5HSRUW1R 5HYLHZDQGDQDO\VHVRI,0()ELUG VXUYH\VIRU/DFKODQ5LYHUYDOOH\ 'DYLG5RVKLHU DQG'DYLG0:DWVRQ Review and analyses of IMEF bird surveys for Lachlan River valley David Roshier and David M Watson Institute for Land, Water and Society Charles Sturt University PO Box 789, Albury NSW 2840 Review and analyses of IMEF bird surveys for Lachlan River valley Summary The data consist of a mixture of counts of varying precision. Moreover, there are notable absences, unlikely occurrences and species that are underrepresented. In addition single counts of one species at particular wetlands account for the majority of individual birds recorded. For these reasons quantitative analyses and exploratory modelling were no pursued as model outputs would be misleading. The survey approach as written may be appropriate but only if implemented in a consistent manner to ensure standardised data. Adding call-playback survey techiques, considering detectability and expanding the survey area will ensure data are more consistent, more representative of underlying patterns and more comparable. 1. Background Water allocation is made on a whole-of-catchment basis by management agencies yet the responses of individuals and populations of mobile species, such as birds and fish, to changes in water regime are poorly known at the catchment scale. This lack of knowledge is due, in part, to our inablity to reconcile local changes in abundance with changes in the broader population and a limited understanding of the ecology of many species. Nonetheless, targeted studies of particular species or functional groups offer some insight into the responses of biodiversity to changes in water regime. For instance, colonial-nesting waterbirds, such as ibis and spoonbills, are iconic species whose population dynamics have been successfully used as a measure of the impacts of water management (Kingsford & Johnson 1998; Leslie 2001, Driver 2004) and engaged the community’s interest. Analyses of the response of other wetland bird species to changes in river flow and wetland inundation point to significant impacts due to river regulation (Kingsford et al. 2004, Briggs et al. 1997, Stapanian et al. 2004). Thus, there remains an imperative to develop surveys and methods to model the responses of birds and others at the same catchment scale at which agencies manage the resource. The Department of Infrastructure, Planning and Natural Resources (DIPNR) has developed the Integrated Monitoring of Environmental Flows (IMEF) program for the Lachlan River valley and other catchments. As stated in DIPNR (2003), the objectives of IMEF are: 1. to investigate relationships between water regimes, biodiversity and ecosystem processes in the major regulated river systems of NSW (and the Barwon-Darling River); 2. to assess responses in hydrology, habitats, biota and ecological processes associated with specific flow events targeted by environmental flow rules; and 3. to use the resulting knowledge to estimate likely long-term effects of environmental flow rules and provide information to assist in future adjustment of rules. Birds are one of several taxa monitored at 12 sites in the Mid and Lower Lachlan. The monitoring is event-based and, since the commencement of monitoring in 2000, there have been 20 inundation events of individual wetlands—mostly in the first year. This monitoring program has produced data on 143 wetland and woodland species from 111 individual surveys. We (Drs Roshier and Watson) undertook to conduct preliminary analyses of the data, and model responses of wetland and woodland birds to changes in river flow and wetland inundation. The agreed analyses, outcomes and priorities (M-medium, H-high, VH-very high) follow: Preliminary analyses. 1. Quantify changes in abundance and diversity of wetland and non-wetland bird species over the survey period and in response to wetland inundation (VH for wetland species, M for non-wetland species) 2. Examine species composition and diversity in response to flow and water parameters (H) 3. Classify wetland species into functional groups and examine patterns within and between groups (H) 4. Evaluate existing data for potential indicator species to enhance our understanding of waterbird responses to environmental flows (VH) Review of current bird survey techniques 5. Comparative evaluation of measures of waterbird responses: community based versus functional groups versus indicator species (H) 6. Summarise advantages and disadvantages of alternative techniques for monitoring waterbirds (M) 7. Assessment of suitability of response variables to desired management options (ensuring the method informs management) (VH) Outcomes 8. review of survey methods for waterbirds (M) 9. discussion of limitations and strengths in the interpretation of analyses using current and alternative survey methods (H) 10. recommendations for alternative methods to enhance current surveys (M) 11. conceptual (non-mathematical) summary of patterns of abundance and species richness in response to environmental flows and other factors (VH) 12. mathematical and/or statistical summaries of patterns of abundance and species richness in response to environmental flows and other factors that can be linked to current wetland flow models (VH) 13. evaluation of trends in abundance of functional groups of waterbirds (H) 14. identification of indicator species for targeted monitoring of waterbird responses in the Lachlan River valley (H) 15. exploration of data requirements and modelling options based on specified management scenarios (H) 2. Data quality The survey data as originally supplied on 24/06/05 were incomplete with 36 of 109 survey dates with no corresponding bird data. These data included separate entries with the same species name (plumed whistling-duck), species that were entered with two different common names (superb fairy wren, sulphur-crested cockatoo), entries with no scientific name and/or an indecipherable common name (black-eared cuckoo). The data were corrected and returned on 8/11/05. The final data set consisted of 1490 records of 143 species. The median and maximum number of species seen on site surveys was 13 and 29, respectively. Both these figures are lower than that expected of a comprehensive survey of wetland or riparian habitats in this region and suggest there are issues with the either the time allocated to complete the survey, the size of the area surveyed or species identification. Among the waterbirds and those that occupy the margins of aquatic habitats there a number of notable absences (table 1). Also, based on the knowledge of the authors there are a number of species that have been recorded at a lower frequency than would be expected (table 2). Table 1. Notable absences of waterbirds and species associated with aquatic habitats from surveys of wetlands in the Lachlan River valley conducted from 1999 to 2003. Common name Scientific name Status Blue-billed duck uncommon (Pizzey & Knight 2004) Oxyura australis Freckled duck uncommon (authors) or rare (PK 2004) Stictonetta naevosa Hardhead common (authors) Aythya australis Cattle egret uncommon inland (authors) Ardea ibis Brolga uncommon (Pizzey & Knight 2004) Grus rubicunda Buff-banded rail Gallirallus philippensis common (authors) Aust. spotted crake common (Pizzey & Knight 2004) Porzana fluminea Latham’s snipe uncommon (Pizzey & Knight 2004) Gallinago hardwickii Common greenshank uncommon inland (Pizzey & Knight 2004) Tringa nebularia Marsh sandpiper uncommon inland (Pizzey & Knight 2004) Tringa stagnatilis Wood sandpiper uncommon (Pizzey & Knight 2004) Tringa glareola Sharp-tailed sandpiper common (Pizzey & Knight 2004) Calidris acuminata Pectoral sandpiper uncommon (Pizzey & Knight 2004) Calidris melanotos Curlew sandpiper uncommon inland (authors) Calidris ferruginea Red-necked avocet common (Pizzey & Knight 2004) Recurvirostra novaehollandiae Red-capped dotterel Charadrius ruficapillus common (authors) Red-kneed dotterel uncommon (authors) Erythrogonys cinctus Little bittern uncommon (Pizzey & Knight 2004) Ixobrychus minutus Caspian tern uncommon (authors) Sterna caspia White-winged black tern Chlidonias leucopterus uncommon (authors) Little grassbird common (Pizzey & Knight 2004) Megalurus gramineus Golden-headed cisticola Cisticola exilis common (Pizzey & Knight 2004) Table 2. Waterbirds and species associated with aquatic habitats recorded at frequencies less than expected in 111 individual surveys of wetlands in the Lachlan River valley conducted from 1999 to 2003. Common name Scientific name Total Status Hoary-headed grebe Poliocephalis poliocephalus 4 common (Pizzey & Knight 2004) Pink-eared duck 6 common (authors) Malacorhynchus membranaceus Musk duck 8 uncommon (authors) Biziura lobata Wood duck 178 abundant (authors) Chenonetta jubata Little egret 1 uncommon (Pizzey & Knight 2004) Egretta garzetta Baillon’s crake 6 uncommon (authors) Porzana pusilla Spotless crake 1 uncommon (Pizzey & Knight 2004) Porzana tabuensis Dusky moorhen 4 common (Pizzey & Knight 2004) Gallinula tenebrosa Purple swamphen 26 common (authors) Porphyrio porphyrio Improbable records The Great-billed heron Ardea sumatrana recorded at Lignum Swamp on 11/01/00 is an improbable record. This species is confined to coastal regions of the tropical north of the continent, and has never been recorded in inland NSW. Given the time of year and location, this record likely relates to a juvenile White-necked heron Ardea pacifica, which is similar in overall size and plumage (although it has a markedly smaller bill). We recommend that this record be deleted unless the observer has corroborating evidence. Other data issues The figures reported for species abundance are a mixture of counts of individuals and group estimates. While this is often a feature of survey data it is not articulated in the sampling protocol at what level of abundance this change in counting method should occur. Hence, some surveys simply report abundance as estimates of abundance to the nearest 10 while others appear to be counts of individuals as high as 54. The data are few and highly skewed with many zeros. The majority of estimated abundances are less than 10 with a few records in the hundreds or thousands. Of the 13,546 individual birds recorded, most records relate to a small number of species on a single occasion at several sites. Among the 8,456 wetland birds recorded, 3,500 are from two counts of ibis. Thus, in looking for responses of wetland birds to changes in flow regime, it is likely that any significant relationship will be due to the high degree of leverage of one or two points in the analysis. Indeed, the data are largely the product of one flooding event in 2000. Earlier flooding events for which data are available on other taxa do not include bird surveys at all stages of the flood cycle. It is possible to analyse data with many zeros using both parametric and nonparametric methods, but we do not think this appropriate at this stage. Alternative approaches include basing the analysis on presence/absence or to combine data into functional groups (see later). However, we believe it is premature to pursue such analyses. Based on the above concerns about data quality, we are unable to model the responses of wetland birds to changes in river flow. At this stage, the modelled outcomes from these data are likely to be unreliable. As a result it is not possible to complete all components of tasks 1, 2, 3, 4, 11, 12, 13 and 15. While issues of detectability and completeness are recognised in the written protocol for bird surveys, IMEF Method 29 – Wetland waterbird survey (Driver et al. 2003), there is no explicit method in the protocol for correcting for biases and errors. In the next section we review current approaches to bird survey and comment on the protocol. 3. Synthesis of current approaches to bird survey The basis of any analysis of biodiversity occurrence is the raw data—which species were present at which locations at which time periods. Whether historic data, opportunistically collected records or a systematically designed sampling protocol, different kinds of data allow statements of varying precision, underpinning our understanding of determinants of diversity and informing management and meaningful conservation. Matching the scope of ecological inference and management decisions to the resolution of the original data is the ultimate aim of effective biodiversity sampling (Verner 1985, Mac Nally and Horrocks 2002, Watson 2003, 2004). This preliminary review focuses on sampling waterbirds, a diverse and prominent group associated with a range of habitats worldwide. This review covers three main areas. Firstly, we highlight a suite of confounding factors that affect the accuracy of waterbird surveys, focusing on ways to minimise or avoid them. Secondly, we review the main methods used to sample waterbirds, both in Australia and internationally—highlighting the inherent advantages and disadvantages of each. Finally, we discuss some alternative approaches to sampling, demonstrating how a sampling protocol can be designed to minimise the influence of confounding effects while maximising the usefulness of the resultant data for informing management. 3.1 Confounding factors There is a large (and growing) literature on sampling birds, but most focuses exclusively on terrestrial groups. While many recent advances have a direct bearing on waterbirds, others have little relevance. Key reviews or discussions on these topics are listed in the references section, with Verner (1985), Watson (2003, 2004), USFWS (1999), Thompson (2002), Krzys et al (2002) and Conway and Gibbs (2005) being most informative. Time of day, observer behaviour and experience, weather and habitat complexity all have individual confounding influences, as well as interacting to diminish accuracy of bird surveys (Recher 1998, Bibby et al. 1992, Craig and Roberts 2001). Rather than discuss each of these individually, the effect of many of these factors are discussed in terms of detectability. Observer effects are discussed separately. Detectability There has been an increasing amount of research in the past five years on the effects of detectability on the accuracy of bird surveys (Thompson 2002). Specifically, it is clear that different species have different probabilities of being recorded under otherwise similar conditions, and that the same species may differ in its likelihood of detection at different times of the year, or in different habitats. Little work has been conducted on detectability issues in waterbirds, and to-date, no research has examined this topic in Australia. So, although no actual data can be brought to bear on the importance of detectability differences to sampling Australian waterbirds, there are several important issues that can be identified. Habitat-based differences in detectability are the most common and can have marked effects on datasets (Rosenstock et al. 2002). For example, Dusky moorhens may be twice as likely to be seen and recorded in wetlands with no fringing vegetation, compared with wetlands surrounded by reed beds. If detectability is not taken into account, most survey methods would yield the result that moorhens avoid wetlands with fringing vegetation. Typically, however, habitat-based differences in detectability are less obvious, and so can be complex to identify, let alone correct for. Thus, some calls are less audible in more open vegetation, some species are more wary in open areas, and some species call less frequently away from cover—all factors that would affect estimated densities or abundances, and undermine any crosssite comparisons. Time-based differences in detectability relate to seasonal and resource-based changes, whereby birds at one time of the year are more easily recorded than at others. While typically measured as breeding/non-breeding in Northern Hemisphere studies, this distinction is less clear-cut in Australian assemblages, where the prevailing climate is irregular, and so seasonality and the timing of breeding vary greatly from one year to the next. This stochasticity, coupled with the lack of obvious breeding plumage for many Australian species makes it difficult to gauge breeding status, and therefore evaluate any effect it may have on detectability. While there are numerous schemes for estimating detectability of birds (e.g., measuring call frequency contemporaneously with the bird survey; Rosenstock et al. 2002), these issues can be largely avoided in sampling design. In regards to estimating abundance, restricting sampling to presence/absence (or nominal estimates like <10, >100) diminishes the influence of detectability differences between sites and between time periods. As long as the sampling intensity is sufficient to detect a species as present in the least detectable site at the least detectable time of year, these issues will have little effect on the overall data-set. Inter-specific detectability differences are more pervasive. While many waterbirds are highly detectable—large bodied, travelling in groups, foraging in the open and easily identified—other species are more secretive. Rails and bitterns are two prominent groups that present significant difficulties for sampling—they are rarely seen in the open, and when they are seen, have cryptic plumage and may be difficult to positively identify to species. Painted Snipe are similarly wary, and it can be difficult to distinguish true absences from a lack of presence data. Implications for Lachlan Catchment data In exploring the current data set, several species were notable for their absence, and while no individuals of these species may have been present at any of the wetlands during the study period, they may also have been present but not detected. Little Grassbirds, Australasian Bittern and Buff-banded Rails are all reasonably common in wetlands throughout this region, but were not recorded once. Rather than being a few additional species that have little effect on the overall pattern, these species are wetland specialists with strict habitat tolerances. As such they might be expected to be highly sensitive to subtle differences in wetland structure and water regime, and so should be targeted specifically (see playback surveys, below). In sum, if comparisons between different seasons and different sites are to be made, these can currently be made using presence/absence data with a low likelihood of detectability differences affecting the comparison. If changes in relative abundance are a priority, and need to be estimated, some measure of detectability should be incorporated into the sampling design. Given that no procedure for Australian waterbirds exists, a scheme like this would have to be carefully developed, using variables such as call frequency, or time until first record to calibrate the data. Finally, inter-specific differences in detectability will have confounding effects on any inter-specific comparisons based on estimates of relative abundance. Even though these can be minimised by restricting analysis to presence/absence, the current sampling approach may not be recording all species, with several groups probably requiring targeted sampling to confirm their presence. Observer effects While a range of observer-related effects are alluded to in the IMEF Method 29 – Wetland waterbird survey (Driver et al. 2003), it is unclear how these issues were guarded against. As discussed earlier, there are several dubious records and notable absences—suggestive of inadequate observer training and/or inconsistent interpretation of the prescribed sampling approach. Time of day and weather effects are critical for many species, affecting both detectabilities and identification (Shields 1979, Craig and Roberts 2001). While problematic with individual observers, these issues are further compounded when there are multiple observers contributing data. Specifically, interobserver differences add another level of confounding effects—artefacts that can outweigh any underlying differences between locations of time periods (Dawson 1981, Remsen 1994, Recher 1998). The only way to diminish inter-observer differences is training, followed up by regular validation. Initially, observers must have a minimum level of proficiency, in terms of identifying species using plumage, behaviour and vocalizations; as well as estimating numbers. They must be trained to ensure they are familiar with the survey procedures and how they should be applied. Subsequent comparisons between observers are then needed to ensure comparable levels of accuracy (USFWS 1999). Implications for Lachlan Catchment data collection Once the survey methods have been refined and established, all observers need to be evaluated for competency, and trained to ensure consistency. In addition to applying the established sampling protocol, more detailed guidelines need to be developed to ensure the protocol is applied uniformly. Hence, colour of clothing and observer behaviour (i.e., talking, walking speed, approach distance etc) should all be standardized. Similarly, time of day (relative to sunrise and sunset) and weather should be standardised, such that all samples are collected at similar times of the day under comparable weather conditions. 3.2 Approaches to waterbird sampling A wide range of techniques are used for sampling waterbirds and analysing patterns of their occurrence (USFWS 1999, Conway and Gibbs 2005). These techniques may relate to particular surveying techniques or specific variables used for analysis, and will be discussed separately. Visual versus auditory records—Visual cues (including plumage, overall size and proportions as well as behaviour) are typically the main means of identifying birds. With appropriate experience, this information is an accurate and reliable basis for identifying birds to species and estimating abundances (Bibby et al. 1992, Stapanian et al. 2002). While appropriate for many waterbirds (and many wetlands), visual cues alone may be inappropriate in some situations. Heavily vegetated wetlands will contain many species partially or completely obscured from view, either from landward or water-based perspectives. In contrast, auditory cues (both vocalizations and splashing/rustling associated with bird activity) are less affected by intervening vegetation. While many calls can be readily ascribed to particular species, others (especially alarm calls) are not reliable for species-level identification. Moreover, abundances can rarely be estimated from bird calls. Hence, rather than advocating one source of data over the other, we strongly recommend using both visual and auditory cues in concert. Call playback—Even when bird calls are used, some species will consistently be missed, and others will be disproportionately overlooked at certain times of the year (many waterbirds are quiet after breeding). To redress this bias and minimise the number of false absences, playing pre-recorded bird calls is a useful technique. A recent review paper (Conway and Gibbs 2005) demonstrated that the addition of playback surveys significantly increased the likelihood of detecting particular species of rail and bittern in the USA and Canada. Despite being standard for waterbird surveys in North America, call playback is not part of the standard complement of techniques used in Australia. The reasons for this are unclear—call playback is routinely used for nocturnal bird surveys throughout the country, and high quality recordings are readily available for all species of interest. Density and abundance—The advantage of limited area techniques is that abundance estimates can be expressed in terms of density, which can then be extrapolated to the area of interest. Verner (1985) discussed this topic in detail and identified a broad range of artefacts, inherent biases and confounding factors associated with density estimates (see also Dawson 1981). For waterbird surveys, two of the most serious of these shortcomings relate to accuracy of counts (i.e., controlling for individuals present but not detected), and the issue of representation (i.e. the degree to which a detailed count within a subsample can be scaled up to reflect the overall site). While density estimates can be a useful and sensitive complement to richness estimates, Verner concluded that most sampling approaches are simply unable to generate estimates that reflect underlying bird numbers. While this means absolute measures are rarely possible, estimates of relative abundance (or scaled to generate relative density estimates) may still be useful and reflect underlying differences (Dawson 1981). For relative measures, it is assumed that the various confounding factors and biases are constant—occasionally possible, but rarely measurable. By controlling for detectability (as mentioned previously), relative abundance measures can become more robust and reliable. Incidence—An alternative to abundance or density estimates (either in absolute or relative terms) is incidence, or reporting rate (Wright 1991, Bibby et al. 1992, Watson 2003). This measure expresses the proportion of samples in which a species was detected. As it is essentially a composite of multiple presence/absence samples, it is not as finely resolved as direct estimates of abundance. While this means it will be less sensitive to change, this also means that incidence is far more robust to changes in detectability. Hence, in a given sample, a flock of 20 birds or a single individual will both be denoted as “present”, and the records treated as equivalent. After ten samples, the species may have been recorded as present in 8 of them, yielding an incidence of 0.8. Rather than relating exclusively to abundance, incidence incorporates site tenacity, territoriality, vagility and other autecological traits. As such, comparing three species (A, B and C) with incidences of 0.8, 0.5 and 0.1 (respectively), the site could be considered most important for species A, of intermediate value to species B, and of little importance for species C. Obviously, incidence can be affected by differing ecologies and detectabilities, so comparisons are best restricted to ecological similar taxa. Total Richness—Most bird surveys collect data on all species encountered within the sampling period, typically summarising these data as species richness (Herzog et al. 2002). While useful to quantify the range of species recorded, overall richness estimates are typically too broad to be of analytical value. Instead, exotic species, non target species and incidental records are typically removed, giving a woodland bird or wetland bird richness estimates. This list of species may be further subdivided into habitat specialists versus generalists, species that breed in the site, migrants or seasonally restricted species. Of these, breeding species richness is one of the more sensitive indices of site quality, and is frequently used in relation to terrestrial avifaunae. Breeding records—Those species breeding in a particular site are necessarily more sensitive to within and between site variation, and can be a useful assay for site-based differences. While some waterbird species are easily designated as breeding versus non-breeding (e.g., colonially nesting egrets or pelicans), others are far more secretive. Some species acquire distinctive nuptial plumage during breeding seasons, enabling trained observes to distinguish not just breeding species from non breeders, but also proportion of breeding vs non breeding individuals with a particular species. Similarly, some species (bitterns, kingfishers, grebes) give distinctive vocalizations during territory acquisition, courtship and incubation, although most species become characteristically silent during chick rearing and fledging. Hence, distinguishing breeding from non-breeding species within a particular site yields valuable data but relies on detailed (and region-specific) knowledge and experience. Functional groups—Another way to subdivide total richness estimates is to use the overall ecology of the species to assign them to functional groups. Unlike total richness, richness within a carefully defined functional group is far more likely to elucidate subtle change or between-site differences. Functional groups should be biologically real (i.e., corresponding to underlying biological similarities), and not just defined relative to obvious differences like diet, but breeding strategies/nesting, movement/migratory approaches and the like. Moreover, these groups should correspond to groups that are sensitive to the key factors of interested. So, “cavity nesting waterfowl” corresponds to a group of ecologically similar taxa, but may have little explanatory value. Conversely, “reed-nesting residents” may include things as varied as Little Grassbirds, Australasian Bittern and Purple Swamphen, but richness of this group may be a highly sensitive indicator of flows, grazing and salinity. One approach to classify birds in the Lachlan catchment is summarised in appendix 1. Indicator species—An alternative to richness-based measures is to select a particular species that reflects overall change (Brooker 2002). There has been considerable interest (both internationally and within Australia) to identify these indicator species. While a well chosen indicator species can be an efficient and repeatable means of measuring response to treatments, or variation across space and time, choosing the right species is critical. The species must be highly detectable—i.e., need to maximise likelihood that a lack of records corresponds to an actual absence. Identification needs to be straightforward, both to minimize chances of misidentification and to allow ease of sampling by a range of observers. In order to be an accurate indicator, variation in the chosen species should reflect overall change in the group of interest. Finally, and most importantly, the species should respond to factors of interest. Selecting an effective indicator species is generally a three step process. Firstly, collect a large, broad-based data-set incorporating all the species of interest across the breadth of sites/treatments available. Secondly, identify a species that reflects overall change in the data set (and is appropriately detectable and distinctive). Thirdly, evaluate change in a complementary data set to determine whether changes in the indicator species reflect broad-based change. 4. Appraisal of current sampling approach The current approach to sampling waterbirds involves four surveys of 12 wetlands each year: pre-flow counts in March/April followed by counts 14, 60 and 90 days after wetland inundation. On any reach, survey area is a 100m section for full width of billabong, including any birds flying in vicinity within 100m. The current protocol is based on a fixed effort and community scale abundance estimates. The main benefit of this approach is its breadth—it gives a broad overview of occurrence patterns of a large number of wetland dependent species across the catchment. Accordingly, it yields useful baseline data on which species occur where, times at which particular species are abundant within the catchment, and highlights particular localities that support high diversities. Other than these broad statements, however, many of the confounding factors discussed earlier prevent more detailed statements, and preclude explicit comparisons across space and time. The sampling effort used is constant, which assumes that the fixed effort is equally representative of large, complex sites supporting high diversities and small, simplistic sites supporting relatively low diversities. This high inter-site variability coupled with known behaviours of waterbirds confounds between-site comparisons. Species rich sites may be relatively under-sampled, while species-poor sites may be relatively over-sampled, diminishing actual differences between the sites. As discussed briefly earlier, there are several notable absences from the overall dataset, and several notable inclusions. This raises questions about the experience of the observers collecting the data, their training, their familiarity with waterbird species in the region, and their ability to identify species by visual, auditory and/or behavioural cues. This is especially the case with rails, bitterns and other cryptic groups—rarely picked up during routine surveys. Even with experienced observers, however, these species are often overlooked, and call playback is now routinely used in North America when conducting wetland surveys. Aside from issues of comparability and representation, the fixed sampling effort represents a small to very small sub-sample of the overall wetland (USFWS 1999). Hence, for a typical billabong 50 m wide, the survey area for wetland birds is a ~0.5 ha section of billabong 1-5 km in length. Regardless of where the sample is located, the resultant survey cannot be considered representative of the overall wetland. Just as with transect counts in terrestrial systems, the resultant data maybe an accurate indication of which species occurred within the sampling area, but these data cannot be extrapolated to the wetland scale, diminishing the utility and relevance of the data for management decisions (decisions that operate at the wetland and whole-of-catchment scale). A strong argument for continuing with the existing sampling protocol is to make the most of work already in hand, making temporal comparisons across the catchment. The alternative is to evaluate how well the current decisions are being informed and supported by the resultant data. So, if long-term temporal comparisons are the priority (i.e., how have bird distribution patterns changed over time), continuing with the existing sampling approach is reasonable. Alternatively, if meaningful and accurate statements about the timing, magnitude and value of differing flow regimes are to be formulated, at the scale of individual wetlands, subregions or the overall catchment, it may be appropriate to consider changing the sampling regime. 5. Suggested alternative sampling approaches Patch-scale sampling In terms of overall sampling design, patch-scale sampling is a priority. In addition to avoiding problems of unequal representation among samples, this approach also yields data at the wetland scale. Using fixed-area approaches, explicit comparisons can be made between samples (Merikallio 1958), yet statements about the waterbird diversity of the overall wetland cannot be made (Watson 2003, 2004). Hence, using the overall wetland as the scale of sampling allows data of comparable representation to be collected, and can be used to make statements at the wetland scale. If sampling is conducted at the wetland scale, careful consideration is needed to determine when samples are complete. This can be done a posteriori with the data in hand using rarefaction, but this assumes data have been collected in a cumulative way, and necessarily involves removing data points in order to bring all inventories to the same levels of completeness. For subsequent data collection, however, application of a well-selected results-based stopping rule is recommended (Peterson and Slade 1998). Rather than using an arbitrary effort-based stopping rule, based on either time or area sampled, results-based stopping rules use the data to determine when sampling is complete. As discussed in detail in Watson (2002, 2003), stopping rules should be designed with data comparability, logistical efficiency and practical applicability in mind. Hence, stop sampling once all species are seen in at least two sampling periods would yield data of high completeness that are comparable across sites and times, yet would be prohibitively time-intensive to collect. we would advocate tailoring a stopping rule that generates data of at least 75% completeness— any less, and representation becomes an issue. Note, that this need not involve exhaustive surveying efforts—three-to-five back to back half hour surveys would readily satisfy many stopping rules for the vast majority of wetlands. Sampling methods Once scale of sampling, stopping rules and sampling frequency have been selected, survey methods require consideration. Passive observation, even complementing visual based observations with auditory cues, will consistently miss some species, and under-represent several key groups (Conway and Gibbs 2005). To address this shortcoming, we would recommend complementing observation-based sampling with the use of call play-back. Rather than for all species, this would be for a predetermined subset of species: species considered possible in the region, and unlikely to be detected from observation alone. Playback location, timing, intensity and response times would all need to be determined, balancing time required with data yielded. As with survey timing and frequency, these decisions are best made on the basis of a small scale, representative pilot study, rather than arbitrarily, or with reference to research conducted elsewhere. Rather than being directly comparable to records of species gathered passively, these proactive detections could only be used to confirm presence or support putative absences—they would not yield incidence or abundance estimates (even in relative terms). Presence, absence and incidence Ideally, survey data should give a range of responses, not simply present or absent; yes or no. One way to achieve this, without significant increases in survey effort, is to record presence/absence data in sequential sampling periods (as advocated previously, and discussed in depth in Watson 2003). This allows incidence estimates to be calculated: the proportion of sampled in which a particular species was recorded. Although lacking the sensitivity of true abundance estimates, incidence is far more robust to changes in detectability, and so is far more likely to yield reliable estimates. While related to abundance, incidence is also affected by site tenacity, territoriality, social structure and home range size of particular species. Hence, it yields a broad index of how important a particular site is for a given species or suite of species. Duck species a, b and c may have been recorded with incidences of 0.1, 0.4 and 0.9, respectively. Regardless of differences in actual or estimated number, this translates to this particular site being most important for species c, (being recorded 9 out of 10 times), and of least importance for species a. This provides a more fine-grained set of responses, allowing the resultant datsa-set to be more sensitive to detect subtle differences between treatments/sites/times. Beyond presence/absence While compiling multiple presence/absence records can yield greater resolution than a simple binary variable, further details may be needed. What are the effects of altered flow regimes on the breeding response of a particular species or group of species? Incidence data would not address this question—additional data are required. For many species of waterbird, external characteristics (typically, plumage pattern, occasionally size and behaviour) can be used to distinguish adults from immatures, females from males, and breeding adults from those not in breeding condition. While not practicable (or even possible) for all species, collecting these supplemental data for selected species would augment the occurrence data, and give far more finely resolved information to inform management. Hence, for a particular species, a range of possible responses are possible. Present—yes or no? If present, breeding—yes or no? If present and breeding, was breeding successful—yes or no. Additional data would yield even greater resolution: number of breeding attempts, number of chicks fledged etc. One potential sampling approach As a worked example, I’ve selected spoonbills. There is more than one species (in this case, two); easily identified to species with minimal training; adults can be readily distinguished from immatures/juveniles; breeding adults can be readily distinguished from non-breeding adults; and nests are relatively easy to locate. These procedural differences reflect underlying ecological differences between the two species—they utilise habitats differently (for foraging, roosting and nesting), they feed on different prey (large invertebrates, compared with small fish). Taking this ecological information into account, a sampling protocol is designed. Standardised searches will be conducted before and after altered flow regimes, using the overall wetland as the sampling scale, and conducting targeted observations coupled with call playback for rails and bitterns at least once. Spoonbill data will be collected separately, noting maximum number of individuals seen, and approximate ratio of adults to immatures, non-breeding adults to breeding adults. Using a conservative stopping rule of 75% completeness, a survey will be deemed completed after (typically) 3 to five 15 minute sampling periods—i.e., a total effort of between 45 minutes and 75 minutes. These will be conducted twice, before and after the event. Resultant data will comprise richness estimates (of known completeness) for all waterbirds before and after the event (including cryptic species), standardised incidence estimates for all species detected, plus detailed demographic indices for two congeneric species of divergent ecologies. As Yellow-billed Spoonbills feed primarily on small waterborne inverts which in turn, respond directly (and rapidly) to water conditions, their presence, numbers and population composition give a sensitive, accurate and broad-based measure of overall water quality and biodiversity value. Given that some flows lead to short-term benefits whereas others entrain longer-term effects, a subset of wetlands may require a third survey (just for spoonbills). This would take less than one hour per wetland, and would focus on the other species, Royal Spoonbill. Unlike its congener, this species is primarily piscivorous, with fish taking longer to respond to changes in flow than small invertebrates. Hence, a flow that yields breeding in both species of spoonbills likely represents a more significant event for a broad range of other species than a flow that stimulates breeding in only Yellow-billed Spoonbills. This approach combines many of the specific recommendations made earlier. ¾ Observers are trained ¾ Sampling is timed relative to events of biological significance and management relevance ¾ Sampling is conducted at the patch scale ¾ Sample completeness is standardised, allowing explicit comparisons between surveys ¾ Call play-back is incorporated to enhance sampling resolution for cryptic groups ¾ Indicator species are chosen, reflecting their ecology and sensitivity to key underlying processes ¾ Focal species that can be readily identified, aged, and scored for breeding status are incorporated. Herons and cormorants could also be used as focal groups, but both would yield differing information—herons being more sensitive to water depth and reeds, cormorants more sensitive to water temperature and presence of dead trees. Moreover, as both groups are primarily piscivorous, their numbers would reflect water changes over a longer period than spoonbills. While the richness of duck species is greater still, their ecological requirements (particularly feeding preferences, habitat preferences for breeding) are less clearly demarcated, making interpretation and management response less straightforward. Community approach Functional groups Call playback Limited-area samples Patch-scale samples Incidence Approach /Method Abundance Most meaningful measure, fine grained and sensitive Estimated in either absolute or relative terms, this is a measure of how many individuals of a particular species were detected Also called reporting rate, this is the proportion of samples in which a species was recorded Using pre-existing biological features to define the sampling area (ie, the lake, the woodland remnant, the catchment) Typically rectangular (transects) or circular (point counts) areas of fixed dimensions, used to subsample larger areas Playing a pre-recorded vocalization of a particular species to elicit a response— typically a territorial call Rather than examining species, using ecological similarities (typically feeding, habitat use, nest site preference) to group similar species together Picks up subtle factors unclear with species-specific analyses, maximises sensitivity to variables of concern—ie match underlying variables to particular groups most liely to show signal Inclusive approach to sampling Includes all species, so ideal for longbiodiversity, incorporating all term studies, useful for comparisons Essential for determining whether cryptic species are present, efficient Can yield density estimates; best when samples are internally homogeneous and subsampled area is large and consistent subset of whole Yields data at patch scale, ideal when management/treatment is also applied at patch scale Less affected by flocking birds, easily calculated, Advantages Description Requires excellent identification skills, lots of noise, so overall patterns rarely Inaccurate when sites vary in area, or when sites exhibit high within-site heterogeneity; yield data at the sample scale—often not the most meaningful scale for inference or management Unable to give definitive data on absences, less accurate than count data for estimating numbers of individuals, ethic concerns during breeding season Assumes functional groups are well defined, simplistic and may not be comparable between regions Fairly coarse grained measure unable to distinguish subtle changes in numbers Can only be used in inherently patchy systems—ie where area of interest is biologically defined Issues of detectability, both between species and observers and also sites and times of year Disadvantages Indicator species species within a class (all between regions birds), or within an ecological subgroup of that class(waterbirds) Rather than examining all Most efficient approach species (either individually or collectively), using one species as an assay for wider patterns May miss other factors, and assumes species have been chosen correctly conclusive References cited Bibby C. J., Burgess, N. D., and Hill, D. A. (1992). Bird Census Techniques. Academic Press, London. Briggs S.V., Thornton S.A. and Lawler W.G. (1997) Relationships between hydrological control of river red gum wetlands and waterbird breeding. Emu 97, 31-42 Brooker, L. (2002). The application of focal species knowledge to landscape design in agricultural lands using the ecological neighbourhood as a template. Landscape and Urban Planning 60, 185–210. Conway, C. J. and Gibbs, J. P. (2005). Effectiveness of call-broadcast surveys for monitoring marsh birds. Auk 122, 26–35. Craig, M. D., and Roberts, J. D. (2001). Evaluation of the impact of time of day, weather, vegetation density and bird movements on outcomes of area searches for birds in eucalypt forests of south-western Australia. Wildlife Research 28, 33–39. Dawson, D. G. (1981). The usefulness of absolute (‘census’) and relative (‘sampling’ or ‘index’) measures of abundance. Studies in Avian Biology 6, 554–548. DIPNR (2003) Integrated monitoring of environmental flows - State summary reort 1998-2000. Driver, P., Chowdhury, S., Wettin, P. and Jones, H. (2004) Models to predict the effects of environmental flow releases on wetland inundation and the success of colonial bird breeding in the Lachlan River, NSW. In: Proceedings 4th Australian Stream Management Conference Herzog, S. K., Kessler, M., and Cahill, T. M. (2002). Estimating species richness of tropical bird communities from rapid assessment data. Auk 119, 749–769. Kingsford R.T. and Johnson W. Impact of water diversions on colonially-nesting waterbirds in the macquarie marshes of arid Australia. Colonial Waterbirds 21[2], 159-170. 1998. Kingsford R.T., Jenkins K.M. and Porter J.L. (2004) Imposed Hydrological Stability on Lakes in Arid Australia and Effects on Waterbirds. Ecology 85, 2478-2492 Krzys, Greg, Thomas A. Waite, Martin Stapanian, and John A. Vucetich. (2002). Assessing avian richness in remnant wetlands: towards an improved methodology. Wetlands 22(1), 186–190. Leslie D.J. (2001) Effect of river management on colonially-nesting waterbirds in the Barmah-Millewa forest, south-eastern Australia. Regulated Rivers-Research & Management 17, 21-36 Mac Nally, R., and Horrocks, G. (2002). Proportionate spatial sampling and equal-time sampling of mobile animals: a dilemma for inferring areal dependence. Austral Ecology 27, 405–415. Merikallio, E. (1958). Finnish birds: their distribution and numbers. Fauna Fennica 5, 1– 81. Peterson, A. T., and Slade, N. A (1998). Extrapolating inventory results into biodiversity estimates and the importance of stopping rules. Diversity and Distributions 4, 95–105. Recher, H. F. (1988). Counting terrestrial birds: use and application of census procedures in Australia. Australian Zoological Review 1, 25–45. Remsen, J. V., Jr. (1994). Use and misuse of bird lists in community ecology and conservation. Auk 111, 225–27. Rosenstock, S. S., Anderson, D. R., Giesen, K. M., Leukering, T., and Carter, M. F. (2002). Landbird counting techniques: current practices and an alternative. Auk 119, 46–53. Shields, W. M. (1979). Avian census techniques: an analytical review. In ‘The role of Insectivorous Birds in Forest Ecosystems’ (Eds J. G. Dickson, R. N. Connor, R.R. Fleet, J. C. Kroll & J.A. Jackson) pp. 23–51. (Academic Press, New York.) Stapanian, Martin A., Thomas A. Waite, Gregory Krzys, John J. Mack, Mick Micacchion. (2004). Rapid assessment indicator of wetland integrity as an unintended predictor of avian diversity. Hydrobiologia 520, 119–126. Stapanian, Martin A., Thomas A. Waite. (2003). Species density of waterbirds in offshore habitats in western Lake Erie. Journal of Field Ornithology 74, 381–393. Thompson, W. L. (2002). Overview: Towards reliable bird surveys: accounting for individuals present but not detected. Auk 119, 18–25. U.S. Fish & Wildlife Service and U.S. Geological Survey. (1999). Proceedings of the Marsh Bird Monitoring Workshop. U.S. Fish & Wildlife Service, Denver. CO. Verner, J. (1985). Assessment of counting techniques. In ‘Current Ornithology. Vol. 2’. (Ed. R. F. Johnstone.) pp. 247–301. (Plenum Press, New York.) Watson D.M. (2004). Comparative evaluation of new approaches to survey birds. Wildlife Research 31, 1–11. Watson, D. M. (2003). The “standardized search”: an improved way to conduct bird surveys. Austral Ecology 28, 515–25. Wright, D. H. (1991). Correlations between incidence and abundance are expected by chance. Journal of Biogeography 18, 463–66. Appendix 1: A Functional classification scheme for birds in the Lachlan catchment For management the two flow parameters most easily adjusted are duration of inundation and water depth. Birds will respond to these both directly and indirectly. For colonialnesting species there is a direct response to duration of inundation and water depth that results in initiation of breeding activity (Kingsford and Johnstone 1996; Leslie 2001). For others, changes in water depth alters access to food resources and/or the availability of prey items. Indirect effects of duration of inundation and water depth include the presence or not of fringing vegetation and the extent of exposed shoreline. These provide protection from predators and disturbance and/or feeding habitat for some species. The survey data were divided into functional groups based on feeding habit and responses to flow parameters. One possible scheme for dividing the wetland birds of the Lachlan River valley follows; Dabblers and probers—feed in the shallow margins of wetlands and include stilts, rednecked avocet, migratory shorebirds and most of the ducks. Colonial-nesting waders—this group are long-legged waders that either dabble in shallow water or probe for food. They distinguish themselves from other dabblers and probers by forming nesting colonies when conditions are suitable—ibis and spoonbills. Deep-water foragers—only occur on the deeper wetlands where water depth is ~1m or more. This group includes the black swan, Eurasion coot, musk duck, blue-billed duck and hardhead. Reed-nesting birds—rely on fringing vegetation for protection from predation and disturbance. This group includes the bitterns, crakes, rails, swamphen and several small passerines that occur exclusively in these habitats. Piscivores—fish eating species. This group includes grebes, terns, gulls, white-bellied sea-eagle and azure kingfisher. Colonial-nesting Piscivores—Piscivores that distinguish themselves from other piscivores by forming nesting colonies when conditions are suitable—cormorants, most herons and the Australian pelican Shoreline foragers—feed mostly on the exposed shore of retreating wetlands and their shallow margins. This disparate group includes Australian wood duck, Australian shellduck, plumed whistle-duck and dotterels. Woodland birds—all species not primarily dependent of aquatic habitats. Other—this group has two species not readily classified with any of the above groups because they occur in a broader range of habitats. These are the masked lapwing and the magpie goose. Table 3. Proposed functional groups for birds observed in the Lachlan River valley. Common Name Emu Peaceful Dove Diamond Dove Common Bronzewing Crested Pigeon Baillon's Crake Spotless Crake Black-tailed Native-hen Dusky Moorhen Purple Swamphen Eurasian Coot Great Crested Grebe Australasian Grebe Hoary-headed Grebe Great Cormorant Little Black Cormorant Pied Cormorant Little Pied Cormorant Darter Australian Pelican Marsh Tern Gull Billed Tern Silver Gull Masked Lapwing Black Fronted Dotterel Black-winged Stilt Banded Stilts Common Sandpiper Painted Snipe Glossy Ibis Sacred (Aust. White) Ibis Straw necked ibis Royal Spoonbill Yellow Spoonbill Great Billed Heron Little Egret Intermediate Egret Great Egret White-Faced heron White-Necked Heron Nankeen (Rufous) Night Heron Australiasian Bittern Magpie Goose Australian Wood (Maned) Duck Black Swan Plumed Whistling-Duck Australian Shelduck Pacific Black duck Chestnut Teal Grey teal Australiasian Shoveler Pink Eared Duck Musk Duck Swamp Harrier White Goshawk Scientific Name Dromaius novaehollandiae Geopelia striata Geopelia cuneata Phaps chalcoptera Ocyphaps lophotes Porzana pusilla Porzana tabuensis Gallinula ventralis Gallinula tenebrosa Porphyrio porphyrio Fulica atra Podiceps cristatus Tachybaptus novaehollandiae Poliocephalus poliocephalus Phalacrocorax carbo Phalacrocorax sulcirostris Phalacrocorax varius Phalacrocorax melanoleucos Anhinga melanogaster Pelecanus conspicillatus Chlidonias hybridus Sterna nilotica Larus novaehollandiae Vanellus miles Elesyornis melanops Himantopus himantopus Cladorhynchus leucocephalus Actitis hypoleucos Rostratula benghalensis Plegadis falcinellus Threskiornis molucca Threskiornis spinicollis Platalea regia Platalea flavipes Ardea sumatrana Egretta garzetta Ardea intermedia Ardea alba Egretta novaehollandiae Ardea pacifica Nycticorax caledonicus Botaurus poiciloptilus Anseranas semipalmata Chenonetta jubata Cygnus atratus Dendrocygna eytoni Tadorna tadornoides Anas superciliosa Anas castanea Anas gracilis Anas rhynchotis Malacorhynchus membranaceus Biziura lobata Circus approximans Accipiter novaehollandiae Functional group woodland woodland woodland woodland woodland reed-nesting reed-nesting reed-nesting reed-nesting reed-nesting deep-water forager piscivore deep-water forager deep-water forager colonial-nesting piscivore colonial-nesting piscivore colonial-nesting piscivore colonial-nesting piscivore piscivore colonial-nesting piscivore piscivore piscivore piscivore other shoreline forager dabblers and probers dabblers and probers dabblers and probers reed-nesting colonial-nesting wader colonial-nesting wader colonial-nesting wader colonial-nesting wader colonial-nesting wader piscivore colonial-nesting piscivore colonial-nesting piscivore colonial-nesting piscivore piscivore piscivore colonial-nesting piscivore reed-nesting other shoreline forager deep-water forager shoreline forager shoreline forager dabblers and probers dabblers and probers dabblers and probers dabblers and probers dabblers and probers deep-water forager woodland woodland Wedged Tail Eagle Little Eagle White Bellied Sea Eagle Whistling Kite Black (fork-tailed) Kite Square-tail Kite Black Shouldered Kite Letter Winged Kite Grey Falcon Brown Falcon Nankeen Kestrel Sulfur crested cockatoo Little Corella Galah Cockatiel Crimson Rosella Yellow Rosella Eastern Rosella Australian Ringneck Red Rumped Parrot Blue Bonnet Blue-winged Parrot Azure Kingfisher Laughing kookaburra Sacred kingfisher Fork-tailed Swift Pallid cuckoo Black eared cuckoo Welcome Swallow Tree Martin Fairy Martin Willie Wagtail Jacky Winter Red Capped Robin Hooded Robin Eastern Yellow Robin Rufous Whistler Gilberts Whistler Grey Shrike-Thrush Magpie Lark Black Faced Cuckoo Shrike Grey Crowned Babbler White Browed Babbler Western Gerygone (Warbler) Weebill Southern Whiteface Yellow (Little) Thornbill Western Thornbill Inland Thornbill Yellow Rumped Thornbill Brown Songlark Rufous Songlark Clamorous Reed-Warbler Golden-headed Cisticola Superb Fairy (Blue) Wren Variegated Fairy Wren Black Faced Woodswallow Brown Treecreeper Aquila audax Hieraaetus morphnoides Haliaeetus leucogaster Haliastur sphenurus Milvus migrans Lophoictinia isura Elanus axillaris Elanus scriptus Falco hypoleucos Falco berigora Falco cenchroides Cacatua galerita Cacatua sanguinea Cacatua roseicapillus Nymphicus hollandicus Platycercus elegans, race elegans Platycercus elegans, race flaveolus Platycercus eximius Barnardius zonarius Psephotus haematonotus Northiella haematogaster Neophema chrysostoma Alcedo azurea Dacelo novaeguineae Todiramphus sanctus Apus pacificus Cuculus pallidus Chrysococcy) osculans Hirundo neoxena Hirundo nigricans Hirundo ariel Rhipidura leucophrys Microeca fascinans Petroica goodenovii Melanodryas cucullata Eopsaltria australis Pachycephala rufiventris Pachycephala inornata Colluricincla harmonica Grallina cyanoleuca Coracina novaehollandiae Pomatostomus temporalis Pomatostomus superciliosus Gerygone fusca Smicrornis brevirostris Aphelocephala leucopsis Acanthiza nana Acanthiza inornata Acanthiza ewingii Acanthiza chrysorrhoa Cincloramphus cruralis Cincloramphus mathewsi Acrocephalus stentoreus Cisticola exilis Malurus cyaneus Malurus lamberti Artamus cinereus Climateris picumnus woodland woodland piscivore woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland piscivore woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland Mistletoe Bird Silvereye Striped Honeyeater Pied Honeyeater White Plumed honeyeater Noisy Miner Yellow Throated Miner Blue Faced Honeyeater Noisy Friarbird Little Friarbird Diamond Firetail Finch Double-barred finch Apostle Bird White winged chough Pied Currawong Pied Butcher Bird Grey Butcher Bird Australian Magpie Little Wattlebird Grey Fantail Restless Flycatcher Australian Raven Little Raven Striated Pardalote European Goldfinch Common Myna Common Starling Dicaeum hirundinaceum Zosterops lateralis Plectorhyncha lanceolata Certhionyx variegatus Lichenostomus penicillatus Manorina melanocephala Manorina flavigula Entomyzon cyanotis Philemon corniculatus Philemon citreogularis Stagonopleura guttata Taeniopygia bichenovii Struthidea cinerea Corcorax melanorhamphos Strepera graculina Cracticus nigrogularis Cracticus torquatus Gymnorhina tibicen Anthochaera chrysoptera Rhipidura fuliginosa Myiagra inquieta Corvus coronoides Corvus mellori Pardalotus striatus Carduelis carduelis Acridotheres tristis Sturnus vulgaris woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland woodland
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