The Role of Shallow Seagrass Meadows As Habitat for Fish
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5-tr-€ The Role of Shallow Seagrass Meadows as Habitat for Fish by Roderick Martin Connolly Thesis submitted for the degree of Doctor of Philosophy m Department of Zoology University of Adelaide August 1994 {wa"rleot gqb Contents List of figures.. Summary D( Declaration )tr Acknowledgements.... .......)í1 2 Comparison of fish catches from a buoyant pop net and a beach seine net 9 2.4 Discussion t6 3 Comparison of fish assemblages from eelgrass and unvegetated areas .......23 3.1 Introduction 23 3.3 Results 29 4 Diet of Síllagínodes punctata ......50 4.1 Introduction .........50 4.2 Methods 53 57 59 5 Comparison of epifauna from eelgrass and unvegetated habitats 78 5.1 Inroduction 78 79 82 5.4 Discussion.. 86 6 Removal of seagrass canopy: effects on small fish........... 100 6.1 Introduction 100 6.2 Methods 102 7 Removal of seagrass canopy: effects on epifauna ll3 7.4 Discussion t2l 8 Altering seagrass canopy height: effects on epifauna of shallow Mediterranean embayments..... ................134 8.1 Introduction ....... 134 8.2 Methods............... ...............135 8.3 Results............... ................. 138 8.4 Discussion.......... .................145 9 The role of seagrass as preferred habitat for juvenile Sillaginodes punctata: habitat selection or feeding?..................... t7l 9.1 Introduction t7t 9.2 Methods r73 176 9.4 Discussion r79 l0 Concludingdiscussion 184 Appendix Published and submitted manuscripts 191 A.1 Comparison of fish catches from a buoyant pop net and a beach seine net in a shallow seagrass habitat. Mar. Ecol. Prog. Ser. 109: 305-309. Chapter 2 r92 4.2 A comparison of fish assemblages from seagrass and unvegetated areas of a southern Australian esturiry. Aust. J. Mar. Freshwat. Res. 45 (in press). Chapter 3 r97 4.3 Removal of seagrass canopy: effects on small fish and their prey. J. Exp. Mar. Biol. Ecol. (in press). Chapter 6 215 4.4 The effecs of removal of seagrass canopy on assemblages of small, motile invertebrates. Mar. Ecol. Prog. Ser. (submitted). ChapterT .......229 4.5 The effects of altering seagrass canopy height on small, motile invertebrates of shallow Mediterr¿nean embayments. Mar.Ecol.(Napoli).(submitted).Chapter8............... ..........244 4.6 The role of seagrass as preferred habitat for juvenile SíIlaginodes punctata (Cuv. & Val.Xsillaginidae, Pisces): habitat selection or feeding? J. Exp. Mar. Biol. EcoI.l80:39-47. Chapter 9 269 4.7 Improved experimental procedure in ecology: what to set, what to forget. (unpublished)............ ...................278 Bibliography 293 List of tables Table 2.1 Number of individuals of main species caught during study 2.2 Comparisons of pop net and seine net catches 2.3 Comparison of lengths of fish from pop net and seine net 2.4 Comparisons of catches from three pulls of collecting net in pop nets 3.1 Number of individuals of the ten most cornmon species (all dates combined) in eelgrass and unvegetated habitat 3.2 Results of Mann-Whitney U-tests for differences in abundance between eelgrass and unvegetated sites 3.3 Comparisons of median length of Sillaginodes punctata at eelgrass and unvegetated sites 3.4 Multiva¡iate comparisons of eelgrass and unvegetated assemblages 3.5 Water temperature and salinity at eelgrass and unvegetated sites separately at each sampling perid 4.L Siltaginodes punctataexamined for stomach contents: number of sites at which S. punctatawere caught,.and number of fish examined 4.2 Size ranges (mm) of prey types matching mesh sizes 4.3 Numbers of fish with no food in stomach, and fullness indices 5.1 List of taxa into which animals were grouped in 5.2 Mean ash-free dry weights (mg) of diffcrent forms of epifaunal invertebrates each sieve size-class 5.3 Results of ANOSIM comparisons between epifaunal assemblages from eelgrass and unvegetated habitats 5.4 Results of two-way ANOSM comparisons of epifaunal assemblages benveen habitats and between the two waterways, Barker Inlet and Port River 5.5 Results of two-way ANOSIM comparisons of epifaunal assemblages between habitats and amongst sampling periods on data from all periods together lv 5.6 Comparisons of abundances and biomasses of total epifauna and key taxa from eelgrass and unvegetated habitats 5.7 Comparisons of abundances and biomasses of cumaceans from eelgrass and unvegetated habitats 5.8 Total epifaunal production and total crustacean production (pgldayl0.0625m2) in eelgrass and unvegetated habitats at each sampling perid 6.1 Fish abundance by habitat 7.1 List of taxa into which animals were grouped 7 .2 Results of ANOSIM comparisons amongst epifaunal assemblages 7 .3 Abundances of total epifauna and key taxa in each habitat 7.4 Biomasses of total epifauna and key taxa in each habitat 7.5 Total epifaunal and crustacean production in each habitat 8.1 List of taxa into which animals were grouped 8.2 Results of ANOSIM comparisons amongst epifaunal assemblages from Embiez 8.3 Abundances of total epifauna and key taxa in each habitat at Embiez 8.4 Biomasses (AFDTW h -g) of total epifauna and key taxa in each habitat at Embiez 8.5 Total epifaunal production and total crustacean production (pglday/0.0625m2)n each habitat atEmbiez 8.6 Results of ANOSIM comparisons ¿rmongst epifaunal assemblages from Diana-Night 8.7 Abundances of total epifauna and key taxa in each habitat at Diana-Night 8.8 Biomasses (AFDW in mg) of total epifauna and key ta"ra in each habitat at Diana-Night 8.9 Total epifaunal production and total crustacean production (pglday/O.O625m2) n each habitat at Diana-Night 8.10 Results of ANOSIM comparisons anþngst epifaunal assemblages from Diana-Day 8.11 Abundances of total epifauna and key taxa in each habitat at Diana-Day v 8.12 Biomass (AFDW in mg) of total epifauna and key taxa in each habitat at Diana-Day 8.13 Total epifaunal production and total crustacean production (¡rg/day/0.0625m2)n each habitat at Diana-Day 8.14 Results of two-way crossed ANOSIM comparisons amongst epifaunal assemblages of locations and habitats 8.15 Total epifaunal abundance and production comparisons across locations 9.1 Predicted results from the two models, habitat selection and simple feeding vl List of figures Figure 2.1 Pop net design 3.1 Map of survey region and habitat locations 3.2 Total number of species from eelgrass and unvegetated habitats caught at each sampling perid 3.3 Mean number of fish per site from eelgrass and unvegetated habitas 3.4 l-ength - frequency of Sillaginodes punctata at each sampling period 3.5 Twedimensional MDS ordination plos of fish assemblages showing habitat differences 3.6 Overlay of distance from site to open water onto MDS plots of Fig. 3.5 3.7 Twodimensional MDS plot of fish assemblages over all periods. 4.1 I-ength - dry weight relationship for juvenile Sillagínodes punctata 4.2 Plot of residuals of linea¡ regression for length - dry weight of Sillaginodes punctata shown in Fig. 4.1 4.3 Length - wet weight relationship for juvenile Sillaginodes punctata 4.4 Plot of residuals of linea¡ regression for length - wet weight of Sillaginodes punctotct shown in Fig. 4.3 4.5 Stomach contents of Sillaginodes punctara in January 1990 by habitat 4.6 Stomach contents of Sillaginodes punctara in April 1990 by habitat 4.7 Stomach contents of Siilaginodes punctarø in August 1990 by habitat 4.8 Stomach contents of Sillaginodes punctar¿ in October 1990 by habitat 4.9 Stomach contents of Sillaginodes punctara in February 1991 by habitat 4.10 Stomach contents of .Si//ag inodes punctata in June 1991 by habitat 4.lI Stomach contents of Sillaginodes punctar¿ collected at night in June l99l 4.12 Stomach contents of Si/løginodes punctata in October 1991 by habitat vll 4.13 Stomach contents of Si//aginodes punctata at night in October 1991 5.1 Two-dimensional MDS plots of epifaunal assemblages based on abundance 5.2 Two-dimensional MDS plots of epifaunal assemblages based on biomass 5.3 Overlay of distance from site to open water onto MDS plots for August 1990 5.4 Twedimensional MDS plot of epifaunal assemblages over all periods combined 6.1 Twodimensional MDS plot of fish assemblages, all species included 6.2 Two-dimensional MDS plot of fish assemblages, pelagic species excluded for the three 7 .I Two-dimensional MDS plot of epifaunal assemblages, averaged samples within each Patch 7.1 7 .2 Overlay of position of patch along shore onto MDS plots in Fig. j.j Relationship of total number of fish and total number of non-pelagic frsh (frsh/net) to epifaunal production (pglday/0.0625ß*) and seagrass cover (mz leaf arealrrÊ sediment area) by habitat 8.1 Map showing location of sites (Mediterr¿nean Sea) g.Z Two.dimensional MDS plots of epifaunal assemblages from Embiez based on abundance 8.3 Twedimensional MDS plos of epifaunal assemblages from Embiez based on biomass 8.4 Twedimensional MDS plots of epifaunal assemblages from Diana - Night based on abundance 8.5 Twodimensional MDS plots of epifaunal assemblages from Diana - Night based on biomass 8.6 Two-dimensional MDS plots of epifaunal assemblages from Diana - Day based on abundance 8.7 Two-dimensional MDS plots of epifaunal assemblages fromDiana - Day based on biomass 8.8 Two-dimensional MDS plots of epifaunal assemblages from all locations 9.1 Mean number of trials in which most flsh were in eelgrass habitat ull Summary Shallow seagrass meadows a¡e considered important as habitat for small f,rsh, including juveniles of economically important species. Surveys of eelgrass (Zostera) and unvegetated patches in the Ba¡ker Inlet region, a shallow, sheltered, ma¡ine-dominated estuary in South Australia, showed that there were more fish over eelgrass than over adjacent unvegetated habitat.