Christian C. Voigt · Tigga Kingston Editors Bats in the Anthropocene: Conservation of Bats in a Changing World Bats in the Anthropocene: Conservation of Bats in a Changing World Christian C. Voigt · Tigga Kingston Editors Bats in the Anthropocene: Conservation of Bats in a Changing World Editors Christian C. Voigt Berlin Germany ISBN 978-3-319-25218-6 ISBN 978-3-319-25220-9 (eBook) DOI 10.1007/978-3-319-25220-9 Library of Congress Control Number: 2015950865 Springer Cham Heidelberg New York Dordrecht London © The Editor(s) (if applicable) and The Author(s) 2016. The book is published with open access at SpringerLink.com. Open Access This book is distributed under the terms of the Creative Commons Attribution Noncommercial License, which permits any noncommercial use, distribution, and reproduction in any medium, provided the original author(s) and source are credited. All commercial rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. Printed on acid-free paper Springer International Publishing AG Switzerland is part of Springer Science+Business Media (www.springer.com) Tigga Kingston Lubbock, TX USA For Thomas H. Kunz and Otto von Helversen for sharing with us their passion for bats. For Silke, Philippa and Florian (CCV) and for Danny (TK) for their inspiration and patience. vii Contents 1 Bats in the Anthropocene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Christian C. Voigt and Tigga Kingston Part I Bats in Anthropogenically Changed Landscapes 2 Urbanisation and Its Effects on Bats—A Global Meta-Analysis . . . . 13 Kirsten Jung and Caragh G. Threlfall 3 Bats and Roads . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 John Altringham and Gerald Kerth 4 Responses of Tropical Bats to Habitat Fragmentation, Logging, and Deforestation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 Christoph F.J. Meyer, Matthew J. Struebig and Michael R. Willig 5 Insectivorous Bats and Silviculture: Balancing Timber Production and Bat Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 Bradley Law, Kirsty J. Park and Michael J. Lacki 6 Bats in the Anthropogenic Matrix: Challenges and Opportunities for the Conservation of Chiroptera and Their Ecosystem Services in Agricultural Landscapes . . . . . . . . 151 Kimberly Williams-Guillén, Elissa Olimpi, Bea Maas, Peter J. Taylor and Raphaël Arlettaz 7 Dark Matters: The Effects of Artificial Lighting on Bats . . . . . . . . . . 187 E.G. Rowse, D. Lewanzik, E.L. Stone, S. Harris and G. Jones Contents viii 8 Bats and Water: Anthropogenic Alterations Threaten Global Bat Populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215 Carmi Korine, Rick Adams, Danilo Russo, Marina Fisher-Phelps and David Jacobs Part II Emerging Disesases 9 White-Nose Syndrome in Bats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 Winifred F. Frick, Sébastien J. Puechmaille and Craig K.R. Willis 10 Zoonotic Viruses and Conservation of Bats . . . . . . . . . . . . . . . . . . . . . 263 Karin Schneeberger and Christian C. Voigt Part III Human-Bat Conflicts 11 Impacts of Wind Energy Development on Bats: A Global Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295 Edward B. Arnett, Erin F. Baerwald, Fiona Mathews, Luisa Rodrigues, Armando Rodríguez-Durán, Jens Rydell, Rafael Villegas-Patraca and Christian C. Voigt 12 Exploitation of Bats for Bushmeat and Medicine . . . . . . . . . . . . . . . . 325 Tammy Mildenstein, Iroro Tanshi and Paul A. Racey 13 The Conflict Between Pteropodid Bats and Fruit Growers: Species, Legislation and Mitigation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Sheema Abdul Aziz, Kevin J. Olival, Sara Bumrungsri, Greg C. Richards and Paul A. Racey 14 Bats and Buildings: The Conservation of Synanthropic Bats . . . . . . . 427 Christian C. Voigt, Kendra L. Phelps, Luis F. Aguirre, M. Corrie Schoeman, Juliet Vanitharani and Akbar Zubaid 15 Conservation Ecology of Cave Bats . . . . . . . . . . . . . . . . . . . . . . . . . . . . 463 Neil M. Furey and Paul A. Racey Part IV Conservation Approaches, Educational and Outreach Programs 16 The Roles of Taxonomy and Systematics in Bat Conservation . . . . . . 503 Susan M. Tsang, Andrea L. Cirranello, Paul J.J. Bates and Nancy B. Simmons Contents ix 17 Networking Networks for Global Bat Conservation . . . . . . . . . . . . . . 539 Tigga Kingston, Luis Aguirre, Kyle Armstrong, Rob Mies, Paul Racey, Bernal Rodríguez-Herrera and Dave Waldien 18 Cute, Creepy, or Crispy—How Values, Attitudes, and Norms Shape Human Behavior Toward Bats . . . . . . . . . . . . . . . . 571 Tigga Kingston Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 597 1 Chapter 1 Bats in the Anthropocene Christian C. Voigt and Tigga Kingston © The Author(s) 2016 C.C. Voigt and T. Kingston (eds.), Bats in the Anthropocene: Conservation of Bats in a Changing World , DOI 10.1007/978-3-319-25220-9_1 Abstract Humans have inadvertently changed global ecosystems and triggered the dawn of a new geological epoch, the Anthropocene. While some organisms can tolerate human activities and even flourish in anthropogenic habitats, the vast majority are experiencing dramatic population declines, pushing our planet into a sixth mass extinction. Bats are particularly susceptible to anthropogenic changes because of their low reproductive rate, longevity, and high metabolic rates. Fifteen percent of bat species are listed as threatened by the IUCN, i.e., they are consid- ered Critically Endangered, Endangered or Vulnerable. About 18 % of species are Data Deficient, highlighting the paucity of ecological studies that can support conservation status assessments. This book summarizes major topics related to the conservation of bats organized into sections that address: the response of bats to land use changes; how the emergence of viral and fungal diseases has changed bat populations; our perception of bats; and drivers of human–bat conflicts and possi- ble resolutions and mitigation. The book ends with approaches that might advance bat conservation through conservation networks and a better understanding of human behavior and behavioral change. C.C. Voigt ( * ) Leibniz Institute for Zoo and Wildlife Research, Alfred-Kowalke-Str. 17, 10315 Berlin, Germany e-mail: voigt@izw-berlin.de C.C. Voigt Institute for Biology, Freie Universität Berlin, Takustr. 6, 14195 Berlin, Germany T. Kingston ( * ) Department of Biological Sciences, Texas Tech University, Lubbock, TX, USA e-mail: tigga.kingston@ttu.edu 2 C.C. Voigt and T. Kingston 1.1 The Emergence of a New Geological Epoch: The Anthropocene The world in which we live is fragile; a small layer of organismic activity covers the planet like a microbial film on top of a large boulder. Nonetheless, humans treat the Earth as if anthropogenic impacts on this delicate biological layer may be absorbed by unfailing natural buffers. Yet, convergent and overwhelming evidence from all over the world underlines that mankind has already changed and contin- ues changing the face of our planet. Among the many transformations humans imposed on our planet, some of the most severe appear to be (1) the addition of more than 550 billion metric tons of carbon to the atmosphere which are the main drivers of global climate change and ocean acidification (Gray 2007; Ciasi and Sabine 2013), (2) the alteration of the global nitrogen cycle by the use of artificial fertilizers (Canfield et al. 2010), (3) the routing of more than one third of global primary production to human consumption (Krausmann et al. 2013), (4) the ongo- ing mass extinction of species (Barnosky et al. 2011), and (5) the globalization of transport which has resulted in the spread of invasive species and pathogens (Lewis and Maslin 2015). It is now widely recognized that global ecosystem ser- vices may be inadvertently suffering from human action, because human-induced environmental impacts are overriding natural process that have dominated our planet for millions of years (Steffen et al. 2011). In the face of lasting human impacts on the Earth’s geological conditions and processes, many scientists, beginning with Paul Crutzen and Eugene Stoermer in 2000, now posit that our actions have brought us to the dawn of a new geologi- cal epoch—the Anthropocene. The pros and cons regarding this definition, which literally means “Human Epoch” and would succeed the Holocene, are still heavily debated (Monastersky 2015). Yet skeptics are declining in number, and much of the current debate focuses on the exact beginning of the Anthropocene, generally considered to be c. 1800. The Anthropocene working group of the Subcommission on Quaternary Stratigraphy reports to the International Commission on Stratigraphy with a proposal to formalize the Anthropocene in 2016. For the pur- pose of this book, we do not refer to an exact starting point of the Anthropocene, but merely acknowledge the fact that humans have an impact on virtually all global ecosystems and that wildlife species such as bats (order Chiroptera) have adjusted to these changes, experienced substantial population declines, or gone extinct. 1.2 Bats in the Anthropocene: The Conservation of a Nocturnal Taxon Bats (order Chiroptera) include more than 1300 extant species, forming the second largest mammalian order, and are unique among mammals in their evolution of powered flight. Although the common ancestor of living bats dates back to the K/T 3 1 Bats in the Anthropocene boundary (c. 70 mya), the most rapid radiation of any mammalian order resulted in all 18 extant families by the end of the Eocene c. 37 mya (Teeling et al.2005). Moreover, although the majority of bat species are insectivorous, trophic diversity is extraordinary for a single order, with frugivores, nectarivores, piscivores, san- guinivores, and carnivores represented. Bats currently inhabit all continents except Antarctica, and in many parts of the world, especially the tropics, are the most species-rich mammalian group at a given locality, with alpha diversity reaching about 70 species in the Paleotropics (Kingston et al. 2010) and over 100 in the Neotropics (Voss and Emmons 1996; Rex et al. 2008). From any perspective, bats are an evolutionary and ecological success story. Nonetheless, bat populations are under severe threat in many regions of the world (Racey and Entwistle 2003). The last recorded case of a bat species driven to extinction is that of the Christmas Island pipistrelle, Pipistrellus murrayi (Lumsden and Schulz 2009; Lumsden 2009; Martin et al. 2012), yet this species is most likely not the last one to vanish from our planet. The IUCN Bat Specialist Group is in the process of reassessing the Red List status of bat species, with the current assessments of 1150 species mostly com- pleted in 2008, with 34 species assessed since. From these assessments, five spe- cies were assessed as Extinct (giant vampire bat ( Desmodus draculae ), dusky flying fox ( Pteropus brunneus ), large Pelew flying fox ( P. pilosus ), dark fly- ing fox ( P. subniger ), and Guam flying fox ( P. tokudae )). The giant vampire bat is known only from the fossil and subfossil records, and the causes of its extinc- tion are unknown. However, the four island Pteropus spp. are all victims of the Anthropocene, with hunting and habitat loss as the main drivers of extinction. Fifteen percent of bat species are listed in the threatened categories [Critically Endangered (CE), Endangered (EN), and Vulnerable (VU)] and 7 % are Near Threatened (Fig. 1.1). Around 18 % of species are Data Deficient (DD), and there have been a wealth of new species discovered since the last assessment. The pat- tern of vulnerability is fairly consistent across families (Fig. 1.2), with the notable exception of the Pteropodidae with 36 % of species extinct or threatened, prob- ably because of their size, their appeal as bushmeat and for traditional medicine, Fig. 1.1 Red List status of the 1150 bat species assessed 2008–2014 (IUCN 2015). IUCN categories are EX Extinct, CR Critically Endangered, EN Endangered, VU Vulnerable, NT Near Threatened, DD Data Deficient, LC Least Concern. Number of species and percentage of all species given as labels 4 C.C. Voigt and T. Kingston and because many form susceptible island populations. Even this depicts only part of the picture; populations are only considered stable in 21 % of all species and increasing in less than 1 %. Of the remaining species, populations are decreasing (23 %) or the trend is unknown (55 %). Moreover, of the 687 species assessed as Least Concern (LC), current specific threats were identified for about 27 % of spe- cies. Declining populations and identified threats suggest a bleak future, and it is probable that more species will satisfy the rigorous criteria of the threatened cat- egories in the coming years. Globally, the major threats to bat species identified by IUCN assessments are land use change (logging, non-timber crops, livestock farming and ranching, wood and pulp plantations, and fire), urbanization, hunting and persecution, quarrying and general human intrusions on bat habitats (Fig. 1.3). Bats are particularly sus- ceptible to these human-induced perturbations of habitats because of their distinct life history. Bats are on the slow side of the slow-fast continuum of life histories (Barclay and Harder 2003). For example, they reproduce at a low rate (Barclay et al. 2004) and are long-lived mammals (Munshi-South and Wilkinson 2010; Wilkinson and South 2002). Thus, bat populations recover slowly from increased mortality rates. Despite their low reproductive rate and longevity, bats have rela- tively high metabolic rates owing to their small size which leads to relatively high food requirements (Thomas and Speakman 2003). Lastly, bats are nocturnal animals with often cryptic habits. Even though they are present in many larger cities of the temperate zone, they often go unnoticed by their human neighbors. It is quite likely that perceptions of bats would be very dif- ferent if Homo sapiens evolved as a nocturnal hominid. Or to put it in the words of Rich and Longcore: What if we woke up one morning and realize that we missed Fig. 1.2 Red List status of bats by family. Abbreviations as for Fig. 1.1 5 1 Bats in the Anthropocene half of the story in our conservation efforts, namely the night part? (modified after Rich and Longcore 2004, p. 1). This brings up an important question: Do noctur- nal animals benefit less from legal protection than diurnal animals? Are we more concerned about animals that we see and interact with during daytime? Do human societies perceive and evaluate, for example, fatalities of birds of prey at wind tur- bines in a different way than bat fatalities when both ought to benefit from the same level of protection? Do we consider recommendations to reduce light pol- lution for the sake of nocturnal animals such as bats, or does the expansion of the human temporal niche into the night come at high costs for all nocturnal animals? In summary, we speculate that bats as nocturnal animals might be particularly exposed to human-induced ecological perturbations because we are driven by our visual system and therefore tend to neglect the dark side of conservation, i.e., the protection of nocturnal animals. 1.3 Why Care About Bat Conservation? The reasoning for the conservation of nature can be manifold, reaching from purely moral to monetary arguments and legal requirements. It may also vary according to the scale of the conservation approach, i.e., whether it is driven by Fig. 1.3 Frequency of threats listed in the IUCN assessments of bat species. a Distribution of major threats across assessments. Land use changes, urbanization. and hunting are aggregations of IUCN listed threats given in b – d . Frequency of threat and percentage contribution are given 6 C.C. Voigt and T. Kingston local, national, or international perspectives. Indeed, ethical considerations for the protection of species—although quite often neglected in modern civiliza- tion—should be the primary motivation; i.e., the obligation of humans to con- serve nature for the simple reason of its existence and for the more selfish reason to make the diversity of biological life accessible and useable to following gen- erations of humans. Lately, economic arguments for the conservation of nature are increasingly used, e.g., the importance of protecting water catchment areas to provide potable water or irrigation in agriculture. So-called ecosystem services of nature are highly valued in modern societies and therefore benefit from increasing protection. Recent attempts to critically review the ecosystem services provided by bats have revealed that many species offer unique and large-scale monetary benefits to agricultural industry (Kunz et al. 2011; Ghanem and Voigt 2012; Maas et al. 2015). For example, flowers of the Durian tree are only effectively pollinated by the Dawn bat, Eonycteris spelaea , in Southeast Asia (Bumrungsri et al. 2009). Durian is a highly valued fruit in Asia with Thailand producing a market value of durians of almost 600 million US$ annually (Ghanem and Voigt 2012). Other bats consume large amounts of pest insects, thereby offering services that could save millions of US$ for national industries (Boyles et al. 2011; Wanger et al. 2014). However, the monetary approach for protecting bat species is a double- edged sword, since bat species without apparent use for human economy may not benefit from protection compared to those that provide some ecosystem services. Moreover, arguments based on economic or utilitarian values of wildlife may appeal to self-interest motivations and suppress environmental concern (Kingston 2016). In this context, it is important to note that we have just started to under- stand the ecological role bats fill in natural ecosystems. For example, bats have been recently documented as top-down regulators of insect populations in forest habitats of the tropics and temperate zone (Kalka et al. 2008; Boehm et al. 2011) and also in subtropical coffee and cacao plantations (Williams-Guillen et al. 2008; Maas et al. 2013). Finally, bats are protected by law in some countries. For exam- ple, they are covered by the Habitat Directive of the European Union and thus strictly protected in E.U. countries. Also, migratory bats benefit from some level of protection because they are covered by the UN Convention for the Protection of Migratory Species. Countries that have signed this convention are obliged to support conservation actions that are beneficial for migratory species. CITES (The Convention on International Trade in Endangered Species of Wild Fauna and Flora) protects threatened species through controls of international trade in speci- mens. The precarious conservation status of the flying foxes is apparent. Currently, Acerodon jubatus and ten Pteropus spp are on CITES Appendix I, with trade only permitted in exceptional circumstances, and the remaining Acerodon and Pteropus species on Appendix II, by which trade is controlled to avoid utilization incompat- ible with their survival. 7 1 Bats in the Anthropocene 1.4 About This Book The idea to publish a book about bat conservation was stimulated by the “3rd International Berlin Bat Meeting: Bats in the Anthropocene” in 2013. The overall goal is to provide a summary of the major threats bats are facing in a rapidly chang- ing world. The book is organized in four major sections: (1) bats in anthropogen- ically-shaped landscapes, (2) emerging diseases, (3) human–bat conflicts, and (4) conservation approaches. The basic concept of chapters in all of these sections is to review the literature that is available in peer-reviewed journals. We are aware that many topics related to bat conservation have also been addressed in brochures or books published by non-governmental or governmental organizations. Sometimes these sources have been cited in the corresponding chapters, yet in most cases authors of this book have focused on the aforementioned sources of information. From our editorial perspective, the chapters cover the majority of relevant top- ics in bat conservation. However, we acknowledge that at least three topics are missing in this book. First, this book misses a chapter on “bats and global cli- mate change,” because Jones and Rebelo (2013) published a recent review on this topic and the body of literature about this topic has not largely increased since then. Second, we did not commission a chapter on “Bats and chemical pollut- ants,” as current knowledge of heavy metals was recently synthesized by Zukal et al. (2015) and information for other pollutants is sparse. That said, the subject is referenced in several chapters (Williams-Guillen et al. 2015; Korine et al. 2015; Voigt et al. 2016). Third, we did not include a chapter on “island bats,” although many of them are endangered and some even are threatened by extinction, as Fleming and Racey (2010) provide a detailed overview of this topic in their recent book. Finally, authors integrate successful interventions into their accounts and make specific recommendations for future research, but additional evidence- based evaluations of the success of conservation interventions per se are found in Berthinussen et al. (2014). The Anthropocene has gained momentum. It is a geological epoch that is not in equilibrium but is constantly changing by the action of mankind. For a handful of bat species anthropogenic changes may prove beneficial, but for the vast majority our actions precipitate drastic population declines that must be slowed if we are to conserve the extraordinary diversity of this unique mammalian order. We hope that this book will stimulate new directions for research and support conservation interventions that will keep the night sky alive with bats in the Human Epoch. Acknowledgements We would like to acknowledge the financial support provided by the Leibniz Institute for Zoo and Wildlife Research in Berlin, Germany, by EUROBATS and a National Science Foundation grant to the Southeast Asian Bat Conservation Research Unit (NSF Grant No. 1051363) that enabled us to publish this book as an open-access electronic book. We thank Mark Brigham, Anne Brooke, Justin Boyles, Gabor Csorba, Brock Fenton, Jorge Galindo-González, Chris Hein, Carmi Korine, Allen Kurta, Pia Lentin, Herman Limpens, Lindy Lumsden, Jörg Müller, Alison Peel, Paul Racey, Hugo Rebelo, DeeAnn Reeder, Scott Reynolds, Danilo Russo, Armando Rodríguez-Durán, N. Singaravelan, Vikash Tatayah, Peter Taylor, and numerous anonymous reviewers for providing constructive comments on chapters of this book. 8 C.C. Voigt and T. Kingston References Barclay RMR, Harder LD (2003) Life histories of bats: life in the slow land. In: Kunz TH, Fenton MB (eds) Bat ecology. 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Springer International AG, Cham, pp 151–178 Zukal J, Pikula J, Bandouchova H (2015) Bats as bioindicators of heavy metal pollution: history and prospect. Mamm Biol-Z Saugertierkd 80:220–227 Part I Bats in Anthropogenically Changed Landscapes 13 Chapter 2 Urbanisation and Its Effects on Bats—A Global Meta-Analysis Kirsten Jung and Caragh G. Threlfall © The Author(s) 2016 C.C. Voigt and T. Kingston (eds.), Bats in the Anthropocene: Conservation of Bats in a Changing World , DOI 10.1007/978-3-319-25220-9_2 Abstract Urbanisation is viewed as the most ecologically damaging change to land use worldwide, posing significant threats to global biodiversity. However, studies from around the world suggest that the impacts of urbanisation are not always negative and can differ between geographic regions and taxa. Bats are a highly diverse group of mammals that occur worldwide, and many species per- sist in cities. In this chapter, we synthesise current knowledge of bats in urban environments. In addition, we use a meta-analysis approach to test if the general response of bats depends on the intensity of urbanisation. We further investigate if phylogenetic relatedness or functional ecology determines adaptability of spe- cies to urban landscapes and if determining factors for urban adaptability are con- sistent worldwide. Our meta-analysis revealed that, in general, habitat use of bats decreases in urban areas in comparison to natural areas. A high degree of urbani- sation had a stronger negative effect on habitat use compared to an intermediate degree of urbanisation. Neither phylogenetic relatedness nor functional ecology alone explained species persistence in urban environments; however, our analy- sis did indicate differences in the response of bats to urban development at the family level. Bats in the families Rhinolophidae and Mormoopidae exhibited a negative association with urban development, while responses in all other fami- lies were highly heterogeneous. Furthermore, our analysis of insectivorous bats K. Jung ( * ) Institute of Evolutionary Ecology and Conservation Genomics, University of Ulm, Ulm, Germany e-mail: kirsten.jung@uni-ulm.de C.G. Threlfall ( * ) School of Ecosystem and Forest Sciences, University of Melbourne, Melbourne, Australia e-mail: caragh.threlfall@unimelb.edu.au 14 K. Jung and C.G. Threlfall revealed that the adaptability of individual families, e.g. Emballonuridae and Vespertilionidae, to urbanisation is not consistent worldwide. These results sug- gest that behavioural and/or morphological traits of individual species may better determine species’ adaptability to urban areas, rather than phylogenetic or func- tional classifications, and that driving factors for species adaptability to urban areas might be regionally divergent. We thus argue that future research should focus on behavioural and morphological traits of bats, to assess if these determine urban adaptability in this species-rich group of mammals. 2.1 Introduction 2.1.1 The Urban Context Urbanisation results in extreme forms of land use alteration (Shochat et al. 2006; Grimm et al. 2008). In the last century, the human population has undergone a transition in which the majority of people now live in urban rather than rural areas (UNPD 2012). The rate of change at which urban areas are evolving due to natural population growth is dramatic, including significant rural-to-urban migration and spatial expansion (Grimm et al. 2008; Montgomery 2008; UN 2012; McDonnell and Hahs 2013). In the last 50 years, the global human population in urban areas increased from 2.53 to 6.97 billion people (UNPD 2012). Yet human pres- sure resulting from urbanisation is not uniformly distributed on the planet. While urbanisation in the developed countries is slowing down slightly, it is increasing rapidly in developing countries of Asia, Africa, Latin America and the Caribbean, many of which harbour hotspots of biodiversity (Myers et al. 2000). In addition, over half of the urban population growth is projected to occur in smaller towns and cities (UN 2012). This implies that urbanisation is not a locally concentrated event, it is rather a fundamentally dispersed process and a happening worldwide (McDonald 2008). The ecological footprint of cities reaches far beyond their boundaries (McGranahan and Satterthwaite 2003; McDonald and Marcotullio 2013). Effects of cities operate from local (e.g. through urban sprawl) to global scales (e.g. through greenhouse gas emission) (McDonald et al. 2008), and act both directly, through expansion of urban areas, and indirectly through growth in infrastructure and changes in consumption and pollution (McIntyre et al. 2000; Pickett et al. 2001). Apart from the obvious loss in natural area, expansion of cities also impacts the surrounding rural and natural habitats through increased fragmentation, and edge effects with increasing temperature and noise levels, which together intro- duce new anthropogenic stressors on fringe ecosystems (Grimm et al. 2008) and nearby protected areas (McDonald et al. 2008; McDonald and Marcotullio 2013). However, despite the radical land transformation and habitat loss incurred through urbanisation, many species (native and introduced) can still persist in urban envi- ronments and some even experience population increases (McKinney 2006). This