Effect of macrophytes on aquatic invertebrates – a literature review P. Papas April 2007 Arthur Rylah Institute for Environmental Research Technical Report Series No. 158 Arthur Rylah Institute for Environmental Research Technical Report Series No. 158 Effect of macrophytes on aquatic invertebrates – a literature review Phil Papas April 2007 Effect of macrophytes on aquatic invertebrates – a literature review Published by: Arthur Rylah Institute for Environmental Research Department of Sustainability and Environment PO Box 137 Heidelberg, Victoria 3094 Australia Telephone: (03) 9450 8600 www.dse.vic.gov.au/ari This publication may be cited as: Papas, P. (2007) Effect of macrophytes on aquatic invertebrates – a literature review. Freshwater Ecology, Arthur Rylah Institute for Environmental Research, Technical Report Series No. 158, Department of Sustainability and Environment, Melbourne; Melbourne Water, Melbourne, Victoria. © The State of Victoria Department of Sustainability and Environment 2007 This publication is copyright. Apart from any fair dealing for private study, research, criticism or review allowed under the Copyright Act 1968, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any forms or by any means, electronic, photocopying or other, without the prior permission of the copyright holder. ISBN 978-1-74152-895-4 (print) ISBN 978-1-74152-901-2 (online) ISSN 1326 6446 Disclaimer This publication may be of assistance to you but the State of Victoria and its employees do not guarantee that the publication is without flaw of any kind or is wholly appropriate for your particular purposes and therefore disclaims all liability for any error, loss or other consequence, which may arise from you relying on any information in this publication. Authorised by the Victorian Government, Melbourne. Effect of macrophytes on aquatic invertebrates – a literature review Contents ABSTRACT ...........................................................................................................................................II 1 INTRODUCTION .......................................................................................................................... 1 2 USE OF MACROPHYTES BY INVERTEBRATES................................................................... 1 2.1 FACTORS AFFECTING INVERTEBRATE COMPOSITION AND DISTRIBUTION ON MACROPHYTES 2 Leaf architecture............................................................................................................................... 3 Nutritional value of the plant tissue .................................................................................................. 4 Chemical inhibitors in the plant tissue.............................................................................................. 4 Plant growth habit ............................................................................................................................ 5 Macrophyte community structure and composition .......................................................................... 5 Other regulators for invertebrates .................................................................................................... 5 3 INVERTEBRATE PREFERENCE FOR NATIVE OR EXOTIC MACROPHYTES ............. 6 4 CONCLUSION ............................................................................................................................... 7 5 ACKNOWLEDGEMENTS ........................................................................................................... 7 6 REFERENCES ............................................................................................................................... 8 APPENDIX 1- SUMMARY OF LITERATURE REVIEWED ........................................................ 12 APPENDIX 2 - EXPERIMENTAL CONSIDERATIONS ................................................................ 22 Effect of macrophytes on aquatic invertebrates – a literature review Abstract Aquatic macrophytes are an important habitat for invertebrates. Invertebrates utilise aquatic plants as a direct food source, shelter from predators, spawning and attachment sites as well as feeding on the periphyton growing on their surfaces. There are many attributes of macrophytes, operating at different scales that may affect invertebrate abundance, diversity and community composition. Examples of such attributes include the architecture of the leaves, the growth habit of the plant and the presence of chemical inhibitors in the plant tissue. A summary of the findings from studies pertaining to these attributes is as follows: • The degree of leaf dissectedness (leaf shape complexity) and surface area appear to influence macroinvertebrate community composition, diversity and abundance. • There is scant information on the presence of chemical inhibitors for E. canadensis, E. densa and M. aquaticum. Chemical effects exist for some related species. • Sturdiness of the plant and position of the leaves of the plant relative to the substratum effects invertebrate community. • There is evidence to suggest both negative and positive impacts on invertebrates with increasing macrophyte bed density. • Few studies compare macrophyte species diversity in macrophyte beds with invertebrate assemblages. There is some evidence to support greater invertebrate species diversity and abundance associated with greater macrophyte species diversity. Dense monospecific stands can negatively impact on water quality and degrade invertebrate habitat. • There are many factors other than those associated with the plant itself that are likely to affect invertebrate abundance, diversity and composition. Variables such as water depth, water quality, predator pressure and water flow have been shown to have a greater influence on invertebrate distribution in wetlands and streams that the macrophyte species present in the waterbody. • Some studies have found greater invertebrate abundance and diversity in native macrophytes compared to exotics, however others have found no difference. Care must be taken in extrapolating these findings to artificial systems as most of the studies are related to natural systems. Additional knowledge gaps are outlined at the end of the review. Some considerations for experimental design are presented. Effect of macrophytes on aquatic invertebrates – a literature review 1 Introduction Melbourne Water Corporation manages many artificial wetlands in the Melbourne metropolitan area. The primary role of these wetlands is to improve water quality (e.g. reduction of nitrogen and particulate matter) in streams that receive large amounts of stormwater. These wetlands also have environmental values, e.g. habitat for birds, frogs, fish and invertebrates as well as social values, e.g. recreation and education. Recently, three exotic aquatic plants have caused concern due to their invasive nature and rapid spread in some artificial wetlands. These are the submersed Egeria densa (egeria) and Elodea canadensis (elodea) and the submersed/emergent Myriophyllum aquaticum (parrots feather). The impact of these plants on the nutrient removal function of the wetlands forms part of a project to provide information useful for the management of the wetlands. A brief consideration of possible effects of these weeds on social, ecological, and economic values is provided in Ainsworth et al. (unpublished a) with some further assessment of impacts on aquatic biodiversity in Ainsworth (unpublished b). This literature review is the first stage of a project which aims to determine whether the properties of exotic macrophytes are inferior to those of natives such that when present at similar densities the native plants support more biodiversity. Aquatic invertebrates were selected as a component of the biodiversity to measure as: • • • • • They are generally less mobile than other fauna and thus more likely to reflect conditions where they are collected; they are generally easily sampled; experimental work does not require the approvals needed for work on vertebrates; changes in invertebrate populations will have flow on effects on other components of biodiversity; invertebrates are widely used to monitor water quality. An initial literature search revealed scant information specific to the effects of E. densa, E. canadensis and M. aquaticum on aquatic invertebrates, hence the scope of the literature search was broadened to incorporate all research pertaining to (principally) submerged macrophytes and their effect on aquatic invertebrate abundance, diversity and/or composition. 2 Use of macrophytes by invertebrates Aquatic macrophytes are an important habitat for invertebrates - invertebrates utilise them as a direct food source (Gregg and Rose 1982, 1985, Rooke 1986) shelter from predators (Harrod 1964, Gregg and Rose 1985), spawning and attachment sites (Keast 1984) as well as feeding on the periphyton growing on their surfaces (Higler 1975, Cattaneo and Kaliff 1980). Feeding groups (also known as feeding guilds and trophic groups – see Table 1) associated with aquatic macrophyte genera such as Myriophllyum, Elodea and Egeria, typically include shredders and grazers, detritovores, filter feeders and predators. The composition and abundance of the groups present varies depending on the Effect of macrophytes on aquatic invertebrates – a literature review location of the waterbody and the plant species that are present (e.g. Dvorak and Best 1982, Talbot and Ward 1987). Table 1 Functional feeding groups associated with aquatic plants (adapted from Cummins 1973, Dvorak and Best 1982). 2.1 Feeding group Type of organism and feeding habit Shredders/Grazers Invertebrates that consume living plant tissue– e.g. some Lepidoptera (moth larvae) and Trichoptera (caddisflies) Detritivores Invertebrates that collect dead organic material and/or scrape periphyton from plants – e.g. some Hempitera (true bugs), Lepidoptera (moth larvae) and Gastropoda (snails) Filter feeders Zooplankton and some insect larvae that filter water to gain detritus from around plants – e.g. Cladocera (small crustaceans) Predators Carnivorous invertebrates that prey on smaller invertebrates – e.g. Odonata (dragonflies), some Hemiptera (true bugs) Factors affecting invertebrate composition and distribution on macrophytes In order to begin to understand why there may be differences in the invertebrates associated with native and exotic macrophyte species or preferences for particular species, it is important to consider the factors affecting the distribution and composition of invertebrates on all aquatic plant species. These include effects operating at a range of scales from the individual leaves to the macrophyte beds (Table 2). Additionally, external factors such as water quality, temperature and water regime may also be a determinant of the macroinvertebrate community present on macrophytes (Strayer et al. 2003). These factors are explored in detail following Table 2. Effect of macrophytes on aquatic invertebrates – a literature review Table 2 Factors affecting the macroinvertebrate abundance and diversity associated with macrophytes at different scales. Scales at which effects are operating Factors affecting macroinvertebrate abundance and richness Individual plant • • • • • Whole plant • • Macrophyte bed • • Architecture of the leaves (also referred to as leaf morphology) (Balci and Kennedy 2003). Dissectedness and surface area of the leaves) (Shiel 1976, Balci and Kennedy 2003). Nutrient content of the leaves (i.e. nutritional value) of the plants (Lodge 1991). Chemical compounds in the plant tissue (Lodge 1991, Erven and Wetzel 2003). Presence and type of epiphytic community on the leaves (e.g. diatoms, algae etc.) (Lodge 1985, Ervin and Wetzel 2003). Plant growth habit: proximity of the leaves of the plant to the substratum (Timms 1981). Location of microhabitats within the water column (Shiel 1976). Plant growth habit: density of the standing crop (Cattaneo et al. 1998). Diversity of the macrophyte community (Quade 1969, Hanson 1990). Leaf architecture Leaf architecture refers to the physical form of the leaves and includes attributes such as surface area of the leaves and the leaf dissectedness, which refers to the complexity of the leaf shape - ovate leaves have low dissectedness and pinnate leaves high dissectedness, for example. The relationships between leaf architecture and surface area and aquatic invertebrate abundance and diversity has been welldocumented (e.g. Krecker 1939, Cheruvelli et al. 2001, Balci and Kennedy 2003). Highly dissected leaves result in a higher surface area to volume ration per unit of biomass and can consequently support a greater biomass of epiphytic growth (Cattaneo et al. 1998) for grazing aquatic invertebrates. Macrophytes with highly dissected leaves are generally recognised as supporting larger aquatic invertebrate populations than plants with broader leaves (Krecker 1939, Cheruvelli et al. 2000), with exceptions (eg. Cyr and Downing 1988). It is also suggested that the increased complexity of the leaf surface can provide better refuge from predators (Balci and Kennedy 2003) and substrate for invertebrates (Keast 1984). Cyr and Downing (1988), in a study of ten lakes in Canada, determined, however, that the abundance of invertebrates was not systematically related to the level of leaf dissection but due to other factors such as Effect of macrophytes on aquatic invertebrates – a literature review plant morphology, surface texture, epiphytic algal growth and community composition, nutrient content of plant tissues and presence of defensive chemicals in the plant tissue. Nutritional value of the plant tissue It has been suggested that the nitrogen (protein) content of vascular macrophytes limits herbivory (Mattson 1980, Lodge 1991). Lodge (1991) reviewed literature to determine if the nitrogen content of aquatic plants varied between species and other plant groups and hence was likely to lead to differences in grazing by invertebrates. Data from the literature indicted little difference in the nitrogen content between algae, aquatic plants and terrestrial shrub and tree leaves. There was some evidence to suggest that floating aquatic plants had higher nitrogen content that other aquatic groups, however Lodge (1991) noted there were many variables likely to be affecting nitrogen concentrations and the results should be interpreted with caution. Lodge (1991) summarised that nutrient concentrations were not likely to be limiting grazing on macrophytes and that typically grazing preference was unrelated to commonly measured plant physical and chemical characteristics (such as cellulose content, micronutrients etc.). Chemical inhibitors in the plant tissue Allelochemicals or allelopathins, are largely secondary plant metabolites that play an important role in allelopathic interactions (an interaction where one plant causes suffering to another plant) and plant defence (Liu and An unpublished). Lodge (1991) found evidence that the phenolic content of some macrophytes may be a factor limiting grazing by invertebrates, however concluded that more work needed to be done to test the hypothesis proposed by Crawley (1983) and Coley et al, (1985) that discuss mechanisms that plants use to defend against herbivory. A review by Ervin and Wetzel (2003) highlights that the allelochemical compounds may be influenced by the availability of nutrients, inorganic carbon and light. The following information is an extract from their review: “Species are presently known to produce anti-algal compounds: Chara spp. (van Donk and van de Bund 2002; Wium-Andersen 1987), Ceratophyllum demersum (Wium-Andersen 1987), Myriophyllum spicatum (Gross et al. 1996; Gross and Sütfeld 1994; Nakai et al. 1996, 1999, 2000), Myriophyllum brasiliense (M. aquaticum) (Saito et al. 1989), Hottonia palustris and the bryophyte Fontinalis antipyretica (Gross 1999). Hydrolyzablepolyphenolic compounds (tannins) from Myriophyllum aquaticum demonstrated significant suppressive activity against cyanobacteria; such compounds were isolated from plant material and from waters surrounding live plants, indicating an active production and release (Saito et al. 1989). Similar properties were shown for hydrolyzablepolyphenolic compounds from Myriophyllum spicatum and sulfur-containing compounds from Ceratophyllum spp. (Gross 1999). Other work by Newman (Newman et al. 1990, 1992, 1996) demonstrated experimentally the deterrent properties of glucosinolate compounds (demonstrated by Siemens et al. 2002) to have dual allelopathic/anti-herbivory properties) in watercress (Nasturtium officinale) to feeding by the omnivorous amphipod Gammarus pseudolimnaeus, the snail Physella gyrina, and a suite of Trichoptera Effect of macrophytes on aquatic invertebrates – a literature review (caddisfly) larvae. The ecological benefit of multiple functionality of such compounds, however, remains to be demonstrated in aquatic systems”. Plant growth habit Cyr and Downing (1988) propose that the macroinvertebrate community composition on the plants is be affected by the sturdiness of the plant, e.g. larger plants might be more suitable to support heavy crawling invertebrates. It has also been found that elodea often supports more gastropods than more fragile plants such as Chara and Myriophyllum (Kufikowski 1974, Soszka 1975). Timms (1981) postulates that differences in aquatic macro and micro invertebrates between different macrophyte beds is driven by the proximity of the leaves to the substratum. In his study involving three lakes in Victoria (Australia), differences in invertebrate assemblages were found between Myriophyllum propinquum and Ruppia sp. beds. – species with similar leaf architecture but different growth habits, i.e. Ruppia grows close to the substratum and M. propinquum has long stalks with most leaves near the water surface. Some macrophyte species have the tendency to form thick beds that can lead to a homogeneous canopy with low light penetration. Dense homogeneous beds can prevent the establishment of ‘understorey’ species, inhibit the growth of epiphytic algae and lead to low macroinvertebrate diversity (Cheruvelil et al. 2001). Unmuth et al. (2000) found significant changes in temperature and dissolved oxygen concentration in dense stands of M. spciatum in a Wisconsin lake. Cheruvelil et al. (2001) suggests these conditions can make it inhospitable for invertebrates. Sandilands and Hann (1996) found however, that dense stands of macrophytes harboured large numbers of invertebrates and considered that protection from predation was the likely mechanism explaining this. Dvorak and Best (1982) also found high numbers of macroinvertebrate species in dense beds and linked this to the colonisable surface area of the macrophyte bed and its density. Macrophyte community structure and composition Whilst there are many studies comparing the invertebrates between macrophyte species (e.g. Krecker 1939, Gerrish and Bristow 1979, Jeffries 1993, Rooke 1986, Nichols and Shaw 1986, Cheruvelli et al. 2000, Strayer et al. 2003), there appear to be few studies involving macrophyte beds that contain a suite of species. Of the few, Hanson (1990) found in a study on two different macrophyte communities in a Canadian lake, that any changes in plant species composition in the macrophyte beds greatly altered the structure of the macroinvertebrate communities. Keast (1984) found greater numbers of invertebrates on macrophyte beds with two native species of Potamogeton and Vallisneria compared to a monospecific bed of an exotic species M. spicatum (Eurasian milfoil) in a Canadian lake. Other regulators for invertebrates Many authors agree that there is a range of factors affecting invertebrate abundance and diversity in aquatic systems – some of which have been covered thus far in the review. Factors other than those associated with the plants themselves are likely Effect of macrophytes on aquatic invertebrates – a literature review determinants (in addition to macrophyte attributes) for macroinvertebrate abundance, diversity and/or community composition, these include: flow (e.g. Greg and Rose 1985), water level (e.g. Timms 1981, Humphries 1996), water quality (e.g. Strayer et al. 2003) and interactions between invertebrates such as predator-prey relationships and fish (e.g. Dibble and Harrel 1997). A study by Humphries (1996) showed water level was a determinant of invertebrate richness and abundance in rivers. Similarly, Timms (1981) found water depth was correlated with invertebrate abundance and community composition in three lakes studied. In systems with water movement, macrophytes can significantly alter flow patterns as demonstrated by Gregg and Rose (1985) who conducted manipulated macrophyte trials in a flowing system and concluded reduced current velocities in the macrophytes appeared to be the reason why invertebrates chose to occupy them. Strayer et al. (2003) proposed that low dissolved oxygen (DO) levels in Trapa beds may have reduced the amount of food available to predatory invertebrates, while Gregg and Rose (1985) proposed that low DO levels at night was a possible cause for reduced planktonic invertebrate numbers observed in dense beds in their manipulated trials. The potential for seasonal die-off of dense macrophyte beds and the consequential accumulation and breakdown of organic matter may also lead to low DO levels and impacts on invertebrates (Smock and Stoneburner 1980). 3 Invertebrate preference for native or exotic macrophytes Few studies have specifically compared the invertebrate communities of native and exotic species of macrophytes – however some related studies have drawn conclusions that compare invertebrate communities associated with native and exotic macrophytes. All of the studies found by this review were from countries other than Australia. Findings from the studies are mixed with some showing clear differences in invertebrate abundance, diversity and/or community composition between native and exotic species and others no difference was demonstrated. In a study of invertebrate communities associated with macrophyte species in a Canadian lake, Keast (1984) found that the emergence of invertebrates was twice as high above native macrophyte beds (i.e. Potamogeton sp. and Vallisneria sp.) than the exotic beds (i.e. M. spicatum) and in the benthos beneath the native macrophyte beds, five major taxa of prey invertebrate were five to seven times more abundant. Also, the foliage of Potamogeton plus Vallisneria supported twice as many invertebrate per square metre than M. spicatum. Keats (1984) concludes that the likely reason for these patterns is the small substrate particle size beneath M. spicatum, which was unfavourable for invertebrates. Other studies where M. spicatum were shown to support fewer invertebrate species that native macrophytes include Soszka (1975) and Cattaneo et al. (1998) Balci and Kennedy (2003) also studied M. spicatum and compared the invertebrate communities of that exotic species with the native water stargrass Heteranthera dubia in a Texan reservoir. They found no significant difference in the species richness between the native and exotic macrophyte beds and concluded that temporal variation was the overriding determinant of invertebrate patterns in abundance. Similarly, in a study on the invertebrate assemblages in a New Zealand Effect of macrophytes on aquatic invertebrates – a literature review lake, Biggs and Malthus (1982) found no clear preference (in terms of either numbers of taxa, abundance or biomass) for native macrophytes, as opposed to exotic species. 4 Conclusion The assessment of the literature relating to macrophytes and aquatic invertebrates revealed that findings from the studies are divergent with respect to invertebrate preference for native versus exotic macrophyte species, different macrophyte species in general and different densities of macrophytes. Hence, the effect of exotic species such as elodea, egeria and parrots feather on invertebrates in artificial wetlands cannot be confidently ascertained from the literature alone. Most studies have been undertaken in natural lakes and wetlands. Caution is needed when extrapolating the results of these studies to artificial systems as artificial systems exhibit altered water regimes and degraded water quality, which is likely to influence the type of invertebrate community in these systems. Importantly, external factors such as water quality and the relatively recent establishment of the wetlands themselves may be overriding the effect of the macrophyte species on the invertebrate communities present in the wetlands. Knowledge gaps The following knowledge gaps need to be overcome before an appropriate assessment/experiment of the impact of the exotic species on aquatic invertebrates can be determined (see also Appendix 2 for experimental design considerations). • The invertebrate community present in the artificial wetlands with and without macrophytes needs to be ascertained (from existing data and/or sampling). • Water quality and hydrological data from the artificial wetlands needs to be examined and its likely impact on invertebrate communities determined. • The value of an experiment to determine effects of exotic species on invertebrates should be debated and pending the outcome of the debate, the most appropriate experimental design should be determined. 5 Acknowledgements Funding for this review was provided by Melbourne Water. David Bryant and Michael Nicol, Freshwater Ecology, Department of Sustainability and Environment are thanked for assisting with the literature search and review. Diane Crowther and David Bryant, Freshwater Ecology, Department of Sustainability and Environment are thanked for proof reading the document. Effect of macrophytes on aquatic invertebrates – a literature review 6 References Ainsworth, N. (unpublished) Elodea canadensis and Egeria densa in Melbourne Water wetlands. Primary Industries Research Victoria. April 2004. Ainsworth, N., Hunt, T., and Weiss, J (unpublished) Elodea candensis and Egeria densa identification and management techniques with an Australian Focus. Primary Industries Research Victoria. April 2004. Balci, P. and Kennedy, J. H. (2003) Comparison of Chironomids and Other Macroinvertebrates Associated with Myriophyllum spicatum and Heteranthera dubia. Journal of Freshwater Ecology 18(2): 235-247. Biggs, B.J.F. and. Malthus, T.J (1982) Macroinvertebrates associated with various aquatic macrophytes in the backwaters and lakes of the upper Clutha Valley, New Zealand. New Zealand Journal of Marine and Freshwater Research 16(1): 8189. Cattaneo, A., Galanti, G., Gentinetta, S. and Romo, S. (1998) Epiphytic algae and macroinvertebrates on submerged and floating-leaved macrophytes in an Italian lake. Freshwater Biology 39: 725-740. Cheruvelil, K.S., Soranno, P.A. and Madsen, J.D. (2001) Epiphytic Macroinvertebrates Along a Gradient of Eurasian Watermilfoil Cover. Journal of Aquatic Plant Management 39: 67-72. Cyr, H.A. and Downing., J.A (1988) The abundance of phytophilous invertebrates on different species of submerged macrophytes. Freshwater Biology 20: 365-374. Cattaneo, W.P and Kaliff (1980) The relative contribution of aquatic macrophytes and their epiphytes to the production of macrophyte beds. Limnology and Oceanography 25:280-289 Coley, P.D., Bryant, J.P., III, F.S. (1985) Resource availability and plant antiherbivore defense. Science 230:895-899 Crawley, M.J. (1983) Herbivory: the Dynamics of Animal-Plant Interactions. University of California Press, Berkley, CA. 437 pp Cummins, K.W. (1973) Trophic relations of aquatic insects. Annual Review of Entomology 18:182-206 Cyr, H. and Downing., J.A (1988). the abundance of phytophilous invertebrates on different species of submerged macrophytes. Freshwater Biology 20:365-374. Dibble, E.D. and Harrel, S.L. (1997) Largemouth Bass Diets in Two Aquatic Plant Communities. Journal of Aquatic Plant Management 35:74-78. Downing, J.,A. and Cyr, H. (1985) Quantitative estimation of epiphytic invertebrate populations. Canadian Journal of Fisheries and Aquatic Sciences 42:1570-1579. Douglas, M.M. and O'Connor R.A. (2003) Effects of the exotic macrophyte, para grass (Urochloa mutica), on benthic and epiphytic macroinvertebrates of a tropical floodplain. Freshwater Biology 48(6): 962-971. Dvorak, J.A. and Best, E.P.H. (1982) Macro-invertebrate communities associated with macrophytes of Lake Vechten: structural and functional relationships. Hydrobiologia 95: 115-126. Effect of macrophytes on aquatic invertebrates – a literature review Ervin, G.N. and. Wetzel, R.G. (2003) An ecological perspective of allelochemical interference in land–water interface communities. Plant and Soil 256: 13-28. Gregg, W.W. and Rose, F.L. (1982) The effect of aquatic macrophytes on the stream microenvironment. Aquatic Botany 14:309-324 Gregg, W. W. and Rose, F.L. (1985) Influences of aquatic macrophytes on invertebrate community structure, guild structure, and microdistribution in streams. Hydrobiologia, 128:45-56. Hann, B.J. (1995) Invertebrate associations with submersed aquatic plants in a prairie wetland. UFS Delta Marsh Annual Report. Winnipeg, Manitoba, Department of Zoology, University of Manitoba. 30: 78-84. Gross, E.M. and Sütfeld, R (1994) Polyphenols with algicidal activity in the submerged macrophyte Myriophyllum spicatum L. Acta Hort. 381:710–716. Gross, E.M. (1999) Allelopathy in benthic and littoral areas: case studies on allelochemicals from benthic cyanobacteria and submersed macrophytes. In Principles and Practices in Plant Ecology: Allelochemical Interactions. Eds. Inderjit, K M M Dakshini and C L Foy. pp. 179–199. CRC Press, Boca Raton, FL. Gross, E.M. and Sütfeld, R. (1994) Polyphenols with algicidal activity in the submerged macrophyte Myriophyllum spicatum L. Acta Hort. 381:710–716. Gross, E.M., Meyer, H. and Schilling, G. (1996) Release and ecological impact of algicidal hydrolysable polyphenols in Myriophyllum spicatum. Phytochemistry 41:133–138. Hanson, J.M. (1990). Macroinvertebrate size-distributions of two contrasting freshwater macrophyte communities. Freshwater Biology 24: 481-491. Harrod, J.J. (1964) The distribution of invertebrates on submerged aquatic plants in a chalk stream. Journal of Animal Ecology 33:335-348 Higler, L.W.G. (1975) Analysis of the macrofauna community on Stratiotes vegetation. Verhand-lungen de r International Vereinigung fur Theoretische und Andewandte Limnologie 19:2773-2777 Humphries, P. (1996) Aquatic macrophytes, macroinvertebrate associations and water levels in a lowland Tasmanian river. Hydrobiologia, 321: 219-233. Keast, A. (1984) The introduced aquatic macrophyte, Myriophyllum spicatum, as a habitat for fish and their invertebrate prey. Canadian Journal of Zoology 62:1289-1303. Krecker, H.F. (1939) A comparative study of the animal population of certain submerged aquatic plants. Ecology 20(4): 553-562. Kufikowski, T. (1974) The phytophilous fauna of the dam reservoir at Goczalkowice. Acta hydrobiologia, Krakow 16:189-207 Liu, D.L and An, M (unpublished) Simulating dynamics of allelochemical production from living plants.Proceedings of the Australian Agronomy Conference, Australian Society of Agronomy. Effect of macrophytes on aquatic invertebrates – a literature review Lodge, D.M. (1985) Macrophyte-gastropod associations: observations and experiments on macrophyte choice by gastropods. Freshwater Biology 15: 695708. Lodge, D.M. (1991) Herbivory on freshwater macrophytes. Aquatic Botany 41: 195224. Marklund, O. (2000) A new sampler for collecting invertebrates in submerged vegetation. Hydrobiologia 432: 229-231. Mattson, Jr., W.J. (1980) Herbivory in relation to plant nitrogen content. Annual Review of Ecological Systematics 11:119-161 Miller, A.C., Beckett, D.C., David, C., Way, M. and Bacon E.J. (1989) The habitat value of aquatic macrophytes for macroinvertebrates. US Army Engineers Waterway Experimental Station. Technical Report A-89, Vicksburg, MS. Nakai, S., Hosomi, M., Okada, M. and Murakami, A, (1996) Control of algal growth by macrophytes and macrophyte-extracted bioactive compounds. Wat. Sci. Technol. 34: 227–235. Nakai, S., Inoue, Y., Hosomi, M. and Murakami, A. (1999) Growth inhibition of bluegreen algae by allelopathic effect of macrophytes. Wat. Sci. Technol. 39:47–53. Nakai, S., Inoue, Y., Hosomi, M. and Murakami, A. (2000) Myriophyllum spicatumreleased allelopathic polyphenols inhibiting growth of blue-green algae Microcystis aeruginosa. Wat. Res. 34:3026–3032. Newman, R.M., Kerfoot, W.C. and Hanscom, Z, III (1990) Watercress and amphipods: potential chemical defenses in a spring stream macrophyte. J. Chem. Ecol. 16:245– 259. Newman, R.M, Hanscom, Z. III and Kerfoot, W.C. (1992) The watercress glucosinolatemyrosinase system: a feeding deterrent to caddisflies, snails, and amphipods. Oecologia 92:1–7. Newman, M.R.M, Kerfoot, W.C. and Hanscom, Z. III (1996) Watercress allelochemical defends high-nitrogen foliage against consumption: effects on freshwater invertebrate herbivores. Ecology 77:2312–2323. Nichols, S. A. and Shaw, B. H. (1986) Ecological life histories of the three aquatic nuisance plants, Myriophyllum spicatum, Potamogeton crispus and Elodea canadensis. Hydrobiologia, 131: 3-21. Quade, H.W. (1969) Cladoceran faunas associated with aquatic macrophytes in some lakes in northeastern Minnesota. Ecology 50:170-179 Rooke, B. (1986) Macroinvertebrates associated with macrophytes and plastic imitations in the Eramosa River, Ontario, Canada. Archiv fuer Hydrobiologie 106(3): 307-325. Saito, K., Matsumoto, M., Sekine, T. and Murakoshi, I. (1989) Inhibitory substances from Myriophyllum brasiliense on growth of bluegreen algae. J. Nat. Prod. 52:1221–1226. Sandilands, K. A. nd Hann, B. J. (1996) Effect of fish and submersed macrophytes on the abundance of zooplankton in a prairie wetland. UFS (Delta Marsh) Annual Report 31: 58-62. Effect of macrophytes on aquatic invertebrates – a literature review Shiel, R.J. (1976) Associations of Entomostraca with Weedbed Habitats in a Billabong of the Goulburn River, Victoria. Australian Journal of Marine and Freshwater Research 27: 533 - 549. Siemens, D.H., Garner, S.H., Mitchell-Olds, T. and Callaway, R.M. (2002) Cost of defense in the context of plant competition: Brassica rapa may grow and defend. Ecology 83:505–517. Smock, A. and Stoneburner, D.L. (1980) The Response of macroinvertebrates to aquatic macrophyte decomposition. Oikos 35:397-403 Soszka, G.J. (1975) The invertebrates on macrophytes in three Masurian lakes. Ekologia Polska 23:371-391 Strayer, D.L., Lutz, C., Malcom, H.M., Munger, K. and Shaw, W.H. (2003) Invertebrate communities associated with a native (Vallisneria americana) and an alien (Trapa natans) macrophyte in a large river. Freshwater Biology 48: 1938-1949. Talbot, J. M. and Ward, J. C. (1987) Macroinvertebrates associated with aquatic macrophytes in Lake Alexandrina, New Zealand. New Zealand Journal of Marine and Freshwater Research 21(2): 199-213. Thorp, A. G., Jones, R.C. And Kelso, D.P. (1997) A Comparison of Water-Column Macroinvertebrate Communities in Beds of Differing Submersed Aquatic Vegetation in the Tidal Freshwater Potomac River. Estuaries 20(1): 86-95. Timms, B.V. (1981) Animal communities in three Victorian lakes of different salinities. Hydrobiologia 81: 181-193. Wollheim, W. M. and Lovvorn, J. R. (1996) Effects of macrophyte growth forms on invertebrate communities in saline lakes of Wyoming High Plains. Hydrobiologia, 323: 83-96. Unmuth, J.M.L., Lillie, R.A., Dreikosen, D.S. and Marshall, D.W. (2000) Influence of dense growth of Eurasian watermilfoil on lake water temperature and dissolved oxygen. Freshwater Ecology 15:497-503 Van Donk, E. and van de Bund, W.J. (2002) Impact of submerged macrophytes including charophytes on phyto- and zooplankton communities: allelopathy versus other mechanisms. Aquatic Botany. 72:261–274. Wium-Andersen, S. (1987) Allelopathy among aquatic plants. Arch. Hydrobiol. 27:167–172. 11 Effect of macrophytes on aquatic invertebrates – a literature review Appendix 1 – Summary of the literature reviewed. Reference Location Balci, P. and Kennedy, J. H. (2003). Comparison of Chironomids and Other Macroinvertebrates Associated with Myriophyllum spicatum and Heteranthera dubia. Journal of Freshwater Ecology 18(2): 235247. USA Waterbody Objective/Aim Findings Experimental ponds • • • • Investigate the influence of different plant species on macroinvertebrate abundance and community composition. Examine the secondary productivity of chironomids, the major macrofauna associated with macrophytes. • • • Biggs, B. J. F. and. Malthus, T. J (1982). Macroinvertebrates associated with various aquatic macrophytes in the backwaters and lakes of the upper Clutha Valley, New Zealand. New Zealand Journal of Marine and Freshwater Research 16(1): 8189. New Zealand Cattaneo, A., Galanti, G., Gentinetta, S. and Romo, S. (1998). Epiphytic algae and macroinvertebrates on submerged and floating-leaved macrophytes in an Italian lake. Freshwater Biology 39: 725-740. Italy Lago Di Candia Backwater Lakes • A study of the macroinvertebrates inhabiting the major aquatic macrophyte communities in the Upper Clutha Valley. • • • • Lake • • • To test the effect of different host architecture on epiphytic algae and invertebrates. Three species tested – submerged Myriophyllum spicatum and Ceratophyllum demersum, Najas marina and floating Trapa natans With the aim of predicting consequences for the lake of changes in predominant vegetation. • • • • • • Experimental ponds 30m x 16m x 50cm Macrophyte morphology explained some of the variation in the abundance of macroinvertebrates. In general, plants with highly dissected leaves supported larger macroinvertebrate populations than plants with broader, undissected leaves. Significant correlation between epiphyton and macroinvertebrate abundance. Suggested that the contributions of macroinvertebrates to the aquatic communities (with respect to secondary productivity) are similar in native and non-native plants. Samples collected by SCUBA divers Invertebrates showed a preference for certain macrophyte taxa as habitat, however, no discussion of architecture or theories on this finding. All communities were dominated by Potamopyrgus antipodarum. There was no clear preference (in terms of either numbers of taxa, abundance or biomass) for native macrophytes, as opposed to adventive species. Plant architecture affected the quantity of epiphyton. Algae and macroinvertebrate density higher on submerged plants. Taxonomic composition of epiphytic algae and macroinvertebrates was similar on the different plants. Floating-leaved T natans had greater diversity of algae but not macroinvertebrates. Significant inverse relationship found between epiphyton biomass and the biomass of the standing crop of the host plant. Replacement of floating-leaved macrophytes with submerged plants will increase the total biomass of epiphytic algae and invertebrates. 12 Effect of macrophytes on aquatic invertebrates – a literature review Reference Cheruvelil, K. S., Soranno, P. A. and Madsen, J. D. (2001). Epiphytic Macroinvertebrates Along a Gradient of Eurasian Watermilfoil Cover. Journal of Aquatic Plant Management 39: 67-72. Location USA Southern Michigan Waterbody Lakes Objective/Aim Findings • • • • Examine how milfoil cover affects the interactions between macrophytes and epiphytic macroinvertebrates. Determine whether macroinvertebrate density and biomass on the dominant plant species in a lake varies predicably with percent of littoral zone covered with milfoil. Assess potential of Fluridone as a means of Milfoil control. • • • Cyr, H. A. and Downing., J.A (1988). The abundance of phytophilous invertebrates on different species of submerged macrophytes. Freshwater Biology 20: 365-374. Lakes, Quebec, Canada Lakes • To test the hypothesis that macrophytes with finely dissected leaves support higher abundance of invertebrates than plants with broad leaves. • • • • • • Snorkelers sampled Macroinvertebrates using a 500 micron mesh bag with individual stems sampled. Macroinvertebrate biomass did not show consistent trends with milfoil cover while it was expected despite milfoil leading to an increased SA:V ratio, macroinvertebrate biomass would decrease due to thick milfoil canopy reducing heterogeneity of macrophyte beds. Hypothesised that as samples were taken from heterogenous macrophyte beds while milfoil often results in homogeneous canopies, a true comparison may not have been achieved. The use of Fluridone reduced milfoil cover while having no long term effects on macroinvertebrate biomass. Ten lakes in Quebec, Canada sampled with Myriophyllum spp., Ceratophyllum spp., Potamogeton spp., Vallisneria sp. Abundance of invertebrates not systematically related to level of dissection. Macrophytes with finely divided leaves did not support more macroinvertebrate per unit plant biomass that plants with larger leaves. Larger plants might be more suitable to support heavy crawling invertebrates. Elodea often supports more gastropods than more fragile plants such as Chara and Myriophyllum. Abundance of epiphytic invertebrates is probably related to a suite of factors including plant morphology, surface texture, epiphytic algal growth and community composition, nutrient content of plant tissues and presence of defensive chemicals. 13 Effect of macrophytes on aquatic invertebrates – a literature review Reference Douglas, M. M. and O'Connor R. A. (2003). Effects of the exotic macrophyte, para grass (Urochloa mutica), on benthic and epiphytic macroinvertebrates of a tropical floodplain. Freshwater Biology 48(6): 962-971. Location Australia (tropical) Waterbody Magela Creek Objective/Aim Findings • • Examine the effect of the exotic macrophyte, para grass, on benthic and epiphytic macroinvertebrates of a tropical floodplain. • • Dvorak, J. A. and Best, E.P.H. (1982). Macro-invertebrate communities associated with macrophytes of Lake Vechten: structural and functional relationships. Hydrobiologia 95: 115-126. Lake Vechten The Netherland s Lake • Study the structure and size of the macroinvertebrate communities associated with eight species of macrophytes in the lake. • • Macroinvertebrate richness, abundance and community similarity showed very few differences among the grass communities, particularly in the epiphytic habitat. Benthic invertebrates showed some differences among grasses, with lower richness and abundance and different community structure associated with Hymenachne. Herbicide control of para grass had no apparent effect on benthic invertebrates but reduced the abundance of epiphytic invertebrates in the short term. The results of this study indicate that para grass has very little impact on macroinvertebrate communities, despite the changes to macrophyte communities. This is probably because para grass has similar physical structure to the native grasses and because none of these grasses contribute directly to aquatic food webs. Control of para grass using herbicide has little impact on aquatic invertebrates. This suggests that predicting the impact of weed invasion in wetlands requires an understanding of both the functional properties of macrophytes and the habitat preferences of the macroinvertebrates SCUBA diving used to sample macrophytes Distribution of macroinvertebrates depended on colonisable plant surface area and vegetation density. 14 Effect of macrophytes on aquatic invertebrates – a literature review Reference Ervin, G. N. and. Wetzel, R.G. (2003). An ecological perspective of allelochemical interference in land–water interface communities. Plant and Soil 256: 13-28. Location Literature Review Waterbody Literature Review Objective/Aim Findings • • Literature review • • • • Among the species presently known to produce anti-algal compounds are: Chara spp., Ceratophyllum demersum, Myriophyllum spicatum, Myriophyllum brasiliense (M. aquaticum), Hottonia palustris and the bryophyte Fontinalis antipyretica. Hydrolyzablepolyphenolic compounds (tannins) from Myriophyllum aquaticum demonstrated significant suppressive activity against cyanobacteria; such compounds were isolated from plant material and from waters surrounding live plants, indicating an active production and release. Similar properties were shown for hydrolyzable polyphenolic compounds from Myriophyllum spicatum and sulfur-containing compounds from Ceratophyllum spp. Other work has demonstrated experimentally the deterrent properties of glucosinolate compounds to have dual allelopathic /anti-herbivory properties in watercress (Nasturtium officinale) to feeding by the omnivorous amphipod Gammarus pseudolimnaeus, the snail Physella gyrina, and a suite of Trichoptera (caddisfly) larvae. The ecological benefit of multiple functionality of such compounds, however, remains to be demonstrated in aquatic systems. In studies of the effects of atmospheric CO2 enrichment on macrophyte growth and decomposition, fungal biomass and growth rates were demonstrated to be markedly lower (25–50%) on the more recalcitrant particulate organic matter of plants grown in elevated CO2 concentrations that also possess lower tissue N concentrations. Bacterial development on detrital surfaces was also appreciably reduced by nearly an order of magnitude on elevated CO2-grown tissues. Detritivorous invertebrates feeding on these more recalcitrant organic materials and their microbial epiphytes consumed less, assimilated less, and grew 12 times more slowly than their counterparts fed on microbes associated with plant tissues grown on ambient CO2 concentrations. 15 Effect of macrophytes on aquatic invertebrates – a literature review Reference Location Gregg, W. W. and Rose, F. L. (1985) Influences of aquatic macrophytes on invertebrate community structure, guild structure, and microdistribution in streams. Hydrobiologia, 128:4556. USA Hann, B. J. (1995). Invertebrate associations with submersed aquatic plants in a prairie wetland. UFS Delta Marsh Annual Report. Winnipeg, Manitoba, Department of Zoology, University of Manitoba. 30: 78-84. USA Winnipeg, Manitoba Waterbody River Prairie Wetland Objective/Aim Findings • Study the influences of macrophytes on invertebrates in streams (Macrophytes previously shown by the authors to alter conditions of current velocity, substrate and detritus/diatom availability in the stream microhabitat). • • Determine if the structure and relative abundances of invertebrates differ between plant taxa (Ceratophyllum demersum, Potamogeton zosteriformis and Chara). • • • • • • • Hanson, J. M. (1990). Macroinvertebrate sizedistributions of two contrasting freshwater macrophyte communities. Freshwater Biology 24: 481-491. Narrow Lake, Canada Lake • Compare size distributions of benthic macroinvertebrates between Chara and rooted plants. • • • • • • Macrophytes were shown to increase stream invertebrate taxa richness and abundance, particularly that of shredders, scrapers and predators, and more so in Autumn than in Spring. Some groups of invertebrates were found to avoid the macrophytes. Reduced current velocities in the macrophytes appears to be what causes the invertebrates to choose to occupy them (bugs are known to have current velocity preferences). However the increase in microhabitats created by plants, and their large surface area, are also considered to benefit invertebrate abundances by adding habitat to the water column that would otherwise not be there (ie. compared to open water, no plants). Macrophytes and Chara sampled using Downing sampler. Species composition generally similar between different plants Many species favoured Ceratophyllum and Potamogeton in preference to Chara even though Chara and Ceratophyllum are more similar morphologically. Very low abundance of many species associated with Chara may be due to allelochemical properties of the taxon. No preference found with gastropods. Fewer planktonic invertebrates associated with dense macrophyte beds – possibly due to low oxygen at night and physical conditions detrimental to filtering and feeding activities. Macrophytes/substrate sampled using an Ekman grab sampler. Macroinvertebrates on macrophytes and sediment not differentiated. Chara had consistently higher and significantly greater invertebrate abundance than rooted plants. Chara beds dominated by chironomids, anisopterans, gastropods and sphaeridae. Rooted plants dominated by amphipods. Species composition of macrophyte beds can greatly influence abundance size structure and composition of macroinvertebrates. 16 Effect of macrophytes on aquatic invertebrates – a literature review Reference Humphries, P. (1996) Aquatic macrophytes, macroinvertebrate associations and water levels in a lowland Tasmanian river. Hydrobiologia, 321: 219-233. Location Tasmania Australia Waterbody River Objective/Aim Findings • • Measure invertebrate abundance/richness and describe the invertebrate assemblages associated with three species of macrophytes, and examine their response to changes in water levels • • • • • Keast, A. (1984). The introduced aquatic macrophyte, Myriophyllum spicatum, as a habitat for fish and their invertebrate prey. Canadian Journal of Zoology 62: 12891303. Lake Opinicon, Canada Lake • Investigate the distribution and abundance of fish species and their prey invertebrates by comparing them (i) before and after the milfoil invasion and (ii) in communities of M. spicatum and Potamogeton-Vallisneria. • • • • • • Other studies have found that different macrophytes support different invertebrates > often attributed to differences in plant architecture resulting in differences in microhabitats available to invertebrates. The three plant species. used in the study each had different dissectedness of its stems/leaves (architecture), and different locations in the littoral zone (depth) of the lowland river studied Greatest abundance was found at all times (or with decreasing flow) with the plant that was the most structurally complex and occupying the shallowest position in the littoral zone. However it often had lower associated invertebrate richness when compared to the other two, structurally simpler and deeper, plant species. Water depth and total plant biomass often correlated with invertebrate abundance/richness, however the relationships were different for each of the three plant species/ Species of inverts are not evenly distributed across the species of macrophytes. Water levels and their influence on macrophytes as invertebrate habitat may play an integral part in determining invertebrate abundance /richness and assemblages in rivers. M. spicatum supported fewer isopods, chironomids, and trichopteran larvae. In the benthos beneath the native macrophyte beds, five major taxa of prey invertebrate were five to seven times more abundant. Foliage of Potamogeton plus Vallisneria supported twice as many invertebrates per square metre than M. spicatum. Emergence of invertebrates was twice as high above native beds. These results may be due to the greater plant biomass and diversity as previously found. Some authors note that plants with finely divided leaves provide better substrate for invertebrates than those with simple leaves, however it has also been found that M. spicatum had the poorest fauna of eight species studied, consistent with the present study. 17 Effect of macrophytes on aquatic invertebrates – a literature review Reference Location Krecker, H. F. (1939). A comparative study of the animal population of certain submereged aquatic plants. Ecology 20(4): 553-562. Lake Eerie, USA Waterbody Lake Objective/Aim Findings • • Compare the invertebrates present on seven different macrophyte species. • • • • • • Lodge, D. M. (1985). Macrophytegastropod associations: observations and experiments on macrophyte choice by gastropods. Freshwater Biology 15: 695-708. Radley Pond England Lake Lodge, D. M. (1991). Herbivory on freshwater macrophytes. Aquatic Botany 41: 195-224. Literature Review Literature Review • • • • Distribution and abundance of several periphyton pulmonate snails described. Associations between macrophyte species tested. • • Sweep sampling method most effective. Preference for macrophytes by snails linked to periphyton preference. Literature review of grazers (moose, snails and carp) Experimental work (crayfish) • Nutrient concentrations not limiting grazing on macrophytes Preference unrelated to commonly measured plant physical and chemical characteristics (such as cellulose content, micronutrients etc.) Some evidence that phenolic content may be a grazing deterrent. • • Nichols, S. A. and Shaw, B. H. (1986) Ecological life histories of the three aquatic nuisance plants, Myriophyllum spicatum, Potamogeton crispus and Elodea canadensis. Hydrobiologia, 131: 3-21. North America Both • Sampled by snipping lengths of macrophytes and standardising data to 20 foot lengths. Macroinvertebrate identified to genus where possible. Species richness: Elodea had the highest number of taxon (6) and Vallisneria the least (4). Abundance: Myriophyllum spictatum had the greatest abundance and Vallisneria the least. Chironomids, caddis larva and flatworm found on all species. Annelids abundant on Myriophyllum and chironomids abundant on Potamogeton crispus. Macrophytes with dissected leaves suit annelids and chironomids, Elodea less suitable but Hydra abundant. Define the life history characteristics of Myriophyllum spicatum, Potamogeton crispus and Elodea canadensis (serious aquatic nuisances in many regions of the world) which allow them to compete so successfully in the aquatic environment and discuss what additional information is needed to better manage the plants. • • • All three species possess a number of adaptations, including an ability to rapidly propagate vegetatively, an opportunistic nature for obtaining nutrients, a life cycle that favours cool weather, and a number of mechanisms which enhance photosynthetic efficiency. The three species provide benefits to the ecosystem through their roles in primary productivity, therefore management should take this into account. The life history information of the species in question is incomplete and better understanding of resource gain and allocation is needed to manage all three species. 18 Effect of macrophytes on aquatic invertebrates – a literature review Reference Rooke, B. (1986). Macroinvertebrates associated with macrophytes and plastic imitations in the Eramosa River, Ontario, Canada. Archiv fuer Hydrobiologie 106(3): 307-325. Location Eramosa River Canada Waterbody River Objective/Aim Findings • • • • To discover if there were differences amongst the invertebrate fauna of several real macrophytes and a few plastic imitations in a short reach of running water. • • • Sandilands, K. A. and Hann, B. J. (1996). Effect of fish and submersed macrophytes on the abundance of zooplankton in a prairie wetland. UFS (Delta Marsh) Annual Report 31: 58-62. Shiel, R. J. (1976). Associations of Entomostraca with Weedbed Habitats in a Billabong of the Goulburn River, Victoria. Australian Journal of Marine and Freshwater Research 27: 533 549. Canada Prairie Wetland • • Victoria Billabong • PVC cylinder used to sample macrophytes. Differences between macrophytes were evident. Greatest abundance on plants with most dissected leaves. Plastic imitations supported very different abundances to real plants hence leaf form not only important factor – also periphyton, surface microstructure, and organic exudates. Differences between forms of plastic plants indicate that form is important. Similar community composition of plastic to real plants. Examine the effects of planktivorous fish on the zooplankton community. Determine if submersed macrophytes provide a refuge for phytophilous zooplankton against predation from fish. • Macrophytes provide refuge for zooplankton from fish predation with fish reducing abundance in more open water before reducing the abundance within macrophyte beds. Describes the composition of microcrustacean communities associated with beds of various hydrophytes in a billabong. • • Infers partitioning into microhabitats within weed beds. Makes mention of different surface texture of different plant species favouring particular taxa. 19 Effect of macrophytes on aquatic invertebrates – a literature review Reference Strayer, D. L., Lutz, C., Malcom, H.M., Munger, K. and Shaw, W.H. (2003). Invertebrate communities associated with a native (Vallisneria americana) and an alien (Trapa natans) macrophyte in a large river. Freshwater Biology 48: 19381949. Location USA Waterbody Freshwater Tidal River Objective/Aim Findings • • Compare the macroinvertebrate faunas (including both epiphytic and benthic animals) of Trapa with those of nearby beds of Vallisneria, the species it is thought to have displaced. • • • • • Talbot, J. M. and Ward, J. C. (1987). Macroinvertebrates associated with aquatic macrophytes in Lake Alexandrina, New Zealand. New Zealand Journal of Marine and Freshwater Research 21(2): 199213. Thorp, A. G., Jones,R.C. And Kelso,D.P. (1997). A Comparison of Water-Column Macroinvertebrate Communities in Beds of Differing Submersed Aquatic Vegetation in the Tidal Freshwater Potomac River. Estuaries 20(1): 86-95. Lake Alexandrina New Zealand Lake USA Virginia Tidal Freshwater River • • • • • • Macroinvertebrates collected when samples of aquatic macrophytes were taken for biomass and productivity studies. Trophic categories identified and patterns in abundance and biomass of macroinvertebrate species in six macrophyte communities and a sub littoral community were studies over a 2-5 year period. Investigates the influence of submerged aquatic vegetation on water-column macroinvertebrate abundance and community composition. Compares open water to macrophyte macroinvertebrate assemblages. Compares macroinvertebrate assemblages between different macrophyte beds. Compares macroinvertebrate assemblages over time in macrophyte beds. • • • • • The density of epiphytic macroinvertebrates was positively correlated with plant biomass. The density of benthic macroinvertebrates was nearly unrelated to plant biomass, except for being low in entirely unvegetated areas. Effects of site and plant species on density (per unit biomass of plant) of both epiphytic and benthic macroinvertebrates were weak and inconsistent. The two macrophytes supported different kinds of invertebrates leading to an increase in biodiversity. Low DO levels in Trapa beds may reduce the amount of food available to predators. Lack of congruence among studies suggests either that the macroinvertebrate communities are so variable across beds or years that all underlying patterns are obscured or that the ecological role of an aquatic plant depends on the environmental context into which it is placed. Mostly species-specific results/discussion. Sampling was performed using a 500 micron mesh around a weighted frame that was lowered over the sample area. Using SCUBA a draw string was tightened 10 cm above the surface and plant shoots severed. Higher richness and abundance were recorded in macrophyte beds when compared to open water. Differences between different macrophyte beds were less striking with temporal influences appearing to complicate findings. Most taxa demonstrated a significant interaction between plant type and month of sampling. In a study utilising artificial plants in the same area fish predation was shown to strongly impact odonate and chironomid numbers. 20 Effect of macrophytes on aquatic invertebrates – a literature review Reference Timms, B.V. (1981). Animal communities in three Victorian lakes of different salinities. Hydrobiologia 81: 181-193. Location Australia Victoria Waterbody Lakes Objective/Aim Findings • • A comparison of the effects of differing salinity in inland lakes on their animal communities (vertebrates, zooplankton, net invertebrates). • • Wollheim, W. M. and Lovvorn, J. R. (1996) Effects of macrophyte growth forms on invertebrate communities in saline lakes of Wyoming High Plains. Hydrobiologia, 323: 83-96. USA Wyoming Saline Lakes • • Explores the effects on invertebrate communities of changes in macrophyte species and growth forms between oligosaline (0.8-4.2mS/cm) and mesosaline (9.3-23.5 mS/cm) lakes of the Wyoming High Plains. Paper not relevant • • Invertebrate communities were compared (average similarities) between Triglochin, Scirpus, Potamogeton, Myriophyllum and Ruppia. Observed differences were attributed to two factors. 1) Basic plant morphology, and 2) closeness of the leafy parts of the plant to the substratum. The emergent Scirpus has round stems while Triglochin leaves are thick and ribbon-like and hence both provide similar gross physical features. Myriophyllum and Ruppia are both fine leaved but do not harbour similar communities because Ruppia grows close to the substratum while Myriophyllum has long stalks and most of its leaves are clustered near the surface. The community in Potamogeton was intermediate between those of Ruppia and Triglochin since a good proportion of the plant is near the substratum, yet its leaves are broad. Conclude weed-bed structure is the dominant factor. N/A 21 Effect of macrophytes on aquatic invertebrates – a literature review Appendix 2 - Experimental considerations Sampling methods and options There are many approaches to sampling macrophytes that vary according the objectives of the study and study location. Such methods include sweep nets (e.g. Lodge 1995), SCUBA divers or snorkellers that use mesh nets or other devices to remove plants (e.g. Biggs and Malthis 1982, Dvorak and Best 1982, Talbot and Ward 1986, Thorp et al. 1997, Cheruvelil 2001), box samplers (Downing and Cyr 1985, Downing 1986), coring tubes that cover plants and press into the substratum (e.g. Strayer et al. 2003) or other forms such as tong-like devices that snip off sections of plant (e.g. Marklund 2000). The appropriateness of a particular method will vary according to the macrophyte species sampled, water depth, water clarity and the target organisms to be collected (e.g. benthic, epiphytic etc.). Mesocosm experiments (e.g. large tubs) have also been used to test differences between macrophyte species with respect to invertebrate preference and community composition (e.g. Balci and Kennedy 2003) – these studies appear less common than studies undertaken in waterbodies. Experiments using artificial macrophytes (i.e. plastic plants) have been used and is worthy of consideration (e.g. Rooke 1986). Temporal considerations The plant community may change during the growing seasons, with a resultant change in community composition and dominance of species. Wetlands are very dynamic environments where macroinvertebrate community structure may vary throughout the growing season. (Thorp et al. 1997). Hence, temporal effects may be significant as has been found by many researchers (e.g. Humphries 1996, Thorp et al. 1997, Balci and Kennedy 2003). It has been postulated this may be due to changes in epiphyton growth (Miller et al. 1989, Cattaneo et al. 1998). Experimental considerations • • • The scale of the experiment, i.e. whether a microcosm, mesocosm or wetland design is appropriate. The target organisms – i.e. whether to target a limited suite of invertebrate species or examine effects at the community level. The timing of the experiment – be mindful of likely temporal effects. ISBN 978-1-74152-895-4 (print) ISBN 978-1-74152-901-2 (online) ISSN 1326 6446