs

the most critical communities,

, (and managed(and interventions ,

though maybe in terms reducing of .

. et al. et In addition, Fur to remain the at in northern in northern given the ther research , 2017) not only

and

do that risks f

but also should should JCU

. .

or

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2018 10.1038/s41561 h physiology ) 2010 , . Iscurrent floodplain

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. Efficacy of

Marine and

) acclimated at

Pardo, I., Pařil,Pardo, I.,

- 27 squaretail 948. - - , 1 018 Erskine, C.,

- ., Hwan, 23. - 0134

Sus - 4

Pettit, N.,Jardine,Pettit, T.,Hamilton, D.,Dav S.,Sinnamon, V.,Valdez, D.,McJannet, Marvanek,S.,Kinsey E.R.,Larson, C.,Wagener,Kelleher, T.,Gooseff,M., McGlynn, S.N. Jones, James, A.E., J.,Pearson, Reside, VanDerWal, C.S., R.G.,D.,Capon,S.J., Burrows, Hurd, L.E.,Sousa, Siqueira R.G., Winter, K.,DeGórski, H.,Khoruzhaya, Leeuw,J., V.,Boldyrev, This articleis protectedbycopyright. Allrightsreserved. Accepted Article Processes Investigating on controls thermal the sensitivity streams.of Pennsylvania conservation. 501. turnover in rivers Australian under climate change. T.D., L. Hodgson, a rapidly deteriorating environment. ( semi Nagelkerke, L. ( cattle ontropical floodplaincattle waterholes. Bunn, S. ( 1144. and potential impacts ofclimate change. Persistence of in crayfish.invasive different displacement: tolerances to stream dryi 2016

- & Bergey, E. & Bergey, segregation A.(2007).Habitat instream crayfish natural large floodplain (Volga,large natural Russian Federation).

) . Amazon floodplainfishcommunities: connectivityandconservation Habitat in Magoulick, C., D.D., Turner,

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- Freshwater BiologyFreshwater , 785. & ) Waltham, N.J.( . Theimportance offlooded terrestrial fishin habitats forlarval a

- Souza, G F.K.,Cooper, - Henderson, Henderson, A., Petheram, C., Biological Conservation & 2017

54 Laycock, K.H.and Disturbance (2009). species Marine and Research Freshwater Marine ResearchMarine and Freshwater , 1899 )

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& 195 Wallace, J.(2014). Marshall, L.( & , 118 , es: implicationses: for

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& 63 , 105 65

Australia Australia - Harwood, 144. , 788 44 2012 , 1131 , 489 - & 110.

- ) 800. . - -

Waltham, S. &Fixler, N. ( Waltham,Wallace, Burrows,D., N., J.,Brod Wallace, J., Waltham, N.J Wallace H.A.,Stewart, Pusey, B.J.,Burrows,D.W., Perna, Kennard,M.J., C.N., Unmack, Pusey, B.J., This articleis protectedbycopyright. Allrightsreserved. Accepted Article Gilbert RiverGilbert Catchment.Atechnical to report the Australianthe Governmentfrom to responses changesin flow andd Freshwater Science of dry Researchand Freshwater waterholes season in the Flinders and catchmentsAustralia. Gilbert innorthern andhybridcatfish.channel M.P. ( 16. and management fish: a review. offreshwater Floodplain Floodplain Wetland. ( erhole logy_in_the_Flinders_and_Gilbert_catchments/links/5405bab30cf2bba34c1d7e38/Wat www.researchgate.net/profile/Nathan_Waltham/publication/260599789_Waterhole_eco Sustainable A Queensland AgricultureIrrigated Strategy andGilbertAgriculturalResourceAssessment,CSIRO ofthe Flinders part North , J., Waltham, N., Eichhornia crassipes

& - - 2017 season waterholesintropical Australia: northern potential effects on fish refugia. ecology Arthington, A.H., & Allen, P.J.( ) . Australia. Freshwater fishes ofnorthern griculture 460pp. Australia, griculture flagships, - in - ., Burrows,D.W., the & 2017

34 Tropical Conservation Science Burrows, D.( - 2014 ): ): Water QualityConcerns Fish Tropical Fronting Occupyinga Flinders , 663 ) . Aerial H (

68 2003 ) North American JournalAquacultureNorth American of . Critical thermalmaxim - 678. , 650 - and ) . Importance of the riparian . Importance oftheriparian zone totheconservation

- 667. erbicide toControl Spray InvasiveWater Hyacinth elivery of pollutant of elivery loadsin River Flinders and - 2017 Gilbert & ie, J.,ie, Thomas, C.,

McJannet, D.( McJannet, ) . Acomparison temperature of regimes dry in . - catchments.pdf CS Marine and IROWater aHealthy for Countryand

a of two geographic strains oftwo geographic a of Available at Available 10 Zootaxa 2015 , & 1 - 10. Butler, B.( Freshwater Research

) P.J., Allsop, Q., Allsop, P.J., . Thetemperature regim

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54

, 1 es - - Supporting information befoundinthe can SUPPORTING INFORMATION Waltham, &Schaffer, J.( N. Waltham, &Schaffer, J.( N. This articleis protectedbycopyright. Allrightsreserved. Accepted Article ent tlands_with_and_without_feral_pig_and_cattle_fence_exclusion_Archer_River_catchm /www.researchgate.net/publication/320079038_Continuing_aquatic_assessment_of_we NumberReport 17/04,James University, Cook feralwithout pigandcattle exclusion, Archerfence River catchment ds_identified_for_fer www.researchgate.net/publication/303667623_Baseline_aquatic_assessment_of_wetlan James University, Australia Cook exclusion, Archerpig fence River catchment

al_pig_fence_exclusion_Archer_River_catchment 2015 2017 ) ) . .

Continuing assessmentContinuing aquatic ofwetlands with and aquatic assessmentBaseline o .

Available at online of version paper. this .

TropWATER Number 15/41, Report Australia .

Available at f wetlands identified forferal

. TropWATER

This articleis protectedbycopyright. Allrightsreserved. Typesetter after (▬) fenced (c) FIG Typesetter www.ecogrow.ca WalesSouth Museum ( time water the thresholds. above effects temperature(AET) types temperature time present pig access access, permanent wetland ( FIG ofFiguresList

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dominant in (2007). f ( y requency distributionrequency logged Oxyeleotris lineolata

- that axis to read Timeaxis toread AET(%). above ) , without submerged, without vegetation, aquatic turbid and eutrophic; is ….. mogumda

, deeper (typically <

of fre etc , permanent; Melanotaenia s. s. Melanotaenia inornata

(d) to (b) (a), shwater fish present inwetlandsshwater illustrate percentage to of Fish imagesofB.Australia Western courtesy Hutchins, the the Archer ,

), Department Government ofPrimary Industry, ofNew

2m clear with deep), submergedaquaticvegetation ); www.fishesofaustralia.net.au ).

- etc oneach panel

: pig(a) modified,

-

- , fenced

River catchment. (

2m deep) , 12

,

– 22 November 2017 . Horizontal showlines minimum

) and ) and 5% effect concentrations ( and (a) pig .

steep side steep shallowwetlands that are ,

(c) - .

modified, (b) permanentmodified,(b) and e

fenced wetland )

Surface (0.2m) water ( Melanotaenia Melanotaenia inomata d, which in three the wetland

limit

that s

prevent ,

pig (b)

acute acute -

-

- ); )

, s

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 5 4 3

Delete DissolvedoxygenfromDelete (%) (c). and (a) On eachx Change Date/time toread (c) time Dateand below (hours). - axis 0:000000. change to

Fig 1. This articleis protectedbycopyright. Allrightsreserved. Accepted Article

This articleis protectedbycopyright. Allrightsreserved. Fig 2. AcceptedC) ArticleB) A)

14/11/2017 0:00 14/11/2017 19/11/2017 0:00 19/11/2017 14/11/2017 0:00 14/11/2017 Dissolved oxygen (%) Dissolved oxygen (%) Dissolved oxygen (%) 100 120 100 120 100 120 20 40 60 80 20 40 60 80 20 40 60 80 0 0 0 15/11/2017 0:00 15/11/2017 15/11/2017 0:00 15/11/2017 20/11/2017 0:00 20/11/2017 16/11/2017 0:00 16/11/2017 Date/time Date/time Date/time 16/11/2017 0:00 16/11/2017 21/11/2017 0:00 21/11/2017 17/11/2017 0:00 17/11/2017 17/11/2017 0:00 17/11/2017 22/11/2017 0:00 22/11/2017 18/11/2017 0:00 18/11/2017

This articleis protectedbycopyright. Allrightsreserved. Accepted Porochilus rendahli Oxyeleotris Pseudomugil tenellus Melanotaenia s. inornata Melanotaenia nigrans Ambassis macleayi Article Hypseleotris compressa Iriatherina werneri Mogurnda mogurnda wherefor eachfishspecies, possible. TABLE 1 Scientific name Scientific

lineolata Thermal

acute acute

effec B N lack bandedlack rainbowfish T E orthern trout gudgeon hreadfin rainbow fish astern rainbow fish D t R E

temperature elicate blue eye endahl’s catfish mpire gudgeon Common name Common G S lass perch leepy cod

(

AET

)

results results Count 10 18 13 2 8 6 6 9 1

for

fish length, range length, Standard Standard 28 75 60 30 33 48 20 40 (mm tested in this tested inthis study 19 – – – – – – – -

90 85 40 40 80 35 51 40

)

A 38.3 37.7 37.7 39.1 37.8 36.8 37.5 ET – –

10

38.65 38.05 38.75 A 38.5 38.5 39.3 38.4 38.3

with the ET –

50

39.72 39.18 39.33 A 38.7 39.4 38.7 38.4 ET – –

AET

90

1 Highest Observed Observed Highest 0th, 50th and90th0th, 50th 38.7 39.9 39.6 38.7 39.4 39.6 39.1 38.3 38.5

A ET

percentile s calculated calculated