Regional Differences in the Feeding of the Ambush Predator Neosebastes Pandus and Comparisons of Diets in the Scorpaenidae, Triglidae and Platycephalidae

Home , Bellator (fish), Cape gurnard, Grey gurnard, Helicolenus hilgendorfii, Neosebastes, Platycephalidae

RESEARCH REPOSITORY

This is the author’s final version of the work, as accepted for publication following peer review but without the publisher’s layout or pagination. The definitive version is available at:

https://doi.org/10.1111/jfb.13677

Greenwell, C.N., Coulson, P.G., Tweedley, J.R. and Loneragan, N.R. (2018) Regional differences in the feeding of the ambush predator Neosebastes pandus and comparisons of diets in the Scorpaenidae, Triglidae and Platycephalidae. Journal of Fish Biology

http://researchrepository.murdoch.edu.au/id/eprint/41646/

[email protected] ‡ Accepted Article Butler Institute uhr to Author 1 Regional differencesin thefeeding of M School of VeterinarySchool of and Sciences Life

pagination andproofreading process, which version undergone full peer review but has not This article hasbeen accepted for publication intheJournalofFishBiology and estern Australia Murdoch

C. N. GREENWELL comparisons dietsin of the Scorpaenidae, Triglidaeand University and the Versionof Record. Please cite this hm orsodne hud e addressed be should correspondence whom , , RUNNINGHEAD:

M School ofVeterinary andLife estern Australia estern ,

, 90 SouthStreet

6150 ,

Australia 1,2

, P.G.COULSON This articleisprotected bycopyright.Allrightsreserved. ,

6150 DIETS OFDIETS LONERAGAN ,

, Murdoch

2 Centre for Sustainable AquaticCentre for Ecosystems ,

Australia , been through the copyediting, typesetting, the

Murdoch University Sciences ambush predator

SOME SOME ,

M may lead to differencesbetweenlead to this may 1,2

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and , J. R. TWEEDLEY, J.R. 1,2,3 SCORPAENIFORMES article as doi: 10.1111/jfb. , 3 ‡

Asia Research Centre Murdoch University

Tel.: Tel.: ,

90 SouthStreet

Neosebastes 6 8 30 6453; 9360 8 +61 ,

6150 Platycephalidae 1,2 ,

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90 South Street ,

pandus Murdoch Murdoch , 13677

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This articleisprotected bycopyright.Allrightsreserved. Accepted Australia.western words: Key most distinct andfeeding the being platycephalids with families, among detected were diet in differences Significant platycephalid Meta prey. mobile more larger, ingest and capture to fish bigger allow that changes morphological reflecting size, Articlehabitat west of diets to important more being teleosts with diet, to contributors overall largest and the were brachyurans teleosts Cephalopods, coast. south the from fish than diet their in diverse more coast west the from identified. contents stomach the of contribution volumetric and frequency percentage south of (Esperance) coasts south and Island) (Rottnest west (1 Platycephalidae and species) predatory benthic predator provides study This

between the two locations. two the between Neosebastes Neosebastes

unr perch; gurnard - coa eeld ht hy ed rdmnnl o tlot ad ag crustaceans. large and teleosts on predominantly feed they that revealed s

families

fish t consumed a

pandus comprehensive assessment of the dietary composition of the of composition dietary the of assessment comprehensive

and more scorpaenids onteleosts than andtriglids. -

nlss of analysis in the Scor the in

aia structure; habitat and oyhee fr south for polychaetes a greater mean number of broad taxonomic broad of number mean greater a 0

species). compares the diets of diets the compares

Dietary composition also changed with increasing body body increasing with changed also composition Dietary

paeniformes h des of diets the A total of 275 of total A

cranie; size Scorpaenoidei;

: Scorpaenidae ( Scorpaenidae : - os fish. coast 49 49 49 pce o sopei, rgi and triglid scorpaenid, of species N. species from species

pandus hs elcs ifrne in differences reflects This - western Australia and the the and Australia western 2 -

0 species), Triglidae (19 Triglidae species), 0 eae cags south changes; related were collected were

39 groups and were were and groups

studies of studies from the the from ambush ambush three three Fish Fish -

This articleisprotected bycopyright.Allrightsreserved. al. et Platycephalidae and Triglidae Scorpaenidae, the of members many for information Acceptedal. is yet pr behavio cryptic camouflage, on relies that behaviour Motomura 1998; Potter & Bra 2008; leaf and ridges spiny cirri, decorative colouration, cryptic fromranging 10to200m Vincent St of Articleis It of species 12 the of al. south the in found typically species) 18 genera, (two Scorpaenidae (Nelson 80 125, with Scorpaeniformes the of 32% up make rockfishes) (searobins (Nelson species 2092 and genera 398 families, 41 suborders, six comprising order diverse a are Scorpaeniformes The ey , ,

2015; French

2016). The bighead gurnard perch gurnard bighead The 2016). endemic to southern Australia (Houtman Abrolhos Islands, Abrolhos (Houtman Australia southern to endemic , (Curio 1976; Keenleyside 1976; (Curio

highly 2007; Consoli 2007; Owing scorpaenoid Many t al. et y, 2016) y,

or effective at catching prey with precision with prey catching at effective

, unrs, ltcpaia (flatheads) Platycephalidae gurnards),

c. o hi eooia ad cnmc motne tee s ex is there importance, economic and ecological their to 2016). The Neosebastinae (gurnard perches) are a small subfamily of the the of subfamily small a are perches) (gurnard Neosebastinae The 2016). et al.et 5° . To capture their prey, these fish rely largely on ambush predation ambush on largely rely fish these prey, their capture To .

Neosebastes et al. et S ; ,

2017) 138° , s (Hutchins &Swainston,(Hutchins 1986;

2010; Horn 2010; et al. et are well are t al. et

. ) ocrig n ok res n mcoht aes t depths at areas macrophyte and reefs rocky on occurring E),

1979; Gerking 1994) Gerking 1979;

Guichenot 1867 Guichenot ,

2006; Consoli 2006; 06. nopsig rud 6 seis te Triglidae the species, 660 around Encompassing 2016).

c INTRODUCTION amouflaged and often characteriz often and amouflaged et al. et Neosebastes pandus Neosebastes ,

2012; Coulson 2012; et al. et

species, reaching lengths of up to 500 mm. 500 to up of lengths reaching species, . u

This feeding strategy uses little energy, little uses strategy feeding This (Schultz & Kruschel, 2010; Coulson 2010; Kruschel, & (Schultz capture to strikes selective rapid, and r ,

and 2010)

Edgar, 2008; Edgar, 2008; - crande (scorpion Scorpaenidae (Richardson, 1842) (Richardson, et al. et appendages like , a specialized form of feeding feeding of form specialized a ,

and c. ,

2015). While these studies studies these While 2015). 29° 454 species, respectively species, 454 - west Pacific (Nelson Pacific west Gomon

ed by a wide range of range wide a by ed S ;

114° tensive dietary dietary tensive (Gomon et al. et

is the largest largest the is ,

to the Gulf the to

, (Boudaya

fishes 2008 (Platell t al. et )

. or et et

This articleisprotected bycopyright.Allrightsreserved. of diet the to prey differ Acceptedal. but molluscs, by dominated is Australia of coast west the on example, diet in differences in result coast south the of those than warmer appreciably thus are and Ocean Indian the in located are coast west the spp impermeable kelp the 1995) Hyndes, & Ayvazian 1989; communities fish their influence significantly which habitat, and geomorphology Article al. et to crustaceans mainly 2001) Potter, gastropod scorpaenoides scabriceps varies diet how investigate al. et (Brodeur fishes these of diets the to significantly crustaceans and teleosts that shown have ,

00. T 2010). are found along the south coast south the along found are , ,

between coasts between 2011; Baeck 2001; different of contributions dietary volumetric the quantify to firstly aimed study This south of coasts south and west The Ecklonia radiata Ecklonia s

Whitley 1935 Whitley and tel and , . he

Demirham

rtrzi granite Proterozoic As some sebastid and scorpaenid species increase in size, their diets shift from from shift diets their size, in increase species scorpaenid and sebastid some As

uceo 1867 Guichenot (Coulson diet of the the of diet eosts constituting the remaining portions ( portions remaining the constituting eosts Neosebastes et al. et

(French primarily ,

& 2013). ate ate

et al. et are a conspicuous feature of the west coast west the of feature conspicuous a are ary among a 20) no 2009), Can labrid mainly large crustaceans, while 50% of the diet of diet the of 50% while crustaceans, large mainly et al.et

composition among conspecifics among composition ,

on the south coast south the on

pandus fish

2010). Differing habitat and environmental conditions environmental and habitat Differing 2010). . oins frenchii Bodianus , opie lre rsaen, ih ml crustaceans, small with crustaceans, large comprised ef structures reef families. Of the two studies of neosebastids of studies two the Of families. Extensive limestone reefs covered with dense growths of growths dense with covered reefs limestone Extensive

2012). ( Harmelin (Ayvazian & Hyndes, 1995) Hyndes, & (Ayvazian , ,

obtained from marine waters off the west and south and west the off waters marine from obtained

the diet of diet the (

e.g. comprehensive - etr Asrla xii mjr ifrne in differences major exhibit Australia western

- euphausiids, mysids and decapods) contribute decapods) and mysids euphausiids, Viven & Bouchon, 1976; Bouchon, & Viven & , oiae b te macroalgae the by dominated

crustaceans are more prevalent more are crustaceans Pearcy 1984; Chess 1984; Pearcy Pseudocaranx georgianus Pseudocaranx

Kuzne 1879) (Klunzinger comparisons have been made to to made been have comparisons Bulman from . In addition, waters along along waters addition, In .

different locations different et al. et , while , et al. et , Murie, 1995; Murie, os not does ,

2001; Platell & Platell 2001; ,

less extensive less 1988; (Cuvier 1833) (Cuvier ,

Maxillicosta Maxillicosta Neosebastes Neosebastes Sargassum

(Platell (Platell (Howard, (Howard, however,

Bulman

Neves

. may may

For et ,

This articleisprotected bycopyright.Allrightsreserved. N.B of coast Australia; Western (south Esperance off operating trawlers scallop commercial from obtained also Table autumn; Acceptedand summer (Austral 2013 and spring) early and winter (Austral 2012 September and July between Australia; ( scallop COLLECTION OF flows groups Article feeding functional identifying for knowledge fundamental comp Platycephalidae and species other on information dietary published with consistent th whether investigate size increasing with items prey mobile more of composition dietary the greater the to due available types habitats of prey complexity of range diverse more a contain will coast west the from hypotheses: following the tested we range, size large a over and south of coasts .

that very little, or no, commercial trawling occurs between October and January due to to due January and October between occurs trawling commercial no, or little, very that sto of osition for trophodynamic mode for trophodynamic of Samples msu balloti Amusium 32.0°S - western Australia. Australia. western N. ;

115.5° E), 115.5° pandus pandus 33.9° S 33.9° NEOSEBASTES PANDUS ag ( 20 m oa length total mm 230 (> large and de of diet e if ) trawlers in waters off Rottnest Island (west coast of Western Western of coast (west Island Rottnest off waters in trawlers ) ; N. diet

n te meta the and 121.9° E 121.9°

MATERIALS ANDMETHODS pandus were collected from fish processors or purchased from a market purchasedfroma or fromprocessors fish were collected l ary composition ary ling ofling the and N. Since this ambush predator was collected fr collected was predator ambush this Since

pandus ),

will undergo a undergo will therefore the dietary compositions will differ; will compositions dietary the therefore between May and September of 2013, 2014 and 2015. and 2014 2013, of September and May between - nlss f pce i te he fmle provides families three the in species of analysis marine ecosystems ofsouth I

). Samples of Samples ). n h mrn wtr o Wsen utai is Australia Western of waters marine the in

differs . Another ,

size L T among families among ) - related N.

N. pandus N. main pandus in the in

change aim of this study was to to was study this of aim first, cuh b commercial by caught ,

(148 Scorpaenidae, Triglidae Triglidae Scorpaenidae, - and western western .

the diet of diet the

This study of dietary dietary of study This shifting to larger and larger to shifting in – siaig trophic estimating 427 mm 427 January and May and January om two coasts coasts two om Australia. N. pandus N. L second, second, T ) were were )

This articleisprotected bycopyright.Allrightsreserved. (Table categories dietary 12 form to combined were II (Table components percentage a as expressed and Acceptedrecorded and Industries Regional Development, speci fish from otoliths of collection reference extensive an using family to prey the identify to used were otoliths their of morphology the of combination a digestion, (2004) (1988) Bryce to often level, taxonomic species possible lowest the to identified were stomach each in items and100% full fullydistended(Platell Articleof equivalent the being 10 with 10, to 1 of scale a on visually estimated was fullness Stomach QUANTIFICATIONOF on caught are scallops of numbers min cod the in mm 45 and wings the in 40 to 16 of depths at occurs mostly trawling (Kangas fishery scallop the of closure seasonal the bare substratumbare to adjacent (Jones (Jones The , , occurrence The

Edgar (2008) Edgar sn mcocpc n mcocpceaiain ad h ecitos in descriptions the and examinations microscopic and macroscopic using as L et al. et T percentage frequency percentage

, of each each of Hutchins & Swainston (1986) Swainston & Hutchins ,

2010)

). For graphical and analytical purposes, these 33 dietary components components dietary 33 these purposes, analytical and graphical For ). N. pandus N. and of each of the 52 prey items found in the stomachs of of stomachs the in found items prey 52 the of each of s n south in es . NEOSEBASTES PANDUS Scallop trawlers operate mostly at at mostly operate trawlers Scallop Gomon reef andreef rocky

volume (Laurenson was measured to the nearest mm and its stomach removed. stomach its and mm nearest the to measured was ( et al. et - % - end that that end etr Asrla ( Australia western F et al. ) Western unpubl. Australia, ( , % and

(2008) V ,

) et al. et 1998; Lek is , the

m, using a trawl net with a m a with net trawl a using m, by order or family, subsequently te subsequently family, or order by Norman & Reid (2000) Reid & Norman habitats.

towed at a speed of speed a at towed . When teleost prey had undergone extensive extensive undergone had prey teleost When . volume ,

1993; Sporer 1993; et al. et

DIETARY remains II

et al.et s M. ). Note that seagrass (Posidoniaceae) seagrass that Note ).

, of the items the of

2011). On both coasts, commercial coasts, both On 2011). Dowling ,

night, 2011; Coulson and the characteristics of their their of characteristics the and et al. et COMPOSITION

data

c. since this is when greater when is this since ,

, 6.5 km 6.5

eatet of Department we 201 Miskelly (2002) Miskelly ).

esh size of 50 mm 50 of size esh ) estimated visually visually estimated . Trawling .

et al. et h – N. 1

for 60 to 180 to 60 for rmed dietary rmed

pandus

2015). The 2015). The el & Wells Primary Primary , occurs Poore was

This articleisprotected bycopyright.Allrightsreserved. ( sizes sample small < fish from samples that mm) Acceptedrespectively of effects main the Anderson three a to These dominant. excessively be to use then were data transformed components dietary those for tendency any avoid down to of composition dietary the in variation STATISTICAL ANALYSES OF Articlepresented clearly be not could they that small and calculated histograms. were as coasts plotted south and west lower the from classes length mm 50 in fish of and volumedietary total the of 1% only up made which material, discarded is (Harmelin was it as selected not was four ,

Season (4 levels; spring, summer, autumn and winter) and autumn summer, spring, levels; (4 Season N. P mean the trends, dietary determine To - reliminary weight the contributions of dietary categories with consistently high values and to to and values high consistently with categories dietary of contributions the weight .

- pandus et al et - Vivien way crossed crossed way ), by commercial fishers commercial by Length

., 2008 .,

from coast. south the et al. et

Location (2 levels; west and south coast south and west levels; (2 Location multivariate n ) to determine whether dietary composition differed significantly differed composition dietary whether determine to )

ls ( lvl; 150 levels; (5 Class ≤ , N.B

1989) 2 150 mm and > and mm 150 emttoa mliait aayi o variance of analysis multivariate permutational ; . probably probably

Table Table ht h cnrbtos f cioemt and Echinodermata of contributions the that . Likewise, scallop roe, which was found in only two fish and fish two only in found was which roe, scallop Likewise, . d to construct a Bray a construct to d

NEOSEBASTES analyses were aimed at exploring the major sources of of sources major the exploring at aimed were analyses I ). ,

ingested as an unintentional consequence of prey capture capture prey of consequence unintentional an as ingested was not included in the analyses, analyses, the in included not was

hs EMNV test PERMANOVA This ersne i te histograms the in represented N.

400 mm 400

pandus – 9, 200 199, % V . The . L

PANDUS

T contributions of the dietary categories for for categories dietary the of contributions - were not included in this analysis due to due analysis this in included not were Curtis resemblance matrix and subjected subjected and matrix resemblance Curtis – 4, 250 249, % V

data were square were data

DIETARYCOMPOSITION and

Rottnest Island Rottnest

– demonstrated that significant significant that demonstrated 9, 300 299, their was

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and Esperance, and

terms

ee so were . Note Note . – with with

399 not ;

This articleisprotected bycopyright.Allrightsreserved. centroid a create to used then was matrix resemblance Location the using matrix centroids among distance a construct to used was which Acceptedmatrix, resemblance Curtis and Gorley 2015). using class length replica of number unequal an in result of numbers unequal from obtained data guts, individual dietary pooled by created bias potential any classes and length locations amongdiffered fish square and averaged was replicates of group (Platell collected Articlefish of number the on depending five, to three of groups into sorted randomly were location each from fish of stomachs effect, potential this reduce To diet. in trends re but subtle mask can same which composition, dietary the their in from markedly differ time may location, same the at collected species, same the of samples two categories, LOCATION ANDSIZE RELATED DIFFERENCES INDIET from analyses. further three or two the of ( differences s h soah o idvda fs my oti ol a ml nme o te dietary the of number small a only contain may fish individual of stomachs the As analysis of similarity ( similarity of analysis all for data averaged The P obntos sc ublne saitcl ein cn e nlsd effectively analysed be can designs statistical unbalanced such combinations,

< i.e.

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- , ×

way interaction terms interaction way 01 a species accumulation effect (Lek effect accumulation species a

Length Class factor Class Length

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2001). The 2001). were detected were ANOSIM oain ad length and locations % V for

) contribution of the dif the of contribution

e ( tes and PERMANOVA (Anderson PERMANOVA and (Anderson l ( ocation P i.e.

- > root transformed. As the number of individual individual of number the As transformed. root

0. (Table I (Table ie o ih) cos the across eight) to five 05

and ). Thus, ). et al. et length c length et al. et ), this averaging approach overcomes this approach averaging ), non classes ,

2008; Lek 2008; - , metric multidimensional scaling scaling multidimensional metric s

ferent dietary categories in each each in categories dietary ferent 2011). While, this approach did approach this While, 2011). eason was removed as a factor factor a as removed was eason

lass

ee sd o rae Bray a create to used were , but not for not but , et al. et et al. et

length class length various ,

2011). T 2011). ,

s 2008; Clarke 2008; eason

l ocation his latter his

at each at or any or al × -

This articleisprotected bycopyright.Allrightsreserved. prior constructingaveraged to shadethe plots. diet Valesini total the to category dietary that of contribution percentage the to proportional linearly is black to grey from shading of depth Acceptedthe while consumed, not that demonstrates category dietary a for space white a where matrix visuali simple a are plots Shade volume. by proportion greatest the contributed each of composition dietary the distinguished and plots greatest the showed factor that Similarity of differences. levels pair which difference, determine to significant used a were comparisons detected ANOSIM where cases those 300 250 south ≠ Articlesame, the in those test. ANOSIM crossed two a with analysed was matrix resemblance same the detected, was interaction no As effects. main the to relative interaction that of extent the determine to si so, if and, interaction a was there whether determine to PERMANOVA Bray a construct to used were replicate five to three of group a representing (each samples these coasts, both among samples. similarity the of representation visual a provided which plot, ordination (nMDS) – – 349 mm and the largest would be 250 be would largest the and mm 349 300 < 299 (Clarke i.e. 250 the in fish As coast,

t al. et level length et al. et i.e. ,

– A 2014) 349 < 350 < 349

= level = they were different were they - , class

2014 Nt ta te ouerc aa for data volumetric the that Note . p N.B retgs analysis ercentages factor were. Thus, while the null hypothesis for both tests were the the were tests both for hypothesis null the while Thus, were. factor – B b 299, 300 299, ) . , the alternative hypothesis for the test for test the for hypothesis alternative the ,

–

that the l the that were employed to identify those dietary categories that typif that categories dietary those identify to employed were 399 mm, 399 - Curtis resemblance matrix and subjected to a two a to subjected and matrix resemblance Curtis – 349 and 350 and 349 , evels in the in evels

but in an unspecified way and that for that and way unspecified an in but i.e.

that the smalles the that – SME; lre Gre, 2015) Gorley, & Clarke (SIMPER; 299

– 399 mm 399 v. l ocation factor were not ordered, whereas whereas ordered, not were factor ocation a priori a

350 – gnificant gnificant 399 mm (Clarke (Clarke mm 399 length length t difference would be 250 be would difference t

group (SIMPER) and those that those and (SIMPER) group oain n lnt class length and location classes were collected from collected were classes location l (Clarke a ocation was west coast west was ocation dietary category was was category dietary et al. et - ie ANOSIM wise length class length × t al. et - egh class length way crossed crossed way ,

N. pandus N. z 2014 n shade and ation data data ation ,

– 2014 299 a - were ). way was was ied

In In b v ) ; .

This articleisprotected bycopyright.Allrightsreserved. ( species studies Acceptedmeta (% number percentage either as recorded were items Dietary prey. stomatopods crustaceans large teleosts, cephalopods, polychaetes, the Oceans. Indian and Pacific Atlantic, 44° at Rise Chatham t from obtained of diet the similar how determine to conducted was Triglidae) and Scorpaenidae Platycephalidae, Article DIETARYCOMPOSITION OF SCORPAENIFORMES THE SIMPER andshadeplots were help then used to describedinterpret as the results, above. determi to ANOSIM 200 199, 150 Esperance from individuals analyses of suite second a coast, south from the collected only were – ulse suis pe ies ee loae t svn itr categories dietary seven to allocated were items prey studies, published - 400 mm 400 nlss Percentage analysis.

N. pandus N. meta this For A of numbers substantial As and thusw six – 249, 250 249, meta N studies), studies), ) ). Only those studies recording data as either % either as data recording studies those Only ). , small crustaceans small , L T - . The differences in mean volumetric data for groups of replicates in the 150 the in replicates of groups for data meanvolumetric in differences The . e ieaue for literature he analysis

was to other species within this suborder globally. Dietary information was information Dietary globally. suborder this within species other to was ere – 299, 300 299, - ne whether dietary composition differed across a broader range of sizes. sizes. of range broader a across differed composition dietary whether ne

analysis, 10 species of platycephalid, including two populations of three of populations two including platycephalid, of species 10 analysis, not included not included in S, to the Irminger Sea at 60° at Sea Irminger the to S, 0

scorpaenids, including two or more populations of six species species six of populations more or two including scorpaenids,

f h de o seis ihn h Sopeodi (families Scorpaenoidei the within species of diet the of

frequency o xmn cag i de ars a ie range wider a across diet in change examine to – 349 and 350 and 349

( 48 e.g N. pandus N.

. Table Table species covering a large geographical area, from the the from area, geographical large a covering species To ensure consistency of taxonomic resolution among resolution taxonomic of consistency ensure To

amphipods ( % F – III )

399 mm length classes were tested by one by tested were classes length mm 399 in the 150 the in f curne aa w data occurrence of

or the the , isopods, mysids and ostracods and mysids isopods, ,

N subsequent

and – 199 and 200 and 199

( e.g mass including the Mediterranean Sea, Sea, Mediterranean the including M

statistical analyses brachy or % or (% were ere M – V 249 mm length classes length mm 249 o vlm (% volume or )

rn, eais and penaeids urans, were included in the in included were not available for all all for available not conducted solely on solely conducted of :

gastropods, ) lengths

. and other other and

and V

- way or ) i.e. N. N. –

This articleisprotected bycopyright.Allrightsreserved. OVERALLDIETARYCOMPOSITION Acceptedfamily toconstructashade andused plot. a and SIMPER to subjected then were data transformed among differed The plot. ordination nMDS an construct to and Platycephalidae) and Triglidae compositionsScorpaenidae, dietary whether determine to Bray a construct square averaged, was groups) (18 replicates of group each in o (platycephalids) two of categorydietary major each ofcontributionpercentage The species. triglids) and (scorpaenids groups into sorted randomly were samples intra of effects confounding a geographic of range a from Articlecolle were samples location, given a in species individual an of diet the of representation 26 and 20 between comprised species a Gulf of Mexico; ( regions of range broadest Pacific the from largely came Scorpaenidae the for those waters; Australian southern in out carried studiesdietary for platycephalids the (predominantly ( species nine pandus

composition dietary The Ocean in the present study present in the

– oain combination; location 18 , Mediterranean Mediterranean , - studies), were included ( included were studies), Table Curtis resemblance matrix. This matrix This matrix. resemblance Curtis

III ) .

- ( Mediterranean family differencesfamily 1 8 Sea es n hbtt (e Tbe III Table (see habitats and reas

studies) and1 studies) s 8 idvda soah ape ad hs rvdd sound a provided thus and samples stomach individual 381

Table by and the Atlantic the and

V % III

RESULTS and

a t a

w ) Sea otal

% 9 ere

, Pacific, Atlantic Pacific, , and triglids, including two, or more populations of of includingtwo,ormorepopulations triglids, M of

oprd ttsial. hl ec samp each While statistically. compared of the of

49 Ocean spatial differences in dietary composition,dietary in differences spatial

Platycephalus species veraged across the replicates within a within replicates the across veraged 49 was subjected to one to subjected was ; while th while ; species in species

from - root transformed and used to used and transformed root ). To remove the potentially potentially the remove To ).

and

39 e Triglidae came from the the from came Triglidae e Bloch 1795 f

amil Indian studies; studies; 77 ‘samples’ ( ‘samples’ ies

fu t five to four r Oceans - Table way ANOSIM ANOSIM way ( three species) were were species)

III i.e. and levels; levels; )

. The . each cted the the le le

This articleisprotected bycopyright.Allrightsreserved. fish mm [ 2 from 250 the for respectively size, body increasing coast with west decreased the polychaetes On and 1). cephalopods (Fig. locations both in greatly changed categories 1% coast west sillaginids monacanthids, ostraciids, clupeids, (47.2 N. on % by 6 only constituting The coast. west of diet the to crustaceans d and groups two volume octop (mainly highest the seven only and coast west the on identified 15 with ingested, teleosts of diversity the in differences (Table coast south the on items with stomachs west the on items prey with stomachs 106 the

i. 1 Fig. Accepted Article endrobranchs

ly on the south coast south the on ly pandus V

% of the diet of of diet the V

south on thesouthcoast (Table (

F As highest the to contributing components dietary The found in excludingunidentifiablewere components, 25components, ofdietary total A a) ( of ) 32% to ]

I cnrs, h rltv cnrbto o tlot duld rm 1 i 250 in 21% from doubled teleosts of contribution relative the contrast, In . N. pandus N. n h ws cat bt ny 6% only but coast, west the on N. , - in the two locations locations two the in 42% in 300 in 42%

coast crustaceans were the most important, constituting 33 and 45% and 33 constituting important, most the were crustaceans ecnae rqec o ocrec ( occurrence of frequency percentage

pandus pandus use y % by

(prawns) contribution

s) and teleosts combined comprised 93% and 79% of the dietary contents dietary the of 79% and 93% comprised combined teleosts and s) N. pandus % fish, respectively. The larger volumetric contributions made by by made contributions volumetric larger The respectively. fish, F

. On the south coast, the gerreids were the only family to contribute to > to contribute to family only the were gerreids the coast, south the On .

) off Rottnest Island and Esperance, respectively (Table respectively Esperance, and Island Rottnest off increased in body size, the volumetric contributions of different dietary dietary different of contributions volumetric the size, body in increased by % by n h suh os, agl de to due largely coast, south the on – . The . – ioos n apios cone fr h oeal otiuin of contribution overall the for accounted amphipods and isopods , 349 mm fish. Decapods fish. mm 349 299 mm 299 N. pandus N. V

frequency and frequency (12.6% (Table of

I ). oyhee i te it dfee markedly differed diets the in polychaetes length

also

on the south coast being 12% greater than i than greater 12% being coast south the on by % by ).

ifrd akdy rpeetn 33 representing markedly, differed

class, to 9 and 1% respectively for 350 for respectively 1% and 9 to class, F volumetric contributions of teleosts to the diet of diet the to teleosts of contributions volumetric )

V of the diet on the west coast compared to 18 to compared coast west the on diet the of and

coast, compared with 19 components in 125 in components 19 with compared coast, (8.8% made similar contributions to contributions similar made gerreids allcomprised > gerreids

I . hs ifrne a mil de to due mainly was difference This ). % F F ) ;

n h suh os. Tetradontids, coast. south the on the the Table Table percentage volume ( volume percentage Aphroditidae being consumed consumed being Aphroditidae II ). Crustaceans, molluscs molluscs Crustaceans,

1% of II

ewe coasts between ). Among these Among ). the the V V the – , %

ad 8.5%, and 7

of the diet of of diet theof n fish on the on fish n 399 mm 399 the diet the of % V % ) V

V also had also

of west west of %

across V – fish fish 299

by of of % ,

This articleisprotected bycopyright.Allrightsreserved. 0.01 0.0 of composition Two 2). Fig. coast; (south before (300 plot the coast of top the to south classes) length (smallest bottom the the from sequentially from those of left the to lie coast VARIATIONS IN forsamewith 42% the size body with increased [ pandus coast, west the on trends the to 350 the in fish of contributions volumetric the of 150 the in 1% < to 27 from respectively 200 the class of size body increasing [ the

Accepted Fig. Article Fig. 1 5 t 39 mm 399 to 250 ), but the but ), and )

1 1 to 5% in the 150 the in 5% to ( (

the 350 the the On b) and isopods of contribution the coast, south the On a) increased with body size, peaking at 300 at peaking bodysize, with increased – 4 m fs to fish mm 249 ] ] . . Although the proportion of teleosts consumed teleosts of proportion the Although . egh classes length

[ i 1 Fig location – N. centroid nMDS ordination, the points representing samples from the west west the from samples representing points the ordination, nMDS centroid 399 mm length class progress horizontally to the left (west coast) and right and coast) (west left the to horizontally progress class length mm 399

pandus pandus DIET DIET ( b) ie lse, otiuig ewe 26 between contributing classes, size – ] × length . 199 and 200 and 199

, length class length N. hl te ouerc otiuin o plcats lo decreased also polychaetes of contributions volumetric the While

LOCATION AND SIZE RELATED DIFFERENCESLOCATION ANDSIZE - the largest largest the

(250 between Location between way PERMANOVA detected a significant difference in the dietary dietary the in difference significant a detected PERMANOVA way 1 n 25 i te 250 the in 2.5% and 31 – pandus 199 and 300 and 199

class – the the 9, 300 299, : isopods declined isopods : on the contributions of cephalopods to the diet of south coast south of diet the to cephalopods of contributions – % 249 mm 249 interaction was not significant ( significant not was interaction V

was 18% was – – west coast (Fig. 1).west coast(Fig. 4 ad 350 and 349 349 mm 349 s

(west and south coast; coast; south and (west length – – 3 399 mm 399

in the 350 the in 49 mm 49 length

– classes 9 ad 350 and 299 from – B . 9 mm; 399

( classes, this group constituted 31% constituted group this classes, length length 50 t groups oth . by by 25

– –

D % 399 mm 399 decapods 5 t te it of diet the to 35% ecapods % N. V

) class [ class

in the 100 the in

pandus and p – seudo 9 mm 399 p p seudo seudo length then declined then

declined Fig. 1 Fig. however, to

- on the south coast coast south the on % F - - – 2 F F

V , class, compared class, 149 mm length mm 149 ( 31 1 2 length

, , 31 31

declined with with declined 2.94, = from ]

. In contrast contrast In . = 5.33, = = 1.44, = – progressed progressed these 349 mm), 349

to classes, classes, 49 20 % in in % P P P fish % N. < < > V

This articleisprotected bycopyright.Allrightsreserved. (Fig. individuals larger < detected were 150 the between differences largest The classes. length consecutive between those except ( ( classes length of range greater ofteleosts4).relatively (Fig. proportions to 350 the of uniqueness The classes. smallest two the in brachyurans with along class, length each of diet the typifying as SIMPER by identified were teleosts ate fish larger classes 350 ( locations both were consumed proportions south onthe coast greater in larger to brachyuran and due teleosts of quantities mainly were locations two the between diet in differences significant The (Fig. location each at diet overall the to contributions volumetric large made categories cephalopods and typif as SIMPER by identified were than composition dietary in variation the of versa 0.05

Acceptedg Article

lobal 0.01). the larger individuals containing lower contributions of brachyurans and cephalopods, but cephalopods, and brachyurans of contributions lower containing individuals larger the – ). 399 mm class399 differed significantly( length . In other words, the length class differences were differences class length the words, other In Two

wider a over length increasing with composition dietary in differences Significant to common classes length three the among significantly differed composition Dietary R (250 This was due to smaller smaller to due was This

- 0.48, way – 299 mm, 299 fewer o – crossed ne typified 199 P -

way ANOSIM way

< brachyurans and octopus and brachyurans mm and both the 300 the both and mm

R .0) Tpcly al pair all Typically, 0.001). 5). ANOSIM demonstrated that that demonstrated ANOSIM

the diet of south of diet the i.e. = 0.17, =

In contrast, larger length classes contained greater proportions of proportions greater contained classes length larger contrast, In

150 – P N. pandas N. 9 m ttl egh wr dtce o te ot coast south the on detected were length) total mm 399

ying the diets of west coast coast west of diets the ying g < s lobal lobal

in 0.0

west - 5 R coast fish coast – ; 300 ; length c length 349 and 350 and 349

= 0.20, = eating more eating - es coast fish, while cephalopods and polychaetes polychaetes and cephalopods while fish, coast

R – and

349 mm, 349 = 0.17, - lass ( lass wise comparisons differed significantly significantly differed comparisons wise , as , more teleosts more P l ocation ( ocation

< 0.01 <

shown on the shade plot where these where plot shade the on shown consistent in both locations both in consistent – (Fig.

̅ 399 mm length classes ( classes mm length 399 P

polychaetes and brachyurans and polychaetes = 0.20). Teleosts and brachyurans and Teleosts 0.20). =

R <

3) ). The diet of diet The ). 0.0 = 0.21, =

N. pandus N. ̅ .

= 0.31) accounted for more for accounted 0.31) =

) from ) from – (Fig. 399 length class was due due was class length 399 P

4) < the twosmallerthe size ,

. while 0.0 Cephalopods and and Cephalopods N. 1

pandus ) brachyurans brachyurans because the the because R

> 0.80, >

and

in the the in than than vice vice

3). P

This articleisprotected bycopyright.Allrightsreserved. triglids and scorpaenids the representing points the where plot nMDS the on demonstrated is This platycephalids. involving comparisons for those than less markedly was difference of compositions dietary ( the significantly differed Although triglids 6). and (Fig. scorpaenids families two other the of those Acceptedfrom right the on group discrete a forming platycephalids representing P famil two other the of each and platycephalids between found were wise significantly platycephalids andtriglids ( of diet Scorpaena of diet the to teleosts Article (Table crustaceans polychaetes and cephalopods by contribution high relatively of diets (Table crustaceans proportions large comprised Platycephalidae DIETARYCOMPOSITION OF SCORPAENIFORMES smallestexcept the (150 fish tele

< osts.

0.0 comparison of diets between families between diets of comparison N. The Overall 1 Cephalopods, on the other hand, were eaten in similar proportions by all size classes classes size all by proportions similar in eaten were hand, other the on Cephalopods, ; triglids triglids ;

Helicolenus pandus

L. 1758 L.

it o fs i te crande Tilde n Payehlde differed Platycephalidae and Triglidae Scorpaenidae, the in fish of diets among families ( families among ,

the dietary composition of members of the Scorpaenidae, Triglidae and and Triglidae Scorpaenidae, the of members of composition dietary the

from locations insouth both

III III species. T species.

0.9 = ). . h hg vlms f ag crustaceans large of volumes high The ). N. pandus N. p. particularly spp.,

Among the scorpaenids, the diet of diet the scorpaenids, the Among 8 Table , – P 199 mm; Figs one he lower contribution of teleosts and small crustaceans to the to crustaceans small and teleosts of contribution lower he

on the south coast is comparable to other other to comparable is coast south the on - III 0.0 way ). 1

. hs s vdn o te MS plot nMDS the on evident is This ). ANOSIM eioeu barathri Helicolenus , the greatest differences in dietary compositions dietary in differences greatest the , 4

- and 5). west ,

by g R lobal lobal ern Australiaern differs

0.2 =

% V

9 r or = N. ,

0.7

P pandus

and % 4 < - (Hector ,

hand side, clearly separated clearly side, hand

M and P i 0. es (scorpaenids es ,

low proportion of small small of proportion low

< 0

was most similar to the the to similar most was

f eess n large and teleosts of 5 0.001) oe cnrbto of contribution lower however, ), the extent of this this of extent the ),

1875) wt te points the with , Neosebastes . From the From , from of that d ue to the the to ue R

= 0.98, =

pair

and -

This articleisprotected bycopyright.Allrightsreserved. Bulman 1998; Potter, & (Platell teleosts and example other Acceptedtwo prey benthic on predominately feeds and carnivore a is species reveal analysis dietary the Overall, teleosts. and polychaetes of contributions the N. OVERALLCONTRIBUTIONS OFTAXATO THEDIETOF twootherfamilies the particularly families, three the among and Australia western of diets Article the of members thedifferencesof indietbetween studies, families the remained consistent teleosts cephalopods, of the between diet in differences crustaceans small of contributions lower and triglids and scorpaenids the both than teleosts of proportions greater contained platycephalids of composition dietary than groups overlapping and discrete less form

pandus of composition dietary the of study This

of diet the to molluscs and crustaceans of contributions volumetric the Although N. , all prey types ingested by ingested types prey all ,

from the the from pandus neosebastid

and Scorpaenidae, Triglidae and Platycephalidae found major differences in the the in differences major found Platycephalidae and Triglidae Scorpaenidae, the large majority of the crustacean prey of prey crustacean the of majority large the on the west coast (Rottnest Island) and south coasts (Esperance) of south of (Esperance) coasts south and Island) (Rottnest coast west the on west and south coast south and west .

i Asrla ( Australia in s

and gastropods latter two families was due mainly to the greater contribution greater the to mainly due was families two latter N. t al. et

DISCUSSION scorpaenoides ltl & otr 19; Bulman 1998; Potter, & Platell

to their diets their to were similar, conspicuous differences were found in found were differences conspicuous similar, were ,

2001). N. hs fr h payehld (Fig. platycephalids the for those

pandus

h hg vlmti cnrbto of contribution volumetric high The (Fig. 7). Despite the geographic spread geographic the Despite 7). (Fig. were benthic invertebrates, crustaceans invertebrates, benthic were

and the meta the and between between M. N.

, paralleling the findings of findings the paralleling , scabriceps

PANDUS the Platycephalidae the - analysis of diets of diets of analysis T .

across regions t al. et

was also benthic also was e es marked less he ,

ed 2001

that this that 6 . The ). ). For For ). .

and

This articleisprotected bycopyright.Allrightsreserved. compared otherspecies the to gape mouth and size body small their to related probably is which mysids, and amphipods as neosebastid 1957 Smith and crustaceans; status [current Schneider & (Bloch 1758) predominantl feeds cirrosa Scorpaenopsis other from distinct is components prey various by contribution the explains components whythedietary highest contributingtothe % thus and feeding of frequency the reduces ( are teleosts Since polychaetes. demersal brachyurans, cephalopods, by made being epifaun infauna, including Harmelin & Bouchon,1976; barathri scorpaenids, other two only of diets the to 10% > of diet the to polychaetes

Accepted ArticleHarmelin probably

consumes fishes and crustaceans in relatively equal amounts; equal relatively in crustaceans and fishes consumes typically prey their that reveal fishes scorpaenid related closely these of Studies the While scorpaen other Like

(Bulman - ie & oco, 1976; Bouchon, & Viven M. )

abl

consumes mainly polychaetes polychaetes mainly consumes scabriceps eatpse strongia Sebastapistes

Scorpaenodes hirsutus Scorpaenodes t cpue n igs rltvl lre ry ee fo a ml size small a from even prey, large relatively ingest and capture to e diet of diet et al. et N. pandus N. y on fishes (78% by (78% fishes on y

1801) (Thunberg ,

2001; N. pandus N. N.

ad etnc ry wt te raet ouerc contributions volumetric greatest the with prey, nektonic and a o consumes mainly consumes is exclusively piscivorous; exclusively is

- ids, pandus Vivien

is a cryptic ambush predator with a relatively large mouth, they mouth, large relatively a with predator ambush cryptic a is (Platell & Potter,(Platell & 1998) Morte et al. N. pandus N.

1793)

is similar to those of other scorpaenids and neosebastids, and scorpaenids other of those to similar is et al. et on ekn, 94 Pael Pte, 1998) Potter, & Platell 1994; Gerking,

the south coast is unusual, with this prey contributing prey this with unusual, is coast south the , previously recognized as recognized previously , Cve 1829)] (Cuvier (Smith mass ,

, 1989;

feeds on a diverse range of prey prey of range diverse a on feeds 2001, Table crustaceans (91%), crustaceans nry xedtr (French expenditure energy )

and (Harmelin 1957) Platell & Potter, 1998; &Potter, Platell Scorpaena notata Scorpaena cephalopods (16%); (16%); cephalopods . Sebastapistes nuchalis Sebastapistes

(formerly (formerly

III feeds predominantly on fishes and and fishes on predominantly feeds ). - ie & oco, 1976) Bouchon, & Viven

particularly

Parascorpaenodes hirsutus Parascorpaenodes V Dendroscorpaena cirrosa Dendroscorpaena and

R also % hadthehighest

Scorpaenopsis gibbosa Scorpaenopsis afinesque, 1810 afinesque, species

La Mesa La Mesa o lse degree, lesser a to Pterois volitans Pterois t al et (Harmelin small taxa such taxa small (Günther . For example, example, For . . This strategy strategy This , 2017) ., et al. et , - and

Viven . 2007) 1874)

The The

and (L. F H. H. . ,

This articleisprotected bycopyright.Allrightsreserved. Ay ideal providing at while waters, these inhabiting species teleost many the for refugia and seas Accepted open 2008) Edgar, 1986; Swainston, and bays sandy exposed moderately in found typically are fish seagr or reef sand, sheltered occupy coast west the from fish of stomachs the in identified prey teleost of diet the 2006) Szedlmayer, & Lingo 2005; Speight, abundance the influence complex more structurally 1998) Hutchings, 1995; Hyndes, floor sea the of nature physical the by explained be may which coast, south than west the from Article BETWEENCOMPARISONSOF LOCATIONS DIET &Potter,1989; Platell 1998 scabriceps of diet the to cephalopods of exception Harmelin studied were species the where zone geographical decapods comprises vazian & & vazian Hyndes, 1995) A

ass habitats, while four of the five fish families found in the stomachs of stomachs the in found families fish five the of four while habitats, ass greater number of taxa of number greater - Vivien ) west ,

is eioeu percoides Helicolenus

- relatively small compared small relatively cover coast t al. et

and

N. for ,

pandus and diversity and 1989; La Mesa Mesa La 1989; to N. other .

;

lse etn tlot ad oyhee, repcie f ie or size of irrespective polychaetes, and teleosts extent lesser a hn h granite the than pandus Morte Morte . The .

, particularly teleosts, particularly , scorpaenids scorpaenids and much less on the south coast. Eight of the 11 families of families 11 the of Eight coast. south the on less much and .

et T

and west and an opportunity for opportunity an and he fact that the limestone reef on the the on reef limestone the that fact he

al ( of teleosts of ihrsn Slne 1842 Solander & Richardson ., 2001 sediment - coast with t al. et . ( Teleosts were a very significant component of component significant very a were Teleosts - e.g. ae res n the on reefs based )

. limestone reef limestone those of those ,

(Friedlander & Parr & (Friedlander Scorpaena 07 Mri & kn 2009) Akins & Morris 2007; composition

were found in the diets of diets the in found were (Harmelin N. pandus N.

in situ in notata differences in reef complexity, reef in differences s offer numerous microhabitats numerous offer s (Howard, 1989; Ayvazian & & Ayvazian 1989; (Howard, - Viven & Bouchon, 1976; 1976; Bouchon, & Viven

foraging south (Harmelin R ish, 1998; Gratwicke & Gratwicke 1998; ish, afinesque ) , the coast (Howard, 1989; (Howard,

contribution the same time time same the - west Vivien

Hthn & (Hutchins 1810 s iey to likely is south . N. pandus N. With the the With os is coast and - et al. et coast

M. M. of ,

This articleisprotected bycopyright.Allrightsreserved. coasts both SIZE nightVolutidae) areactive Octopoda Aphroditidae, the of prey the of number A night. at consumed prey the to differ may which environments these within refuge takes that fauna the day the during numbers greater in consumed were which brachyurans, for found was pattern reverse shrimp polychaetes, (1976) Bouchon of composition dietary the in differences of coast. fishthis from gastropods and polychaetes of contribution greater the for accounting types, prey ( time over of diets the granite regional from eroded has Short 2003; (Commander, which sand, quartz fine of composed typically are coast south the on those while limestone, from derived quartz detrital and sands N. of diet the to gastropods and polychaetes brachyurans, by made contributions in disparity

Accepted Article1995) Hyndes, & Ayvazian

pandus - The d The RELATED CHANGES INDIETARYCOMPOSITION . As scorpaenids hide in reef holes during the day, they may fe may they day, the during holes reef in hide scorpaenids As . be also may patterns feeding in variation Nocturnal in Differences . On the west coast, coast, west the On . , shifting from small prey such as amphipods and isopods in smaller fish smaller in isopods and amphipods as such prey small from shifting , ietary composition of composition ietary south

ouetd il aito i scorp in variation diel documented - coast s

and teleosts being consumed at night than during the day. However, the However, day. the during than night at consumed being teleosts and micro

fish

- et al. et time feeders habitat , .

N. and f eess r less are teleosts If ,

eiet cmrs a i o mdrtl cas carbonate coarse moderately of mix a comprise sediments

pan 2009) N. and sediment composition composition sediment and other

dus pandus .

(Edgar Polychaetes and gastropods were more important in important more were gastropods and Polychaetes may pce fo te olsa (Polyplacopho Mollusca the from species

differed markedly with increasing body size body increasing with markedly differed

. pandus. N. be feeding opportunistically on other abundant other on opportunistically feeding be ,

2008) ies and diverse . aenid diets, with greater volumes of of volumes greater with diets, aenid

(Harmelin

o example, For may also explain the locational the explain also may

abundant on the south coast coast south the on abundant - Viven & Bouchon, 1976) Bouchon, & Viven a atr otiuig to contributing factor ed opportunistically ed Harmelin N.

pandus

to the diets diets the to - ie & Viven , such as such ,

a and ra (< 2 (<

on 49 on ,

This articleisprotected bycopyright.Allrightsreserved. Barnes 1998; Potter, & (Platell crabs and teleosts substratum the below just burrow Gomon, 2001; Potter, & Platell 1996; (Gosline, eyes located sub Acceptedlarge with head flattened behaviour. feeding betw significant interspecificis typically dietary that variation Pottershowed (2001), who & Platell with consistent are findings These families. two latter these between occurring differences platycephalids DIETARYVARIATIONWITHIN THE SCORPAENIFORMES significant transformations Article Consoli 1998; Wootton, N. larger of ability the as in increase well an in as resulting abilities, sensory gape, and locomotory mouth improved increased an as such changes maturational and morphological 2013) Motlagh, Taghavi & Hashemi 2013; show also decapods and teleosts ingest fish Scorpaena example, For species. scorpaenid other individuals larger length) in mm

pandus een families due to distinct differences in body shape, head and mouth morphology mouth and head shape, body in differences distinct to due families een

The

ing

to capture and ingest larger, more mobile prey prey mobile more larger, ingest and capture to meta species feed mainly on small crustaceans and amphipods, while the larger adult larger the while amphipods, and crustaceans small on mainly feed species

hs pattern this t lre pe sc a lre rcyrn rb, ehlpd ad eess in teleosts and cephalopods crabs, brachyuran large as such prey larger to , differed strikingly differed - analysis of analysis

(≥

nie h tild ad sco and triglids the Unlike 250

mm) t al. et ( in feeding

lbr Bla, 1987; Bulman, & Blaber 49 - . Similar Similar . terminal jaws that open downwards, a large gape large a downwards, open that jaws terminal

, species of scorpaeniformes demonstrated that the the that demonstrated scorpaeniformes of species

2010) , from those of the scorpaenids and triglids, with more subtle subtle more with triglids, and scorpaenids the of those from

and and with occur oncea of occur fishlength reach have a tendency to feed on large benthic prey such as such prey benthic large on feed to tendency a have . Harmelin size te pce nteSbsia n Platycephalidae and Sebastidae the in species other Size related dietary changes suggest that the most most the that suggest changes dietary related Size - related Te hf t lre prey larger to shift The . - r Vivien and, p paenids,

et al. et changes in diet have been documented in in documented been have diet in changes ltl & otr 1998 Potter, & Platell ,

et al. et (Keenleyside, 1979; Gerking, 1994; Gerking, 1979; (Keenleyside, 2011, Coulson 2011, ayehld hv a distinctly a have latycephalids ,

(1989)

t al. et

, found that juveniles of juveniles that found c.

2008). They usually usually They 2008).

300 mm.300 et al. et probably ; ,

Baeck, 2016)

and

diet of diet

dorsally dorsally . Their . reflects t al. et

and the ,

This articleisprotected bycopyright.Allrightsreserved. Acceptedobtaining samples. Financial support was provided byMurdoch University. B. types. gape scorpaenids of diet the (Fig triglids the with categories (Harmelin cryptofauna on Articlepre they where cavities, reef around and within day the during refuge take to scorpaenids Gomon 2001; Potter, & (Platell outwards opens that mouth but 2003), Eschmeyer, & (Poss triglids Manderson ( prey epibenthic out flush and excavate detect, enable which receptors, chemosensory tactile with equipped Gomon 2001; Potter, & (Platell eyes developed and triglids ventrally prey large ingesting for capacity

McGovern and his crew his and McGovern (Gomon pca tak ae u t M Jcsn SLih ie efos Apers) cmeca fisherman commercial Applecross), Seafoods, Fine (SELfish Jackson M. to due are thanks Special

sub small a have contrast, in Triglids, ,

due to a clear separation in the pelvic girdle; a feature a girdle; pelvic the in separation clear a to due

(Douglas t al. et et al. et ,

99. uefcal, h sopeis r mrhlgcly iia t the to similar morphologically are scorpaenids the Superficially, 1999). ,

2008), which allows them to capture and ingest a larger range of prey prey of range larger a ingest and capture to them allows which 2008), &

and than triglids triglids than Lanzing, S. Flynn (All Seas Fish Supply, Supply, Fish Seas (All Flynn S. - ie & oco, 96, a epan the explain may 1976), Bouchon, & Viven .

6 1981; Gosline, 1996). ). T ). s accommodated is may he greater contributions of cephalopods and teleosts to teleosts and cephalopods of contributions greater he References ae mr rmnn ha wt a ag terminal large a with head prominent more a have however, however, -

Bardach & Case, 1965; & Case, Bardach terminal mouth that opens downwards, well downwards, opens that mouth terminal et al. et be explained by their significantly larger larger significantly their by explained be

y h aiiy o xad hi stomach their expand to ability the by Esperance) for their very generous assistance in in assistance generous very their for Esperance) ,

2008) these bottom these

t al. et

and not found in found not modified pectoral fin pectoral modified ,

08. h tnec of tendency The 2008). Silver & Finger, 1984; Finger, & Silver ir -

dwelling species to species dwelling overlap of dietary dietary of overlap

the scorpaenids the

y on y rays rays -

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g Tweedley, J. R. & Valesini, F. J. ( J. F. Valesini, & R. J. Tweedley, enus

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aie clg Pors Series Progress Ecology Marine Othos dentex Othos Journal of Experimental of Journal

Ecology marine life: the plants and animals of temperate waters, 2nd waters, temperate of animals and plants the life: marine

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nde n Sopeia) rm uer (Madagascar). Tulear from Scorpaenidae) and anidae

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etr Australian Western

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Cybium - aie clg Pors Series Progress Ecology Marine 0414 nat yac species. bycatch undant ltcpau laevigatus Platycephalus

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108 The impact of impact The vial at Available 34

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745 , 365 , 211. . - Per NY , Perth: 1320. - - - 381. 381. 754. 288. th: th: :

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core.ac.uk/download/pdf/11984538.pdf

Odr cranfre, aiy rgia) n ot Arc. atr Thesis Masters Africa. South in Triglidae) Family Scorpaeniformes, (Order , 1913 F., Nunes, D., Ávila, M., Vieira, R. P. & Barreiros, J. P (2014). Feeding Feeding (2014). P J. Barreiros, & P. R. Vieira, M., Ávila, D., Nunes, F., 10.1007/s10641 r l o tohc n hbtt segregation. habitat and trophic of ole

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This articleisprotected bycopyright.Allrightsreserved. Acceptedof habits Feeding (2000). C. Papaconstantinou, & G. Petrakis, A., Terrats, gurnard tub of Feeding (2012). M. Vallisneri, & S. Montanini, M., Stagioni, Article Kangas, E., Sporer, non a of examination Electrophysiological (1984). E. T. Finger, & L. W. Silver, A. Short, predation chase and ambush of success and Frequency (2010). C. Kruschel, & T. S. Schultz, sad (. Mediterranean). (E. Islands lastoviza Trigloporus gurnard), (red 1758) Kingdom (L., United the of Association doi.org/10.1017/S0025315411000671 Biological Marine lucerna www.fish.wa.gov.au/Documents/sofar/status_reports_of_the_fisheries_and_aquatic_resou Fisheries. Australia Western Fishery Managed A Physiology non Press. University Hydrobiologia fish in Research punctatus dx .doi.org/10.12681/mms.280 rces_2011 -

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This articleisprotected bycopyright.Allrightsreserved. Accepted Article (1998). R. Wootton, (1988). W. C. Bryce, & E. F. Wells, of importance The (2014). C I. Potter, & R. K. Clarke, R., J. Tweedley, J., F. Valesini, sexual size, Body (2011). S. Tommasini, & S. Montanini, M., Stagioni, M., Vallisneri, Publishers. Australian Museum. communities. system regional, eastern Mediterranean.Sea, north in diet and maturity

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4 ad 350 and 349 to – .

– 399 mm length classes from the west coast (Rottnest Island) (Rottnest coast west the from classes length mm 399 – ategories to the stomach contents of contents stomach the to ategories 9 m lnt cass Wie ersns n bec of absence an represents White classes. length mm 399

- western Australia, combined. White represents an an represents White combined. Australia, western - - - root transformed percentage volumetric volumetric percentage transformed root volumetric percentage transformed root root transformed percentage volumetric volumetric percentage transformed root - western Australia in the 150 the in Australia western of south of Neosebastes pandus Neosebastes Neosebastes pandus Neosebastes Neosebastes pandus Neosebastes - western Australi western ory to the totalthe to ory – 199, 200 199,

in a. – 2

This articleisprotected bycopyright.Allrightsreserved. Acceptedincreasing acategory dietary contribution of ( platycephalids main seven of contributions FIG. Articlereproduced from withpermission Nelson morphology. their in differences and similarities the highlight to provided ( triglids prey main seven the of contributions volumetric among FIG. Typesetter 1

6 7 . -

. centroid matrix constructed from Bray from constructed matrix centroid Non

 hd po soig h ma square mean the showing plot Shade

) and platycephalids ( platycephalids and ) Change - metric multidimensional scaling ( scaling multidimensional metric  ) - .

to ht rpeet a asne f ry ad ry o lc rpeet the represents black to grey and prey, of absence an represents White – .

prey categories prey  ). A ). line drawing line et al.

to the total diet. to th to , nMDS – 2016). Curtis similarity matrices that employed the the employed that matrices similarity Curtis e categories - ot transfo root diets of diets )

of ordination plot, derived from distance from derived plot, ordination a typical member of member typical a

scorpaenids ( scorpaenids

to the diets of scorpaenids ( scorpaenids of diets the to rmed percentage volumetric volumetric percentage rmed  ), triglids ( triglids ),

( ie drawings Line each family is is family each  ) and )  ), 3 -

This articleisprotected bycopyright.Allrightsreserved. Accepted winter Wi, Su, summer; spring; autumn; Sp, Au, Article south of (Esperance) 50 sequential I TABLE Total l 400 350 300 250 200 150 100 . ength (mm) Total number of stomach samples of of samples stomach of number Total – – – – – – –

449 399 349 299 249 199 149

length Total -

western

class in each season season each in class ––––––––––––––––––––––––––––––––––– Su 27 12 12 1 2

Australia.

Au 39 16 16 7

West Coast

Wi 17 11 34 6

from

Sp 2 2 2 6 esbse pandus Neosebastes

the the

Location and Season

west Total 106 17 47 41 1

coast (Rottnest Island (Rottnest coast

–––––––––––––––––––––––––––––

Au 10 12 33 4 5 2

that

Wi 18 26 60 contain 3 5 6 2

South Coast

)

Sp and and 11 32 7 8 4 1 1

prey in each each in prey south coast south

Total 125 14 21 16 32 39 2 1

This articleisprotected bycopyright.Allrightsreserved. Accepted Article groups (Esperance) southcoastof ( histograms stacked and components dietary TABLE , shading denotes Crustacea (total) Crustacea component/category Dietary Echinodermata Posidoniaceae Teleostei (total) Teleostei (total) Polychaeta Cephalopoda (total) Mollusca Crustacea

II . Mean volumetric contribution (% contribution volumetric Mean . Unidentifiable Unidentifiable Congridae Callionmidae Syngnathidae Leptoscopidae Pleuronectidae Carangidae Sillaginidae Gerreidae Monacanthidae Pinguipedidae Ostraciidae Clupeidae Tetraodontidae polychaetes Other Aphroditidae (total) Cephalopods molluscs (total) Other Gastropoda (total) Decapoda Stomatopoda Isopoda Amphipoda Unidentifiable Unidentifiable Teuthida Octopoda Polyplacophora Bivalvia Unidentifiable Dendrobranchiata Brachyura

including the amalgamated dietary categories used for multivariate analysis ( analysis multivariate for used categories dietary amalgamated the including

dietary contributions >



western Australiawestern ) of of )

Neosebastes pandus Neosebastes

                  

–––––––––––––––––––– 26.82 26.76 29.46 32.58 11.08 33.47 22.72 23.66 2.31 0.38 1.65 1.27 0.20 0.05 0.67 0.47 0.19 0.94 0.94 4.25 1.51 1.89 0.19 7.50 3.20 1.32 5.66 5.66 0.94 0.94 0.94 1.89 1.27 %V . – – – – Bold denotes

15%

)

West coast West and frequency of occurrence of frequency and

10.26 ± for 6.98 6.14 5.94 6.31 1.94 1.00 3.89 6.85 4.86 1.94 9.71 9.71 7.85 7.98 9.67 1.94 7.88 8.27 4.51 5.67 7.08 from S – – – – – – – – – – – – – – – .

E %V .

the the

total total

33.96 44.34 50.00 57.55 16.98 47.17 12.26 12.26 29.25 30.19 and > 20% and >20% for 4.72 0.94 2.83 2.83 1.89 0.94 3.77 0.94 0.00 0.94 0.94 0.94 5.66 1.89 1.89 0.94 9.43 3.77 2.83 0.94 0.94 0.94 1.89 1.89 %F – – – west coast (Rottnest Island) (Rottnest coast west

and ––––––––––––––––––––– 29.12 25.28 40.05 44.49 17.75 19.56 24.32 8.57 6.20 3.28 1.16 1.76 0.80 0.32 0.80 1.60 0.36 5.64 8.12 9.63 4.04 0.72 0.28 0.28 4.52 %V %F – – – – – – – – – – – –

South coast South

contribution of contribution .

( %

± 4.31 4.30 5.37 4.01 4.61 4.28 8.94 3.58 8.94 8.91 4.02 6.08 5.19 7.57 8.05 6.99 1.92 8.31 F S – – – – – – – – – – – – – – – – – –

. )

E .

fte different the of

35.20 21.60 14.40 40.00 76.00 88.00 12.80 19.20 32.00 23.20 28.80 8.80 3.20 4.00 0.00 0.80 0.00 0.00 0.80 0.80 0.00 1.60 0.00 0.00 0.00 0.80 0.00 8.80 4.80 0.80 1.60 1.60 4.80 %F – – – –

and

12 major12

south  )

This articleisprotected bycopyright.Allrightsreserved. Accepted Article Unidentified material Unidentified Scallop Elasmobranchii Stomachs with( prey Stomachs Stomachs examined ( examined Stomachs

roe

n :

%) n )



106 0.75 122 : – –

87%

– – –

0.94 – –

125 1.40 0.80 0.80 153 :

82%

4.48 8.94 –

3.20 0.80 –

Scorpaenidae Helicolenus dactylopterus dactylopterus Helicolenus dactylopterus dactylopterus Scorpaena Scorpaena maderensis Scorpaena Scorpaena notata Scorpaena porcus Scorpaena porcus Scorpaena porcus Scorpaena porcus Scorpaena scrofa Scorpaena scrofa Sebastes mentella Sebastes maliger Sebastes caurinus Sebastes jordani Sebastes crameri Sebastes diploproa Sebastes alutus Sebastes pinniger Sebastes flavidus Helicolenus hilgendorfii Helicolenus barathri Helicolenus percoides Helicolenus percoides

This articleisprotected bycopyright.Allrightsreserved. Accepted Article

maderensis notate published (O prey other III TABLE

Species

data were

. The The .

;

a offal; offal; mass (%M) mass

available b echinoderms;

and and

volume (% volume c ascidians; ascidians; Data % % % % % % % % % % % % % % % % % % % % % % % %V %V V M M M M M M M M M M M M M M M M M M M M M M M )

of the seven main prey groups, gastropods (G), polycheates (P), cephalopods (C), teleosts (T), large crustaceans (LC), small (LC), crustaceans large (T), teleosts (C), cephalopods (P), polycheates (G), gastropods groups, prey main seven the of d tunicates;

8 2 1 1 1

unidentified material; unidentified P 12 11

1 1 1 3 2 4

Main preycategories C 10 32 12 17

2 5 9 1 5 5

T 10 68 78 15 54 29 16 45 11 43 16 24 29 71 70

1 8 5 7 3 4 7

amphioxiforms LC 19 20 43 60 69 51 74 65 48 65 90 72 69 73 24 12 5 1 8 4 2 9 9

SC 11 89 98 90 95 92 57 21 15 10 22

2 1 3 1 8 7 2 6 2 2 1 1 5

), in the diets of the species within the Platycephalidae, Scorpaenidae and Triglidae and Scorpaenidae Platycephalidae, the within species the of diets the in ),

O 73 13 5 2 1

d b a a c

26,38 285 602 977 368 164 494 848 455 619 182 245 230 576 715 277 321 230 n

30 62 73 93 74 39 64

330 ± 77 ( 265 ± 42 ( 365 ± 60 ( Length range 370 300 ~ 250 ~ 110 ~ 110 ~ 110 ~ 110 111 35 99 49 27 22 116 ~1 33 52 48 87 85 – – – – – – – – – – – – – 135 – na na 272 135 176 240 – 600 580 230 249 469 109 246 225 320 – – – – – 490

450 350 350 150 150 L L L L S.D S.D S.D L L

S S S S L L S

F F

.) .) .)

NWMediterranean NWMediterranean CN Mediterranean CN CN Mediterranean CS Mediterranean W MediterraneanW C Mediterranean C Mediterranean C Mediterranean NE Atlantic NE Atlantic NWPacific SW SW Pacific SW Pacific SW Pacific NE PacificNE PacificNE PacificNE PacificNE PacificNE PacificNE PacificNE N Atlantic Mediterranean E Pacific Location

Rubble Sand, mud, rock Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Rocky reefs Rocky reefs Rocky reefs Rocky reefs Seagrass Seagrass Seagrass Habitat sediment –

seagrass crustaceans (SM) and and (SM) crustaceans

Gonzalez Gonzalez Murie, Murie, Chess & Pearcy, Bodeur & Pearcy, Bodeur & Pearcy, Bodeur & Pearcy, Bodeur & Pearcy, Bodeur Baeck Bulman Horn Bulman Consoli Neves Mesa La Machado 2 Morte Rafrafi 2 Arcuelo Morte 2 Cabiddu which for Harmelin Harmelin Harmelin 1989 1989 1989 et al et al et al et al et et al et al –

Nouira Nouira et al et al et al et al Reference et al 1995 1995 et al et et al et al ., ., – – – ., ., ., ., ., ., vivien vivien vivien vivien

., ., .,

., .,

., 2010 ., 2010 ., 2012

., ., ., .,

1988

2001 2001

2013 2011

et al

2010 2001 2001 1993 2007 2014 2000

., 1984 1984 1984 1984 1984

t al et al et al et

2016

., ., ., .,

Triglidae Chelidonichthys lucerna Trigloporus lastoviza Chelidonichthys Chelidonichthys Chelidonichthys capensis Maxillicostascabriceps Neosebastes pandus Neosebastes pandus Neosebastes scorpaenoides Prionotus Eutrigla gurnardus Eutrigla gurnardus Eutrigla gurnardus Chelidonichthys cuculus Chelidonichthys cuculus Chelidonichthys cuculus Trigloporus lastoviza Trigloporus lastoviza Trigloporus lastoviza Lepidotrigla Lepidotrigla mulhalli Lepidotrigla mulhalli Lepidotrigla Lepidotrigla papilio Lepidotrigla modest Lepidotrigla modesta Lepidotrigla modesta Lepidotriglacavillone Lepidotrigla Trigla lyra Trigla lyra Chelidonichthys kumu Chelidonichthys kumu Chelidonichthys lucerna Chelidonichthys lucerna

This articleisprotected bycopyright.Allrightsreserved. Accepted Article roseus (

Vanessa Vanessa cavillone obscures obscures scitulus

(southcoast) (west coast)

( ( (

Chelidonichthys Chelidonichthys Chelidonichthys

( ( (

) Aspitrigla Aspitrigla Aspitrigla

) ) )

% % % % % % % % % % % % % % % % % % % % %V %V %V %V %V %V %V %V %V %V %V %V %V %V %V M M M M M M M M M M M M M M M M M M M M

23 5 2 3 4 1

14 18 7 6 2 1 1 1 2 4 3 1 2 4 2 1

24 24 14 2 4 5 3 6 4 3 1 1 2 1 1 6 3 9 3

33 18 18 30 25 27 26 20 15 44 88 12 90 40 23 40 34 1 1 1 4 1 1 1 6

16 33 46 50 41 55 36 59 91 59 39 12 68 11 20 50 63 15 56 44 44 37 58 73 58 60 53 30 62 47 40 31 50 7

44 47 19 25 20 15 39 17 24 17 50 27 73 27 10 80 44 93 48 15 17 33 39 60 4 2 8 7 3 1

35 60 b

5,031 4,569 1,096 1,073 1,114 1,092 1,416 469 589 449 193 209 296 396 203 162 106 170 181 129 133 694 203 246 807 135 860 555 207 153 122

20 25 30 86

171 ± 0.1 (S.E.) 189 ± 0.1 ( 203 ± 0.1 ( 295 ± 0.1 ( 225 ± 0.1 ( ~ 21 116 ~ 85 ~ 85 65 98 106 119 103 105 139 160 113 148 254 60 50 50 50 50 50 64 40 63 30 – – – – – – – – – – – – – na na na na na – 295 165 – – – – – – – – – 193 – – 255 300 250 250 175 180 176 140 415 145 140 260 248 207 218 197 267 415 416 418 235 235

L L S.E S.E S.E S.E L

F S L

.) .) .) .)

NE Gulf ofMexico NW NWMediterranean NE MediterraneanNE MediterraneanNE MediterraneanNE MediterraneanNE MediterraneanNE MediterraneanNE CS Mediterranean CS Mediterranean CS Mediterranean E Mediterranean E Mediterranean E Mediterranean NE Atlantic NE Atlantic NE Atlantic NE Atlantic NE Atlantic SW SW Pacific SW Pacific SW SW Pacific SW Pacific SW Pacific SW Pacific SW Pacific SW SW pacific SW SW Indian SE Indian SE SE Indian SE Indian SE Indian Mediterranean

Indian Pacific

Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Sand Sand – – rubble rubble

Ross, 1978 Ross, 1978 Moreno Lopez Montanini Moreno Lopez Terrats Machias Jrad Ben & Labropoulou Terrats Park Park Park Bulman Platell & Potter, Park Bulman & Machias Platell & Potter, & & Labropoulou Terrats Caragitsou Caragitsou Park Bulman Stagioni Lopez Lopez Boudaya Lopez Boudaya McPhail, Platell & study Present study Present Bulman 1998 1998 1998 Papaconstantinou, Papaconstantinou, et al et al et al et al et al – – – – –

Lopez Lopez Lopez Lopez Lopez Lopez et al et al et al – – et al et al et al et al

et al

Amich, 1994 1994 Amich, 1992 Amich, et al et al et al ., ., ., 2017 ., ., ., Potter,

et al 2017 1998 ., 2000 2000 ., 2000 ., ., ., ., ., ., ., 2017 2017 2017 ., . ., ., .,

et al et al et al et al et al

., 2010 2010 .,

2000

b b

2001 2001 2001 2001

2010 2012 2007 2007

a a a

1999 1999 1998 ., 2011 ., 2011 ., 2011 ., ., .,

2011 2011 2011

1994 1994

Platycephalidae Platycephalus caeruleopunctatus Leviprora inops Platycephalus bassensis Platycephalus Ratabulus diversidens Bellator Pterygotrigla polyommata Prionotus punctatus Prionotus nudigula Platycephalus grandispinis Platycephalus speculator Platycephalus laevigatus Platycephalus laevigatus Platycephalus grandispinis Platycephalus westraliae Ambiserrulajugosa Platycephalus richardsoni

This articleisprotected bycopyright.Allrightsreserved. Accepted Article brachychir

7).(Fig. studyeachwere samples in which over type the benthos refersto habitat and wasconducted study the whichin region length from ascertained was range n , richardsoni

sample size. Length range is provided as total length ( length total as provided is range Length size. sample

- frequency data. The mean len mean The data. frequency % % % % % % % % % % % % %V %V %V %V %V M M M M M M M M M M M M

4 1 L

T , mm), unless otherwise specified ( specified otherwise unless mm), ,

20 1 4 1 1 8 gth ( gth

̅ 29 10 ) and and ) 1 5 8 1 2 1 6

S.D 100 77 37 33 63 64 51 75 78 83 79 92 91 96 1 1 . or .

S.E 26 63 32 27 15 20 19 13 10 19 23 33 13 . is presented for those studies which only provided this information. Location refers to the broad geographic broad the to refers Location information. this provided only which studies those for presented is . 7 1 2

L 16 18 54 2 1 1 3 3 4 1 1 S , standard length in mm; in length standard ,

64 48 32 e e e

304 413 415 296 406 362 270 140 350 126 537 248 74 83 42 22 67

collected. T collected. ~ 200 ~ 280 103 171 154 138 41 49 83 83 95 54 54 34 L – – – – – – – na na na F – – – – – 275 600 197 197 409 305 166 , fork length, mm) and ~ is used to denote those studies in which length which in studies those denote to used is ~ and mm) length, fork , – – 83 380 545 470 631

500 510

able ordered to match the order defined by seriation in a shade plot shade a inseriation by defined order the matchto ordered able SW SW Atlantic SW Atlantic SW Atlantic SW SW Pacific SW Pacific SW Pacific SW Pacific SW Pacific SW Pacific SW Pacific SW Pacific SW Pacific SE Indian SE Indian SE Indian SE Indian SE Indian

Sand, soft sediment Sand, seagrass Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Soft sediment Seagrass Seagrass Seagrass Sand

Barnes Barnes Barnes Bulman Bulman Barnes Barnes Coulson 1983 Nichols, and Klumpp Barnes Coulson 1983 Nichols, and Klumpp Coulson Coulson SaoClemente de Bulman SaoClemente de SaoClemente de Platell et al et al et al et al et al et al et al et al et al et al et al et al et t al et ., 2011 2011 ., 2011 ., 2011 ., 2011 ., 2011 ., ., ., ., ., ., 2015 ., 2015 ., 2015 ., 2015 ., 2006 ., 2006

, 05 ; 2015 ., 2001 2001 2001 et al et al et al et

., 2014 ., 2014 ., 2014 ., 2014