Global invasion by Anthidium manicatum (Linnaeus) (Hymenoptera: Megachilidae): assessing potential distribution in North America and beyond James P. Strange, Jonathan B. Koch, Victor H. Gonzalez, Lindsay Nemelka & Terry Griswold Biological Invasions ISSN 1387-3547 Volume 13 Number 9 Biol Invasions (2011) 13:2115-2133 DOI 10.1007/s10530-011-0030-y 1 23 Author's personal copy Biol Invasions (2011) 13:2115–2133 DOI 10.1007/s10530-011-0030-y ORIGINAL PAPER Global invasion by Anthidium manicatum (Linnaeus) (Hymenoptera: Megachilidae): assessing potential distribution in North America and beyond James P. Strange • Jonathan B. Koch • Victor H. Gonzalez • Lindsay Nemelka • Terry Griswold Received: 8 July 2010 / Accepted: 20 May 2011 / Published online: 22 June 2011 Ó Springer Science+Business Media B.V. (outside the USA) 2011 Abstract The wool carder bee, Anthidium manica- model predicted a much broader distribution of A. tum, is the most widely distributed unmanaged bee in manicatum (214% increase); whereas, the South the world. It was unintentionally introduced to North American model predicted a narrower distribution America in the late 1960s from Europe, and subse- (88% decrease). The poor predictive power of the quently, into South America, New Zealand and the latter model in estimating suitable habitats in the Canary Islands. We provide information on the local invasive South American range of A. manicatum distribution, seasonal abundance and sex ratio of suggests that the bee may still be limited by the A.manicatum from samples collected in an intensive bioclimatic constraints associated with a novel envi- two-year survey across Utah, USA. Anthidium man- ronment. Estimates of niche similarity (D) between icatum was detected in 10 of the 29 Utah counties, the native and invasive models find that the North largely in urban and suburban settings. Combining America bioclimatic niche is more similar to the presence-only and MaxEnt background data from bioclimatic niche of the native model (D = 0.78), literature, museum databases and new records from whereas the bioclimatic niche of the South America Utah, we constructed three species distribution mod- invasion is relatively dissimilar (D = 0.69). We els to examine the potential distribution of A. manic- discuss the naturalization of A. manicatum in North atum in its native Eurasian range as well as invaded America, possibly through punctuated dispersal, the ranges of North and South America. The A. manic- probability of suitable habitats across the globe and atum model based on locality and background data the synanthropy exhibited by this invasive species. from the species’ native range predicted 50% of the invasive records associated with high habitat suit- Keywords Anthidium manicatum Á Invasion ability (HS C 0.90). The invasive North American dynamics Á Species distribution modeling Á Synanthropy Á MaxEnt background data J. P. Strange (&) Á J. B. Koch Á V. H. Gonzalez Á T. Griswold USDA-ARS Pollinating Insects - Biology, Management and Systematics Laboratory, Utah State University, Introduction 261 BNR, Logan, UT 84322-5310, USA e-mail: [email protected] Bees (Hymenoptera: Apoidea) are among the most important pollinators of many flowering plants J. B. Koch Á L. Nemelka Biology Department, Utah State University, including agricultural crops through which they 5305 Old Main Hill, Logan, UT 84322-5305, USA contribute an estimated one in three bites of food 123 Author's personal copy 2116 J. P. Strange et al. consumed by humans (Buchmann and Nabhan 1996). manicatum (Linnaeus), is a solitary bee in the family While several bee species have been intentionally Megachilidae that has been unintentionally introduced transported around the globe for pollination services into several regions of the world, and has recently [e.g., A. mellifera L., B. terrestris L., M. rotundata shown rapid expansion in its non-native geographic (Fabricius)], many other species across multiple distribution (Smith 1991; Miller et al. 2002; Hoebeke families are finding their ways into novel environ- and Wheeler 2005; Maier 2005; Zavortink and Shanks ments through accidental and indirect introductions 2008; Gibbs and Sheffield 2009). Females and males [e.g., M. sculpturalis Smith, A. oblongatum (Illiger), of A. manicatum, like many other anthidiines, are H. hyalinatus (Smith)]. There is growing concern that conspicuous because of their black and yellow-striped the shuffling of bee pollinators across habitats may abdomen and robust body form. In its invasive range facilitate the spread of novel bee pathogens (Goka males can be distinguished from native anthidiines by et al. 2001; Goulson 2004; Colla et al. 2006) and the distinctive series of protruding spines on the provoke competitive interactions with native bees for posterior segments of their abdomen (Fig. 1). Males floral resources and habitat (Roubik 1980; Goulson are territorial and aggressively defend mating sites 2003; Schmid-Hempel et al. 2007). While the from intruders by employing the abdominal spines to Africanized honey bee invasion of the New World break or disable the intruders’ wings during aerial may be the most dramatic example of the unintended battery (Pechuman 1967; Wirtz et al. 1992). Male A. consequences of bee movement (Roubik 1980), focus manicatum do not discriminate between conspecific has not been placed on the impacts of invasions by and heterospecific intruders and are often seen unintentionally introduced bees, nor have the under- patrolling flowers and attacking other bees that enter lying factors that facilitate their range expansion been into their territory (Kurtak 1973; Severinghaus et al. studied. 1981). This behavior is of particular interest because To date, most research has focused on the impacts some bees, particularly bumble bees, have been of managed bees on native bee communities documented to be deterred from foraging by A. (reviewed in Goulson 2003), while virtually nothing manicatum (Pechuman 1967; Comba et al. 1999). is known about the ecological ramifications of unin- Because of their distinctive body form and aggressive tentionally introduced bees (but see Severinghaus behavior A. manicatum are easily recognizable inhab- et al. 1981). The wool carder bee, Anthidium itants of residential gardens and are widely reported Fig. 1 Anthidium manicatum (#). Dorsal view of abdominal spines (right) used in aerial battery 123 Author's personal copy Global invasion by Anthidium manicatum (Linnaeus) 2117 on internet identification sites such as ‘‘Bug Guide’’ the bioclimatic profile associated with its native and ‘‘Discover Life’’, where pictures of the bees are distribution. posted along with location and date of sighting. Alternatively, the distribution of A. manicatum The native distribution of A. manicatum spans may be related to nesting behavior or foraging diet most of Europe, western Asia and coastal North (Comba et al. 1999; Corbet et al. 2001). Anthidium Africa. Anthidium manicatum was first observed in manicatum often nests in holes and cavities in wood or North America in 1963 in Ithaca, New York (Jaycox hollowed stems of plants, thus, facilitating dispersal 1967). Initial range expansion by A. manicatum was (Kurtak 1973). Female A. manicatum card fibers off of apparently not rapid; it was subsequently documented the leaves and stems of plants such as wooly hedge from Ontario, Canada in 1991 (Smith 1991), and nettle (Stachys byzantina K. Koch) and use the Pennsylvania in 1996 (Miller et al. 2002; Hoebeke material to line nest cells in cavities (Mu¨ller et al. and Wheeler 2005). However, since 1996 A. manic- 1996). The host-plants associated with the carding atum detection has increased dramatically; it is now behavior of A. manicatum seem to be restricted to the well documented throughout the eastern USA (e.g., family Lamiaceae, whereas pollen and nectar foraging Miller et al. 2002; Matteson et al. 2008; Tonietto and occurs on a greater array of plants, primarily plants in Ascher 2008), and most recently in California the family’s Lamiaceae, Fabaceae and Scrophularia- (Zavortink and Shanks 2008), Colorado, Idaho and ceae (Kurtak 1973, reviewed in Zavortink and Shanks western Canada (Gibbs and Sheffield 2009). In less 2008). However, it is still unknown to what extent than 50 years A. manicatum has dispersed across A. manicatum will utilize novel plants for nest- North America, primarily documented in major urban building or larval provisions in invaded environments, areas. Concurrently, it has colonized large portions of or whether the species range is limited by the presence southern Brazil, Peru, Suriname, Argentina, Para- of introduced European plant species. guay, Uruguay, New Zealand, and the Canary Islands During 2007 and 2008 the Utah Department of (Hoebeke and Wheeler 2005; Gibbs and Sheffield Agriculture and Food (UDAF) conducted a statewide 2009), making it the most widely distributed Anthi- monitoring and eradication program of the invasive dium in the world and the most widespread unman- Japanese beetle (Popillia japonica Newman Coleop- aged bee species. Records of A. manicatum from tera: Scarabaeidae) in response to beetle detections in central Asia are known from the 1960s (Wu 2005), Orem, Utah, USA. The network of baited beetle traps but it is unclear if these records are due to a recent deployed throughout the state, particularly concen- range expansion, or are from poorly documented trating on populated areas, resulted in large captures areas of their native range. of non-target insects. These unintentionally trapped Despite the widespread distribution of A. manic- insects, termed ‘‘by-catch’’, were primarily composed atum, little is known about the invasive potential of of Coleoptera, Lepidoptera and Hymenoptera (espe- this bee to colonize new areas. As recently as 2009 cially bees) (the authors, pers. obs.). Anthidium A. manicatum was called ‘‘adventive’’ (Gibbs and manicatum was found among the by-catch, providing Sheffield 2009), indicating that it was newly colo- an opportunity to survey from large portions of Utah. nizing and not yet well established in North America. Here we report on the distribution, seasonal While the bee occupies a large and diverse habitat in abundance and sex ratio of A.