CHAPTER 8

GENERAL DISCUSSION

This chapter will discuss the factors that influence the distribution of Fuscous

Honeyeaters within Eastwood State Forest. These include the abundance of foliage

insects, their major food source, and the interactions among themselves and other

species. I will also synthesize the role of the Fuscous in the bird community. A

wider view will then be taken to make comparisons between this and other Australian

bird communities and to suggest reasons why honeyeaters so frequently dominate the

avifauna.

The structure of the vegetation at Eastwood is simple ( Chapter 2 ), with most

of the available foraging substrates for foliage-using birds being provided in the

canopy level of trees. Fuscous Honeyeaters were associated with areas of gums

rather than stringybarks ( see Chapter 3 ). Previous work ( Woinarski and Cullen

1984, Recher 1985, etc. ) suggested that differences in type and abundance of foliage

insects between eucalypts in the subgenus M ono calyptus ( stringybarks ) and

Symphyomyrtus ( gums and boxes, hereafter called gums ) could account for such

bird preferences. Hence, it was important to study the foliage insects on trees in order

to explain the distribution of birds at Eastwood.

8.1 INSECTS ON PLANTS, AND THE EFFECT ON BIRDS

The arthropod fauna was studied in two ways. Chapter 4 gave the results of

sampling the canopy arthropods, that is, the standing crop, on stringybarks and gums.

Chapter 5 reported the results of a bird exclusion experiment, in order to assess the

impact of foraging by birds on the foliage arthropods. The numbers inside the

exclusion cages gave some idea of the productivity of the insects. Stringybarks had

the highest standing crop, that is most available insects, in autumn and winter ( see

Chapters 4 and 5 ), whereas the different gums peaked in numbers of insects at

different times and showed some year to year variation in the timing of peaks. For

instance, during the exclusion experiment ( Chapter 5 ), gums had the greatest

numbers in spring and summer but in the canopy sampling ( Chapter 4 ), E. viminalisand blakelyi peaked in autumn and winter, the melliodora in summer. The exclusion

experiment was mostly done in 1986, a rather dry year, and dry conditions can affect

the arthropods ( see Chapter 5 ). Therefore, this may not be the usual pattern of insect

abundance. Gums had the highest productivity ( see Chapter 5 ) in spring, the

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stringybarks were most productive in the same seasons as the peak of standing crop.

All peaks in numbers in the exclusion experiment were mostly due to one tree having

an outbreak of bugs ( coccids, pseudococcids or psyllids ), so caution should be

exercised when interpreting these results. The basic pattern of depletion of insects by

the birds in most seasons, however, appears to be one that can be accepted. This

suggests that foliage insects, as food for birds, were in short supply in most seasons.

The pattern of insect distribution should be noted, with individual branches

having mostly low numbers of insects, but with some branches having many insects

due to local outbreaks. At the level of individual plants, the same distribution of

insects occurred, with some trees having more insects due to flowering or growth or

stress ( see Chapters 4 and 5 ), but with most trees having fewer insects. Similarly,

differences will occur between tree species, because they vary in the amount of

nutrients and secondary chemicals ( Lambert and Turner 1983, Rhoades 1983 ). This

pattern of distribution was found both in the canopy sampling ( Chapter 4) and in the

exclusion experiments ( for the uncaged samples, see Chapter 5 ) although, in the

latter, the distribution was skewed to the right, indicating that saplings generally had

more arthropods than mature trees. This could be due to saplings growing more

actively ( as opposed to spending energy on reproductive activities or defensive

chemicals ), with a greater proportion of new leaves probably resulting in higher

nutrient levels and perhaps less secondary chemicals ( Mattson 1980, Jones 1983,

White 1984 etc. ).

Overall, few consistent differences in numbers of insects between the tree

types were found. As well as being generally sparse and significantly depleted in

most seasons ( Chapter 5 ), individual insects were mostly small. Insects were found

to be larger on gums than on stringybarks in the exclusion experiment ( Chapter 5 )

but not during the canopy sampling ( Chapter 4 ). There were also few significant

differences between gums and stringybarks in the types of arthropods that were

available, although gums had more 'sugary' lerps ( Family Psyllidae ) and

stringybarks had more larvae. With so few differences and with insects sparse, it

might be expected that there was little reason for birds to specialize, that is, the

foliage-gleaning insectivorous birds would show a weak tree preference. This

appears to be true for many species ( see Chapter 5 and Ford et al. 1986 ). The

Fuscous Honeyeater shows some overall preference for feeding on gums, particularly

E. viminalis, and against stringybarks ( E. caliginosa ), but this can vary, probably

according to the availability of food.

The greatest depletion of insects on gums ( see Chapter 5) was in spring, on

stringybarks in winter, and hemipterans were reduced more than other insect types.

Hemipterans make up a large proportion of the sample of foliage insects, and appear

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to be an important item in the diets of many honeyeaters and other small insectivores

( see Chapters 4, 5 and 6, and Paton 1980, Woinarski 1984a, 1985a ). Gums appear

to be more heavily utilized by birds as feeding substrates than stringybarks, as

supported by the finding of more abundant insects outside the cages than inside only

on stringybarks ( Chapter 5 ). Woinarski and Cullen ( 1984) found many more bugs

on gums than on stringybarks, which was not found here, but both studies found that

gums had more sugary lerps. Stringybarks had their seasonal peak in numbers in

autumn and winter and it might be expected that, as food was short for birds, they

might utilize the stringybarks more in the cooler seasons. The bird distribution data

( see Chapter 3) appear to support this with Fuscous Honeyeaters and several other

species found more in stringybark areas in autumn and winter. However, many birds

were more noticeable in these seasons because of their participation in mixed-species

feeding flocks which occurred more frequently in the stringybark areas. The areas

with fewer insectivorous birds appeared to have less depletion of insects ( Chapter 5 ).

8.2 BIRDS AND THEIR FOOD

Flowering mistletoe and trees are likely to attract both birds and insects.

However, mistletoe occurs at low density throughout the Forest ( see Chapter 2) and

the plants are mostly small. Eucalypts in flower would represent a large source of

food, but few eucalypts flowered during this study, and none of them abundantly.

The most common food increases for foliage-gleaners ( from insect sampling ),

therefore, were localized outbreaks of hemipterans. So, when patches of abundant

food occur, they are likely to be small, relative to the requirements of the birds.

Estimates of the daily energy requirements of Fuscous are not available, however,

McFarland ( 1985a ) calculated the daily estimated energy expenditure of Yellow-faced

Honeyeaters ( Lichenostornus chrysops, a species similar to Fuscous in weight and

diet ) to be about 75kJ. Calculations based on the average size of food items, in

particular lerps, and the energy available from these insects, suggests that a minimum

requirement by a Fuscous might be around 8000 lerps per day. The highest count of

lerps in the canopy sampling was about 10 per 100 leaves. So, a Fuscous might need

to search nearly 100,000 leaves to obtain its daily requirements, which would

probably represent most of the foliage of a mature tree. Observations of Fuscous

foraging suggest they search many trees rapidly, rather than one tree thoroughly.

Time-budgeting studies by Ford have found that Fuscous foraged for up to 480

minutes per day, so Fuscous would need to search around 200 leaves per minute!

Furthermore, the outbreaks of hemipterans can occur on any branch of any tree type at

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any time, that is, they are unpredictable. It is probably uneconomic to defend an area

without an outbreak, and the usually small size of outbreaks, relative to requirements,

means defense would be short term and highly localized. Birds, therefore, should

search for and quickly exploit these patches. This appears to be the case with the

Fuscous, as shown in the food supplementation experiment ( see Chapter 6 ).

Fuscous visited saplings with added food more frequently and for longer than control

trees. Usually, most of their feeding visits to a plant are short ( see Chapter 6 ), with

a lower frequency of longer foraging visits. The Fuscous appear to have adapted to

the distribution of food on eucalypts, by searching for insects rapidly and possibly

avoiding food items that are large ( see Chapter 5 ). Large food items took a long time

to handle and Fuscous were occasionally kleptoparasitic ( more likely to occur when

handling times are long, Barnard and Thompson 1985 ). It appears that Fuscous

continuously adjust their foraging behaviour, in particular visit length, in the light of

the rate at which they are finding food. This is additional to sampling the available

food resources and Fuscous have been seen to return to a tree that has been visited

already, after feeding in other plants, in the course of one foraging bout.

Such sampling is necessary for a bird to learn about its environment. This is

one of the major areas where optimal foraging models break down, as one of the

assumptions is 'perfect knowledge' of the expected food resources by the birds. In

addition, most tests deal with two or three prey types or densities ( see, for example,

references in Kamil et al. 1987 ). In reality, birds are exposed to a continuous

distribution of food abundance, as mentioned above. It is possible that many of the

responses of birds in tests, to changes in abundance of the less profitable food types

or patches ( which, from the models, ought to be ignored except when more profitable

food is very scarce or highly variable ), may be due to the birds having a relative view

of food abundance, which is continuously modified in the light of encounter and

intake of food items. These factors would account for the existence of 'partial

preferences', which have been frequently observed in experiments testing optimal

foraging ( Krebs and McCleery 1984, Pyke 1984, references in Kamil et al. 1987 ).

Another criticism ( see, for example, Gray 1987) is that energy intake may not always

be the currency that is being optimized - most tests are on non-breeding birds, and

breeding birds may have special nutrient requirements for themselves or their young,

and it might not be always the case that short-term energy gain is linked with increased

evolutionary fitness.

Fuscous Honeyeaters display considerable flexibility in their foraging

behaviour ( see Chapter 5 ), foraging on most substrates, at most heights, and by

several methods. Calculations of niche breadth ( see Ford et al. 1986) show that the

Fuscous are, overall, one of the most generalized species of bird in Eastwood

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( exceeded only by Noisy Friarbirds, who are only present for spring and summer,

and Restless Flycatchers, which are in low numbers ). A close relationship between

numbers of Fuscous and insect abundance was not found, but is unlikely to be

detected, due to the great variability associated with arthropod sampling. However,

Fuscous were associated with areas of many gums, rather than stringybarks,

presumably because of the more acceptable food they can expect on the gums. They

also displayed considerable aggression towards other small insectivores, and may

have influenced the distribution of these species. This will be discussed in the next

section.

8.3 INTERACTIONS AMONG BIRDS

Being the dominant species of small insectivore in the Forest means the

Fuscous is probably little restricted in its fundational niche. Other small, leaf-gleaning

insectivores, such as thornbills and other small honeyeaters, are chased by Fuscous

and may avoid areas where Fuscous are common. Certainly, their distribution within

Eastwood overlaps little with that of the Fuscous Honeyeater. The relative structural

simplicity of the vegetation at Eastwood precludes avoidance of Fuscous aggression

by partitioning the habitat by height. In particular, the almost complete lack of a well-

developed shrub-layer, means that small birds generally cannot avoid attacks by

Fuscous ( as can happen in other areas, for example Smith and Robertson 1978 ).

Not only are the small insectivores restricted in range which must reduce their

numbers, but also they may be further disadvantaged by having to forage in less

productive areas ( Chapter 5 ) or on less desirable tree species ( if they prefer lerps,

for instance ). Some species, however, may prefer stringybarks such as some

thornbills and bark-foragers ( see Bell 1983a, Noske 1985, Woinarski 1985a, Ford et

al. 1986, etc. ).

Some other evidence supports the argument that such small insectivorous

species are restricted in their foraging behaviour. Ford et al. ( 1985 ) found that

White-naped and Brown-headed Honeyeaters were reduced in numbers during the

1979-1980 drought ( which also affected numbers of insects, see Bell 1985a ),

whereas Fuscous Honeyeaters were not, the implication being that the latter species

had access to sufficient resources during a time of shortage. Furthermore, Ford

( 1989 ) compared the amount of time spent foraging by several species, from time-

budgeting work. All species foraged for longer in autumn and winter, but the

Fuscous Honeyeater spent less time feeding in any season than the White-naped

Honeyeater. This suggests that the former, more dominant, species was obtaining

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food more efficiently than the latter although the White-napes, being smaller, may

need higher intake rates of food.

It was suggested above that there were no good reasons for strong selection to

specialize in foraging on a particular tree type, and that patches of abundant food were

not usually defensible. However, it was found that Fuscous Honeyeaters were

significantly positively associated with numbers of gums ( Chapter 3 ). Furthermore,

during the breeding season, Fuscous defended territories around the nest that were

used for some feeding ( Chapter 7 ). Other reasons, in addition to the distribution of

food, must be sought to explain the observed distribution of Fuscous.

Reproductive success of Fuscous, as measured by the number of nests

successfully fledging young, was higher in areas where Fuscous nests were clumped,

relative to areas where the nests were isolated ( see Chapter 7 ). The areas of high

Fuscous density, and where nests were close, occurred in parts of Eastwood that had

many gums ( mostly E. viminalis and blakelyi, and see Chapters 3 and 7 ).

Reproductive success was generally low, as found in many other Australian

passerines ( Chapter 7 ). The fewest nests fledged in a dry year ( see Chapter 7 ) and

bad weather caused many nests to be abandoned, presumably after the death of the

nestlings. However, predation was also thought to cause the loss of many nests. By

nesting close to other Fuscous, nest success was increased by up to 40% of nests in

some years and 20% overall.

Nesting in groups is usually advantageous for Fuscous probably because in

groups, greater vigilance for, and co-operation in mobbing of, predators and co-

operative chasing of unknown conspecifics and intruding birds of other species is

possible. These would decrease costs of defense against intruders ( both predators

and competitors ) for the individual and increase foraging efficiency through less

interspecific competition and fewer interruptions. Costs associated with higher

density of birds from grouping are more competition for ( generally ) sparse food

insects and increased costs of defence of the territory and nest against conspecifics.

The density of other species of foliage-feeding insectivores was less in Fuscous-

dominated areas ( Chapters 3 and 7 ), and these species were chased more frequently

by Fuscous than non-competing species ( Chapter 3 ). The overall density of

insectivores was, however, higher due to the abundance of the Fuscous and intrusion

rates into territories were quite high ( Chapter 7 ). The increase in reproductive

success did not occur in the dry year suggesting that, if food was short, the higher

density of Fuscous due to grouping, which presumably increased the competition for

that food, overwhelmed any advantage of nesting in groups.

The process of habitat selection for the Fuscous is, therefore, thought to

involve selection of an area by tree type, because gums, on average, offer more of the

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food that they prefer. Fuscous congregate in these areas and, by grouping, can

dominate the food resources and increase their reproductive success. Behaviours such

as toleration of close neighbours, and co-operation with them, then evolve to decrease

the costs of the high density of Fuscous. The selection of habitat is re-inforced by the

increased reproductive success of Fuscous nesting in groups. The mechanisms for

habitat preference are thus behavioural, social and evolutionary.

8.4 THE S'T'RUCTURING OF COMMUNITIES

The distribution and abundance of food for birds in Australian eucalypt forests

and woodlands will have an enormous impact on the community organization. Where

nectar is abundant, which tends to be in infertile, often mesic, environments, the

aggressive dominants in the community are nectarivorous honeyeaters ( Paton 1979,

Rooke 1979, Loyn 1985, McFarland 1985a ), and most of the honeyeaters will

concentrate on this resource if it is not defended by a more dominant species ( Ford

and Paton 1977, 1982, Ford 1979, Wykes 1982, McFarland 1985a ). In sites on

richer soils, insects appear to become the more significant food source, and

insectivorous honeyeaters predominate in the community ( Dow 1977b, 1979, Clarke

1988, Wykes 1982, 1985, present study ). Insectivorous honeyeaters are generally

uncommon in rainforests, and in other habitats where there are few eucalypts ( see

references in Keast et al. 1985 ). This is partly because a preferred food, psyllids (

Paton 1980, Woinarski 1984a, 1985a ), are rare on non-eucalypts ( Woinarski and

Cullen 1984 ).

Honeyeaters, particularly the dominant ones in a community, can be highly

aggressive. They are usually most aggressive towards conspecifics, but will attack

most species, particularly other birds with similar foraging niches ( Dow 1977b,

Clarke 1984b, McFarland 1984b, present study ). Nectarivorous honeyeaters will

defend feeding territories while it is economic to do so, often for relatively short

periods of time ( Paton 1979, Rooke 1979, Ford and Paton 1982, McFarland 1986d

), but will feed with relatively little aggression when food is super-abundant. Some

species, particularly the insectivorous ones such as the Noisy Miner ( Dow 1977b ),

will exclude or attempt to exclude all other birds from the areas that they are

defending. This super-aggression may require a large expenditure of energy and

suggests that the increased access to food outweighs the cost of aggression. Although

there are no detailed studies on the relationships between food abundance and the

ecology of miners, it is possible that the argument outlined for Fuscous may also

apply to other species. If food increases are unpredictable, instead of honeyeaters

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being able to develop specialized foraging niches to avoid competition, priority of

access to the food resources, that is dominance, must be re-asserted with all changes

in food abundance. This necessitates a continuously elevated level of aggression.

Several studies have found that the numbers of honeyeaters varied through

nomadism or migration in response to food abundance, or that the birds changed their

diets seasonally ( Halse 1978, Paton 1979, 1982, 1985, Collins and Briffa 1982,

Wykes 1982, 1985, Collins et al. 1984, Pyke 1985 but see Pyke 1983, McFarland

1986b etc. ). These were mostly changes shown by nectarivorous honeyeaters and

were probably adaptations to the variable distribution of this food source. Where

there were large changes in food abundance in a shorter time span ( McFarland 1985a,

1986a) birds could only respond behaviourally. It appears that nectarivorous

honeyeaters more often have sufficient excess resources to be able to deposit and store

fat rapidly ( Paton 1979, Ford and Pursey 1982, McFarland 1985a, 1986b ). This

will act as a buffer against large short-term fluctuations in food abundance.

Honeyeaters that feed more on insects do not appear to track food levels to the

same extent as more nectarivorous honeyeaters, being more likely to defend, or

attempt to retain exclusive use of an area, thus having most access to its resources

( Dow 1977b, 1979, Smith and Robertson 1978, Wykes 1982, present study ).

These are presumably also adaptations to the less variable, but often sparse

distribution of insects. Woinarski and Cullen ( 1984) found sites with intermediate

rainfall to have the lowest seasonal fluctuations in numbers of insects ( and also the

greatest abundance, on average ). This implies that food for insectivorous birds is

most predictable in areas of intermediate rainfall, which should mean that more species

can co-exist in these areas, as the food resource base is able to be partitioned. More

species do appear to co-occur in intermediate areas ( see, for example, Loyn 1985 ),

and a higher proportion of these species are co-operative breeders ( Ford et al. 1988,

Ford 1989 ), relative to sites of more or less rainfall. This is not surprising as, the

more predictable the food supply, the more sedentary a population of birds can be.

The lack of necessity to move leads to more continuous association between birds,

which can result in a more complex degree of social organization. One development

of such social behaviour is the domination of resources in an area, with some degree

of co-operation to reduce the costs associated with higher intra-specific density. In

fact, the vast majority of co-operatively breeding birds are sedentary insectivores

( Ford et al. 1988 ). The generally more variable supply of nectar as a food source

means that less sociable behaviour might be expected because most birds will have to

move in search of food and, in support of this, no nectarivores have been found to be

regular co-operative breeders.

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With increasing unpredictability and abundance of food, it has been suggested

that the size of the most abundant species should decrease ( Pirnm 1978 ). McFarland

( 1985a, 1986a) found that, in an environment with large day-to-day fluctuations in

nectar abundance, the most numerous species of honeyeater was the smallest. This

species ( Eastern Spinebill ) was subordinate to the larger New Holland Honeyeater

( the 'organiser' species, see McFarland 1985a ) which defended territories that were

not adjusted to short term resource changes. It is likely that, in the present study, the

community was dominated by such a small species because the insect food was sparse

and locally variable. The smaller honeyeaters that are rarely dominant can be expected

to be more nomadic and move in flocks to find undefended food sources or swamp

the areas defended by larger honeyeaters ( Wykes 1982, McFarland 1985a etc. ).

In summary, the factors responsible for structuring the bird community at

Eastwood, and more generally, appear to be patterns of variability in the abundance of

food in time and space and the interactions between species. Predictability of food is

likely to be more important in determining bird community organization than actual

food abundance. At stable, but low, levels of food, bird species can specialize in

foraging in a similar way to when food abundance is stable but high, although in the

former case, it might be expected that fewer species and individuals could co-exist.

These communities may show a high degree of interaction between species, which

may compete. As food changes in abundance, so too will the levels of aggression and

defense of resources ( Carpenter and MacMillen 1976, McFarland 1985b, 1986d ). If

food changes in abundance unpredictably, though, species will more likely be

generalized feeders or nomadic or migratory, and interactions between species may be

less important in structuring the community ( see Wiens 1983 for an illuminating

discussion of the methodologies and conclusions of some community structure studies

although his is an extreme view, see also Holmes et al. 1986 ).

8.5 COMPARISONS WITH OTHER COMMUNITIES

In the community at Eastwood, Fuscous Honeyeaters are numerically the

dominant species of honeyeater. They occur in highest numbers in areas where gums

such as Eucalyptus viminalis and E. blakelyi are common. They choose these areas

because of higher abundance of preferred food and because the presence of other

Fuscous means they can, as a group, dominate the food resources. In these densely

occupied areas, they have better average reproductive success from increased

protection against predators and, possibly, higher foraging efficiency. Aggression of

the Fuscous is primarily directed at competitors for food ( same and other species )

and in nest or territory defence. Other bird species may be divided into those that are

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affected by interactions with the Fuscous, and those that are not. Into the former

group fall species that are excluded from Fuscous-dominated areas because they are

competitors for food ( such as other smaller honeyeaters, White-naped, Yellow-faced

and Brown-headed and thornbills ), and those that may associate with Fuscous for

protective nesting benefits ( see Chapter 3 ). Species that are not affected by Fuscous

are larger and/or forage differently, and are likely to occur in areas with a necessary

resource. The bird community, therefore, tends to have two species suites ( see Table

8.1 ). The Fuscous and associated species occur in gum-rich areas and other small

leaf-gleaners are found in areas dominated by stringybarks. Most of this latter group

of birds associate to feed in mixed-species feeding flocks in the cooler seasons. Their

choice of habitat may be considerably modified by avoidance of Fuscous

Honeyeaters. Some bird species change seasonally from one suite to another,

depending on the availability of resources.

Previous to this project there has been only one major study on the community

organization of insectivorous honeyeaters, that of Wykes ( 1982, 1985 ). He worked

at three sites in Victoria, studying the honeyeater community with emphasis on the

endangered Helmeted Honeyeater ( Lichenostomus melanops cassidix), a subspecies

of the more common Yellow-tufted Honeyeater ( L. melanops, subspecies meltoni inthis area ). He encountered seven species of honeyeaters, all in the genus

Lichenostomus except the Bell Miner. Amongst these species, Wykes found three

sorts of social organization. Yellow-tufted Honeyeaters ( including Helmeted ) and

Bell Miners were semi-colonial aggressive dominants and defended areas of reliable

food year-round. The same occurred to a lesser extent in the Fuscous Honeyeaters

which, however, also had a nomadic/migratory component in the population. It was

not dominant to the Yellow-tufted Honeyeaters or Bell Miners, probably because it

was smaller and had a looser social structure, but usually occurred in different

habitats, thus avoiding aggressive interference. The other species of honeyeaters,

Yellow-faced and Yellow-plumed, were not highly aggressive and congregated in

large numbers only where food was abundant, which in this case was nectar. In

addition, the White-eared Honeyeater, a species subordinant to most other

honeyeaters, specialized in feeding on bark substrates, which are poorly utilized by

the other honeyeaters. It has been found to be abundant only in habitats avoided by

other honeyeaters as being poor sources of food ( for instance, in E. pauciflorawoodlands, Loyn 1985, see Ohmart et al. 1983 for insect abundance ).

The community organization found at Eastwood ( see also Ford et al. 1986 )

can be compared with that of Bondi State Forest, near Bombala on the Southern

Tablelands of N.S.W. ( see Recher et al. 1983, 1985, Recher and Holmes 1985 ). Of

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TABLE 8.1: Species suites at Eastwood State Forest. Species are given as

associating with Fuscous Honeyeaters if they were frequently seen in the high-density

Fuscous areas and if their numbers were positively correlated with that of Fuscous

( see Tables 3.7 and 3.8 ). Some species were equally commonly associated with

either suite ( in middle of table ) or weakly tended to be seen more in the areas of

either suite ( shown displaced towards Fuscous or non-Fuscous end ). Abbreviations

are: HE honeyeater, GST Grey Shrike-thrush, BFCS Black-faced Cuckoo-shrike,

CrST Crested Shrike-tit, Large includes choughs, magpies, currawongs and ravens,

seasons are spr, sum, aut, win.

FUSCOUS DENSITY

HIGH LOW

Fuscous HE

White-naped FIE

Brown-headed HE

Yellow-faced HE

Red Wattlebird Noisy Friarbird

Thornbills

Pardalotes

Brown Treecreepers White-throated

Varied Sittella

Rufous Whistler Golden Whistler

( in sum & aut )

( in spr & sum ) <--GST--> ( in aut & win )

BFCS

CrST

Speckled Warbler

Superb Fairy-wren

Willie Wagtail Grey Fantail

Restless Flycatcher Leaden Flycatcher

Dusky Woodswallow

Eastern Yellow Robin

Scarlet Robin

Rose Robin

Rosellas

Magpies Large

Choughs

Sacred Kingfisher

Kookaburras

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the more common species, Recher and Holmes and Recher et al. ( 1985 ) presented

detailed foraging data on 41 species from Bondi. Ford et al. ( 1986 ) collected similar

data on 40 species at Eastwood. Twenty-five species ( about 60% ) were found in

both areas. Bondi is wetter than Eastwood ( see Recher et al. 1983 and Chapter 2 )

and has more undergrowth in some areas. Most of the differences in the avifauna

seemed to relate to these factors, with species such as Superb Lyrebirds ( Menuranovaehollandiae ), Ground Thrushes ( Zoothera dauma ), Rufous Fantails

( Rhipidura rufifrons ) and Eastern Whipbirds ( Psophodes olivaceus ) being

confined to the wetter forests such as that studied by Recher and Holmes ( 1985) and

scrub-wrens being found where the undergrowth is well-developed ( Blakers et al.1984, Cameron 1985, Ambrose 1986 ). Recher et al. ( 1983) sampled the insects at

Bondi, and found a pronounced seasonality, with low numbers and biomass in winter

( which is colder at Bondi than Eastwood ), although different plant types varied in

both timing and height of peak abundances. These samples were taken during and

after the 1980 drought, which could have greatly affected the results. Numbers of

birds were also highly seasonal, with migrant or locally nomadic birds increasing the

spring-summer totals to nearly double the winter figures. Seasonality of arthropods

was slight at Eastwood ( Chapter 4) and summer totals of birds were only about 20%

higher than in winter ( Chapter 3 ).

Recher et al. ( 1985) and Ford et al. ( 1986) found that the bird species at

both sites formed foraging guilds primarily divided by food type and foraging

substrate, with foraging method an important secondary division. The range of

foraging behaviours was not very different from those found amongst insectivorous

birds in a Northern Hemisphere forest ( Robinson and Holmes 1982, Recher and

Holmes 1985, Holmes and Recher 1986 ). There were differences due to the

structure of the vegetation ( eucalypts are more open and irregular than the dominant

trees at the North American site, eucalypt leaves are pendant, and there can be strips of

peeling bark ). Also, northern forests generally lack small foliage insects providing

sources of carbohydrates such as lerps, manna and honeydew, but have more large

lepidopteran larvae available to insectivorous birds ( at least in spring, see Holmes etal. 1979, Robinson and Holmes 1982, Holmes and Recher 1986). These differences

can be expected to influence the bird community structure.

Comparisons between bird communities in Australia and other continents

support the suggestion above that availability of food can have a profound effect on

the types of birds present and the ways in which the bird communities are organized.

Many Australian birds are wholly or partially dependent on carbohydrates ( nectar

and/or lerps, manna and honeydew, see Paton 1980, Woinarski 1985a, Recher et al.

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1985 ). Few temperate communities elsewhere are dominated by nectarivores, and the

organization of those in the tropics is similar to that found in Australia, with larger and

aggressive species excluding smaller species from areas of richer nectar sources

( Feinsinger 1976, Wolf et al. 1976, Carpenter 1978, references in Paton 1986 etc. ).

Ford et al. ( 1986 ) compared the foraging guild structure of birds in the eucalypt

woodland studied here, with that of evergreen oak woodland in North America and

southern Spain. He found a similar percentage to be partial or complete insectivores

( about 75% ), but whereas the alternative food sources were nectar and alternative

carbohydrates in Australia, many species of birds in the other communities consumed

fruits or seeds. These food sources are eaten by a few specialized species in

Australia, and fleshy fruits are not abundantly produced in eucalypt forests, possibly

due to nutrient deficient soils ( particularly potassium, Milewski 1986 ). Recher and

Holmes ( 1985 ) found similar differences when comparing eucalypt and Northern

Hemisphere forests.

Support for the contention that food abundance is relatively aseasonal in

Australia is given by the different breeding seasons and clutch sizes between Northern

Hemisphere and Australian birds. The pattern in Europe and North America is

generally for one clutch per season of rather large size ( Perrins and Birkhead 1983 ).

In Australia, most smaller passerines lay several clutches over a longer season

( Wyndham 1986 ), whether or not previous attempts have been successful, and these

clutches are small ( mostly 2 to 3 eggs, see Woinarski 1985b, Yom-Tov 1987, Ford

1989 ).

Another major difference between Australian and overseas bird communities is

the relative lack of long distance passerine migrants in Australia. In Northern

Hemisphere forests up to 80% of species may migrate ( Willson 1976, Herrera 1978,

Rabenold 1978, Nilsson 1979, Alatalo 1982, Gauthreaux 1982, Holmes et al. 1986,

etc. ). This will have a major impact on the bird communities in the overwintering

areas ( see references in Keast and Morton 1980 and Gauthreaux 1982 ). Northern

Hemisphere species migrate to avoid winter crashes in arthropod numbers and suffer

high over-winter mortality ( Graber and Graber 1979, Perrins 1979, Gauthreaux

1982, Perrins and Birkhead 1983 ). Of the foliage-gleaning species at Eastwood,

only Yellow-faced and White-naped Honeyeaters and Silvereyes are considered to

migrate for a long distance ( as opposed to local or regional nomadism, Keast 1968

etc. ) and these species also have local breeding populations. McClure ( 1975 )

suggested that the difference was due to the relative isolation of Australia from other

land masses until quite recently, geologically speaking ( see also review by

Gauthreaux 1982 ). It is also possible that the relative lack of extreme cold means that

food for birds is little reduced in Australian winters ( see Chapters 4 and 5, and

178

Woinarski and Cullen 1984, Wykes 1982 etc. but see Recher et al. 1983 ), rendering

long migration unnecessary. Keast ( 1968 ) found the percentage of resident

honeyeater species in a community increased with increases in the reliability and

amount of rainfall, and thought that most movement was in response to changes in

nectar availability, which could vary markedly between places and years. He

concluded that honeyeaters were quite plastic in their movement patterns and feeding

behaviour. Ford ( 1989 ) suggested that the proportion of resident bird species of all

types was correlated with rainfall. Shureliff ( 1986 ) reached similar conclusions

when comparing bird communities from some arid sites, but the nectarivorous and

frugivorous species were more likely to move.

Predominance by one family of birds is relatively rare in bird communities on

other continents. In some studies, families may represent a similar percentage of

species to that found in the present study and elsewhere in Australia ( about 40% here,

can be up to 60% or more, Paton 1979, 1985, Wykes 1982, 1985, McFarland

1985a ). Holmes et al. ( 1986) and Rabenold ( 1978 ), working in North America

found species in the family Parulidae to comprise around 40% of total numbers,

whereas in European forests, the Family Sylviidae forms a similar percentage of the

community ( Nilsson 1979, Cody 1978 ). The communities are, however, rarely

aggressively dominated by one, or a few species, although some species may be

numerous. The aggression of honeyeaters ( Dow 1977b, Rooke 1979, Wykes 1982,

Loyn et al. 1983, McFarland 1985b ) seems to be a contributing factor to their

dominance of Australian bird communities.

8.6 CONCLUDING REMARKS

Honeyeaters are the dominant, in terms of abundance and aggression, group

of birds in many Australian eucalypt forests and woodlands. Nectarivorous

honeyeaters will be dominant if nectar ( or alternatives ) is abundant and reliable,

otherwise a more generalized species will predominate. Increases in both food

supplies can be relatively unpredictable. Nectar can be economic to defend if it is

sufficiently abundant, but it will be more ephemeral than insects and honeyeaters must

often be nomadic to take advantage of local increases. Insects are often less abundant,

but will always be present, which means honeyeaters can be resident. The sparseness

of insects ( and relatively slower renewal rates ) may preclude individual defence of an

area but, by grouping or flocking, honeyeaters can dominate an area that has a

reasonable expectation of providing sufficient food. By defence, they may maintain

an adequate food abundance. The unpredictability of food increases, and the

179

necessary constant aggression of dominants means that dominant honeyeaters are

unable to become specialized foragers and so, are usually separated by habitat. This

can have a large impact on subordinate species of birds, which may avoid areas where

the dominant honeyeater(s) are common and tend to flock and/or utilize habitats or

substrates not occupied by dominants. Honeyeaters have probably evolved

concurrently with eucalypts, and seem to be adapted to utilizing the food provided by

eucalypts. The dominance of one family of birds in Australia is, therefore, closely

related to the dominance of one genus of plants.

180

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196

APPENDIX 1

PLANT SPECIES OF EASTWOOD STATE FOREST

Compiled by J. B. Williams, Department of Botany, University of New England,

1986.

TREES

Angoplzora floribunda (Rough-barked Apple )

Eucalyptus blakelyi (Blakely's Red Gum )

E. bridgesiana (Long-leaved Box )

E. caliginosa (New England Stringybark )

E. melliodora (Yellow Box )

E. viminalis (Manna Gum )

SHRUBS

Acacia filicifoliaA. ulicifoliaBrachyloma daphnoidesCassinia quinquefariaDaviesia latifoliaHibbertia linearisIndigofera australisLespedeza junceaMelichrus urceolatusO. viscidula

MISTLETOES

Amyema miquelliiA. pendulaMuellerina eucalyptoides

A. implexaBossiaea buxifoliaCasuarina littoralisCryptandra amaraExocarpos cupressiformisHovea linearisJacksonia scopariaLissanthe strigosaOlearia ellipticaPultenaea microphylla

197

HERBS - GENERAL DICOTYLEDONS

Acaena ovinaAsperula confertaDesmodium variansEpilobium billardierianumGalium sp.Goodenia bellidifoliaGonocarpus tetragynusHaloragis heterophyllaH. tripartitaMentha diemenicaOreomyrrhis eriopodaPhyllanthusPlantago debilisPoranthera microphyllaRhagodia hastataSolanum nigrumTrifolium repensViola betonicifolia

HERBS - ASTERACEAE

Brachycome sp.Centipeda minimaCo ny za sp.Cymbonotus lawsonianusHelichrysum semipapposumHypochoeris radicataSenecio sp.Taraxacum officinaleV. sp. 2

Ajuga australisCentaurium erythraeaDichondra repensEuphorbia drummondiiGeranium solanderiG. hederaceaGratiola larifoliaHydrocotyleHypericum gramineumOpercularia asperaOxalis corniculataPimelea curvifloraPolygala veronicaPrunella vulgarisScleranthus biflorusStackhousia monogynaVeronica calycinaWahlenbergia sp.

Calotis cuneifoliaCirsiurn vulgareCrepis capillarisGnaphalium involucratum

H. sp.Lagenifera sp.Sonchus oleraceusVittadinia sp. 1

198

HERBS - GRASSES

Aristida ramosaDanthonia linkiiDichelachne micrantha

Microlaena stipoidesPoa sieberiana

Bothriochloa macraD. sp.Echinopogon caespitosus

Panic= effusumSorghum leiocladurn

199

Sporobolus sp. Stipa sp.Therneda australis

HERBS - GENERAL MONOCOTS

Bulbine bulbosaCyperus sanguinolentusD. revolutaHypoxis hygrometricaLomandra longifoliaLuzula sp.Tricoryne elatius

HERBS - FERNS

AspleniumflabellifoliumPellaea fakata var. falcata

Carex inversaDianella laevisFirnbristylis dichotomyJuncus sp.L. multifloraSchoenus ericetorum

Cheilanthes tenuifolia

TRAILERS

Glycine clandestina G. tabacinaHardenbergia violacea Rubus parvifolius

APPENDIX 2

COMMON AND SCIENTIFIC NAMES OF BIRDS

Names follow Blakers et al. 1984.

COMMON NAME SCIENTIFIC NAME

200

Family Meliphagidae:

Fuscous Honeyeater

Yellow-faced Honeyeater

White-eared Honeyeater

White-naped Honeyeater

Brown-headed Honeyeater

Noisy Miner

Red Wattlebird

Noisy Friarbird

Eastern Spinebill

Scarlet Honeyeater

Regent Honeyeater

Family Acanthizidae:

Buff-rumped Thornhill

Striated Thornhill

Yellow-rumped Thornhill

White-throated Gerygone

Speckled Warbler

Family Pardalotidae:

Spotted Pardalote

Striated Pardalote

Family Zosteropidae:

Silvereye

Lichenostomus fuscusL. chrysopsL. leucotisMelithreptus lunatusM. brevirostrisManorina melanocephalaAnthochaera carunculataPhilemon corniculatusAcanthorhynchus tenuirostrisMyzomela sanguinolentaXanthomyza phrygia

Acanthiza reguloidesA. lineataA. chrysorrhoaGerygone olivaceaSericornis sagittatus

Pardalotus punctatusP. striates

Zosterops lateralis

Family Climacteridae:

White-throated Treecreeper Clirnacteris leucophaea

Brown Treecreeper C. picumnus

201

Family Neosittidae:

Varied Sittella Daphoenositta chrysoptera

Family Muscicapidae:

Golden Whistler

Rufous Whistler

Grey Shrike-thrush

Crested Shrike-tit

Satin Flycatcher

Restless Flycatcher

Leaden Flycatcher

Grey Fantail

Willie Wagtail

Eastern Yellow Robin

Scarlet Robin

Rose Robin

Family Campephagidae:,

Black-faced Cuckoo-shrike

White-winged Triller

Family Dicaeidae:

Mistletoebird

Family Maluridae:

Superb Fairy-wren

Pachycephala pectoralisP. rufiventrisColluricincla harmonicaFalcunculus frontatusMyiagra cyanoleucaM. inquietaM. rubeculaRhipidurafuliginosaR. leucophrysEopsaltria australisPetroica multicolorP. rosea

Coracina novaehollandiaeLalage sueurii

Dicaeum hirundinaceum

Malurus cyaneus

Family Artamidae:

Dusky Woodswallow

White-browed Woodswallow

Family Cuculidae:

Horsfield's Bronze-cuckoo

Fan-tailed Cuckoo

Artamus cyanopterusA. superciliosus

Chrysococcyx basalisCuculus pyrrhophanus

Family Sylviidae:

Rufous Songlark

Cinclorhamphus mathewsi

Family Coraciidae:

Dollarbird Eurystomus orientalis

Family Grallinidae:

Australian Magpie-lark Grallina cyanoleuca

Family Ploceidae:

Diamond Firetail Emblema guttataRed-browed Firetail E. temporalis

Family Oriolidae:

Olive-backed Oriole

Family Hirundinidae:

Welcome Swallow

Tree Martin

Family Columbidae:

Peaceful Dove

Family Tumicidae:

Painted Button-quail

Family Alcedinidae:

Laughing Kookaburra

Sacred Kingfisher

Oriolus sagittatus

Hirundo neoxenaCecropis nigricans

Geopelia placida

Turnix varia

Dacelo novaeguineaeHalcyon sancta

Family Platycercidae:

Crimson Rosella Platycercus elegansEastern Rosella P. eximiusRed-rumped Parrot Psephotus haematonotus

Family Cacatuidae:

Galah

Cacanta roseicapilla

Yellow-tailed Black-Cockatoo Calyptorhynchus funereus

202

Family Coracoracidae:

White-winged Chough Corcorax melanorharnphos

Family Cracticidae:

Grey Butcherbird Cracticus torquatusAustralian Magpie Gymnorhina tibicenPied Currawong Strepera graculina

Family Corvidae:

Australian Raven Corvus coronoides

203

APPENDIX 3

DIS'T'RIBUTION MAPS OF OTHER GROUPS OR SPECIES OF BIRDS

204

FIGURE 1: Maps of Eastwood showing seasonal distribution of Red Wattlebirds.

For these maps seasons are, in order, autumn 1986, winter, spring, summer, autumn

1987 and autumn 1988. Relative densities of wattlebirds are: 0 ( empty circle ), >0-1

( vertical lines in circle ), >1-2 ( black grid on white in circle ), >2 ( white grid on

black in circle ).

c),9to

00

0 0 0 0• O•

I

0 0° 0 00 0 0

0 0 0co

0IM

lip

0•••

••0 •

•• III0 • •

0O

(III0

0 O

NI 00 •O(1111"i•Ili•00•

0 0 0 0 0 0 0o0_O 0 0

O • O•0

000 0

•••

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0 00S0SUAga

SPOOd SLUDa

SpQ0

- RoadsCD Dams 1 . . . N

4 T500

Roads

C3 Dams P . . . N

• 4 af500 0

metres 0

metres

- Roads

0 Dams

- Roads

C) DamsN

0 500metres0 500

metres

0

o0 0

FIGURE 2: Maps of Eastwood showing seasonal distribution of Noisy Friarbirds.

For these maps seasons are, in order, spring and summer. Relative densities of

friarbirds are: 0 ( empty circle ), >0-1 ( vertical lines in circle ), >1-2 ( black grid on

white in circle ), >2 ( white grid on black in circle ).

205

0 00 0 0 0 0 0

0 0 0 0 0

0 0 00 0 0 0 0

0000

Roads

CD Dams--- Roads

CD Dams 0 500metres

NP . I . 1 4 TN

P ' ■ ■ • I

0 500metres

FIGURE 3: Maps of Eastwood showing seasonal distribution of pardalotes. For

these maps seasons are, in order, autumn 1986, winter, spring and autumn 1987.

Pardalotes were extremely rare in summer and autumn 1988, and so maps are not

included. Relative densities of pardalotes are: 0 ( empty circle ), >0 ( vertical lines in

circle ), >1 ( black grid on white in circle ), >2 ( white grid on black in circle ).

206

0 0 c)---00 0 0 0 0 0 0

0 0 0 0

I0

metres

Roads

CD DamsN

4 Tr • • • .

Roads

CD DamsN

- ■ T5000 500

metres

0 S 0 0 0

0 0 0001111►

0 0 0 0 00 0 0 0 0 0

0Q,0 00

0 0

(III •

101 III'

00 0 0 0 0

0 0 0 0 0 0 0,0Q,0 0 0

1

00

ill

I. al

OS

000

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0

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ills

O loll 00 0 0

ED 0 00 0

000

000000

•00

0

1111'

00

saJlaw

saJlaw00S

N

0 00S

N

0swea ,c)

spood

... S W 0 a

spend

CD...

FIGURE 4: Maps of Eastwood showing seasonal distribution of treecreepers (

White-throated and Brown Treecreepers ). For these maps seasons are, in order,

autumn 1986, winter, spring, summer, autumn 1987 and autumn 1988. Presence of

unidentified treecreepers and of White-throated Treecreepers indicated by vertical lines

within the circle, that of Brown Treecreepers by horizontal lines within the circle.

Split circles indicate that both species were seen at that point within a season.

207

- RoadsC) Dams

N■ a a ■ 4

p0 500metres

RoadsCD Dams

0. . . N

• 4 T500

metres

'III►S 'III►

'llhiil^S0

lIIi III'S'III'I- •

411Iu 000,III0

IIII •'IIIII III 000

a a'III'

aas

015O'III'ffD0S III

004111liii0 IIh

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•

00

4111

III•

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S

S

saaiaw0sweq Q

speo?J

sai}aw0swag

spooel

N

00+IIM S

00 ,IIIO S+II► S

OQQIDIllO

SS

lip S+III.0S00 'III' 4111llD

000Q

4111^4111''III'0+III

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SS

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i••

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III

IIIM•

Op

110

•

•110

O 0,11)0 0 0

I0 0

lip1110

0111

•

lip I•

• IIII

• I

0metres

RoadsC) Dams

RoadsCD Dams

• 4 al500

metres

FIGURE 5: Maps of Eastwood showing seasonal distribution of whistlers. For these

maps seasons are, in order, autumn 1986, winter, spring, summer, autumn 1987 and

autumn 1988. Relative densities of whistlers are: 0 ( empty circle ), >0-1 ( vertical

lines in circle ), >1-2 ( black grid on white in circle ).

208

0 0 0 • 00 0 0 0 0 0 0

0 0 0 0 0

00 0-400 0 0 0 0 0

0 OD 0 aioo

Roads

CD Dams

Roads

CD Dams0 500

metres0 500

metres

NP ' • • ' 4 TNP . ' " • 4 T

- RoadsCD Dams N• I ■ • ■

500II .1

- Roads

CD Dams N

• 1 al500

Po

0metres

0metres

0 0 0OD 0 0 0oonuoo

00 0 -so 00 0 0 0 0 0000

II)

0 0 0G) 0 0 illo 0 0

0 0 0 0 0

. I ■1

0metres

- RoadsCD Dams N

0 500 T

- RoadsCD Dams

N

• 4 T500

. . . .

metres

FIGURE 6: Maps of Eastwood showing seasonal distribution of Grey Shrike-

thrushes ( GST ), Black-faced Cuckoo-shrikes ( BFCS ) and Crested Shrike-tits

( CrST ). For these maps seasons are, in order, autumn 1986, winter, spring,

summer, autumn 1987 and autumn 1988. Presence of GST indicated by vertical lines

in the circle, that of BFCS by horizontal lines in the circle and that of CrST by

diagonal lines within the circle. Split circles indicates more than one species was seen

at a point in that season.

209

0 a 0.---0 O0 0 0 0

oOD uuoo

NSo ■ ■ ' . 4 T0 500

metres

- Roads

(=) Dams

- Roads

c Dams P • ' ■ . 40

metres500

(ED 0 0 • 00 0 IIIP 0 0 0 0

0 0 0 0

00 0-ID 00 0 f: 0 0 0 @

• 0 CID 0 0 0 •

Roads

CD' DamsN

. Tr • • • .

- Roads

• DamsN

• 1 T500

I . . .

0metres

0 500metres

0 0 , III0 --0 00 0 0 0 0 0 0

0 0 0 0 0

0 0 00000

00 040

Roads

C) DamsN

/ ■ ■ ■ °0 500

metres

Roads

CD Dams I . . . N

• 4 T500 0

metres

FIGURE 7: Maps of Eastwood showing seasonal distribution of aerial feeders, Grey

Fantails and Willie Wagtails. For these maps seasons are, in order, autumn 1986,

winter, spring, summer, autumn 1987 and autumn 1988. Presence of Grey Fantails

indicated by vertical lines in the circle and that of Willie Wagtails by horizontal lines in

the circle.

210

0 0 0 • 00 0 0 0 0 0 0

0 0 0 0 0

RoadsCD Dams

- RoadsC) Dams P-

N

- RI T500 0 500metres

0 metres

N41 TP • • • .

Roads

• Dams0 500

metres

0 0 o---0 e0 0 0 0 1DP 0 0

0 0 0 0 0

Roads

CD Dams0 005

metres

•0 '10 •

0 0

0 0 0---00 0 0 0 0 0

0 0 a 0•

1111►

40III

I000

6 o alp0 0 0o o 000 II 0 0 0

•0

OP

4 Ir,

o

IEY 0

0

4111'

00000

000

0o 0

(ID o o00

• MP

IMI

000

0•III, I

III

II► 0 0

•

. VIIIII

III

- RoadsCD DamsN/ ■ ■ . . 41 T

0 500metres

- RoadsC) Dams / . . . N

• 4 al500 0

metres

FIGURE 8: Map of Eastwood showing seasonal distribution of Eastern Yellow

Robins and Scarlet Robins. For this map, presence of Yellow Robins at a point for

one season indicated by vertical lines in the circle and for >1 season by black grid on

white in a circle. Presence of Scarlet Robins in one season indicated by black

diagonal lines on white circle and for >1 season by white diagonals on black.

211

Roads

CD Dams54000

metres

FIGURE 9: Maps of Eastwood showing seasonal distribution of rosellas. For these

maps seasons are, in order, autumn 1986, winter, spring, summer, autumn 1987 and

autumn 1988. Relative densities of rosellas are: 0 ( blank circle ), >0-1 ( vertical

lines ), >1-2 ( black grid on white circle ), >2-4 ( white grid on black circle ) and >4 (

black circle }.

212

0 0 00 0 0 0 0 0

0000

(5-0---0 0 0 0 0 0

0 0 0 0 0

Roads

CD Dams

Roads

CD Dams I0 500

me tres 0 500

metres

00 0 0 00 0 0 0 0 0 0

0 0

saslaw00S0

...N 1 00Sk N

sealaw0

SWCK) c)

SPQ08 -

SWOa c

SPE/Od

0 0 • ---0 o0 0 0 0 0 0

ooamoo

- Roads

CD Dams P

Roads

CD DamsN■ . .

ometres 900

0metres

500

FIGURE 10: Maps of Eastwood showing seasonal distribution of White-winged

Choughs. For these maps seasons are, in order, autumn 1986 and winter combined,

spring, summer, autumn 1987 and autumn 1988. Relative densities of choughs are: 0

( blank circle ), >0-5 ( vertical lines ), >5-10 ( black grid on white circle ) and >10

( black circle ).

213

0 0 0 0 0 00 0 0 0

0 0 0 0

Roads

'CD Dams ■ ■ ■

- Roads

Dams p • 500 0 500

metres 0

metres

0 0 0'0 00 0 0 0 0 0 0

0 0 0 0 0

- Roads

CD Dams- Roads

CD Dams P 0

metres500 0

metres500

0 0 0-s-c) 00 0 0 0 0 0 0

0 0 0 0 0

NP l II 1

a 4 T

Roads

' Dams0

500metres

FIGURE 11: Maps of Eastwood showing seasonal distribution of Australian

Magpies. For these maps seasons are, in order, autumn 1986 and winter combined,

spring, summer, autumn 1987 and autumn 1988. Relative densities of magpies are: 0

( blank circle ), >0-1 ( vertical lines ), >1-2 ( black grid on white circle ) and >2-4

( white grid on black circle ).

214

0 0 00 0 0 0 0 0 0

0 00

00 0 0 0

0 0 0 0 0

- Roads

C) Dams . . . . N500

RoadsC2) Dams P

n. . . • I

Ae1....) V' V

NI0

metres metres

n: O0 0 00 0 0 0 0 0 0

0 0 0 0I 4111IIO0 0 0

O 110I1111

0o 0

0 0 00

I• I I00I •••i

••••••LP:f

000

0 00 0 00 0 00 0

III

I

• 0 0O•

• •I• II

O Roads

CD DamsRoads

CD Dams0

500 NPO

■

500mPtrPq metres

0 0 00 0

0 0 0 00

0 0 --o0 0 0 0 0 0 e

0 0 0 0 0

Roads

CD Dams P ■ •' . N 4 T0

metres500