Avian functional group responses to rainfall across four

vegetation types in the Simpson Desert, central Australia

MAX TISCHLER,
1,2
* CHRISTOPHER R. DICKMAN
2,3
AND GLENDA M. WARDLE
2,3
1
Science and Monitoring, Bush Heritage Australia, PO Box 1090 Byron Bay, NSW 2481, Australia
(Email: mtischler@bushheritage.org.au),
2
Desert Ecology Research Group, School of Biological Sciences,
University of Sydney, Sydney, New SouthWales, and
3
Long-term Ecological Research Network,
Multi-Scale Plot Network, Terrestrial Ecosystem Research Network, LTERN, St Lucia, Queensland,
Australia
Abstract Temporal and spatial variability in rainfall and subsequent resource ows often drive large changes in
bird demography and the structure of avian assemblages in arid regions. Here, we describe broad avian functional
group responses to rainfall across four vegetation classes in the Simpson Desert, central Australia, over a period
when both dry and wet conditions prevailed. Birds were surveyed in eucalypt woodlands, gidyea woodlands, mixed
Acacia (mulga) shrublands and spinifex-dominated hummock grasslands in this region between 2006 and 2008
using a spatially nested hierarchical sampling design with xed radius point counts as replicates. Predictably, the
composition of bird communities varied across vegetation classes as well as over time. Generalist feeders and
insectivorous birds dominated assemblages during dry periods, whereas granivorous species were most abundant
and widespread following heavy rains. Carnivores and specialist nectarivores responded variably throughout the
sampling period regardless of habitat. The dynamics of granivores were especially interesting; large numbers of
nomadic species moved into the study area in response to rainfall, highlighting a key role for this group in shaping
community structure. This study represents the rst attempt to investigate avian functional group dynamics across
differing vegetation types in a variable but low productivity landscape. The ndings conrm that community
composition is spatially and temporally dynamic, and prompt further investigation into the roles that habitat and
resource limitation play in dening bird communities in arid Australia.
Key words: central Australia, desert bird, functional group, rainfall.
INTRODUCTION
One of the enduring questions in ecology is to under-
stand the factors that drive the assembly of local com-
munities from the taxa available in the broader species
pool (Tilman & Kareiva 1997; Lawton 1999; Hubbell
2005). Equally compelling is to ask whether patterns
in species composition and functional groups are con-
sistent over multiple scales of space and time. Birds are
ideal for answering such questions as their diversity,
mobility and iconic migrations (Kingsford & Norman
2002) make them highly responsive to ephemeral
resource shifts in the environment (Wiens 1989).
The search to meet energy and moisture require-
ments is a key imperative for avifauna in many habi-
tats, especially those in arid regions (Serventy 1971).
Both resources vary spatially and temporally, primarily
with respect to rainfall (Noy-Meir 1973; Whitford
2002), and their availability is linked further to
patterns of vegetative response (Schwinning & Sala
2004). Consequently, birds need to cope with, and
track, dynamic and often ephemeral resources contex-
tually within extended periods of low abundance
and during unpredictable pulses of high abundance
(Serventy 1971; Dickman & Tischler 2010; Letnic &
Dickman 2010). As such, there is much evidence that
resource availability acts in a regulatory manner in
habitat selection by birds (e.g. Pulliam & Mills 1977;
Morton & Davies 1983). Birds are often common and
ubiquitous components of faunal diversity in arid envi-
ronments (Dean 2004) and, while some specialist bird
species can maintain residency during lean periods
(Maclean 1996), the mobility of many others allows
them to move large distances in search of more favour-
able conditions (Davies 1983). Arid zone bird com-
munities therefore tend to be temporally dynamic
assemblages, often with a mix of persistent or resident
species and a high proportion of nomadic species
whose presence and distribution are dictated by pulsed
resource events. This is especially so in arid regions
of inland Australia where the proportions of nomads
(or partial nomads) comprise 3046% of breeding bird
species (Dean 2004).
Such high gures for nomads have prompted debate
about the strength and repeatability of patterns within
bird communities in arid Australia, with the roles
played by habitat selection and resource availability
*Corresponding author.
Accepted for publication May 2013.
Austral Ecology (2013) 38, 809819
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2013 The Authors doi:10.1111/aec.12065
Austral Ecology 2013 Ecological Society of Australia
forming pivotal points in the discussion (Pavey &
Nano 2009). As rainfall is acknowledged to be the
dominant factor driving arid Australian ecosystems
(Morton et al. 2011), investigating bird responses to
exceptional rain events across diverse vegetation types
has the potential to shed light on the roles these factors
play in shaping bird communities in variable, low
productivity landscapes.
While single species dynamics can be useful in
exploring bird responses to limiting conditions (e.g.
Marone 1991), there has long been support for group-
ing species based on ecological similarities or niche
parameters (Root 1967; Cody 1974). By assigning
bird species to functional groups or guilds a better
understanding of community structure and function,
as well as how animals utilize their environment, can
be gained (Simberloff & Dayan 1991; Marone 1992;
Recher & Davis 1997). Furthermore, identifying envi-
ronmental components that are important to many
bird species, as well as the potential response of a large
proportion of those species to perturbation, can guide
conservation management decisions (Mac Nally et al.
2008).
Here, we use broadly dened functional groups to
investigate bird responses to rainfall across a spatially
variable desert landscape. The structure and com-
position of vegetation often strongly inuence arid
bird communities (Rotenberry 1985; Marone 1991;
Marone et al. 1997; Paltridge & Southgate 2001), and
recent work in central Australia suggests that rainfall
induces signicant changes in species richness, com-
munity composition and abundance (Burbidge &
Fuller 2007). However, predictable patterns can still
exist across a range of vegetation types under such
conditions (Pavey & Nano 2009). Hence, investigating
how functional groups are distributed across the
landscape with respect to vegetation type, and their
responses to episodic rainfall events, may assist in
developing a better understanding of the factors that
shape desert bird communities. This is important as
the need for effective management and conservation of
biodiversity in arid regions is becoming more critical
under projected climatic models and intensied land
use (Reynolds et al. 2007), especially in Australia
(Morton et al. 2011; Greenville et al. 2012).
In this study we surveyed birds across four distinct
vegetation types in the north-eastern Simpson Desert
in 2006, 2007 and 2008. As signicant rain fell across
the study area at the start of 2007 we were able to
investigate avian responses to rainfall following a dry
year (2006), and then again a year after the rainfall
event in 2008. We predicted that functional groups
would be evenly distributed regardless of vegetation
type, although each group would contain unique
assemblages that respond differently to rainfall. We
tested this hypothesis by designating species as gener-
alist foragers, insectivores, granivores, nectarivores,
carnivores or other, and then examined avian
responses to both dry and wet conditions according to
vegetation type.
METHODS
Study area
The study area is located across two conservation reserves
situated near the north-eastern margin of the Simpson
Desert, in Queensland, Australia (Fig. 1). These are
Ethabuka (2345S, 13828E) and Cravens Peak (2316S,
13817E).The study area is bounded to the north, south and
east by the Mulligan River oodplain, to the west by the Field
River and Gnallan-a-gea Creek, and lies along a north-south
rainfall gradient between the 150 mm and 100 mm median
annual rainfall isopleths (Purdie 1984). Rainfall is spatially
patchy with a subtle rainfall gradient across the study region;
northern areas receive higher annual rainfall than those in
the south (Haythornthwaite & Dickman 2006). The mean
annual temperature is 2123C with maxima of 4649C in
summer and minima of 6C in winter (Purdie 1984).
The dominant landforms in the region are sandplains and
dune elds that are characterized by sand ridges 810 m high
and spaced 1001000 m apart. The sand ridges run NNW-
SSE. Landforms punctuating the dune elds are expanses of
clay ats, often topped with gravel or gibber. Several drainage
lines run through the area, with numerous ood-outs and
claypans turning into ephemeral water bodies after rain.
This study focused on recording birds in four of the most
distinct and dominant vegetation types: eucalypt woodlands
conned to the riparian zones and fringing plains of the Field
River and Gnallan-a-gea catchment; gidyea woodlands
within the inter-dune corridors or swales of the dune elds;
mixed Acacia (mulga) shrublands on scattered sandplains;
and hummock (spinifex) grasslands dominated by Triodia
basedowii and associated with sand ridges.
Study design
We surveyed birds using a spatially nested hierarchical
design. A total of 16 sites was established across the study
area, each representing one of the four vegetation classes;
this provided a total of four sites within each class (Fig. 1).
Within each site four replicate points, spaced independently
at 300-m intervals on a straight-line trajectory, were used to
survey birds using the point-count method with xed radius
(100 m). On each sampling occasion two observers recorded
all birds seen or heard at each point within a 15-min period
over two consecutive days. Surveys took place over six sam-
pling periods: March, June and October 2006; March, June
2007; and June 2008.
Environmental conditions
Before this study began weather conditions were typically
dry, with ve consecutive years experiencing average or
810 M. TISCHLER ET AL.
2013 The Authors doi:10.1111/aec.12065
Austral Ecology 2013 Ecological Society of Australia
below average rainfall in the range of 56199 mm per
annum. Average conditions were experienced in 2006, with
Ethabuka and Cravens Peak Reserves receiving 141 mm and
143 mm, respectively. In early 2007 the region received wide-
spread above average rainfall (>300 mm) with 224 mm and
214 mm at Ethabuka and Cravens Peak falling over just
several days. This rainfall resulted in a substantial amount of
free-standing water and a consequent resource pulse across
the landscape; it also restricted our sampling to only half the
study sites in March and June that year. Follow-up rain
occurred in March (1660 mm), but conditions remained
dry for the rest of the year. Despite patchy summer rain in
late 2007 (442 mm), conditions continued to dry in 2008,
with only 13 mm and 22 mm falling in August at Ethabuka
and Cravens Peak, respectively.
Data analysis
Bird abundance was estimated using the mean count of each
recorded species over the two sampling days, giving a mean
species abundance per point count per site within the four
habitat types. This approach was used to ensure independ-
ence of each replicate count, as resident birds could conceiv-
ably be counted twice over both days.
We then classied all birds into broad functional groups:
generalists, insectivores, granivores, nectarivores, carnivores
and others. This gave total bird abundance in each func-
tional group at each site per sampling period. Classication
of species into functional groups was based on expert knowl-
edge of foraging behaviours of birds in the study area, and
veried by the literature (e.g. Barker & Vestjens 1990).
Fig. 1. Map of the study area in the north-eastern Simpson Desert showing the distribution of sites used to survey birds in
2006, 2007 and 2008.
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Although the groups are coarse and further partitioning of
each functional category may be possible, we used them
to investigate broad patterns and used individual species to
elucidate ner scale dynamics over the course of the study.
To investigate differences in functional groups between
and within each vegetation class, for each sampling period,
data were rst log (x + 1) transformed to reduce the effects of
highly abundant ocking species and then analysed
and plotted using non-metric multi-dimensional scaling
(nMDS). We used BrayCurtis similarity coefcients for the
resemblance matrix with 50 iterations to congure minimum
stress in the nMDS plots (Anderson et al. 2008). Two-way
nested analyses of similarities (ANOSIM) were used to iden-
tify differences between vegetation types and sites, except
in March and June 2007 when only one-way analysis was
possible, before making pairwise comparisons between veg-
etation types in the resemblance matrix. When signicant,
SIMPER analyses were then used to identify the functional
guild and main species contributing to similarities and dif-
ferences between groups. All analyses were carried out using
PRIMER (V6.1.11) (Anderson et al. 2008).
RESULTS
Overall, 83 species of birds were identied and 8303
individuals counted from 2999 observations during
the study. The number of species in each func-
tional group was: insectivores (29), generalists (19),
granivores (13), carnivores (raptors) (14), specialist
nectarivores (5), and three others (including two
piscivores and one frugivore) (Appendix S1). More
species were detected in the eucalypt woodlands
than the other three vegetation types; insectivores and
generalists were the most prevalent functional
group across all classes (Table 1), although this shifted
during the study in response to rainfall.
Generalists and insectivores dominated the counts
in 2006 (Table 2), producing signicant yet inconsist-
ent differences both within and across the four vege-
tation types (Fig. 2, Table 3). SIMPER analysis of
functional groups produced dissimilarities ranging
from 50.4% to 70.9% in March, and 53.8% to 60.4%
in October for all signicant pairwise comparisons.
However, SIMPER analysis of species produced dis-
similarities >91% in March and >87.3% in October
in all pairwise comparisons. Overall, large percentage
contributions were made by generalist species such
as singing honeyeaters (Lichenostomus virescens) and
white-plumed honeyeaters (Lichenostomus penicillatus),
and insectivorous species such as masked wood-
swallows (Artamus personatus), white-backed swallows
(Cheramoeca leucosterna) and crimson chats
(Epthianura tricolor) (Table 4). In October, the arrival
of rainbow bee-eaters (Merops ornatus) and white-
winged trillers (Lalage sueurii), both seasonal migrants,
Table 1. Number of species detected in each functional group across the four vegetation types
Functional group Eucalypt woodlands Gidyea woodlands Mulga shrublands Spinifex grasslands
Generalists 15 11 9 13
Insectivores 20 22 19 20
Granivores 11 6 9 7
Nectarivores 5 1 2 1
Carnivores 11 6 5 5
Other 2 1 2 0
TOTAL 64 47 46 46
Table 2. Top ranked SIMPER results for functional group contributions (%) for similarities between each vegetation type
during sampling periods
Eucalypt woodlands Gidyea woodlands Mulga shrublands Spinifex grasslands
Mar 06 Gen (45.2), Gran (31.7),
Ins (17.8)
Ins (69.6), Gen (16.7),
Gran (12.2)
Ins (76.1), Gen (12.5),
Gran (10.1)
Gen (52.9), Ins (41.1)
Jun 06 Gen (50.7), Ins (46.9) Gen (46.5), Ins (41.8),
Gran (11.7)
Gen (55.8), Ins (41.9) Gen (46.9), Ins (44.3)
Oct 06 Gen (55.8), Ins (34.7) Ins (88.5), Gen (5.8) Ins (76), Gen (21.9) Ins (65.8), Gen (19.9),
Gran (14.3)
Mar 07 Gran (36.7), Gen (33.3),
Ins (14.9), Car (14.2)
Ins (49), Gran (39.4),
Gen (10)
Gran (48.7), Ins (41.9) Gran (49.9), Ins (24.5),
Gen (14.9), Car (10.7)
Jun 07 Gen (51), Gran (38.7),
Ins (8.3)
Gran (49.5), Ins (28.6),
Gen (20.1)
Gran (66), Ins (18.3),
Gen (12.8)
Gran (69.4), Ins (19.5),
Gen (11.1)
Jun 08 Gen (74.4), Ins (20.6) Gran (43.9), Ins (34.7),
Gen (21.4)
Ins (65.1), Gen (28.3) Gen (64.5), Ins (24.1),
Gran (11.2)
Car, carnivores; Gen, generalists; Gran, granivores; Ins, insectivores.
812 M. TISCHLER ET AL.
2013 The Authors doi:10.1111/aec.12065
Austral Ecology 2013 Ecological Society of Australia
further strengthened the dominance of insectivores
and generalist species throughout. The prevalence of
generalists and insectivores was not unexpected as
prevailing dry conditions saw very little vegetative
growth or oristic activity that would otherwise
provide favourable foraging conditions for granivores
or nectarivores. Some granivores were recorded in
2006, with zebra nch (Taeniopygia guttata) the most
common species (Table 4). As zebra nches are resi-
dent in the study area they were expected to occur
across most vegetation types, albeit in low numbers.
Following the signicant rainfall event of early 2007
a large inux of birds moved into the study area to
exploit free-standing water in ephemeral water bodies
and a ush of owering and seeding plants. This
resulted in clear differences across all four vegetation
types, except in pairwise comparisons between the
spinifex and mulga (Fig. 2, Table 3). Not surprisingly,
granivores dominated many of the counts in March,
with most records attributable to the arrival of large
numbers of nomadic species not previously observed
in the study. SIMPER analysis of functional groups
a)
b)
c)
d)
e)
f)
2D Stress: 0.18
2D Stress: 0.16
2D Stress: 0.17
2D Stress: 0.15
2D Stress: 0.17
2D Stress: 0.14
Fig. 2. Non-metric multi-dimensional scaling (nMDS) plots of functional groups across four vegetation types- eucalypt
woodlands (), gidyea woodlands (X), mulga shrublands (+) and, spinifex grasslands (O) in (a) March 2006, (b) June 2006,
(c) October 2006, (d) March 2007, (e) June 2007 and, (f) June 2008.
Table 3. Summary of signicant ANOSIM results (in bold) for each sampling period
Date
Vegetation type Site
Signicant pairwise tests Global R P Global R P
Mar 06 0.503 <0.01 0.22 <0.01 All
Jun 06 0.102 0.15 0.259 <0.01
Oct 06 0.371 0.04 0.269 <0.01 Euc/Gid, Euc/Mul, Euc/Spin
Mar 07 0.418 <0.01 All except Mul/Spin
Jun 07 0.175 <0.01 Euc/Gid, Euc/Mul, Euc/Spin
Jun 08 0.21 0.04 0.289 <0.01 Euc/Gid, Euc/Mul
Euc, eucalypt woodlands; Gid, gidyea woodlands; Mul, mulga shrublands; Spin, spinifex grasslands.
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produced dissimilarities of <36.3% in all pairwise
comparisons, while analysis of species produced dis-
similarities ranging from 54.7% to 74.8%. Diamond
doves (Geopelia cuneata) were the most notable
granivore species to arrive and made large contribu-
tions across the four vegetation types (Table 4).
Despite the arrival of large numbers of granivores,
many insectivorous and generalist species also made
large percentage contributions throughout the survey
period (Table 2).
As conditions dried towards June 2007 the strongest
differences detected were between the eucalypt wood-
lands and the other three vegetation classes (Fig. 2,
Table 3). Granivore numbers remained high, although
the percentage contributions of generalists and insec-
tivores were also considerable (Table 2). SIMPER
analysis of functional groups produced dissimilarities
ranging from 39.647.1%, although analysis of species
produced dissimilarities >90.7% in all pairwise com-
parisons with the eucalypt woodlands. Zebra nches
were the most abundant and widespread granivore
during this time, with budgerigars (Melopsittacus
undulatus) common in the spinifex grassland habitat
and diamond dove numbers dramatically reduced and
conned more to the eucalypt woodlands (Table 4).
The landscape returned to widespread dry con-
ditions by June 2008; differences in bird species
composition between vegetation types were then less
pronounced, with the strongest differences again exist-
ing between the eucalypt woodlands and the other
three vegetation classes (Fig. 2, Table 3). Generalists
and insectivores once again dominated (Table 2),
although high contributions by zebra nches boosted
the granivore contribution in the gidyea and spinifex
Table 4. Top ranked SIMPER results for species contributions (>10%) for similarities between each vegetation type during
sampling periods
Eucalypt woodlands Gidyea woodlands Mulga shrublands Spinifex grasslands
Mar 06 White-plumed
Honeyeater (54.7)
Masked Woodswallow (43.2) White-backed Swallow
(38.1)
Singing Honeyeater
(41.7)
Zebra Finch (10.6) Crimson Chat (22.3) Singing Honeyeater
(21.8)
Variegated Fairy-wren
(13.6)
Singing Honeyeater (11.6) Zebra Finch (20.3) Black-faced Woodswallow
(12.1)
Variegated Fairy-wren
(13)
Grey-headed Honeyeater
(10.9)
Jun 06 White-plumed
Honeyeater (38.2)
Singing Honeyeater (46.7) Singing Honeyeater
(70.1)
Singing Honeyeater
(39.4)
Fairy Martin (32) Zebra Finch (19.5) Variegated Fairy-wren
(13.5)
Zebra Finch (14.2)
White-winged Fairy-wren
(11)
Oct 06 White-plumed
Honeyeater (29.2)
Rainbow Bee-eater (24.2) White-browed Babbler
(38.4)
White-backed Swallow
(27)
White-winged Triller
(17)
Black-faced Woodswallow
(20.5)
Singing Honeyeater
(26.6)
Zebra Finch (20.5)
Spiny-cheeked
Honeyeater (10.3)
Crimson Chat (14.8) Crested Bellbird (15.2) Rainbow Bee-eater
(18.4)
White-browed Babbler
(13.5)
White-backed Swallow
(13.9)
Variegated Fairy-wren
(15.9)
Mar 07 White-plumed
Honeyeater (27.6)
Masked Woodswallow (33.3) Diamond Dove (31.4) Diamond Dove (37.4)
Diamond Dove (26.5) Crimson Chat (23) Crimson Chat (22.7) Zebra Finch (24.3)
Diamond Dove (17.5) Zebra Finch (19.1) Masked Woodswallow
(10.5)
Zebra Finch (10.4)
Jun 07 White-plumed
Honeyeater (42.9)
Zebra Finch (54.3) Zebra Finch (83.38) Zebra Finch (51.2)
Diamond Dove (38) Singing Honeyeater (18.1) Budgerigar (21.4)
Jun 08 White-plumed
Honeyeater (73.2)
Zebra Finch (24.4) Black-faced Woodswallow
(33.6)
Singing Honeyeater
(36.4)
Crested Pigeon (17.5) Singing Honeyeater
(29.6)
Grey-headed Honeyeater
(19.4)
Singing Honeyeater (14.6) Zebra Finch (13.1) Zebra Finch (12.7)
Black-faced Woodswallow
(11.6)
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Austral Ecology 2013 Ecological Society of Australia
habitats, especially (Table 4). SIMPER analysis of
functional groups produced dissimilarities of 49.6%
and 54.8%, and analysis of species >92.1% for all
signicant pairwise comparisons with the eucalypt
woodlands.
Carnivores rarely contributed to the analysis of simi-
larities between vegetation groups during dry condi-
tions, but did show a response to rainfall during March
2007 in the eucalypt woodlands and spinifex grasslands
(Table 2). Black kite (Milvus migrans) was the only
species with a noticeable increase during this period,
with SIMPERanalysis producing percentage contribu-
tions of 58% similarities in the eucalypt and spinifex
associations, as well as the mulga, increasing the overall
contribution of carnivores during this period (Table 2).
Most other raptor species sighted during the study were
individuals, and despite some such as brown falcon
(Falco berigora) and nankeen kestrel (Falco cenchroides)
being common across all four vegetation types, their
low contribution to counts may be explained by overall
low abundances throughout the study.
Nectarivores were in similarly low numbers through-
out the study. Grey-headed honeyeaters (Lichenostomus
keartlandi) were the most common species recorded,
especially at those spinifex sites containing stands
of owering mallee (Eucalyptus pachyphylla and
E. gamophylla) and corkwood (Hakea eyreana)
(Table 4). Surveys in March and June 2007, hindered
by restricted access, failed to record any response by
this species to rainfall. Overall, most nectarivore species
were recorded in the eucalypt woodlands (Table 1,
Appendix S1), with nomadic honeyeaters such as
pied (Certhionyx variegatus), black (Sugomel niger),
white-fronted (Purnella albifrons) and black-chinned
(Melithreptus gularis) honeyeaters observed in low
numbers, and only when isolated owerings of peren-
nials such as mistletoe and Eremophila spp. were noted.
DISCUSSION
We chose four distinct and replicated vegetation types
in this study as a novel approach to investigating the
dynamics and composition of avian communities over
a spatially variable desert landscape, and our results
clearly show differential temporal responses by birds
across that landscape. Rainfall appeared to be a key
driver of community change. Generalist and insectivo-
rous species dominated during dry conditions in 2006
and 2008, but granivores arrived following a heavy and
widespread rainfall event in 2007 and remained preva-
lent for some months thereafter.
We expected low dissimilarity scores for functional
groups across all vegetation types as most bird commu-
nities have proportional niche opportunities (Wiens
1989). Our results support this, with all functional
groups represented in each vegetation type. Composi-
tional shifts within the vegetation types occurred over
time in response to both rainfall and probably other
more discrete resource dynamics that were not meas-
ured.The high dissimilarity scores for species contribu-
tions offer further support to this pattern, but highlight
the variability that bird communities can display within
vegetation types and across landscapes. Despite this,
dened communities were still detected throughout the
study, especially in the eucalypt woodlands, often in
the gidyea, but only occasionally in the mulga and
spinifex.This suggests that at times predictable patterns
still exist within bird communities regardless of species
turnover the ooding of most communities by
granivores following rain supports this notion, at least
in the two woodland vegetation classes. This has im-
plications for how bird utilize their environment, as
inconsistent variability across vegetation types through
time implies dynamic and episodic resource shifts, with
habitat selection decisions needing to be made accord-
ingly. If the woodland habitats provide the most pre-
dictable habitats for birds we might assume that these
most likely act as dry-period refugia.
Functional groups across vegetation types
The differences in species richness and community
composition between the vegetation types over time
were not unexpected, as past studies have demon-
strated predictable assemblage patterns of association
with vegetation types in arid Australia (Cody 1994;
Recher & Davis 1997; Pavey & Nano 2009). It was
within the eucalypt woodlands that the most dened
patterns were recorded, and these were responsible for
many of the differences detected in the analyses. Tall
woodlands, especially those associated with riverine
habitats, are well known to harbour a higher species
richness and abundance of birds than surrounding
areas (Shurcliff 1980; Mac Nally et al. 2008), espe-
cially where water availability is limited across the
landscape (Schneider & Griesser 2009). Our results
support the utility of these areas as dry-period refugia.
Generalists and insectivorous species, especially those
with high dietary plasticity, dominated counts within
the more structurally complex eucalypt woodlands
when conditions were dry. In October 2006 the arrival
of seasonal migrants into the study area strengthened
this pattern, with high contributions by white-winged
trillers and rainbow bee-eaters (8.4%). Unfortunately,
surveys were not conducted in October 2007, prevent-
ing a temporal comparison of seasonal migrant distri-
butions following rainfall.
In a similar result to that of Pavey and Nano (2009),
white-plumed honeyeaters were the dominant species
in the eucalypt woodlands. They were consistently the
most common species recorded in this vegetation
type, and were rarely observed in the gidyea, mulga or
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Austral Ecology 2013 Ecological Society of Australia
spinifex. This species takes advantage of the canopy
strata in Eucalyptus-dominated woodlands where it
feeds on a variety of foods like insects (especially sugar-
secreting lerps) andnectar (Ford&Paton1976).White-
plumed honeyeaters are gregarious birds and, typical of
their congeners in more temperate areas, tend to avoid
moment-to-moment competition by excluding poten-
tial competitors through aggression (Piper & Catterall
2003).The dominance of white-plumed honeyeaters at
some sites correlated with the absence of most species
of small birds (<50 g). For example, other small foliage-
dependent species such as red-browed pardalotes
(Pardalotus rubricatus) and weebills (Smicrornis
brevirostris) were recorded only rarely. In contrast,
common species in the eucalypt woodland sites like
zebra nches and willie wagtails (Rhipidura leucophrys)
avoid competition with white-plumed honeyeaters by
utilizing different microhabitats, thus avoiding niche
overlap. In cases where niche overlap potentially
exists, most rivals like spiny-cheeked honeyeaters
(Acanthagenys rufogularis) are substantially larger and
are less threatened by the aggression of the smaller
honeyeater (Ford 1979).
Functional group patterns in gidyea woodlands were
variable. There was a high degree of species turnover,
and habitat specicity at any time was difcult to
detect. Aerial and ground foraging insectivores such as
woodswallows and crimson chats were the dominant
species most of the time; however, resident granivores
such as zebra nches were recorded frequently,
as well as generalists like singing honeyeaters. High
numbers of nomadic masked woodswallows and resi-
dent black-faced woodswallows (Artamus cinereus)
were commonly sighted. It is not unusual for a number
of woodswallow species to co-occur despite them
sharing similar ecological niches and exhibiting little
obvious segregation (Loyn 2002). The sparse open
canopies of gidyea woodland suit these birds as they
often perch on dead tree limbs and hawk or pounce
on insects.The lack of any distinct shrub canopy layers
in much of the gidyea favoured ground foraging
species such as crimson chats, but surprisingly did not
appear to reduce the incidence of shrub-dwelling
insectivorous species such as variegated fairy-wrens
(Malurus lamberti) or generalist crested bellbirds
(Oreoica gutturalis).
An unexpected result was that despite quite obvious
structural and oristic differences between the mulga
shrublands and spinifex grasslands the bird communi-
ties in each were remarkably similar. They generally
contained low numbers of generalist and insectivorous
species such as singing honeyeaters, white-backed
swallows, variegated fairy-wrens and black-faced
woodswallows, as well as the granivorous zebra nch.
It was not surprising then that they occupied much of
the same space in the ordinations, but intrinsic vari-
ability and shared species, with each other and with the
gidyea, explain why denitive patterns were rarely
found across these three vegetation types.
The mulga sites were spaced at considerable dis-
tances from each other in the study area (Fig. 1);
consequently the sites, although supercially similar,
were variable enough to often contain different bird
species. It is likely that differences in the composition
of subdominant plant species, substrate, or a combi-
nation of habitat attributes might explain this. Some
endemic species, for example cinnamon quail thrush
(Cinclosoma cinnamomeum) were recorded only at
mulga sites with a stony substrate, a preferred habitat
for this species (Ford 1970). Similar patterns were also
found for other common species such as white-browed
babblers (Pomatostomus superciliosus). As such, intrinsic
variability might explain the overall patterns detected
for this vegetation type. However, that many sites often
recorded nil counts suggests that mulga shrublands
overall are marginal habitats for birds in the study area.
This result is in contrast to past studies in mulga
shrublands (e.g. Reid et al. 1993; Cody 1994; Recher
& Davis 1997; Pavey & Nano 2009). These past
studies have focused on large contiguous stands of
mulga whereas our study area contains more discrete
and sometimes isolated groves typical of the eastern
Simpson Desert. The size and extent of mulga
shrublands might explain some differences between
our results and those obtained further west in central
Australia and elsewhere.
Similarly, variability in bird abundance and species
richness between spinifex sites might explain the lack
of dened communities in this vegetation type. As
noted, the presence of owering mallee at the two
northern sites resulted in frequent sightings of grey-
headed honeyeaters (Lichenostomus keartlandi). This
species actively seeks mallee and corkwood blossoms
(Ford & Paton 1976), and all records in this vegetation
type during the study were associated with these trees.
Despite this, it was still the most commonly recorded
nectarivore throughout the study, and a common
sighting in this vegetation type.
Functional group responses to rainfall
Episodic rainfall events are the principal drivers
of many ecological processes in arid landscapes
(Schwinning & Sala 2004), and the response of
animals to rain-induced resource pulses can shape the
structure of their populations and the communities to
which they belong (DSouza et al. 2013; Greenville
et al. 2013; Pavey & Nano 2013). In our study we
recorded patterns across the landscape during dry
periods, as described, and then again following a sig-
nicant and widespread rain event. Our results show
a clear shift in the functional composition of bird
communities in response to rainfall.
816 M. TISCHLER ET AL.
2013 The Authors doi:10.1111/aec.12065
Austral Ecology 2013 Ecological Society of Australia
The most notable response was the inux of large
numbers of new granivores into the study area follow-
ing rainfall in early 2007.The arrival of diamond doves
to all four vegetation types contributed to the overall
dominance of granivores during this period. Diamond
doves are free-ranging endemic arid zone specialists,
possessing a range of physiological and behavioural
traits that adapt them well to arid conditions
(Schleucher 1993). Flocks roam inland Australia
seeking favourable foraging conditions, and exemplify
the peripatetic nature of much of Australias arid avi-
fauna (Davies 1983). In June 2007, 5 months after rain,
this species was almost completely absent frommuch of
the study site, although we did nd reasonable numbers
in the eucalypt woodlands. At this time, waterholes
along the riverine habitat still provided free-standing
water in an otherwise drying landscape. By June 2008
no diamond doves were sighted in the study area. Our
results conrmthemas opportunistic nomads continu-
ally searching for favourable conditions, and reafrm
the eucalypt woodlands as important dry-period
refugia. Similarly, lownumbers of other granivores were
recorded for the rst time in March 2007 in all vegeta-
tion types. Little button-quail (Turnix velox), painted
nches (Emblema pictum) and vagrant pictorella man-
nikins (Heteromunia pectoralis) all arrived in the study
area following rain but had left it by June 2007.
Zebra nches increased in numbers following rain,
although as a resident species it was unclear initially
whether immigration from elsewhere or opportunistic
breeding was responsible. Zebra nches have the
ability to respond quickly to rainfall, with breeding
events synchronized with the most protable stages
of seed maturation (Zann et al. 1995). Persistently
high numbers of this species recorded in both March
(only weeks after rain) and June 2007 suggest that
both strategies were in evidence. By contrast, many
other resident granivores such as galah (Eolophus
rosiecapillus) and crested pigeon (Ocyphaps lophotes)
did not show marked increases after rain.
Granivory is one of the dominant trophic strategies
used by nomadic birds in arid lands worldwide (Dean
2004). Seeds are diverse and sometimes abundant,
providing a nutritious albeit variable resource for
birds (Morton & Davies 1983; Reichman 1984). The
disadvantage of this foraging strategy is the need to access
free-standing water (MacMillen 1990). Some specialist
granivore species, such as the zebra nch, have the physi-
ological ability to form and use metabolic water (Zann
1996), but most need to travel frequently and often over
large distances to meet their moisture requirements
(MacMillen 1990). Our results are consistent with mois-
ture being a limiting factor for granivorous birds, with the
presence and distribution of many species appearing to be
dictated by the availability of water.
Insectivores also increased across the study area fol-
lowing rain, but it was two species that were common
during dry conditions masked woodswallows
and crimson chats that contributed most. Others
species recorded following rain were also common
during dry conditions, including seasonal migrants
such as rainbow bee-eaters and brown songlarks
(Cincloramphus cruralis). Of particular interest was that
these species were not conned to one vegetation type
as was evident throughout 2006, but recorded in
higher numbers across all vegetation classes. Our
results suggest that the response of this functional
group to rain is to emigrate from dry-period refugia
where they persist in low numbers year-round to more
marginal habitat when conditions become more
favourable, perhaps following episodic irruptions of
preferred food items (e.g. Farrow & McDonald 1987).
The relative paucity of nectarivorous birds through-
out the study, and their lack of response to rainfall,
was unexpected.We had anticipated a similar pattern to
that found by Burbidge and Fuller (2007) with at least
some rain-responsive species being detected in high
numbers.The patchy, asynchronous nature of owering
by many Australian arid-dwelling plants has led to a
high degree of nomadism in nectarivorous bird species
(Keast 1968). Some birds do respond opportunistically
to rain-induced owering events while others track
more predictable seasonal episodes of owering (Ford
&Paton 1976). For example, two species of honeyeater,
both detected in this study in low numbers the black
and pied are highly nomadic and depend largely
on the owering of species within several perennial
plant genera, including Eremophila, Grevillea and Hakea
(Ford 1979; Schodde 1982). Similar patterns of speci-
city are common in more mesic environments else-
where in Australia (McGoldrick & Mac Nally 1998),
with patterns of behaviour and competition regulated
by nectar availability that likely reect patterns in arid
regions (Dean 2004). As the seasonal timing and extent
of rainfall can inuence owering events in arid lands
(Schwinning et al. 2003) it may be that many of the
preferred nectar-producing plants did not ower
during the study, which in turn offers an explanation for
the lack of response by nectarivores.
Dietary plasticity, as a means of coping with highly
variable food resources and avoiding competition, is
also a common feature of nectarivorous birds, with
many species using a range of nectar sources and
including insects in their diet (Ford 1979; Schodde
1982). Some species of the Melaphagidae, categorized
as generalists in this study, use this adaptation;
examples are spiny-cheeked honeyeaters and yellow-
throated miners (Manoria avigula). These species
at times feed on nectar (Keast 1968) and the contri-
bution of nectarivores may have been higher had
these species been so classied. Regardless, generalist
species overall did not display notable responses to
rainfall either. Discounting the continued domin-
ance of white-plumed honeyeaters in the eucalypt
BIRD RESPONSES TO RAINFALL 817
2013 The Authors doi:10.1111/aec.12065
Austral Ecology 2013 Ecological Society of Australia
woodlands, other birds such as singing honeyeaters all
but disappeared from counts in March and June 2007.
A common feature of arid land raptors is the move-
ment of birds into areas where prey has become abun-
dant following rain, often leading to synchronous
breeding events (Galushin 1974; Dean 2004). The
movement and increase in numbers of letter-wing kites
(Elanus scriptus) in the Lake Eyre Basin, in response to
irruptions of the long-haired rat (Rattus villosissimus),
is a good example of this dynamic (Hollands 1979;
Pavey et al. 2008). We found black kites to be the only
species to respond to rain, and even then their nume-
rical contribution was low. By June 2007 they had
moved out of the study area, suggesting that the
resource pulse following rain had not been sufcient to
sustain a localized population irruption for them or
any other raptor species.
The results of this study show that broadly dened
functional groups can be used to elucidate patterns in
avian community dynamics in spatially and temporally
variable landscapes. Despite high species turnover we
were still able to describe community composition
during dry periods and after a signicant rainfall
event. Of particular interest was the movement of
nomadic birds, most notably granivores, into and
around the landscape following rain. This result dem-
onstrates the dynamics of arid-land birds and high-
lights resource limitation as a key driver in habitat
selection, prompting further investigation into the role
this factor plays in shaping communities. We did not
measure resource abundance or availability directly,
nor did we record breeding activity and success, and
acknowledge that these would be useful descriptors
in any future studies. Long-term studies are required
to continue investigating bird responses to dry and
wet conditions, subsequent resource pulses, as well as
associated determinants such as competition, preda-
tion and variations under differing land usage.
ACKNOWLEDGEMENTS
MT was supported by an Australian Postgraduate
Award and funding support was provided by the
Australian Research Council. During the preparation
of the manuscript GW and CD received support
from the Australian Governments Terrestrial Ecosy-
stems Research Network (http://www.tern.gov.au), an
Australian research infrastructure facility established
under the National Collaborative Research Infrastruc-
ture Strategy and Education Infrastructure Fund
Super Science Initiative through the Department of
Industry, Innovation, Science, Research and Tertiary
Education. We are grateful to the landowners and
managers of the reserves and properties on which we
conduct our research and would like to thank Gordon
and Col McDonald, Jo and Len Rule, Mark and Nella
Lithgow all formerly from Cravens Peak Reserve;
Al Dermer and Karen Harland, Scott Morrison and
Sajidah Abdullah all formerly from Ethabuka Reserve;
and Greg, Shae, Dean and Emma Woods from Carlo
Station, for their continued support and hospitality.
We thank the reviewers and editors for their many
suggestions to improve the manuscript.
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SUPPORTING INFORMATION
Additional Supporting Information may be found in
the online version of this article at the publishers
web-site:
Appendix S1. List of bird species recorded during
the study in each vegetation type.
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