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Home , Australian hobby, Cacatua, Cacatua (subgenus), Calyptorhynchus, Cockatiel, Crimson rosella

J. Raptor Res. 42(2):125–137 E 2008 The Raptor Research Foundation, Inc.

DIETARY SHIFTS BASED UPON PREY AVAILABILITY IN PEREGRINE AND AUSTRALIAN HOBBIES BREEDING NEAR ,

JERRY OLSEN1 AND ESTEBAN FUENTES Institute for Applied Ecology, University of Canberra, ACT, Australia 2601

DAVID M. Avian Science and Conservation Centre of McGill University, 21111 Lakeshore Road, Ste. Anne de Bellevue, Quebec, Canada H9X 3V9

A. B. ROSE2 The Australian Museum, 6 College Street, , 2010

DAVID JUDGE Australian Public Service Commission, 16 Furzer Street, Phillip ACT, Australia 2606

ABSTRACT.—We collected prey remains and pellets at 16 Peregrine (Falco peregrinus) nest territories (975 prey items from 152 collections) and one Australian (F. longipennis) territory (181 prey items from 39 collections) during four breeding seasons in two time periods: 1991–1992 and 2002–2003, a total of 60 peregrine nest-years and three hobby nest-years. By number, European Starlings (Sturnus vulgaris) were the main prey taken by both falcons in 1991–1992 and 2002–2003, but starlings made up a smaller percentage of the diet by number in the latter period, apparently because their numbers had declined in the wild. Although the geometric mean of prey weights and geometric mean weights were similar in the two time periods, both falcons compensated for the decline in European Starlings in the latter period by taking a greater variety of bird species, particularly small numbers of mostly native , rather than taking more of one or two other major prey species. Peregrines took 37 bird species in the latter period not found among their prey remains in the earlier period, and more individuals of some large species such as Gang-gang (Callocephalon fimbriatum), ( roseicapilla), and Rock Pigeons (Columba livia). Prolonged drought and competition from increasing numbers of Common Mynas (Acridotheres tristis) may have caused European Starlings to decline, but neither falcon species compensated by catching Common Mynas, even though they are of optimal prey size. Though both falcons, especially peregrines, took the more common bird species found in surveys in the Australian Capital Territory, both tended to avoid large or dangerous species, or agile species that foraged close to cover, such as Common Mynas.

KEY WORDS: ; Falco peregrinus; ; Falco longipennis; European Starling; Stur- nus vulgaris; Australia; Canberra; diet.

CAMBIOS EN LA DIETA CON BASE EN LA DISPONIBILIDAD DE PRESAS EN POBLACIONES REPRO- DUCTIVAS DE FALCO PEREGRINUS Y F. LONGIPENNIS EN LAS CERCANI´AS DE CANBERRA, AUSTRA- LIA

RESUMEN.—Recolectamos restos de presas y egagro´pilas en 16 territorios de anidacio´n de Falco peregrinus (975´tems ı presa provenientes de 152 recolecciones) y en un territorio de F. longipennis (181´tems ı prove- nientes de 39 recolecciones) durante cuatro e´pocas reproductivas en dos perı´odos: 1991–1992 y 2002–2003, correspondiendo a un total de 60 an˜os-nido para F. peregrinus y a tres an˜os-nido para F. longipennis.En nu´mero, Sturnus vulgaris fue la principal presa cazada por ambas especies de halcones tanto en el per´odoı

1 Email address: [email protected] 2 Present address: 61 Boundary Street, Forster, New South Wales, Australia 2428

125 126 OLSEN ET AL. VOL. 42, NO.2

de 1991–1992 como en el de 2002–2003, aunque el porcentaje contribuido a la dieta en te´rminos de nu´meros por S. vulgaris fue menor durante el segundo perı´odo debido a que aparentemente esta especie ha disminuido en los ambientes silvestres. A pesar de que la media geome´trica de los pesos de las presas y la media geome´trica de los pesos de las especies fueron similares en los dos periodos de tiempo, ambas especies de halcones compensaron la disminucio´n de S. vulgaris durante el segundo periodo cazando una mayor variedad de especies de aves. Lo hicieron cazando particularmente nu´meros pequen˜os de aves principalmente nativas, en vez de cazar mayor cantidad de una o dos especies de presas principales. Los individuos de F. peregrinus cazaron 37 especies de aves durante el segundo periodo, las cuales no se encontraron entre los restos de presas recolectados durante el primer periodo, y ma´s individuos de algunas especies de gran taman˜o como Callocephalon fimbriatum, Cacatua roseicapilla y Columba livia. La sequı´a prolongada y la competencia debido al nu´mero en aumento de Acridotheres tristis, pueden haber causado la disminucio´n de S. vulgaris, pero ninguna de las especies de halcones compenso´ cazando individuos de A. tristis a pesar de que e´stos tienen un taman˜o de presa o´ptimo. A pesar de que las dos especies de halcones, especialmente F. peregrinus, cazaron la especie de ave ma´s comu´n que se registra en los muestreos del territorio de la capital australiana, ambas especies tendieron a evitar especies grandes o peligrosas, y a especies a´giles que forrajean cerca de a´reas con mayor cobertura como A. tristis. [Traduccio´n del equipo editorial]

The Peregrine Falcon (Falco peregrinus) and Aus- ble (Olsen 1992). The same nest cliff may be occu- tralian Hobby (F. longipennis) breed throughout pied for many years and these cliffs are ideal sites Australia and are typical fast-flying, bird-catching for repeated annual collections of prey remains. falcons (Marchant and Higgins 1993). There have Peregrines near Canberra tend to lay eggs in Au- been no major studies of the diet of the Australian gust–September and fledge young in November– Hobby outside the Northern Territory, and no stud- December. Hobbies lay eggs about November and ies comparing the diet of Australian Hobbies and fledge young in January–February (Olsen 1992). Peregrine Falcons breeding in the same region, or Because there have been no long-term studies comparing their diets over time (Marchant and Hig- (.10 yr) of trends in diet for peregrines or hobbies gins 1993). The diet of the Peregrine Falcon has in Australia, our purpose in this study was to com- been well studied in rural and coastal parts of south- pare the diets of breeding Peregrine Falcons during eastern Australia, particularly in and in two time periods, 1991–1992 and 2002–2003. We open, lower-elevation parts of the Australian Capital also examined diet differences at one Australian Territory (ACT) and New South Wales (Pruett-Jones Hobby breeding territory in 1991, 2002, and 2003 et al. 1981, Olsen 1992, Marchant and Higgins 1993, in order to look for similar patterns in the prey Olsen et al. 1993, Olsen and Stevenson 1996, Rose captured, as both falcons eat primarily birds. 2001, Olsen and Tucker 2003, Olsen et al. 2004). They eat primarily flocking birds, particularly pi- METHODS geons, , and European Starlings (Sturnus vul- Study Area. We monitored 16 Peregrine Falcon garis), but also larger species up to the size of herons, territories, mostly within 60 km of Canberra. At low- ibises, and waterfowl (Debus 1998). The diet of the er elevations this area comprised urban areas, rivers, Australian Hobby has been less well studied (Cze- grazing land, open forest, tall woodland with dom- chura and Debus 1986, Debus et al. 1991, Marchant inants of Scribbly Gum ( rossii), Brittle and Higgins 1993, Aumann 2001). Outside the Gum (E. mannifera), Red Stringybark (E. macro- Northern Territory (Aumann 2001), there has been rhyncha), Blakely’s Red Gum (E. blakelyi) with Red only one diet study of a single nest of Australian Box (E. polyanthemos) and Yellow Box (E. melliodora), Hobbies (Debus et al. 1991, N 5 100 prey items). and Casuarina in river bottoms (NCDC 1988). Four Peregrines and hobbies are sympatric in Australia territories were in Namadgi National Park in the (Simpson and Day 1999). They compete for some southern half of the ACT, a 106 000-ha reserve cov- prey species, like European Starlings and small par- ering much of the Brindabella Range, with peaks up rots, but hobbies specialize in smaller birds such as to 1900 m. Namadgi National Park consists primar- larks, pipits, grassfinches, doves, and swallows, as well ily of tall wet sclerophyll forest and dense shrub as insectivorous bats and flying (Debus 1998). understory, with alpine woodland at the highest el- Near Canberra, there is a shortage of nest cliffs evations, and open, drier forest with open grassy for peregrines and pair densities are remarkably sta- valleys at the lower elevations (Taylor and COG JUNE 2008 DIETS OF PEREGRINES AND HOBBIES 127

1992, Olsen and Rehwinkel 1995). Eight of the low- one prey bird, because nestlings share prey items er-elevation cliff nests were above rivers and dams, and more than one species was often found in each and four were in deserted or semi-active quarries, all pellet. at approximately 500 m above sea level (asl); the We also included observations of kills and prey four nests on cliffs in Namadgi National Park were consumed in situ, if these items were not reflected at around 1000 m asl. in the prey remains and pellets collected. For exam- Collection of Prey Remains and Pellets. Pere- ple, if we saw a European Starling carried to the grine Falcons. Sixteen Peregrine Falcon nest cliffs brood, but did not find European Starling feathers were monitored during the 1991, 1992, 2002, and in the collection for that day or the following day, 2003 nesting periods (total 5 60 nest-years; 4 nest- we counted that European Starling as a prey item. years not sampled). We attempted to collect prey However, if we did find European Starling in the remains and pellets at the nest cliffs two or three pellets or prey remains for that day or the following times during the nestling period. However, logistics day, we did not include the observation. restricted our collections to once per season at the Estimation of Prey Weights. Calculating mean Namadgi National Park sites. We collected prey re- prey weight is one straightforward method of esti- mains and pellets from nest ledges and cliff tops mating raptor prey size, but the frequencies of prey above or within 5 m either side of the nest after weights for most raptor species do not follow nor- young hatched. mal distributions, and are usually skewed to one Australian Hobbies. Three hobby nests in the same side of the mean. An alternate measure, geometric territory were monitored in 1991, 2002 and 2003. mean prey weight (GMPW) has been used ( Jaksic´ The hobbies used the same nest in 2002 and 2003, and Braker 1983, Marti 1987). Geometric mean but a different one in 1991. We collected pellets and prey weight is calculated by summing the products prey remains after young hatched, approximately of the number of prey items in each category times twice per week during the three breeding seasons its loge-transformed weight and dividing it by the (incubation to post-fledging). total number of prey individuals in all categories. Measurement of Reproductive Success. Brood To determine the prey weight class most favored size and the number of fledged young were recorded by each raptor species during the two time periods, for each monitored territory. Peregrine and hobby we used two alternative measures: GMPW and geo- young were considered as fledged when they reached metric mean weight of prey species (GMSW). The banding age, ca. 4 wk. Successful and failed nests latter considers only the weights of each prey type, were included, and a successful territory was defined whereas the former considers these weights and the as one that raised at least one chick to banding age. number of captured in each category, thus Analysis of Pellets and Prey Remains. We ana- reflecting the differential contribution of each prey lyzed both regurgitated pellets and prey remains species. To avoid pseudoreplication, items that were to minimize potential biases in estimates of overall not determined to species (such as ‘‘unidentified diet and pooled the data for analysis (Collopy 1983, birds’’ or ‘‘unidentified ’’) were not includ- Marti 1987, Simmons et al. 1991, Seguin et al. ed in most GMPW and GMSW calculations, the only 1998). To identify prey species we compared feath- exception being when there was no species of that ers, bones, feet and beaks with those in private col- group identified. That is, if in any given year we lections and, when necessary, to skins at the Austra- found Crimson (Platycercus elegans), Eastern lian Museum, Sydney. Bones were identified by Rosella (P. eximius) and unidentified rosella, the morphological comparison with museum specimens latter was not included. If we found only unidenti- and a photographic key of humerus bones of fami- fied rosella, then it was included. To determine dif- lies of nonpasserine birds. (The humerus is typically ferences in diet between the two falcon species, and the strongest bone, because it bears the stress of between the two time periods, we scored the most flight, and is often the only unbroken bone in per- common prey species taken. Because peregrines egrine prey remains; W. Boles pers. comm.). To and hobbies feed predominantly on birds, and we estimate the minimum number of prey items were most interested in the differences in bird spe- (MNI) in the pooled sample of pellets and prey cies taken, two analyses for both prey size estimators remains we counted bones, feet, beaks, primary were undertaken: (1) all prey, and (2) avian prey and tail feathers as in Olsen and Tucker (2003). only (to eliminate the confounding influences of We did not assume that one pellet represented the few mammal and species taken). 128 OLSEN ET AL. VOL. 42, NO.2

Table 1. Main Prey Items (MPIs) by capture rank in the first time period 1991–1992 and mass of species captured by Australian Hobbies and Peregrine Falcons. Scientific names of prey species in Appendix 1.

FALCON SPECIES PREY SPECIES CAPTURE RANK SPECIES MASS (g) Australian Hobby European Starling 1 75 Red-eyed 2 2 House Sparrow 3 27 4 135 5 28 6 106 Peregrine Falcon European Starling 1 75 2 335 Eastern Rosella 3 106 Rock Pigeon 4 308 Silver Gull 5 303

Estimation of Prey Abundance. To compare (1999) and applied a Chi-square test following the changes in the diet of the two falcons during Haber method. Post hoc contrasts were subjected to a the two time periods against a broad measure of Bonferroni correction. The Shannon Diversity In- bird prey abundance in the area, we used the dex (Shannon and Weaver 1949) was used to esti- Canberra Ornithologists Group (COG) Garden mate the dietary diversity: Bird Surveys conducted from 1991–2003 (COG 0 H ~{Spilog pi, 2003, 2005). These indices of abundance are done mainly by amateurs throughout the Australian where pi represents the proportional contribution Capital Territory using a system of weekly tallies by number of each species in the sample. We used developed by Henry Nix (Veerman 2003). The the Pianka Index (Pianka 1973) to explore prey surveys did not provide exact data for prey avail- overlap between peregrines and hobbies, and be- able at each falcon nest, but did give a ranking for tween prey caught by the falcons in the two differ- all bird species found in the area where falcons ent time periods: nested. Rankings of bird prey in the falcons’ diets . ~ 2 2 1=2 were compared to rankings found in COG surveys. O Spiqi Spi Sqi One limitation of this comparison was that surveys of prey bird species close to peregrine nests may All analysis and calculations were performed using not provide valid measures of prey availability, be- SAS 8.0 (SAS Institute Inc. Cary, NC) and Excel cause peregrines are known to forage great dis- 2000 (Microsoft Corporation). tances from their nests (up to 79 km; Henderson and Craig 1997). RESULTS Statistical Analysis. For the purposes of statistical Reproductive Success. For peregrines, the mean analysis, we used the Main Prey Items (MPIs) con- number of young fledged per territory, was slightly sumed by each falcon species. An MPI was any spe- lower during the second time period (1.47 in 1991– cies that contributed 5% or more, by number, to the 1992 and 1.27 in 2002–2003 (nonsignificant, x2 5 overall diet in at least one of the time periods (Ta- 1.38; df 5 3, P 5 0.71); young per successful terri- ble 1). Other prey items contributing ,5% by num- tory was 2.24 in 1991–1992 and 2.00 in 2002–2003). ber to the total diet were grouped under the follow- For hobbies, the mean young per territory was 3.0 in ing categories: other birds, mammals, and insects 1991 and 2.5 in 2002–2003. (Appendices 1 and 2). Diet. A total of 975 prey items were obtained in To contrast the differences in the proportion of 152 collections of prey remains and pellets at the prey items between 1991–1992 and 2002–2003 we peregrine cliff nests, and 181 prey items in 39 col- used a Chi-square analysis on contingency tables. lections from the hobby nest site. In both periods, For differences between each prey class, we subdi- peregrines and hobbies caught predominantly birds vided the contingency tables as suggested by Zar (Appendices 1 and 2). In the 2002–2003 period, JUNE 2008 DIETS OF PEREGRINES AND HOBBIES 129

Table 2. Species taken by Australian Hobbies and Peregrine Falcons in the region of Canberra, Australia, in both study periods and in each period exclusively. Scientific names of prey species in Appendix 1.

NO.SPECIES TAKEN IN EACH PREY CATEGORY

FALCON EXOTIC SPECIES YEARS OF STUDY NATIVE BIRDS BIRDS MAMMALS INSECTS TOTAL Peregrine In 1991–92 and 2002–03 16 2 1 0 19 Falcon In 1991–92 but not in 2002–03 8 0 0 0 8 In 2002–03 but not in 1991–92 37 2 1 4 44 Total 1991–92 24 2 1 1 28 Total 2002–03 53 4 2 4 63 Australian In 1991–92 and 2002–03 4 2 0 2 8 Hobby In 1991–92 but not in 2002–03 6 0 0 2 8 In 2002–03 but not in 1991–92 11 0 1 4 16 Total 1991–92 10 2 0 4 16 Total 2002–03 15 1 2 7 25

both falcon species captured more species than in Peregrine Falcons. The percentage contribution of the 1991–1992 period and both captured propor- each of the MPIs to the diet of the Peregrine Falcon tionally more species that were not taken in 1991– differed between the two periods of time (x2 5 1992 (Table 2 ). 78.21, df 5 7, P , 0.001; Fig. 1). These differences

Figure 1. Comparative contribution of the main prey items (MPIs) by number to the diet of the Peregrine Falcon in the Canberra Region, 1991–1992 and 2002–2003. 130 OLSEN ET AL. VOL. 42, NO.2

Figure 2. Comparative contribution of the main prey items (MPIs) by number to the diet of the Australian Hobby in the Canberra Region, 1991–1992 and 2002–2003.

were mainly the result of a decrease in Silver Gulls diverse in the 2002–2003 survey (H991–92 5 0.99 vs. (Larus novaehollandiae; 7.5% of total diet in 1991– H902–03 5 1.25). 1992 vs. 0.2% in 2002–2003; x2 5 35.25, df 5 1, P , Australian Hobbies. As with Peregrine Falcons, 0.001) and European Starlings (36.1 vs. 23.2%; x2 5 measures of the MPIs showed that the diet of the 19.15, df 5 1, P , 0.001), as well as an increase in Australian Hobbies differed between the two peri- the number of other birds (20.5 vs. 33.6%; x2 5 ods (x2 5 31.68, df 5 8, P , 0.001; Fig. 2, Appen- 20.52, df 5 1, P , 0.001). dix 2). There was a decrease in the number of Eu- In the latter period peregrines took fewer exotic ropean Starlings taken compared to the previous birds, especially European Starlings, and increased survey (32.6 vs. 15.1%; x2 5 7.71, df 5 1, P , numbers of native species (Fig. 1, Appendix 1, Ta- 0.005), coupled with an increase in other birds ble 2). Though the 2002–2003 sample was only (12.6% vs. 41.9%; x2 5 19.21, df 5 1, P , 0.0001; slightly larger than the 1991 sample (521 vs. 454 Fig. 2). Similarly, the proportion of exotic species in items), more than two times the number of native their diet, mainly European Starlings, decreased bird species (53 vs. 24) were recorded. Considering and the numbers of native species increased (Ta- all groups, there were more than twice as many spe- ble 2, Appendix 2). Nine more species, and five cies in the 2002–2003 samples than in the 1991– (50%) more species of native birds, were taken in 1992 samples (i.e., 63 species compared with 28; the 2002–2003 period. All other categories showed Table 2). In 2002–2003, peregrines caught 37 bird no significant changes. Though there was a slightly species that were not recorded in the 1991–1992 smaller sample size in the 2002–2003 period, hob- samples, but only eight bird species taken in bies took 16 species in the 2002–2003 period that 1991–1992 were not found in the 2002–2003 sam- were not recorded in the 1991 sample, but they took ples (Table 2). The diet of the peregrines was more only eight species in the 1991 period that were not JUNE 2008 DIETS OF PEREGRINES AND HOBBIES 131

Table 3. Geometric Mean of Prey Weights (GMPW), Geometric Mean of Prey Species Weights (GMSW) in grams and percentage of large and small prey taken by Peregrine Falcons and Australian Hobbies near Canberra, Australia. Data are presented as mean 6SE.

PEREGRINE FALCON AUSTRALIAN HOBBY

MEASURE OF PREY MASS 1991–1992 2002–2003 1991–1992 2002–2003 All Prey GMPW 140.1 6 1.0 132.1 6 1.0 22.1 6 1.2 21.9 6 1.2 GMSW 119.6 6 1.3 82.6 6 1.2 18.3 6 1.4 17.7 6 1.3 Avian Prey GMPW 140.3 6 1.0 143.2 6 1.0 55.8 6 1.0 46.84 6 1.0 GMSW 120.2 6 1.2 102.3 6 1.2 43.0 6 1.3 41.8 6 1.2 present in the 2002–2003 samples (Table 2). The banded racing pigeons so these were not compared diet of the hobbies was also more diverse in 2002– to COG estimates of feral Rock Pigeons. 2003 (H951.31) than in 1991 (H951.02). When the relative abundances of the avian prey Both the Australian Hobbies and Peregrine Fal- species were considered (Table 4), Peregrine Fal- cons took a significantly higher proportion of native cons took common species like Silver Gulls, Rock species in 2002–2003 compared to 1991–1992 Pigeons, Galahs, Crimson Rosellas, Eastern Rosellas (x2 54.4, df 5 1, P , 0.05; x2 55.8, df 5 1, P , and Red-rumped Parrots (Psephotus haematonotus)in 0.05, respectively). Pianka’s Index showed a 92.1% both time periods, but tended to avoid other com- overlap in diet for peregrines 1991–1992 vs. 2002– mon species like Sulphur-Crested Cockatoos (Caca- 2003. Hobbies showed a 67.2% overlap between the tua galerita), Australian Magpies (Gymnorhina tibi- two periods. In 1991–1992 peregrines and hobbies cen), and House Sparrows (Passer domesticus). showed an 82% overlap in diet, and in 2002–2003 Hobbies avoided these common large prey species, the overlap was 66.1%. and captured common smaller ones like House Prey Weights. The GMPW and GMSW for avian Sparrows and Crimson Rosellas. Neither falcon took prey were different for the two falcon species (Ta- Common Mynas (Acridotheres tristis) in either period ble 3). Hobbies captured avian prey 33–40% smaller despite their increase in numbers in the COG sur- by mass than that taken by peregrines. When all veys (x2 5 99.6, P , 0.001). prey was considered, including insects, the hobbies captured animals that were 16.2–17.8% of the mass DISCUSSION of that taken by peregrines. The productivity of Peregrine Falcons was similar There were no significant differences in prey size to that found in other Australian studies (Marchant (GMPW and GMSW) between the two periods for and Higgins 1993). Near Canberra, on average, per- either falcon species for both avian and all prey (t - egrines fledged a mean 1.4 young per territory, and test; all P . 0.05). 2.14 per successful pair (Olsen 1992). These means Comparison to Measures of Avian Prey Abun- are greater than those reported for Victoria (1.1 dance. Some changes in the falcon diets from young per territory and 2.0 young per successful 1991–1992 to 2002–2003 were reflected in data from pair; Pruett-Jones et al. 1981). Canberra Ornithologists Group Garden Bird Sur- Overall, the diets were similar to those previously veys. European Starlings and Silver Gulls decreased reported for peregrines in Australia (Pruett-Jones et as peregrine prey (Fig. 1) and decreased in the al. 1981, Olsen 1992, Marchant and Higgins 1993, COG surveys (x2 5 1 6132.7, P , 0.001 and Olsen et al. 1993, Rose 2001, Olsen and Tucker 1447.8, P , 0.001, respectively). Crested Pigeons 2003, Olsen et al. 2004) and hobbies (Czechura (Ocyphaps lophotes), Gang-gang Cockatoos (Calloceph- and Debus 1986, Debus et al. 1991, Marchant and alon fimbriatum), and Galahs (Cacatua roseicapilla) Higgins 1993, Debus 1998, Aumann 2001). The size increased as peregrine prey, and in the COG surveys of avian prey (based on GMPW and GMSW) of hob- (x2 5 4323.1, P , 0.001, 1379.9, P , 0.001, and bies was approximately one-third that of peregrines 504.5, P , 0.001, respectively). Almost all of the (Table 3), reflecting the difference in the two spe- Rock Pigeons found as prey in peregrine nests were cies’ masses, as local hobbies are ca. 33% of the 132 OLSEN ET AL. VOL. 42, NO.2

Table 4. Ten most abundant species in COG surveys in 1991–1992 and 2002–2003 (from Veerman 2003, COG 2003, 2005), number of each species captured by Australian Hobbies and Peregrine Falcons in the two time periods, and the species capture rank (in parentheses). Scientific names of prey species in Appendix 1.

NO.CAPTURED BY FALCONS (RANK)

MOST ABUNDANT SPECIES IN COG SURVEY COG ABUNDANCE RANK AUSTRALIAN HOBBIES PEREGRINE FALCONS 1991–1992 European Starling 1 31 (1) 164 (1) House Sparrow 2 8 (2) 0 3 0 0 Galah 4 0 70 (2) Pied 5 0 3 (10) Sulphur-crested 6 0 0 7 0 0 Crimson Rosella 8 7 (3) 22 (6) Australian Magpie 9 0 5 (8) Yellow-faced Honeyeater 10 0 0 2002–2003 Galah 1 0 101 (2) Sulphur-crested Cockatoo 2 0 2 (14) Silvereye 3 1 (6) 3 (13) Common Myna 4 0 0 European Starling 5 13 (1) 121 (1) 6 0 1 (15) House Sparrow 7 7 (2) 0 Crimson Rosella 8 0 22 (5) Australian Magpie 9 0 4 (12) 10 0 10 (8) mass of local peregrines ( J. Olsen unpubl. data). change was in the category of native birds (Table 2). This was not the case when all prey were considered, In the second time period, peregrines took more because hobbies caught large numbers of insects. than 100% more bird species and hobbies took For all prey, hobby GMPW and GMSW were about 50% more bird species. The decline in the number 16.2% and 17.8% of that of peregrine prey. of European Starlings in the falcons’ diet and the Peregrines often capture prey that is most abun- increase in native species may be a functional re- dant, and European Starlings were important in the sponse to the decline of this exotic species, and diet in both time periods. Starlings are also impor- relative abundance of similar-sized native species tant prey for peregrines elsewhere in the world (in in the area. Europe [Ratcliffe 1993], and North America [White Despite the lack of significant differences be- et al. 2002]), and important for hobbies in this tween the two time periods in GMPW and GMSW, study. When starlings were abundant in 1991– there were some interesting patterns. With the re- 1992, there was a greater overlap in peregrine/hob- duction of European Starlings in the latter period, by diet (Pianka’s Index 82% vs. 66.1%) than in peregrines caught a larger number of species and 2002–2003 when they were less available. Peregrines many of these were similar in mass or smaller than may have taken more larger birds such as Galahs, European Starlings. However, they captured more Rock Pigeons, and Gang-gang Cockatoos that hob- individuals from a few species of large birds, includ- bies could not capture. Both falcon species ap- ing Galahs, and this offset any potential difference peared to have a more diverse diet during the sec- in GMPW and GMSW between the two periods (Ta- ond study period when European Starlings were less ble 3). That is, peregrines seem to have adjusted to abundant. To compensate, they captured small the decrease in European Starlings in the latter pe- numbers of a large number of native species. Both riod by capturing a larger number of smaller spe- took more species in 2002–2003, but the greatest cies, which decreased the overall prey size, but offset JUNE 2008 DIETS OF PEREGRINES AND HOBBIES 133 this with greater numbers of a few larger species, like ed some species, both large and small, but they tended Rock Pigeons, Galahs, and Gang-gang Cockatoos. to avoid agile species that foraged close to cover. Veerman (2003) related the decline in European Postscript—In the four breeding seasons after this Starlings over the last 13 yr in the Australian Capital study we found only one Common Myna in prey re- Territory to loss of foraging , and a sharp mains at these peregrine sites, but several Common increase in the population of Common Mynas, Mynas in prey remains from hobby nest sites. which displaced them. The decrease in European ACKNOWLEDGMENTS Starling numbers may be related to an extended drought in the Southern Tablelands of New South Thanks to Alison Rowell for analysis of some of the 1991– Wales (M. Clayton pers. comm.). Common Mynas 1992 prey, to Susan Trost, David Mallinson, Greg Hayes, Frank Barnes, Paul Higginbotham, Brett McNamara, Don would have been available to adults at all nests in Fletcher, Jeff and Angie Young, Simon Tozer, Darren Rosso, this study but were not taken. A larger study of the Michael McGonicky, Bernard Morris, Christie Gould, Tony diets of raptors near Canberra, ( J. Olsen and A. Bell, Monica Muranyi, and the ACT Parks and Conservation Rose unpubl. data) found no evidence that pere- Service for assistance in the field. We gratefully acknowl- edge the Australian Museum for enabling ABR to compare grines or hobbies captured Common Mynas. To prey samples with reference material, especially Walter our knowledge, this species has not been recorded Boles who developed the photographic key for humerus as prey for either of these falcon species in the pub- identification, and John Disney and Ian McAllan for assis- lished literature, possibly because mynas forage tance with identifying feathers and bones. Thanks also to Arthur Georges, Jim Hone, Stephen Saare, Richard Norris, near cover and are too agile. Terry Dennis, Jeff Jolly, Stephen Debus, and Les Boyd for Peregrines and hobbies are almost exclusively logistic support and advice. Special thanks to Martin Butter- bird-catchers, and these pairs tended to take some field, David Rosalky, and Barbara Allan for their assistance of the most abundant species in the Canberra area with the COG data. EF was supported by Graduate Scholar- ships from CONACYT in Mexico and the University of Can- (Appendices 1, 2, Table 4). The COG data are only berra. Dick Dekker, Robert DeCandido, Cheryl Dykstra and a broad measure of abundance; however, some spe- an anonymous referee considerably improved the manu- cies appeared to be selected more often than oth- script. The study methodology was approved by the Univer- ers. Peregrines regularly took some large species sity of Canberra Ethics Committee Registration like Galahs, Magpies, and Silver Gulls, but hobbies Number CEAE 02/10. in this study took none of them. European Starlings, LITERATURE CITED Eastern Rosellas, and Red-rumped Parrots often for- AUMANN, T. 2001. An intraspecific and interspecific com- age in the open in flocks, and were specifically tar- parison of raptor diets in the south-west of the North- geted by both falcons. Crimson Rosellas often for- ern Territory, Australia. Wildl. Res. 28:379–393. age in woodlands and were targeted less often by CANBERRA ORNITHOLOGISTS GROUP. 2003. Annual bird re- peregrines but were targeted by hobbies. House port: 1 July 2002 to 30 June 2003. Canberra Bird Notes Sparrows in the COG surveys were the second most 28:125–188. abundant species in 1991–1992 and the seventh ———. 2005. Annual bird report: 1 July 2003 to 30 June most abundant species in 2002–2003 (COG 2003, 2004. Canberra Bird Notes 30:1–64. 2005); hobbies in this study took them, but pere- COLLOPY, M.W. 1983. A comparison of direct observations and collections of prey remains in determining the diet grines did not, and they have not been taken by of Golden Eagles. J. Wildl. Manage. 47:360–368. peregrines during our long-term monitoring CZECHURA,G.V.AND S.F.S. DEBUS. 1986. The Australian (1990–2004, N 5 2035 prey items, J. Olsen and A. Hobby Falco longipennis:areview.Aust. Bird Watcher Rose unpubl. data). 11:185–207. Conclusions. GMPW and GMSW for avian prey DEBUS, S. 1998. The birds of prey of Australia: a field guide. species reflected the different weights of the two Oxford Univ. Press, , Australia. falcons, and were consistent for both falcons in each ———, A.J. LEY,S.TREMONT, AND R. TREMONT. 1991. study period. By number, European Starlings were Breeding behaviour and diet of the Australian Hobby the main prey taken by Australian Hobbies and Per- Falco longipennis in northern New South Wales. Aust. egrine Falcons in the two time periods, but by 2002– Bird Watcher 14:123–137. HENDERSON, J.H. AND G.R. CRAIG. 1997. Wide ranging by 2003 the percentage of European Starlings had de- nesting Peregrine Falcons (Falco peregrinus) determined creased. The falcons compensated for the decline of by radiotelemetry. J. Raptor Res. 31:333–338. European Starlings by taking small numbers of a wider HIGGINS, P.J. 1999. Handbook of Australian, New Zealand variety of native species, many that were not taken in and Antarctic birds, Vol. 4. Oxford Univ. Press, Mel- the earlier time period. Peregrines specifically target- bourne, Australia. 134 OLSEN ET AL. VOL. 42, NO.2

——— AND S.J.J.F. DAVIES. 1996. Handbook of Australian, OLSEN, P., J. OLSEN, AND I. MASON. 1993. Breeding and non- New Zealand and Antarctic birds, Vol. 3. Oxford Univ. breeding season diet of the Peregrine Falcon Falco pere- Press, Melbourne, Australia. grinus near Canberra, prey selection, and the relation- ——— AND J.M. PETER. 2002. Handbook of Australian, ship between diet and reproductive success. Pages 55–77 New Zealand and Antarctic birds, Vol. 6. Oxford Univ. in P.D. Olsen [ED.], Australian raptor studies. Australa- Press, Melbourne, Australia. sian Raptor Association, RAOU, Melbourne, Australia. ———, ———, AND W.K. STEELE. 2001. Handbook of Aus- PIANKA, E.R. 1973. The structure of lizard communities. tralian, New Zealand and Antarctic birds, Vol. 5. Ox- Annu. Rev. Ecol. Syst. 4:53–74. ford Univ. Press, Melbourne, Australia. PRUETT-JONES, S.G., C.M. WHITE, AND W.R. DEVINE. 1981. JAKSIC´ , F.M. AND H.E. BRAKER. 1983. Food-niche relation- Breeding of the Peregrine Falcon in Victoria, Australia. ships and guild structure of diurnal birds of prey: com- Emu 80:253–267. petition versus opportunism. Can. J. Zool. 61:2230–2241. RATCLIFFE, D. 1993. The Peregrine Falcon, Second Ed. T. MAGRATH, M.L., L. BROUWER,A.VAN PETERSEN., M.L. BERG, and A.D. Poyser, London, U.K. AND J. KOMDEUR. 2004. Breeding behaviour and ecology ROSE, A.B. 2001. Supplementary records of the food of of the sexually size-dimorphic Brown Songlark, Cinclo- some terrestrial non- in New South Wales. rhamphus cruralis. Aust. J. Zool. 51:429–441. Aust. Bird Watcher 19:60–68. MARCHANT,S.AND P.J. HIGGINS. 1990a. Handbook of Aus- SEGUIN, J.F., P. BAYLE,J.C.THIBAULT,J.TORRE, AND J.D. tralian, New Zealand and Antarctic birds, Vol. 1a. Ox- VIGNE. 1998. A comparison of methods to evaluate ford Univ. Press, Melbourne, Australia. the diet of the Golden Eagle in Corsica. J. Raptor Res. ——— AND ———. 1990b. Handbook of Australian, New 32:314–318. Zealand and Antarctic birds, Vol. 1b. Oxford Univ. SHARP, A., L. GIBSON,M.NORTON,B.RYAN,A.MARKS, AND Press, Melbourne, Australia. L. SEMERARO. 2002. The breeding-season diet of the ——— AND ———. 1993. Handbook of Australian, New Wedge-tailed Eagle (Aquila audax)inwesternNew Zealand and Antarctic birds: Vol. 2. Oxford Univ. South Wales and the influence of rabbit calicivirus dis- Press, Melbourne, Australia. ease. Wildl. Res. 29:175–184. MARTI, C.D. 1987. Raptor food habits studies. Pages 67–80 SHANNON, C.E. AND W. WEAVER. 1949. The mathematical in B.A. Giron Pendleton, B.A. Millsap, K.W. Cline, and theory of communication. Univ. of Illinois Press, Ur- D.M. Bird [EDS.], Raptor Management Techniques bana, IL U.S.A. Manual. Natl. Wildl. Fed., Washington, DC U.S.A. SIMMONS, R.E., D.M. AVERY, AND G. AVERY. 1991. Biases in MCDONALD, P.G., P.D. OLSEN, AND D.J. BAKER-GABB. 2003. diets determined from pellets and remains: correction Territory fidelity, reproductive success and prey choice factor for a mammal- and bird-eating raptor. J. Raptor in the Brown Falcon, Falco berigora: a flexible bet-hedg- Res. 25:63–67. er? Aust. J. Zool. 51:399–414. SIMPSON,K.AND N. DAY. 1999. Field guide to the birds of NATIONAL CAPITAL DEVELOPMENT COMMISSION. 1988. Sites Australia, Sixth Ed. Penguin, Ringwood, Victoria, Aus- of significance in the ACT Volume 2: inner Canberra, tralia. Technical Paper 56. Canberra, Australian Capital Ter- STRAHAN,R.[ED.]. 1988. The complete book of Australian ritory, Australia. mammals. William Collins, Auckland, Australia. OLSEN, J. 1992. Raptors in Namadgi, Canberra Nature TAYLOR,M.AND CANBERRA ORNITHOLOGISTS GROUP. 1992. Parks, the Murrumbidgee River Corridor and Googong Birds of the Australian Capital Territory, an atlas. Can- Foreshore. Report to ACT Parks and Conservation Ser- berra Ornithologists Group and the National Capital vice, Canberra, Australia. Planning Authority, Canberra, Australia. ———, S. DEBUS, A.B. ROSE, AND G. HAYES. 2004. Breeding VEERMAN, P. 2003. Canberra birds: a report on the first success, cliff characteristics and diet of Peregrine Fal- 21 years of the Garden Bird Survey. Canberra Ornithol- cons in Namadgi National Park, Australian Capital Ter- ogists Group, Canberra, Australia. ritory. 29:33–37. WHITE, C.M., N.J. CLUM, T.J. CADE, AND W.G. HUNT. 2002. ——— AND R. REHWINKEL. 1995. Peregrines and Powerful Peregrine Falcon (Falco peregrinus). In A. Poole and F. Owls in Namadgi and Tidbinbilla. Report for the Na- Gill [EDS.], The birds of North America, No. 660. The tional Estates Grants Program, Canberra, Australia. Academy of Natural Sciences, Philadelphia, PA and the ——— AND E. STEVENSON. 1996. Female Peregrine Falcon American Ornithologists’ Union, Washington, DC replaces an incubating female and raises her young. U.S.A. Aust. Bird Watcher 16:205–210. ZAR, J.H. 1999. Biostatistical analysis. Prentice Hall, Engle- ——— AND A.D. TUCKER. 2003. A brood-size manipulation wood Cliffs, NJ U.S.A. experiment with Peregrine Falcons, Falco peregrinus, near Canberra. Emu 103:127–132. Received 16 April 2007; accepted 25 January 2008 JUNE 2008 DIETS OF PEREGRINES AND HOBBIES 135

Appendix 1. Number of items, percentage by number and biomass, and individual masses for the prey captured by the Peregrine Falcon in 1991–1992 and 2002–2003.

PREY SPECIES 1991–1992 2002–2003

PERCENT PERCENT PERCENT PERCENT MASS COMMON NAME SCIENTIFIC NAME N ITEMS BIOMASS N ITEMS BIOMASS (g)1

MAMMALS Gould’s Wattled Bat Chalinolobus gouldii 0 1 0.2 0.01 14 European Rabbit Oryctolagus cuniculus 1 0.2 1.8 2 0.4 2.9 1500 European Rabbit juvenile 1 0.2 0.6 0 500 Mammal subtotals 2 0.4 2.3 3 0.6 2.9

BIRDS Painted Button quail Turnix varia 1 0.2 0.1 0 93 Pacific Black Duck Anas superciliosa 0 1 0.2 1.0 1036 Grey Teal Anas gracilis 0 2 0.4 0.9 475 Maned Duck Chenonetta jubata 4 0.9 3.8 4 0.8 3.1 808 Cattle Egret Bulbulcus ibis 0 1 0.2 0.3 332 Little Egret Egretta garzetta 0 2 0.4 0.6 300 Nankeen Night Heron Nycticorax caledonicus 0 1 0.2 0.8 800 Masked Lapwing Vanellus miles 1 0.2 0.4 0 360 Pacific Golden-Plover Pluvialis fulva 0 1 0.2 0.1 148 Silver Gull Larus novaehollandiae 34 7.5 12.1 1 0.2 0.3 303 Falco cenchroides 0 1 0.2 0.2 168 Rock Pigeon Columba livia 42 9.3 15.2 65 12.5 19.5 308 Rock Pigeon juvenile 0 1 0.2 0.3 290 Bar-shouldered Dove Geopelia humeralis 0 1 0.2 0.1 130 Peaceful Dove Geopelia placida 0 1 0.2 ,0.05 44 Common Bronzewing Phaps chalcoptera 4 0.9 1.6 1 0.2 0.3 331 Brush Bronzewing Phaps elegans 0 1 0.2 0.2 184 Crested Pigeon Ocyphaps lophotes 0 7 1.3 1.4 205 Yellow-tailed Black-Cockatoo funereus 0 1 0.2 0.7 766 Gang-gang Cockatoo Callocephalon fimbriatum 0 17 3.3 4.2 251 Galah Cacatua roseicapilla 70 15.4 27.6 101 19.4 32.9 335 Cacatua sanguinea 0 1 0.2 0.5 500 Sulphur-crested Cockatoo Cacatua galerita 0 2 0.4 1.6 804 Little Lorikeet Glossopsitta pusilla 0 2 0.4 0.1 50 Australian King- Alisterus scapularis 2 0.4 0.5 1 0.2 0.2 211 Nymphicus hollandicus 1 0.2 0.1 0 91 Budgerigar Melopsittacus undulatus 1 0.2 ,0.05 0 28 Crimson Rosella Platycercus elegans 22 4.8 3.5 19 3.6 2.5 135 Crimson Rosella juvenile 0 3 0.6 0.4 131 Eastern Rosella Platycercus eximius 46 10.1 5.7 41 7.9 4.2 106 Eastern Rosella juvenile 2 0.4 0.2 2 0.4 0.2 100 Unidentified rosellas Platycercus sp. 3 0.7 0.4 1 0.2 0.1 115 Ringneck Parrot Barnardius zonarius 0 1 0.2 0.2 157 Red-rumped Parrot Psephotus haematonotus 8 1.8 0.6 12 2.3 0.7 61 Bluebonnet Northiella haematogaster 1 0.2 0.1 1 0.2 0.1 90 Unidentified parrots Fam. Psittacidae 2 0.4 0.5 1 0.2 0.2 230 Pallid Cuckoo Cuculus pallidus 0 2 0.4 0.2 83 Black-eared Cuckoo Chrysococcyx osculans 0 1 0.2 ,0.05 30 Sacred Kingfisher Todiramphus sanctus 0 2 0.4 0.1 40 Asian Dollarbird Eurystomus orientalis 0 1 0.2 0.1 150 Spotted Pardalotus punctatus 0 2 0.4 ,0.05 9 136 OLSEN ET AL. VOL. 42, NO.2

Appendix 1. Continued.

PREY SPECIES 1991–1992 2002–2003

PERCENT PERCENT PERCENT PERCENT MASS COMMON NAME SCIENTIFIC NAME N ITEMS BIOMASS N ITEMS BIOMASS (g)1 Pardalotus striatus 0 3 0.6 ,0.05 12 Yellow-rumped Thornbill Acanthiza chrysorrhoa 0 1 0.2 ,0.05 9 Red Wattlebird Anthochaera carunculata 2 0.4 0.3 10 1.9 1.1 108 Noisy Miner Manorina melanocephala 0 5 1.0 0.5 107 Yellow-faced Honeyeater Lichenostomus chrysops 0 2 0.4 ,0.05 17 White-eared Honeyeater Lichenostomus leucotis 0 1 0.2 ,0.05 22 White-naped Honeyeater Melithreptus lunatus 1 0.2 ,0.05 0 0.0 0.00 14 Eastern Spinebill Acanthorhynchus tenuirostris 0 1 0.2 ,0.05 11 White-browed Babbler Pomatostomus superciliosus 0 1 0.2 ,0.05 39 Eastern Yellow Robin Eopsaltria australis 0 1 0.2 ,0.05 20 Unidentified robin Fam. Petroicidae 0 1 0.2 ,0.05 20 Grey Shrike-thrush Colluricincla harmonica 2 0.4 0.2 0 65 Magpie-lark Grallina cyanoleuca 5 1.1 0.5 8 1.5 0.7 90 Olive-backed Oriole Oriolus sagittatus 0 1 0.2 0.1 100 Black-faced Cuckoo-shrike Coracina novaehollandiae 0 2 0.4 0.2 105 White-bellied Cuckoo-shrike Coracina papuensis 0 2 0.4 0.1 70 White-browed Woodswallow Artamus superciliosus 1 0.2 ,0.05 2 0.4 0.1 35 Dusky Woodswallow Artamus cyanopterus 0 2 0.4 0.1 39 Australian Magpie Gymnorhina tibicen 5 1.1 1.9 4 0.8 1.3 329 Pied Currawong Strepera graculina 3 0.7 1.0 1 0.2 0.3 270 Strepera versicolor 0 1 0.2 0.3 300 Corvus coronoides 2 0.4 1.5 1 0.2 0.6 645 Little Raven Corvus mellori 1 0.2 0.6 0 541 Unidentified raven Corvus sp. 3 0.7 2.1 0 593 Welcome Swallow Hirundo neoxena 1 0.2 ,0.05 2 0.4 ,0.05 15 Australasian Pipit Anthus novaeseelandiae 3 0.7 0.1 0 23 Brown Songlark Cincloramphus cruralis 0 2 0.4 0.1 54 European Goldfinch Carduelis carduelis 0 1 0.2 ,0.05 18 Silvereye Zosterops lateralis 0 3 0.6 ,0.05 10 Eurasian Blackbird Turdus merula 0 2 0.4 0.2 95 European Starling Sturnus vulgaris 161 35.5 14.2 113 21.7 8.2 75 Starling juvenile 3 0.7 0.2 8 1.5 0.5 60 Unidentified small passerines Order Passeriformes 4 0.9 0.2 0 46 Unidentified birds 10 2.2 2.4 21 4.0 4.1 201 Bird subtotals 451 99.3 97.6 507 97.3 97.0

INSECTS Brown Beetle Fam. Scarabidae 0 1 0.2 ,0.05 2 Diamond Weevil Chrysolophus spectabilis 0 1 0.2 ,0.05 2 Unidentified Christmas Anoplognathus sp. 0 2 0.4 ,0.05 2 Beetles Unidentified Beetles Order Coleoptera 1 0.2 ,0.05 1 0.2 ,0.05 2 Cicada moerens 0 6 1.2 ,0.05 2 Insect subtotals 1 0.2 ,0.05 11 2.1 ,0.05

1 Biomass sources: Higgins 1999, Higgins and Davies 1996, Higgins and Peter 2002, Higgins et al. 2001, McDonald et al. 2003, Magrath et al. 2004, Marchant and Higgins 1990a, 1990b, 1993, Olsen et al. 2004, Olsen and Tucker 2003, Sharp et al. 2002, Strahan 1988. JUNE 2008 DIETS OF PEREGRINES AND HOBBIES 137

Appendix 2. Number of items, percentage by number and biomass, and individual mass of the different prey captured by the Australian Hobby in 1991–1992 and 2002–2003.

PREY SPECIES 1991–1992 2002–2003

PERCENT PERCENT PERCENT PERCENT MASS COMMON NAME SCIENTIFIC NAME N ITEMS BIOMASS N ITEMS BIOMASS (g)1

MAMMALS Gould’s Wattled Bat Chalinolobus gouldii 0 1 1.2 0.4 14 Mammal subtotal 0 1 1.2 0.4

BIRDS Stubble Quail Coturnix pectoralis 0 1 1.2 2.7 104 Budgerigar Melopsittacus undulatus 6 6.3 3.7 2 2.3 1.4 28 Crimson Rosella Platycercus elegans 6 6.3 17.9 1 1.2 3.5 135 Crimson Rosella juvenile 1 1.1 2.9 0 131 Eastern Rosella Platycercus eximius 3 3.2 7.0 2 2.3 5.4 106 Eastern Rosella juvenile 0 5 5.8 13.6 106 Red-rumped Parrot Psephotus haematonotus 0 1 1.2 1.6 61 Unidentified parrots Fam. Psittacidae 0 3 3.5 6.9 90 White-throated Needletail Hirundapus caudacutus 2 2.1 5.1 0 116 Sacred Kingfisher Todiramphus sanctus 0 1 1.2 1.0 40 Spotted Pardalote Pardalotus punctatus 0 3 3.5 0.7 9 Striated Pardalote Pardalotus striatus 0 2 2.3 0.6 12 Noisy Miner Manorina melanocephala 0 4 4.7 10.9 107 Red-capped Robin Petroica goodenovii 1 1.1 0.2 0 9 Golden Whistler Pachycephala pectoralis 1 1.1 0.6 0 26 Dusky Woodswallow Artamus cyanopterus 0 1 1.2 1.0 39 White-browed Woodswallow Artamus superciliosus 0 4 4.7 3.6 35 Tree Martin Hirundo nigricans 0 1 1.2 0.4 14 Eurasian Skylark Alauda arvensis 1 1.1 0.9 0 39 Rufous Songlark Cincloramphus mathewsi 1 1.1 0.6 0 27 Brown Songlark Cincloramphus cruralis 1 1.1 1.2 2 2.3 2.7 54 House Sparrow Passer domesticus 8 8.4 4.7 7 8.1 4.8 27 Unidentified sparrows Passer sp. 0 2 2.3 1.4 27 Red-browed Finch Neochmia temporalis 3 3.2 0.7 0 11 Silvereye Zosterops lateralis 0 1 1.2 0.3 10 European Starling Sturnus vulgaris 31 32.6 51.4 10 11.6 19.2 75 Starling juvenile 0 3 3.5 4.6 60 Unidentified small birds 1 1.1 0.7 3 3.5 2.3 30 Unidentified birds 1 1.1 1.3 7 8.1 10.2 57 Bird subtotal 67 70.5 98.8 66 76.7 98.7

INSECTS Christmas Beetle Anoplognathus porosus 3 3.2 0.1 0 2 Unidentified Christmas Beetles Anoplognathus sp. 2 2.1 0.1 4 4.7 0.2 2 Diaphonia Beetles Diaphonia dorsalis 0 1 1.2 0.1 2 Scarab Beetles Heteronychus aratora 0 1 1.2 0.1 2 Stag Beetle Lamprisma latreille 1 1.1 0.0 1 1.2 0.1 2 Cerambycidae Beetles Phorocantha sp. 0 1 1.2 0.1 2 Unidentified Beetles Order Coleoptera 1 1.1 0.0 3 3.5 0.2 2 Cicada Psaltoda moerens 16 16.8 0.7 7 8.1 0.4 2 Mole Cricket Order Orthoptera 1 1.1 0.0 0 2 Grasshoppers Order Orthoptera 4 4.2 0.2 0 2 Cockroach Blattoidea 0 1 1.2 0.1 2 Insect subtotal 28 29.5 1.2 19 22.1 1.0

1 Biomass Sources: Higgins 1999, Higgins and Davies 1996, Higgins and Peter 2002, Higgins et al. 2001, McDonald et al. 2003, Magrath et al. 2004, Marchant and Higgins 1990a, 1990b, 1993, Olsen et al. 2004, Olsen and Tucker 2003, Sharp et al. 2002, Strahan 1988.