Int J Biometeorol (2006) 50: 292304 DOI 10.

1007/s00484-005-0019-2

ORIGINA L ARTI CLE

Lynda E. Chambers . Richard H. Loyn

The influence of climate variability on numbers of three waterbird species in Western Port, Victoria, 19732002

Received: 1 April 2005 / Revised: 9 November 2005 / Accepted: 14 November 2005 / Published online: 25 January 2006 # ISB 2006

Abstract Seasonal and annual movements of Australian waterbirds are generally more complex than those of their Northern Hemisphere counterparts, and long-term data are needed to understand their relationships with climatic variables. This paper explores a long-term (19732002) set of waterbird counts from coastal Victoria and relates them to climatic data at local and continental scales. Three species (Black Swan Cygnus atratus, White-faced Heron Egretta novaehollandiae and Grey Teal Anas gracilis) were chosen for this analysis. Black Swans have large local breeding populations near the study region; White-faced Herons have smaller local breeding populations and Grey Teal breed extensively in ephemeral inland floodplains, such as those in the Murray-Darling Basin. All showed significant relationships with streamflow, regional rainfall and the Southern Oscillation Index (SOI) at appropriate scales and time-lags, with streamflow explaining the most variance. Black Swans showed a strong seasonal cycle in abundance and local climate variables had the greatest influence on the counts. Numbers were positively correlated with streamflow in southern Victoria three to six seasons before each count. Broader-scale climatic patterns were more important for the other two species. Numbers of White-faced Herons were positively correlated with streamflow or rainfall over various parts of Australia seven to nine seasons before each count. Numbers of Grey Teal showed weak seasonal cycles, and were negatively correlated with rainfall in Victoria or the Murray-Darling Basin in the seasons before or during each count, and positively with streamflow in the Murray-Darling Basin 1518 months before each count.
L. E. Chambers (*) Bureau of Meteorology Research Centre, G.P.O. Box 1289 Melbourne, Vic 3001, Australia e-mail: L.Chambers@bom.gov.au Tel.: +613-9669-4784 Fax: +613-9669-4660 R. H. Loyn Arthur Rylah Institute for Environmental Research, Department of Sustainability & Environment, P.O. Box 137 Heidelberg, Vic 3084, Australia

Keywords Australia . Climate . Rainfall . Streamflow . Waterbirds

Introduction
Climatic variations can have profound effects on ecological and human environments, and many of the ecological effects remain poorly understood despite enormous advances in meteorology (Nicholls 2001; Stevermer 2001; Pearce 2002). Seasonal and annual movements of Australian waterbirds are generally more complex and less predictable than those of their Northern Hemisphere counterparts (Frith 1982; Kingsford and Norman 2002). This is partly related to the physiography and erratic climate of Australia (Pittock 2003; Wright 2004) and, in particular, to the highly variable wetland and floodplain systems. Much of Australia experiences an arid climate, and wetlands may be dry for many years before filling with flushes of water originating from rainfall events hundreds of kilometres away. These events can occur at any time of year, as a consequence of weather patterns in the tropical north (where most rain falls in summer) or the temperate south (where more rain falls in winter and spring, and snow melts from the mountains in spring). For waterbirds, population boom periods can be initiated by such flooding, and bust periods can follow months or years of drought. Even in a dry year, ephemeral floodwaters from distant rains may allow arid parts of Australia to support many millions of waterbirds (Roshier et al. 2001a,b). In the Northern Hemisphere, most waterfowl undertake regular and predictable migrations, in which most of the population moves between breeding areas and wintering areas each year in response to regular changes in day-length and temperature (Cramp and Simmons 1977; del Hoyo et al. 1992). Seasonal freezing and thawing of wetlands can also induce large-scale movements in the Northern Hemisphere. Winters in Australia are generally benign, with temperatures rarely dropping below freezing point except in southern mountains where there are few wetlands. Most endemic Australian waterbirds are nomadic or known to move irregularly and, in the past, it was thought

293

that waterbirds used the temporary wetlands of Australias arid interior opportunistically from their preferred habitats in more mesic regions of southern and eastern Australia (Roshier et al. 2001a). For the ducks of Australia, as well as various other Australian birds that use the arid zone, the timing and length of the breeding season can be quite erratic, although underlying seasonal patterns are evident in more temperate areas (Briggs 1977, 1992; Halse and Jaensch 1989; Marchant and Higgins 1990; Geering et al. 1998; Roshier et al. 2001a; Kingsford and Norman 2002). During summer in southern Australia, waterbirds tend to concentrate in coastal regions with more regular and constant rains, and hence more permanent wetlands (Frith 1982). Mass movements to coastal regions are also known to occur during prolonged dry periods (Gentilli and Bekle 1983; Gosper et al. 1983; Loyn et al. 1994; Kingsford and Norman 2002). One such coastal refuge is Western Port, a coastal embayment in Victoria, south-eastern Australia. This paper provides a case study documenting effects of climatic fluctuations over a 29-year period on the abundance of selected waterbird species in Western Port. The purpose of this paper is to explore the influence of broadscale and local climate on the abundance of selected waterbird species in Western Port, over a longer time-frame than has been attempted previously for this or other Australian wetlands. Studies, such as this one, provide quantitative data on the effects of changes in climate on waterbird numbers and provide useful information for assessing how waterbirds may respond to future climate change. Recent studies suggest that many eastern Australian wetlands may experience reduced rainfall, increased evaporation, and decreased surface-water flows in the future (e.g. Chambers et al. 2005).

hollandiae) and the Grey Teal (Anas gracilis). All three species are commonly seen in Western Port, and previous studies indicate that they respond in different ways to season and climate (Loyn et al. 1994). The Black Swan has been defined as nomadic and prone to erratic dispersal (Pringle 1985). However, Marchant and Higgins (1990) note that when suitable habitat is permanently available the adults are sedentary, but the young, and adults from ephemeral habitats, can move far and often. The species breeds commonly on wetlands close to Western Port and is classed as a local breeding species (Loyn et al. 1994). White-faced Herons are found in Western Port throughout the year and some breed locally, although there is a general decrease during the spring breeding season (Loyn et al. 1994). White-faced Herons are classified as sedentary or locally nomadic, and may move away from the coast during the breeding season (Pringle 1985; Marchant and Higgins 1990). The Grey Teal is widely distributed across Australia and generally considered to be nomadic, responding rapidly to changes in rainfall and hydrology (Frith 1982; Gentilli and Bekle 1983; Marchant and Higgins 1990; Loyn et al. 1994; Kingsford and Norman 2002). It is an important game species in states where duck hunting is permitted (including Victoria) (Norman and Powell 1981; Briggs et al. 1985a; Loyn 1991). Few Grey Teal breed in the Western Port region, and most breeding is believed to occur in ephemeral swamps away from the coast. Waterbird counts Teams of observers made simultaneous counts of all waterbirds on selected days, usually at high tide, at strategic sites around Western Port, including French and Phillip Islands (Fig. 1). The 17 most important sites for waterbirds were counted relatively consistently across survey dates, and this paper focuses on data from these 17 sites. From October 1973 to June 1976, counts were made every month with the day selected so that high tide fell within 4 h of midday and the day was on a weekend (Loyn et al. 1994). Since June 1976, counts were made five times per year, twice in summer (December and February) and once in autumn (April or May), winter (June, July or early August) and spring (September or October). The spring and autumn counts were discontinued from 1996. Over 500 voluntary observers have taken part in the study since its inception (Heislers 2003). The fieldwork is described more fully in previous papers (Loyn 1978; Dann et al. 1994; Loyn et al. 1994, 2001; Heislers 2003). This paper uses seasonal count data from December 1973 to February 2002 (127 dates). Summer count values were taken as the maximum of the December and February counts, but almost identical results were obtained when the mean of the December and February counts was used.

Materials and methods

Study area Situated to the south east of Melbourne, the large coastal bay known as Western Port (Fig. 1) covers an area of around 67,000 ha. Western Port is one of the most important wader and waterfowl sites in Australia, in terms of numbers of birds that it supports, and is listed under the Ramsar Convention (Loyn et al. 2001). Since October 1973, regular counts of waterbirds have been made in this region by the Bird Observers Club of Australia, making it one of the first large sites in Australia with a long record of waterbird abundance (Loyn 1978; Dann et al. 1994; Loyn et al. 1994, 2001; Heislers 2003). Bird species considered Three species were chosen for this study: the Black Swan (Cygnus atratus), the White-faced Heron (Egretta novae-

294 Fig. 1 Location of Western Port and other sites mentioned in the text

Allowance for variable coverage The analysis was restricted to 17 sites that were included in the regular survey and counted on most dates. Nevertheless, some of these sites were missed occasionally, for example when high winds prevented the use of small boats. Allowance for variable coverage was made using an initial Analysis of Variance, as follows. First, data were examined for each species across the 17 sites, with a view to excluding any sites rarely used by that species. This proved necessary for Grey Teal (6 sites retained), but not for Black Swan and White-faced Heron, which occurred commonly at most sites. Secondly, a two-way Analysis of Variance was conducted for each species, with date (n=127) and site (n=17 for White-faced Heron or Black Swan, 6 for Grey Teal) as factors. This approach deliberately paid no attention to seasonal or climatic variables, or interactions between variables. Data were transformed to meet the assumptions of this analysis (fourth root for Grey Teal; square root for White-faced Heron; no transformation for Black Swan). After appropriate back-transformation, tables were generated for the means of each species for each of the 127 dates, corrected for variable coverage. These corrected means were then used as the basis for further analysis.

Climatic variables considered Three types of climatic variables are considered, operating at different spatial scales. The Southern Oscillation Index (SOI) operates at broad global scales, affecting weather through the Pacific region (Allan et al. 1996). Rainfall data were obtained for various regions of Australia, along with streamflow data for selected catchments. All variables are expected to influence the extent and productivity of wetlands in various parts of Australia (Norman and Nicholls 1991; Roshier et al. 2001a,b; Kingsford and Norman 2002). Rainfall and the Southern Oscillation Index (SOI) Area-averaged (all Australia) monthly rainfall was obtained from the Australian Bureau of Meteorologys National Climate Centre. This index is a measure of how wet Australia is as a whole, with data available from 1890 to the present (Jones and Weymouth 1997). In addition, gridded monthly rainfall data were extracted from the Australian Bureau of Meteorology, National Climate Centres 0.25 degree spatial data set (Jones and Weymouth 1997). Based on these data regional rainfall indices were also considered:

295

1. Victoria - rainfall averaged over the state of Victoria 2. Murray-Darling Basin - rainfall averaged over the Murray-Darling Basin 3. East Gippsland - averaged over the region 147.00E to 148.25E and 37.75S to 38.50S. 4. Western District (SW) - averaged over 141.00E to 144.00E, 36.50S to 38.50S 5. Western District (NW) - averaged over 141.00E to 143.00E, 34.00S to 36.00S 6. Western Port - averaged over 145.00E to 145.50E, 38.00S to 38.75S. Rainfall in these regions is thought to influence the abundance of some species of waterbirds in Western Port (Loyn et al. 1994), based on known movements and breeding distributions of waterbirds in south-eastern Australia (Norman 1971; Marchant and Higgins 1990; Barrett et al. 2003). The Southern Oscillation Index (SOI) was calculated by the National Climate Centre from monthly fluctuations in air pressure difference between Tahiti and Darwin (Troup 1967).

western Tasmania), the Western Plateau division (where there were very few streams and even fewer available data), or the southern Northern Territory (where few data were available). The drainage divisions and river basins are shown in Fig. 3. Seven regions were considered, and streamflow averaged over all gauges in the region, in relation to waterbird counts in Western Port: Gulf of Carpentaria (Division IX): 35 gauges used Northeast Coast (Division I): 74 gauges used Murray-Darling Basin (Division IV): 345 gauges used South (Division II - basins 1439, Division IV - basins 115): 337 gauges used. (These drainage basins are in the southern parts of the divisions; see Fig. 3) 5. South of Divide (Division II - basins 2139): 135 gauges used 6. Inland (Divisions X and XI): 7 gauges used 7. Tasmania (Division III): 29 gauges used These regions cover distinct climatic provinces over most of eastern Australia. This broad-scale approach is justified by the wide continental distributions of many waterbirds, including the three species under consideration (Marchant and Higgins 1990; Barrett et al. 2003). Many catchments cover vast areas, especially in arid Australia, and may receive water from thousands of kilometres distant. Recent work has highlighted that ephemeral floodwaters in arid Australia are key drivers of continental populations of many waterbird species (Kingsford et al. 1999a). Although some streamflow data were available for the western divisions (VI, VII and VIII), waterbird populations in these regions are generally considered to be isolated from those in the eastern parts of the continent, with the 1. 2. 3. 4.

Streamflow data Monthly total volume (megalitres) streamflow data were obtained, from the relevant state authorities, for a number of gauging sites around Australia (Fig. 2). The location of the stream gauges reflects the distribution of predominantly permanent or semi-permanent rivers and streams, most located along coastal areas and southern regions of the continent. No data were collected for the northern New South Wales coastal region, Hydro Tasmania sites (south
Fig. 2 Locations of stream gauge sites for which data were obtained. Plotted over the region 10S to 45S, 110E to 155E

Northern Territory WesternAustralia

Queensland

South Australia New South Wales

Victoria Tasmania

296 Fig. 3 Australian drainage divisions and drainage basins mentioned in the text

central arid zone inhibiting regular movement (e.g. Gentilli and Bekle 1983). Subsequent analysis of all available streamflow data (not shown) confirmed that streamflow in eastern Australia was more highly correlated with waterbird numbers in Western Port than streamflow in the west of the continent. Correlations between the species counts were compared, both with and without adjustments for season in all analyses. Initially, relationships between the climate parameters and waterbird counts were investigated using correlation analysis. To allow for possible seasonal variations in counts, independent of climate, counts for each season were considered separately. As climate is unlikely to have an instantaneous effect on the number of waterbirds counted in Western Port, particularly in more distant regions of Australia, lags of up to 16 seasons were considered in the correlation analysis. This allows for the climatewaterbird system to have a memory of up to 4 years. However, little credence was placed on correlations involving long time lags and distant regions, especially for those involving data from northern Australia, as most of the known movements of the three species considered are concentrated within the temperate and arid parts of Australia and not the tropical north (Marchant and Higgins 1990). There are several reasons for considering a range of time lags (Briggs 1977; Norman and Nicholls 1991). Water can take many months to flow from the tropics or the Great Dividing Range, where it falls as rain, to inland wetlands where it provides breeding habitat for waterbirds (Kingsford et al. 1999a). Waterbirds may use these habitats

for several years until they dry out, forcing birds to seek refuge elsewhere including coastal habitats such as Western Port. The seasonal count data were also combined for each species to form an annual series with four values per year (summer, autumn, winter and spring). This data set was analysed using General Linear Models with an indicator for season and a climate covariate (for example, AllAustralian Rainfall). General Linear Models were used to look for relationships between climate variables, such as rainfall, and waterbird counts, while taking into account the seasonal cycle in the count data. This allowed the modelling of both seasonal and climate effects on the waterbird counts and gives an estimate of how much of the variation in the waterbird counts can be explained by seasonal and climate effects. Again, lags of up to 16 seasons were considered.

Results
Waterbird count data: seasonal and year-to-year variation There was considerable interseasonal and interannual variability in the Western Port waterbird counts. Numbers of all three species showed distinct seasonal patterns (p<0.001), generally with minimum numbers in spring and maximum numbers in summer, autumn or winter (Fig. 4). However, the pattern for Grey Teal varied greatly between years (Fig. 5), and season only explained 3.2% of variance,

297

a
10000

b
1000 8000 White-faced Heron

800

Black Swan

6000

600

4000

400

2000

200

0 Summer Autumn Winter Spring

0 Summer Autumn Winter Spring

c
700 600 500 Grey Teal 400 300 200 100 0 Summer Autumn Winter Spring

Fig. 4 Box plots of season counts for each of the three species, showing median values, the interquartile range (values within the box, the middle 50% of all counts) and extremes. a Black Swan, b White-faced Heron, c Grey Teal

compared with 18.2% for White-faced Heron and 41.7% for Black Swan. Numbers of White-faced Herons and Black Swans showed a positive correlation with each other (r=0.271, p=0.007) when considered on the basis of 127 individual counts with no regard to season (reflecting the similar seasonal cycles for these species, with peaks in autumn and troughs in spring). Numbers of White-faced Herons and Grey Teal showed a weaker positive correlation when considered on the basis of standardised counts (allowing for seasonal effects) (r=0.229, p=0.023). No other significant correlations were found between seasonal or annual counts of the three species. On an annual basis there was little relationship between the species counts. Because the seasonal effect was so strong for two of the three species, subsequent analyses are based on generalised linear models after allowing for the effects of season. Black Swans were particularly numerous in 19731977 and 19941995. White-faced Herons were most numerous near the start of the survey, particularly in 1976. Grey Teal showed an erratic pattern of peaks, with the major ones occurring in 1975, 1977, 1982, 1992 and, to a lesser extent, 2001. The lowest annual averaged counts of Grey Teal occurred in 1974, 1988, 1996 and 2000.

Relationships of waterbird counts with climatic data All three species showed significant relationships with the SOI (Tables 1, 2, 3), regional rainfall (Tables 1, 2, 3) and streamflow (Tables 4, 5, 6), with appropriate seasonal lags. In each case, streamflow explained more of the count variance than the SOI or regional rainfall. Streamflow data explained up to 10.2% of the variance for Black Swan, 14.5% of the variance for White-faced Heron and 34.5% of the variance for Grey Teal, having allowed for seasonal effects in each case (Tables 4, 5, 6). The SOI was the poorest predictor of waterbird numbers, explaining a maximum of 2.8% of the count variance for Black Swan, 9.1% of the variance for White-faced Heron and 4.7% for Grey Teal (Tables 1, 2, 3). Regional rainfall explained up to 6.2% of the variance for Black Swan, 12.2% for Whitefaced Heron and 9.9% for Grey Teal, having allowed for seasonal effects in each case (Tables 1, 2, 3). The SOI, regional rainfall and streamflow predictors performed quite similarly to each other for White-faced Heron, but very differently for the other two species (with SOI and regional rainfall being markedly inferior to streamflow data). These relationships are explored further below.

298

10000 600 8000 GreyTeal 400 White.facedHeron 1975 1980 1985 1990 1995 2000 BlackSwan

1000 800 600 400 200

6000

4000

200 2000

0 1970 1975 1980 1985 1990 1995 2000

0 1970

0 1970 1975 1980 1985 1990 1995 2000

b
4 Black Swan Anomaly 4 Grey Teal Anomaly 3 2 1 0 -2 1970 1975 1980 1985 1990 1995 2000 -1 1970 1975 1980 1985 1990 1995 2000 White-faced Heron Anomaly 4 2

-2 1970 1975 1980 1985 1990 1995 2000

Fig. 5 a Adjusted species counts over time. b Seasonal anomalies (seasonal mean subtracted and result divided by seasonal standard deviation) of species counts over time

SOI - Waterbird relationships The SOI for the current season made a negative contribution to models for numbers of Black Swans (p=0.032), but the relationship was weak and explained only 2.8% of variance (Table 1). SOI for 13 seasons before the count made positive contributions to models for numbers of White-faced Herons, explaining 9.1% of variance with the 3-season lag (p=0.001; Table 2). SOI for 58 seasons before the count made positive contributions to models for numbers of Grey Teal, explaining 4.7% of variance with the 7-season lag (p=0.030; Table 3).

Similar results were obtained when counts for each season were considered separately (not shown), with the spring species counts having the strongest relationships with the SOI (for up to six seasons lag). When the SOI was high, spring waterbird counts tended to be low in all three species, particularly the Grey Teal and White-faced Heron. Rainfall - Waterbird relationships Rainfall data from various parts of Victoria made significant though weak contributions to models for numbers of

Table 1 Partial correlations between Black Swan count data and seasonal values of the Southern Oscillation Index (SOI) or regional rainfall; after accounting for season Lag SOI Rainfall All-Australia Victoria 0 1 2 3 4 5 6 7 8 9 10 0.219# 0.158 0.137 0.157 0.094 0.016 0.043 0.024 0.034 0.114 0.174 0.131 0.157 0.094 0.097 0.087 0.033 0.005 0.042 0.045 0.000 0.071 0.223# 0.053 0.100 0.018 0.068 0.202# 0.264# 0.122 0.135 0.287# 0.051 Murray-Darling East Gippsland Western Dist. (NW) Western Dist. (SW) Western Port 0.160 0.060 0.038 0.106 0.091 0.107 0.061 0.024 0.047 0.148 0.056 0.256# 0.038 0.151 0.068 0.014 0.226# 0.254# 0.091 0.029 0.180 0.044 0.186 0.064 0.071 0.047 0.081 0.168 0.193 0.093 0.151 0.261# 0.053 0.197 0.078 0.080 0.026 0.100 0.136 0.229# 0.095 0.128 0.283# 0.045 0.176 0.047 0.196 0.108 0.020 0.252# 0.324## 0.173 0.102 0.295# 0.056

Lag is the number of seasons lagged, for example, 0 is simultaneous and 1 uses the previous seasons rainfall. Hashes indicate partial correlations significant at the 5% level; a single # indicating a percentage of variance explained by the SOI or rainfall in the range 2.55.0%; ## in the range 5.110.0%; and ### over 10%

299 Table 2 Partial correlations between White-faced Heron count data and seasonal values of the Southern Oscillation Index (SOI) or regional rainfall; after accounting for season Lag SOI Rainfall All-Australia Victoria 0 1 2 3 4 5 6 7 8 9 10 0.158 0.230# 0.305## 0.333## 0.231# 0.210# 0.177 0.233# 0.232# 0.249## 0.174 0.197 0.322## 0.123 0.109 0.174 0.267## 0.081 0.251## 0.386### 0.263## 0.144 0.028 0.105 0.059 0.089 0.082 0.215# 0.045 0.248## 0.238# 0.303## 0.125 Murray-Darling East Gippsland Western Dist. (NW) Western Dist. (SW) Western Port 0.175 0.154 0.060 0.047 0.084 0.169 0.055 0.165 0.226# 0.274## 0.116 0.067 0.014 0.055 0.118 0.092 0.140 0.223# 0.195 0.066 0.105 0.013 0.086 0.196 0.062 0.059 0.079 0.196 0.029 0.297## 0.260## 0.307## 0.162 0.090 0.102 0.072 0.071 0.040 0.177 0.004 0.169 0.168 0.293## 0.112 0.090 0.046 0.033 0.008 0.041 0.163 0.015 0.183 0.133 0.167 0.016

Lag is the number of seasons lagged, for example, 0 is simultaneous and 1 uses the previous seasons rainfall. Hashes indicate partial correlations significant at the 5% level; a single # indicating a percentage of variance explained by the SOI or rainfall in the range 2.55.0%; ## in the range 5.110.0%; and ### over 10%

Black Swans (Table 1). Weak negative relationships were found for rainfall in the season of the count, and positive relationships with rainfall five or six seasons before each count (Table 1). Rainfall data from Western Port six seasons before each count made a positive contribution, explaining 6.2% of the variance (p=0.001; Table 1). For White-faced Herons, plausible models could be developed using rainfall data from a broader range of regions including All-Australia (Table 2). Relationships were generally positive for rainfall seven to nine seasons before each count. Rainfall data from All Australia seven seasons before each count made a positive contribution, explaining 12.2% of the variance (p<0.001; Table 2). For Grey Teal, significant negative relationships were found with rainfall in various parts of Victoria or the Murray-Darling Basin 01 seasons before each count

(Table 3). Positive relationships were found with rainfall in the same regions, or more widely in Australia, 5 or 6 seasons before each count. Rainfall data from All-Australia six seasons before each count made a positive contribution, explaining 9.9% of variance (p=0.002; Table 3). For all three species, counts during the spring showed some strong negative correlations with rainfall in Australia, particularly in south-eastern Australia in the winter of the year of the count (not shown). For the Black Swan the highest correlations incorporated the Western Port region; while for the White-faced Heron, correlations were stronger than those for the other species, and were highest over the Murray-Darling Basin. In contrast to the other species, counts of Grey Teal during autumn and winter had strong positive relationships with rainfall in north-eastern and central Australia in the summer of the year prior to the count.

Table 3 Partial correlations between Grey Teal count data and seasonal values of the Southern Oscillation Index (SOI) or regional rainfall; after accounting for season Lag SOI Rainfall All-Australia Victoria 0 1 2 3 4 5 6 7 8 9 10 0.040 0.069 0.001 0.115 0.138 0.215# 0.216# 0.221# 0.150 0.109 0.044 0.099 0.122 0.034 0.072 0.122 0.209# 0.319## 0.158 0.084 0.112 0.087 0.245## 0.207# 0.095 0.027 0.124 0.264## 0.037 0.084 0.005 0.188 0.027 Murray-Darling East Gippsland Western Dist. (NW) Western Dist. (SW) Western Port 0.201# 0.230## 0.078 0.013 0.127 0.172 0.127 0.061 0.014 0.078 0.012 0.145 0.189 0.050 0.202# 0.243## 0.111 0.051 0.020 0.071 0.015 0.010 0.224# 0.186 0.176 0.011 0.088 0.280## 0.091 0.104 0.042 0.204# 0.007 0.255## 0.130 0.098 0.024 0.069 0.216# 0.019 0.076 0.009 0.187 0.058 0.204# 0.107 0.031 0.010 0.044 0.180 0.080 0.038 0.080 0.149 0.023

Lag is the number of seasons lagged, for example, 0 is simultaneous and 1 uses the previous seasons rainfall. Hashes indicate partial correlations significant at the 5% level; a single # indicating a percentage of variance explained by the SOI or rainfall in the range 2.55.0%; ## in the range 5.110.0%; and ### over 10%

300 Table 4 Partial correlations between Black Swan count data and seasonal values of the regional streamflow; after accounting for season Lag 0 1 2 3 4 5 6 7 8 9 10 Gulf of Carpentaria 0.001 0.251# 0.076 0.001 0.082 0.135 0.033 0.107 0.053 0.120 0.000 NE Coast 0.064 0.131 0.032 0.087 0.042 0.003 0.102 0.010 0.061 0.005 0.035 Inland 0.062 0.123 0.074 0.058 0.035 0.028 0.025 0.041 0.024 0.050 0.022 Murray-Darling 0.024 0.104 0.054 0.087 0.108 0.156 0.313## 0.108 0.085 0.095 0.005 South 0.036 0.030 0.192 0.238# 0.234# 0.264# 0.417### 0.125 0.109 0.144 0.022 South of Divide 0.236# 0.300## 0.402## 0.393## 0.295## 0.295## 0.295## 0.014 0.041 0.119 0.048 Tasmania 0.080 0.035 0.112 0.061 0.092 0.262# 0.333## 0.139 0.232# 0.055 0.012

Lag is the number of seasons lagged, for example, 0 is simultaneous and 1 uses the previous seasons streamflow. Hashes indicate partial correlations significant at the 5% level; a single # indicating a percentage of variance explained by the streamflow in the range 2.5 5.0%; ## in the range 5.110.0%; and ### over 10%

Streamflowwaterbird relationships For the Black Swan, streamflow in the region South of Divide (the smallest region incorporating Western Port) up to six seasons prior to the count made positive contributions to the models and explained 3% to around 10% of the variance in the counts (Table 4). Streamflow in the broader South region (which included most of Victoria and parts of southern New South Wales), three to six seasons prior to the count, made similar positive contributions (Table 4). The best relationship was with data from the South region six seasons before each count, explaining 10.2% of variance (p<0.001; Table 4). For the White-faced Heron, similar patterns were found, but they were generally stronger for streamflow data applied to broad areas, including the Murray-Darling basin, than for local streamflow data (Table 5). Streamflow South of Divide explained little of the variance in the seasonal White-faced Heron counts, except when lagged by six or seven seasons, when it made a small positive contribution (Table 5). Streamflow in the

Murray-Darling basin and South regions, made positive contributions over a broader range of time-lags and explained up to 14.5% of the count variance (for South Region six seasons before each count, p<0.001; Table 5). Grey Teal showed an even stronger tendency than White-faced Heron for relationships to be improved when streamfall data came from broad areas of Australia, including parts of the Murray-Darling Basin. Negative relationships were found with streamflow from various regions at the time of each count (Table 6). Stronger positive relationships were found with rainfall from the Murray-Darling basin and the South Region three or four seasons before each count (Table 6). The strongest relationships were found with streamflow from the three distant parts of Australia six seasons before each count, namely the North-east Coast, Inland Australia and the Gulf of Carpentaria (p<0.001; Table 6). The streamflow data from the Gulf of Carpentaria explained 34.5% of variance and streamflow data from Inland Australia explained a similar 28.8% of variance (Table 6).

Table 5 Partial correlations between White-faced Heron count data and seasonal values of the regional streamflow; after accounting for season Lag 0 1 2 3 4 5 6 7 8 9 10 Gulf of Carpentaria 0.199 0.344## 0.076 0.046 0.173 0.266## 0.083 0.222# 0.601### 0.166 0.026 NE Coast 0.179 0.183 0.016 0.008 0.079 0.097 0.025 0.100 0.205# 0.071 0.016 Inland 0.037 0.307## 0.032 0.016 0.036 0.209# 0.016 0.001 0.578### 0.216# 0.060 Murray-Darling 0.167 0.323## 0.264## 0.214# 0.148 0.331## 0.284## 0.175 0.282## 0.200 0.014 South 0.032 0.271## 0.328## 0.247# 0.186 0.411### 0.421### 0.249## 0.207# 0.241# 0.157 South of Divide 0.073 0.048 0.058 0.171 0.112 0.150 0.202# 0.263## 0.081 0.009 0.143 Tasmania 0.039 0.129 0.270## 0.329## 0.250## 0.261## 0.124 0.197 0.131 0.148 0.070

Lag is the number of seasons lagged, for example, 0 is simultaneous and 1 uses the previous seasons streamflow. Hashes indicate partial correlations significant at the 5% level; a single # indicating a percentage of variance explained by the streamflow in the range 2.5 5.0%; ## in the range 5.110.0%; and ### over 10%

301 Table 6 Partial correlations between Grey Teal count data and seasonal values of the regional streamflow; after accounting for season Lag 0 1 2 3 4 5 6 7 8 9 10 Gulf of Carpentaria 0.014 0.033 0.074 0.016 0.107 0.324### 0.596### 0.137 0.019 0.009 0.060 NE Coast 0.048 0.056 0.024 0.004 0.097 0.133 0.380### 0.145 0.013 0.011 0.144 Inland 0.082 0.116 0.124 0.054 0.009 0.179 0.545### 0.040 0.014 0.131 0.040 Murray-Darling 0.241## 0.137 0.067 0.334### 0.252## 0.159 0.093 0.121 0.012 0.046 0.018 South 0.211# 0.151 0.083 0.405### 0.308## 0.142 0.040 0.098 0.003 0.105 0.047 South of Divide 0.276## 0.180 0.028 0.153 0.152 0.008 0.195 0.114 0.079 0.034 0.015 Tasmania 0.118 0.004 0.109 0.134 0.145 0.220# 0.021 0.114 0.017 0.139 0.048

Lag is the number of seasons lagged, for example, 0 is simultaneous and 1 uses the previous seasons streamflow. Hashes indicate partial correlations significant at the 5% level; a single # indicating a percentage of variance explained by the streamflow in the range 2.5 5.0%; ## in the range 5.110.0%; and ### over 10%

Discussion
Seasonal patterns The study showed a marked difference in consistency of seasonal pattern between the three species: Black Swans could be described as having a strong seasonal cycle, White-faced Herons a moderate seasonal cycle and Grey Teal a weak seasonal cycle. Gosper et al. (1983) also found that counts of Black Swans were more seasonal than those of White-faced Heron and Grey Teal, in a 4-year study of waterbirds in the Richmond Valley, New South Wales. The erratic movements of Grey Teal are well known: this highly mobile species breeds mainly in shallow swamps created by erratic inland flooding (Frith 1982; Gentilli and Bekle 1983; Marchant and Higgins 1990). Grey Teal disperse over long distances once these swamps dry out, appearing in coastal regions of Australia and New Zealand (Pringle 1985; Marchant and Higgins 1990). Grey Teal are rapid dispersers with individuals capable of covering 180 km per day and up to 3,200 km in a few weeks (Roshier et al. 2001a). They are usually the first ducks to reach a newly flooded area (Pringle 1985). Minimum numbers of all three species tended to occur in spring in Western Port, or winter and spring for Black Swan. This period corresponds to the main breeding season for many waterbirds in south-eastern Australia (Frith 1982; Marchant and Higgins 1990). The two locally breeding species have been observed breeding in nearby farmland and swamps, outside the study area, in these respective seasons (spring for White-faced Heron; winter and spring for Black Swan) (Loyn et al. 1994). Fewer birds are seen during counts when they have dispersed to breed in nearby wetlands. Similar conclusions have been reached from other studies in south-eastern Australia, most of which have shown that minimum numbers of waterbirds occur on specific wetlands during spring (e.g. Gosper et al. 1983; Kingsford and Norman 2002). Peak numbers of waterbirds have been reported on large waterbodies in south-eastern Australia, at various

times from late summer through autumn into winter (Frith 1982; Gosper et al. 1983; Marchant and Higgins 1990). Grey Teal can breed at almost any time of year when water is available (Frith 1982; Marchant and Higgins 1990), but in southern Australia wetlands are most likely to fill and provide good breeding conditions in spring (Dexter et al. 1986; Briggs and Thornton 1999; Roshier et al. 2001b; Kingsford and Norman 2002; Wright 2004). This is a consequence of high rainfall in winter and spring in much of south-eastern Australia, along with spring snow-melt from the high mountains. Hence the average abundance pattern for Grey Teal showed similar spring minima to the other species (Fig. 5), but it varied far more between years (Fig. 5). Loyn et al. (1994) considered that inland breeding species, such as the Grey Teal, are less likely to have a stable seasonal pattern in abundance between years than local breeders, but they also observed a similarity in average patterns when data were combined across many years. Relationships with climatic variables Many of the climatic variables were inter-correlated, and observed relationships with waterbird numbers do not necessarily provide strong evidence that birds were responding directly to a particular region, rather than another region with similar rainfall patterns. Interpretation needs to consider both the movements of the birds and the movement of the water, both of which are extensive at the continental scale (Frith 1982; Kingsford and Norman 2002). For example, we observed correlations between numbers of Grey Teal and rainfall in the Gulf of Carpentaria in far north Queensland, over 3,000 km away from Western Port. Grey Teal are known to travel as far as this (and further), but shorter movements are more common (Norman 1971). A more plausible mechanism for such correlation lies with the movement of water from the tropical north during the summer wet season into inland

302

floodplains where Grey Teal breed in large numbers for short periods (Kingsford et al. 1999a). Rainfall data from the Gulf of Carpentaria may act as a surrogate for rainfall generally in the tropical north. Water actually used by Grey Teal is more likely to have fallen as rain further south on tropical parts of the Great Dividing Range, within the catchments of vast inland floodplains and river systems. Australias weather systems are driven by global air and water currents, and the Southern Oscillation Index (SOI) has proved a useful indicator of long-term patterns (Allan et al. 1996; Scott 1997). Some relationships were found in this study between the SOI and waterbird numbers, as in other studies in south-eastern Australia (Norman and Nicholls 1991; Roshier et al. 2001a,b; Kingsford and Norman 2002). However, in this study it did not prove as good a predictor of waterbird numbers in Western Port as regional rainfall or streamflow data. This may be because the SOI is an imperfect and spatially coarse indicator of weather patterns that actually eventuate. Indeed, the data used in this study showed that the influence of the SOI on rainfall in southern Australia was generally less strong than it was for rainfall in other regions (Fig. 6), reflecting the seasonal predictability of rainfall in temperate southern Australia in comparison with more northern or arid regions. The best predictors for all three species examined were the streamflow data, which integrate effects of previous rainfall at the level of specific catchments, despite imperfections and anomalies arising from artificial regulation of many of the rivers. Wetland processes and waterbird numbers may relate more directly to streamflow data than rainfall data, because much rain feeds terrestrial and groundwater systems with little impact on wetlands used by waterbirds. Streamflow data have also been used in other studies (e.g. Briggs and Holmes 1988). Loyn et al. (1994) used River Murray flows as an integrating index of recent wetness in the Murray-Darling Basin. Briggs and Holmes (1988) developed a wetness index for various regions of Australia, and Roshier et al. (2001a,b; 2002) developed a process for estimating wetland areas from satellite imagery. These approaches clearly have great potential for further understanding waterbird dynamics at local and continental scales. The present study confirms expectations from previous work that waterbirds in Western Port are least common when there is plentiful water in breeding habitats for the respective species (locally or inland) and most common at various periods of time after wet periods in those habitats. The study quantified those lag periods as three to nine seasons (up to 4 years) for the three species considered in this time-span (19732002). Conservation planning is typically conducted on a local basis, recognising the values of local habitats and perhaps their immediate context in the local landscape. But waterfowl appear to respond to climatic events at a broad continental scale, and need habitats in widely different places and times for different purposes. These fundamental points are well recognised by waterfowl biologists (e.g. Frith 1982; Marchant and Higgins 1990; Briggs 1992; Kingsford et al. 1999b; Roshier et al. 2001a; Kingsford and Norman 2002) but perhaps less so by the general public or the planning

Fig. 6 Simultaneous correlations between seasonal rainfall and the Southern Oscillation Index (SOI) for a winter (JuneJulyAugust) and b spring (SeptemberOctoberNovember), using data from 1950 to 1999. The contour interval is 0.1 with shading starting at 0.3. Solid lines indicate positive correlations and dashed lines negative correlations

profession. The present study supplements the body of knowledge on these matters, and illustrates how numbers of birds in one area (Western Port) may be influenced by climatic events in disparate and distant parts of the continent. These responses are only possible if suitable habitat has been conserved in these disparate areas. Ephemeral swamps are particularly important as breeding habitat for Australian waterbirds for many reasons (Kingsford and Norman 2002). Such swamps provide rich supplies of invertebrate food for waterfowl, often based on decaying terrestrial vegetation as the primary source of organic matter (Briggs et al. 1985b; Crome and Carpenter 1988). Waterfowl are able to access a large share of the available food, with few fish to compete for food or prey on ducklings. Many ephemeral swamps support stands of trees such as River Red Gums Eucalyptus camaldulensis, providing nest-sites on branches and in hollows (Briggs and Thornton 1999; Loyn et al. 2002).

303

Ephemeral swamps of this sort can support large numbers of breeding waterfowl, whether they are situated in temperate parts of Australia or far inland (Roshier et al. 2001a). Conservation of ephemeral swamps must involve appropriate protection or management of vegetation and water regimes (Kingsford and Norman 2002). Maintaining water regimes is a special challenge in view of their enormous natural variation in the arid zone, the complicating effects of climate change and the competing demands for water in settled parts of Australia. This challenge will intensify as human populations increase.
Acknowledgements The data on waterbird numbers could not have been collected without the dedicated efforts of numerous voluntary observers over 29 years. Many thanks to them all, and to the Bird Observers Club of Australia for supporting the survey. Streamflow data were kindly provided by a number of state authorities: Department of Infrastructure, Planning and Environment (Northern Territory); Department of Primary Industries, Water and Environment (Tasmania); Department of Natural Resources and Mines (Queensland); Department of Water, Land and Biodiversity Conservation (South Australia); Rubicon Systems Australia Pty Ltd. (Victoria); Sydney Water (New South Wales); Water and Rivers Commission (Western Australia). Mike Matheson, of the Bureau of Meteorology, provided assistance in obtaining information on stream flows, Wasyl Drosdowsky for the provision of the SOI-rainfall plots, Drte Jakob for assistance in producing the drainage division plot, and the National Climate Centre provided the SOI and rainfall data. The authors wish to acknowledge the insightful comments made by Dr Ian Norman of the Arthur Rylah Institute for Environmental Research, and Drs Scott Power and Bertrand Timbal, both of the Bureau of Meteorology, on an earlier draft of this paper. The paper was further improved by comments from two anonymous reviewers.

References
Allan R, Lindesay J, Parker D (1996) El Nino Southern Oscillation and Climate Variability. CSIRO Publishing, Collingwood, Australia Barrett G, Silcocks A, Barry S, Cunningham R, Poulter R (2003) The New Atlas of Australian Birds. Royal Australian Ornithologists Union, Hawthorn East Briggs SV (1977) Variation in waterbird numbers at four swamps on the northern tablelands of NSW. Aust Wildl Res 4:301309 Briggs SV (1992) Movement patterns and breeding characteristics of arid zone ducks. Corella 16:1522 Briggs SV, Holmes JE (1988) Bag sizes of waterfowl in New South Wales and their relation to antecedent rainfall. Aust Wildl Res 15:459468 Briggs SV, Thornton SA (1999) Management of water regimes in River Red Gum Eucalyptus camaldulensis wetlands for waterbird breeding. Aust Zool 31:187197 Briggs SV, Maher MT, Davey CC (1985a) Hunter activity and waterfowl harvests in New South Wales, 19771982. Aust Wildl Res 12:515522 Briggs SV, Maher MT, Tongway DS (1985b) Dry matter and nutrient loss from decomposing Vallisneria spiralis L. Aquat Bot 22:387392 Chambers LE, Hughes L, Weston MA (2005) Climate change and its impact on Australias avifauna. Emu 105:120 Cramp S, Simmons KEL (eds) (1977) The Birds of the Western Palearctic, Vol. 1. Oxford University Press, Oxford Crome FHJ, Carpenter SM (1988) Plankton community cycling and recovery after drought - dynamics in a basin on a flood plain. Hydrobiologia 164:193211

Dann P, Loyn RH, Bingham P (1994) Ten years of waterbird counts in Western Port, Victoria, 197383. II. Waders, Gulls and Terns. Aust Bird Watch 15:351365 Del Hoyo J, Elliott A, Sargatal J (eds) (1992) Handbook of the Birds of the World, Vol. 1. Lynx Editions, Barcelona Dexter BD, Rose J, Davies N (1986) River regulation and associated forest management problems in the River Murray red gum forests. Aust For 49:1627 Frith HJ (1982) Waterfowl in Australia. Angus & Robertson, Sydney Geering DJ, Maddock M, Cam G, Ireland C, Halse SA, Pearson GB (1998) Movement patterns of Great, Intermediate and Little Egrets from Australian breeding colonies. Corella 22:3746 Gentilli J, Bekle H (1983) Modelling a climatically pulsating population: grey teal in south-western Australia. J Biogeogr 10:7596 Gosper DG, Briggs SV, Carpenter SM (1983) Waterbird dynamics in the Richmond Valley, New South Wales, 197477. Aust Wildl Res 10:319327 Halse SA, Jaensch RP (1989) Breeding seasons of waterbirds in south-western Australia - the importance of rainfall. Emu 89:232249 Heislers A (2003) Wings over Western Port. Bird Observers Club of Australia, Nunawading Jones DA, Weymouth G (1997) An Australian monthly rainfall dataset. Technical Report 70. Bureau of Meteorology, Melbourne, Australia, pp 19 Kingsford RT, Norman FI (2002) Australian waterbirds - products of the continents ecology. Emu 102:4769 Kingsford RT, Curtin AL, Porter J (1999a) Water flows on Cooper Creek in arid Australia determine boom and bust periods for waterbirds. Biol Conserv 88:231248 Kingsford RT, Wong PS, Braithwaite LW, Maher MT (1999b) Waterbird abundance in eastern Australia, 198392. Wildl Res 26:351366 Loyn RH (1978) A survey of birds in Westernport Bay, Victoria, 197374. Emu 78:1119 Loyn RH (1991) Assessing and managing the impact of duck hunting in Victoria - a new approach. Wildfowl 42:155161 Loyn RH, Dann P, Bingham P (1994) Ten years of waterbird counts in Western Port, Victoria, 197383. I Waterfowl and large wading birds. Aust Bird Watch 15:333350 Loyn RH, Dann P, McCulloch E (2001) Important wader sites in the east Asian-Australasian flyway: 1. Western Port, Victoria, Australia. The Stilt 38:3953 Loyn RH, Lumsden LF, Ward KA (2002) Vertebrate fauna of Barmah Forest, a large forest of River Red Gum Eucalyptus camaldulensis on the floodplain of the Murray River. Vic Nat 119:114132 Marchant S, Higgins PJ (eds) (1990) Handbook of Australian, New Zealand and Antarctic Birds, Volume 1, Part B. Oxford University Press, Melbourne Nicholls N (2001) Atmospheric and climatic hazards: improved monitoring and prediction for disaster mitigation. Nat Hazards 23:137155 Norman FI (1971) Movement and mortality of Black Duck, Mountain Duck and Grey Teal banded in South Australia, 19531963. Trans Royal Soc S Aust 95:17 Norman FI, Powell DGM (1981) Rates of recovery of bands, harvest patterns, and estimates for Black Duck, Chestnut Teal, Grey Teal and Mountain Duck shot during Victorian open seasons, 195377. Aust Wildl Res 8:659664 Norman FI, Nicholls N (1991) The Southern Oscillation and variations in waterfowl abundance in southeastern Australia. Aust J Ecol 16:485490 Pearce RP (2002) Meteorology at the Millennium. International Geophysics Series, Volume 83, Academic Press, London Pittock B (2003) Climate Change: an Australian Guide to the Science and Potential Impacts. Australian Greenhouse Office, Canberra

304 Pringle JD (1985) The Waterbirds of Australia. Angus and Robertson and the National Photographic Index of Australian Wildlife, Sydney Roshier DA, Robertson AI, Kingsford RT, Green DG (2001a) Continental-scale interactions with temporal resources may explain the paradox of large populations of desert waterbirds in Australia. Landsc Ecol 16:547556 Roshier DA, Whetton PH, Allan RJ, Robertson AI (2001b) Distribution and persistence of temporary wetland habitats in arid Australia in relation to climate. Austral Ecol 26:371384 Roshier DA, Robertson AI, Kingsford RT (2002) Responses of waterbirds to flooding in an arid region of Australia, and implications for conservation. Biol Conserv 106:399411 Scott A (1997) Relationships between waterbird ecology and river flows in the Murray-Darling Basin. CSIRO Land and Water Technical Report No. 5/97, pp 41 Stevermer AJ (2001) Recent Advances and Issues in Meteorology. Oryx Press, Westport, Conn. Troup AJ (1967) Opposition of anomalies in upper tropospheric winds at Singapore and Canton. Aust Meteorol Mag 15:3237 Wright WJ (2004) Drought, dust and deluge: a century of climatic extremes in Australia. Bureau of Meteorology, Melbourne