Supporting Information
Table S1. Occupancy model pathway descriptions, supporting evidence, and level of confidence for each process contributing to the probability
of a site being within an interspecific exclusion zone (IEZ). High confidence indicates published work supports the existence of the process; Low
confidence indicates the process is plausible based on published work on similar processes or repeated field observations.
Path
Pathway description
way
Evidence & reasoning
Confidence
References
1a
IEZ requires sufficient and temporally
reliable small prey items
High density of birds engaged in
relatively energetically-expensive
territorial behaviour; must be able to
meet energy requirements year-round
within territory
High
Dow, 1977; Ford, 1979; Ford,
1981, 1983; Collins & Paton,
1989; Woinarski et al., 1989;
Newton, 1992
1b
IEZ must be able to be effectively and
efficiently defended against competitors
Competitors may deplete availability of
required prey items and can be
energetically costly to expunge
High
Taylor et al., 2008
1c
Predators of nests and fledged birds
reduce noisy miner (NM) productivity
Sites with high predation pressure from
avian, mammalian and reptilian predators
may produce fewer NMs and breeding
groups may be less able to persist
Low
Arnold, 2000a, b
1d
Sites must be able to be reached by
colonising NMs
Some apparently suitable sites
unoccupied when far from source of
colonists
Mod
R. Major pers. comm.
2a
Sites with dominant tree genera:
Eucalyptus/Corymbia/Angophora and
blade-like leaf shape (as opposed to
needle-like or reduced leaves) preferred
NMs are less likely to be present in sites
where few trees have blade-like leaves;
NMs forage predominantly by gleaning
invertebrates from foliage and such
High
Catterall et al., 1997; Watson et
al., 2000; Major et al., 2001a;
Martin & Catterall, 2001;
Hastings & Beattie, 2006; Maron,
Path
Pathway description
way
Evidence & reasoning
Confidence
leaves have a larger surface area. Sites
dominated by eucalypts and allies more
likely to be occupied
References
2007
3a
High-productivity sites are more likely
to supply sufficient and reliable food
Higher biomass of foliage-dwelling
invertebrates in more-productive sites
High
Majer et al., 1990; Majer et al.,
1992; Majer et al., 1994; Recher
et al., 1996; Majer et al., 1997;
Oldland et al., 2009
3b
Vegetation architecture must facilitate
access to food resources
Dense and stiff or and feathery foliage
may reduce the ability of NMs to access
prey items
Mod
Keast, 1985; Recher et al., 1985;
Ford et al., 1986; Landsberg &
Cork, 1997; Woinarski et al.,
1997
4a
More stable climate results in higher
and more reliable productivity
Productivity is higher and more stable
where rainfall is less variable among and
within years
Mod
Pittock & Nix, 1986; Neave et al.,
1996
4b
Higher insolation increases productivity
Productivity may be higher at lower
latitudes with higher insolation
Mod
Christopherson, 1997
4c
More-fertile soils result in greater
productivity
Higher biomass of invertebrate prey and
higher abundance of NMs in sites on
better/nutrient enriched soils
High
Matson, 1980; Recher et al.,
1996; Keith, 1997; Keatley &
Hudson, 2006; Oldland et al.,
2009
4d
Larger trees are more productive
NM abundance and invertebrate biomass
positively associated with large trees
High
Ashton, 1975; Goldingay, 1990;
House, 1997; Wilson & Bennett,
1999; Kath et al., 2009
5a
Larger trees tend to have denser
Understorey density declines and canopy
canopies and reduce understorey density density increases as density of large trees
increases
High
Specht & Morgan, 2006
Path
Pathway description
way
Evidence & reasoning
Confidence
References
5b
More-productive sites tend to have a
more open, grassy understorey
Grassy woodlands occur on more-fertile
soils and are relatively highly productive
High
Lunt, 1997; McIvor & McIntyre,
2002
5c
Denser understorey reduces
architectural accessibility
Relatively large-bodied species such as
NMs may be less-able to manoeuvre in
dense foliage while foraging
Mod
Butler & Gillings, 2004
5d
Denser understorey reduces
defensibility
Potential competitors or predators are less Mod
visible in sites with high density of
vegetation
Whittingham et al., 2004
6a
Sites are more defensible if close to
open areas, particularly if the site itself
is denser
One or more open areas adjacent to a
High
territory improve visual detectability of
intruders and reduce the frequency of
intruders approaching from that direction.
This is likely to be of relatively greater
benefit in denser vegetation (see 5d)
Catterall et al., 2002; Piper &
Catterall, 2003; Clarke &
Oldland, 2007; Grey, 2008;
Taylor et al., 2008
7a
The more competitors present, the lower Sites accessed by many competitors,
the defensibility of the site
particularly larger-bodied species, require
more effort to defend
Mod
McFarland, 1986b; Mac Nally &
Timewell, 2005
7b
Nectar availability increases the
presence of competitors
Abundance of highly-mobile
nectarivorous bird species, including
large-bodied competitors, is positively
related to nectar availability
Mod
Mac Nally & McGoldrick, 1997;
McGoldrick & Mac Nally, 1998
7c
The more competitors, the less
colonizable is the site
Establishment of a colony requires the
exclusion of most competitors, and this is
likely to be more difficult at sites with
many competitors
Mod
Clarke, 1984; Grey et al., 1997
7d
Sites far from other occupied sites are
Nearby source populations increase the
Mod
Clarke & Schedvin, 1997
Path
Pathway description
way
7e
Evidence & reasoning
less likely to be colonised
chance of colonisation of suitable sites,
although NMs can move large distances
Sites surrounded by large areas of
unsuitable habitat are less colonisable
Movement of potential colonisers may be
facilitated by suitable habitat
Confidence
References
Low
Clarke & Schedvin, 1997
Table S2. Miner effect model pathway descriptions, supporting evidence, and level of confidence for each process. High confidence indicates
published work supports the existence of the process; Low confidence indicates the process is plausible based on published work on similar
processes or repeated field observations.
Path
Pathway description
way
Evidence and reasoning
Confidence
References
1a
Anthropogenic and other disturbance
reduces habitat structural complexity
through common habitat changes
(CHC)
Temperate forests and woodlands have
been extensively modified in the past two
centuries (clearing, logging, thinning
grazing, introductions, and altered fire
regimes). Common habitat changes
(CHC) are a more open canopy, reduced
shrub layer and conversion of continuous
forest to isolated trees or forest fragments
High
Yates & Hobbs, 1997
1b
Noisy miner (NM) abundance responds
positively to most CHC
See IEZ model; NMs more abundant in
more structurally open, grazed sites and
at forest and woodland edges in
fragmented landscapes
High
Loyn, 1987; Clarke et al., 1995;
Ford et al., 1995; Green &
Catterall, 1998; Sewell &
Catterall, 1998; Mac Nally et al.,
2000; Martin & Catterall, 2001;
Catterall et al., 2002; MacDonald
& Kirkpatrick, 2003; Catterall,
2004; Hastings & Beattie, 2006;
Martin et al., 2006; Piper &
Catterall, 2006; Hannah et al.,
2007; Catterall, 2009. See also
IEZ model.
Path
Pathway description
way
Evidence and reasoning
Confidence
References
1c
Smaller nectarivores and insectivores
show more negative than positive
responses to CHC
Many of these species forage in or near
dense foliage – a source of food, nest
sites and protection from predation.
Various assemblage-wide studies show
lower abundance (either individually or
collectively), and species richness, with
CHC
High
Loyn, 1987; Ford & Recher,
1991; Robinson, 1993; Ford et al.,
1995; Catterall et al., 2002;
Catterall & Woinarski, 2003;
Catterall, 2009
1d
Large insectivores/ vertebrate feeders
show more positive than negative
responses to CHC
These species typically nest in taller trees
and feed in more open areas; group
includes several ground-foragers. They
are generally more common in areas
subject to CHC
High
Catterall et al., 2002; Woinarski
& Catterall, 2004; Catterall, 2009
1e
Some large nectarivores show more
positive than negative responses to
CHC.
Open eucalypt woodlands with large trees Low
are associated with more large-bodied
nectarivores
Loyn, 1987; Loyn et al., 2011
2a
NM becomes hyper-aggressive under
CHC
Open canopy and reduced shrubs make it Mod
easier for NMs to locate, target and attack
other birds, potentially resulting in more
frequent attack initiations per miner
Loyn, 1985; Clarke & Oldland,
2007; Taylor et al., 2008; Oldland
et al., 2009
2b
NM is more aggressive when at high
local abundance
Co-occurrence of breeding groups
provides more opportunities for
cooperative attacks; individual birds may
be more likely to initiate attacks when
conspecifics are present
Mod/Low
Grey et al., 1997
2c
Other birds experience more attacks in
sites with high NM abundance
Attack frequency increases because there
are more NMs
High
Piper & Catterall, 2003
Path
Pathway description
way
Evidence and reasoning
Confidence
High
References
2d
Other birds experience more attacks
when NM is in hyper-aggressive mode
Attack frequency increases due to altered
NM behaviour
2e
NM attacks cause decreased local
abundance of small-bodied birds
NM aggression causes reduced local
High
abundance or diversity of other birds
(many birds move away, either on
hearing the NMs or when attacked).
Smaller birds show this negative response
more strongly than larger birds
Dow, 1977; Clarke, 1984; Loyn,
1987; Ford et al., 1995; Grey et
al., 1997, 1998; Catterall et al.,
2002; Piper & Catterall, 2003;
Catterall, 2004; Martin et al.,
2006; Maron et al., 2011; Mac
Nally et al., 2012
2f
Large insectivores/ vertebrate feeders
are more common when NM present
Some large insectivores/ vertebrate
feeders, especially butcherbirds, are
positively associated with NM; causal
mechanisms are unclear but mutual
benefit through cooperative mobbing of
predators is plausible
Mod
Catterall, 2004; Fulton, 2008;
Maron, 2009; M. Maron,
unpublished data.
3a
Abundant large nectarivores in
flowering trees temporarily disrupt
interspecific territorial defence by NMs
Not systematically studied but has been
frequently observed
Low-Mod
D. Oliver (pers obs New South
Wales), A Kutt (pers obs northern
Queensland), M Grey (pers obs
northeast Victoria).
3b
Abundant large nectarivores in
flowering trees cause temporary
decreases in small nectarivore
/insectivore abundance
Size-based aggressive hierarchies often
High
form among honeyeaters within
flowering trees, resulting in fewer smaller
birds
Ford, 1979; Paton, 1985;
McFarland, 1986a; Armstrong,
1991; Mac Nally & Timewell,
2005
4a
Nest predation by NMs reduces
reproductive success of small-bodied
birds
Experiments with artificial nests indicate
that NMs are minor egg predators
Major et al (2001) (Major et al.,
1996; Major et al., 1999, 2001b;
Piper & Catterall, 2004)
Mod
Path
Pathway description
way
Evidence and reasoning
Confidence
References
4b
Nest predation by large insectivores/
vertebrate feeders reduces reproductive
success of small-bodied birds
Butcherbirds, currawongs and corvids are
frequent predators of eggs and nestlings
of smaller birds. Nest predation reduces
seasonal reproductive output even if
parents re-nest; this may contribute to
overall population decline, especially in
sedentary species
Mod
Major et al., 1996; Major et al.,
1999; Fulton & Ford, 2001;
Remes et al., 2012
5 all
Flower visits by nectarivorous birds
increase the success of eucalypt
reproduction (seed set)
Birds are likely pollinators of more than
half of all eucalypt species. Many
eucalypts depend on birds for effective
outcrossing. Birds also pollinate a range
of other tree/ shrub genera in
forest/woodland
High
Ford et al., 1979; Paton & Ford,
1983; House, 1997; Paton et al.,
2004; Phillips et al., 2010
5a
Large nectarivores contribute to
pollination of eucalypt trees
Lorikeets may pollinate in spite of some
Mod
flower destruction. Large honeyeaters
with feeding territories within single trees
assist pollination but are probably
inefficient outcrossers
Paton & Ford, 1983; House, 1997
and references therein
5b
Small nectarivores are the most
effective pollinators of many eucalypt
species
Small honeyeaters make shorter visits to
Mod
more flowers in more trees, creating more
opportunities for pollen transfer
Paton & Ford, 1983; House, 1997
6a
Insectivory by small birds reduces the
abundance of predatory insects
Some small insectivores are likely to
consume predators and parasitoids of
herbivorous insects (e.g., some
Hymenoptera and Diptera)
Gruner, 2004
Low-Mod
Path
Pathway description
way
Evidence and reasoning
Confidence
References
6b
Small insectivore feeding reduces the
abundance of herbivorous insects
The small insectivore guild includes
High
many leaf-gleaners (e.g., small
honeyeaters, pardalotes, silvereyes)
which consume insect herbivores
(Homoptera, Lepidoptera, small
Coleoptera, and other taxa). Experimental
exclosure supports population limitation
by birds
Clark, 1964; Otvos, 1979; Ford,
1983; Loyn et al., 1983;
Woinarski, 1985; Wyndham &
Cannon, 1985; Ford, 1989; Kirk et
al., 1996; Recher et al., 1996;
Woinarski et al., 1997; Clarke &
Schedvin, 1999; Greenberg et al.,
2000; Christie & Hochuli, 2005;
Van Bael & Brawn, 2005; Van
Bael et al., 2008; Morrison &
Lindell, 2012
6c
Predatory invertebrates reduce
abundance of herbivorous insects
Insect predators and parasitoids are
known to limit abundance of insect
herbivores in many ecosystems
de Bach, 1974; Rogers &
Hubbard, 1974; Frazer & Gilbert,
1976; Hassell & May, 1986; New,
1991
6d
Feeding by large numbers of
herbivorous insects leads to tree decline
or death
Associations between extremely high
High
density of herbivorous insects,
defoliation, and “dieback” (partial tree
crown death) have been observed at a
variety of eucalypt forest/ woodland sites.
Repeated outbreak cycles can lead to tree
death
High
Day, 1981; Landsberg & Wylie,
1983; Heatwole & Lowman,
1986; Landsberg et al., 1990;
Landsberg, 1993; Landsberg &
Cork, 1997
Path
Pathway description
way
Evidence and reasoning
Confidence
References
7a
The supply of seed influences trees’
reproductive output
Recruitment of young trees depends in
part on the volume of seed rain, which in
turn depends on pollination success.
Successful reproduction and high
survival of mature trees maintain the
typical floristics and structure of eucalypt
forest/ woodland
Low
Andersen, 1988
7b
Severe herbivory and associated tree
crown dieback can cause tree death
Excessive herbivore-induced defoliation
leads to partial tree crown death. Trees
recover by re-sprouting, but repeated
defoliation cycles exhaust energy
reserves, leading to death
High
Landsberg, 1988, 1990a, b, c;
Ohmart & Edwards, 1991;
Lowman & Heatwole, 1993;
Farrow & Floyd, 1995; Marsh &
Adams, 1995; Stone & Bacon,
1995; Collett, 2001
8
Lowered forest/ woodland “condition”
create positive feedback, causing
increased CHC (and vice versa)
The causal pathways in 1-7 above create
a positive feedback cycle of forest
decline, accompanied by a reduction in
bird diversity, especially of small-bodied
species. Ultimately a hard-to-reverse
transition to a different ecosystem state
may be expected. Its probability of
occurrence would vary in space and time
as other contingencies influence the
operation of each pathway
Mod
Landsberg & Wylie, 1983;
Landsberg, 1988; Jurskis &
Turner, 2002
Path
Pathway description
way
9
Other environmental stressors
associated with human effect feed into
the cycle
Evidence and reasoning
Other human effects resulting from
clearing eucalypt forest and woodland
may also cause decreased tree survival
and reproduction, leading to
complementary positive feedback cycles
(e.g., those involving increased soil
salinity)
Confidence
References
Old et al., 1981; Wylie &
Landsberg, 1987; Landsberg,
1988; Hobbs & Hopkins, 1990;
Landsberg, 1990a, b, c; Old, 2000
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Appendix S1
To evaluate recent trends in the reporting rate of the noisy miner across its range, we analysed data from the BirdLife Australia Atlas dataset. We
used all standard 2 ha, 20 minute surveys conducted from 1998 through 2012 within the range of the noisy miner, yielding presence-absence data
from 69 718 surveys from 51 980 unique sites in 37 bioregions.
We defined the reporting rate as the probability of observing a noisy miner at a specific site in a single survey in a given year:
𝑦𝑖𝑗 ~𝐵𝑖𝑛𝑜𝑚𝑖𝑎𝑙(𝑛𝑖𝑗 , 𝑝𝑖𝑗 ), where yij is the number of surveys at site i in year j that recorded noisy miner as present, nij is the total number of
surveys, and pij is the reporting rate. We used a hierarchical Bayesian model to estimate temporal trends in reporting rates while accounting for
landscape context and other spatial variation. Our measure of landscape context was the distance from the survey point to the nearest habitat
edge (DISTEDGE), where habitat was defined as native vegetation (forest or woodland) as mapped in the National Vegetation Information
System (ESCAVI 2003). Survey locations falling outside of mapped native vegetation were assigned a zero DISTEDGE value.
The model included ‘random slope’ parameters, allowing for variation in temporal trends (and effects of DISTEDGE) among bioregions. The
model also included interactions between time (YEAR) and DISTEDGE, to determine whether temporal trends were dependent on landscape
context. The full model was:
𝑦𝑖𝑗 ~𝐵𝑖𝑛𝑜𝑚𝑖𝑎𝑙(𝑛𝑖𝑗 , 𝑝𝑖𝑗 );
𝑙𝑜𝑔𝑖𝑡(𝑝𝑖𝑗 ) = 𝛼0 + (𝛽𝑟(𝑖) + 𝛿𝑟(𝑖) . 𝐷𝐼𝑆𝑇𝐸𝐷𝐺𝐸𝑖 ) × 𝑌𝐸𝐴𝑅𝑗 + 𝛾𝑟(𝑖) × 𝐷𝐼𝑆𝑇𝐸𝐷𝐺𝐸𝑖 + 𝜀𝑖𝑗 ;
1
1
1
); 𝛿𝑟 ~𝑁(∆, 𝜎𝑖𝑛𝑡
); 𝛾𝑟 ~𝑁(Γ, 𝜎𝑑𝑖𝑠𝑡
);
𝛽𝑟 ~𝑁(Β, 𝜎𝑡𝑟𝑒𝑛𝑑
𝜀𝑖𝑗 = 𝑟𝑒𝑔𝑖𝑜𝑛𝑟(𝑖) + 𝑠𝑢𝑏. 𝑟𝑒𝑔𝑖𝑜𝑛𝑠(𝑖) + 𝑠𝑢𝑏. 𝑟𝑒𝑔. 𝑐𝑎𝑟𝑠(𝑖) + 𝑠𝑖𝑡𝑒𝑖 + 𝑦𝑒𝑎𝑟𝑗 + 𝑠𝑖𝑡𝑒. 𝑦𝑒𝑎𝑟𝑖𝑗 ;
Here, 𝛼0 is the overall intercept, 𝛽𝑟 is the temporal trend for bioregion r (r(i) indicates the bioregion that site i falls within), 𝛾𝑟 is the regionspecific effect of DISTEDGE (landscape context), and 𝛿𝑟 is the region-specific interaction between YEAR and DISTEDGE. Thus, (𝛽𝑟(𝑖) +
𝛿𝑟(𝑖) . 𝐷𝐼𝑆𝑇𝐸𝐷𝐺𝐸𝑖 ) yields a point-specific trend, which is a function of the bioregion trend 𝛽𝑟 and site-specific landscape context. The regionspecific coefficients were modelled hierarchically, with exchangeable normal prior distributions and group means (Β, ∆, Γ) assigned independent
normal prior distributions, N(0,1000). The model included spatial and temporal random intercepts, with nested spatial effects corresponding to
region, subregion and site. Subregion-level random effects were partitioned into separate random (𝑠𝑢𝑏. 𝑟𝑒𝑔𝑖𝑜𝑛𝑠 ) and spatially structured
(𝑠𝑢𝑏. 𝑟𝑒𝑔. 𝑐𝑎𝑟𝑠 ) components. The spatially structured components were modelled using conditional autoregressive prior distributions (Besagand
and Kooperberg 1995). All other spatial random effects, including the site × year components, were assigned exchangeable normal prior
distributions, N(0, 2);  ~ Uniform(0,10). The year random effects were modelled with a first-order autoregressive model (Fahrmeir and Lang
2001).
Models were fitted by Markov chain Monte Carlo (MCMC) using WinBUGS software (Lunn et al. 2000). Posterior distributions were estimated
using 3 concurrent MCMC chains of 100 000 iterations each, following 50 000 iteration ‘burn-in’ periods, with every 15th iteration retained to
reduce autocorrelation. Chain history plots and Gelman-Brooks-Ruben statistics confirmed adequate MCMC mixing and convergence.
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