Distribution and community composition of demersal

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Dendrogram of Bray-Curtis similarities derived from demersal fish abundance data (double square root ..... In particular we are grateful to Robert Evans (skipper) for efficiently ... Currie, D.R., and Small, K.J. (2004). ..... Type III Sum of. Squares.
Distribution and community composition of demersal fish in shallow nearshore waters of Port Curtis

2004

David R. Currie AB and Rod M. Connolly AC

A

Cooperative Research Centre for Coastal Zone, Estuary and Waterway Management, Indooroopilly, 80 Meiers Road, Indooroopilly, Queensland 4068, Australia. B

Centre for Environmental Management, Central Queensland University, P.O. Box 1319, Gladstone, Queensland 4680, Australia. C

Centre for Aquatic Processes and Pollution, School of Environmental & Applied Sciences, Griffith University, Bundall, QLD 9726, Australia.

Port Curtis Demersal Fish

Table of Contents

TABLE OF CONTENTS ....................................................................................................................................................... I LIST OF FIGURES ...............................................................................................................................................................II LIST OF TABLES .................................................................................................................................................................II LIST OF APPENDICES .......................................................................................................................................................II 1. INTRODUCTION ..............................................................................................................................................................3 2. MATERIALS AND METHODS .......................................................................................................................................3

2.1 Trawl sampling ...............................................................................................................................3 2.2 Macrobenthos..................................................................................................................................3 2.3 Water chemistry ..............................................................................................................................4 2.4 Data analysis ...................................................................................................................................4 3. RESULTS ............................................................................................................................................................................5

3.1 Faunal characteristics......................................................................................................................5 3.1.1 General distribution of fish ......................................................................................................5 3.1.1 Regional and depth related differences in fish abundance and richness..................................5 3.1.2 Community structure................................................................................................................5 3.1.3 Environmental variables ..........................................................................................................6 4. DISCUSSION......................................................................................................................................................................6 REFERENCES .......................................................................................................................................................................7

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List of Figures Fig. 1. Hydrographical map of Port Curtis showing the locations of 105 sampling stations (small filled circles) surveyed for demersal fish during 2003. ........................................................................................................................................9 Fig. 2. Map of Port Curtis showing the locations of 149 sampling stations (small filled circles) surveyed for macrobenthos, sediment structure and water quality during winter 2002. 105 of these stations (Fig. 1) were subsequently sampled for demersal fish during winter 2003. ...........................................................................................................................10 Fig. 3. Map of Port Curtis showing a) total species richness, and b) total species abundance of demersal fish collected from 3 replicate beam trawl samples (200m length) at 105 sampling stations. .....................................................................11 Fig. 4. Plots of a) mean species abundance and b) mean species richness of demersal fish in Port Curtis. Means (± s.e.) are derived from replicate 200m trawl shots at 105 sampling stations. These stations have been classified here according depth (intertidal, littoral fringe and subtidal) and regional location (upper estuarine, central estuarine an open coastal). ................................................................................................................................................................12 Fig. 5. Dendrogram of Bray-Curtis similarities derived from demersal fish abundance data (double square root transformed). Five station groups are identified............................................................................................................13 Fig. 6. Map showing the locations of 105 demersal fish sampling stations and their classification into 5 groups following cluster analysis of species abundance data. ...................................................................................................................14

List of Tables Table 1. Total abundance (N/31.5 ha) and frequency of occurrence of fish species in beam trawl samples (3*5m*200m) collected at 105 sampling stations in Port Curtis. .........................................................................................................15 Table 2. Results of two-way ANOVA on differences in a) richness, and b) abundance of demersal fish species collected from trawl sampling in three regions (upper estuarine, central estuarine, open coastal) and three depth strata (intertidal, littoral fringe, subtidal) of Port Curtis. All data were transformed (Log10N+1) prior to analysis to remove heterogeneity of variance. .............................................................................................................................................16 Table 3. Pearson correlation coefficients between depth, temperature, salinity, dissolved oxygen concentration, pH, turbidity (secci depth), total chlorophyll concentration, sediment structure, organic carbon, content seagrass density, infaunal species richness (S), infaunal species abundance (N), demersal fish abundance (N) and demersal fish richness (S). Significant correlations are denoted at the: **1% level and *5% level. ................................................................17 Table 4. Abundance (N/3000m2) of demersal fish in five regions of Port Curtis identified from hierarchical classification of species/abundance data. Species listed were identified from SIMPER analyses as contributing ≥ 5% to the similarity within and dissimilarity between regional groupings. Those species indicative of each regional grouping (contributing ≥ 5% to the total similarity within a group) are highlighted in bold. Species are ranked in order of decreasing abundance across all stations.......................................................................................................................18

List of Appendices Appendix 1. Distributional maps of demersal fish abundance determine from replicate beam trawl shots (3*200m) at 105 station in Port Curtis......................................................................................................................................................19 Appendix 2. Predictive maps of Port Curtis showing distributions of key water quality parameters including: (a) temperature, (b) salinity, (c) ph, (d) dissolved oxygen content, (e) turbidity and (f) total chlorophyll concentration. Note all estimates presented are derived from winter 2002 depth-profiled averages at 149 separate sampling stations (small filled circles).......................................................................................................................................................62 Appendix 2 cont. Predictive maps of Port Curtis benthos showing distributions of: (a) infaunal richness, (b) infaunal abundance, (c) sediment size-structure, and (d) organic carbon content. Note estimates presented are derived from replicate samples (n=3) collected at 149 separate sampling stations (small filled circles)............................................63

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1. Introduction Estuaries are widely recognised as important nursery grounds for juvenile fish, and are thought to provide a high availability of food resources and offer shelter from predators (Blaber, 1980; Robertson and Duke, 1987; Blaber et al., 1989).

Human alterations to estuaries, including loss of habitat, may therefore have profound effects on fishery

productivity and other ecological processes (Nichols et al., 1986; Hutchings, 1999). Land reclamation of intertidal areas along the Forth estuary in Scotland have, for example, resulted in a loss of over 50% of the invertebrate benthic production, and reportedly led to large declines in fish and bird populations (McLusky et al., 1992). Characterising and evaluating the relative importance of benthic habitats is therefore a fundamental requirement for the conservation and better management of these waterways. This study concentrates on assessing species composition and distribution of demersal fish in a marine dominated estuary, and their utilization of different sedimentary facies. In particular, we examine 1) spatial patterns in the community structure of fish and their relationships with prey availability, 2) the importance of structural complexity (i.e. seagrass availability) in influencing species composition, and 3) the relative importance of a range of environmental variables (including sediment structure, depth , salinity and seagrass biomass) in governing the abundance and diversity of fish.

2. Materials and methods 2.1 Trawl sampling Fish were collected from 105 intertidal and shallow subtidal sampling stations (1m above MLWS), “littoral fringe” (1m ± MLWS), and “subtidal” (>1m below MLWS). Prior to these analyses, homogeneity of variance was examined by Cochran’s test and heterogeneity removed by a log10 (N+1) transformation. Variations in community structure of demersal fish between the 105 Port Curtis stations were examined using Bray-Curtis dissimilarity measures (Bray and Curtis, 1957). This dissimilarity measure was chosen because it is not affected by joint absences, it gives more weighting to abundant than rare species, and it has consistently performed well in preserving ecological distance in a variety of simulations on different types of data (Field et al., 1982; Faith et al., 1987). Double square root transformations were applied to the data to prevent abundant species from influencing the Bray-Curtis dissimilarity measures excessively (Clarke and Green, 1988; Clarke, 1993). Multivariate analyses were conducted using the computer package PRIMER (Clarke and Gorley, 2001). Hierarchical agglomerative clustering was used to group sites according to their community composition, and a similarity percentage test (SIMPER) conducted to determine those species contributing most to within and between site groupings. The extent to which measured environmental variables (temperature, salinity, pH, DO, turbidity, chlorophyll, sediment structure, organic carbon, seagrass, macrobenthic abundance, and macrobenthic richness) could account for observed community groupings was further tested using the BIOENV routine of Clarke and Ainsworth (1993). A geographical information system (GIS) was employed to characterise and display spatial trends in environmental data. Physical, chemical and biological attributes for each sampling station were interpolated using an inverse distance weighting (IDW) algorithm (Cressie, 1993), and series of predictive maps constructed. These maps were used to visualise

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discontinuities between homogeneous regions and highlight patterns of similarity between variables. Relationships between each environmental variable were subsequently tested using Pearsons correlation coefficients.

3. Results 3.1 Faunal characteristics 3.1.1 General distribution of fish A total 88 species of fish and 2994 individuals were collected from the 315 replicate trawl shots (3*105 stations) undertaken in Port Curtis. The Ponyfish Leiognathus decora was the most abundant and widespread species found during the study, and occurred at half of the stations sampled (50%), and accounted for more than a third of the total abundance (34%) (Table 1). Another schooling fish, the Herring Herklostichthys castelnaui, was also common, and occurred at 36% of the sampling stations, and accounted for over 16% of the total catch. All other species had restricted distributions and were found at fewer than 30% of the stations sampled, and individually contributed less than 9% to the total abundance (Appendix 1). 3.1.1 Regional and depth related differences in fish abundance and richness Patterns in fish richness broadly mirrored patterns in fish abundance across the estuary, and in most locations where richness was high, the abundance was also high (Fig. 3). Similarly, in areas where fish richness was low, abundance was also low (Fig. 3). Analysis of variance showed that there were no significant differences in the number of species or individuals found between the three regions of the port (upper estuarine, central estuarine, open coastal) (Tables 2a and 2b). Despite this, plots of mean abundance and richness suggest a general decline in both parameters towards the open coastal waters of the estuary (Figs. 4a and 4b). Furthermore, these plots also suggest that both fish abundance and richness was lower in the intertidal than the littoral fringe and subtidal, even though the ANOVA tests between depth strata showed no significant differences for these parameters. 3.1.2 Community structure Five station groupings were separated by cluster analysis of species abundance data (Fig. 5), and their corresponding distributions plotted (Fig. 6). These groupings closely reflected differences in depth and habitat type within Port Curtis. Group I was intimately associated with the seagrass beds (Pelican banks, South Trees, Tannum). SIMPER analysis was undertaken to assess which species contributed most to similarities within and differences between the nine station groupings. Abundances of the 14 species contributing ≥5% to within-group similarity or between-group dissimilarity for at least one of the five groupings are given in Table 4. The average within-group similarity ranged from 9 to 35%, and between group dissimilarity ranged from 76 to 99%. Two station group (I & V) were independently characterised by small subsets of species with restricted distributions. The other three groups (II, III & IV) had generally higher diversity, and were differentiated by varying proportions of co-occurring species. The PRIMER routine BIOENV was used to assess the correspondence and significance of environmental data to the nine station groupings. The best fit was with turbidity (ρw = 0.22), which in combination with depth, infaunal richness and seagrass biomass and gave a best fit of ρw = 0.34. The remaining variables (sediment, pH, salinity, temperature, dissolved oxygen, organic carbon, chlorophyll and infaunal abundance) were apparently unrelated to any pattern in station groupings (ρw < 0.14).

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3.1.3 Environmental variables Fish abundance and richness were found to be highly (positively) correlated (Table 3), and consistent with the similarities demonstrated in the distributional maps for both parameters (Fig. 3). Fish abundance and richness were also correlated salinity, but this association was not particularly strong and less than 30% of the variation was explained. It is notable that salinity was highest in the upper estuary, but gradually declined through the central estuary towards the open coast (Appendix 2). All other environmental parameters measured were unrelated to either the abundance or diversity of fish in the estuary (Table 3).

4. Discussion Fish assemblages of Port Curtis are diverse, with 88 species of fish recorded. The dominance of the fish fauna by two small schooling species is typical of the fauna recorded in similar studies elsewhere in Queensland (Blaber et al., 1989; Thomas and Connolly, 2001). In this case, the dominant species were ponyfish (Leiognathus equulus) and herring (Herklotsichthys castelnaui), both species common in inshore coastal waters elsewhere in tropical and subtropical waters of Australia. Fish assemblages change gradually from the more estuarine Narrows to open coastal waters. This is essentially an estuarine gradient, known to occur elsewhere in Australia (West and King, 1996) but not previously reported in subtropical waters. The influence of salinity per se is not strong. If anything, turbidity is more important, since the correlation between fish and turbidity was the strongest measurable relationship. Alternatively, the degree of exposure to open coastal waters might itself be influencing fish distributions (Connolly, 1994), but this is difficult to separate from turbidity since open coastal waters are typically less turbid in any case. In addition to the estuarine gradient, five assemblage types were defined, reflecting differences in habitat type and depth. This comes as no surprise since both habitat and depth have been shown to be important in determining fish assemblages in several other places (Jackson et al., 2001). The group of fish associated with seagrass is of particular note, since it confirms in Port Curtis the importance of this habitat type, as reported elsewhere in Australia (Connolly, 1994) and internationally (Jackson et al., 2001).

Acknowledgements Many people have made significant contributions to this project. In particular we are grateful to Robert Evans (skipper) for efficiently undertaking all trawl sampling associated with this project. We are also most grateful to the following Centre for Environmental Management staff for their assistance in collecting and processing samples: Damon Shearer, Vikki Rogers, Kirsty Small, Shona Marks, Andrew Davis, and Nathalie Callanan. Hydrographic survey data for the port was made available by the Queensland Department of Transport and the Australian Department of Defence, and its provision is greatly acknowledged.

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References Blaber, S.J.M. (1980). Fish of the Trinity Inlet system of North Queensland with notes on the ecology of fish faunas of tropical Indo-Pacific estuaries. Australian Journal of Marine and Freshwater Research 31, 137-146. Blaber, S.J.M., Brewer, D.T., and Salini, J.P. (1989). Species composition and biomasses of fishes in different habitats of a tropical northern Australian estuary: their occurrence in the adjoining sea and estuarine dependence. Estuarine, Coastal and Shelf Science 29, 509-531. Bray, J.R., and Curtis, J.T. (1957). An ordination of the upland forest communities of southern Wisconsin. Ecological Monographs 27, 325-349. Clarke, K.R. (1993). Non-parametric multivariate analysis of changes in community structure. Australian Journal of Ecology 18, 117-143. Clarke, K.R., and Ainsworth, M. (1993). A method for linking multivariate community structure to environmental variables. Marine Ecology Progress Series 92, 205-219. Clarke, K.R., and Gorley, R.N. (2001). ‘PRIMER v5 Users Manual / Tutorial.’ (PRIMER-E: Plymouth.) Clarke, K.R., and Green, R.H. (1988). Statistical design and analysis for a ‘biological effects’ study. Marine Ecology Progress Series 46, 213-226. Connolly, R.M. (1994). A comparison of fish assemblages from seagrass and unvegetated areas of a southern Australian estuary. Australian Journal of Marine and Freshwater Research 45, 1033-1044. Cressie, N. (1993). ‘Statistics for Spatial Data.’ (John Wiley & Sons: New York.) Currie, D.R., and Small, K.J. (2004). Physico-chemical characteristics of Port Curtis and their influence on the community composition of bottom dwelling fauna. Centre for Environmental Management Technical Report. (Central Queensland University: Gladstone.) Faith, D.P., Minchin, P.R., and Belbin, L. (1987). Compositional dissimilarity as a robust measure of ecological distance. Vegetatio 69, 57-68. Field, J.G., Clarke, K.R., and Warwick, R.M. (1982). A practical strategy for analysing multispecies distribution patterns. Marine Ecology Progress Series 8, 37-52. Hutchings, P. (1999). Taxonomy of estuarine invertebrates in Australia. Australian Journal of Ecology 24, 381-394. ISO 10260 (1992). Water quality - Measurement of biochemical parameters - Spectometric determination of the chlorophyll a concentration. (International Organization for Standardization: Geneva) Jackson, E.L., Rowden, A.A., Attrill, M.J., Bossey, S.J., and Jones, M.B. ( 2001). The importance of seagrass beds as a habitat for fishery species. Oceanography and Marine Biology – an Annual Review 39, 269-303. McLuskey, D.S., Bryant, D,M., and Elliott, M. (1992). The impact of land-claim on macrozoobenthos, fish and shorebirds on the Forth estuary, eastern Scotland. Aquatic Conservation: Marine and Freshwater Ecosystems 2, 211-222. Nichols, F.H., Thompson, J.K., and Schemel, L.E. (1986). The modification of an estuary. Science 231, 567-573. Robertson, A.I., and Duke, N.C. (1987). Mangroves as nursery sites: comparisons of the abundance and species composition of fish and crustaceans in mangrove and other nearshore habitats in tropical Australia. Marine Biology 96, 193-205. Thomas, B.E. and Connolly R.M. (2001). Fish use of subtropical saltmarshes in Queensland, Australia: relationships with vegetation, water depth and distance onto the marsh. Marine Ecology Progress Series 209, 275-288.

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West, R.J. and King, R.J. (1996). Marine, brackish, and freshwater fish communities in the vegetated and bare shallows of an Australian coastal river. Estuaries 19,:31-41.

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Fig. 1. Hydrographical map of Port Curtis showing the locations of 105 sampling stations (small filled circles) surveyed for demersal fish during 2003.

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Fig. 2. Map of Port Curtis showing the locations of 149 sampling stations (small filled circles) surveyed for macrobenthos, sediment structure and water quality during winter 2002. 105 of these stations (Fig. 1) were subsequently sampled for demersal fish during winter 2003.

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Fig. 3. Map of Port Curtis showing a) total species richness, and b) total species abundance of demersal fish collected from 3 replicate beam trawl samples (200m length) at 105 sampling stations. (a) ! (

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Fig. 4. Plots of a) mean species abundance and b) mean species richness of demersal fish in Port Curtis. Means (± s.e.) are derived from replicate 200m trawl shots at 105 sampling stations. These stations have been classified here according depth (intertidal, littoral fringe and subtidal) and regional location (upper estuarine, central estuarine an open coastal). (a)

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Fig. 5. Dendrogram of Bray-Curtis similarities derived from demersal fish abundance data (double square root transformed). Five station groups are identified.

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Fig. 6. Map showing the locations of 105 demersal fish sampling stations and their classification into 5 groups following cluster analysis of species abundance data.

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Table 1. Total abundance (N/31.5 ha) and frequency of occurrence of fish species in beam trawl samples (3*5m*200m) collected at 105 sampling stations in Port Curtis. Species

Common name

Leiognathus decora Herklostichthys castelnaui Ambassis marianus Megalaspis cordyla Atherinomorous ogilbyi Siganus rivulatus Mugil georgii Pomadasys kaakan Gerres subfasciatus Saurida undosquamis Pelates quadrilineatus Acanthopagrus australis Meuschenia sp. 2 Apogon fasciatus Leptobrama mulleri Liza dussumieri Tripodichthys angustifrons Arrhamphus sclerolepis Gerres filamentosus Pseudorhombus arsius Gerres oyena Acanthopagrus sp. 2 Nematalosa come Pelates sexlineatus Selaroides leptolepis Sillago maculata maculata Trepaon jarbua Drepane punctata Scomberoides commersonianus Thryssa aestuaria Tetractenos hamiltoni Marilyna pleurosticta Triacanthus brevirostris Upeneus tragula Arothron manilensis Hyporhamphus quoyi Leiognathus sp. 3 Leognathus sp. 2 Lethrinus variegatus Platycephalus indicus Sphyraena obtusata Polydactylus multiradiatus Leiognathus bindus Meuschenia trachlepis Monodactylus argenteus Rachycentron canadus Sillago ciliata Caranx ignobilis Istigobius hoesei Lagocephalus sceleratus Leiognathus sp. 1 Lutjanus russelli Rhadbosargus sarba Selenotoca multifasciata Sillago (Parasillago) analis Sygnathidae sp.1 Absalom radiatus Apogon sp. 1 Argyrosomus japonicus Carangoides caeruleopinnatus Chaetodon tricinctus Dasyatis fluviorum Hippocampus sp. 1 Hyperolophus translucidus Meuschenia sp. 3 Nemipterus theodorei Parachaetodon ocellatus Tylosurus crocodilus Achlyopa nigra Auxis thazard Choerodon cephalotes Dasyatis kuhlii Diodor nichthemerus Eleutheronema tetradactylum Epinephelus coioides Gymnura australis Himantura uarnak Hyporhamphus australis Meuschenia sp. 1 Meuschenia sp. 4 Muraenesox cinevus Pentapodus paradiseus Periopthalmus koelreuteri Platycephalus arenarius Pseudorhombus argent Scomberomorus queenslandicus Scorpaenopsis sp. Ulua mentalis Total

Common Ponyfish Southern Herring Glass Perch Finny Scad Common Hardyhead Happy Moments Fantail Mullet Barred Grunter Common Silverbelly Large-scaled Grinner Trumpeter Silver Bream Leatherjacket Striped Cardinalfish Steelback Salmon Flat-tail Mullet Yellow-fin Tripod-fish Snub-nosed Garfish Threadfin Silverbelly Large Toothed Flounder Oceanic Silverbelly Bream Bony Bream Eastern Striped Trumpeter Smooth-tailed Trevally Winter Whiting Crescent Perch Sicklefish Giant Leatherskin Southern Anchovy Common Toadfish Banded Toadfish Short-nosed Tripod-fish Bar-tailed Goatfish Narrow-lined Toadfish Short-nosed Garfish Ponyfish Ponyfish Variegated Sweetlip Bar-tailed Flathead Striped Sea Pike Blind Threadfin Orange-tipped Ponyfish Yellow-tailed Leatherjacket Butter Bream Black Kingfish Sand Whiting Giant Trevally Sloth Goby Giant Toadfish Ponyfish Moses Perch Tarwhine Striped Butterfish Rough Scaled Whiting Pipefish Fringe-finned Trevally Cardinalfish Jewfish Onion-ring Trevally Three-band Coralfish Brown Stingray Seahorse Glassy Sprat Leatherjacket Yellow-lip Butterfly-bream Ocellate Coralfish Crocodile Long-tom Black Sole Frigate Mackeral Purple Tuskfish Bluespot Stingray Porcupine Fish Blue Threadfin Salmon Orange-spotted Cod Rat-tailed Ray Long-tailed Ray Sea Garfish Leatherjacket Leatherjacket Pike Eel Blue-faced Whiptail Mud-skipper Sand Flathead Flounder School Mackeral Scorpionfish Cale Cale Trevally

Abundance (n/31.5 ha) 1030 489 244 121 97 83 80 77 65 65 53 47 42 30 30 27 26 24 23 23 20 18 18 17 15 15 14 12 10 10 9 8 7 7 6 6 6 6 6 6 6 5 4 4 4 4 4 3 3 3 3 3 3 3 3 3 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2994

Relative abundance (% ) 34.40 16.33 8.15 4.04 3.24 2.77 2.67 2.57 2.17 2.17 1.77 1.57 1.40 1.00 1.00 0.90 0.87 0.80 0.77 0.77 0.67 0.60 0.60 0.57 0.50 0.50 0.47 0.40 0.33 0.33 0.30 0.27 0.23 0.23 0.20 0.20 0.20 0.20 0.20 0.20 0.20 0.17 0.13 0.13 0.13 0.13 0.13 0.10 0.10 0.10 0.10 0.10 0.10 0.10 0.10 0.10 0.07 0.07 0.07 0.07 0.07 0.07 0.07 0.07 0.07 0.07 0.07 0.07 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03 0.03

Frequency of occurrence 53 36 7 15 17 9 12 14 25 29 10 13 4 12 14 8 18 7 10 18 1 1 9 2 3 10 6 8 8 2 7 5 6 4 5 1 3 3 2 4 4 4 2 3 3 1 3 2 1 3 2 2 1 1 2 2 1 2 2 2 1 2 2 1 2 2 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 105

Occurrence (%) 50.48 34.29 6.67 14.29 16.19 8.57 11.43 13.33 23.81 27.62 9.52 12.38 3.81 11.43 13.33 7.62 17.14 6.67 9.52 17.14 0.95 0.95 8.57 1.90 2.86 9.52 5.71 7.62 7.62 1.90 6.67 4.76 5.71 3.81 4.76 0.95 2.86 2.86 1.90 3.81 3.81 3.81 1.90 2.86 2.86 0.95 2.86 1.90 0.95 2.86 1.90 1.90 0.95 0.95 1.90 1.90 0.95 1.90 1.90 1.90 0.95 1.90 1.90 0.95 1.90 1.90 0.95 1.90 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95 0.95

Centre for Environmental Management CQU - 15

Port Curtis Demersal Fish

Table 2. Results of two-way ANOVA on differences in a) richness, and b) abundance of demersal fish species collected from trawl sampling in three regions (upper estuarine, central estuarine, open coastal) and three depth strata (intertidal, littoral fringe, subtidal) of Port Curtis. All data were transformed (Log10N+1) prior to analysis to remove heterogeneity of variance. (a) - Species richness Source Corrected Model Intercept Strata Region Strata * Region Error Total Corrected Total

Type III Sum of Squares 1.038 16.072 .272 .295 .156 7.811 55.545 8.848

df 8 1 2 2 4 96 105 104

Mean Square .130 16.072 .136 .147 .039 .081

F 1.594 197.546 1.671 1.810 .480

Sig. .137 .000 .193 .169 .750

Power .675 1.000 .344 .370 .160

Type III Sum of Squares 2.996 42.646 .527 2.155 .631 33.719 166.064 36.715

df 8 1 2 2 4 96 105 104

Mean Square .375 42.646 .264 1.078 .158 .351

F 1.066 121.416 .750 3.068 .449

Sig. .393 .000 .475 .051 .773

Power .470 1.000 .174 .580 .151

(b) - Species abundance Source Corrected Model Intercept Strata Region Strata * Region Error Total Corrected Total

Centre for Environmental Management CQU - 16

Port Curtis Demersal Fish

Table 3. Pearson correlation coefficients between depth, temperature, salinity, dissolved oxygen concentration, pH, turbidity (secci depth), total chlorophyll concentration, sediment structure, organic carbon, content seagrass density, infaunal species richness (S), infaunal species abundance (N), demersal fish abundance (N) and demersal fish richness (S). Significant correlations are denoted at the: **1% level and *5% level. Depth Depth Temperature Salinity D0 pH Turbidity Chlorophyll Sediment Carbon Seagrass Infauna S Infauna N Fish S Fish N

0.08 0.45** -0.06 -0.44** -0.65** -0.02 -0.27** -0.13 0.08 -0.19* 0.04 -0.04 0.09

Temperature 0.01 -0.27** -0.15 -0.13 -0.02 -0.02 -0.13 -0.04 -0.07 0.14 -0.06 -0.09

Salinity 0.04 -0.88** -0.56** -0.29** -0.09 0.04 0.02 0.01 -0.01 0.20** 0.30**

DO 0.08 0.04 -0.17* 0.10 -0.08 0.02 0.13 0.01 -0.07 0.05

pH 0.57** 0.26** 0.04 -0.01 0.06 -0.09 -0.09 -0.10 -0.08

Turbidity 0.05 -0.02 0.17 0.06 -0.05 -0.09 -0.07 -0.11

Chlorophyll 0.02 -0.03 0.09 -0.01 0.06 -0.23* -0.18

Sediment -0.07 -0.09 0.44** 0.34** -0.02 -0.06

Carbon -0.08 0.34** -0.02 0.11 0.03

Seagrass -0.09 -0.02 -0.05 -0.06

Infauna S 0.37** -0.02 -0.14

Infauna N -0.10 -0.09

Fish S 0.64**

Centre for Environmental Management CQU - 17

Port Curtis Demersal Fish

Table 4. Abundance (N/3000m2) of demersal fish in five regions of Port Curtis identified from hierarchical classification of species/abundance data. Species listed were identified from SIMPER analyses as contributing ≥ 5% to the similarity within and dissimilarity between regional groupings. Those species indicative of each regional grouping (contributing ≥ 5% to the total similarity within a group) are highlighted in bold. Species are ranked in order of decreasing abundance across all stations. Species

Common Name 1

Leiognathus decora Herklostichthys castelnaui Ambassis marianus Megalaspis cordyla Siganus rivulatus Saurida undosquamis Apogon fasciatus Liza dussumieri Tripodichthys angustifrons Pseudorhombus arsius Sillago maculata maculata Scomberoides commersonianus Upeneus tragula Hippocampus sp. 1

Common Ponyfish Southern Herring Glass Perch Finny Scad Happy Moment Large-scaled Grinner Striped Cardinalfish Flat-tailed Mullet Yellow-fin Tripod Fish Large-toothed Flounder Winter Whiting Giant Leatherskin Bar-tailed Goatfish Seahorse

2 1.43 0.88 0.29 0.62 0.29 0.29

0.05

Station Grouping 3 4 28.28 12.40 7.68 0.32 0.08 0.80 0.36 0.24 0.24 0.12

5

9.47 4.38 1.53 2.53 6.75 1.15 0.50

0.33

0.38 0.38 0.21

0.67

0.24 0.50

0.50

Centre for Environmental Management CQU - 18

Port Curtis Demersal Fish

Appendix 1. Distributional maps of demersal fish abundance determine from replicate beam trawl shots (3*200m) at 105 station in Port Curtis.

³

( !

Total abundance (n/600m)

Absalom_radiatus ( !

2

0 1 2

4

6 Km

! ( ( !

³

( !

! (!( ! ( ( !

! (

( !

( !

Total abundance (n/600m) ( !

Acanthopagrus_australis 1 2 ! ( 3 ( 4-7 ! ! ( 8 - 13

( !

( !

( !

( !

0 1 2

4

6 Km

Centre for Environmental Management CQU - 19

Port Curtis Demersal Fish

³

( !

Total abundance (n/600m)

Acanthopagrus_sp2 ( 18 !

³

0 1 2

4

6 Km

0 1 2

4

6 Km

( !

Total abundance (n/600m)

Achlyopa_nigra ( !

1

Centre for Environmental Management CQU - 20

Port Curtis Demersal Fish

³

! (

! ( ( !

( !

( !

( !

! (

Total abundance (n/600m)

Ambassis_marianus 1 2-5 ! ( 6 - 19 ( 20 - 22 ! ! ( 23 - 99 ( !

( !

³

0 1 2

4

6 Km

( !

( !

( !

( !

( !

( !

( !

( !

! ( ( !

( !

Total abundance (n/600m)

Apogon_fasciatus 1 2 ( 3-4 ! ! ( 5 - 14 ( !

( !

( !

0 1 2

4

6 Km

Centre for Environmental Management CQU - 21

Port Curtis Demersal Fish

³

( !

( !

Total abundance (n/600m)

Apogon_sp1 ( !

1

³

0 1 2

4

6 Km

4

6 Km

( !

( !

Total abundance (n/600m)

Argyrosomus_japonicus ( !

1

0 1 2

Centre for Environmental Management CQU - 22

Port Curtis Demersal Fish

³

( ! ( ! ( !

( !

Total abundance (n/600m)

Arothron_manilensis ( ! ( !

( !

1 2

0 1 2

4

6 Km

( !

³

( !

( !

! (

( !

( !

Total abundance (n/600m)

Arrhamphus_sclerolepis 1 2 ! ( 3 ( 4 ! ! ( 5 - 12 ( !

! (

( !

0 1 2

4

6 Km

Centre for Environmental Management CQU - 23

Port Curtis Demersal Fish

( !

³

! ( ! ( ! (

( !

( !

( !

( !

( !

( !

( ! ( !

( !

! ( Total abundance (n/600m) ( !

Atherinomorous_ogilbyi 1 2-3 ! ( 4-7 ( 8 - 18 ! ! ( 19 - 26 ( !

( !

( !

( !

0 1 2

4

6 Km

³ ( !

Total abundance (n/600m)

Auxis_thazard ( !

1

0 1 2

4

6 Km

Centre for Environmental Management CQU - 24

Port Curtis Demersal Fish

³ ( !

Total abundance (n/600m)

Carangoides_caeruleopinnatus ( !

1

( !

0 1 2

³

4

6 Km

4

6 Km

( !

( !

Total abundance (n/600m)

Caranx_ignobilis ( !

( !

1 2

0 1 2

Centre for Environmental Management CQU - 25

Port Curtis Demersal Fish

³ ( !

Total abundance (n/600m)

Chaetodon_tricinctus ( !

2

0 1 2

4

6 Km

³ ( !

Total abundance (n/600m)

Choerodon_cephalotes ( !

1

0 1 2

4

6 Km

Centre for Environmental Management CQU - 26

Port Curtis Demersal Fish

³

( ! ( !

Total abundance (n/600m)

Dasyatis_fluviorum ( !

1

³

0 1 2

4

6 Km

0 1 2

4

6 Km

( !

Total abundance (n/600m)

Dasyatis_kuhlii ( !

1

Centre for Environmental Management CQU - 27

Port Curtis Demersal Fish

³ ( !

Total abundance (n/600m)

Diodor_nichthemerus 1

( !

0 1 2

4

6 Km

( !

³ ( !

( !

! (

( !

( ! ( !

( !

Total abundance (n/600m)

Drepane_punctata 1 2 ! ( 3-4 ( !

( !

0 1 2

4

6 Km

Centre for Environmental Management CQU - 28

Port Curtis Demersal Fish

³ ( !

Total abundance (n/600m)

Eleutheronema_tetradactylum ( !

1

³

0 1 2

4

6 Km

0 1 2

4

6 Km

( !

Total abundance (n/600m)

Epinephelus_coioides ( !

1

Centre for Environmental Management CQU - 29

Port Curtis Demersal Fish

³

( !

( !

! ( ( !

( ! ( !

( !

( !

Total abundance (n/600m)

Gerres_filamentosus 1 ( ! 2-3 ! 4 - 12 (

( !

( !

( !

0 1 2

4

6 Km

³ ( !

Total abundance (n/600m)

Gerres_oyena ( !

2

0 1 2

4

6 Km

Centre for Environmental Management CQU - 30

Port Curtis Demersal Fish

! ( ( !

³

( !

! ( ( !

! (

! (

( !

( ! ( ! ( !

( !

( !

( ! ( !

( !

( ! ( !

! ( ( !

! (

( !

( !

( !

! (

Total abundance (n/600m)

Gerres_subfasciatus 1 2-3 ! ( 4 - 11 ( !

( !

0 1 2

4

6 Km

0 1 2

4

6 Km

³ ( !

Total abundance (n/600m)

Gymnura_australis ( !

1

Centre for Environmental Management CQU - 31

Port Curtis Demersal Fish

( ! ( !

( ! ( !

³

! ( ! (

( ! ( !

! ( ( !

! (

! ( ! (

( !

( !

( !

! (

! (

! (

! ( ( !

( !

( !

! (

( !

( !

( !

! (

( !

( !

! (

( !

( !

Total abundance (n/600m) ( !

Herklostichthys_castelnaui

! (

1-2 3-5 ! ( 6 - 12 ( 13 - 25 ! ! ( 26 - 55 ( !

( !

³

( ! 0 1 2

4

6 Km

0 1 2

4

6 Km

( !

Total abundance (n/600m)

Himantura_uarnak ( !

1

Centre for Environmental Management CQU - 32

Port Curtis Demersal Fish

³ ( !

( !

Total abundance (n/600m)

Hippocampus_sp1 ( !

1

0 1 2

4

6 Km

0 1 2

4

6 Km

³ ( !

Total abundance (n/600m)

Hyperolophus_translucidus ( !

2

Centre for Environmental Management CQU - 33

Port Curtis Demersal Fish

³

( !

Total abundance (n/600m)

Hyporhamphus_australis ( !

1

0 1 2

4

6 Km

³ ( !

Total abundance (n/600m)

Hyporhamphus_quoyi ( !

6

0 1 2

4

6 Km

Centre for Environmental Management CQU - 34

Port Curtis Demersal Fish

³

( !

Total abundance (n/600m)

Istigobius_hoesei 3

( !

0 1 2

4

6 Km

³ ( !

( !

( !

Total abundance (n/600m)

Lagocephalus_sceleratus ( !

1

0 1 2

4

6 Km

Centre for Environmental Management CQU - 35

Port Curtis Demersal Fish

³ ( !

( !

Total abundance (n/600m)

Leiognathus_bindus ( !

2

0 1 2

4

6 Km

( ! ! ( ( ! ( ! ( !

³

( !

( !

( ! ( ! ( !

( !

( ! ( !

( ! ( !

! (

( !

! (

! ( ( !

( !

( !

( !

( !

! (

( !

( !

! ( ! ( ( !

( !

( !

( !

( !

( !

( !

( !

( ! ( !

( !

( ! ! ( ( !

! (

( !

( !

! (

( !

! (

( ! ( !

Total abundance (n/600m) ( !

Leiognathus_decora 1-7 8 - 18 ! ( 19 - 32 ( 33 - 56 ! ! ( 57 - 83 ( !

( !

( !

0 1 2

4

6 Km

Centre for Environmental Management CQU - 36

Port Curtis Demersal Fish

³

( !

( !

Total abundance (n/600m)

Leiognathus_sp1 ( !

( !

1 2

³

0 1 2

4

6 Km

0 1 2

4

6 Km

( !

( !

( !

Total abundance (n/600m)

Leognathus_sp2 ( !

( !

1 2-4

Centre for Environmental Management CQU - 37

Port Curtis Demersal Fish

³

( !

( !

Total abundance (n/600m)

Leiognathus_sp3 ( ! ( !

( !

1 2-3

0 1 2

4

6 Km

4

6 Km

( ! ( !

³

( !

( ! ( !

( !

! (

( !

( !

( !

( !

( ! ( !

( !

Total abundance (n/600m)

Leptobrama_mulleri 1 2-3 ! ( 4-6 ( !

( !

0 1 2

Centre for Environmental Management CQU - 38

Port Curtis Demersal Fish

³

( !

! (

Total abundance (n/600m)

Lethrinus_variegatus ( !

! (

2 3-4

0 1 2

4

6 Km

( !

³

( ! ( ! ( !

( !

( !

! (

Total abundance (n/600m)

Liza_dussumieri 1-2 3-5 ! ( 6-8

( !

( !

( !

0 1 2

4

6 Km

Centre for Environmental Management CQU - 39

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( !

³

( !

Total abundance (n/600m)

Lutjanus_russelli 1 2

( !

( !

³

0 1 2

4

6 Km

4

6 Km

( ! ( !

( !

( !

( !

Total abundance (n/600m)

Marilyna_pleurosticta ( !

( !

1 2

0 1 2

Centre for Environmental Management CQU - 40

Port Curtis Demersal Fish

³

( ! ( ! ( !

( !

( !

! (

( ! ( ! ( !

( !

( !

! ( ( !

( ! ( !

Total abundance (n/600m)

Megalaspis_cordyla 1-2 3-6 ! ( 7-8 ( 9 - 22 ! ( !

( !

³

0 1 2

4

6 Km

4

6 Km

( !

( !

( !

Total abundance (n/600m)

Meuschenia_trachlepis ( !

( !

1 2

0 1 2

Centre for Environmental Management CQU - 41

Port Curtis Demersal Fish

³ ( !

Total abundance (n/600m)

Meuschenia_sp1 ( !

1

0 1 2

4

6 Km

³ ( !

( ! Total abundance (n/600m)

Meuschenia_sp2 ( !

1 ( 2 - 39 !

( !

( !

0 1 2

4

6 Km

Centre for Environmental Management CQU - 42

Port Curtis Demersal Fish

³ Total abundance (n/600m) ( !

Meuschenia_sp3 ( !

1

0 1 2

4

( !

6 Km

³ ( !

Total abundance (n/600m)

Meuschenia_sp4 ( !

1

0 1 2

4

6 Km

Centre for Environmental Management CQU - 43

Port Curtis Demersal Fish

³

( !

( !

( !

Total abundance (n/600m)

Monodactylus_argenteus ( !

( !

1 2

³

0 1 2

4

6 Km

0 1 2

4

6 Km

! ( ( ! ( !

( !

! ( ( !

( !

( !

( !

( !

( !

( !

Total abundance (n/600m)

Mugil_georgii 1-3 4-6 ! ( 7 - 11 ( 12 - 19 ! ( !

( !

Centre for Environmental Management CQU - 44

Port Curtis Demersal Fish

³

( !

Total abundance (n/600m)

Muraenesox_cinevus 1

( !

0 1 2

³

4

6 Km

( !

( !

( !

( ! ( !

( !

( !

( !

Total abundance (n/600m) ( !

Nematalosa_come ( !

( !

1-2 3-9

0 1 2

4

6 Km

Centre for Environmental Management CQU - 45

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³

( !

( !

Total abundance (n/600m)

Nemipterus_theodorei ( !

1

0 1 2

4

6 Km

³ Total abundance (n/600m) ( !

Parachaetodon_ocellatus ( !

2

0 1 2

4

6 Km

Centre for Environmental Management CQU - 46

Port Curtis Demersal Fish

! (

³

! (

( !

! ( ! ( ( ! ( !

( !

( !

Total abundance (n/600m)

( !

Pelates_quadrilineatus 1-2 3-8 ( 9 - 17 ! ( !

! (

0 1 2

4

6 Km

³ ( !

( ! Total abundance (n/600m)

Pelates_sexlineatus ( !

1

( 2 - 16 !

0 1 2

4

6 Km

Centre for Environmental Management CQU - 47

Port Curtis Demersal Fish

³ ( !

Total abundance (n/600m)

Pentapodus_paradiseus ( !

1

³

0 1 2

4

6 Km

0 1 2

4

6 Km

( !

Total abundance (n/600m)

Periopthalmus_koelreuteri ( !

1

Centre for Environmental Management CQU - 48

Port Curtis Demersal Fish

³

( !

Total abundance (n/600m)

Platycephalus_arenarius ( !

1

0 1 2

4

6 Km

4

6 Km

( !

³

( !

( ! ( !

Total abundance (n/600m)

Platycephalus_indicus ( !

( !

1 2-3

0 1 2

Centre for Environmental Management CQU - 49

Port Curtis Demersal Fish

³

( !

( ! ( !

( !

Total abundance (n/600m)

Polydactylus_multiradiatus 1 2

( !

( !

0 1 2

4

6 Km

4

6 Km

( !

( !

! (

³

( ! ( !

( ! ( !

( !

( ! ( !

( !

( !

( !

( !

Total abundance (n/600m)

Pomadasys_kaakan 1-2 3-6 ! ( 7 - 11 ( 12 - 35 ! ( !

( !

0 1 2

Centre for Environmental Management CQU - 50

Port Curtis Demersal Fish

³ ( !

Total abundance (n/600m)

Pseudorhombus_argent ( !

1

³

0 1 2

4

6 Km

( ! ( !

( !

( !

( !

( !

( !

( !

( !

( !

( !

( ! ( ! ( !

( !

( !

( !

Total abundance (n/600m) ( !

Pseudorhombus_arsius ( !

( !

1 2

0 1 2

4

6 Km

Centre for Environmental Management CQU - 51

Port Curtis Demersal Fish

( !

³ Total abundance (n/600m)

Rachycentron_canadus ( !

4

0 1 2

³

4

6 Km

4

6 Km

( !

Total abundance (n/600m)

Rhadbosargus_sarba ( !

3

0 1 2

Centre for Environmental Management CQU - 52

Port Curtis Demersal Fish

( !

³

( !

( ! ( !

( !

( !

( !

( ! ( !

! (

( ! ( ! ( !

( !

( !

( !

( !

( !

! (

( !

( !

( !

( !

( ! ( ! ( !

( ! ( !

Total abundance (n/600m)

Saurida_undosquamis ( !

1-2 ( ! 3-5 ! 6-9 ( ( !

0 1 2

4

6 Km

( !

( !

³

( !

( !

( !

( !

( !

( !

Total abundance (n/600m)

Scomberoides_commersonianus ( !

( !

1 2

0 1 2

4

6 Km

Centre for Environmental Management CQU - 53

Port Curtis Demersal Fish

³ ( !

Total abundance (n/600m)

Scomberomorus_queenslandicus 1

( !

0 1 2

4

6 Km

³ Total abundance (n/600m) ( !

Scorpaenopsis_sp ( !

1

0 1 2

4

6 Km

Centre for Environmental Management CQU - 54

Port Curtis Demersal Fish

³ Total abundance (n/600m)

Selaroides_leptolepis

( !

2 ! ( 3 - 11 ( !

³

! (

( ! 0 1 2

4

6 Km

0 1 2

4

6 Km

( !

Total abundance (n/600m)

Selenotoca_multifasciata ( !

3

Centre for Environmental Management CQU - 55

Port Curtis Demersal Fish

( !

³

( !

( !

( !

! ( Total abundance (n/600m)

( !

Siganus_rivulatus

( !

1 2 ( 3-8 ! ! ( 9 - 66

( !

( !

( !

³

( !

0 1 2

4

6 Km

0 1 2

4

6 Km

( !

( !

Total abundance (n/600m)

Sillago_analis ( !

( !

1 2

Centre for Environmental Management CQU - 56

Port Curtis Demersal Fish

³ ( !

( !

Total abundance (n/600m)

Sillago_ciliata ( !

1 2

( !

( !

³

0 1 2

4

6 Km

( !

( !

( !

( !

( ! ( !

( !

( !

Total abundance (n/600m)

Sillago_maculata ( !

( !

1 2

( !

0 1 2

4

6 Km

( !

Centre for Environmental Management CQU - 57

Port Curtis Demersal Fish

³ ( !

( !

( !

Total abundance (n/600m)

Sphyraena_obtusata ( !

( !

1 2

( !

0 1 2

³

4

6 Km

( !

( !

Total abundance (n/600m)

Sygnathidae_sp ( !

( !

1 2

0 1 2

4

6 Km

Centre for Environmental Management CQU - 58

Port Curtis Demersal Fish

( !

³

( !

( ! ( !

( !

( !

Total abundance (n/600m)

Tetractenos_hamiltoni ( !

1 2-3

( !

( !

³

0 1 2

4

6 Km

( !

( !

Total abundance (n/600m)

Thryssa_aestuaria ( !

( !

1 2-9

0 1 2

4

6 Km

Centre for Environmental Management CQU - 59

Port Curtis Demersal Fish

³

( !

! ( ( !

! (

( !

( !

Total abundance (n/600m)

Trepaon_jarbua 1 2 ! ( 3-5 ( !

( !

0 1 2

4

6 Km

4

6 Km

( !

³

( ! ( !

( !

( !

( !

Total abundance (n/600m)

Triacanthus_brevirostris ( !

( !

1 2

0 1 2

Centre for Environmental Management CQU - 60

Port Curtis Demersal Fish

³

( !

( !

( !

( !

( !

( !

( ! ( !

( ! ( !

( !

( !

( !

( ! ( !

( !

( !

Total abundance (n/600m)

Tripodichthys_angustifrons ( ! ( !

( !

1 2-4

³

0 1 2

4

6 Km

( !

( !

Total abundance (n/600m)

Tylosurus_crocodilus ( !

1

0 1 2

4

6 Km

Centre for Environmental Management CQU - 61

Port Curtis Demersal Fish

Appendix 2. Predictive maps of Port Curtis showing distributions of key water quality parameters including: (a) temperature, (b) salinity, (c) ph, (d) dissolved oxygen content, (e) turbidity and (f) total chlorophyll concentration. Note all estimates presented are derived from winter 2002 depth-profiled averages at 149 separate sampling stations (small filled circles). (b) (a) !

! !

!

Temperature (ºC)

!

!

Salinity (ppt)

!

!

4. 7

1. 3

!

!

-2

9. 6

-2

!

21

.3

8. 8

-1 19

.6

-1

.4

18

18

.8

8. 2

-1

-1

17

16

!

.8

.9

-1

!

!

8. 4

!

7. 8

!

.2

!

18

!

!

35 .7 9 36 - 3 .0 6. 0 1 36 - 3 1 .1 6.1 8 8 36 - 3 6 .3 0 .30 36 - 3 .3 6. 3 9 36 - 3 9 .4 6.4 6 36 - 3 6 .5 6.5 5 5 36 - 3 .6 6.6 7 36 - 3 7 6 .8 4 .8 37 - 3 4 .0 7.0 6 -3 6 7. 36

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(c)

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(d)

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pH

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!

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81 ! !

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Dissolved Oxygen (mg/L)

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Secci Depth (m)

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Centre for Environmental Management CQU - 62

Port Curtis Demersal Fish

Appendix 2 cont. Predictive maps of Port Curtis benthos showing distributions of: (a) infaunal richness, (b) infaunal abundance, (c) sediment size-structure, and (d) organic carbon content. Note estimates presented are derived from replicate samples (n=3) collected at 149 separate sampling stations (small filled circles). (a)

(b) !

! !

Organic Carbon (%)

!

Median Sediment (mm)

!

!

!

!

-2

19

2. 87

.7 1

.1 9

-1

!

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71 0.

0.

!

1.

.2 6

.5 0

-0

50

.1 7

-0

-0

-0

26

17 0.

11

08

0.

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0.

.0 8

-0

-0

.0 6

05

04

0.

0.

!

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-0

-0

!

.1 1

!

.0 5

!

06

!

0.

!

!

5. 5. 26 26 6. 6.8 89 9 8. 8.0 00 0 8. 8. 75 75 9. - 9 26 .2 10 - 1 6 .0 0. 0 1 11 - 1 1 .1 1.1 2 12 - 1 2 .7 2.7 4 15 - 1 4 .1 5.1 4 -1 4 8. 66

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(c)

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Species Richness (N/0.3m²)

Species Abundance (N/0.3m²)

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48 8 91 91 18 18 9 9 40 406 6 -8 90

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Centre for Environmental Management CQU - 63