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Hall & Ascher: Bees in Florida Natural Areas 609

SURVEYS OF BEES (: APOIDEA: ANTHOPHILA) IN NATURAL AREAS OF ALACHUA COUNTY IN NORTH-CENTRAL FLORIDA

H. GLENN HALL1 AND JOHN S. ASCHER2 1Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611-0620 E-mail; hgh@ufl.edu

2Division of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024-5192 E-mail; [email protected]

ABSTRACT

Bee surveys were conducted in Alachua County, Florida, at 1 to 6 sites within each of 4 nat- ural areas including 2 large state preserves. Bees were collected passively with colored cups and actively with nets. A total of 2,590 bees were captured belonging to 34 genera and 111 species. Of the 5 bee families found, was represented by the most species, whereas the 6 most numerous species were Halictidae. Six species are new state records for Florida. Males of the rarely-collected Florida endemic bee Stelis ater Mitchell were discovered, and this species is reported for the first time as a cleptoparasite of Osmia chalybea Smith, con- firmed by rearing from trap nests. Other potential new host-parasite associations are dis- cussed. Bee species lists and ecological patterns are compared with those from previous surveys in southern Florida. Distinctive characteristics of the north-central Florida bee fauna are discussed including the presence of both northern and peninsular species, subspe- cies, and populations.

Key Words: native bees, bee bowls, trap nests, cleptoparasitic bees

RESUMEN

Información acerca de la diversidad de abejas nativas en áreas naturales de la región Nor- Central del estado de Florida es limitada. Con el objetivo de disminuir este vacío, se llevó a cabo muestreos de abejas en 1 a 6 sitios dentro de cuatro áreas naturales, incluyendo dos re- servas estatales, localizadas en el condado de Alachua. Se utilizó un método pasivo de colecta con contenedores plásticos de colores y un método activo con redes entomológicas. Se captu- raron 2590 abejas en total, pertenecientes a 34 géneros y 111 especies. De las 5 familias do- cumentadas, la familia Apidae fue la más especiosa, mientras que las seis especies más abundantes pertenecieron a la familia Halictidae. De las especies atrapadas seis no han sido reportadas anteriormente en el estado de Florida. Además, fueron descubiertos machos de la especie endémica y raramente colectada Stelis ater Mitchell. Por primera vez se reporta a esta especie como cleptoraparásito de Osmia chalybea Smith; hallazgo verificado a través de su crianza en trampas nido. Se enumeran en esta publicación otras asociaciones parásito- hospedero, las cuales podrían ser nuevas. Las listas de especies de abejas y sus rangos de dis- tribución son comparados con muestreos previos realizados en el sur de Florida. Las carac- terísticas distintivas de las especies del área Nor-Central de Florida son discutidas, incluyendo la presencia de especies norteñas y peninsulares, subespecies, y poblaciones.

Translation provided by the authors.

To understand ecosystems, the biodiversity re- are largely dependent on the honey bee Apis mel- sponsible for their function must be documented. lifera L. Beyond their vital importance in natural Pollinators are an essential component of terres- communities, native bees contribute significantly trial ecosystems and of agricultural production. to agricultural pollination (Hurd & Linsley 1964; They are needed for the reproduction of three- Cane & Payne 1988; Thorp 2003; Torchio 2003; quarters of the world’s flowering plants, which in- Pascarella 2007; Winfree et al. 2008). Native bees cludes a large proportion of the food for wild ver- can be diverse in agricultural systems (Tuell et al. tebrates and two-thirds of the crop species (Com- 2009) and can potentially have a much greater mittee on the Status of Pollinators in North role in crop pollination, especially in light of re- America 2007). Because bees collect pollen as cent honey bee losses (Winfree et al. 2007). their source of protein, they are overall the most Natural bee populations and even species, es- efficient and important pollinators (Michener pecially endemic species, are threatened by sev- 2007). Agricultural crops that require pollination eral factors. The major threat is the loss of natu-

610 Florida Entomologist 93(4) December 2010 ral habitat, including nesting sites and floral re- tains a summary of bee species from the Ever- sources, due to land development and agricul- glades and from Dade and Monroe counties out- tural intensification (Buchmann & Nabhan 1996; side the park including those found by Graen- Allen-Wardell et al. 1998; Kearns et al. 1998; Kre- icher, those listed in other published records, and men et al. 2002; Klein et al. 2007; Goulson et al. present in regional museum collections. Deyrup 2008). The expansion of large-scale industrial ag- et al. (2002) list the bee species and their floral riculture eliminates natural areas, along with the hosts found at Archbold Biological Station on the associated bee diversity responsible for pollina- Lake Wales Ridge, Highlands County, south-cen- tion services that provide stability for the human tral Florida, and discuss the Florida bee fauna. food supply. Thus, conservation and restoration Unpublished surveys at the Tall Timbers Land are “crucial to the preservation of pollinator pop- Conservancy (TTLC) Research Station and the ulations and diversity” (Committee on the Status St. Marks National Wildlife Refuge, north and of Pollinators in North America, 2007). south of Tallahassee, respectively, and the Os- More than 19,500 described species of bees are ceola National Forest, west of Jacksonville, have known world-wide, about 3,500 of which occur in been done by John Pascarella (Georgia Southern the United States and Canada (Ascher & Picker- University, personal communication). Despite ing 2010). Much remains to be learned about bee these multiple surveys, the bees of large areas of , distribution, species richness, abun- Florida, including major regions such as north- dance, natural history, and floral hosts. The need central Florida, have not been studied ade- and importance of thorough systematic surveys to quately. increase our knowledge of bee faunas has been This report documents 111 bee species in 34 emphasized (Committee on the Status of Pollina- genera captured in natural areas of Alachua tors in North America 2007). Existing natural ar- County in the north-central region of peninsular eas, restored natural areas, farms, and other an- Florida. This survey represents the first in a se- thropogenic landscapes are all relevant sites for ries that will include additional areas and differ- such studies. Ongoing monitoring is required to ent plant communities in northern Florida. Bees detect changes over time, including decline or loss move between natural and cultivated areas (Kre- of species, which might occur as a consequence of men et al. 2002, 2004; Kohler et al. 2008; Winfree environmental disturbances and land use et al. 2008), so both types of landscapes are being changes (Deyrup et al. 2002). Given the ability of studied. bees to disperse and occupy newly available hab- itats, bee communities may serve as quality bio- MATERIALS AND METHODS indicators when comparing restored natural land with established natural areas. Collecting Methods and Preparation Florida is a unique place for biological studies, being a large peninsular extension of the eastern Most bees were caught in colored cups filled United States that extends from temperate into with soapy water after being attracted to the color subtropical climates. Florida has a large diversity and drowned in the water. Translucent plastic of natural plant communities and a major agri- 3.25 oz soufflé cups (Solo, Highland Park, Illinois) cultural industry. The state has had an increasing were used, painted either white, fluorescent yel- human population and intense development, low, or fluorescent blue as described in the which have slowed recently but will likely resume “Handy Bee Manual” (Droege 2008), with several in the future. Thus, there is urgency to document modifications. Spray paint was used (yellow and and preserve biodiversity in its remaining natu- blue, respectively, Krylon® 3104 and 3107 or ral areas. About 320 described and several unde- Ace® Brand, made by Krylon, I17052A00 and scribed bee species have been recorded from Flor- I19716A00; white, Krylon® Fusion 2320) (the in- ida (Mitchell 1960, 1962; Deyrup et al. 2002; up- side bottom rim was first lined with a water-based dated by Pascarella 2008; additional state records paint to prevent dissolving it with the spray and updated nomenclature in Ascher & Pickering paint) for cups used in 2007 and later, rather than 2010). The state list for Florida is relatively well the non-spray paint recommended in the manual. documented, but county records are fragmentary The cups were hung 10 to 20 cm above the ground (Pascarella 2008), and only a few systematic site vegetation on custom hand-bent wires stuck into surveys of the bee fauna have been conducted. the ground. Organic-based dishwashing soap was Graenicher (1930) studied the bees along the At- used, (Seventh Generation™, Free and Clear, lantic coast of southeastern Florida, mainly the Burlington, Vermont), 2 tablespoons per gallon of Miami area, extending north to Jupiter, south to water. The cups were placed about 5 m apart Homestead, and southwest to the former Royal along curved or straight lines, alternating 4-cup Palm State Park on the eastern side of what is groups of each color. For each collection, 24 to 48 now Everglades National Park. Pascarella et al. cups were placed at a site for about 30 h. Bees (2000) conducted a broader survey of bees at 4 re- were also collected by net in flight, usually while gions within the Everglades. Their report con- foraging on flowers, or caught with a hand-held

Hall & Ascher: Bees in Florida Natural Areas 611 vacuum (http://tech.groups.yahoo.com/group/bee- complexes border 2 sides of the lab area. Collec- monitoring/files/). Collected bees were frozen, tions were in an area of about 2 ha of open field of washed, and dried later as described in the herbaceous and woody vegetation. Half-hectare “Handy Bee Manual” with minor modifications. plots represent different succession time periods The bees were washed by vigorous shaking in a between prescribed burns and cutting. Adjacent large capped tube with soapy water for 2 min, fol- are upland mixed forest, upland pine forest, and a lowed by shaking in 95% ethanol for 2 min. After retention basin. brief blotting of the ethanol, the bees were placed Paynes Prairie Preserve State Park (PP) is a in a large small-mesh tea strainer, with a fash- 8,500-ha area, predominantly a large basin ioned metal lid, and vigorously shaken for 1.5 min marsh which periodically floods and drains over a hair-dryer at a low heat setting. through Alachua Sink. Two sites on the south Trap nests were either bundles of blocks (3/4” side, TW and BB (Tower and Bolen Bluff), are on square 6” long) with drilled holes ranging in di- the wet prairie margin between the lower basin ameter from 1/8” to 3/8” or commercially available marsh and the upland mixed forest. Site NE 14-hole, 5/16” diameter, “Binderboard®” nests (Northeast) on the north side and site LT (Lake (www.pollinatorparadise.com/). Trail) on the south side, both about 2 km from the basin boundary, are former pasture, cleared from Collection Areas and Sites what was originally sandhill and upland pine for- est, adjacent to existing upland mixed forest and This study surveyed bees in 4 natural areas of upland pine forest, respectively. Site PW (Pine Alachua County, Florida: Kanapaha Prairie (3 Woods) is within mesic flatwoods. Site RS (Re- sites), Paynes Prairie Preserve State Park (6 stored), about 3 km from the south edge of the ba- sites), San Felasco Hammock Preserve State Park sin marsh, is being restored to its former sandhill (6 sites), and the University of Florida Natural community and is adjacent to upland pine forest, Area Teaching Lab (1 site). Generally, most of the floodplain forest, and mesic flatwoods and near sites were sampled once a month from May upland mixed forest, baygall, and a small basin through Oct, 2006, and Mar and Apr, 2007. Some marsh. sites were sampled late Feb and mid Nov. A few San Felasco Hammock Preserve State park other sites were determined to be less favorable (SF) is about a 2,800-ha, mostly forested, natural and sampled only once or twice. Table 1 lists the area. Sites SH (Sandhill) and PL (Pine Land) are sites within the areas, their abbreviations, their in sandhill communities, adjacent to surrounding geographical coordinates (from Google Earth), upland pine forest and near upland mixed forest and the months and numbers of times when sam- beyond. Sites NS and EW are along clearings for pled. The areas and sites therein are described powerlines running north-south and southeast- below as the natural communities defined by the northwest, respectively, through upland mixed Florida Natural Areas Inventory (www.fnai.org/ forest. Site EW is adjacent to upland pine forest. natcomguide_update.cfm). In the following de- Site NS is on a slope descending to a nearby lower scriptions, “adjacent” means within 50 m; “near” basin swamp and bottomland forest to the north or “nearby” means within 500 m. and near site PL to the west. Site BA (Old Barn) Kanapaha Prairie (KP) is about a 300-ha basin is in a former pasture and ruderal area being re- marsh largely owned by the Conservation Fund. stored as a long-leaf pine forest, adjacent to up- It is surrounded by mesic hammock dominated by land mixed forest. Site IB (Itchy Bottom Lake) is large Live Oaks (Quercus virginiana Mill.) in in former pasture adjacent to upland mixed for- which there are homesites of 5 to 10 ha. Portions est, upland pine forest, and near a floodplain of the prairie periodically flood. Much of the prai- marsh. rie had been pasture of imported Bahia Grass (Paspalum notatum Flüggé) and is still used for RESULTS cattle grazing. Site W (West) is along the higher and drier elevations of the prairie adjacent to the In this study, a total of 2,590 bees were cap- hammock, distant from the cattle grazing area, tured in the Alachua County natural areas, be- and that has largely returned to a natural state. longing to 34 genera and 111 species, 1 of which is Site R (Road) is deeper within the basin marsh believed to be undescribed. Six percent of the spe- along a road and ditch between the grazed and a cies were in the family Colletidae, 15% in An- more natural part of the prairie. Site S (South) is drenidae, 23% in Halictidae, 25% in Megachil- along a road south and outside of the prairie, ad- idae, and 31% in Apidae. Eighty-eight percent of jacent to the mesic hammock and former pasture. individuals were female; 12% male. The list of the Ruderal areas and some homesites are nearby. species, including the earliest and latest dates NATL (The University of Florida Natural Area when captured, the areas and sites where cap- Teaching Laboratory, http://natl.ifas.ufl.edu/) is a tured (abbreviations from Table 1), and the num- 24-ha area at the southwest corner of the main bers of females and males captured in cups and on Gainesville campus. Major roads and building each of the plant species (names and abbrevia-

612 Florida Entomologist 93(4) December 2010

TABLE 1. BEE COLLECTION AREAS AND SITES.

Area-Site Geographical Collections Months (# from bee Abbreviation Area Site Name Coordinates bowls—# from flowers)/ Year1

KP-W Kanapaha Prairie West 29°33’13”N 5, 4, 8 (0-2), 9, 10 (0-2), 11 (0-1)/ 06 82°26’10”W 3, 4 (1-1), 6 (1-1), 10 (0-1)/ 07 4 (0-1), 7 (0-1), 9 (0-1)/ 08 KP-R Kanapaha Prairie Road 29°33’07”N 11 (0-1)/06, 10 (0-1)/ 07 82°25’42”W 3 (0-1), 4 (0-1), 5 (0-1)/ 08, 9 (0-1), 11 (0-2) / 09 KP-S Kanapaha Prairie South 29°32’03”N 5 (0-2), 6 (0-1), 9 (0-1)/ 08, 82°26’08”W 9 (0-1)/ 09 NATL UF Natural Area Teaching Lab 29°38’02”N 4, 5, 6, 9, 10 (2-1)/ 06 82°22’08”W 2 (1-1), 3, 4/ 07, 10 (0-1)/ 09 PP-BB Paynes Prairie Bolen Bluff 29°33’36”N 5/ 06 82°19’34”W PP-LT Paynes Prairie Lake Trail 29°32’20”N 4, 8/ 06 82°17’39”W PP-NE Paynes Prairie Northeast 29°36’32”N 6, 9 (1-1), 10 (2-3)/ 06 82°17’03”W 3, 5/ 07, 10 (0-1)/ 09 PP-PW Paynes Prairie Pine Woods 29°31’58”N 9/ 06 82°17’08”W PP-RS Paynes Prairie Restored Area 29°31’18”N 4, 8, 9, 11 (1-1)/ 06, 82°17’19”W 3 (2), 5, 6/ 07, 11 (0-1)/08, 3 (0-1)/ 09 PP-TW Paynes Prairie Tower 29°33’00”N 10/ 06 82°17’30”W 3, 5, 6/ 07 SF-BA San Felasco Old Barn 29°45’28”N 4/ 06 82°27’40”W 3, 4/ 07 SF-EW San Felasco East-West Powerline 29°43’36”N 5, 6, 8, 9, 10/ 06 82°26’01”W 3, 4/ 07 SF-IB San Felasco Itchy Bottom Lake 29°45’59”N 4 (1-1), 6/ 06 82°27’02”W 4/ 07 SF-NS San Felasco North-South Powerline 29°44’35”N 5, 6, 8, 9, 10/ 06 82°27’19”W 3, 4/ 07 SF-PL San Felasco Pine Land 29°44’34”N 5/ 06 82°27’41”W SF-SH San Felasco Sandhill 29°42’56”N 5, 6, 8, 9, 10/ 06 82°27’28”W 3, 4/ 07

1Parentheses indicate months when either more than 1 collection was made, or when collections were made also, or only, from flowers. tions in Table 2), is in Table 3. Eight species were about 25 bee species not found in this study, captured over a span of 6 months but were far which will be reported in a forthcoming article. more abundant during a segment of that time. From the spring of 2006 to the spring of 2007, For these species, the 2 or 3 months and the per- bee collecting was most consistently and system- centage of bees caught during that period that ex- atically done with colored cups, which were par- ceed 70% of the total are indicated in the “Re- ticularly useful at locations and at times with lit- marks” column. The species found in the county tle herbaceous flowering. Eighty percent of the previously or reported to be found throughout bees were caught in cups. As others have found Florida, according to Pascarella (2008), are indi- (Cane et al. 2000; Roulston et al. 2007; Wilson et cated by an “AC” in the “Remarks” column. In or- al. 2008), bees caught in the cups were not fully ganic farms of Alachua County, we have captured representative of the resident fauna. Some bee

Hall & Ascher: Bees in Florida Natural Areas 613

TABLE 2. PLANTS FROM WHICH BEES WERE COLLECTED. lieved to be a new, undescribed, species (Jason Gibbs, personal communication). The bee species Ba — Bidens alba (L.) DC. previously recorded from Alachua County to- Bl — Bidens laevis (L.) Britton et al. gether with those listed as “throughout Florida” Ca — Cirsium altissimum (L.) Spreng. (Pascarella 2008) include 73 species found in this Cd — Croptilon divaricatum (Nutt.) Raf. study and 67 not found. Thirty-seven additional Ch — Cirsium horridulum Michx. species found in this study are new county Io — Ilex opaca Aiton records, including the 6 new state records. Thus, a Ec — Euthamia caroliniana (L.) total of 177 bee species are now reported from Ex Porter and Britton Alachua County, which does not include those Em — Eupatorium mikaniodes Chapm. among the additional species found in organic Eq — Erigeron quercifolius Poir. farms, mentioned above. Hf — Helianthus floridanus A. Gray ex Chapm. In descending order, the following species were Hs — Heterotheca subaxillaris (Lam.) the most abundant, the first 6 of which were Hal- Britton and Rusby ictidae: Lasioglossum (Dialictus) reticulatum Pg — Pityopsis graminifolia (Michx.) Nutt. (Robertson) (527F 1M); Augochlorella aurata Rc — Rhus copallinum L. (Smith) (268F 1M); Lasioglossum (Dialictus) Rr — Raphanus raphanistrum L. Not native. puteulanum Gibbs (180F 1M); Lasioglossum (Di- Sd — Symphotrichum dumosum (L.) G. L. Nesom alictus) apopkense (Robertson) (146F); Agaposte- Ss — Solidago spp. L. mon (Agapostemon) splendens (Lepeletier) (114F Vac — Vitex agnus-castus L. Not native. 11M); Halictus poeyi (109F 11M); Melissodes (Me- Vb — Verbena brasiliensis Vell. Not native. lissodes) communis communis Cresson (78F Vg — Vernonia gigantea (Walter) 25M); Melissodes (Melissodes) bimaculata bimac- Trel. ex Branner and Coville ulata (Lepeletier) (78F 17M); Lasioglossum (Dial- Authorities from Wunderlin & Hansen (2003). ictus) nymphale (Smith) (80F 1M); Lasioglossum (Dialictus) pectorale (Smith) (53F 5M). Two Megachile Latreille species were caught species were seen on flowers but not in cups emerging from or flying over ground nests. Mega- placed nearby. Although the species are not chile (Megachiloides) rubi Mitchell had clustered, equally attracted to the colored cups, for each spe- but well-separated, nests. A single female of M. cies that is caught, the cups provide an objective (Acentron) albitarsis Cresson came from an iso- measure of their abundance at different locations. lated nest opening (see images by Tim Lethbridge From 2006 through 2009, bees also were captured of a similar nest believed to be of this species from in flight over flowers or nests, but this collecting Archbold Biological Station, bugguide.net/node/ was opportunistic, inconsistent, and done mainly view/375132). in the fall, when herbaceous flowers were most Traps nests were located at Kanapaha and abundant. The plants on which bees were caught Paynes Prairies primarily for a separate study, are listed in Table 2 (plant authorities from but the bee species that occupied the nests are Wunderlin & Hansen 2003). Most of the sampling mentioned here. Osmia (Helicosmia) chalybea sites were open areas near the edges of mixed Smith was the main bee species to construct cells pine-hardwood forests and wetlands, at bound- in the nests (the 5/16” and 3/8” diameter holes). aries between or near 2 or more natural commu- For the first time, this species was confirmed as a nities. Thus, the different collections of bees from host of Stelis ater which parasitized about a third the sites are not necessarily characteristic of sin- of the cells. Both species are univoltine and gle plant communities. emerge in the spring. The first males of S. ater Six species are new state records for Florida: were discovered earlier in this survey, captured in Andrena (Callandrena s.l.) asteroides Mitchell, cups. More detail about these species’ nests, im- Andrena (Iomelissa) violae Robertson, Xenoglossa mature stages, and other aspects of their associa- (Eoxenoglossa) kansensis Cockerell (discovery re- tion will be discussed in a separate report (Rozen ported by Hall (2010) included specimens from & Hall, in preparation). Megachile (Litomega- this study), Sphecodes antennariae Robertson, chile) mendica mendica Cresson constructed cells Osmia (Melanosmia) collinsiae Robertson, and in 1 to 2 burrows of about 10% of the trap nests. Nomada annulata Smith. The last 3 had been From 1 nest that had been placed in an emer- found previously only as far south as North Caro- gence cage while the cells were still sealed, the lina (Mitchell 1960, 1962). Three species and 1 first brood emerged within about 2 months after subspecies are Florida endemics: Lasioglossum cell construction began (timing was not closely (Dialictus) robertsonellum Michener, Stelis (Ste- monitored). Bee cleptoparasites emerged from lis) ater Mitchell, Epeolus floridensis Mitchell, the same nest, viz. 2 females of Coelioxys (Boreo- and Ceratina (Zadontomerus) dupla floridana coelioxys) sayi Robertson (established hosts Mitchell. The specimens listed as Lasioglossum Megachile (Litomegachile) brevis Say and M. (Dialictus) aff. raleighense (Crawford) are be- mendica) and 2 males of C. (Acrocoelioxys) doli- 614 Florida Entomologist 93(4) December 2010

Remarks4,5 AC; AC; 88% X-XI AC AC AC

3 Males 1Cd 1Em 2Ec 1Hs 1Sd 1Io AC 1Em 4Ec 3Sd 4Ss 4Em 3Ec 2Sd 14Ss 1Io 1Pg AC

Females 1Hs 1Pg 20Sd 1Sd 1Ec 1Pg 23Sd 1Ss 10Em 8Pg 6Sd 7Ss 1Ba 1Ba AC

Individuals collected Males 3 2Ba 1Ec 1 82 9 1 10Pg 4Pg

In cupsFemales On flowers 264 1Ca 2 Pg

SF-SH SF-PL

SF-NS SF-IB SF-EW SF-BA 2

PP-TW

PP-RS PP-PW Areas - Sites PP-NE , 2006-2009. PP-LT PP-BB see text. ; Osmia chalybea LORIDA , F NATL KP-S

OUNTY KP-R C

KP-W

LACHUA Late date of collection A OF

Early date of collection 8-III 7-XI 8-III 7-XI 7-III 1-VI 10-X 11-XI 13-IV 13-IV 19-X 11-XI 11-X 11-X 12-III 12-III AREAS

NATURAL

Say 1 IN

—new Florida record reported previously (Hall 2010). Mitchell 13-IV 13-IV Species CAUGHT Cockerell

(Cockerell) —first report of males this species and host association with EES 3. B Colletes simulans miamensis Mitchell Hylaeus ( Prosopis ) modestus Dieunomia ( ) heteropoda (Say) aurata Augochlorella (Smith) Colletes mandibularis Smith 6-VI 11-XI Colletes thysanellae Mitchell ) Hylaeus ( Paraprosopis georgicus Hylaeus ( Prosopis ) schwarzii ) pura ( Augochlora Augochlora pura (Say) Colletes brimleyi Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 3 6 8 10 2 4 5 7 9 1 T Hall & Ascher: Bees in Florida Natural Areas 615

4,5

Remarks Florida C C AC record AC AC AC AC

3 Males 2Ca 1Pg 1Sd 3Vac 1Vb 1Vg

Females 1Vb 1Rr 2Vac 1Rr 2Vac

Individuals collected Males 9A 41 1A 133 New 1 1Rc 1Sd AC In cups On flowers Females 10 1 22 1Io 2Rc 2Sd 108 2 2Pg 1Rc SF-SH SF-PL

SF-NS

SF-IB SF-EW SF-BA 2 , 2006-2009. PP-TW PP-RS LORIDA PP-PW , F Areas - Sites PP-NE

OUNTY PP-LT C

PP-BB see text. ; Osmia chalybea NATL

LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection

Early date of NATURAL

collection 10-III 16-IX 14-IV 14-IV 14-IV 6-V 14-IV 1-XI 12-III 4-XI 15-V 16-IX 23-II 11-XI IN

CAUGHT

EES 1 Mitchell 6-V 6-V —new Florida record reported previously (Hall 2010). ) B ( Paraugochlo- ( Paraugochlo- ( Paraugochlo- Species (Lepeletier) —first report of males this species and host association with ONTINUED 3. (C ropsis ) anonyma (Cockerell) Augochloropsis Augochloropsis Agapostemon ( ) splendens Sphecodes antennariae Robertson Sphecodes brachycephalus Mitchell Sphecodes heraclei ignitus Cockerell ropsis ) metallica (Fabricius) ropsis ) sumptuosa (Smith) Augochloropsis Augochloropsis Augochloropsis Sphecodes atlantis Augochlorella gratiosa (Smith)Augochlorella 27-IX 27-IX Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 12 15 17 18 19 13 14 16 11 T 616 Florida Entomologist 93(4) December 2010

Remarks4,5 C C AC AC; 95% III-V AC; 81% III-V AC 83% IV-V AC; AC; 86% IX-X 90% III-V

3 Males 2Ba 1Ec 2Pg 3Sd 1Vb 1Bl 3Rc AC

Females 2Eq 1Pg 7Sd 2Vb 6Vg 1 over ground

Individuals collected Males 3A 2A In cups On flowers Females 86 2 3Ba 1Ca 20 1 79 153 1Vac 5 11 1 146 180 1 523 1

SF-SH SF-PL SF-NS SF-IB

SF-EW SF-BA 2 , 2006-2009. PP-TW PP-RS LORIDA PP-PW , F

Areas - Sites PP-NE

OUNTY PP-LT C

PP-BB see text. ; Osmia chalybea

NATL

LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection 22-IX Early date of NATURAL

collection 8-III 2-XI 8-III 8-VI 7-III 2-XI 7-III 22-IX 9-III 11-XI 8-III 27-IX 14-VIII 14-IV 31-X 23-II 2-XI 14-IV 27-X IN

ra- CAUGHT

) EES 1 —new Florida record reported previously (Hall 2010). ) B (Robertson) Species (Crawford) (Mitchell) —first report of males this species and host association with ONTINUED 3. (C Lasioglossum ( Dialictus ) macoupinense Lasioglossum ( Dialictus ) nymphale (Smith) Lasioglossum ( Dialictus ) apopkense (Robertson) Lasioglossum ( Dialictus ) leviense Lasioglossum ( Dialictus ) pectorale (Smith) Lasioglossum ( Dialictus ) puteu- lanum Gibbs Lasioglossum ( Dialictus ) reticulatum (Robertson) leighense Lasioglossum ( Dialictus ) creber- rimum (Smith) Lasioglossum ( Dialictus ) aff. Halictus ( Odontalictus poeyi Lepeletier Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 24 25 21 23 26 27 29 22 28 20 T Hall & Ascher: Bees in Florida Natural Areas 617

Remarks4,5 C C C C AC New Florida record AC AC AC New Florida record

3 Males

Females 2Pg 26Sd 1Hs 13Pg 1Hs 1Pg AC Individuals collected Males 1A 2A 1A 11 1 21 1A In cups On flowers Females 4736 1Rc 10Bl

SF-SH SF-PL

SF-NS SF-IB

SF-EW

SF-BA 2 , 2006-2009. PP-TW

PP-RS LORIDA PP-PW , F Areas - Sites PP-NE

OUNTY PP-LT C

PP-BB see text. ; Osmia chalybea

NATL

LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection

Early date of NATURAL

collection 8-III 11-XI 8-III 27-III 9-III 9-III 6-V 27-X 12-III 2-XI 12-III 12-III 23-II 23-II 12-III 12-III 27-X 11-XI 28-III 28-III 22-X 7-XI IN

CAUGHT ) banksi

) macra EES 1 Robertson —new Florida record reported previously (Hall 2010). ) B Michener Species ) as- ( Callandrena s.l. ( Iomelissa ) violae Rob- ( Larandrena ) ( Leucandrena ) kri- ( Callandrena s.l. ( Holandrena ) ( Archiandrena ) fulvi- ( Callandrena s.l. Mitchell (Mitchell) —first report of males this species and host association with Smith ONTINUED 3. (C Lasioglossum ( Dialictus ) tar- ponense Andrena teroides Andrena ertson Andrena miserabilis Cresson Andrena Lasioglossum ( Evylaeus ) nelum- bonis (Robertson) Andrena Andrena pennis Andrena giana Robertson Lasioglossum ( Dialictus ) robertsonellum Andrena cressonii Mitchell Malloch Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 31 34 38 39 40 32 33 35 36 30 37 T 618 Florida Entomologist 93(4) December 2010

Remarks4,5 C AC AC

3 Males

Females 1Io 28Hs 1Pg 14Hs AC Individuals collected

Males 2 2Pg In cups On flowers Females 1A 1 1Io1 3Io11 AC 2 2 4 1Ch 2Ch 2

SF-SH SF-PL SF-NS SF-IB

SF-EW SF-BA 2 , 2006-2009. PP-TW PP-RS LORIDA PP-PW , F Areas - Sites PP-NE

OUNTY PP-LT C

PP-BB see text. ; Osmia chalybea

NATL

LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection

Early date of NATURAL

collection 6-IV 6-IV 6-V 6-V 13-IV 14-IV 21-X 27-X 19-IV 8-VI 27-X 27-X 27-X 27-X 14-IV 29-IV 30-III 30-III 29-IV 29-IV IN

CAUGHT

) barbara ) EES 1 —new Florida record reported previously (Hall 2010). ) B Species Viereck ( Lithurgopsis ) ( Scrapteropsis ) ( Scrapteropsis ) ( Melandrena ( Melandrena Viereck Viereck —first report of males this species and host association with Timberlake ONTINUED 3. (C Andrena atlantica Mitchell Andrena imitatrix Cresson ( Hexaperdita ) Perdita bishoppi Cockerell Lithurgus Smith gibbosus ) gerardiae ( Perdita Perdita Crawford Pseudopanurgus nebrascensis Michener muesebecki ( Hexaperdita ) Perdita nubila Andrena Andrena Perdita ( Alloperdita ) Perdita bradleyi Bouseman and LaBerge confederata Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 43 44 47 50 49 45 48 41 42 46 T Hall & Ascher: Bees in Florida Natural Areas 619

Remarks4,5 7 AC AC 6,7 New Florida record

3 Males 1Sd 1Vb AC 1Pg AC

Females 3Hs 3Vb 6Vg 1 from ground nest 1Vac1Pg 1Pg 1Vac Individuals collected

Males 2 2 In cups On flowers Females 13 3 152 2Ch 1Ch4 AC; 1Cd 2Ec 36 7

SF-SH SF-PL

SF-NS SF-IB

SF-EW

SF-BA 2 , 2006-2009. PP-TW PP-RS LORIDA PP-PW , F

Areas - Sites PP-NE

OUNTY PP-LT C

PP-BB see text. ; Osmia chalybea

NATL LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection

Early date of NATURAL

collection 7-III 28-III 1-XI 1-XI 7-III 10-IV 14-VIII 4-XI 27-X 27-X 29-V 2-XI 12-III 15-IV 14-IV 14-IV 23-II 6-V IN

CAUGHT

) ) albitarsis EES 1 ) —new Florida record reported previously (Hall 2010). ) B ( Loyolanthidium ) Mitchell Species Robertson ( Neotrypetes —first report of males this species and host association with ONTINUED 3. (C Osmia ( Melanosmia ) collinsiae Osmia ( Helicosmia ) Smith chalybea Cresson Megachile ( Acentron Megachile Stelis ( Dolichostelis louisae Cockerell Stelis ( ) ater Mitchell 9-III 9-III Anthidiellum perplexum (Smith) Hoplitis ( Alcidamea ) pilosifrons (Cresson) Osmia ( Melanosmia ) atriventris Cresson Heriades Osmia ( Melanosmia ) sandhouseae leavitti Crawford Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 58 56 60 52 53 51 55 57 54 59 T 620 Florida Entomologist 93(4) December 2010

Remarks4,5 AC 7 AC AC

3 Males 1Vac 4Ba 2Bl 1Pg 1Sd 16Vac 2Vb 1Vg 1Pg 1Vac 1Ba 1Vg 1Bl, 1 flying 1Bl, over ground

Females 5Rc 4Sd 5Vg 1Vac 2Vac 1Vac 1Ca 2Rc 1Vg ground ground nest 1 1 flying over 2 4 flying over Individuals collected Males 2 1 2 21 6Vac 1 5 2Ba 1Cd 1 1 1Vg 1Vac 1Rc 4Vac AC 24 5 In cupsFemales On flowers

SF-SH SF-PL SF-NS SF-IB SF-EW SF-BA 2 , 2006-2009. PP-TW PP-RS LORIDA PP-PW , F

Areas - Sites PP-NE

OUNTY PP-LT C

PP-BB see text. ; Osmia chalybea

NATL LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection

Early date of NATURAL

collection 8-VI 8-VI 6-V 11-XI 2-VI 2-XI 6-V 3-IX 2-VI 2-VI 28-III 23-IV 18-V 27-IX 30-III 11-XI 18-V 7-X 7-III 2-XI IN

) ) ) rubi ) CAUGHT

EES 1 —new Florida record reported previously (Hall 2010). ) B Say Species ) brevis ( Litomegachile —first report of males this species and host association with ONTINUED 3. (C Mitchell Megachile ( Chelostomoides Megachile ) con- ( Eutricharaea Megachile cinna Smith ) ( Leptorachis Megachile petulans Cresson ) tex- ( Litomegachile Megachile ana Cresson ( Megachiloides Megachile georgica Cresson ) men- ( Litomegachile Megachile dica mendica Cresson Megachile ( Chelostomoides Megachile campanulae (Robertson) ) ( Callomegachile Megachile sculpturalis Smith pseudobrevis Megachile Megachile ( Melanosarus Megachile xylocopoides Smith Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 63 64 65 68 69 67 62 61 66 70 T Hall & Ascher: Bees in Florida Natural Areas 621

Remarks4,5 7,9 C C AC AC 7,8 8 8 AC

3 Males 2Ba 1Sd AC;

Females 1Vac 2 from over Megachile rubi nests 1Vb 1Vac 3 from over Megachile rubi nests 1Ba

Individuals collected Males In cups On flowers Females 1A 2 1Vac 1Vac 1A 10

SF-SH SF-PL SF-NS

SF-IB

SF-EW SF-BA 2 , 2006-2009. PP-TW

PP-RS LORIDA PP-PW , F Areas - Sites PP-NE

OUNTY PP-LT C

PP-BB see text. ; Osmia chalybea

NATL LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection

Early date of NATURAL

collection 2-VI 2-IX 6-V 6-V 2-VI 2-VI 14-IV 15-V 13-IV 13-IV 29-V 2-XI 30-X 30-X 13-IV 23-IV 30-X 4-XI 22-IX 22-IX IN

) sayi CAUGHT ) ) doli- )

EES 1 L. —new Florida record reported previously (Hall 2010). ) B Species —first report of males this species and host association with ONTINUED Fox 3. (C Megachile ( Xanthosarus ) Megachile addenda Cresson Coelioxys ( ) Baker mitchelli Coelioxys ( Acrocoelioxys chos Robertson micans Lepeletier Coelioxys ( Boreocoelioxys Coelioxys ( Haplocoelioxys ) mexi- cana Cresson Coelioxys ( Xerocoelioxys galactiae Mitchell Xylocopa ( Schonnherria Xylocopa ( Xylocopoides ) virginica Megachile ( Sayapis ) Megachile policaris Say Ceratina ( Ceratinula ) Smith cockerelli Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 72 75 73 74 76 77 78 79 71 80 T 622 Florida Entomologist 93(4) December 2010

Remarks4,5 8 AC AC AC; New Florida record

3 Males 9Vb

Females over ground 1 flying over ground 1Vg 1Vac 2Sd 1Vac 1Sd AC 2Sd1Vb 1Ec1Vb AC Individuals collected

Males 1 In cups On flowers Females 11 2 flying 2 41 6Pg 1Sd 1Hs 1Pg AC 1Bl 25 4 1Sd

SF-SH SF-PL

SF-NS SF-IB SF-EW SF-BA 2 , 2006-2009. PP-TW PP-RS LORIDA PP-PW , F Areas - Sites PP-NE

OUNTY PP-LT C PP-BB see text. ; Osmia chalybea NATL LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection

Early date of NATURAL

collection 2-VI 1-XI 2-IX 2-IX 7-III 11-XI IN

) CAUGHT

Mitchell 25-X 1-XI EES 1 Mitchell 4-V 18-V Mitchell 27-X 11-XI Mitchell 22-X 2-XI —new Florida record reported previously (Hall 2010). ) B Species Mitchell —first report of males this species and host association with ONTINUED 3. (C Epeolus australis MitchellEpeolus bifasciatus Cresson 14-IV 2-VI 14-IV Epeolus floridensis 2-VI Epeolus glabratus Cresson 30-V 2-VI Nomada annulata Smith 14-IV 14-IV Nomada fervida Smith Nomada texana Cresson georgicus Triepeolus quadrifasciatus at- Triepeolus 4-XIlanticus 4-XI Epeolus carolinus Ceratina ( Zadontomerus dupla floridana Mitchell rugosus Triepeolus Nomada australis Mitchell 24-IV 24-IV Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 89 90 92 93 82 84 85 86 87 91 81 88 83 T Hall & Ascher: Bees in Florida Natural Areas 623

Remarks4,5 C C AC AC 71% IX-X 92% IV-VI

3 Males 3Hs 1Pg AC 1Vb

Females 3Pg 6Sd 2Vg 1Hf Individuals collected

Males 1 1A In cups On flowers Females 1A 11 3 1 3Ba 6Hs 21 2 1 4Vg 2Vac 5Vb AC 77 16 1Pg69 15 1Vg 8Vac 1Vac 3Vb 7Vac AC; AC;

SF-SH SF-PL

SF-NS SF-IB SF-EW

SF-BA 2 , 2006-2009. PP-TW PP-RS LORIDA PP-PW , F Areas - Sites PP-NE

OUNTY PP-LT C

PP-BB see text. ; Osmia chalybea

NATL LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection

Early date of NATURAL

collection 7-III 1-XI 8-VI 8-VI 19-V 19-V 23-II 7-XI 14-IV 31-X 30-VII 3-IX 14-IV 3-IX 19-IV 1-VI 22-IX 11-XI 14-VIII 16-IV IN

) bolto- CAUGHT

) ) tepan- ) ) comp- ) EES 1 —new Florida record reported previously (Hall 2010). Cresson 6-V 6-V ) B Species bimaculata ( Melissodes ( Melissodes ( Melissodes ( Melissodes ( Apomelissodes ) api- ( Eumelissodes (Cresson) ( Florilegus communis Cresson —first report of males this species and host association with ONTINUED Robertson 3. (C Florilegus condignus bimaculata (Lepeletier) eca Cresson Melissodes Melissodes Melissodes Svastra ( Epimelissodes ) aegis (LaBerge) communis toides Melissodes Melissodes Svastra ( Epimelissodes ) atripes georgica (Cresson) Epeolus pusillus Melissodes cata Lovell and Cockerell Melitoma taurea (Say) niae Robertson Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 96 98 99 102 103 100 101 104 94 97 95 T 624 Florida Entomologist 93(4) December 2010

Remarks4,5 10 11 C C AC; AC; New Florida record AC AC

3 Males 1Ca AC

Females 5Rr

Individuals collected Males In cups On flowers Females 2 1A 2 61 1A

SF-SH SF-PL

SF-NS SF-IB SF-EW SF-BA 2 , 2006-2009. PP-TW

PP-RS LORIDA PP-PW , F

Areas - Sites PP-NE

OUNTY PP-LT C PP-BB see text. ; Osmia chalybea

NATL

LACHUA KP-S A

OF KP-R

KP-W AREAS Late date of collection

Early date of NATURAL

collection 28-IV 28-IX 27-V 2-XI 23-II 23-II 27-V 27-V 12-III 12-III 19-V 22-VI IN

CAUGHT

EES 1 —new Florida record reported previously (Hall 2010). ) B Species (DeGeer) (DeGeer) Cockerell —first report of males this species and host association with ONTINUED 3. (C Bombus ( Pyrobombus ) bimaculatus Cresson Apis ( ) mellifera L. Bombus ( Pyrobombus ) impatiens Cresson ) pen- Bombus ( Thoracobombus sylvanicus Bombus ( Cullumanobombus ) griseocollis Xenoglossa ( Eoxenoglossa ) kansensis Habropoda laboriosa (Fabricius) The totals for the study are 35 genera and 112 species. missed. was at SF-BA ( Synhalonia ) rosae (Robertson) caught on 14-IV-07 Xenoglossa kansensis not collected or counted. Apis mellifera —caught in several areas, Species are listed in phylogenetic sequence by family-group, genus-group taxa, and alphabetically within the least inclusive applicable genus-group taxon. and alphabetically within the least inclusive genus-group taxa, Species are listed in phylogenetic sequence by family-group, 1. Table Area and site abbreviations in 2. Table Abbreviations for plant species in (2008). Florida according to Pascarella “throughout” County records or found Alachua AC—previous Indicated are the percentages of bees caught during a 2- or 3- month period that exceed 70% total. Stelis ater see text. Also emerged from trap nests, see text. new host association, Potential but were not captured. Xylocopa virginica —many were seen at KP-W in the spring, 1 2 3 4 5 6 7 8 9 10 11 ABLE 108 111 109 110 107 105 106 112 T Hall & Ascher: Bees in Florida Natural Areas 625 chos Fox. Most of the leaf-cutter bee cells were drena F., and Perdita Smith were found either in heavily infested by tiny parasitic eulophid wasps, the spring or fall, as has been well-documented Melittobia digitata Dahms (the biology of this ge- previously (Mitchell 1960, 1962). Colletes were nus was reviewed by Matthews et al. 2009). Rem- captured almost exclusively on Asteraceae flow- nants of a female and a male M. m. mendica and ers. Lithurgus gibbosus Smith was found at the 2 male C. dolichos were found in the destroyed sites with abundant Opuntia humifusa (Raf.) cells along 1 burrow. There was no evidence of M. Raf., reflecting an oligolectic association with that (Melanosarus) xylocopoides Smith, the recognized plant genus (Hurd 1979). Only 2 females of Xeno- host of C. dolichos. Although these results suggest glossa kansensis, an oligolege of L. and that M. m. mendica may be an alternate host of C. an important squash pollinator, were found in the dolichos, the possible prior presence of M. xyloco- natural areas, in contrast to large numbers found poides, which might have been destroyed by the during this survey period in squash-growing or- Melittobia, cannot be ruled out. Xylocopa (Xyloco- ganic farms in Alachua County (Hall 2010). In poides) virginica virginica L. occupied and wid- this survey, only 5 females of the southeastern ened the burrows of a few trap nests of a different blueberry bee Habropoda laboriosa (Fabricius) design, with U-shaped burrows with a plexiglass were captured from non-native wild radish side. Offspring began to emerge in late Jun. The Raphanus raphanistrum L. However, this bee bee species that emerged from trap nests are foot- species is probably more abundant in natural ar- noted on our list (Table 3), but the number of in- eas than indicated here, as we have captured this dividuals is not included in the total bee count. bee along roadsides and other locations from sev- The 25 cleptoparasitic bees species found in eral native plants, such as Eastern Redbud, Cer- this study included 4 Sphecodes Latreille, 5 Coe- cis canadensis L., and Carolina Jessamine, lioxys Latreille, 2 Stelis Panzer, 5 Nomada Sco- Gelsemium sempervirens (L.) (Pascarella 2007) poli, 6 Epeolus Latreille, and 3 Triepeolus Robert- that are also found in the natural areas. Osmia son. Coelioxys (Coelioxys) mitchelli Baker were chalybea was found foraging on Cirsium Mill. in captured flying over Megachile rubi nests, sug- Alachua County, but we also captured this species gesting a potential new host association for this along with both sexes of Stelis ater on Cirsium in species. Coelioxys (Xerocoelioxys) galactiae Mitch- Highlands County (4 miles SW of Old Venus on 1 ell were also flying over M. rubi nests. The latter Apr 2009 by J. S. Ascher and D. Webber; speci- new potential host association is not without pre- mens deposited in the American Museum of Nat- cedent, as another species of Xerocoelioxys, C. ural History), which further reinforces their asso- soledadensis Cockerell, is reported as a possible ciation. parasite of Megachile (Megachiloides) soledaden- sis Cockerell (Hurd 1979), which like M. rubi be- DISCUSSION longed to the former subgenus Xeromegachile (in- cluded in an expanded subgenus Megachiloides Previously published Florida bee surveys were by Michener 2007). In the Kanapaha Prairie area, from southern Florida. Pascarella et al. (2000) re- we found several Svastra (Epimelissodes) atripes corded species they had captured in the Ever- georgica (Cresson) and Triepeolus quadrifascia- glades National Park along with those that tus atlanticus Mitchell, suggesting a new host- Graenicher (1930) and others had found in the parasite association. Rightmyer (2008) reported 3 Everglades and in Dade and Monroe Counties specimens of this Triepeolus cleptoparasite from a outside the Park. In total, Pascarella listed 99 nesting site of S. atripes atrimitra (LaBerge), and species and, for each of 5 of these, 1 additional Cane (1995) observed adults inspecting and en- subspecies. Graenicher had found 61 of these spe- tering host nests. Both of these S. atripes subspe- cies. Two species, Augochlorella gratiosa (Smith) cies are recorded from Alachua County (Pas- and Coelioxys mexicana Cresson and 1 additional carella 2008). subspecies of Megachile brevis (Say) were incor- Only 3 non-native species, Megachile (Eutri- rectly attributed to Graenicher. Deyrup et al. charaea) concinna Smith, Megachile (Callomega- (2002) found 107 described and 5 undescribed chile) sculpturalis Smith, and Apis mellifera L. species at Archbold Biological Station, not count- were caught. Honey bees were seen consistently ing Augochlorella striata (Provancher) now in the cups, but only in small numbers, even at a placed in synonomy with A. aurata (Smith) (Co- location (KP-W) where managed colonies were lo- elho 2004). The morphospecies referred to as cated nearby. They were not collected or counted. Caupolicana sp. has since been described as Both previously reported and new associations C. (Caupolicana) floridana (Michener & Deyrup were found between bee species and the sites or 2004). Pascarella and Deyrup together reported a floral resources therein. Hylaeus schwarzii Cock- total of 142 species, 69 of which were shared, and erell, a species associated with wetlands (Graen- which included 15 of Florida’s 20 endemic bee icher 1930) was found in the wet prairie commu- species and 7 of the 9 endemic subspecies. In Ala- nity of Paynes Prairie. Most of the Colletes La- chua County natural areas, we captured 49 of the treille and associated cleptoparasitic Epeolus, An- species shared between these 2 studies, plus 6 626 Florida Entomologist 93(4) December 2010 species listed only in the Pascarella report and 12 northern elements and between Atlantic and Gulf only in the Deyrup report. Thus, of the 142 total Coast elements. Compared with subtropical species in the 2 reports, we found 67. Two of the 3 southern Florida, temperate Alachua County is endemic species and the 1 endemic subspecies notably enriched in Andrena and associated No- that we found were in either or both of the 2 re- mada, Osmia Panzer, and Bombus Latreille. On ports. Of these endemic taxa, absence of Stelis the other hand, Florida’s endemic bee species and ater from the Archbold list is noteworthy, espe- subspecies are concentrated toward the southern cially in light of our subsequent collections of this end of the peninsula. A few species in southern species elsewhere in Highlands County and in Florida have West Indian affinities and also are Alachua County. The paucity of Andrena and No- found in the Bahamas and Cuba (Deyrup et al. mada, a cleptoparasitic genus of Andrena, in 2002; Pascarella et al. 2000), but none of these is southern Florida and Archbold was noted in the present in north-central Florida. Thus, the entire earlier reports, compared with greater numbers Alachua County bee fauna can be considered con- in the northern part of the state, and especially in tinental. Some species best known from the states to the north. Only 3 Andrena species were southwestern United States also are present as found at Archbold, none at all in the collections disjunct populations in Florida. One of the most from southern Florida, and only 1 species of No- distinctive, Centris lanosa Cresson, was not found mada (N. fervida Smith, a cleptoparasite of Aga- in this survey but has been reported from Ala- postemon splendens (Lepeletier) rather than of chua County (Pascarella 2008), and we found it Andrena) from both areas. By contrast, we cap- recently in adjacent Putnam County. The new tured 13 Andrena and 5 Nomada species. Sixteen state records resulting from this study indicate percent of the species from southern Florida were that more remains to be learned about state-level cleptoparasitic and 27% from Archbold, compared distribution of bee species, even in a relatively with 23% we found in Alachua County. In compar- well-known Eastern state. ing our study with the others, we updated species Alachua County has a long native bee flight identities as described below. season but with a short hiatus in Dec and Jan, un- Compared with southwestern states such as like the Keys, Everglades, and other subtropical California and Arizona, and to a lesser degree At- areas of the state where native bees fly year- lantic coastal states to the north such as North round. Conspicuous flowering in natural areas is Carolina, Florida has a relatively low bee diver- concentrated in the spring (initially of woody sity (Mitchell 1960, 1962; Michener 1979; Ascher plants followed by herbaceous plants) and espe- & Pickering 2010) which decreases further in the cially the fall (largely Asteraceae). At these times, southern part of the state (Graenicher 1930). large numbers of bees, including specialists such Deyrup et al. (2002) and Pascarella et al. (2000) as various Andrena, Colletes, , and their discuss possible reasons which include the abun- cleptoparasites can be net-collected on flowers. dant rainfall flooding ground nests, to which at Bees are present in significant numbers in natu- least 1 Florida bee species has become adapted ral areas even at times and places where flower- (Norden et al. 2003), frequent fires characteristic ing is inconspicuous. In such situations, bees are of some plant communities, spoilage of pollen most efficiently collected with the colored cups stores because of the high summer heat and hu- and consist largely of rather cryptic, generalist, midity, reduced land topography thereby lacking halictids. The numerical dominance of certain vertical banks available for ground nests, absence halictine species may be a function of their euso- of many temperate plant species in the subtropi- ciality, their small size, and their use of a variety cal region, and a peninsular effect (Schwartz of sparse, scattered, floral food resources. Signifi- 1988). cant crop pollinators such as Habropoda and Xe- Although Florida does not have an especially noglossa are present in the natural areas of Ala- large bee fauna relative to other states, it is a chua County, but not in great numbers. Nonethe- unique area, evident from the endemic bee spe- less, these natural areas can be viewed as poten- cies and regional specialties present in the fauna. tial sources of such bees for colonization of new Because Florida extends as a peninsula as the acreage of relevant crops, possibly even across southern-most region of the eastern United large dispersal distances. States, almost all of the bee species are derived The bee fauna of Alachua County includes taxa from farther north on the Atlantic coastal plain or characteristic of the southeastern coastal plain from the west along the Gulf Coast. A relatively that have recently changed status, or soon will, as high diversity and abundance of Colletes and as- a result of modern revisionary studies. Southeast- sociated Epeolus, Lasioglossum (Dialictus) and ern populations of Halictus ligatus Say, including associated Sphecodes, Perdita, Megachilinae es- all present in Florida, are now considered a sepa- pecially Megachilini, and Eucerini and associated rate, cryptic species Halictus poeyi (Packer 1999). Triepeolus is characteristic of the Florida bee Many Florida specimens identified as Lasioglos- fauna as a whole. Alachua County is an interface sum (Dialictus) coreopsis (Robertson) pertain to between the peninsular-Floridian fauna and L. robertsonellum. From recent integrative stud- Hall & Ascher: Bees in Florida Natural Areas 627 ies of DNA and morphology, Lasioglossum (Dial- viding permits and broad access to Paynes Prai- ictus) tegulare (Robertson) has been divided into 5 rie and San Felasco State Parks, particularly species, 2 of which, L. puteulanum and L. lepidii Jim Weimer, Paynes Prairie Biologist. We thank (Graenicher), are in Florida (Gibbs 2009, 2010). Sam Droege, United States Geological Survey, The subspecies Ceratina dupla floridana is now Beltsville, Maryland, and Molly Rightmyer, thought to be deserving of specific rank, as DNA United States Department of Agriculture, Agri- analyses readily separate it from typical Ceratina culture Research Service, Logan, Utah, for assis- dupla Say (Cory Sheffield, personal communica- tance identifying specimens, mainly Lasioglos- tion). Lasioglossum puteulanum and C. dupla sum and Nomada, and Jim Wiley, Florida State floridana are widely distributed in Florida and Collections of , Florida Department occur north to North Carolina. Other subspecies of Agriculture and Consumer Services, for facili- present in Alachua County such as Colletes simu- tating access to the collection and providing ref- lans miamiensis Mitchell, Sphecodes heraclei ig- erence specimens. Jason Gibbs and Cory Shef- nitus Cockerell, and Megachile (Litomegachile) field, York University, Toronto, Ontario, Canada, brevis pseudobrevis Say are distinctive southeast- shared unpublished data from DNA barcoding ern elements that may prove to be phylogenetic studies including Florida bee species that in- species and are therefore deserving of further tax- formed our discussion of southeastern bee sub- onomic study. species and cryptic species. Jason Gibbs further Diagnostic features of Florida’s cryptic species, assisted with the identification of Lasioglossum putative subspecies, and visually distinctive re- specimens. John Sivinski, United States Depart- gional populations lacking formal taxonomic sta- ment of Agriculture, Agriculture Research Ser- tus, include geographic color patterns. Au- vice, Gainesville Florida, and Jorge González, glochlora pura pura (Say) is green in Alachua Texas A&M University, identified the Melittobia. County, but the southern Florida subspecies A. p. Heather Campbell, American Museum of Natu- mosieri Cockerell is blue or even purple in the ral History (AMNH), commented on the manu- Keys (Mitchell 1960; Pascarella 2008). Likewise, script. Laura Ávila, University of Florida, trans- Augochloropsis anonyma (Cockerell) is green in lated the abstract into Spanish. Two anonymous the northern part of the state and blue or purple reviewers offered thoughtful suggestions which farther south (Pascarella 2008). Ceratina dupla included the format for Table 3. Glenn Hall’s dupla and Lasioglossum tegulare north of Florida work was supported by the University of Florida are dark green, whereas in Florida C. d. floridana Agricultural Experiment Station. John Ascher’s is blue (Michell 1962) and L. puteulanum is blue- work at the AMNH was supported by Robert Go- black (Gibbs 2009, 2010). In Alachua County, spe- elet. cies such as Stelis lousiae Cockerell and Anthidi- ellum notatum (Latreille) have maculae that in- clude more red than in bees north of Florida, in REFERENCES CITED which the maculae are almost entirely yellow. 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