Intraspecific Morphological and Reproductive Trait Variation in Mouth Almighty Glossamia Aprion (Apogonidae) Across Different Flow Environments

Charles Darwin University

Intraspecific morphological and reproductive trait variation in mouth almighty Glossamia aprion (Apogonidae) across different flow environments

Abecia, Janine E.D.; Luiz, Osmar J.; King, Alison J.

Published in: Journal of Fish Biology

DOI: 10.1111/jfb.13821

Published: 01/11/2018

Document Version Peer reviewed version

Link to publication

Citation for published version (APA): Abecia, J. E. D., Luiz, O. J., & King, A. J. (2018). Intraspecific morphological and reproductive trait variation in mouth almighty Glossamia aprion (Apogonidae) across different flow environments. Journal of Fish Biology, 93(5), 961-971. https://doi.org/10.1111/jfb.13821

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Download date: 07. Oct. 2021 varied Significant variationvaried infunctionalacrossintraspecific flowhabitattypes. traitsrelated Five ofthe morphologicalhabitats. out nine andreproductive studiedtraits significantly acrosscontrastingmorphological andreproductive the traits wouldvaryamong individuals collected Australia, northern the mouth almighty (intermittent billabong creek, andriver) investigated intraspecific traitvariation across three contrastingaquaticflowtypes habitat Thisto environmentalimportant study toolunderstanding gradients. population in responses Intraspecific variability, comparably lessstudiedalthough than interspecific isan variation, ABSTRACT Email: [email protected] 0810,University, Darwin,Northern Territory Drive, Australia Ellengowan forJ. E.ABECIA,ResearchEnvironment Institute Charles the Darwin andLivelihoods, Correspondence Australia Ellengowan Darwin,Northern Territory, Drive, Research Institute for theandLivelihoods,CharlesDarwin Environment University, This articleisprotected bycopyright.Allrightsreserved. Accepted Article

Intraspecific morphological and reproductive trait variation and reproductive morphological trait in Intraspecific

pagination andproofreading process, which version undergone full peer review but has not This article hasbeen accepted for publication intheJournalofFishBiologyand at various sites, various with Glossamia aprion Glossamia

and the Versionof Record. Please cite this J. E.ABECIA, O.A. J.KING J.LUIZ, (Apogonidae) acrossflow different (Apogonidae) environments in the catchment. Daly River It was predictedanumber that of in a commonmouth-broodingfreshwater fish in Glossamia aprion

been through the copyediting, typesetting,

may lead to differencesbetweenlead to this may . Samples of article as doi: 10.1111/jfb.13

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. 2018) g Pakkasmaa& ., lightconditions) and factors ( biotic

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size gradients

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fish SchmitzWainwright, 2011 & at

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., ., al

this Lamouroux, food

2001 1995).

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diversity level that

the 1842)

species and and suggest

analysi Glossamia or

gillii and shown

recent

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1991;

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1993)

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and al End varying

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moderate 2004).

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2002

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this

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anck -

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igh

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season contained

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This articleisprotected bycopyright.Allrightsreserved. of creek Acceptedconsiderably whilst and creek investment from vertical inconsistent and across traits Articletraits This 4 samples gonadal billabong dry aprion

L DISCUSSION

period, S therefore swimming

; study

studies revealed of

habitat. and

Pusey

replicate individuals spawning The Glossamia

position G stages where

fish . billabong

suggests aprion there in

proportion

responses

et at

lower

the

intermittent compared

marked

dominated.

al performance a the numbers

and

higher is ., could number

collected are aprion

interspecific 2004). a

compared that males

relative higher

associated

variation to

proportion

intraspecific

with

were

the

of - gonadal

has

occurring creek having This

in proportion is eggs

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with low, mostly different

deep

level. result among

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late that

larger stages

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throughout

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contrast habitat tage

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III the

maxillary

hold) also where

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with

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zero

included body evidence

some body gonads

than male

found

river se

a . the

Five to habitat

higher velocity in ason

Traits lengths

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morphological riverine

lateral foraging

late

collected male environments, line

in in of two

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Figure

riverine

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male

proportion types the that relatively

with higher

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males.

intermittent

nine

and proxy

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revealed 4).

the males reproductive

habitat ,

morphological and

predictions to

female During which

and

large Additionally,

for with

intermittent manoeuvrability

utiliz

significantly

were

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that tage - -

intermittent head use creek

showed

riverine the es

observed.

G I (eye

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This articleisprotected bycopyright.Allrightsreserved. commonly preferences associated Acceptedfield riverine recorded Magnan, revealed studies not this (Webb, thought elongated Articlecaudal those presented Drucker ventral found (Webb, mode morphology where forms

show combination

observations

collected

adapted to of - it

peduncle

position (Aguirre,

1984: 1984;

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This articleisprotected bycopyright.Allrightsreserved. collection and sharing the knowledgeandprotocolsaspartoftheircultural of cultural agreements andsteering committees. been Theyhave involved inproject fieldplanning, data Acceptedongoing collaboration with WagimanOwner Traditional ofsystemof part theDalyRiver the the study tookplace. that acknowledge Wagiman, Wardaman the the andJawoynpeople, traditionalindigenous owners University Animal EthicsCommittee (Permit Number A15003,A1300 (LP150100388).Project was research This project by Charles approved the Darwin we gratefullyacknowledge. O fromassistance particularly ofpeopleWe longlist a acknowledge who during helpedwithfield theproject, assistance ACKNOWLEDGEMENTS Article showing that factors environmental can influence ecological atintraspecifictraits levels. prospective 2012 diffe furtherbe tested validity toexaminethe ofthisnew hypothesis. suchvelocity habi asstructural differentiation inswimming of interplay variables The performance. other withwater previous studies,opposed to was flowhabitat type weakpredictor a ofphenotypic ofattributingtraitdifficulty interspecific level. rentiation ofrentiation newlyrecogniz the , 2017) acr P. for area

Novak, oss a oss a range ofcatchments differentin with D.

Though were there significant differences, findings the reiterate Wedd. ThisprojectfundingWedd. received fromani

future investigation. L.

Sutherland Sutherland and - enviro

. L tat tat pressurecomplexity, predation andfoodavailability could .

was supported by Research Linkagewas supported anAustralian Council nment a to single factorin relationships ecosystems. As ed Glossamia

E. This study a addsto growing bodyofknowledge

Sullivan. WeSullivan. also acknowledge lab

aprion aprion s through a series s through aseries of research candidate species ( - channel flowchannel regimes

We also want to highlight We wantto the also Moreover,morphological nternal nternal CDU which grant, 2). We 2). gratefully also Cook oratory is a et al

This articleisprotected bycopyright.Allrightsreserved. G. Allen, Accepted G.R.Allen, (1991) G.R.Allen, (1989) B.R., A.F.,Alexandre, Ferreira, C. M.,Quintella, Romão, F. W.Aguirre, ArticleREFERENCES funding A.J.K O.J.L. J.E.A CONTRIBUTIONS andwaters. land managingresponsibilities to country and to ., ., ,

Australia. Research Institute, Madang, PapuaNewGuinea. Jersey. Freshwater Fish bocagei Swimmingperformance andecomorphology barbel of Iberian the 8312.2009.01267.x of Journal the Linnean Society shape ideas, ideas, ideas, .

R., S.

E. (2009) -

data data habitat small correlations ina ecologically Alaskan diverse drainage. data

(Steindachner, 1864) onpermanent andtemporary rivers. H. Midgley, &Allen, M.(2002 H. Midgley,

Western Australian Australian Western Museum,Perth.

analysis, gathering gathering . .

Field of GuidetotheFreshwater Fishes New Guinea Freshwater Australia Fishes of

Microgeographical diversificationof threespine

, 244

manuscript

(measurements), and -

258. doi.org/10.1111/eff.12073 field

. collection,

98 preparation , 139 ensure safetravels for onWagiman researchers

data – ) 151. .

manuscript Field Guide FreshwaterField to the Fishes of

analysis, .

. T.F.H. Publications, . T.F.H.Neptune City,New Publications,

doi.org/10.1111/j.1095

manuscript

preparation, A., &Almeida, P.R.(2014)

stickleback: body preparation

Luciobarbus . Christensen original Ecology of -

concept, . Biological

This articleisprotected bycopyright.Allrightsreserved. AcceptedBrinsmead, &Fox,M. J. G.(2002) Bl Bl ArticleBishop, Bishop, K.A.,Pidgeon,W. R. Walden, D. J.,& A., Barnett, & Bellwood, D.R. (2005) Barneche, ake, R. W.ake, R.(2004) anck, A.&Lamouroux,N.(2007) dynamics floodplain tropical ina prerequisites forriver: procedure a to monitor the doi.org/10.1007/s00227 mouth energy 1638. dwelling rock bassandpumpkinseed populations. biology doi:10.1111/j.1365 ofEuropeantraits freshwaterfish. northern scientist/publications/ssr/ecological http://www.environment.gov.au/science/supervising Department Supervising freshwater 9993.1995.tb00524.x impactsmining of

A.,

doi:10.1111/j.1095 R., -

brooding cardinalfishes (Pisces: Apogonidae). output

Allen, 65 -

territory Robertson,

, 1193 fishes

of Scientist .

Fish andswimming functional design performance. increases S.

Environment

A., - of 1222. doi:10.1111/j.0022 . autecology

- Austral Ecology Austral

2699.2006.01654.x

the Pollard,

D. Report

Alligator disproportionately R., - - 005 8649.2002.tb02502.x

. White,

and Morphological variationMorphological lake between . No. D.

0052 Large .

A., Sexual dimorphism buccalcavity the ofpaternal in

Energy. 145. Rivers Journal ofBiogeography

C. - -

& 20 ‐ studies z

scale intraspecific variation inlife R., Supervising Cook, , 81

Region,

J. (1995). StudiesonfishmovementJ. (1995).

Availa & -

- with 107. doi:10.1111/j.1442 -

1112.2004. freshwater Marshall,

ble body

G. Northern Journal of Journal Fish Biology

Scientist,

(2001). at -

size.

Marine Biology D. - 00568.x fishes

Territory: Science

Ecological (2018)

34 Darwin - alligator , 862

Journal of Journal fish .

- 360, - Fish -

. and stream

875.

Autoecology. Canberra,

148 - studies rivers

‐ 642 reproductive history ,

61 205. – - 645. , 1619 region

on -

NT:

the

–

- This articleisprotected bycopyright.Allrightsreserved. Gatz, Accepted N.Franssen, R.(2011) R.&Fisher, Hogan,J.D.(2007) Dumay, P.S.,Tomasini,Tari, J.A., O.,D. &Mouillot, E. Drucker, G., &Lauder, Article T.J.,Drinan, McGinnity,P.,Coughlan, J.P.,Cross, Domenici, P.(2010) Devlaming,V., Grossman, G.,&Chapm Costa

‐ A. Press. andPhysiology A Comparative Biochemistry Pantanalthe wetlands. morphology ofthetetra fish Zoology doi:10.1111/j.1752 divergence inanative stream fish. 2436 o 970 inLanguedocfish species Roussillon,France. southern Biology interpreting patternsin evolutionary finpectoral design. doi:10.1111/j.1600 trutta variabilityMorphological of salmon Atlantic Pereira, R.,Araújo, M.S.,

f pre J.

– (1979) – 983. doi:10.1111/j.1095

- in different in different river environments. 2443. settlement fishes. coral juvenile reef

42 & .

Botany

Ecological 997 doi: .

Fish aneco locomotion: . –

Anthropogenic habitat Anthropogenic habitat alteration induces rapid morphological 10.1242/jeb.004275 1008.

- - 21 G. V. (2002) 4571.2011.00200.x 0633.2012.00561.x ,

91 Journal ofFishBiology morphology doi.org/10.1093/icb/42.5.997 . –

Morphological ofpredictors swimmingacase speed: study 124. Paiva, F., & Tavares,Paiva, F.,& L.E. R. (2016) Astyanax lacustris - 8649.2004.00365.x

Wake andlocomotor dynamics function infishes: an, F.(1982) Evolutionary Evolutionary

Ecology of Ecology of Freshwater Fish freshwater -

ethological perspective Journal Experimental of Biology T. F.,&Harrison,S.(2012) Salmo salar 73 .

89 On the use ofthegonosomatic index. differs between divergent habitats in , 31 (2004) Applications

stream , 1450 -

39. .

Journal FishBiology of

- Functional groups of Functional groups lagoon

Integrative and Comparative 1458. fishes.

and browntrout

4 . , 791 doi:10.1111/jfb.13026

. Boca FL: Raton, Tulane

Functional

21 – , 420 804.

Studies – Salmo .

432.

210

in 64 , CRC

This articleisprotected bycopyright.Allrightsreserved. AcceptedKennard, M. J.,Pusey,B.Olden,J.D., S.J., Stein, J.L.&Marsh,N.(2010) Mackay, Ebert,Kawecki, T.J.& D. (2004) C.,Mondy,Jung, V.,Violle, C.,Hoffmann, S.(2010) Muller, L.& B.&Jonsson,N.Jonsson, (2009) Article T.Johnston, W. A.&Leggett, C.(2002) &Pouilly, Hugueny, B. M. (1999) Male mouthHess, H.C.(1993). jawfishes Constraints in (Opistognathidae): on brooding J. Herler, (2007) West Africanfishes. freshwater polygyny. doi:10.1111/j.1439 Gobiidae) fish(Teleostei: gobiid in the northern Red Sea. 2427.2009.02307.x management. ofClassification flow support natural inAustraliato regimes environmental flow 1225 doi:10.1111/j.1365 and trait 2447. doi:10.1111/j.1095 reference towaterparticular temperature andflow. anadromous Atla 9658(2002)083[1777:MAEGIT]2.0.CO;2 ofaniteroparous fish.size doi:10.1111/j.1095 – 1241. ‐ .

based community assembly. Microhabitats andecomorphology of coral Bulletin of Bulletin Marine Science doi:10.1111/j.1461 Freshwater Biology ntic salmon - - -

0485.2007.00165.x 2745.2010.01687.x 8649.1999.tb02057.x - 8649.2009.02380.x . .

Ecology .

A review ofA review the

Conceptual issues in local Conceptual issuesin adaptation. Morphological correlatesMorphological an ofdietin assemblage of Salmo salar Salmo -

Journal FishBiology of 0248.2004.00684.x .

Maternal andenvironmental gradientsinthe egg 55

83 , 171

52 , 1777

Journal of Journal Ecology

, 806

and brown trout – 193. probable –

– 1791. doi:10.1890/0012 818. doi:10.1111/j.1365

Journal of Fish Biology ofFish Journal

effects ofclimate on change ‐ and coralrock

Marine Ecology Marine

Salmo trutta 98 , 1310 .

Intraspecific variability , 1134 – Ecology Letters 1325. - – 1140. ‐ - , with associated

75 , 82 , 2381 . –

94. 7 , – This articleisprotected bycopyright.Allrightsreserved. Langerhans, R.B.A.M.(2009) &Makowicz, AcceptedLangerhans, R.B. (2009) Langerhans, R.B. (2008) Langerhans, R.B.,C. A.,Shokrollahi, Layman, ArticleLangerhans, R.B.,C. A.,La Layman, Kristjansson, (1993)Kolding, J. F.(2006)Kerfoot, J.R.&Schaefer, Fishes niloticus 9000 species. Evolutionary Biology differentiation betweenpredator regimes in 1057 predator fishes. doi.org/10.1554/04 phenotypicdiversification in driven 8312.2003.00266.x species. Habitat 1112. relation parallel – -

0 doi:10.1002/ece3.235 1075. doi:10.1111/j.1420 B.

Integrative andComparative Biology 37

‐

patterns to ‐ Environmental Fishes of Biology Biological Journal the of LinneanSociety , in Ferguson's Gulf, Lake Kenya. Turkana,

K., associated morphological divergenceassociated morphological fish in twoNeotropical , 25

driven divergence in the .

Population dynamics andlife

Skulason, –

ecology 46.

in . . doi.org/10.1007/BF00000710

Predictability ofphenotypic - diversity Trade

127

of S.,

, 2231 ‐

lava/groundwater

Snorrason, off steady between swimming andunsteady underlies

of ngerhans, A. K. & Dewitt, T.J.(2003)ngerhans, A.K. &Dewitt,

– . small Gambusia affinis

- 2242. doi:10.1111/j.1420 Ecomorphology and habitat utilization of 9101.2009.01716.x Gambusia affinis

benthic

S. .

Shared anduniquefeaturesofmorphological S. A. M.&DeWitt, T.J.(2004) -

76 history styles ofNile tilapia,

& habitats. Gambusia caymanensis Gambusia

, 1

Arctic Noakes, – differentiation across flow regimes in , 750 13. .

doi.org/10.1007/s10641 urnal ofEvolutionaryurnal Biology charr

Environmental Biology of – Ecology , . 768.

689 D. Evolution

- L. ( 9101.2009.01839.x – Salvelinus doi.org/10.1093/icb/icn092 698. doi:10.1111/j.1095

(2012)

Evolution

58 . . , 2305

Fine Journal of

. alpinus

Oreochromis Predator - scale – Cottus

2 2318. ,

- ) 1099

006

- –

22 - ,

This articleisprotected bycopyright.Allrightsreserved. AcceptedMiyazaki,Okuda, N., Yanagisawa, M.& Y.(2002) Tayasu, I., Okuda, N.,& Yanagisawa,Y.(1998) Morrongiello, N.R.,D.A.&Wong, J.R.,Bond, (2012) B. Crook, Article D.L.,Hambleton,S.J. Morgan, H.S.&Beatty, (2002) S.J.,Gill, J.D.McPhail, (1984) Lobon Langerhans, - Cervia, J.,Utrilla,Cervia, C.,Rincon, P.&Amezcua, F.(1997) doi:10.1046/j.1365 trade spatio Eco performance 19 of mouth paternal the capacity. fishwhosereproductive successismouthbrooding limited by buccal fish. andeggsize number reflects trade doi.org/10.1071/MF02047 Australia. likely impacts oftheintroduced redfin( perch Columbia. morphological andgeneticevidencefor species inEnos a pair Lake, British , 801 -

ethological

Journal ofAnimalEcology ‐ R. ‐ offs between eggsizeandnumber. –

temporal fecundity variations the in ofbrown trout B. 807.

Evolutionary Ecology

Marine andFreshwater Research

Canadian Journal ofZoology &

doi.org/10.2108/zsj.19.801 Reznick, .

Ecology and evolution ofsympatricEcology andevolution sticklebacks ( fishes:

Perspective - 2427.1997.00217.x - brooding cardinalfishbrooding

predicting D.

200 (2010)

12 – 81

evolution 248. ‐ , , 806

681 . offs andbet

Ecology

– –

699. . 817. 62

Freshwater BiologyFreshwater Determinate growthin apaternal

with , 1402 .

Apogon doederleini

53 Sexual differencemorphology in buccal

doi.org/10.1023/A:1006533531 Perca fluviatilis and doi:10.1111/j.1365 , 1211 ‐

biomechanics. hedging in a hedging in freshwater

– evolution 1408. – 1221. .

Environmentally induced do . Salmo trutta .

Distribution, biology and i.org/10.1139/z84 Spatial variation inegg of ) (Percidae) ) (Percidae) in Western

swimming Fish . , 277 Zoological Science - Gasterosteus 2656.2012.01961.x

Locomotion: – 288.

L.:

): 201

This articleisprotected bycopyright.Allrightsreserved. AcceptedSmerdon Schmitz, L.&Wainwright, P.C. (2011) Pusey, Pusey, Article M., J. Pouilly, Lino, F.,Bretenoux, G. &Rosales,C.(2003) I.(2001)Plaut, Peres Pakkasmaa,S. &Piironen, J.(2000) Pakkasmaa,S., - Neto, P. R. P.(2004) &Magnan,

B., B. plasticity andmorphological plasticity integration: twoa comparisonof charr polymorphic Ecology jstor.org/stable/23735601 salmonidspecies.locked doi:http://dx.doi.org/10.1016 Australia).River, contributionthe of groundwater regional baseflowto the ofatropical river (Daly doi.org/10.1186/1471 diversityfunctional in the ofreef eyes fishes. Freshwater an Australia. Fish Biology relationships inafishassemblage Bolivian Amazonian floodplain. ofthe Biochemistry andPhysiologyA species. , B. D., Payton Gardner, W., Gardner, S.J. , B.D.,Payton Harrington,G.A.,(2012). &Tickell, Identifying

J., intermittent Kennard,

Kennard, .

Piironen,Ranta, E.& January) J.(1998, Oecologia 14 Critical swimming its relevance. ecological speed:

Collingwood, , 721

Fish

M. 62

river M. , 1137

– , &

730. n/a Journal Hydrology of

J., Arthington, 140

of -

- n/a. Douglas, – 2148 doi.org/10.1023/A:1011691810801

, 36 the 1158.

Vic:

Annales ZoologiciAnnales Fennici

doi:10.1111/e

northern – - 45. 11 . /j.jhydrol.2012.06.058

doi: CSIRO Water velocity shapesWater velocity juvenile salmonids.

M.,

A. 338 . doi.org/10.1007/s00442 131

. The influence ofswimming onphenotypic demand

10.1046/j.1095

Nocturnality constrains morphological and H.

, 41 Australian

(2004) Publishing. Allsop, – , ff.12325

50. 464

doi.org/10.1016/S1095 – Freshwater

BMC Evolutionary Biology Q. 465

wet

. - (2016).

8649.2003.00108.x A morphometric onfour study land , 107 –

dry

. 35

Dietary - , 131

115. tropics.

- Fish Fishes 004

Comparative –

– 140. - assemblage 1562 morphological

Ecology of

North - - 6433(01)00462 y

- of

Evolutionar

eastern Journal of 11 dynamics

, 338.

- in 7 y -

This articleisprotected bycopyright.Allrightsreserved. Accepted C.,Enquist,Violle, B. Villéger, Utne Utne, A.C.W.(1997) Article Biogeography Unmack,J. (2001) of P. Townsend, S.A. A.,Toussaint, Charpin,N., Villéger, Brosse,S.& S.(2016) Kjosnes, Solem,Berg, O.K.& A.J.(2006).Inter O., - Palm, A.C. (2002) widespread. widespread. fish isconcentratedfreshwater in the vulnerabilityNeotropics while functional is 1466 differences wildandfarmedbetween salmon. Atlantic doi.org/10.1016/j.tree.2011.11.014 ecology. (2012) doi.org/10.1007/s00027 of doi.org/10.1080/10236240290025644 behavioural aspects. 50 search time ofthemarine planktivore 28 Freshwater Research intheDaly anoligotrophic(Spirogyra) River, in river tropical Australia.

, 926 , 1053 fish:

Brosse, – 1481. .

– current The ofthereturn variance: intraspecific variability in community

938. doi:10.1111/j.1095 – Trends Ecology in &Evolution & Padovan, A.V.(2005) & Padovan, 1089.

doi:10.1111/j.1095 Scientific reportsScientific S.,

J., B. McGill, approaches

Mouchet, The effect ofturbidity andillumination onthereaction distance and doi:10.1046/j.1365 .

Visual feedingVisual offishaturbid environment: in physicaland

Marine andFreshwater Behaviour andPhysiology

56 , 317 -

017 M.,

and

- –

J., Jiang, L., J., Jiang, C. Albert, J. H., Hulshof, C....&Messier,

Mouillot, 0546 6 327. doi.org/10.1071/MF04079

- future , Australian freshwater fishes freshwater Australian 8649.2006.01208.x - 22125. 8649.1997.tb01619.x

. - -

2699.2001.00615.x The seasonal accrual and loss ofalgae benthic z

Gobiusculus challenges.

D doi:10.1038/srep22125

& , 244

Vanni, -

and intra –

252. Aquatic flavescens

. M.

Global functional diversityGlobal functional of

Journal of FishBiology of Journal

J. - population morphological

(2017) Science .

Journal Biogeography of .

Journal ofJournal FishBiology

Functional 79 , 783

35 Marine andMarine – , 111 801.

ecology

– 128. ,

This articleisprotected bycopyright.Allrightsreserved. Accepted Supporting information online befoundinthe of can version paper. this SUPPORTING Wootton, ArticleWinemiller, K.O.(1991) Wikramanayake, E.D.(1990) Webb,W. (1984) P. Walker, J.A.(1997) assemblage: evolutionofassemblage structure. Zoologist Linnean Society Gasterosteus aculeatus Sons, doi: assemblages from five biotic regions. doi:10.2307/1937583

R. 10.2307/2937046

J. Oxford,

INFORMATION

24 C. , 107

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Body form, and locomotion foragingvertebrates. in aquatic Ecological morphology ofstickleback lacustrine threespine UK.

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Ecomorphological (2015).

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doi.org/10.1093/icb/24.1.107 L. Ecomorphology

(Gasterosteidae) body(Gasterosteidae) shape.

Reproductive

Ecological Monographs diversification diversification fishin freshwater lowland

and biogeographyof atropicalstream fish

Biology - 8312.1997.tb01777.x

Ecology

Teleost

Biological JournalBiological the of 71 , 1756

Fishes.

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This articleisprotected bycopyright.Allrightsreserved. Accepted Article used Table Intermittent Intermittent

Habitat 2

Billabong Billabong Location of River River River River

type

creek creek

Glossamia aprion Upstream Sawmill Shillinghole Oolloo Galloping Ferguson

Edith Site (Jagard) Claravale

Oolloo #2 Crossing Name

River

Billabong

Jacks River

Billabong sampling sites sampling sites inthe catchmentDaly River (Figure1)

crossing

14° 14° 14° 14° 14° 14° 14° 14° Latitude

07 32 21 21 20 11 04 04 ′ ′ ′ ′ ′ ′ ′ ′

31.56 59.26 43.66 32.21 25.43 04.16 16.58 23.20

(S) ′′ ′′ ′′ ′′ ′′ ′′ ′′ ′′

131° 131° 131° 131° 131° 132° 131° 131° Longitude

17 07 33 33 34 02 58 15 ′ ′ ′ ′ ′ ′ ′ ′

12.89 36.27 25.01 07.21 33.18 01.78 37.27 14.70

(E) ′′ ′′ ′′ ′′ ′′ ′′ ′′ ′′

This articleisprotected bycopyright.Allrightsreserved. Accepted Article asthepredictortype variable Table Pectoral Relative Body Relative B B River Billabong Billabong River Billabong Billabong River Billabong Billabong R

lateral 3 iver illabong illabong

Pair Morphological eye maxillary fin – – – – creek creek creek creek

ventral shape size - wise one – – – – – – – – river river river river creek creek creek creek

length

position

- Trait

way ANOVAof (male)

Estimate – – – – – – 0.040 0.037 0.032 0.062 0.030 0.010 0.031 0.041 0.092 0.052 0.039 0.075 morphological traits

0.011 0.015 0.031 0.022 0.031 0.028 0.020 0.029 0.015 0.011 0.015 0.015 SE

of t – – – – – – –

1.813 1.322 2.160 5.684 1.951 0.646 Glossamia aprion 2.864 2.676 2.941 1.678 1.895 2.644 value

P < 0.01 <0.001 < < > 0.05 > 0.05 > > < > > <0.01 –

value 0.01 0.01 0.05 0.05 0.05 0.05 0.05

with habitat This articleisprotected bycopyright.Allrightsreserved. Accepted Article across Table

4 all three NA, Non available in samplesavailable NA, in Non

M F M emale S ean ex ale

gonado flow Ma habitat - somatic ( index turity IV IV IV III III V II II I I

types stage

I G I g 0.13 0.80 0.18 0.06 0.00 9.44 1.50 0.46 0.04 ) NA (

of %)

male andfemale

0.05 0.07 0.03 0.00 5.86 0.60 0.21 0.08 NA NA S D

Glossamia 4.86 0.11 0.03 0.00 0.89 0.22 0.00 0.0

aprion Range NA NA – – – – – – – – 19.08 0.21 2.61 0.90 0.18 0.31 0.10 0.00

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