The avifauna of severely fragmented, • Buloke Allocasuarina luehmanni woodland In western Victoria, Australia

DAVID M. WATSON,I RALPH Mac NALLY 2 and ANDREW F. BENNETT3

Buloke Allocasuarina luehmanni woodland is an endangered habitat type that was once widespread in southern Australia but now is restricted to a series of remnants, many of which are located in the Wimmera region of western Victoria. The bird communities inhabiting 27 remnants were sampled on transects of 1.0 ha at 6-week intervals over one year. Ninety-five species of birds were observed in Buloke woodlands, of which 66 species were recorded during transect counts. The total species richness, the richness of groups of birds based on their habitat use, and the composition of assemblages on transects, were examined in a series of analyses with respect to habitat type (which probably reflects "quality"), remnant area, shape, isolation and geographic location. There was little evidence for attributes of remnants significantly influencing species richness per transect, but the composition of avifaunal assemblages varied in relation to habitat type, size and geographic location. The Buloke woodlands supported a diverse assemblage, including numerous species believed to be experiencing a regional decline' in southern Australia. The composition of the Buloke assemblage has similarities to those of dry eucalypt woodlands across the plains of central Victoria, but elements contributing to a distinctive avifauna include species associated with semi-arid and mallee environments, and a high frequency of occurrence of a group of small insectivores (thornbills, robins) that favour dry She-oak and Callitris woodlands. A likely reason for the rich representation of small insectivores in woodland stands, even in small degraded remnants grazed by stock, was the scarcity of the Noisy Miner Manorina melanocephala that are often common in eucalypt remnants and known to aggressively exclude other species. The results of this study add weight to the recommendation that protection and restoration of Buloke woodlands is a priority for conservation in the Wimmera bioregion of Victoria. Key words: Allocasuarina luehmanni, Australia, Birds, Buloke, Habitat degradation, Habitat fragmentation, Woodlands.

INTRODUCTION and Traill 1996; Lunt and Bennett, in press). There has been a documented decline in the THE loss and fragmentation of natural environ­ local occurrence of numerous woodland­ ments in areas primarily committed to agri­ dependent species of birds and in their cultural land uses, and the impact of these distribution and status across woodland habitats processes on the avifauna, is an issue of concern of southern Australia (Saunders 1989, 1993; throughout the world (Lynch and Whigham Webster and Ahern 1992; Menkhorst 1993; 1984; Opdam et al. 1985; Diamond et al. 1987; Robinson 1993; Traill et al. 1996). In many Newmark 1991; McCollin 1993; Barrett et al. areas, small remnants ofwoodland now comprise 1994; Newton 1994; Wiens 1995). In Australia, the main habitats upon which such species particular concern has been expressed about depend. Consequently, increased attention has the effects of landscape change on bird species been given to assessing the value of remnants and communities in temperate and semi-arid of different sizes, vegetation types, management woodland ecosystems (Saunders 1989; Robinson history and landscape position for maintaining 1993; Saunders and Ingram 1995; Robinson and woodland bird populations in agricultural Traill 1996). These woodland ecosystems have regions (Loyn 1987; Arnold and Weeldenburg been preferentially cleared for agriculture because 1990; Barrett et al. 1994; Fisher and Goldney they generally occur on flat, relatively fertile 1997; Bennett and Ford 1997). soils with the greatest agricultural potential. A number of regions in southern Australia In this study, we examine the bird communities now retain less than 15% native vegetation associated with Buloke (also known as "Bull­ (e.g., Northern Plains, Victoria; Western Slopes, oak") woodlands in the Wimmera region of New South Wales; southern Brigalow Belt, western Victoria. Extensive clearing has much Queensland; Wheatbelt, Western Al,lstralia), and reduced the extent of this vegetation, which was particular types of woodland vegetation such formerly widespread across western Victoria as that dominated by White Box Eucalyptus (Connor 1966; Land Conservation Council 1985). alhens, Grey Box E. microcarpa, Salmon Gum Present Buloke woodlands are most appro­ E. salmonophloia, Brigalow Acacia harpophylla and priately viewed as a system of small isolated Buloke Allocasuarina luehmanni are now severely remnants set within a dominant agricultural depleted (Prober and Thiele 1993; Saunders matrix rather than a fragmented woodland et al. 1993; Biodiversity Unit 1995; Robinson per se. For a set of woodland remnants, we

IEnvironmental Studies Unit, Charles Sturt University, Bathurst, New South Wales, Australia 2794. 2Corresponding author: Section of Ecology. Department of Biological Sciences, Monash University, Clayton. Victoria. Australia 3168. 'School of Ecology and Environment, Deakin University - Rusden Campus, 662 Blackburn Road. Clayton. Victoria. Australia 3168. PACIFIC CONSERVATION BIOLOGY Vol. 6: 46-60. Surrey Beatty & Sons, Sydney. 2000. WATSON, Mac NALLY and BENNETT: AVIFAUNA OF BULOKE WOODLANDS 47

collected information on remnant area, isolation, contributing factors include other introduced shape and habitat structure, as well as the set herbivores (e.g., European Rabbit Oryctolagus of bird species associated with each remnant. cuniculus), timber removal and the spread of weed species (Land Conservation Council 1985). From these data, we address two main objectives. First, we document the assemblage Preliminary surveys located 50 remnants of of birds utilizing Buloke woodlands. There Buloke woodland in the study region. For each have been several surveys of the avifauna in the remnant, the area, canopy composition, tree area, some of which have recorded rare and density, understorey characteristics and manage­ uncommon species (e.g., Land Conservation ment status (intensity of sheep grazing) were Council 1985), but none of these surveys were recorded. From this set, 27 remnants were restricted to this habitat and the composition selected for study to provide combinations of of the Buloke woodland avifauna is not well size, management status and habitat complexity. known. Second, Buloke itself has been classed Preference was given to remnants > 2 ha in 1 as a depleted species (Gullan et al. 1990), and area and with a tree density > 50 stems ha- • All with the few remaining patches still threatened, remnants were on the same landform (Land it is listed under the Victorian Flora and Fauna Conservation Council 1985) and were located Guarantee Act. Thus, there is a need to evaluate within an area of 3200 km2 (Fig. 1). Twenty-two the relative conservation value of different remnants were on freehold land while the types of remnants if the system and its fauna remaining five were within Flora and Fauna are to be preserved. Consequently, we examine Reserves (remnants 1 and 15) or small blbCks of the relationship between the richness and Crown land (remnants 2, 5 and 22). composition of bird assemblages and the characteristics of Buloke remnants to further Censuses understand patterns of distribution of birds in the Buloke system. Bird populations were sampled by transect counts (Recher 1989). Fixed transect dimensions were used - 200 X 50 m - yielding a sampling METHODS area of 1 ha. For consistency, transects were located > 20 m from the edge of the remnant, Study area the maximum value possible in the smallest The study region is in northwestern Victoria remnant. Transects were randomly located in adjoining South Australia, and is dominated remnants apart from this restriction. All censuses by the extensive Wimmera plains. Primarily were conducted by DMW. Census time was of aeolian origin, this landform comprises standardized to 20 min., with 5 min. taken to low, elongate sand-ridges or linear dunes with traverse each 50 m section following a path sandy clay soils (Land Conservation Council down the centre of the transect. All birds 1985). The climate is temperate with hot, dry observed within the transect were recorded, as summers and mild winters. About 60% of were individuals outside the transect but within rainfall occurs during the winter months (Land the remnant. Unless otherwise specified, the Conservation Council 1985). Severe frosts are avian data discussed here are the accumulated common during winter. species richness within the 1 ha transect. The Wimmera plains were originally covered Over the course of the study, each remnant with open woodlands dominated by Grey Box, was censused nine times at six-weekly intervals Yellow Gum Eucalyptus leucoxylon, Black Box between April 20, 1994 and March 22, 1995. E. largiflorens and Buloke, interspersed with Three trips were conducted prior to the first expanses of native grassland (Connor 1966; date for familiarization with sites and bird Land Conservation Council 1985). Virtually species and to refine methods. Censuses were all of the grasslands were alienated by early conducted in early morning (beginning later European settlers and the woodlands were than 30 min. after sunrise) and late afternoon subsequently cleared. Agricultural cropping for (completed earlier than 30 min. before sunset), wheat is now the major land use in the region. with at least four morning censuses in each Surviving Buloke woodlands occur on freehold remnant. Censuses were conducted only in still, farmland, roadside strips and in small reserves. dry conditions. A constrained randomization Most are degraded to varying degrees. In technique was developed to determine the order severely degraded remnants, floristic richness in which transects within remnants were is low and there is little regeneration of Buloke, censused. The 27 remnants were grouped into with most trees being of the same age and size. five sectors according to their geographic The understorey is dominated by exotic grasses proximities. The order in which these sectors and there may be areas of bare, compacted soil. were visited was randomized, and the order in While the main agent of degradation appears which the transects within remnants within each to be grazing by domestic stock, further sector were censused was also randomized. 48 PACIFIC CONSERVATION BIOLOGY

25

Little Desert National Park 22 21

Rupanyup N - Victoria 10km 1

Fig. 1. Map of the study area in western Victoria, showing the location of 27 remnants selected as study sites; main roads are shown as straight lines.

Vegetation the entire transect. Since the sampling unit was 1 ha, these data are measures of density in Differences in vegetation have often been 1 individuals ha- . A stratified sampling protocol found to have a profound effect on the com­ was employed to measure the understorey. The position and dynamics of bird communities transect was subdivided into four 50 X 50 m (Kitchener et ai. 1982; Loyn 1987; Lynch 1987; squares and within each square, a 2 X 2 m Mac Nally 1990). The description of vegetation quadrat was randomly located (Kent and Coker in ornithological studies has usually been 1992). For each quadrat, the percentage-covers divided into two different elements: habitat of introduced grasses, native grasses, leaf litter structure (or "physiognomy") and floristic and bare ground were estimated. composition (Mac Nally 1995: 270-271). For this study, aspects of both were measured. Of the 21 variables measured, many related to Vegetation at each site was surveyed on the physiognomic characteristics of the vegetation same transects as those used to sample the (Table 1). To aid interpretation, the data were avifauna. All trees located within the transect ordinated by using multidimensional scaling were identified and counted. Buloke trunks were (MDS, see Kruskal and Wish 1978). Prior to measured by using a diameter tape at a height calculating the dissimilarity matrix for use of 1.5 m (referred to as diameter at breast height in the MDS, all proportionate variables were or DBH) and assigned to one of five size classes arcsine-square-root transformed (Winer 1971). (see Table 1 for dimensions). Individuals of None of the other variables required trans­ other tree species were identified and tallied. formation but they were standardized to lie Shrubs within the transect were also counted, between 0 and 1. This ensured that all 21 and assigned to one of three size classes habitat variables were weighted equally and (small, intermediate and large) based on height prevented the ordination from being unduly (Table 1). Counts of logs, ant nests, mistletoe influenced by extreme values and highly skewed plants and dead trees were also made for distributions. The dissimilarity matrix was WATSON, Mac NALLY and BENNETT: AVIFAUNA OF BULOKE WOODLANDS 49

Table 1. List of the 21 habitat variables measured for remnant Buloke woodlands with definitions and Spearman rank-correlations of each variable with the first two MDS dimensions. Variables are listed in order of decreasing correlation with MDS 1. Correlations significant at the 0.01 (25 d.f.) level are in bold type.

Variable MDSI MDS2 Total number of shrubs within transect 0.908 0.084 Total number of eucalypt trees within transect 0.806 0.226 Percent cover of bare ground 0.787 0.071 Proportion of shrubs 30-80 cm tall 0.776 0.024 Number of eucalypts with a DBH > 17 cm 0.763 0.252 Proportion of Bulokes with a DBH of 5-8 cm 0.647 0.210 Proportion of eucalypts that are E. leucoxylon 0.612 0.179 Proportion of shrubs 80-130 cm tall 0.500 0.500 Proportion of Buloke trees with a DBH of 1-4 cm 0.487 0.356 Proportion of large eucalypts that are E. largiflorens 0.443 0.561 Proportion of shrubs > 130 cm tall 0.422 0.371 Proportion of Bulokes with a DBH of 9-16 cm 0.419 -0.620 Total number of Buloke trees within transect 0.375 -0.393 Percent cover of leaf litter 0.130 -0.412 Number of large ant nests within transect 0.128 0.224 Number of standing dead trees within transect -0.121 0.009 Number of fallen logs within transect -0.303 0.110 Proportion of Bulokes with a DBH >32 cm -0.409 0.695 Percent cover of all grasses -0.445 0.409 Proportion of Bulokes with a DBH of 17-32 cm -0.504 0.509 Percent cover of exotic grasses -0.581 0.001 calculated by using the Czekanowski measure index of 0.88, with a higher number indicating of dissimilarity (Minchin 1987). Ordinations greater edge to area ratio. were performed in one to four dimensions, with 20 random initial configurations for each Degree of isolation is frequently used as a dimension to deal with degenerate solutions variable in fragmentation and remnant studies (Wilkinson 1989). Unlike other ordination (e.g., Kitchener et al. 1982), and has been techniques, the dimensions of an MDS solution typically viewed as the distance of a habitat are not expressed as functions of the measured fragment from the nearest large tract of variables. To interpret the ordination, pair-wise comparable vegetation (e.g., Forman et al. 1976; Spearman rank correlations were conducted Lynch and Whigham 1984). No such large between the scores for each axis and the expanse of Buloke woodland remains. Instead, 21 habitat variables (Table 1). This process an index of isolation was calculated for each site whereby the combined area of all wood­ permitted the interpretation of the vegetation 2 variables that were most strongly correlated land within 2.5 km (19.6 km ) of the centre of with each of the MDS dimensions. We used a a site (but excluding the area of the remnant) conservative probability level of 0.01 because of was calculated by using digitized, large-scale the large number of correlations calculated on vegetation maps and an image analyser (see the one data set in this context. Otherwise, we Table 2 for values of area, isolation and shape use the conventional a = 0.05 significance level. for the 27 remnants).

Other independent variables Other analyses In addition to the habitat descriptors, we On the basis of observed characteristics of measured the area of each remnant by direct bird species, the avifauna at each remnant was measurements from recently compiled, large­ partitioned into four groups of: "open-country", scale vegetation maps. A shape index (S) similar "woodland-nesting, open-country feeders", to an algorithm used by Schumaker (1996) was "woodland (widespread)" and "woodland also calculated for each remnant according to (restricted)" species (see Results for definitions). the formula: Each remnant could be allocated to classes based on characteristics such as vegetation S (A/P)/(R/2), = structure, area and isolation. These partitions where: A is the area of a remnant, P is the allowed us to analyse the species composition perimeter, and R is the radius of the circle that of sets of remnants in greater detail than just has an area of A. This index relates remnant gross species richness, and to explore how shape to a circle of equal area and is thus local diversity was generated. For example, all allometrically independent of area. According transects within remnants were allocated to to this formula, a perfect square has a shape a size class, and the numbers of each of the 50 PACIFIC CONSERVATION BIOLOGY

Table 2. Species richness of woodland birds, and areas, habitat-type classification (see text), shapes and isolation (see text) for each remnant of buloke woodland.

Remnant No. Richness Area (ha) Shape Habitat type Isolation (ha) I 20 467 1.83 I 4 2 14 13.5 1.18 I 0 3 9 9.0 1.28 I 0 4 13 5.4 1.13 I 8 5 12 50 1.19 1 7 6 23 12 1.15 3 5 7 9 3.7 1.20 4 4 8 11 27 1.38 2 4 9 16 3.2 IAI 3 10 10 13 6.1 1.13 3 II 11 8 4.5 1.13 3 II 12 14 2.5 1.50 4 7 13 15 5.4 1.68 2 5 14 18 2.6 1.52 2 5 15 10 207 1.04 I 6 16 10 3.4 1.15 4 5 17 15 13.5 1.15 3 17 18 10 14.4 0.83 3 4 19 15 18 1.20 4 5 20 12 7.5 1.14 4 5 21 15 4.9 1.27 3 0 22 11 10.7 1.13 2 10 23 8 49 1.13 2 0 24 17 28.8 1.17 3 0 25 12 10.8 1.13 3 0 26 17 15 1.38 2 10 27 17 9 1.28 2 15 groups of species could be calculated. This between groups. All compositional analyses were allowed the data in each such partition to based on presence/absence data and Bray-Curtis be treated as a repeated-measures ANOVA of indices. species richness, where the "between" factor was area (or isolation, etc.) and the "within" (or RESULTS repeated) factor was group of bird. Means Vegetation comparisons were conducted by using the KKS procedures (Keselman and Keselman 1993). The 27 remnants of Buloke woodland exhibited We note that the treatment of the independent broad variation in the variables examined, variables in this way potentially occludes inter­ ranging from dense, almost mallee-like vegetation actions between the variables vis-a-vis the avian (i.e., many thin stems) to sparse open woodland. groups, but note that the distribution of values For example, total Buloke density varied from of these variables among the others is not 50 to 1090 stems ha-1 (215.5 ± 214.4 SD). "random" and so would likely violate analytical In addition to absolute number, the size-class approaches in any event. Thus, our deductions distribution of Buloke trees also varied, with should be interpreted in light of potential inter­ some transects within remnants having dependence among the independent variables. representatives of all five size classes and others having only one. The only other canopy trees We used one-way analyses of similarities recorded were two species of eucalypt, Black (ANOSIMs, Clarke 1993) to compare remnants Box and Yellow Gum. In all transects within of different characteristics for compositional remnants, the density of eucalypts was less differences/similarities. There are N(N - 1)/2 than that of Bulokes, and varied from 0 to combinations for N groups in pair-wise compari­ 53 stems ha-1 (9.15 ± 14.6). One of the key sons, so we employed a sequential Bonferroni structural variables was the presence and nature procedure (Holme 1979) to test for statistical of an understorey, with some transects within significance. While ANOSIMs address global and remnants having a well-developed and species­ group-wise differences/similarities in avifaunas, another technique is needed. The similarity rich shrub layer while others had an open grassy layer. Total shrub density ranged from 0 to percentage (SIMPER, Carr 1994) was used 114 ha-1 (93.15 ± 228.2). to identify those species contributing most to similarities within groups and to differences The MDS ordinations indicated that two among groups of remnants. Within-group dimensions were sufficient to provide a good SIMPERS indicate species "characterizing" the overall "fit" for the vegetation data (stress <0.1, group, while among-group SIMPERs identify Kruskal and Wish 1978). Of the 20 random those species contributing most to the differences initial configurations, 16 produced the same WATSON, Mac NALLY and BENNETT: AVIFAUNA OF BULOKE WOODLANDS 51

low stress value indicating that these almost the regeneration of Buloke and the growth of certainly represented the globally best solution. shrubs. Transects within remnants in "Type 3" One of these solutions was randomly selected as were degraded with no regeneration of Buloke, the basis for further analysis (see Fig. 2). no shrub layer, no eucalypts and many dead standing trees and logs. The understorey was Many of the ordination variables were signifi­ composed of native and exotic grasses. "Type 4" cantly correlated with the first MDS dimension consisted of highly disturbed remnants typically (Table 1). Variables positively correlated with supporting a single size-class of Buloke. Tree scores on MDS 1 were indicative of transects density usually was great, with stunted trees within remnants with many eucalypts (especially clumped into copses separated by areas of bare of Yellow Gum), a dense layer of small shrubs ground and exotic grasses. Sheep were present and Bulokes, and with high cover of bare in all of these remnants. ground. Variables negatively correlated with MDSI were proportions of mature Bulokes and Bird communities of exotic grasses. MDS2, on the other hand, represented a gradient from transects within Ninety-five species of birds were recorded remnants with many mature Bulokes (but with during this study. Sixty-six were recorded on little regeneration), substantial numbers of larger transects during censuses, 15 were seen outside shrubs and a higher fraction of Black Box, to the transect area during censuses and 14 remnants with many smaller Bulokes (Table 1). were recorded from incidental observations in the study sites (Table 3). Within the entire assemblage of birds recorded in the wood­ MDS2 lands, seve--al phylogenetic groups were well 2 represented including: diurnal raptors (11 species), parrots and cockatoos (8), robins and • allied species (11) and honeyeaters (14). Type 1 No species was recorded in all transects within • remnants, although the Yellow-rumped Thornbill • • Type 2 (see Table 3 for Latin binomial names) was 00 the most widespread species (18 remnants), 0 0 followed by the Yellow Thornbill (17), Red­ capped Robin and Australian Magpie (both 16). a -1.5 -1 0.5 Other common species, found in at least half a of the remnants were the Red-rumped Parrot, MDS1 a -0.5 Hooded Robin, Willie Wagtail and the White­ ~ plumed Honeyeater. Species richness varied • from 8 to 23 (mean = 13.5) species per remnant. a • Several rare and locally uncommon species were -1.5 • Type 3 also recorded, including Blue Bonnet, Black­ Type 4 a eared Cuckoo, Gilbert's Whistler, Southern -2 Scrub-robin and Black Honeyeater. Fig. 2. Plot of the positions of 27 study remnants and four Many species commonly observed in eucalypt classes of Buloke habitat in ordination space: Type woodlands to the south and east were either I (solid circles); Type 2 (open circles); Type 3 (solid squares); Type 4 (open squares); habitat types are rare (e.g., Peaceful Dove, Common Bronzewing, described in the text. Golden Whistler, Red Wattlebird, New Holland Honeyeater) or were not recorded (e.g., Crimson Rosella Platycercus elegans, Eastern Yellow Robin Based on the MDS ordination space, transects Eopsaltria australis, Superb Fairy-wren Malurus within remnants could be partitioned into four eyaneus, Buff-rumped Thornbill Acanthiza reguloides, "types" based on habitat similarity (Fig. 2), which Pied Currawong Strepera versicolor) in Buloke we believe largely reflects habitat "quality". woodlands. Transects within remnants in "Type 1" (Fig. 2) were woodlands undergoing vigorous regener­ Based on distribution and frequency of ation, with well-developed shrub layers and occurrence within the study area, most of the abundant eucalypts. All of these remnants have 66 species recorded on transects fell readily been fenced (and thus protected from sheep into one of four groups (refer to Table 3 for grazing), most for at least ten years. "Type 2" numbers of records for each species and group is an intermediate category with transects within allocation). Note that these groupings do not remnants typified by a sparse shrub layer, reflect the conservation status of the species in presence of Buloke saplings and associated general, but indicate status within the study Black Box. These remnants had been grazed area. Moreover, all analyses refer only to these but have been fenced recently (~3 yr), allowing 66 species and exclude the 29 off-transect and 52 PACIFIC CONSERVATION BIOLOGY

Table 3. Bird species recorded in Buloke woodland remnants in the Wimmera. "Sites" are numbers of remnants in which the species was recorded; "records" are the total numbers of individuals observed from transect counts. "Group" refers to the class to which the species was allocated (Or, OW, WC or WU, see text). * denotes records within remnants during censuses but not on transects; t indicates incidental observations within study remnants.

Common name Linnean name Sites Records Group * Nankeen Night Heron Nycticorax caledonicus White-faced Heron Egretta novaehollandiae OP *Australian Shelduck Tadorna tadornoides t Australian Wood Duck Chenonetta jubata * Collared Sparrowhawk Accipiter cirrhocephalus t Brown Goshawk Accipiter fasciatus * Spotted Harrier Circus assimilis t Black Kite Milvus migrans t Whistling Kite Haliastur sphenurus * Little Eagle Hieraaetus morphnoides *Wedge-tailed Eagle Aquila audax Black-shouldered Kite Elanus axillaris I I OP Brown Falcon Falco berigora 3 4 OP Nankeen Kestrel Falco cenchroides 2 3 OP Australian Hobby Falco longipennis 1 3 OP t Bush Stone-curlew Burhinus grallarius * Painted Button-quail Turnix varia t Stubble Quail Corturnix pectoralis * Masked Lapwing Vanellus miles Crested Pigeon Ocyphaps lophotes 11 30 OW Common Bronzewing Phaps chalcoptera 1 2 WU * Peaceful Dove Geopilea striata Galah Cacatua roseicapilla 12 62 OP Long-Billed Corella Cacatua tenuirostris 1 1 OW *Yellow-tailed Black Cockatoo Calyptorhynchus funereus Purple-crowned Lorikeet Glossopsitta porphyrocephala 2 7 WU t Australian Ringneck Barnardius zonarius Eastern Rosella Platycercus eximius 5 8 OW Red-rumped Parrot Psephotus haematonotus 13 39 OW Blue Bonnet N orthiella haematogaster 3 8 OW Fan-tailed Cuckoo Cacamantis flabelliformis I 1 WU * Pallid Cuckoo Cuculus pallidus Black-eared Cuckoo Chrysococcyx osculans 1 I WU Horsfield's Bronze-cuckoo Chrysococcyx basalis 1 1 WU Barn Owl Tyto alba 1 2 OW Southern Boobook Ninox novaeseelandiae 4 5 WC * Sacred Kingfisher Todiramphus sancta Laughing Kookaburra Dacelo novaeguineae 3 7 OW Welcome Swallow Hirundo neoxena 2 3 OP Tree Martin Hirundo nigricans 1 5 OW Rufous Songlark Cinclorhamphus mathewsi 2 2 WU Black-faced Cuckoo-shrike Coracina novaehollandiae 3 6 WU White-winged Triller Lalage sueurii 1 I WU Red-capped Robin Petroica goodenovii 16 61 WC Hooded Robin Melanodryas cucullata 14 36 WC Jacky Winter Microeca fascinans 8 14 WC t Southern Scrub-robin Drymodes brunneopygia Golden Whistler Pachycephala pectoralis I 1 WU Rufous Whistler Pachycephala rufiventris 8 14 WC Gilbert's Whistler Pachycephala inornata 1 1 WU Grey Shrike-thrush Colluricincla harmonica 3 5 WU Restless Flycatcher Myiagra inquieta 3 11 WU Grey Fantail Rhipidura fuliginosa 8 9 WC Willie Wagtail Rhipidura leucophrys 15 47 OP White-Browed Babbler Pomatostomus superciliosus I 3 WU Variegated Fairy-wren Malurus lamberti 3 9 WU Weebill Smicrornis brevirostris I 2 WU Southern Whiteface Aphelocephala leucopsis 2 4 WU Yellow Thornbill Acanthiza nana 17 74 WC Yellow-rumped Thornbill Acanthiza chrysorrhoa 18 54 OW Chestnut-rumped Thornbill Acanthiza uropygialis 12 25 WC Varied Sitella Daphoenositta chrysoptera II 15 WC Brown Treecreeper Climacteris picumnus 7 47 WC Spiny-cheeked Honeyeater Acanthagenys rufogularis 6 31 WC Red Wattlebird Anthochaera carunculata I 2 WU t Yellow-throated Miner Manorina jlavigula Noisy Miner Manorina melanocephala 3 15 WU t White-naped Honeyeater Melithreptus lunatus WATSON, Mac NALLY and BENNETT: AVIFAUNA OF BULOKE WOODLANDS 53

Table 3 - continued.

Common name Linnean name Sites Records Group Brown-headed Honeyeater Melithreptus brevirostris 2 4 WU White-eared Honeyeater Lichenostomus leucotis I I WU Singing Honeyeater Lichenostomus virescens 6 28 WC Yellow-plumed Honeyeater Lichenostomus ornatus I I WU White-plumed Honeyeater Lichenostomus penicillatus 14 67 WC Fuscous Honeyeater Lichenostemus fuscus I I WU White-fronted Honeyeater Phylidonyris albifrons 12 14 WC * New Holland Honeyeater Phylidonyris novaehollandiae Black Honeyeater Certhionyx niger 6 8 WC Silvereye Zosterops lateralis 2 2 WU Mistletoebird Dicaeum hirundinaceum 10 18 WC t Spotted Pardalote Pardalotus punctatus Striated Pardalote Pardalotus striatus 10 41 WC t Diamond Firetail Stagonpleura guttata European Goldfinch Carduelis carduelis 5 12 OP House Sparrow Passer domesticus 14 70 OP Common Starling Sturnus vulgaris 10 42 OP t Olive-backed Oriole Griolus sagittatus Australian Magpie-lark Grallina cyanoleuca 2 5 OP *White-winged Chough Corcorax melanorhamphos Dusky Woodswallow Artamus cyanopterus 10 21 WC Black-faced Woodswallow Artamus cinereus I I ow White-browed Woodswallow Artamus superciliosus 3 5 OW * Grey Butcherbird Cracticus torquatus Australian Magpie Gymnorhina tibicen 16 94 OP t Little Raven Corvus mellori Australian Raven Corvus coronoides 3 16 OP incidentally recorded species, which were not (2) area; (3) shape; (4) degree of isolation; and surveyed consistently and may produce biased (5) geographic location. For each attribute, results (e.g., species with louder calls or of larger the 27 remnants were grouped into classes size being more easily observed off-transect). and mean values of species richness were then "Open-country species" (mnemonic OP): 14 calculated for each class. For example, transects species that have little reliance on woodlands for within remnants were allocated into four classes feeding and breeding, including the Australian vis-a-vis remnant area (small, medium, large, Magpie, Galah and Willie Wagtail (Table 3). very large) and the mean numbers of Op, Ow, WC and WU species for each size class were "Woodland-nesting, open-country feeders" then calculated. (OW): 11 species that often or usually feed in open country but depend on woodland Habitat types patches for breeding. The group included the Eastern Rosella, White-browed Woodswallow and The 27 transects within remnants were Red-rumped Parrot (Table 3). partitioned into four classes according to the habitat ordination (Fig. 2): Type 1 ("vigorously "Common (widespread) woodland species" regenerating"; six remnants), Type 2 ("inter­ (WC): 18 species that are woodland-dependent mediate"; seven remnants), Type 3 ("degraded"; for breeding and feeding and occurred in nine remnants) and Type 4 ("highly disturbed"; four or more remnants. The Yellow Thornbill, five remnants). There was no significant White-plumed Honeyeater, Rufous Whistler and difference in species richness among transects Singing Honeyeater were included in this group within remnants in the four habitat types (Table 3). (Table 4), with the mean varying little from "Uncommon (restricted) woodland species" 12.0 (Type 4) to 14.3 species (Type 3) (Fig. 3a). (WU): 23 species of woodland-dependent birds In this and all subsequent analyses, there that were found in three or fewer transects was highly significant difference in richness within remnants. The Grey Shrike-thrush, Golden among bird classes (Table 4), with mean Whistler, Weebill and Fuscous Honeyeater were richnesses being 1.4 (WU), 2.2 (OW), 3.2 (OP) some of these taxa (Table 3). and 6.6 (WC). However, this is of little interest because of way in which we partitioned species Distribution patterns among groups. It is the "between-factor" and In this section we analyse the distribution of the "groups X between-factor" terms that hold groups of avian species (i.e., Op, Ow, WC and greater ecological interest. WU; see previous section) with respect to five There was a non-significant interaction between remnant attributes: (1) habitat characteristics; habitat type and bird groups (P - 0.06; Table 54 PACIFIC CONSERVATION BIOLOGY

9 II) 4). The major contributor to this border-line II) a interaction was the decline in species richness ~ 8 of WU species from an average of 3.2 (fype 1) ~ U to none (fype 4). Treated as a one-way analysis ";: 7 II) of variance, the decline with respect to habitat CD type of WU species was significant (P - 0.02) "u 6 and there were significantly more of these

8-II) uncommon woodland species in Type 1 habitats C 5 than in Type 4. Z There were significant differences in species E 4 composition among habitat types (ANOSIM, 3 Table 5). Habitat Type 1 differed significantly from habitat Types 3 and 4. The Yellow and 2 Yellow-rumped Thornbills and Red-capped and Hooded Robins were more prevalent in the more open, degraded habitats with little shrub cover (Types 3 and 4), while the Striated o Pardalote and Crested Pigeon were associated Type 1 Type 2 Type 3 Type 4 with regenerating remnants with numerous habitat type eucalypts (fable 5).

Remnant area 9 b II) Based on remnant area, four size classes were II) 8 CD used: "very small" (2.5-5 ha, seven remnants); C ~ "small" (5-10 ha, six remnants); "medium" U 7 ";: (10-20 ha, eight remnants); and "large" (>20 ha, II) six remnants). There were no significant "uCD 6 differences in richness relative to remnant size CD (Table 4). On average, mean species richness Q. 5 II) per 1 ha transect was greatest in medium­ C sized remnants (14.6), with fewer species found Z 4 in the very small (mean = 12.9), small (13.2) E and large remnants (13.0) (Fig. 3b). There was 3 no significant interaction between bird groups and remnant area, and the generally increasing 2 richness of WU species with remnant size (Fig. 3b) also was non-significant. There were significant differences in species o composition among groups of different-sized very small small medium large remnants (fable 5). The only significant pair­ remnant area wise comparison was between the very small and large remnants. The major contributors to Fig. 3. (a) Histograms depicting mean species richness of the groups of birds (OP (open). OW (dense hatch). this difference were the Red-capped Robin, WC (stipple). WU (light hatch)) for the four types of Australian Magpie, Yellow and Yellow-rumped remnants determined by ordination, with habitat type Thornbills - all more prevalent in very small decreasing from Type I, or less-disturbed woodland, remnants - and the Brown Treecreeper, White­ to Type 4 (open degraded woodland). (b) Histograms plumed Honeyeater and Dusky Woodswallow, depicting mean species richness in remnants of different areas (see text). Values are means with which appeared to prefer the larger remnants standard error bars. (fable 5).

Table 4. Probabilities for main factors and interactions in repeated measures ANOVAs of species richness among groups of birds in relation to various classifications of remnants.

Probabilities

Between-factor Between-factor Bird groups Groups X Factor Habitat type (1-4) 0.723 <0.001 0.065 Remnant area 0.803 <0.001 0.185 Remnant shape 0.023 <0.001 0.002 Remnant isolation 0.780 <0.001 0.578 Remnant location 0.748 <0.001 0.278 WATSON, Mac NALLY and BENNETT: AVIFAUNA OF BULOKE WOODLANDS 55

Table 5. Analysis of similarities (ANOSIMs) and contributions to similarities within groups and dissimilarities among groups by bird species (SIMPERs) for five characteristics of remnant Buloke woodlands (only statistically significant - by using the method of Holme (1979) - group-wise comparisons are listed).

Comparison/group t Groups Bird Species t, §

Habitat type (0.156*) Type 1 v Type 3 0.442* Yellow Thornbill (-); Yellow-rumped Thornbill (-); Red-capped Robin (-); Striated Pardalote (+); Hooded Robin (-); Crested Pigeon (+); Chestnut-rumped Thornbill (-); Willie Wagtail (-) lype 1 v Type 4 0.640* Yellow Thornbill (-); Red-capped Robin (-); White-plumed HE (+); White-fronted HE (-); Yellow-rumped Thornbill (-); Striated Pardalote (+); Hooded Robin (-); Galah (+); Crested Pigeon (+); Black HE H Remnant area (0.138*) Very small v Large 0.376* Red-capped Robin (+); Brown Treecreeper (-); Yellow-rumped Thornbill (+); White-plumed HE (-); Dusky Woodswallow (-); Yellow Thornbill (+); Australian Magpie (+); Willie Wagtail (+); Red-rumped Parrot (-) Remnant shape (0.056) Isolation (0.080) Location (0.369*) West v Central 0.732* Yellow-rumped Thornbill (-); Yellow Thornbill (-); Striated Pardalote (+); Red­ capped Robin (-); White-fronted HE (-); Mistletoebird (-); Crested Pigeon (+); White-plumed HE (+); Variegated Fairy-wren (+); Galah (+) West v East 0.480* Yellow Thornbill (-); Crested Pigeon (+); White-Plumed HE (+); Spiny-cheeked HE (+); Varied Sitella (-); Variegated Fairy-wren (+); Australian Magpie (-); Striated Pardalote (+); Hooded Robin (-) Central v North 0.598* Brown Treecreeper (-); Galah (-); Dusky Woodswallow (-); Mistletoebird (+); Red­ rumped Parrot (-); White-plumed HE (-); White-fronted HE (+); Hooded Robin (-); Australian Magpie (+); Red-capped Robin (+) t figure in brackets denotes global test statistic, R; * denotes statistically significant; t HE denotes "honeyeater" (see Table 3 for species' names); § in comparisons: "+" indicates species more prevalent in first category while "-" denotes species more frequent in second category.

Remnant shape remnants; mean richness in transects of isolated Remnants were grouped into three classes remnants was 12.5 species, while the values based on shape: "regular" (remnants with S ~ 1.13, for the transects of < 8 ha and ~8 ha remnant eight remnants), "intermediate" (1.14 < S < 1.2, classes were both 13.8 (Fig. 4b, Table 4). There nine remnants) and "complex" (S > 1.2, ten was a non-significant interaction between remnants). Mean richness increased significantly with remnant-shape complexity, from a low of isolation and bird groups, with little evidence for differences between groups of birds (Fig. 4b). 10.6 in the transects in regular remnants to a The compositions of the avifaunas of transects high of 15.2 in the transects within complex remnants (Fig. 4a). However, only the difference of remnants surrounded by differing amounts between transects in regular- and complex­ of native vegetation were not significantly shaped remnants was significant. There was a different (Table 5). significant bird groups by remnant-shape Geographic location interaction (Table 4), which was due mainly to the depauperate we composition in transects of Remnants were grouped into four classes regularly shaped remnants (Fig. 4a; means: according to geographic location (see Fig. 1): regular 4.0, intermediate 8.1, and complex western (remnants 1-5); central (remnants 7.4). There were no significant differences in 6-14); eastern (remnants 15-22); and northern the composition of avifaunas of transects in (remnants 23-27) regions. There was little remnants of differing shape (Table 5). evidence of differences in mean richness among transects of remnants in the four regions, nor Remnant isolation for interactions between region and bird groups The 27 remnants were grouped into three (Fig. 5, Table 4). The apparently larger number classes based on isolation: "isolated" (six of uncommon woodland-dependent (WU) species remnants with no surrounding woodland, in the western region (Fig. 5) can be mainly excluding the remnant itself, within the attributed to a disproportionate representation 2 of this group (9 'species) in a transect of a single circumscribed area of 19.6 km ), "<8 ha" (13 remnant (remnant 1). remnants) and ~8 ha (8 remnants). There was no evidence for differences in mean richness Geographic position of remnants had a among transects of the isolation classes of pronounced effect on the composition of the 56 PACIFIC CONSERVATION BIOLOGY

tn 9 tn C1l E 8 U "t:; tn 7 "uC1l 8. 6 tn i 5 C1l E 4

3 3

2 2

o o regular intermediate complex western central eastern northern remnant shape geographic location

9 Fig. 5. Histograms depicting mean species richness of tn groups of birds for remnants based on geographic ~ 8 location. Western refers to remnants I to 5, central C to remnants 6--14, eastern to remnants 15-22, and ~ northern to remnants 23-27 (see Fig. I). Legend: see U 7 "t:; Figure 3. tn "~ 6 U remnants compared with a prevalence of White­ a-C1l tn fronted Honeyeaters, Mistletoebirds, Australian C Magpies and Red-capped Robins in the central m4 remnants (Table 5). E 3 DISCUSSION

2 Avifauna of Buloke woodlands The composition of the Buloke assemblage reflects the geographic position and climatic regime of these woodlands, which occur on o isolated < 8 ha ~ 8 ha the relatively dry plains inland of the Great Dividing Range, located between mesic eucalypt remnant isolation forests to the south and semi-arid shrublands Fig. 4. (a) Histograms depicting mean species richness of to the north-west. A number of the species groups of birds for remnants of different shape (see recorded are common and widespread in eucalypt text). (b) Histograms depicting mean species richness forests (e.g., Laughing Kookaburra, Black-faced for remnants of different degrees of isolation (see Cuckoo-shrike, Grey Shrike-thrush, Grey Fantail, text). Legend: see Figure 3. Southern Boobook, Rufous Whistler, Varied Sitella). The majority of the avifauna comprises avifaunas (Table 5). Remnants situated in the species that are widespread in dry open wood­ western part of the study area differed signifi­ lands across central Victoria (e.g., Jacky Winter, cantly from central and eastern remnants, while Restless Flycatcher, White-winged Chough, the central and northern sets of remnants also Eastern Rosella, Red-rumped Parrot, Crested differed (Table 5). In the west, the Crested Pigeon, Brown Treecreeper, White-browed Pigeon, White-plumed Honeyeater, Striated Babbler, White-plumed Honeyeater). However, Pardalote, Galah and Variegated Fairy-wren were a distinctive element in the Buloke avifauna is more widespread, while the central and eastern the relative abundance of a group of these remnants were characterized by the Yellow and species that characteristically occur with high Yellow-rumped Thornbills, Red-capped Robin frequency in woodlands dominated by She-oaks and White-fronted Honeyeater. The Brown and Callitris (Red-capped Robin, Hooded Robin, Treecreeper, Galah, Dusky Woodswallow, Red­ Yellow Thornbill, Chestnut-romped Thornbill, rumped Parrot, Hooded Robin and White­ Singing Honeyeater; Land Conservation Council plumed Honeyeater frequented northern 1987; Emison and Bren 1989). WATSON, Mac NALLY and BENNETT: AVIFAUNA OF BULOKE WOODLANDS 57

The distinctive nature of the Buloke avifauna litter between tussock grasses provide suitable is enhanced by species that typically occur in foraging substrates. The growth-form of Buloke, semi-arid and arid environments, such as the with canopy foliage extending down the trunk nearby Mallee region to the north-west (e.g., (d. distinct upper canopy for most eucalypts), Australian Ringneck, Blue Bonnet, Variegated provides sheltered perches for these ground Fairy-wren, Spiny-cheeked Honeyeater, Yellow­ foragers and foraging substrates for foliage plumed Honeyeater, Black-faced Woodswallow, feeders (e.g., Yellow and Chestnut-rumped Black Kite). This group also includes nomadic Thornbills). In contrast, several species distinctive species that occur in the study area at irregular of the structurally complex, regenerating intervals. The Black Honeyeater, while woodlands (Striated Pardalote, White-plumed historically rare in the region and Victoria Honeyeater) (Table 5) forage in eucalypt canopies generally, irrupted into the study area in the (Ford et al. 1986) and do not appear to favour summer of 1994/95 with more than 30 Buloke foliage. Eucalypts were most common individuals observed over two censusing periods in the regenerating woodlands (Types 1 and 2), (Watson 1995). Similarly, the White-fronted but scarce or absent in open Buloke stands Honeyeater is not a permanent resident, but was (Types 3 and 4). recorded at many transects within remnants Several studies in which vegetation has been when mistletoe was flowering (Watson 1997). found to be important in structuring bird community composition in remnant vegetation Habitat characteristics and avifauna! have not been restricted to a single habitat type, assemblages so that variation between communities can Four "types" of Buloke woodland were identified be attributed to differences in habitat type as from the habitat ordination procedure, and these well as habitat quality (e.g., Kitchener et al. were readily related to the disturbance regime 1982; Lynch and Whigham 1984; Freemark and and management history of remnants. Thus the Merriam 1986; Blake and Karr 1987). In the six remnants that were classed as Type 1 were current study, however, all of the selected not grazed by sheep and four of the remnants remnants were dominated by the same tree had been fenced for ~10 yr (D. M. Watson, species (i.e., Buloke) occurring at densities of at 1 unpubl. data). While other introduced herbivores least 50 stems ha- • Despite this relatively uniform were generally present, the absence of sheep habitat, subtle differences in the presence and grazing seemed to allow Buloke seedlings to abundance of eucalypt trees and in the structural establish and for a complex and species-rich complexity of the vegetation resulted in major understorey to develop. Type 2 remnants also differences in avian communities. were fenced but had been grazed by sheep There was no evidence that size of remnants relatively recently (as little as 2 yr prior to sampling) and there was still evidence of sheep had a significant impact on overall species richness of birds recorded from transects. presence (faecal pellets, bones). Nevertheless, However, the methodology used in this study the processes of regeneration had commenced differed from that used in many other studies and all remnants contained some shrubs, while where a significant relationship between species many supported the youngest cohort of Buloke indicating active regeneration. Those remnants richness and area was found (e.g., Kitchener et al. 1982; Ambuel and Temple 1983; Howe classified as Type 3 were all grazed at the 1984; Freemark and Merriam 1986; Bolger et al. time of sampling, had no shrubs and little 1991). We used a fixed rather than proportional grass cover, while Type 4 remnants could be sampling strategy in each remnant (see also considered overgrazed, with high densities of sheep and extremely simple habitat structure Forman et al. 1976; Barrett et al. 1994). Where proportional sampling is used (i.e., sampling with no understorey and a single-age cohort of Buloke. While other factors doubtless play a effort is proportional to, or increases with, remnant area), much greater time is spent role, it appears that management history ­ and greater area covered in a large remnant particularly grazing intensity by sheep - is a compared with a small one. Numbers of dominant force shaping habitat structure in individual animals detected (and hence numbers Buloke remnants. of species) are usually greater in larger or more The composition of bird assemblages differed intense surveys, so a positive association between among habitat types (Table 5). Species distinctive species richness and area is hardly surprising. of the more open, degraded woodlands (Type Mac Nally and Watson (1997) reported on how 4) included a number of birds that forage richness increased as a function of numbers of from the ground layer (Red-capped and Hooded transects in the four largest remnants considered Robins, Yellow-rumped Thornbill, Willie here, in which two transects were constructed Wagtail). The structurally simpler form of these and surveyed. Numbers of species "shared" woodlands does not appear to disadvantage between the two transects within remnants were these species. Open areas of bare ground and <40%. 58 PACIFIC CONSERVATION BIOLOGY

Fixed-sampling methods ask a different (Grey et at. 1997, 1998). The apparent unsuit­ question: does the spatial context of the ability of Buloke trees for foraging by Noisy sampling unit (in this case, the 1 ha transect) Miners, and the consequent scarcity of the determine the species richness associated with miners in this habitat, means that even small that unit? For example, does the richness of bird Buloke woodlands may have a high conservation species in a 1 ha transect set in a 3 ha remnant potential for the woodland avifauna. Moreover, differ from that in a similar transect set in a the occurrence of many species of concern (e.g., 467 ha remnant (see Westman 1983; Kelly the Hooded Robin, Red-capped Robin, Southern et at. 1989; Kohn and Walsh 1994)? Our analyses Whiteface, Bush Stone-curlew, Blue Bonnet, do not describe complete remnants but refer Gilbert's Whistler, Diamond Firetail, Painted to the surveyed transects within remnants. Button-quail) points to the conservation value of Therefore, deductions are pertinent to the these woodlands at a regional scale. contextual location of the transects and should be interpreted in only this way. When selecting study sites, all of the relatively intact Buloke-dominated remnants in the study area were considered. Thus, the six Implications for conservation remnants identified by the habitat ordination to Despite the extensive clearing and fragment­ be the least degraded (Type 1) represent the ation of Buloke woodlands, remnants in western only remaining, approximately intact Buloke Victoria support a surprisingly high richness vegetation, with other remnants degraded to of woodland birds and, consequently, have a various degrees. However, significant differences critical role in maintaining a distinctive wood­ in composition of assemblages in remnants in land avifauna in the rural landscape. Massive relation to vegetation type raise questions clearing of grassy woodlands across the plains about the components of habitat "quality" for of northern and western Victoria, including the avifauna. Intuitively, woodlands with an more than 90% of most types of dry woodland intact ground flora and active regeneration vegetation (Lunt and Bennett, in press), means would be expected to provide higher-quality that all remaining vegetation has potentially habitat for the avifauna, and such habitats have high conservation value for the maintenance of been shown to support the greatest richness the existing fauna, and also as a template for of reptiles and amphibians (Hadden and future restoration. Westbrooke 1996). However, the occurrence of species such as Hooded and Red-capped Robins While the Buloke avian assemblage has many in grazed, open woodlands indicates the value species in common with those of eucalypt of these habitats also, although their long-term woodlands across central and northern Victoria persistence depends upon adequate regeneration (e.g., Environment Conservation Council of Buloke trees. The most obvious and pervasive 1997; Bennett et at. 1998), at least two factors agent of vegetation degradation in the Wimmera emphasize the importance of the Buloke is grazing by sheep. By compacting soil ecosystem. First, the geographic location of and grazing on emerging seedlings, sheep this ecosystem in northwestern Victoria means effectively prevent whole ecosystems from that Buloke habitats are utilized by some species regenerating (Pigott 1983). Observations suggest commonly associated with mallee vegetation or that Buloke woodland can regenerate well other semi-arid environments. The location of once this grazing pressure is removed. There­ these woodlands between the Little Desert and fore, management of grazing is a key to the the Big Desert raises the possibility that the maintenance and restoration of conservation remnants may enhance connectivity through the values of these woodlands. rural landscape by functioning as "stepping stones" for birds reticent to cross large expanses of cleared land. This could include dispersal of ACKNOWLEDGEMENTS resident mallee species or movements of seasonal migrants to the mallee vegetation. We are most grateful to Lisa Morcom, formerly from the Stawell office of the Department of Second, the Buloke avifauna includes a large Natural Resources and Environment (DNRE), assemblage of small, insectivorous species for sharing her extensive knowledge of the (thornbills, robins, pardalotes, sitellas etc.). area, to Ian Voigt and Charles Hajek of the This rich representation in Buloke woodlands Catchment and Land Management team at may be due to the scarcity of Noisy Miners in the Horsham office, DNRE, for the use of a these remnants. Noisy Miners dominate the vehicle, and to the Flora and Fauna Branch, avifauna of small isolated eucalypt woodlands in DNRE, for providing financial assistance. rural environments across southeastern Australia Further financial assistance was provided by the (Loyn 1987; Barrett et at. 1994; Bennett et at. Australian Research Council and the Depart­ 1998) and aggressively exclude small insectivores ment of Biological Sciences, Monash University. when present in moderate to great numbers Dennis O'Dowd, Alan Lill and Vivienne Turner WATSON, Mac NALLY and BENNETT: AVIFAUNA OF BULOKE WOODLANDS 59 made important contributions to the study. We Forman, R. T T, Galli, A. E. and Leck, C. F., 1976. Forest thank also Richard Major, Gary Luck and an size and avian diversity in New Jersey woodlots with anonymous referee for greatly improving certain some land use implications. Oecol. 26: 1-8. aspects of the manuscript. Freemark, K. E. and Merriam, H. G., 1986. Importance of area and habitat heterogeneity to bird assemblages in REFERENCES temperate forest fragments. Bioi. Cons. 36: 115-41. Grey, M. Clarke, M. F. and Loyn, R. H., 1997. Initial Ambuel, R. and Temple, S. A, 1983. 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