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Article

The Bird Assemblage of the Darwin Region (Australia): What Is the Effect of Twenty Years of Increasing Urbanisation?

by
Sarah E. Fischer
1,*,
Andrew C. Edwards
1,
Patrice Weber
1,
Stephen T. Garnett
2 and
Timothy G. Whiteside
3
1
Research Institute for the Environment and Livelihoods, College of Engineering, IT & Environment, Charles Darwin University, Darwin, NT 0909, Australia
2
College of Engineering, IT & Environment, EIE Environment, Charles Darwin University, Darwin, NT 0909, Australia
3
Environmental Research Institute of the Supervising Scientist, Darwin, NT 0820, Australia
*
Author to whom correspondence should be addressed.
Diversity 2021, 13(7), 294; https://doi.org/10.3390/d13070294
Submission received: 30 April 2021 / Revised: 14 June 2021 / Accepted: 21 June 2021 / Published: 28 June 2021

Abstract

:
There has been considerable urban development in the Darwin region over the last twenty years; as for most fauna in Australia since colonisation, the potential effects to the bird assemblage were expected to be disastrous. To provide a broad overview of changes, bird survey data from 1998 and 2018 were extracted from BirdLife Australia’s ‘Atlas of Australian Birds’ database. A total of 165 species were categorised into primary food source feeding guilds and levels of food specialisation. This was integrated into ArcGIS along with land use change mapping from 1998 and 2018 to investigate its impact on bird assemblages. There was no significant change in overall species numbers when all sites were analysed. However, when sites were separated into those with increased urbanisation or decreased greenspace, several sites showed a significant change in the number of species. For the majority of species, analysis of primary food types found no difference in the proportion of species within the assemblages between 1998 and 2018, regardless of the level of urbanisation or greenspace; the exception being those species that primarily feed on insects, where the difference was just significant. An analysis using bird community data sorted into levels of food specialisation also found no difference between 1998 and 2018 despite habitat changes. These findings suggest that although there has been considerable urban development in the Darwin region, bird communities are remaining relatively stable.

1. Introduction

Global population trends show that humans are moving from rural areas into cities at a considerable rate, and once sparsely populated regions are being transformed to cope with the influx of people [1,2]. Subsequently, urbanisation is now widely considered a major threat to biodiversity conservation [3,4,5].
Species’ responses to urbanisation can vary significantly [6]; however, prior to the 1990s, very few ecological studies were undertaken in urban areas due to them being considered unviable habitats for faunal populations and, ergo, immaterial from a conservation standpoint [7]. It is now shown that urban areas support an array of species that have been able to tolerate or adapt to the highly fragmented new environments but, due to a lack of appropriate knowledge, the success of urban conservation programmes may be hampered [4]. There is little doubt that avian assemblages are greatly altered by urbanisation; many bird species unable to adapt to the suddenly changed environment will often move out of an area resulting in diminished biodiversity and allowing invasive species to move in [4,8,9]. Other species can benefit from urbanisation; these are often birds with a more generalist diet and life-history traits that are conducive to living in a fragmented habitat [10].
Whilst studies of bird populations in urban areas abound in the temperate zones of both the Northern and Southern hemispheres, research into tropical urban avian communities is scant [11]. This trend is continued in Australian studies with the majority of research being undertaken in the temperate regions, predominantly in the two heavily populated states of New South Wales and Victoria. In contrast, this study is situated in the monsoonal tropics of northern Australia; specifically, in Darwin, the capital of the Northern Territory (NT). There has been targeted research on a range of species and thorough overviews of bird distributions within the region [12,13,14], with more detailed studies of shorebirds [15,16,17,18] and mangrove assemblages [19,20,21]. To date, however, there have been no studies of trends in terrestrial bird assemblages as the city has grown. Given the increased rate of urbanisation in the global tropics [22], coupled with high levels of biodiversity in the zone [11,23,24,25], there is an increasing need to gather data to better understand how bird communities are coping with the rising encroachment of human habitation. This is particularly interesting in Darwin as, unlike all other urban centres in Australia, Darwin has no introduced birds, so all the adaptation is being undertaken by native species as their environment is changed.
In the nearly 45 years since Darwin was severely damaged by Cyclone Tracy, where 70–80% of dwellings were destroyed [26,27], the city has grown from a perceived ‘frontier town’ to be a modern capital city. This development has seen an increase of over 20 new suburbs in the greater Darwin area and surrounds, and land once considered bush or rural properties is now being subdivided into urban blocks. One of the most common effects of urbanisation is the increasing prevalence of exotic species [28,29,30]; however, this has not occurred in the Darwin region with only four species listed as ‘foreign invaders’, none of which have established permanent breeding populations [31].
In this paper, we shall test this theory by investigating the response of the terrestrial avian assemblages in the Darwin region to urban expansion and land use change over the past 20 years.

2. Materials and Methods

2.1. Study Area

Darwin (12.4634° S, 130.8456° E) is situated on the north coast of Australia, a landscape dominated by tropical savanna (Figure 1).
The population of approximately 140,000 [32] constitutes an increase of nearly 60,000 people in twenty years [33]. The climate is monsoonal and experiences distinct annual dry (May to September) and wet (November to April) seasons with transitional periods in between. Mean annual rainfall is approximately 1700 mm; the mean minimum and maximum temperatures range from 19.3° to 25.3° and 30.6° to 33.3°, respectively [34]. Compared to other major Australian capital cities, housing density is low; under 20 private dwellings per square kilometre as opposed to between 150 to 200 per square kilometre in Sydney and Melbourne [35]. A combination of urban and periurban environments in the Darwin region provides resources for avian populations that are typically unavailable outside of this area in the dry season.

2.2. Spatial Data

To investigate broad-scale changes in urbanisation, Landsat satellite imagery of the study area from April 1998 and 2018 was obtained from the United States Geological Survey’s Global Visualization Viewer (GloVis), with 1998 imagery obtained from the Landsat 4 and 5 Thematic Mapper (TM) satellite and 2018 imagery taken by the Landsat 8 Operational Land Imager (OLI) and Thermal Infrared Sensor (TIRS) (Table A1 and Table A2). April marks the end of the wet season in Darwin, allowing for images with minimal cloud cover and maximum vegetation growth. Images were clipped in ArcGIS version 10.4.1 [36] to a shapefile of Darwin region localities provided by the Northern Territory Government Department of Environment and Natural Resources and then imported into SAGA GIS version 7.3.0 [37]. Classification of land use types was carried out via geographic object-based image analysis (GEOBIA). In traditional pixel-based image classification, classes are assigned per pixel; however, GEOBIA uses segmentation and classification to better replicate what the human eye perceives [38,39]. The segmentation process combines pixels of similar spectral properties into objects in the form of polygons, and these can then be classified using either supervised or unsupervised techniques [40]. After automated segmentation in SAGA GIS, the two clipped images were classified by assigning ‘training sites’ (essentially selecting a minimum number of polygons and ascribing them a land use type) and then running a supervised classification. The resulting vector layers were then manually edited using the original satellite image to reassign any misclassified polygons. The initial uncorrected GEOBIA and user-corrected images were then re-imported back into ArcGIS, where fifty accuracy assessment points were randomly generated and an error matrix was constructed to assess both the producer (SAGA) and user (human) accuracy when assigning classification. For all map classifications satellite imagery, aerial photographs and Google Earth Pro version 7.3.2.5776 (64 bit) were used to assist in the accuracy assessment; however, due to the retrospective nature of the earlier imagery, only the 2018 images could be further checked, if required, using ground control points.

2.3. Bird Survey Data

Survey data were extracted from the BirdLife Australia ‘Atlas of Australian Birds’ database (hereafter referred to as the ‘Bird Atlas’) for the years 1998 and 2018. Several types of surveys compiled these data: systematic bird surveys of 2 ha, 5 km and 500 m; unstandardised bird surveys, either along a fixed route or incidental; bird list surveys and the Shorebird 2020 surveys (a record of shorebird sightings not necessarily in coastal habitats). All records include a location, latitude and longitude, dates and species common names. In most records, a time is recorded and whether there is any breeding activity. Sighting notes of interest are sometimes included.
As the focus of the project was on terrestrial, predominantly diurnal species, Bird Atlas records were excluded if the species was almost exclusively nocturnal, was a waterbird or seabird (except Magpie Geese, Anseranas semipalmata), or the species was considered ‘vagrant’. Using information from BirdLife Australia [41], the Atlas of Living Australia [42] and Australian Bird Data Version 1.0 [43], the feeding preferences of species were categorised from most preferred to occasional.
To give a general overview of any assemblage changes, the records of 1998 and 2018 were categorised into the following primary food sources: fruit, insects, invertebrates, nectar, omnivore, raptor, scavenger, seed, vegetation or vertebrate. If a species was considered to feed on two types of food source equally, both were considered the primary food source. Species were sorted by their level of specialisation: whether they had one, two, three or more food sources (Table A3).

2.4. Integration of Data

Following Hahs and McDonnell [44] and Conole and Kirkpatrick [30], the final edited spatial images were re-imported into ArcGIS and a 1 × 1 km grid was overlaid. The modified Bird Atlas data from 1998 and 2018 were then added as point layer files. As with Conole and Kirkpatrick [30], the locations were taken from the coordinates provided. The grid cells that contained records from both 1998 and 2018 were extracted for each year, and the level of land use type in each grid cell was calculated. Land use types were combined and simplified into greenspace (woodland, grass and forest), coastal (mangrove, sea and sand), urbanised (suburban, periurban, building and road), water and bare earth. Percentage change of greenspace and urbanisation between 1998 and 2018 was calculated for the relevant grid cells.

2.5. Statistical Analysis

Data were analysed using R version 3.6.1 [45].
Analysis of variance (ANOVA) was used to investigate whether the site and year had an effect on overall species numbers; paired t-tests were then used to examine the significance of any changes between 1998 and 2018 in those sites where urbanisation had increased or greenspace had decreased. The analysis was repeated to assess species’ primary food source categories.
Finally, effects of land change on feeding specialisation numbers in bird assemblages were also explored. To do this, assemblage species proportions of one, two, three or more food sources were analysed for grid cells that were determined to have had an increase in urbanisation, a decrease in urbanisation, no change in the amount of urbanisation, an increase in greenspace or a decrease in greenspace.

3. Results

3.1. Spatial Data

Results of the GEOBIA found that urbanised areas increased from 114 to 151 km2, a percentage increase of almost 3% of the total land mass measured. Greenspace, too, increased by just under 5%; 986 to 1045 km2.
For the 1998 imagery, overall accuracy was 75% (producer accuracy 72%; user accuracy 84%). The 2018 imagery returned an overall accuracy of 87% (84% producer and 90% user accuracies). This difference in accuracy was expected due to the quality differences in the Landsat 5 versus Landsat 8 imagery. Final corrections were made to the classified imagery before extraction of the relevant land use data for analysis (Figure 2).

3.2. Bird Atlas Data

Overall, there were 23 grid cells that contained Bird Atlas records from both 1998 and 2018. Of these, nine showed an increase in urbanisation ranging from 0.9 to 58.2%; eight indicated a decrease in urbanisation and six were recorded as no change. A decrease in greenspace was found in 15 grid cells, and 8 were determined to have had an increase. No species considered primarily a scavenger was found in the 23 grid cells.

3.3. Overall Change in Species Numbers

In total, 165 individual species were represented in the bird records used. Of these, 23 species were recorded in 1998 only and 19 in 2018 only (Table A4). All sites showed a change in species numbers between 1998 and 2018; however, when all sites were included, there was no significant change in total species numbers (F = 1.1909, p = 0.3999 and F = 1.9395, p = 0.1707, respectively).
Focussing on those sites where urbanisation had increased or greenspace had decreased over the 20-year period, changes in species numbers were more distinctive. Although sites are listed numerically, this is for brevity, and they are not necessarily the same location.
Where urbanisation had increased, of the nine sites analysed, five sites showed an increase in overall species numbers with four of these increases found to be statistically significant (p < 0.05). Examination of sites where species numbers decreased found two where the decrease was significant and two where there was no statistical significance (Table 1).
In sites where there was a decrease in greenspace, the majority (10 out of 15) displayed an increase in species numbers, but of these, only three were found to be significant. The remaining five sites where a decrease in species was recorded again found three with statistical significance (Table 2).

3.4. Changes to Species Numbers with Regard to Primary Food Sources

Species for each site were categorised by their primary food source, and changes in numbers initially were record as either an increase, decrease or no change for each food source.
In areas where urbanisation had increased, site 6 (increase in urbanisation of 5.8%) showed an increase in every species type, whereas site 4, which had the highest increase in urbanisation of any site (58.2%), displayed a decrease in almost every species type bar one, in which there was no change (Table 3).
Subsequent analysis found that there was no significant change in species numbers by primary food source overall (Table 4).
When investigating those sites where greenspace had decreased, one site (7) showed an increase in every species type despite having a greater reduction in greenspace (14.2%) than either site 6 (13.4%) or site 8 (5.3%), both of which showed a reduction in every species type except those that feed primarily on vegetation (Table 4).
Unlike those sites where urbanisation had increased, analysis of changes in species numbers in the sites where greenspace had decreased found a significant change in those species whose primary food source was insects (p < 0.05). Some change was noted for raptors too, but this was not quite statistically significant (p = 0.06) (Table 5).

3.5. Changes to Bird Assemblages with Regard to Primary Food Sources

Alongside changes to the number of species within bird communities over time, it was also pertinent to investigate whether the proportion of feeding guild types was altered within assemblages due to changing habitats, specifically in those sites where urbanisation increased or greenspace decreased.
A comparison of the proportions of species type for each site where urbanisation was found to have increased is shown in Figure 3.
Subsequent analysis showed an increase in urbanisation resulted in no significant difference in the proportion of species of different primary food sources between 1998 and 2018.
However, in grid cells that had a decreased amount of greenspace, a slight significant difference was shown in those species that chiefly feed on insects, and there is a suggestion of change in those species that primarily feed on fruit, although this was not found to be significant (Table 6).
A comparison of species proportion by primary food source for those grid cell sites where greenspace had decreased over the 20-year period is shown in Figure 4.

3.6. Levels of Specialisation

The levels of feeding specialisation within bird communities between 1998 and 2018 showed no significant change despite varying levels of urbanisation and greenspace (Table 7).

4. Discussion

Of the 258 birds species recorded in the Darwin area (escaped or introduced species and vagrants excluded), approximately 40% are considered resident [46] with the remaining species described as mobile. It has previously been thought that mobile species cope better with urbanisation than sedentary species [47,48], although other studies have suggested that highly mobile species may be more vulnerable to habitat changes due to their dependence on larger habitat patches and generally far-reaching home ranges [49,50,51]. Furthermore, there is also evidence to suggest that the fragmentation of habitats caused by urbanisation is going some way to changing the status of some birds from ‘visitor’ to ‘resident’ [52,53,54].
That there is a significant difference, albeit slight, in the proportion of those species that primarily feed on insects in areas where greenspace has decreased may possibly be attributed to the establishment of urban gardens. Although the size of the greenspace has decreased overall in these locations, in 73% of the grid cells analysed, the proportion of species that preferred insects increased; in contrast, only 47% of grid cells contained an increase in fructivores. It may be that the plants in these gardens attract a larger array of insects, thus providing more appealing spaces for insectivores. Numerous studies have indicated that the bird species that tend to thrive in urban environments are those that nest in cavities or canopies (sites less likely to be disturbed by human activity) and are more likely to be granivorous or omnivorous, whereas those species that avoid urban areas are those that predominantly nest in shrubs or trees or at ground level and have a more specialised diet, frequently insectivores [29,30,55]. However, these characteristics have been determined from research undertaken in either the Northern Hemisphere or the temperate zones of the Southern Hemisphere, places where garden vegetation is often vastly different to that of the original habitat. Suburban gardens of the Darwin region often contain native plant species alongside exotics, and this mixture may be enough to maintain, or even increase, insect numbers.
That there is no significant difference in the levels of feeding specialisation in bird assemblages is something of a surprise, as it would be fair to assume that more generalised species would be increasing and those species with one preferred food source would be forced out, particularly in areas of urban increase. Homogenisation of habitat is an oft-quoted result of urbanisation [5,56,57], but from this broad overview, it appears not to be affecting the composition of bird communities in the Darwin region. Further research into the characteristics of species making up urban assemblages will go some way to better understanding why this may be.
Invasion by feral species is a frequent result of an increased human presence in an area [29,30,58]. Many bird species that are unable to adapt to the suddenly changed environment will often move out of an area resulting in diminished biodiversity and allowing invasive species to move in [4,8,9]. Other species can benefit from urbanisation; these are often birds with a more generalist diet and life-history traits that are conducive to living in a fragmented habitat [10].
The only feral species in the records was Columba livia—the common pigeon, rock dove or rock pigeon. This species was recorded in the 1998 records only, most likely due to an eradication programme undertaken by the Parks and Wildlife Commission, NT, beginning in 1996, that saw the species considered eradicated from the Darwin area by 2004 [59,60]. The lack of established feral bird populations in the Darwin region could, potentially, suggest that native bird species are filling the ‘urban exploiter’ or ‘suburban adapter’ niche that is commonly filled by exotic species in other cities, but there is little or no research to show what these species may be. Additionally, a unique aspect of the Darwin region is that it is the only capital in Australia where urban fires are a regular annual occurrence. The city is located within the monsoonal tropics; a landscape dominated by tropical savannas, experiencing frequent fires during the prolonged dry season from May to October, often within just metres of suburban housing areas [61]. In addition to wildfires, many of these fires are prescribed, being undertaken by various land-management bodies as part of a yearly landscape management approach [62,63]. Whether these annual fires are in some way supporting and maintaining the local bird assemblages whilst preventing invasive species from establishing populations is an intriguing question. Research into the combined impact of urban fires and human modification of the environment on local bird populations over time is ongoing; it is hoped that the results will go some way towards answering this question.

5. Conclusions

Whilst very transient, the population of Darwin and its surrounds has grown significantly between the years 1998 and 2018, and urbanisation has expanded more than 50 km from the city centre. Despite this, this general overview of bird assemblages shows that although species numbers fluctuate, avian communities appear to be stable. The increase in the insectivorous species may be due to the establishment, maturity and vegetative composition of suburban gardens. This may potentially signify a shift in the species diversity of the area; however, further research is needed to investigate this.

Author Contributions

S.E.F. conceived, designed and performed the experiments. S.E.F. analysed the data. S.E.F. and A.C.E. wrote the paper. P.W., S.T.G. and T.G.W. provided guidance and comments on the manuscript. All authors have read and agreed to the published version of the manuscript.

Funding

This study is part of a larger Ph.D. project that is supported by Charles Darwin University and the Australian Government via the Research Training Programme scholarship.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The authors confirm that the data supporting the findings of this study are available within the article, its Appendices, and Australian Bird Data Version 1.0 (https://doi.org/10.6084/m9.figshare.1499292_D8, access date 10 September 2019) at http://www.birdsaustralia.com.au/ (access date 19 June 2019) and https://www.ala.org.au/ (access date 1 September 2019).

Acknowledgments

The authors would like to thank Andrew Silcox and BirdLife Australia for access to bird data records.

Conflicts of Interest

The authors declare no conflict of interest.

Appendix A. Landsat Satellite Specifications

Table A1. Landsat 4 and Landsat 5 Thematic Mapper (TM) 1998 imagery (Data Taken from USGS [64,65]).
Table A1. Landsat 4 and Landsat 5 Thematic Mapper (TM) 1998 imagery (Data Taken from USGS [64,65]).
BandWavelength (Micrometres)Resolution (Metres)
1—Blue0.45–0.5230
2—Green0.52–0.6030
3—Red0.63–0.6930
4—Near Infrared0.76–0.9030
5—Short-wave Infrared1.55–1.7530
6—Thermal Infrared10.40–12.50120 (30) *
7—Short-wave Infrared2.08–2.3530
* Band 6 has thermal infrared resolution of 120 metres but is re-sampled to 30-metre pixels.
Table A2. Landsat 8 and 9 Operational Land Imager (OLI) and Thermal Infrared Sensor (TIRS) 2018 imagery (Data Taken from USGS [64,65]).
Table A2. Landsat 8 and 9 Operational Land Imager (OLI) and Thermal Infrared Sensor (TIRS) 2018 imagery (Data Taken from USGS [64,65]).
BandWavelength (Micrometres)Resolution (Metres)
1—Coastal aerosol0.43–0.4530
2—Blue0.45–0.5130
3—Green0.53–0.5930
4—Red0.64–0.6730
5—Near Infrared (NIR)0.85–0.8830
6—SWIR 11.57–1.6530
7—SWIR 22.11–2.2930
8—Panchromatic0.50–0.6815
9—Cirrus1.36–1.3830
10—Thermal Infrared (TIRS) 110.6–11.19100
11—Thermal Infrared (TIRS) 211.50–12.51100

Appendix B. Bird Species

Table A3. Total bird species recorded.
Table A3. Total bird species recorded.
Species NameYear RecordedPrimary Food SourceNumber of Food Sources
19982018
Australasian Figbird (Sphecotheres vieilloti ashbyi)Fruit2
Australian Magpie (Cracticus tibicen)XOmnivore3+
Australian Pied Oystercatcher (Haematopus longirostris)Invertebrates2
Australian White Ibis (Threskiornis moluccus)Omnivore3+
Azure Kingfisher (Ceyx azureus ruficollaris)XInvertebrates2
Bar-breasted Honeyeater (Ramsayornis fasciatus)XNectar/Insects2
Bar-shouldered Dove (Geopelia humeralis)Seed1
Bar-tailed Godwit (Limosa lapponica)Invertebrates3+
Beach Stone-curlew (Esacus magnirostris)XInvertebrates1
Black Bittern (Ixobrychus flavicollis)Vertebrates1
Black Butcherbird (Cracticus quoyi)Vertebrates3+
Black Kite (Milvus migrans)Raptor2
Black-faced Cuckoo-shrike (Coracina novaehollandiae melanops)Invertebrates3+
Black-faced Woodswallow (Artamus cinereus melanops)Insects1
Black-fronted Dotterel (Elseyornis melanops)XInvertebrates2
Black-necked Stork (Ephippiorhynchus asiaticus)Vertebrates2
Black-shouldered Kite (Elanus axillaris)XRaptor2
Black-tailed Godwit (Limosa limosa)Invertebrates3+
Black-winged Stilt (Himantopus himantopus)Invertebrates2
Blue-faced Honeyeater (Entomyzon cyanotis albipennis)Insects3+
Blue-winged Kookaburra (Dacelo leachii leachii)Vertebrates/Invertebrates2
Brahminy Kite (Haliastur indus)Raptor2
19982018
Broad-billed Flycatcher (Myiagra ruficollis)XInsects2
Broad-billed Sandpiper (Limicola falcinellus)XInvertebrates2
Brolga (Grus rubicunda)Omnivore3+
Brown Falcon (Falco berigora)XRaptor1
Brown Goshawk (Accipiter fasciatus didimus)Raptor1
Brown Honeyeater (Lichmera indistincta indistincta)Nectar2
Brown Quail (Coturnix ypsilophora)Seed3+
Brown-capped Emerald-Dove (Chalcophaps longirostris)Seed2
Brush Cuckoo (Cacomantis variolosus)Insects1
Buff-banded Rail (Hypotaenidia philippensis)XOmnivore3+
Bush Stone-curlew (Burhinus grallarius)XInvertebrates2
Cattle Egret (Ardea ibis)XInsects3+
Channel-billed Cuckoo (Scythrops novaehollandiae)XFruit3+
Chestnut Rail (Eulabeornis castaneoventris)Invertebrates1
Chestnut-breasted Mannikin (Lonchura castaneothorax)Seed2
Cicadabird (Coracina tenuirostris)XInsects2
Collared Kingfisher (Todiramphus chloris sordidus)Invertebrates2
Common Greenshank (Tringa nebularia)Insects3+
Common Sandpiper (Actitis hypoleucos)Insects2
Crimson Finch (Neochmia phaeton phaeton)Seed2
Curlew Sandpiper (Calidris ferruginea)Insects2
Dollarbird (Eurystomus orientalis)Insects1
Double-barred Finch (Taeniopygia bichenovii annulosa)Seed2
Dusky Honeyeater (Myzomela obscura obscura)Nectar2
Eastern Curlew (Numenius madagascariensis)Invertebrates1
19982018
Eastern Koel (Eudynamys orientalis subcyanocephala)Fruit3+
Eastern Reef Egret (Egretta sacra)Vertebrates/Invertebrates2
Eurasian Coot (Fulica atra)XOmnivore3+
Forest Kingfisher (Todiramphus macleayii macleayii)Invertebrates2
Galah (Eolophus roseicapilla)Seed1
Glossy Ibis (Plegadis falcinellus)Invertebrates2
Golden-headed Cisticola (Cisticola exilis)Insects2
Great Bowerbird (Ptilonorhynchus nuchalis nuchalis)XOmnivore3+
Great Egret (Ardea modesta)Vertebrates2
Great Knot (Calidris tenuirostris)Invertebrates1
Greater Sand Plover (Charadrius leschenaultii)Invertebrates2
Green-backed Gerygone (Gerygone chloronota)Insects1
Grey Butcherbird (Cracticus torquatus colletti)XInvertebrates3+
Grey Plover (Pluvialis squatarola)Insects3+
Grey Shrike-thrush (Colluricincla harmonica)Omnivore3+
Grey Whistler (Pachycephala simplex simplex)Insects1
Grey-crowned Babbler (Pomatostomus temporalis rubeculus)Invertebrates3+
Grey-tailed Tattler (Tringa brevipes)Invertebrates1
Helmeted Friarbird (Philemon buceroides ammitophila)Nectar3+
Horsfield’s Bushlark (Mirafra javanica)Seed2
Intermediate Egret (Ardea intermedia)Vertebrates1
Jacky Winter (Microeca fascinans pallida)XInsects1
Large-billed Gerygone (Gerygone magnirostris magnirostris)Insects1
Large-tailed Nightjar (Caprimulgus macrurus)XInsects1
Leaden Flycatcher (Myiagra rubecula concinna)Insects1
19982018
Lemon-bellied Flycatcher (Microeca flavigaster flavigaster)Insects1
Lesser Sand Plover (Charadrius mongolus)Invertebrates1
Little Bronze-cuckoo (Chalcites minutillus minutillus)Insects1
Little Corella (Cacatua sanguinea sanguinea)Seed2
Little Curlew (Numenius minutus)Insects3+
Little Egret (Egretta garzetta)Invertebrates2
Little Friarbird (Philemon citreogularis sordidus)Nectar3+
Little Kingfisher (Ceyx pusilla ramsayi)XVertebrates/Invertebrates2
Little Shrike-thrush (Colluricincla megarhyncha parvula)Insects2
Long-tailed Finch (Poephila acuticauda hecki)Insects1
Long-toed Stint (Calidris subminuta)XInsects3+
Magpie Goose (Anseranas semipalmata)Seed2
Magpie-lark (Grallina cyanoleuca neglecta)Insects2
Mangrove Gerygone (Gerygone levigaster levigaster)Insects1
Mangrove Golden Whistler (Pachycephala melanura melanura)XInsects1
Mangrove Grey Fantail (Rhipidura phasiana)XInsects1
Marsh Sandpiper (Tringa stagnatilis)Invertebrates2
Masked Finch (Poephila personata personata)XSeed2
Masked Lapwing (Vanellus miles miles)Insects2
Mistletoebird (Dicaeum hirundinaceum)Fruit2
Nankeen Kestrel (Falco cenchroides)XRaptor2
Nankeen Night-heron (Nycticorax caledonicus)XVertebrates2
Northern Fantail (Rhipidura rufiventris)Insects1
Olive-backed Oriole (Oriolus sagittatus affinis)Fruit3+
Orange-footed Scrubfowl (Megapodius reinwardt)Insects3+
19982018
Oriental Plover (Charadrius veredus)Insects1
Osprey (Pandion cristatus)Raptor1
Pacific Golden Plover (Pluvialis fulva)Insects3+
Peaceful Dove (Geopelia striata placida)Seed2
Pectoral Sandpiper (Calidris melanotos)XInsects1
Pheasant Coucal (Centropus phasianinus melanurus)Invertebrates2
Pied Butcherbird (Cracticus nigrogularis picatus)XInvertebrates3+
Pied Heron (Egretta picata)Vertebrates3+
Purple Swamphen (Porphyrio porphyrio melanotus)XVertebrates/Invertebrates3+
Rainbow Bee-eater (Merops ornatus)Insects1
Rainbow Pitta (Pitta iris iris)Invertebrates2
Red Knot (Calidris canutus)Insects3+
Red-backed Button-quail (Turnix maculosus)XSeed2
Red-backed Fairy-wren (Malurus melanocephalus cruentatus)XInsects1
Red-backed Kingfisher (Todiramphus pyrrhopygius)Vertebrates2
Red-capped Plover (Charadrius ruficapillus)Invertebrates2
Red-collared Lorikeet (Trichoglossus haematodus rubritorquis)Vegetation3+
Red-headed Honeyeater (Myzomela erythrocephala erythrocephala)Nectar2
Red-kneed Dotterel (Erythrogonys cinctus)XInvertebrates2
Red-necked Stint (Calidris ruficollis)Omnivore3+
Red-tailed Black-Cockatoo (Calyptorhynchus banksii macrorhychus)Seed2
Red-winged Parrot (Aprosmictus erythropterus coccineopterus)Seed3+
Restless Flycatcher (Myiagra inquieta nana)Insects2
Rock Dove (Columba livia)XSeed2
Rose-crowned Fruit-Dove (Ptilinopus regina)Fruit1
19982018
Royal Spoonbill (Platalea regia)Vertebrates/Invertebrates2
Ruddy Turnstone (Arenaria interpres)Invertebrates1
Rufous Fantail (Rhipidura rufifrons)XInsects1
Rufous-banded Honeyeater (Conopophila albogularis)Invertebrates2
Rufous-throated Honeyeater (Conopophila rufogularis)Invertebrates3+
Sacred Kingfisher (Todiramphus sanctus)Invertebrates2
Sanderling (Calidris alba)XInvertebrates3+
Sharp-tailed Sandpiper (Calidris acuminata)Invertebrates1
Shining Flycatcher (Myiagra alecto melvillensis)Invertebrates2
Silver-crowned Friarbird (Philemon argenticeps argenticeps)XNectar2
Sooty Oystercatcher (Haematopus fuliginosus)Invertebrates2
Spangled Drongo (Dicrurus bracteatus baileyi)Omnivore3+
Square-tailed Kite (Lophoictinia isura)XRaptor1
Straw-necked Ibis (Threskiornis spinicollis)Insects2
Striated Heron (Butorides striatus stagnatilis)Vertebrates2
Striated Pardalote (Pardalotus striatus uropygialis)Insects1
Sulphur-crested Cockatoo (Cacatua galerita fitzroyi)Fruit3+
Tawny Frogmouth (Podargus strigoides phalaenoides)XInvertebrates2
Terek Sandpiper (Xenus cinereus)Invertebrates1
Torresian Crow (Corvus orru)Omnivore3+
Torresian Pied Imperial-pigeon (Ducula bicolor)Fruit1
Tree Martin (Petrochelidon nigricans)Insects1
Varied Lorikeet (Psitteuteles versicolor)Nectar1
Varied Triller (Lalage leucomela rufiventris)Insects2
Weebill (Smicrornis brevirostris flavescens)XInsects1
19982018
Whimbrel (Numenius phaeopus)Invertebrates2
Whistling Kite (Haliastur sphenurus)Raptor2
White-bellied Cuckoo-shrike (Coracina papuensis hypoleuca)Insects3+
White-bellied Sea-eagle (Haliaeetus leucogaster)Raptor2
White-breasted Woodswallow (Artamus leucorynchus)Insects2
White-browed Crake (Amaurornis cinerea)Invertebrates3+
White-faced Heron (Egretta novaehollandiae)XInvertebrates3+
White-gaped Honeyeater (Stomiopera unicolor)Nectar3+
White-necked Heron (Ardea pacifica)Vertebrates/Invertebrates2
White-throated Gerygone (Gerygone olivacea rogersi)XInsects1
White-throated Honeyeater (Melithreptus albogularis)Insects2
White-winged Triller (Lalage sueurii)Insects1
Willie Wagtail (Rhipidura leucophrys picata)Insects1
Yellow Oriole (Oriolus flavocinctus flavocinctus)Fruit1
Wood Sandpiper (Tringa glareola)Invertebrates1
Yellow Wagtail (Motacilla flava tschutschensis)XInsects1
Yellow White-eye (Zosterops luteus luteus)Insects3+
Zitting Cisticola (Cisticola juncidis leanyeri)Insects1
Table A4. Bird species recorded in 1998 only or 2018 only.
Table A4. Bird species recorded in 1998 only or 2018 only.
Species NamePrimary Food Source
1998
Azure Kingfisher (Ceyx azureus ruficollaris)Invertebrates
Black-fronted Dotterel (Elseyornis melanops)Invertebrates
Broad-billed Flycatcher (Myiagra ruficollis)Insects
Cattle Egret (Ardea ibis)Insects
Cicadabird (Coracina tenuirostris)Insects
Eurasian Coot (Fulica atra)Omnivore
Great Bowerbird (Ptilonorhynchus nuchalis nuchalis)Omnivore
Large-tailed Nightjar (Caprimulgus macrurus)Insects
Long-toed Stint (Calidris subminuta)Insects
Mangrove Golden Whistler (Pachycephala melanura melanura)Insects
Nankeen Kestrel (Falco cenchroides)Raptor
Nankeen Night-heron (Nycticorax caledonicus)Vertebrates
Pectoral Sandpiper (Calidris melanotos)Insects
Purple Swamphen (Porphyrio porphyrio melanotus)Vertebrates/Invertebrates
Red-backed Button-quail (Turnix maculosus)Seed
Red-backed Fairy-wren (Malurus melanocephalus cruentatus)Insects
Red-kneed Dotterel (Erythrogonys cinctus)Invertebrates
Rock Dove (Columba livia)Seed
Sanderling (Calidris alba)Invertebrates
Silver-crowned Friarbird (Philemon argenticeps argenticeps)Nectar
Weebill (Smicrornis brevirostris flavescens)Insects
White-throated Gerygone (Gerygone olivacea rogersi)Insects
Yellow Wagtail (Motacilla flava tschutschensis)Insects
2018
Australian Magpie (Cracticus tibicen)Omnivore
Bar-breasted Honeyeater (Ramsayornis fasciatus)Nectar/Insects
Beach Stone-curlew (Esacus magnirostris)Invertebrates
Black-shouldered Kite (Elanus axillaris)Raptor
Broad-billed Sandpiper (Limicola falcinellus)Invertebrates
Brown Falcon (Falco berigora)Raptor
Buff-banded Rail (Hypotaenidia philippensis)Omnivore
Bush Stone-curlew (Burhinus grallarius)Invertebrates
Channel-billed Cuckoo (Scythrops novaehollandiae)Fruit
Grey Butcherbird (Cracticus torquatus colletti)Invertebrates
Jacky Winter (Microeca fascinans pallida)Insects
Little Kingfisher (Ceyx pusilla ramsayi)Vertebrates/Invertebrates
Mangrove Grey Fantail (Rhipidura phasiana)Insects
2018
Masked Finch (Poephila personata personata)Seed
Pied Butcherbird (Cracticus nigrogularis picatus)Invertebrates
Rufous Fantail (Rhipidura rufifrons)Insects
Square-tailed Kite (Lophoictinia isura)Raptor
Tawny Frogmouth (Podargus strigoides phalaenoides)Invertebrates
White-faced Heron (Egretta novaehollandiae)Invertebrates

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Figure 1. Study area of the Greater Darwin region (map data sources: Esri, DeLorme, HERE, USGS, Intermap, iPC, NRCAN, Esri Japan, METI, Esri China (Hong Kong), Esri (Thailand), MapmyIndia, Tomtom).
Figure 1. Study area of the Greater Darwin region (map data sources: Esri, DeLorme, HERE, USGS, Intermap, iPC, NRCAN, Esri Japan, METI, Esri China (Hong Kong), Esri (Thailand), MapmyIndia, Tomtom).
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Figure 2. Final classified images of land use change with combined land classes.
Figure 2. Final classified images of land use change with combined land classes.
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Figure 3. Proportion of species by primary food source at sites where urbanisation has increased.
Figure 3. Proportion of species by primary food source at sites where urbanisation has increased.
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Figure 4. Proportion of species by primary food source at sites where greenspace has decreased.
Figure 4. Proportion of species by primary food source at sites where greenspace has decreased.
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Table 1. Results of paired t-test for difference in the overall number of bird species per site between 1998 and 2018 in areas where urbanisation has increased (df = 8).
Table 1. Results of paired t-test for difference in the overall number of bird species per site between 1998 and 2018 in areas where urbanisation has increased (df = 8).
Site Percentage Increase in UrbanisationPercentage Change in Species Numberstp-Value
115.0537.5−3.7390.006
20.992.0−2.8420.022
335.1164.7−3.5630.007
458.2−74.53.1620.013
57.4−42.12.1770.061
65.8336.4−4.3310.003
74.2−16.20.6030.563
88.2−84.62.4750.038
939.117.2−1.5520.159
Table 2. Results of paired t-test for difference in the overall number of bird species per site between 1998 and 2018 in areas where greenspace has decreased (df = 14).
Table 2. Results of paired t-test for difference in the overall number of bird species per site between 1998 and 2018 in areas where greenspace has decreased (df = 14).
SitePercentage Decrease in GreenspacePercentage Change in Species Numberstp-Value
12.9537.5−3.7390.006
22.6155.6−1.9740.084
31361.8−1.4520.185
41.772.7−0.8280.431
527.2164.7−3.5630.007
613.4−74.53.1620.013
714.2336.4−4.3310.003
85.3−90.54.1220.003
931.1−16.20.6030.563
1015.6−84.62.4750.038
110.1−17.20.5330.609
1215.563.6−0.9720.359
1310.717.2−1.5520.159
140.656.3−1.4550.184
151.7192.3−1.9540.087
Table 3. Change in species numbers across sites where urbanisation has increased; I = increase, D = decrease and NC = no change.
Table 3. Change in species numbers across sites where urbanisation has increased; I = increase, D = decrease and NC = no change.
Primary Food SourceSite
123456789
FruitIIIDDIIDI
InsectIIIDIIIDI
InvertebrateIDIDDIDDD
NectarIIIDNCIINCI
OmnivoreIINCDNCIDDD
RaptorIIIDDIDNCI
SeedIIIDDIIDI
VegetationDINCNCNCINCNCD
VertebrateIIIDDIDNCNC
Table 4. Change in species numbers across sites where greenspace has decreased; I = increase, D = decrease and NC = no change.
Table 4. Change in species numbers across sites where greenspace has decreased; I = increase, D = decrease and NC = no change.
Primary Food SourceSite
123456789101112131415
FruitIIDIIDIDIDNCIIDI
InsectIIIIIDIDIDDIIII
InvertebrateIIIIIDIDDDIDDII
NectarINCIDIDIDINCDIIII
OmnivoreIINCDIDIDDDIDDNCI
RaptorIIIDIDIDDNCDIIII
SeedIIIDIDIDIDNCIIII
VegetationDNCDNCINCINCNCNCDDDNCNC
VertebrateIDDDIDIDDNCNCINCNCI
Table 5. Results of paired t-test for difference in the number of bird species by primary food source between 1998 and 2018 in areas where greenspace has decreased (df = 14).
Table 5. Results of paired t-test for difference in the number of bird species by primary food source between 1998 and 2018 in areas where greenspace has decreased (df = 14).
Primary Food Sourcetp-Value
Fruit−0.5560.589
Insect−2.5190.025
Invertebrate−1.0170.326
Nectar−1.1670.263
Omnivore −0.6540.524
Raptor−2.0490.060
Seed−1.3110.211
Vegetation1.7400.104
Vertebrate−0.6880.503
Table 6. Results of paired t-test for difference in the proportion of bird species by primary food source between 1998 and 2018 in areas where greenspace has decreased (df = 14).
Table 6. Results of paired t-test for difference in the proportion of bird species by primary food source between 1998 and 2018 in areas where greenspace has decreased (df = 14).
Primary Food Sourcetp-Value
Fruit1.8490.086
Insect−2.1560.049
Invertebrate0.0720.943
Nectar0.5470.593
Omnivore1.5200.151
Raptor−1.3460.200
Seed0.4580.654
Vegetation1.5170.152
Vertebrate1.3410.201
Table 7. Results of paired t-test for difference in the proportion of feeding specialisation in bird assemblages with changing land use.
Table 7. Results of paired t-test for difference in the proportion of feeding specialisation in bird assemblages with changing land use.
Habitat Changedftp-Value
Urbanisation increase2601
Urbanisation decrease23−1.0481
No change in urbanisation170.0180.986
Greenspace increase230.0020.999
Greenspace decrease440.0050.999
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Fischer, S.E.; Edwards, A.C.; Weber, P.; Garnett, S.T.; Whiteside, T.G. The Bird Assemblage of the Darwin Region (Australia): What Is the Effect of Twenty Years of Increasing Urbanisation? Diversity 2021, 13, 294. https://doi.org/10.3390/d13070294

AMA Style

Fischer SE, Edwards AC, Weber P, Garnett ST, Whiteside TG. The Bird Assemblage of the Darwin Region (Australia): What Is the Effect of Twenty Years of Increasing Urbanisation? Diversity. 2021; 13(7):294. https://doi.org/10.3390/d13070294

Chicago/Turabian Style

Fischer, Sarah E., Andrew C. Edwards, Patrice Weber, Stephen T. Garnett, and Timothy G. Whiteside. 2021. "The Bird Assemblage of the Darwin Region (Australia): What Is the Effect of Twenty Years of Increasing Urbanisation?" Diversity 13, no. 7: 294. https://doi.org/10.3390/d13070294

APA Style

Fischer, S. E., Edwards, A. C., Weber, P., Garnett, S. T., & Whiteside, T. G. (2021). The Bird Assemblage of the Darwin Region (Australia): What Is the Effect of Twenty Years of Increasing Urbanisation? Diversity, 13(7), 294. https://doi.org/10.3390/d13070294

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