Fifteenth Australian Weeds Conference

Different weeds, different habitats, same effects: exotic grass invasion in tropical woodlands and wetlands

Michael M. Douglas1, Samantha A. Setterfield1, Ruth A. O’Connor1,2, Keith Ferdinands1, Natalie A. Rossiter1, Kristine J. Brooks1, Ben Ryan1 and Catherine Parr3 1 Charles Darwin University and Savannas CRC, Darwin, Northern Territory 0909, 2 Present address: CRC Freshwater Ecology, University of Canberra, Canberra, Australian Capital Territory 2601, Australia 3 CSIRO Sustainable Ecosystems and Bushfire CRC, Winnellie, Northern Territory 0822, Australia

Summary There are more than 160 species of natu- and causes an obvious and dramatic change in ralised exotic grasses in northern Australia and these structure, such as the invasion of native grassland by have invaded a range of freshwater and terrestrial woody shrubs and trees. The spread of Melaleuca environments. We have been studying the impacts of (Melaleuca quinquinervia Cav.) in ’s Everg- two exotic grasses, gamba grass (Andropogon gayanus lades (Gordon 1998) and mimosa (Mimosa pigra L.) Kunth.) and para grass ( mutica Forssk.), on the floodplain grasslands of the Northern Territory which occur in tropical savanna and floodplains, (Braithwaite et al. 1998) are good examples of plant in- respectively. Despite the very different habitats they vasions causing structural change of a native grassland. occupy, there are striking similarities in their impacts Far less attention has been given to the effects of weed on native vegetation and invertebrates. In both habi- invasions where the invader is a functionally similar tats, exotic grass invasion results in a shift in plant species, such as the invasion of native grasslands by composition and a reduction of native plant species exotic grasses (Reed et al. 2005). richness. These changes in plant communities have the Displacement of native grasses by exotic grasses potential to undermine biodiversity conservation goals; is particularly relevant across northern Australia where however, our findings suggest that not all biota are native grasses traditionally dominate the understorey affected in the same way. Although there was a clear but the search for improved pasture species for cattle change in the composition of wetland and terrestrial production has left more than 150 species of natural- flora (diversity and biomass) following weed invasion, ised exotic grasses (van Klinken 2004). Some of these freshwater macroinvertebrate communities and ter- exotic grasses have become well established in areas restrial ant communities showed little or no response. outside pastoral lands, including national parks and This is particularly interesting given that both these rural residential areas. The degree to which these exotic invertebrate groups are considered to be sensitive indi- grasses threaten the natural ecosystems of northern cators of environmental change. Although these exotic Australia is very contentious but until very recently has almost completely displace the native vegetation probably been underestimated for two reasons. Firstly, in both environments, in some respects it is simply exotic grass invasion in the woodland and floodplain one grass being replaced by another and the change in ecosystems in the region does not lead to an obvious habitat structure appears to be insufficient to affect the and immediate structural change; superficially it is invertebrate communities. We suggest that, although simply a case of one sort of grass being displaced by weeds may pose a clear threat to plant biodiversity, the another. Secondly, until very recently there was little response of some faunal groups will be determined scientific data documenting the effects of exotic grass largely by the degree of structural change. invasion in these ecosystems. Keywords Weed risk management, Northern To address this gap in knowledge, we have been Territory, stakeholder engagement. studying the impacts of two exotic grass species which are native to Africa, but which occupy very INTRODUCTION different habitats. Gamba grass forms very tall clumps Although weed invasion is widely recognised as a ma- and it invades terrestrial habitats mostly in savanna jor threat to global biodiversity (Vitousek et al. 1996) woodlands and para grass is a semi-aquatic grass that there are still relatively few studies that have actually occurs in rivers, floodplains and riparian zones. This documented the impacts of weeds on biodiversity, paper reviews some of the findings of this work and particularly the effects on fauna (Groves 2002). Many considers the effects of grass invasions on native plant of the studies describing weed impacts have been in communities, habitat structure and invertebrates. situations where the invader is functionally distinct

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METHODS not consistently different to that of the native grasses. Over the past seven years we have taken a comprehen- Para grass has similar total surface area to wild rice sive approach to examine the effect of gamba grass and similar biomass to hymenachne. Secondly, stable and para grass invasion on aspects of biodiversity isotope analysis has revealed that the aquatic food webs and ecosystem processes in savanna woodland and in this system are driven largely by epiphytic algae floodplain habitats. Table 1 summarises the focus of growing on the grasses (Douglas et al. 2002, 2005). these studies. Details of the methods are given in the The grasses themselves are not directly contributing references cited in Table 1, but our work has gener- to the food web and all three grasses represent similar ally been done as comparative studies in replicated environments to support algal growth. areas with and without the exotic grasses. Work on The finding for aquatic invertebrates is in stark gamba grass was done mostly in Wildman Reserve contrast to that found for the terrestrial invertebrates and Litchfield National Park and work on para grass inhabiting the emergent parts of these grasses in the was done on the Magela Creek floodplain in Kakadu wet season. Douglas and O’Connor (2004a) found National Park. that invertebrate species richness in hymenachne was about double that of both para grass and wild rice. They RESULTS AND DISCUSSION attributed this difference to the fact that although all Para grass impacts Plant community composition three grass species had similar emergent cover, they shows major changes in response to para grass inva- differed in their nutritional quality (with hymenachne sion (Douglas et al. 2002). Para grass communities having much higher Nitrogen content) and that the ter- are distinct from the native hymenachne (Hymenachne restrial invertebrate fauna was dominated by groups acutigluma Steud.) and wild rice (Oryza australiensis that were directly consuming the grasses, such as Domin.) communities which they displace. Para grass grasshoppers and caterpillars. dominates where it invades and has consistently lower plant species richness than either of the native plant Gamba grass impacts Gamba grass invaded areas communities (Douglas et al. 2002). have distinctive plant communities that have about Despite the marked effects on plant communities, half the plant species richness of native grass sites (K. there is no evidence that para grass invasion affects Brooks unpublished data). Aspects of habitat structure aquatic invertebrates (Douglas and O’Connor 2003). are altered with gamba grass invasion as it has much Aquatic invertebrate richness in para grass was simi- higher biomass than native grasses and, unlike the lar, and certainly no lower, than that in wild rice or native annual grasses, it remains vertical throughout hymenachne sites and there was no evidence of any the dry season (Rossiter et al. 2003). differences in invertebrate community composition Despite these major changes in the plant commu- between the three grasses. Douglas and O’Connor nities, the composition and species richness of the ant (2003) attributed this apparently paradoxical finding to fauna in sites with high and low density gamba grass is two things. Firstly, the habitat structure of para grass is not substantially different to that in native grass sites.

Table 1. Studies on the impacts of para grass and gamba grass invasion. Focus of study Para grass Gamba grass Plant community Douglas et al. 2002 Setterfield et al. in prep. Douglas et al. in prep. K. Brooks (unpublished) Ferdinands et al. 2005 Invertebrate community Douglas and O’Connor 2003 (Aquatic) Ryan 2005 (Ants) Douglas and O’Connor 2004b (Terrestrial) Vertebrate community Douglas et al. 2002 (Fish) Ferdinands et al. 2005 (Birds) Habitat structure Douglas and O’Connor 2003 (Surface area) Rossiter 2001 (Surface area) Douglas and O’Connor 2004a (Biomass) Rossiter et al. 2003 (Biomass) Ecosystem processes Douglas et al. 2002 (Nutrient cycling) Rossiter et al. 2003 (Fire regime) Douglas et al. 2002, 2005 (Carbon flow) Rossiter (unpublished) (Nutrient cycling) Rossiter et al. 2004 (Water cycling)

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Ryan (2005) attributed this to the fact that whilst the REFERENCES vertical habitat structure had been altered by gamba Braithwaite, R.W., Lonsdale, W.M. and Estbergs, J.A. grass invasion, there was no difference in canopy (1989). Alien vegetation and native biota in tropi- cover and components of the ground cover (e.g. litter, cal Australia: the spread and impact of Mimosa bare ground, and grass cover of some kind) were very pigra. Biological Conservation 48, 189-210. similar between native and exotic grass sites. The simi- D’Antonio, C.M. and Vitousek, P.M. (1992). Biologi- larity in ground cover variables is particularly relevant cal invasions by exotic grasses, the grass/fire cycle, given that the pitfall trapping used in the study would and global change. Annual Review of Ecology and be biased towards collecting epigaeic ants. Systematics 23, 63-87. Douglas, M.M., Bunn, S.E., Pidgeon, R.J.W., Davies, CONCLUSION P.M., Barrow, P., O’Connor, R.A. and Winning, M. The studies reviewed above add to a growing body (2002). Weed management and the biodiversity of evidence demonstrating that exotic grass invasion and ecological processes of tropical wetlands. has a number of negative impacts on native plant com- Final report to Environment Australia and Land munities. These include the immediate effects such as and Water Australia. the loss of native plant biodiversity and a shift in plant Douglas, M.M. and O’Connor, R.A. (2003). Effects composition, but there also likely to be longer-term of the exotic macrophyte, para grass (Urochloa effects brought about by the increase in dry season mutica), on benthic and epiphytic macroinverte- biomass and hence fuel loads and the negative ef- brates of a tropical floodplain. Freshwater Biology fects of this on the tree canopy cover (Douglas and 48, 962-71. O’Connor 2004b, Rossiter et al. 2003). These effects Douglas, M.M. and O’Connor, R.A. (2004a). Ef- of exotic grass invasion on the native plant commu- fects of para grass (Urochloa mutica (Forssk.) nities were consistent in both the floodplain and the T.Q.Nguyen) invasion on terrestrial invertebrates savanna woodland. of a tropical floodplain. Proceedings of the 14th The effects of exotic grasses on invertebrate Australian Weeds Conference, eds B.M. Sindel fauna were variable and appear to depend on the ef- and S.B. Johnson, pp. 153-6. (Weeds Society of fect of grass invasion on habitat structure and on the New South Wales, Sydney). functional relationship between the invertebrates and Douglas, M.M. and O’Connor, R.A. (2004b). Weed the grass community. The shift from native grasses to invasion changes fuel characteristics: para grass exotic grass had no effect on ants or benthic or aquatic (Urochloa mutica (Forssk.) T.Q.Nguyen) on a invertebrates, whereas invertebrates inhabiting emer- tropical floodplain. Ecological Management and gent floodplain vegetation were more abundant and Restoration 5, 147-8. diverse on one of the native grasses. Para grass invasion Douglas, M.M., Bunn, S.E. and Davies, P.M. (2005). had little effect on submerged biomass or surface area River and wetland food webs in Australia’s wet-dry and gamba grass had little effect on ground cover. tropics: general principles and implications for So, from an invertebrate’s point of view, the dis- management. Marine and Freshwater Research placement of native grass by exotic grass probably 56, 329-42. represents a fairly subtle change in habitat structure. Ferdinands, K., Beggs, K. and Whitehead, P. (2005). Unless the exotic grass is nutritionally poorer than Biodiversity and invasive grass species: multi- the native species and the invertebrates are directly ple-use or monoculture? Wildlife Research 32, consuming grass, it appears that many invertebrates 447-57. will be insensitive to this sort of weed invasion. Gordon, D.R. (1998). Effects of invasive, non-in- Therefore, despite dramatic negative effects on digenous plant species on ecosystem processes: the native flora, it cannot be assumed that exotic lessons from Florida. Ecological Monographs grass invasion will lead to direct flow-on effects on 8, 975-89. all components of the fauna. Groves, R.H. (2002). The impacts of alien plants in Australia. In. ‘Biological invasions: economic ACKNOWLEDGMENTS and environmental costs of alien plant, animal The research in this paper has been funded by Land and and microbe species’, ed. D. Pimentel, pp. 11-23. Water Australia, the Tropical Savannas Management (CRC Press, New York). CRC, Charles Darwin University and CSIRO. Access Reed, H.E., Seastedt, T.R. and Blair, J.M. (2005). Eco- to sites was provided by the Parks and Wildlife Com- logical consequences of C4 grass invasion of a C4 mission of the Northern Territory and the Traditional grassland: a dilemma for management. Ecological Owners of . Applications 15, 1560-9.

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Rossiter, N.A. (2001). Comparative ecophysiology and Ryan, B. (2005). The effects of Gamba grass (Andropo- fire ecology of native and exotic savanna grasses. gon gayanus) invasion on ant communities. Hon- Honours thesis, Northern Territory University, ours thesis, Charles Darwin University, Darwin. Darwin. van Klinken, R., Panetta, D.F., Ross, B. and Wilson, C. Rossiter, N.A., Setterfield, S.A., Douglas, M.M. and (2004). A pain in the grass: what’s the diagnosis? Hutley, L.B. (2003). Testing the grass-fire cycle: Proceedings of the 14th Australian Weeds Confer- alien grass invasion in the tropical savannas of ence, eds B.M Sindel and S.B. Johnson, pp. 480-3. northern Australia. Diversity and Distributions (Weeds Society of New South Wales, Sydney). 9, 169-76. Vitousek, P.M., D’Antonio, C.M., Loope, L.L. and Rossiter, N.A., Setterfield, S.A., Douglas, M.M., Hut- Westbrooks, R. (1996). Biological invasions as ley, L.B. and Cook, G. (2004). Exotic grass inva- global environmental change. American Scientist sion in the tropical savannas of northern Australia: 84, 468-78. Ecosystem consequences. Proceedings of the 14th Whitehead, P. and Wilson, C. (2000). Exotic grasses Australian Weeds Conference, eds B.M. Sindel and in northern Australia: species that should be sent S.B. Johnson, pp. 168-71. (Weeds Society of New home. Proceedings of the Northern Grassy Land- South Wales, Sydney). scapes Conference. (Tropical Savannas CRC, Darwin).

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