First records of Hylaeus ( Paraprosopis ) pictipes Nylander , 1852 ( Hymenoptera : Colletidae ) in North America

Cavity-nesting bees, such as members of the genus Hylaeus Fabricius (Hymenoptera: Colletidae), are prone to accidental introductions by human-mediated transportation. The first records of Hylaeus (Paraprosopis) pictipes Nylander, 1852 in North America are reported. Hylaeus pictipes is a European species newly discovered from Mississauga, Ontario, Canada; Meadville, Pennsylvania, USA; and Cleveland, Ohio, USA. Notes on its identification and a discussion of exotic bees in North America are provided.

Collections and photographs of bees were made in a suburban area of Mississauga,Ontario,Canada,(43.5883,) on the afternoon of 17 August 2015 (Fig. 2).Specimens of Hylaeus were collected from Daucus carota L. in the backyard of a private residence using an aerial net and cyanide kill jar.A single male of H. pictipes was collected and the specimen is deposited at the J. B. Wallis/R.E. Roughley Museum of Entomology (JBWM), University of Manitoba (JBWM0363000).Twentyone additional specimens were collected on 5 August 2016 in an adjacent parking lot (43.5885, -79.6037), using identical techniques.These specimens are also deposited at the JBWM (JBWM0363001-JBWMJ0363019) with the exception of 1 male and female deposited at the A. J. Cook Arthropod Research Collection, Michigan State University (MSUC; no voucher numbers).Five additional specimens, 1 female and 4 males, from Meadville, Pennsylvania (41.6516, -80.1461) are also at JBWM (JBWM0363020-JBWM0363024).These specimens were brought to the attention of the lead author by Sam Droege  the R package maps (Becker et al. 2016) with historical specimen records from GBIF.org (28 May 2016), GBIF Occurrence Download http://doi.org/10.15468/dl.logdbd,and analyzed in R (R Core Team 2015) using the packages ggplot2 (Wickham 2009) and rgdal (Bivan et al. 2016).
Images of the live bee (Fig. 3) were taken in the field using a Canon T3i with a 100 mm macro lens and 25 mm extension tube.Subsequently, auto-montaged photos of preserved specimens were taken using a Canon 70d with MPE-65 mm macro lens using a Stackshot system and combined with Zerene Stacker.Additional material collected from the native range was used for comparison.
A single male specimen was collected and photographed (Fig. 3) in 2015.At the time of collection, the novelty of the specimen was not recognized, so additional collections were not attempted.Only when the specimen was examined at a later date was it noted that it did not match any known specimen from the region (Mitchell 1960;Snelling 1966Snelling , 1970;;Romankova 2007).The original male specimen was tentatively identified as H. pictipes using Koster (1986).Owing to uncertainty in the identification, images of the specimen were shared with local experts before the specimen, with genitalia dissected (Fig. 4A), was ultimately sent to HHD for confirma-   To aid in identification, the genitalia and hidden sternites of the original male collected were dissected out and glued to a small card below the specimen (Fig. 4A).The pointed structure of the gonoforceps are characteristic of this species (Dathe 1980;Dathe et al. 2016).Males of H. floridanus differ in the shape of S8, having the paired apical lobes less even in size, the posterior most lobe being much larger (Mitchell 1960;Snelling 1970).In H. pictipes, the paired lateral lobes of S8 are similar in size (Fig. 4).
The female would run to H. floridanus in Mitchell (1960), but European specimens with a black pronotal collar and maculated tegula would likely run to H. (Hylaeus) mesillae (Cockerell, 1896) cressoni (Cockerell, 1907) (as H. cressoni) or near H. (H.) verticalis (Cresson, 1869).In Romankova (2007), the male would likely run to H. (Prosopis) modestus Say, 1837, but the female would not key out well to any species.The female can be separated from other Ontario species by the facial fovea curving strongly away from the compound eye, ending at about the midpoint between the eye and lateral ocellus.Most North American females examined have distinct yellow maculations on the clypeus (Fig. 7A), but this is variable for the species (Fig. 7B).The female can be distinguished from H. floridanus using the following characters: paraocular maculation truncated below, not reaching to mandible; mesopleuron and mesoscutellum appearing smoother due to less microsculpture.(Mitchell 1960;Snelling 1970).The species does not match any of the species of H. (Paraprosopis) included in Snelling (1970).The specimens were compared directly to 10 species of North American H. (Paraprosopis) identified by Roy Snelling deposited at MSUC, including all species previously known from Canada and the eastern United States.In the male, the pale off-white maculations of the face in combination with the maculations on the scape (Fig. 5A) are reminiscent of H. (Hylaeus) leptocephalus (Morawitz, 1871), an apparently naturalized species in North America that originated in the  The first records of Hylaeus pictipes in the New World, collected from Mississauga,Ontario,Canada,(43.5883,43.5885,,and Meadville, Pennsylvania and Cleveland Ohio, USA are presented here (Fig 1).The native range of H. pictipes is largely restricted to more northern parts of Europe, up to 60° N in Sweden, extending east to the Caucasus (Dathe et al. 2016) and south to the Iberian Peninsula.These new records are more than 4000 km from the nearest record in the Azores (Weissmann et al. 2017).The wide distribution of H. pictipes in Europe (Fig. 2) suggests it may thrive under a breadth of climate conditions in North America.Hylaeus pictipes is polylectic in its native range, foraging on at least seven plant families (Koster 1986;Westrich 1990)     a broad range of existing cavities, including beetle burrows, hollow and pithy stem, earthen holes, and man-made materials (Scheuchl & Willner 2016).Exotic Hylaeus (Spatulariella) species that have been recently recorded from eastern North America (Ascher 2001;Ascher et al. 2006), have expanded their range extensively in less than a decade (Sheffield et al. 2011;Ksiazek et al. 2014).Hylaeus leptocephalus has been in Canada and the United States for at least a century and has spread across the continent (Snelling 1970).Based on other exotic cavity-nesting bees introduced to North America (Hinojosa-Díaz 2008; Gibbs & Sheffield 2009;Strange et al. 2011), these range expansions of exotic Hylaeus are expected to continue.The first recognized specimen of H. pictipes was in 2015, but it is now known from two additional sites south of Lake Erie separated by more than 200 km from the Mississauga records.The major trade route from the St. Lawrence River through Lake Ontario and Lake Erie is an important source for non-native species (Pagnucco et al. 2015), and other new records of adventive bees have been first discovered in this region (Sheffield et al. 2010(Sheffield et al. , 2011;;Martins et al. 2017;Normandin et al. 2017).A DNA-barcoded specimen of H. pictipes from southern California (www.boldsystems.org)requires additional verification, but if validated would indicate a much more extensive range for this species in North America.
The abundance of exotic species in collections has risen (Bartomeus et al. 2013), and this may come at the expense of native species.The ecological impacts of introduced bees remain poorly studied (Goulson 2003), although there have been reported cases of direct competition for nesting sites between exotic and native species (Laport & Minckley 2012;Roulston & Malfi 2012).Competition for floral resources and nesting sites creates potential for negative impact on native species (Goulson 2003).Introduction of pests and pathogens with exotic species is of concern (Hedtke et al. 2015;Cameron et al. 2016), but there are no studies of pathogens in introduced Hylaeus species.Should they occur, closely related species are most likely to be susceptible to introduced pathogens.Additional monitoring in the Great Lakes Region and urban areas of neighbouring states is needed to monitor the potential spread of this introduced species and to better document the current state of the native Hylaeus fauna.

Figure 2 .
Figure 2. New North American distribution records of Hylaeus (Paraprosopis) pictipes Nylander in southern Ontario, Pennsylvania, and Ohio.New York State and Michigan included for reference.
Palaearctic region over a century ago(Snelling 1970;Sheffield et al. 2011).However, in H. pictipes the structure of the face and fine punctation of the mesopleuron are quite unlike the longer face and coarse punctation of H. leptocephalus.The male would run to H. (Prosopis) affinis (Smith, 1853) inMitchell (1960) based on the pale scape.However, some European H. pictipes have the scape dark (Fig.5B), which would run to H. (Paraprosopis) floridanus.
tion.In 2016, after the identification of the original specimen, 13 additional males (Figs.5A, 6) and 8 females (Figs.7A, 8) were collected from a neighbouring lot to the original collection site.Both sexes key to the subgenus H. (Paraprosopis) in subgeneric keys to the Hylaeus of the Western Hemisphere and Palaearctic Region (Michener 2007).Sixteen members of the subgenus Hylaeus (Paraprosopis) are native to Canada and the United States, but are primarily restricted to the western United States (Snelling 1970; Ascher & Pickering 2016).Three native species of H. (Paraprosopis) occur in Canada, H. coloradensis (Cockerell, 1896), H. nevadensis (Cockerell, 1896), and H. wootoni (Cockerell, 1896), but none of these are found in Ontario or the eastern United States.In eastern North America, the native species H. (Paraprosopis) floridanus (Robertson, 1893) ranges north to states adjacent to Ontario, including Michigan, and H. (Paraprosopis) georgicus (Cockerell, 1898) is native to southeastern United States