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The distribution patterns of exploited girellid, kyphosid and sparid fishes on temperate rocky reefs in New South Wales, Australia

MICHAELKINGSFORD School of Marine Biology and Aquaculture,James Cook University, Townsville,Queensland 4811, Australia ([email protected])

SUMMARYGirellids and sparids rank highest as the groups of fishes taken by rock fishers in New South Wales, Australia.Rapid visual counts that were stratifiedby depth and transects that were stratifiedby habitat (e.g. forestand urchingrazed barrens) showed that girellidswere most abundant in shallowwater (< 3m deep). A few aggregations(primarily tricuspidata) were found in waters up to 15 m deep. No juvenile G. tricuspidata were observed on reefs, all of the fish in counts were greater than 200mm SL. In contrast, G. elevata of all size lasses (30-300mm SL) were found in water less than 3 m deep. Almost all Kyphosus (98%) were in shallow water.Sparids (57%) were found in shallow water, although some (Pagers auratus) were only found in deep or mid-depthstrata. Girellids,kyphosids and some sparids used the intertidalat high fide. Girellidswere observed to feed in the shallows.Juvenile G. elevata (30-70 mm SL) fed in the intertidalmostly on rocky surfaces, tubeworrns and the backs of large limpets (Cellana spp.) where algae were abundant. Shallow waters and the intertidal of rockyreefs have probablybeen underestimated interms of importance to fisheries.

KEY WORDS: Girellidae, Sparidae, Kyphosidae, temperate reef, depth, habitat

INTRODUCTION The confounding influence of depth and habitat can only be resolved if habitats are found in a mosaic over the same Girellids are the most important recreational fish species depth range, as for New South Wales,11)and/or habitats are taken by fishers on reefs in New South Wales, Australia1) experimentally manipulated It is also well known that Although girellids are found on temperate reefs m Australia, different approaches may be required to look for secretive thePacific coast of NorthAmerica, South America and juveniles versus highly mobile adults. Multiple approaches Japan(pers. obs.), there are few data on their ecology. were used in this study. Girellatricuspidata are knownto recruitto estuariesin The objective of this study was to describe the largenumbers and are thoughtto moveto open coastal distributionpatterns of girellid,kyphosid and sparid fishes rockyreefs as subadultsand juveniles, as forother fishes on rockyreefs in New SouthWales, particularly the Sydney suchas Achoerodusviridis (Labridae;6)). In contrast, region. My specificaims were to: (1) describethe depth Girellaelevata are rarelyfound in estuaries.These fish relatedpatterns of highlymobile fishes using rapid visual havebeen collected on rocky reefs7) and 600% or moreof the counts, because girellids, kyphosids and sparids are dietfor fish of all sizesis algae.8)Kyphosids (Kyphosds generallydiver negative; (2) describepatterns of abundance sydneyanus)and sparids (Acanthopagrusaustralis , of fishes over a broad depth range where samplingwas Rhabdosargussarba and Pagrus auratus) also rank highly stratifiedaccording to habitat; (3) intensivelysample the in thecatches of fishersand were also considered in this intertidalat high tide to describethe utilizationof this area study.Girellids consume considerable quantities of algae, by fishes.Some girellids(eg. ,2);Girella butare omnivorous, Kyphosus are obligate herbivores9) and laevifons,3))use shallowareas includingrock pools, but sparidsare predators. samplingwas not done at hightide in. these studies. Studies in temperate waters of California and New Zealand have found that abundance patterns of fishes are MATERIALS AND METHODS often influenced by the presence of different habitat types (review:10).Habitats are generallydistinguished by the was according to Hutchins and Swainston.13) typeand cover of algae. Sea urchinsand limpetshave importantroles in maintaining areas devoid of algae[sensu: Rapid visual counts of girellids and sparids Barrens'habitat; 1) in temperatewaters of Australiaand NewZealand (review:12)]. In additionto habitattype, the Rapid visual counts are useful for estimating the abundance distributionof fishes often varies with depth on a reef(eg.10) . of large fishes that are typically diver negative or are found 132 in large groups that are well separated (eg.10). In New South G..elevata; these fish were followedfor 35 to 94 seconds. Wales large herbivores (eg. Girella tricuspidata) and some The bite rates of fish (per minute)were recordedand the carnivores (eg. Acanthopagnrs australis) behave in one or substratumon which they were feeding. Observations both of these ways. Counts were stratified by depth weremade within an areaof 1ha.

[shallow (0-3m), mid (4-1Om), deep (11-20m)] and 5x3min replicate counts were done at each depth In a three-minute Table 1 Abundance of reef fish in the intertidal at high tide (125m2, n=6 transects); mean (SE). ND denotes no data period I swam a distance of approximately 45m. Fishes collected due to rough seas. were counted at four random times between May 1987 and May 1988 at three locations along the coast of NSW: Botany Bay (34•‹0'S, 151•‹10'E), Bass Point (34•‹30'S, 151•‹50'E) and Jervis Bay (35•‹10'S, 150•‹30'E). Two sites separated by 0.5-1km were sampled at each location Data were analysed using a 4-factor partially-hierarchical ANOVA; factors: depth (fixed), time, location and site

(nested within location). The latter factors were treated as random Data were log (x+1) transformed.

Fish counts stratified by habitat

Most of the habitats were described in") and were classified as follows: 'Shallow broken rock1 (0.5-2m), the 'Fringe'

(2-3m), urchin grazed 'Barrens' (4-12m), Ecklonia radiata dominated kelp forest (4-12m) and 'Deep reef (15-22m). Fishes, especially secretive juveniles, were counted using the transect method Transects measured 25•~5m (n=6). A 25•~5m transect size was chosen because they fitted easily RESULTS within habitats and gave usable average values for a variety of species of fishes. Transects were non-overlapping. A Rapid Visual Counts of Sparidae, Girellidae and detailed search was made under all boulders and/or amongst Kyphosus kelp for more cryptic fishes (eg. juveniles).

Fishes were counted in 5 habitats and at two sites Girellids(90%, n=1093) and Kyphosussycbeyanus (98%, separated by 200m at Cape Banks (34•‹0'S, 151•‹10'E). n=43) were most abundantin very shallowwater (Fig. 1). Counts were done at two times: January 1988 and May Thispattern was consistentamong times, locations and sites. 1988. Patterns of abundance for fishes in shallow broken On someoccasions few fish of a givenspecies were found rock habitat were extracted from maps of the intertidal and in the shallows. This suggestedthat all of these species immediate subtidal as described below. The standard moveover spatial scales greater than those addressed in this length (SL) of fishes was estimated by eye, this method was study (ie. >1km). Variationin abundanceresulted in a calibrated periodically when fishes were collected for other depth x locationx time interaction. Girellatricuspidata reasons. Fishes were considered to be juveniles if they (n=602,200-350mm SL), G. elevata (n=489,30-300mm were less than 100mm SL. SL) and G. zebra (n=9, 100-340mmSL) were observedin counts.

Fish counts in the intertidal Sparids were found at all depths on reefs, but the greatestpercentage of fish was found in shallowwater

Two areas of the intertidal were mapped near Cape Banks (57%,n=490, Fig. 1). ANOVAdetected differences in the and the position of fishes were plotted onto the maps: area rank of sparid abundanceat locationsamong times and

A = 8000m2 and area B = 6250m2. Transect data (as for significantdifferences were foundamong sites within times. The percentagesof fish found at shallow,mid and deep previous section) were extracted from maps using an acetate overlay of a 25•~5m transect; the start point of the trrnsect strata were as follows:Acanthopagrus australts (67, 16, was determined using random numbers on the gridded 17%, n=353),Rhabdosargus sarba (42, 25, 34%, n=101), maps. All transects were non-overlapping. The low tide Pagrus auratus(0, 29, 71%,n=35). mark determined the lowest side of each study area. The Most of the A. australis and R sarba were juveniles positions of fishes were plotted at area A on 7 occasions and (SL: 240-350mm,200-300mm respectively),while the 5 times at area B, due to bad weather (Table 1). P. auratuswere primarily juveniles (70-140mm SL). Behavioural observations were made on snorkel for small 133

A highproportion (30-70%) of the catch by recreational observedin shallowwater in this study(for samplingdesign fishermenin the Sydney area were G. tricuspidata and seer). A.australis. 1) Thefishes they caught were of a similarsize Abundance patterns of fishes in subtidal habitats

Girellidswere abundantin the study area In general, patternsof abundancewere consistentamong habitatsand times. Girellaelevata were only foundin shallowhabitats (Fig.2). Very high numbersof thesefish werefound and a high proportionof them werejuveniles (75%< 70mm SL). Girellatricuspidata were observedoutside of transects(see section:Rapid visual counts ofgirellids and sparids).

Abundance of fishes in the intertidal at high tide Greatnumbers of girellidswere observed grazing and swimmingin the intertidalat hightide (Table 1). Large groupsof Girellatricuspidata and Girellaelevata were observedfeeding on tuftingalgae, particularly where solitaryascidians (?cunjei? ) and/or Ulva covereda largepercentage of thesubstratum (pers. obs.). All ofthe G. tricuspidata were over 250mm SL and large G.elevata were 150-300mm SL. JuvenileG. elevata (Fig. 3)were observed feeding at mean bite rates of 6.9bites per min(SE=1.6; n=29 fish; 30-70mm SL). Fifty-eightpercent of biteswere on rock,28% on tubeworms(Galeolaria), 9.7%on the backsof limpets(Cellana spp.) and 4% on solitaryascidians and algal turf. Recruitmentof juvenile G.elevata varied greatly among times; a meandensity of 9 recruitsper 125m2was recorded in 1988(Fig. 2) andwas muchhigher than mean densities of recruitsrecorded in 1990,1991 and 1992(1.0, 2.2, 1.0 recruits per 125m2 respectively). Large and small predators were observed in the intertidal. Acanthopagrusaustralis (250-300mmSL) were seen in groups of up to six individualsin <0.5m of water,but onlysmall numbers of thesewere seen

Fig. 1 Abundanoe of girellid and sparid fishes at different depths, locations, sites andtimes; mean•}SE Results of ANOV As: Girellidae: adepth x location x time interaction was significant (df 12 ,18, F=3,p<0.025); Sparidae: a location x time (df6, 9F=898, P<0 .01) and site (locaticn) was significant (df3,9,F= 10.8, p<0.01).

range[mean SL (range):Girella spp . 330mm(130-480mm, Fig. 2 Abundance of two species of fishes in 5 habitats, two sites and at two n=181),A. australis240mm (100-400mm , n=91)]to those times. S=shallow broken rock, F=fringe, B=barrens, E=Ecklonia laminarian forest. D=deen reef: mean•}SE. 134

DISCUSSION REFERENCES

There were great differencesin the representationof 1. KingsfordMJ, Undevwod AJ, Ke nelly SJ. Hmnansas pr girellids,kyphosids and spandswith depth. The herbivores on rocky reefs in New SouthWales, Australia.Mar Ecol Prog. Ser. 1991;72:1-14. (girellidsand kyphosids)predominated in shallowwater. 2. Norris K The functions of temperature in the ecologyof the Although herbivorousfishes are common at temperate percoid fish Girella nigncaw (Ayers).EcoL Monogr. 1963;33: latitudes,14,15,16)descriptions of their distributionpatterns are 24-62. limited despite their importance to recreational and 3. Berrios VivianaL, VargasMauricio E. Intertidalfish assemblages commercialfisheries. Some taxa showedgreat variation in structure of the rocky shore at the north of Chile. Revistade abundance among times (eg. Girella tricuspidata) BiologiaMarinay Oceanografia2000; 35:73-81. indicatingthat they have poor site fidelity and appearto 4. McNeill SE, WorthingtonDG, Ferrell DJ, Bell JD. 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Studyingtemperate this has been my experienceover 14 years), adults and marine environments:A handbook for ecologists.University of Canterburypress, Christchurch,New Zealand,1998, pp. 84-114. juvenilesare commonon reefsin New Zealand.") 13. Hutchins B, Swainston R Sea fishes of southern Australia In New SouthWales, girellids and sparids(especially SwainstonPublishing, Perth, 1986, pp. 180. A. australis)rank as the greatestpart of the catchtaken by 14. Hum MH. Biologyof Marine Herbivorousfishes. Oceanography recreationalrock-fishermen.1) , these groups abound in and Marine Biolog an Annual Review1989; 27:167-272. shallowwater (<3m deep). Hence, managementpolicies 15. ChoatJH The biologyof herbivorousfishes on coralreefs. In Sale drat are concernedwith the protectionof coastalresources PF (ed) The ecologyof fishes on coralreefs. AcademicPress, San Diego,1991, pp. 120-155. shouldconsider the importanceof shallowwater habitats. 16. 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