The Arrival and Spread of the European Firebug (Pyrrhocoris apterus) in Australia as Documented by Citizen Scientists
Luis Mata1#*, Blythe Vogel2#, Estibaliz Palma1, and Mallik Malipatil3,4
#Joint first authors. 1School of Ecosystem and Forest Sciences, The University of Melbourne, Parkville 3010, Australia. 2Environmental Sustainability and Urban Design, Department of Transport, Kew 3101, Australia. 3Agriculture Victoria, AgriBio, Bundoora 3083, Australia. 4La Trobe University, Bundoora 3083, Australia. *Corresponding author.
Urban Naturalist Notes, No. 3 (2022)
Abstract
We report the recent introduction and spread of the European Firebug (Pyrrhocoris apterus) in Australia, as documented on the citizen science platform iNaturalist. The species was first observed in December 2018 in the City of Brimbank (Melbourne, Victoria) and has quickly expanded into 15 adjacent municipalities, including areas in Metropolitan Melbourne and regional Victoria. Records of the European Firebug in Victoria are rapidly rising, with a current tally of almost 100 observations, as of July 31, 2021.
The case of the European Firebug in Australia showcases the key role that citizen scientists are playing in the early detection of introduced species and in documenting their expansion across their non-native range. Citizen science presents an exciting opportunity to complement biosecurity efforts carried out by government agencies, which often lack resources to sufficiently fund detection and monitoring programs given the overwhelming number of current and potential invasive species.
While the European Firebug has the potential to become a nuisance pest, negatively impacting both ornamental and indigenous plants, the ecological consequences of the rapid range expansion of the species in Australia and other parts of the world are currently unknown. We propose that future studies should be conducted to understand and quantify the effects of the species on Australian ecological communities and production industries.
Given the range expansion patterns of the European Firebug worldwide, their adaptation ability, and future climate scenarios, we suspect the European Firebug will continue to expand its range beyond Victoria to other urban environments and regional areas of Australia, New Zealand, and the South Pacific. We believe that most of the knowledge about how this expansion continues to unfold will be provided by citizen scientists contributing observations in or around urban areas.
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Volume 9, 2022 Urban Naturalist Notes No. 3
The Arrival and Spread
of the European Firebug
(Pyrrhocoris apterus) in
Australia as Documented
by Citizen Scientists
Luis Mata, Blythe Vogel,
Estibaliz Palma, and Mallik Malipatil
Urban Naturalist
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Cover Photograph: European Firebug in Westgate Park (Melbourne, Australia). Photograph © Luis Mata.
Urban Naturalist Notes
L. Mata, B. Vogel, E. Palma, and M. Malipatil
Vol. 9, 2022 N3:1–7
1
2021
The Arrival and Spread of the European Firebug (Pyrrhocoris
apterus) in Australia as Documented by Citizen Scientists
Luis Mata1#*, Blythe Vogel2#, Estibaliz Palma1 and Mallik Malipatil3,4
Abstract - We report the recent introduction and spread of the European Firebug (Pyrrhocoris apterus)
in Australia, as documented on the citizen science platform iNaturalist. The species was first
observed in December 2018 in the City of Brimbank (Melbourne, Victoria) and has quickly expanded
into 15 adjacent municipalities, including areas in Metropolitan Melbourne and regional Victoria.
Records of the European Firebug in Victoria are rapidly rising, with a current tally of almost 100
observations, as of July 31, 2021.
The case of the European Firebug in Australia showcases the key role that citizen scientists are
playing in the early detection of introduced species and in documenting their expansion across their
non-native range. Citizen science presents an exciting opportunity to complement biosecurity efforts
carried out by government agencies, which often lack resources to sufficiently fund detection and
monitoring programs given the overwhelming number of current and potential invasive species.
While the European Firebug has the potential to become a nuisance pest, negatively impacting
both ornamental and indigenous plants, the ecological consequences of the rapid range expansion of
the species in Australia and other parts of the world are currently unknown. We propose that future
studies should be conducted to understand and quantify the effects of the species on Australian ecological
communities and production industries.
Given the range expansion patterns of the European Firebug worldwide, their adaptation ability,
and future climate scenarios, we suspect the European Firebug will continue to expand its range
beyond Victoria to other urban environments and regional areas of Australia, New Zealand, and the
South Pacific. We believe that most of the knowledge about how this expansion continues to unfold
will be provided by citizen scientists contributing observations in or around urban areas.
Pyrrhocoris apterus Linnaeus (European Firebug) [Hemiptera: Pyrrhocoridae] is an herbivorous,
gregarious, and vividly aposematic species (Fig. 1). European Firebugs present
distinct wing polymorphism, with most individuals expressing the brachypterous (Fig. 1)
over the macropterous form (Socha 1993). The European Firebug diet is mainly associated
with seeds of Linden Trees (genus Tilia); however, the species is known to be widely polyphagous,
feeding on plant species in the families Asparagaceae (Asparagus), Betulaceae
(Birch), Boraginaceae (Borage), Cupressaceae (Cypress), Asteraceae (Daisy), Cornaceae
(Dogwood), Ulmaceae (Elm), Malvaceae (Mallow), Lamiaceae (Mint), Fagaceae (Oak), Fabaceae
(Pea), Rosaceae (Rose), and Sapindaceae (Soapberry) (Kristenová et al. 2011). The
species occurs in a wide array of terrestrial habitats (including on leaflitter, rock crevices,
under bark, and on vegetation) and has been documented extensively in urban environments
(Endrestøl and Roth 2020, Hodgson 2008, Mata et al. 2013, Oviedo Rojas and Jackson
2018). The species is well adapted to cold climates and presents adaptations that allow it
to survive freezing temperatures (Koštál and Šimek 2000). European Firebugs normally
develop one generation per year, but may also produce a second generation under warmer
conditions (Saulich and Musolin 1996). Easily reared under experimental conditions, Euro-
#Joint first authors 1School of Ecosystem and Forest Sciences, The University of Melbourne, Parkville
3010, Australia. 2Environmental Sustainability and Urban Design, Department of Transport, Kew
3101, Australia. 3Agriculture Victoria, AgriBio, Bundoora 3083, Australia. 4La Trobe University,
Bundoora 3083, Australia. *Corresponding author: lmata@unimelb.edu.au.
Associate Editor: Katalin Szlavecz, Johns Hopkins University.
Vol. 9, 2022 URBAN NATURALIST NOTES N3:1–7
Urban Naturalist Notes
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Vol. 9, 2022 N3:1–7
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pean Firebugs have been widely used as model organisms in a wide range of morphological,
developmental, ecological, ethological, physiological, biochemical, genetic, and evolutionary
studies (Kristenová et al. 2011, Socha 1993).
The European Firebug has been traditionally known as a Palearctic species, ranging
from central and southern Europe to western China (Kerzhner 2001). Ample evidence,
however, shows that it has been rapidly spreading outside its traditional native range (Fig.
2; Gardiner 2020). While, in some instances, it might be difficult to disentangle natural from
anthropogenic expansion—for example, in the Azores (Cardoso et al. 2011), Malta (Cassar
2019), and northern Europe (Endrestøl and Roth 2020)—it is evident that the European
Firebug has reached the American (Hodgson 2008, Oviedo Rojas and Jackson 2018) and
Australian continents, assisted by humans.
Here, we report the arrival and spread of the European Firebug in Australia. On 10 March
2019, the first authors were facilitating a citizen science outdoor activity targeted at documenting
ecological interactions between indigenous plants and insect pollinators in urban environments.
The activity was part of a research and community engagement project entitled “Pollinator
Observatories”, which took place in Westgate Park (Melbourne, Victoria, Australia), between
April 2017 and March 2019 (Mata et al. 2020, Vogel 2019). During the activity, a citizen scientist
asked for assistance to identify an insect they had observed on the ground. To our surprise,
the insect was the European Firebug, a species that one of the first authors had researched in its
native range in the Iberian Peninsula and had experience in surveying and identifying (Mata et
al. 2013). We, therefore, had no difficulties in recognizing the species as non-native to Australia.
We set to assess whether we had documented the first record of the European Firebug
in Australia. We began by searching the literature for evidence of the European Firebug occurring
in Australia but could not find any references documenting this. We then searched
the Atlas of Living Australia for European Firebug records but found none. Next, we asked
Figure 1. The European Firebug Pyrrhocoris apterus (Westgate Park, Melbourne, Victoria, Australia).
Photo by Luis Mata.
Urban Naturalist Notes
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colleagues across our networks whether they had any relevant information on the European
Firebug and used this as an opportunity to communicate the discovery to the relevant biosecurity
authorities. We learned the European Firebug had, indeed, been previously recorded
in Melbourne’s western suburbs as early as April 2018 by the Border Surveillance Group of
the Australia Government’s Department of Agriculture and Water Resources (Adam Broadley
pers. comm., later published in DAWE [2019]). Finally, having experienced first-hand how
straightforward it had been for a citizen scientist to find the European Firebug, we consulted
the online biodiversity citizen science platform iNaturalist (https://www.inaturalist.org/) hoping
to find further evidence of the arrival and potential spread of the species in Australia.
We found that, at the time of our inquiry (April 2019), 4 citizen scientists had contributed
6 iNaturalist observations of the European Firebug in Australia. The earliest was from
21 December 2018, the latest from 9 March 2019; all were from the suburb of Sunshine in
the City of Brimbank (Melbourne, Victoria). We monitored subsequent iNaturalist observa-
Figure 2. Current biogeographical distribution of the European Firebug Pyrrhocoris apterus based
on data provided by (a) the Global Biodiversity Information Facility (https://www.gbif.org/) and (b)
iNaturalist (https://www.inaturalist.org/). The species was, until recently, restricted to the Palearctic
but has now expanded its range to the African, Australian, and American continents. The colored cells
in each map represent the occurrence and density of European Firebug records.
Urban Naturalist Notes
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Figure 3. (a-d) Past and current distribution of the European Firebug Pyrrhocoris apterus in Australia.
The heat maps highlight the Local Government Areas (i.e. municipalities and shires) across the Melbourne
Metropolitan Area and regional Victoria in which the species has been documented by citizen
scientists through iNaturalist as of 31 July 2021. (f) Number of accumulated observations by month of
the European Firebug in Australia from December 2018 to July 2021. CBD: Central Business District;
LGA: Local Government Area.
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tions to track the potential spread of the species throughout Melbourne, regional Victoria,
and likely other urban and regional areas across Australia. We found that, as of July 2021,
over 30 citizen scientists have contributed approximately 100 observations of the European
Firebug in Australia (Fig. 3f). These observations distinctly indicate that the species has
been expanding its range, spreading from its original introduction enclave in the City of
Brimbank (Fig. 3a) to 15 municipalities and shires in the Melbourne Metropolitan Area and
regional Victoria (Fig. 3d) in less than 3 years.
Taken together, our findings highlight the key role that citizen scientists—whether
prompted by academic-led activities, government-initiated programs, or personal enthusiasm—
are playing in detecting newly introduced species and documenting their establishment
and expansion in their non-native ranges. The engagement of citizen scientists can
effectively complement government-funded detection and monitoring programs, ensuring
that introductions or outbreaks of potentially harmful species do not remain undetected
(Ricciardi et al. 2017, Turrini et al. 2018). With careful project design, volunteer training,
and professional validation, citizen science observations can be as reliable and robust
as those undertaken by professional scientists (Kosmala et al. 2016). Citizen scientists
around the world have successfully detected invasive species early, significantly contributing
to the rapid management of potential pest species (Azzurro et al. 2013, Cohen
et al. 2011, Eritja et al. 2019, Scyphers et al. 2015, Walther and Kampen 2017). Citizen
scientists, who can cover much larger areas and longer time periods than experts alone
can (Turrini et al. 2018), have also made critical contributions in the ongoing monitoring
of invasive species following their detection, documenting their long-term spread and impacts
on native species and ecosystems, and in informing management plans and actions
(Maistrello et al. 2016, Pyšek et al. 2020, Ricciardi et al. 2017, Weed and Schwarzländer
2014). Citizen scientists participating in national, continental, and global projects (e.g.
eButterfly, eBird, iNaturalist) often excel at detecting range expansions or shifts of both
native and introduced species much earlier than professional scientists do (Crall et al.
2015, Chandler et al. 2017, Prudic et al. 2017). These and other examples emphasise the
ability for citizen scientists to reliably study species of interest—including actual or potential
threats—across wide geographical ranges.
We argue that the contributions of citizen scientists are highly valuable, as they complement
efforts of biosecurity agencies in charge of detecting and monitoring introduced species
during their early invasion stages or during outbreaks. This is particularly relevant for
species with documented detrimental impacts in their non-native regions, if left unmanaged.
While Oviedo Rojas and Jackson (2018) note that the European Firebug could become a
nuisance pest in Canada, negatively impacting both ornamental and indigenous plants, the
potential ecological consequences, whether detrimental or beneficial, of the rapid range
expansion of the European Firebug in Australia and other parts of the world are currently
unknown. Future studies on the current topic are therefore recommended in order to understand
and quantify the potential detrimental impacts of the European Firebug on Australian
ecological communities, as well as on its grains, horticultural, and nursery industries.
Given the global range expansion patterns of the European Firebug, the reported feeding
and thermal adaptations of the species, and future climate scenarios, we would like
to put forward the following two ideas about the future spread of the European Firebug
across Australia and beyond. First, we believe that, if left unmanaged, the species will
likely continue to spread across metropolitan Melbourne and regional Victoria, eventually
reaching other urban and rural areas across continental Australia, and also Tasmania,
New Zealand, and other South Pacific islands, and likely, in the not-too-distant future,
Urban Naturalist Notes
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Vol. 9, 2022 N3:1–7
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the Antarctic continent. Second, we are convinced that the majority of the future range
expansion of the European Firebug across Australia and beyond will be documented by
citizen scientists contributing observations in or around urban areas. As such, we would
like to thank them in advance for their efforts and call for biosecurity and other related
agencies to continue to support their invaluable contributions to science and society.
Acknowledgements
The authors would like to acknowledge the Traditional Custodians of the land and waterways on
which this research took place, the Woi wurrung and Boon wurrung peoples of the eastern Kulin Nations.
We pay our respects to their Elders, past, present, and emerging. Thanks to Janet Bolitho and
everyone at Westgate Biodiversity: Bili Nursery & Landcare, Adam Broadley at DAWE, the citizen
scientist that first discovered the European Firebug in Westgate Park, all of those that have contributed
observations of the European Firebug and other species to iNaturalist, and last but not least, the
iNaturalist team for developing and maintaining such a powerful scientific and engagement resource.
Data and Code Accessibility Statement
The data and codes to reproduce the analysis shown in Fig. 2 are already published and publicly
available in Zenodo: https://doi.org/10.5281/zenodo.5355347
Author Contribution Statement
Luis Mata: Conceptualization (co-lead); Investigation (equal); Writing-original draft (co-lead);
Writing-review and editing (equal). Blythe Vogel: Conceptualization (co-lead); Investigation (equal);
Writing-original draft (co-lead); Writing-review and editing (equal). Estibaliz Palma: Data curation
(lead); Formal analysis (lead); Investigation (equal); Writing-review and editing (equal). Mallik
Malipatil: Investigation (equal); Writing-review and editing (equal).
Literature Cited
Azzurro, E., E. Broglio, F. Maynou, and M. Bariche. 2013. Citizen science detects the undetected:
The case of Abudefduf saxatilis from the Mediterranean Sea. Management of Biological Invasions
4:167–170.
Cardoso, P., T.L. Erwin, P.A.V. Borges, and T.R. New. 2011. The seven impediments in invertebrate
conservation and how to overcome them. Biological Conservation 144:2647–2655.
Cassar, T. 2019. Pyrrhocoris apterus (Linnaeus , 1758) – a new record of firebug (Hemiptera, Pyrrhocoridae
from Malta. 10:106–107.
Chandler, M., L. See, K. Copas, A.M.Z. Bonde, B.C. López, F. Danielsen, J.K. Legind, S. Masinde,
A.J. Miller-Rushing, G. Newman, A. Rosemartin, and E. Turak. 2017. Contribution of citizen science
towards international biodiversity monitoring. Biological Conservation 213:280–294.
Cohen, C.S., L. McCann, T. Davis, L. Shaw, and G. Ruiz. 2011. Discovery and significance of the
colonial tunicate Didemnum vexillum in Alaska. Aquatic Invasions 6:263–271.
Crall, A.W., C.S. Jarnevich, N.E. Young, B.J. Panke, M. Renz, and T.J. Stohlgren. 2015. Citizen science
contributes to our knowledge of invasive plant species distributions. Biological Invasions
17:2415–2427. Springer International Publishing.
DAWE. 2019. Environmental biosecurity risk management in Australia. Canberra.
Endrestøl, A., and S. Roth. 2020. The firebug Pyrrhocoris apterus (Linnaeus, 1758) (Hemiptera,
Heteroptera) new to the Norwegian fauna–with an explosive expansion in Northern Europe. Norwegian
Journal of Entomology 67:81–90.
Eritja, R., I. Ruiz-Arrondo, S. Delacour-Estrella, F. Schaffner, J. Álvarez-Chachero, M. Bengoa, M.Á.
Puig, R. Melero-Alcíbar, A. Oltra, and F. Bartumeus. 2019. First detection of Aedes japonicus in
Spain: An unexpected finding triggered by citizen science. Parasites an d Vectors 12:1–9.
Gardiner T. (2020) Great Balls of Firebugs! The Global Range Expansion of Pyrrhocoris apterus. Pp.
577–588, In D.A. DellaSala and M.I. Goldstein (Eds.). Imperiled: The Encyclopedia of Conservation,
1st Edition. Elsevier, Amsterdam, The Netherlands. 2608 pp.
Urban Naturalist Notes
L. Mata, B. Vogel, E. Palma, and M. Malipatil
Vol. 9, 2022 N3:1–7
7
Hodgson, E.W. 2008. Utah pests fact sheet: Red fire bug. Utah State University Extension and Utah
Plant Pest Diagnostic Laboratory. Available online at https://digitalcommons.usu.edu/cgi/viewcontent.
cgi?article=1945&context=extension_curall.
Kerzhner, I.M. 2001. Superfamily Pyrrhocoroidea Amyot & Serville, 1843. In B. Aukema and Ch.
Rieger (Eds.). Catalogue of the Heteroptera of the Palaearctic Region. Cimicomorpha I. The Netherlands
Entomological Society, Amsterdam 4:245–258.
Kosmala, M., A. Wiggins, A. Swanson, and B. Simmons. 2016. Assessing data quality in citizen science.
Frontiers in Ecology and the Environment 14:551–560.
Koštál, V. and Šimek, P. 2000. Overwintering strategy in Pyrrhocoris apterus (Heteroptera): The
relations between life-cycle, chill tolerance and physiological adjustments. Journal of Insect
Physiology, 46:1321–1329.
Kristenová, M., Exnerová, A., and Štys, P. 2011. Seed preferences of Pyrrhocoris apterus (Heteroptera:
Pyrrhocoridae): Are there specialized trophic populations? European Journal of Entomology,
108:581–586.
Maistrello, L., P. Dioli, M. Bariselli, G.L. Mazzoli, and I. Giacalone-Forini. 2016. Citizen science and early
detection of invasive species: Phenology of first occurrences of Halyomorpha halys in Southern Europe.
Biological Invasions 18:3109–3116. Springer International Publishing.
Mata, L., B. Vogel, and J. Bolitho. 2020. Pollinator Observatories–Citizen science to engage people
with nature in cities. Report prepared for Westgate Biodiversity: Bili Nursery & Landcare. 82 pp.
Mata, L.M., J.M.G.G. Silva, and M.G. Goula. 2013. Pyrrhocoridae from the Iberian Peninsula (Hemiptera
: Heteroptera). Heteropterus Revista de Entomologia 13:175–189.
Oviedo Rojas, P.J., and M.D. Jackson. 2018. Pyrrhocoris apterus L. (Hemiptera: Pyrrhocoridae), a
newly introduced family, genus, and species to Ontario and Canada. The Journal of the Entomological
Society of Ontario 149:27–32.
Prudic, K.L., K.P. McFarland, J.C. Oliver, R.A. Hutchinson, E.C. Long, J.T. Kerr, and M. Larrivée.
2017. eButterfly: Leveraging massive online citizen science for butterfly consevation. Insects
8:1–12.
Pyšek, P., P.E. Hulme, D. Simberloff, S. Bacher, T.M. Blackburn, J.T. Carlton, W. Dawson, F. Essl,
L.C. Foxcroft, P. Genovesi, J.M. Jeschke, I. Kühn, A.M. Liebhold, N.E. Mandrak, L.A. Meyerson,
A. Pauchard, J. Pergl, H.E. Roy, H. Seebens, M. van Kleunen, M. Vilà, M.J. Wingfield, and D.M.
Richardson. 2020. Scientists’ warning on invasive alien species. Biological Reviews 95:1511–1534.
Ricciardi, A., T.M. Blackburn, J.T. Carlton, J.T.A. Dick, P.E. Hulme, J.C. Iacarella, J.M. Jeschke,
A.M. Liebhold, J.L. Lockwood, H.J. MacIsaac, P. Pyšek, D.M. Richardson, G.M. Ruiz, D. Simberloff,
W.J. Sutherland, D.A. Wardle, and D.C. Aldridge. 2017. Invasion Science: A Horizon
Scan of Emerging Challenges and Opportunities. Trends in Ecology and Evolution 32:464–474.
Saulich, A.K. and Musolin, D.L. 1996. Univoltinism and its regulation in some temperate true bugs
(Heteroptera). European Journal of Entomology, 93:507–518.
Scyphers, S.B., S.P. Powers, J.L. Akins, J.M. Drymon, C.W. Martin, Z.H. Schobernd, P.J. Schofield,
R.L. Shipp, and T.S. Switzer. 2015. The Role of Citizens in Detecting and Responding to a Rapid
Marine Invasion. Conservation Letters 8:242–250.
Socha, R. 1993. Pyrrhocoris apterus (Heteroptera)–an experimental model species: A review. European
Journal of Entomology 90:241–286.
Turrini, T., D. Dörler, A. Richter, F. Heigl, and A. Bonn. 2018. The threefold potential of environmental
citizen science–Generating knowledge, creating learning opportunities and enabling civic
participation. Biological Conservation 225:176–186.
Vogel, B. 2019. Predicting plant-insect pollinator interactions to engage urban residents with nature.
The University of Melbourne, Melbourne, Australia. 66 pp.
Walther, D., and H. Kampen. 2017. The Citizen Science Project “Mueckenatlas” Helps Monitor the
Distribution and Spread of Invasive Mosquito Species in Germany. Journal of Medical Entomology
54:1790–1794.
Weed, A.S., and M. Schwarzländer. 2014. Density dependence, precipitation and biological control
agent herbivory influence landscape-scale dynamics of the invasive Eurasian plant Linaria dalmatica.
Journal of Applied Ecology 51:825–834.