ABUNDANCE, BIOMASS AND ESTIMATED PRODUCTION OF INVERTEBRATE FAUNA ASSOCIATED WITH SEAGRASS, HETEROZOSTERA TASMANICA, IN SWAN BAY AND AN ADJACENT AREA OF PORT PHILLIP BAY, VICTORIA Fiona L. Bird'’* & Gregory P. Jenkins* 'Marine and Freshwater Research Institute, and Zoology Department, University of Melbourne, PO Box 138, Quecnscliff, Victoria 3225, Australia ^Present address: Crustacea laboratory. Museum of Victoria, 71 Victoria Cresent, Abbotsford, Victoria 3067, Australia Bird, F. L. & Jenkins, G. P., 1999:07:31. Abundance, biomass and c.stimated production of invertebrate fauna associated with seagrass, Heierozostera uisntanica. in Swan Bay and an adjacent area of Port Phillip Bay. Victoria. Proceedings of the Royal Society of Victoria 111(1): 1-13. ISSN 0035-9211. Abundance, biomass and production of invertebrates were compared between Swan Bay, a seagrass system in a low energy sheltered environment, and an adjacent area of Port Phillip Bay, which had greater exposure to wave action and tidal currents. Seagrass biomass and abundances of macrofauna were not significanlly different between the two areas, but organic content of sediments, biomass and production of macrofauna were significantly higher in Swan Bay. Faunal composition was markedly different, with Port Phillip Bay sites dominated by gammaridcan and eaprellidean amphipods, while a variety of often larger macrofaunal groups characterised Swan Bay sites. In contrast to macrofauna, the epibenihic meiofauna, dominated by harpaclicoid copepods, was significantly more abundant in Swan Bay. The results support the contention that sheltered .seagrass areas with elevated organic content of sediment can provide increased production of food for juvenile fish. SEAGRASS beds are known to be highly environment with less accumulated debris. This produetive environments (Hillman et al. 1989), area of Port Phillip Bay is adjacent to strong tidal supporting diverse and extensive communities of currents associated with exchange of water through invertebrates. Studies comparing seagrass and un- the narrow entrance at Port Phillip Heads (Black vegetated habitats have consistently found that ct al. 1993). Shaw & Jenkins (1992) showed that vegetated areas sustain greater abundance and unvegetated areas of Swan Bay had finer sediments diversity of benthic invertebrates (Rainer & with higher organic content than unvegetated areas Fitzhardinge 1981; Poore 1982; Lewis 1984; of the adjacent coast of Port Phillip Bay. Jenkins Howard ct al. 1989; Edgar 1990a) and fish (Bell et al. (in review) found that fish communities in & Pollard 1989; Lubbers ct al. 1990; Blaber et al. these scagra.ss beds were dominated by juveniles, 1992). Seagrass is known to trap organic debris and the abundance and biomass of fishes in Swan and detritus (Norwcll & Jumars 1984), with the Bay was significantly higher than in the adjacent organic material thought to enhance the secondary coast of Port Phillip Bay. Juvenile fishes at these production (Pearson & Rosenberg 1978; Mann sites consumed small invertebrates, dominated by 1988; Spies et al. 1988). From this observation it Crustacea (Bird 1990). Edgar & Shaw (1993) found is hypothesi.sed that exposed habitats accumulate that fish and invertebrate production in Western less detritus than sheltered habitats and would Australia was much higher in sheltered than in con.scqucntly have lower overall production levels. exposed sites, and postulated that fishes were Edgar (1990a) showed by comparing a sheltered attracted to seagrass beds with high production to an exposed area, that organic enrichment of of invertebrates. We postulated that the more sediments (by debris and detritus) enhances sheltered environment of Swan Bay would result invertebrate production. in correspondingly higher organic content of Swan Bay, Australia, has extensive beds of the sediments and elevated production of invertebrates. subtidal seagrass, Hetcrozostera lasmaiiica, in a In this paper, we examine this hypothesis by low energy, sheltered environment with large comparing organic content of sediments, meiofauna accumulations of debris and drilt algae. The abundance, and macrofaunal abundance, biomass adjacent coast of Port Phillip Bay has smaller, and production in Swan Bay and an adjacent more discrete H. lasmanica beds in a higher energy area of Port Phillip Bay. FIONA L. BIRD AND GREGORY P. JENKINS 2 INVERTEBRATE FAUNA ASSOCIATED WITH SEAGRASS, HETEROZOSTERA TASMANICA 3 METHODS for 48 hours at 60°C. Organic content of the sediment was estimated from the small-core Study area samples. Total contents of each sample was The study area included Swan Bay and an dried for 48 hours at 60°C and then burned in adjacent area in Port Phillip Bay, situated on a combustion oven for two hours at 500°C. the Bellarine Peninsula, Victoria (Fig. 1). Tidal The change in weight due to the burning was currents in Swan Bay are less than 0.05 ms"' recorded, and this change was considered pro¬ compared with currents between 0.1 and 0.5 m.s”' portional to the organic content. on the adjacent coast of Port Phillip Bay (Black et al. 1993). Moreover, maximum wind fetch is Data analysis less than 10 km in Swan Bay, but greater than 40 km on the adjacent coast of Port Phillip Bay Analysis of macrofauna was restricted to in¬ (Fig. 1). Two sites were sampled in Swan Bay; vertebrates passing through the 5.6 mm sieve Tin Can and North Jetty, and two sites were because larger animals were not consumed by sampled in Port Phillip Bay; St Leonards and juvenile fish in the study area (Bird 1990), and Beacon. Samples were taken .seasonally over two in the case of mobile animals, would have a days beginning on 1 May, 1 August, 29 October greater ability to avoid the corcr. Core data were 1990 and 20 January 1991. One additional site initially checked for homogeneity of variances in each area. Mid Jetty and Edwards Point, was using Cochran’s test, and if necessary log(x-^l) included in October for sampling of organic or square-root transformed. The data were then content of sediment and meiofauna abundance analysed using an analysis of variance with two (Fig- 1). bays, two sites nested within each bay, five replicates at each site, on each of four dates. Time and bay were treated as fixed factors, while Field methods sites were considered to be random factors. The invertebrate community was sampled using The biomass of macrofaunal groups was esti¬ two sizes of corers. A 90 mm diameter PVC mated from abundance and previously determined core, was used to take five haphazardly placed mean weights of these groups in individual replicates, from each of the four sites on all sieve-size classes (Edgar 1990b). The production four dates, to estimate abundance, biomass and of a specific size-class and type of cpifaunal production of macrofauna. A 32 mm diameter invertebrate was calculated using the equation corcr was used to obtain eight replicate samples developed and tested by Edgar (1990b) which at each of the six sites in October. Five replicates relates daily macrobenlhic production (pg/d) to from each site were used for analysis of organic ash-free dry weight (pg) and water temperature content and the remaining three for analysis of (°C). The estimated production of a specific macro¬ meiofauna. Both of the corers were pushed into faunal group in a given size class was multiplied the sediment to a depth of 10 cm to extract a by the number in that size class and summed for constant volume of sediment. The samples were all size classes. Total daily production was preserved in 99% ethanol mixed with Rose Bengal estimated from the combined produetion values to make the animals more visible while sorting. for each macrofaunal category. Laboratory methods RE,SULTS Each large-core sample was washed through a series of nested sieves of 5.6, 4.0, 2.8, 2.0, 1.4, 1.0, 0.71 and 0.5 mm mesh. .Small cores were Bay and site characteristics washed through a sieve series of 0.5, 0.355, 0.25, Water temperatures tended to greater extremes in 0.18, 0.125 and 0.063 mm mesh. Each category Swan Bay than in Port Phillip Bay (Table 1). of invertebrate in each size class was counted. Temperatures conform to those found by Jenkins Seagrass biomass was estimated by drying the (1986) except in October when we encountered seagrass material remaining in the largest sieve an unseasonally warm sampling day. When Fig. 1. Location of the sampling sites in Swan Bay and an adjacent area of Port Phillip Bay. 4 FIONA L. BIRD AND GREGORY P. JENKINS calculating production estimates for that date, Source Mean square DF F-ratio P the temperature 170°C (from Jenkins 1986) was used for all sites. No significant difference in Date 18.727 3 1.681 0.269 seagrass biomass was found between bays Bay 18.075 1 3.342 0.209 (Table 2). The seagrass biomass values ranged Date*Bay 9.902 3 0.889 0.499 from 4 to 10 g per core (628 to 1572 g.m~^) SB sites 10.816 1 1.405 0.240 (Fig. 2). More detritus, seagrass debris and drift PPB sites 0.001 1 0.000 0.993 algae was observed inside Swan Bay than at SB silcs*Dalc 5.164 3 0.671 0.573 the sites in Port Phillip Bay. Swan Bay sediment PPB sites*Date 17.123 3 2.225 0,094 was shown to have a greater organic content Error 7.696 64 than sediment in Port Phillip Bay (Table 3). Table 2. Nested analysis of variance comparing the A difference in organic content was also found estimated seagrass biomass in Swan Bay (SB) and an between the Swan Bay sites (Fig. 3). adjacent area of Port Phillip Bay (PPB). DF—degrees of freedom; P—probability. Sites Date May August October January Source Mean square DF F-ratio P 1990 1990 1990 1991 Bay 0.228 1 8.545 0.043 North Jetty 15.5 12.1 25.0 22.3 SB sites 0.051 2 7,385 0.005 Tin Can 14.7 10.9 25.0 21.4 PPB sites 0.002 2 0.306 0.741 St Leonards 16.4 11.5 18.9 20.2 Error 0.007 16 Beacon 17.0 11.5 18.2 19.8 Table 3. Nested analysis of variance comparing the Table 1. Water temperature measurements taken at four organic content of sediments in Swan Bay (SB) and an sites in Swan Bay and an adjacent area of Port Phillip adjacent area of Port Phillip Bay (PPB). DF—degrees Bay (°C). of freedom; P—probability. Fig. 2. Mean seagrass biomass at sites in Swan Bay and Port Phillip Bay on four sampling dates. Solid, 30 May 1990; open, 31 August 1990; diagonal, 30 October 1990; stipples, 31 January 1991. Error bars are standard error. North Jetty Tincan St Leonards Beacon Swan Bay | Port Phillip Bay Site Macrofaunal communities No significant difference in macrofaunal abun¬ biomass of macrofauna was highcsl for May dance was found between bays (Table 4, Fig. 4). and August samples in Swan Bay (Fig. 4). There Swan Bay had a significantly higher estimated was also a significant interaction between biomass of maerofauna than Port Phillip Bay abundances at sites in Swan Bay with date (Table 5). Macrofauna biomass also varied signi¬ (Table 5). Estimated production of macrofauna for ficantly amongst dates (Table 5). In general. Swan Bay was also significantly higher than for INVERTEBRATE FAUNA ASSOCIATED WITH SEAGRASS, HETEROZOSTERA TASMANICA 5 Fig. 3. Mean percentage of organic content in sediments at sites in Swan Bay and Port Phillip Bay. North J—North Jetty; Mid J—Mid Jetty; St Leo—St Leonards. Error bars are standard error. North J Tincan Midi St Leo Beacon Edwards Pt Swan Bay Port Phillip Bay Site Source Mean square DF F-ratio P Source Mean square DF F-ratio P Date 19173.7 3 2.338 0.173 Date 0.046 3 1.578 0.290 Bay 52173.1 1 5.600 0.142 Bay 2.126 1 55.900 0.017 Date* Bay 25052.4 3 3.055 0.113 Date* Bay 0.093 3 3.204 0.105 SB sites 270.4 1 0.041 0.839 SB sites 0.044 1 1.972 0.165 PPB sites 18361.2 1 2.814 0.098 PPB sites 0.032 1 1,435 0.235 SB sites*Date 14458.5 3 2.214 0.095 SB sites*Date 0.048 3 2.160 0.101 PPB sitcs*Date 1944.0 3 0.298 0.827 PPB sites*Date 0.010 3 0.432 0.731 Error 65.30.4 64 Error 0.022 64 Table 4. Nested analysis of variance comparing the Table 6. Nested analysis of variance comparing the abundance of macrofauna in Swan Bay (SB) and an estimated production of macrofauna in Swan Bay (SB) adjacent area of Port Phillip Bay (PPB). DF—degrees and an adjacent area of Port Phillip Bay (PPB). DF— of freedom; P —probability. degrees of freedom; P—probability. Port Phillip Bay (Table 6). The highest estimated Source Mean square DF F-ralio P production of approximately 170 mg.m"^.d“' was recorded at Tin Can in May (Fig. 4). Production Date 9525.3 3 6.459 0.026 Bay 64412.6 1 104.515 0.009 estimates were also significantly different between Date*Bay 3381.5 3 2.293 0.178 sites in Swan Bay, however, this varied with date SB sites 703.9 1 1.006 0.320 (Table 6). Annual production in g.m’^.yr' at PPB sites 528.7 1 0.756 0.388 each site was estimated to be 45.5 for North SB sites*Date 2640.9 3 3.774 0.015 Jetty, 39.7 for Tin Can, 18.6 for vSt Leonards and PPB sites*Dale 308.7 3 0.441 0.724 21.2 for Beacon. Error 699.7 64 Macrofaunal communities in seagrass beds were distinctly different between the two bays (Fig. 5). Table 5. Nested analysis of variance comparing the biomass of macrofauna in Swan Bay (SB) and an Amphipods dominated in Port Phillip Bay sites; adjacent area of Port Phillip Bay (PPB). DF—degrees caprellid amphipods were found only in Port of freedom; P—probability. Phillip Bay, and gammaridean amphipods were at 6 FIONA L. BIRD AND GREGORY P. JENKINS e r o c r e p a n u a f o r c a m f o e c n a d n u b a n a e M ) g m ( e r o c r e p a F/g. 4. Mean abundance, n u biomass, and estimated a of production of macrofauna cr at sites in Swan Bay and a m Port Phillip Bay on four f sampling dates: o s (A) abundance; as (B) biomass; m (C) production. o bi Key to dates in caption n for Fig. 2. Error bars a e are standard error. M d ^ m g. m ( a n u a f o r c a m f o n o cti u d o r p n a e M North Jetty Tin Can St Leonards Beacon Swan Bay Port Phillip Bay Site INVERTEBRATE FAUNA ASSOCIATED WITH SEAGRASS, HETEROZOSTERA TASMANICA 1 I U • a> o cu cu O y. a B p hilli P ort P d n a o cu y o a B can B wan Tin S !¥ in s e sit at s e g a bl m e s s a al n u a of cr a m of n o siti o p m o c nt e c etty Per J h rt o N uouisodiuoo oSbjuoojoj FIONA L. BIRD AND GREGORY P. JENKINS 40 North Jetty 30 20 10 0 40 Tincan 30 20 10 Fig. 6. Per cent frequency of 0 macrofauna in sieve-size classes at sites in Swan Bay and Port 40 St Leonards Phillip Bay. 30 20 10 0 40 30 20 llh^ 10 0 0.5 0.71 1 1.4 2 2.8 4 Sieve sizes (mm) INVERTEBRATE FAUNA ASSOCIATED WITH SEAGRASS, HETEROZOSTERA TASMANICA 9 least three times more abundant. Cumaceans were In contrast to macrofauna, abundance of also more abundant in Port Phillip Bay. Other meiofauna was significantly higher in Swan Bay invertebrates such as tanaids, gastropods and (Table 7). On average, meiofaunal abundance bivalves were more important at sites in Swan in Swan Bay was approximately four-fold that Bay. The January samples from Beacon were on the adjacent coast of Port Phillip Bay (Fig. 7). unusual in the high representation of tanaidaceans The major difference in the meiofaunal abundances and isopods (Fig. 5). The October samples from between the two areas occurred in the 0.063 to Tin Can had unusually high numbers of ophiuroids 0.18 mm sieve-size range (Fig. 8). This was due (Fig. 5). to the presence of high abundances of epibenthic The size structure of the macrofaunal assem¬ harpacticoid copepods in this size range in Swan blages showed some variation amongst sites Bay (Fig. 8). (Fig. 6). Swan Bay sites generally had a higher proportion of larger invertebrates (greater than DISCUSSION 1.4 mm sieve size). Port Phillip Bay sites appeared Scagrass biomass and organic matter arc two to have a higher proportion of invertebrates in the physical parameters known to be correlated with 1 mm sieve-size class. invertebrate abundance and diversity in sca¬ grass beds. Heck & Whetstone (1977) found that invertebrate species number and abundance increased with plant biomass. Mann (1988) suggested that organic matter trapped in a sea- Source Mean square DF F-ratio P grass bed would provide food for detritovores and Edgar (1990a) showed that organic enrichment Bay 101.2 1 21.463 0.010 of sediments increased invertebrate production. SB sites 6.0 2 2.006 0.142 Edgar & Shaw (1993) found organic matter to be PPB sites 3.4 2 1.289 0.311 greater in sheltered scagrass beds than unsheltered Error 2.6 16 beds, and this correlated with a higher production of invertebrates in the sheltered environments. Table 7. Nested analysis of variance comparing the Scagrass biomass in Swan Bay and Port Phillip abundance of meiofauna in Swan Bay (SB) and an adjacent area of Port Phillip Bay (PPB). DF—degrees Bay did not significantly differ, but the organic of freedom; P—probability. content of sediment was higher in Swan Bay. a 8 150 oO . C 3 o 100 us o Fij’. 7. Mean abundance of meiofaunal organisms at sites in Swan Bay and Port Phillip Bay on 29 October 1990. Error bars arc standard error. North J Tin Can MidJ St Leo Beacon Edwards Pt Swan Bay Port Phillip Bay Site 10 FIONA L. BIRD AND GREGORY P. JENKINS 2 o o >-¥ Q< a c :3 .<4a^ fig. S. Mean abundance of o nieiofauna of different sieve- *6S size components in Swan <*- Bay and Port Phillip Bay. o 0) Solid region denotes mean CJ abundance of cpibcnthic § harpacticoids. Error bars •a c are standard error. 3 I Sieve sizes (mm) Drift algae was seen to accumulate and decompose protect it from the direct effect of wave and in Swan Bay and this would enhance organic current stress. content of the sediment. Macrofaunal abundance gave little indication of The macrofaunal communities of the two the differences between the bays, but Swan Bay bays were distinctly different. The macrofaunal had a significantly higher biomass and estimated assemblages at Port Phillip Bay sites were production of macrofauna than the adjacent area dominated by amphipods, whereas the Swan Bay of Port Phillip Bay. This difference was probably sites supported a wider range of invertebrate because larger invertebrates, especially tanaids types. The dominance of amphipods in Port Phillip and molluscs, were more abundant inside Swan Bay may have been due to the greater exposure Bay than Port Phillip Bay, thus contributing more to wave action and currents compared with Swan to the total biomass and production. Temperature Bay. Fenwick (1976) found that fauna in an algal would appear not to have been an important community was influenced by wave exposure; factor infiucncing production. The highest pro¬ with a high energy environment characterised by duction estimates for Swan Bay were recorded in low species diversity and very high densities of May when temperatures were lower than in Port amphipods. This distribution is consistent with Phillip Bay. High temperatures in Swan Bay over the suggestion that animals with strong grasping summer may have had a negative effect on cpi- appendages, such as amphipods, often dominate faunal production by causing migration to deeper algal communities at exposed sites (Takeuchi el al. water. Differences in macrofaunal production 1987; Hagerman 1966). The secure hold an estimated here may be underestimates if there is individual has on a blade of seagrass would a ditlerence in the available food resources in