Native Pinelands Bees

Native Pinelands Bees

April Hamblin Senior Project Professor W. J. Cromartie December 3rd, 2012

Native Pinelands Bees 1

Table of Content

Abstract...page 2

Introductionpages 2

Study Sitespage 3-6

Methods and Materials...pages 6-9

Resultspages 9-12

Figurespages 12-15

Analysis and Discussion.pages 15-21

Further Research and Implications.pages 21-23

Conclusion.....page 23-24

Acknowledgements...page 24

References..pages 25-29

Native Pinelands Bees 2

ABSTRACT: Even though concern is spreading about worldwide pollinator decline, much is still unknown about native bee populations and their density, abundance, and diversity. The New Jersey Pine Barrens are a highly distinctive ecosystem whose native bees have not been thoroughly sampled. We fifteen transects in New Jersey Pinelands monthly from May to October, 2012. To conduct this survey, thirty bee bowls were placed at each site for about a twenty-four hour period during each sampling. Transects of ten blue, ten white, and ten yellow bee bowls were filled with water and a few drops of blue dawn dish detergent were placed outside for ten to twelve hours. Specimens were collected in 80% ethanol, then washed in water plus detergent, dried, and pinned. The majority of diversity was found during June at the Richard Stockton College site. There were 57 species found at the current stage of research. Over half belong to the family Halictidea. The survey suggests that the Pinelands areas are best suited for the family Halictidae and that future studies should be conducted with an earlier starting month and compared to these collected data.

Introduction The majority of plants80% of angiosperms, which are the largest and most diverse phyla (Bidlack & Jansky, 2011)require insect pollinators for fertilization, which means that without them, this process will not occur naturally. Apoidea, or bees, are the main group of insects that pollinate angiosperms (Brady, 2006). Although there is not research on which group of pollinators is the most vital, most entomologists will agree that bees are the most dominant pollinators in the majority of ecosystems, for they visit flower more often than other insects and both their larval and adult stages feel on floral products (Bartomeus, Cariveau, & Winfree, 2011). Not only are most angiosperms pollinated by bees, but bees pollinate 1/3 of the entire worlds food crops. Since the 1900s bees have been recorded declining (Stevens, 2011), but the past few decades have had the largest declines recorded, one example the genus Bombus is four species have declined as much as 96% (Donavan, 2011). This is urgent because many creatures, including humans, rely on bees to pollinate their food. Most national and regional surveys are still at their initial stages. It is generally hypothesized that many factors such as pesticides and land-use change affect bees negatively, but surveys are only the first step toward definitive solutions declining bee populations (Kearns,

Native Pinelands Bees 3

Inouve, & Waser, 1998). To help provide information to the entomologists involved in this research, we began a survey was conducted to determine which native bees live in the Pine Barrens, their seasonal occurrence, and there specific habitats. Study Sites The Pinelands in New Jersey are about 2,000 to 2,250 square miles and most are forested (Boyd, 1991), yet the study sites chosen were generally in n opened area. The Pinelands were chosen because they are a protected, rare ecosystem, adapted to fire, with many species of insects which are much less common in adjacent forest types. According to McCormick (1970) only 2% of these insects are pollinators. This small percentage is vital for the ecosystem, for many herbaceous plants and shrubs require them for fertilization. The Pinelands generally divides into lowlands and uplands (McCormick, 1970). There are also dwarf forests and cedar swamps (Boyd, 1991). The lowlands are generally categorized into three different habitats: pitch pine lowlands, cedar swamps, and hardwood swamps (Boyd, 1991). Lowlands main species are Pinus rigida (pitch pine), Thuja occidentalis (white cedar), Nyssa sylvatica (black gum), Acer rubrum (red maple), and Magnolia virginiana (sweet bay magnolia). One may also find Liquidambar straciflua (sweet gum), Betula populifolia (gray birch), Quercus palustris (pin oak), Quercus phellos (willow oak), and other species. There are also southern white cedar swamps that have many Thuja occidentalis, Pinus rigida, Liquidambar straciflua, and Betula populifolia. Understory of these lowlands include: Gaylussacia frondosa (dangleberry), Rhododendron viscosum (swamp azalea), Vaccinium corymbosum (high-bush blueberry), Myrica pensylvanica (bayberry), and other shrubs. Utricularia purpurea (bladderwarts), Woodwardia virginica (chain fern), Sarracenia purpurea (pitcher plants), Mitchella repens (partridgeberry), and others also

Native Pinelands Bees 4

occur here. Lowland pitch pine forests also occur with vegetation such as Gaylussacia frondosa, Gaylussacia baccata (black huckleberry), Kalmia angustifolia (sheep laurel), and other shrubs (McCormick, 1970). The lowlands also have water tables close to the surface, usually less than two feet. The cedar swamps mentioned earlier are scattered throughout the lowlands. Their most dominant tree is Chamaecyparis thyoides (Atlantic white cedar). The species that occur here are some listed in the lowlands areas but also include a number of species of mosses and lichens due to the moisture and water over the soil (Boyd, 1991). Uplands have many species, but the most dominant are Pinus rigida (pitch p)ines, Pinus echinata (short-leaf pines), and various species of Quercus (oaks) (Boyd, 1991). Other species that occur include: Vaccinium angustifolium (low-bush blueberry), Rubus fruticosus (blackberries), and many other species already mentioned but with a more complex composition. Some specific forests in the uplands include pine-blackjack oak forests, oak-pine forests, and pine-oak forests. Pine-blackjack oak forests have many species but are special because of their Quercus marilandica (blackjack oak), and various other members of the Quercus genus along with some Pinus. Oak-pine forests have Quercus velutina (black oak), Quercus alba (white oak), and Quercus prinus (chestnut oak). Pine-oak forests have the same Quercus along with Quercus marilandica, Pinus rigida, and other species. Pinelands have more swamps and forests than actual grass, but some grasses can be around (McCormick, 1970). The uplands water table is generally two or more feet below the ground level as well. The dwarf forests generally support the growth of an unusually short forest of mature trees. Generally the dominant trees are Pinus rigida (pitch pines) as well as other species of oaks. The stunted growth may be due to the fact that reproduction is mainly vegetative than seed dispersal (Boyd, 1991).

Native Pinelands Bees 5

There were a total of fifteen study sites chosen for this project within the Pinelands areas. Within the Richard Stockton College of New Jersey site (where the rarest species were found that will be mentioned later on) there were the following siteswith the following habitat: W of Observatory (39.4850 N 74.5565 W)scattered trees, thick grass Hospital Field (39.4780 N 74.5440 W)scattered oaks, dry grass, lichens Baptisia Site Zinckgraf farm (39.4850 N 74.5435 W)sparse pines, dry grass, lichens Sand Road off Delaware Ave. (39.4890 N 74.5410 W)pine oak woodland Parkway Ponds Borrow Pit (39.4820 N 74.5260 W)shallow borrow pit, meadow Powerline R-O-W (39.4930 N 74.5190 W)mowed right of way

Within the S. Vienna Ave site there were the following sites: Orchard (39.5235 N 74.5990 W) orchard and garden Back Field (39.5247 N 74.6000 W)old field

Within the Franklin Parker Preserve site there were the following sites: ~5 km S. Chatsworth Ten Trunks Oaks (39.7736 N 74.5344 W)un-mowed dikes edge of former cranberry bog ~5.5 km S. Chatsworth vic. Cedar Swamp Ten Trunks (39.7776 N 74.5321 W)sand path across restored cranberry bog ~4.25 km S. Chatsworth Ten Trunks Red Pit (39.7818 N 74.5342 W)dry clearing ~4.25 km S. Chatsworth Ten Trunks Bee Yard (39.7807 N 74.5338 W)wet clearing, dry sand road in pine woodland ~3.2km SSW Chatsworth Middle Rd. (39.7869 N 74.5532 W)wet clearing, dry sand road in pine woodland

Native Pinelands Bees 6

~1.5 km SSW Chatsworth Sand SE End Airstrip (39.8060 N 74.5443 W)damp clearing and dry sand pit

Ringler Ave. Chatsworth (39.8141 N 74.5699 W)cripple wetland NW of road, sphagnum The Pinelands are known to be mosaics of the above habitats, generally in relatively

small areas (Robichaud, 1980). Since the Pine Barrens were chosen, the study hopes to reveal some sort of data that gives a basis of what species occur in the Pinelands and how these species relate to season. This data is being collected in hopes that another student in the future hopes to study bees in the Pinelands and compare them to what was found throughout this research. Methods and Materials This survey used Sam Droeges method of sampling with bee bowls to collect the specimens. Bee bowls are small (generally 2-4 oz) plastic cups brightly colored blue, white, and yellow with inflorescence to attract bees. Each site has thirty cups (ten of each color) placed in a transect about five meters apart from one another. These bowls are filled water and blue dawn dish detergent (not citrus or the bees will be repelled) so that the bees cannot break the surface of the water and are collected by the cups. It has been noted that the bees stop moving, apparently dead, within sixty seconds so that they do not suffer. These cups are left out for periods of twenty-four hours, but should not be left our longer for their bodies may start to decompose. For this survey, we placed the bowls out for about ten to twelve hours due to time limitations. To collect the bees, some sort of net such as a brine shrimp net may be used so that the bowls may easily be collected by pour its contents into the net while walking the transect. Once the specimens are brought back to the lab and the bees are sorted out from the collection, they are stored in at least 70% ethanol until they are ready to be washed (Droege, 2012).

Native Pinelands Bees 7

Before identifying bees, since they are generally covered in pollen and matted from floating in the bee bowls for hours, they are generally washed. To do this, one needs two glass jars any size (usually salsa or jam jars are fine for this), an empty jar, one jar of 95% ethanol, one empty jar labeled 95% ethanol, a paper towel, a bee dryer which is a glass jar with some sort of mesh on the top to prevent them from falling (generally the mesh is metal because of the hot temperatures from the blow dryer), a blow dryer, a funnel, mesh for the funnel, and forceps. First, the bees must be dumped from their containers through the funnel and into the empty jar that is not going to be used for anything else. This will take the bees out of the ethanol. One of the glass jars should be half to two thirds filled with warm water and a couple drops of soap, just enough so that there are not many suds. Using the forceps, one should pick the bees out from the funnel and drop them in the warm water. The lid should then be put on the jar and the jar should be swirled around in a circular pattern where the water almost looks like a tornado. After swirling for about thirty seconds to two minutes, depending on the number and size of the sample of bees, the bees should then be poured through the funnel and into the other empty glass jar. Generally for larger bees and more bees they have to be swirled for longer periods of time. Bees bodies are robust, so it is a good idea to swirl harder than softer. After this, the same process happens but in the 95% ethanol. Once the bees are done in the ethanol, they should be picked out with the forceps and put onto the middle of the paper towel. Using ones hands, pick up the paper towel by two of the opposite corners. Then, take the other two corners in the same hand, making sure there are no openings that the bees could fall through. Basically one hand should be in a fist with the corners of the paper towel and the other should be blocking any tine openings that may persist. Shake the bees up and down for about a minute in this position, watching to see if any fall out and need to be rewashed. Once this is

Native Pinelands Bees 8

completed, put the paper towel back onto the table and put the bees into the glass jar that has a mesh lid with the forceps. After putting the mesh lid on, make sure that none of the bees stick to the sides of the glass jar. Using the blow dryer, blow the bees for a minute to two minutes depending on the number and size of the bees. Generally larger bees or more bees require longer periods of blow drying (Droege, 2011). One must remember, after every step, to check that none of the bees are left behind in their previous containers so that the results of the experiment are valid. After washing and drying the bees, they are then ready to be pinned, labeled, and eventually identified. Any members of Bombus, Apis mellifera, or bees of similar sizes were pinned through the thorax with size three or four pins. Smaller bees were glued to the pins on their left side in the area between their middle and last legs on the thorax. Pin sizes varied, but generally any size under one was avoided because of their lack of sturdiness. Once pinned and counted, labels were made, printed, cut, and placed on the specimens. Specimens without labels are considered useless because they contain no geographical information, therefore their site locations and dates are unknown. Because of this, specimens were handled with great care and concern. After labeling the bees, they were then sorted in groups, first by family, and then by genus. Main websites used for this was Discover Life (2012) at http://www.discoverlife.org/mp/20q?search=Apoidea#Identification and Bug Guide (2012) at http://bugguide.net/node/view/8267. Along with these websites, Bees of the Eastern United States by Mitchel (1960) was read to give an overview of the differences of the specific families of bees as well as BugGuide.net (2012) and other resources such Borror and Delongs Intro to the Study of Insects (Triplehorn & Johnson, 2005). After the bees were sorted to genus, they

Native Pinelands Bees 9

were then taken to The American Museum of Natural History in New York, New York for Dr. John S. Ascher to identify to species. These specimens were taken November 16th, 2012 to be identified. Over half of the collection was identified that day, yet the other specimens had to stay at the museum for further identification. Because of this, this report only includes the readily identified specimens. While the other specimens will be taken into consideration later, this report does not talk about them. Not only were many species identified, but this visit also allowed the project to consider being recorded within the museums database so that the information may be updated when materials such as species names change. Although it is unsure what method of databasing will be used eventually, this is something taken into consideration because it would be highly beneficial for the study as well as the museum. Results Of the specimens that were identified, there was a total of 57 different species, which are in table 1. This raw data shows that the majority of the bees were Halictidae, while the majority of the Halictidae were Lasioglossum. Of the species identified so far, the rarest were Augochlorella gratiosa, Lasioglossum arantium, and Lasioglossum sopinci of Halictidae and Osmia felti of Megachilidae. These species did not have many accessible works and seem understudied, but there were a few mentions of them within scholarly journals. Augochlorella gratiosa are known to live in ranges from Michigan to Nova Scotia, south to Texas (Discover Life, 2012), and across to Mississippi on plants such as Chrysopsis microcephala (narrowleaf silkgrass) (Michener, 1947), within Pennsylvania (Donovall & vanEngelsdorp, 2010), and Florida in areas such as Evergreens National Park (Ascher & Hall; Neal, Pascarella, & Waddington, 1999). They are believed to be around from March to September, but year round in Florida. They have floral records of interacting with Berteroa,

Native Pinelands Bees 10

Bidens, Cirsium, Citrus, Clethra, Erigeron, Eryngium, Hypericum, Ilex, Lepidium, Melilotus,Oenothera, Polygala, Polygonum, Rhus, Rubus, and Taraxacum (Discover Life, 2012). A. gratiosa are also known to pollinate plants such as watermelon (Goff, 1937) and Lysimachia (Brownie) fairly well, but are vaguely described and can be one of the harder bees to identify (Coelho, 2004). Within the site at Richard Stockton College of New Jersey, there are known to be Bidens arisfosa (tickseed-sunflower) and other species of Bidens as well as Cirsium arvense (Canada thistle), Cirsium discolor (field thistle), Clethra alnifolia (sweet pepperbush), Erigeron canadensis (horseweed), Erigeron annuus (daisy fleabane), Hypericum boreale (St. Johnswort), Hypericum canadense (lesser Canadian St. Johnswort), Hypericum gentianoides (orange grass), Hypericum perforatum (common St. Johnswort), Ilex galbra (inkberry), Ilex opaca (American holly), other species of Ilex, Melilotus alba (white sweet clover), Melilotus officinalis (sweet clover), Oenothera biennis (common evening primrose), Oenothera laciniata (sinuate-leaved evening primrose), other species of Oenothera, Polygala brevifolia (short-leaved milkweed), Polygala lutea (yellow milkwort), Polygonum aviculare (knotweed), Polygonum caespitosum (oriental ladys thumb), Polygonum lapathifolium (smartweed), Polygonum pensylvanicum (Pennsylvania smartweed), Rhus copallinum (winged sumac), Rhus copallina (dwarf sumac), Rhus glabra (smooth sumac), Rubus hispidus (bristly dewberry), Rubus occidentalis (black raspberry), other species of Rubus, Taraxacum laevigatum (red-seeded dandelion), and Taraxacum officinale (common dandelion) (Cromartie, 2012). Since these genuses were listed as hosts for Augochlorella gratiosa, so it appears as though many of them could be where the species forages at Richard Stockton College. This may be why Augochlorella gratiosa was found at the Richard Stockton College of New Jersey and it was also found at S. Vienna Ave.

Native Pinelands Bees 11

Unfortunately, Lasioglossum arantium did not have as many articles covering the species as A. gratiosa, yet the articles were more based on scientific research. Gibbs (2011) explains that L. arantium is a social parasite. These social parasites are believed to have dual lineages, originating from their combined DNA barcode and specific traits of their morphology (Albert, Gibbs, & Packer, 2011). There are not any listed host plants of this species or much information about it, so it would be interesting to think about why it was found in the Pinelands. Perhaps its hosts were, since it is considered a parasite. The other Lasioglossum, L. sopinci is a large species known to occur from March to July in New Jersey, North Carolina (Discover Life, 2012), and Georgia in areas such as Piedmount (Hanula & Horn, 2011). Davis et al (2009) speaks about micro-deserts and states that L. sopinci relies heavily on the soil structures. L. sopinci tends to be in areas along with L. viercki because this species also relies on sandy soils. This species also has no list of plant host species know yet, but perhaps the species relies more on nesting areas available due to the sandy soil than the specific available flowers. This species was found in all three of the locations for sites, so this common soil could be the limiting factor. Specifically, this species was found at the following sites: Richard Stockton College (W of Obervatory, Hospital field, Baptisia site Zinckgraf farm, Sand Road off Delaware Ave), S. Vienna Ave (Orchard), and Franklin Parker Preserve (Ten Trunks Bee Yard and Middle Road). Only the Stockton site had multiple day collections of this specimen. Out of a total of thirteen specimens, ten were collected from the Richard Stockton College of New Jersey, one was from S. Vienna Ave, and two were from the Franklin Parker Preserve. So far, the rarest species seem most common at the Richard Stockton College of New Jersey sites.

Native Pinelands Bees 12

The final rare species that will be discussed is Osmia felti, which is distributed from Minnesota to New England states such as Pennsylvania (Donovall & vanEngelsdorp, 2010) and most south to West Virginia from June to August (Discover Life, 2012). O. felt is known to heavily pollinate blueberries as well (Cane et al, 1985). Even though other sources do not have a plant host species list, Cane reveals that Osmia felti is known to pollinate Vaccinium (blueberries), which are very common at the Richard Stockton College of New Jersey, where the only specimen was found (at the specific site called Baptisia site Zinckgraf farm, which appears to be the most diverse out of all the Stockton College sites). It is highly likely that this species is in the Pinelands because of the blueberries, yet there may be other undiscovered factors that influence Osmia feltis presence at Stockton College. Table 1
Family Andrenidae Andrenidae Andrenidae Andrenidae Andrenidae Apidae Apidae Apidae Apidae Apidae Apidae Apidae Apidae Apidae Apidae Halictidae Halictidae Halictidae Halictidae Genus Species Andrena (Melandrena) carlini Andrena (Melandrena) vicina Andrena (Simandrena) nasonii Andrena (Trachandrena) rugosa Calliopsis andreniformis Apis mellifera Bombus (Cullumanobombus) griseocollis Bombus (Pyrobombus) bimaculatus Bombus (Pyrobombus) impatiens Bombus (Pyrobombus) perplexus Ceratina (Zadontomerus) calcarata Ceratina (Zadontomerus) strenua Peponapis pruinosa Ptilothrix bombiformis Xylocopa (Xylocopoides) virginica Agapostemon splendens Agapostemon texanus Agapostemon virescens Augochlora pura Total 1 1 4 1 11 Sites May SV X SV X RSC&SV X RSC X RSC&SV&PP RSC&SV 42 &PP X 14 5 6 10 4 2 1 3 2 22 22 11 4 RSC&SV&PP RSC&SV&PP RSC&SV&PP RSC&SV RSC&SV RSC SV RSC&SV RSC&SV SV&PP X RSC&SV X RSC&SV RSC&PP June July Aug. Sep. Oct.

X X X X X X X

X X X X

X X X X X X X X X

X X X

X X X X X X X

X X X X X X X

X X

Native Pinelands Bees 13 RSC&SV 23 &PP 8 SV&PP 3 1 1 32 1 4 39 6 5 10 1

Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Halictidae Megachilidae Megachilidae Megachilidae Megachilidae Megachilidae Megachilidae Megachilidae

Augochlorella aurata Augochlorella gratiosa Augochloropsis (Paraugochloropsis) metallica Halictus (Nealictus) parallelus Halictus (Nealictus) xarallelus Halictus (Pachyceble) confusus Halictus (Protohalictus) rubicundus Halictus (Seladonia) confusus Halictus ligatus or poeyi Lasioglossum (Dialictus) bruneri Lasioglossum (Dialictus) cressonii Lasioglossum (Dialictus) illinoense Lasioglossum (Dialictus) imitatum Lasioglossum (Dialictus) leucocomum Lasioglossum (Dialictus) oblingum & subuiridatum & relatives Lasioglossum (Dialictus) pectorale Lasioglossum (Evylaeus) nelumbonis Lasioglossum (Evylaeus) sopinci Lasioglossum (L.) leucozonium Lasioglossum arantium Lasioglossum fuscipenne Lasioglossum tegulare Lasioglossum vierecki Lassioglossm (Dialictus) coeruleum Sphecodes brachycephalus Sphecodes coronus Sphecodes fattigi Sphecodes pimpinellae Heriades (Neotypetes) carinata Hoplitis producta Megachile (Litomegachile) b. brevis Megachile (Litomegachile) texana Megachile (Xanthosarus) addenda Osmia (Melanosmia) felti Osmia (Melanosmia) pumila

X X

X X X

X X X

X X

RSC X SV X SV X RSC&SV X RSC RSC&SV RSC&SV X RSC&SV RSC&SV&PP RSC RSC

X X X X X X X

X X X X

X X X

X X

X X X

1 RSC 21 RSC&PP 5 RSC&SV X X X X X X X

X X X X X

29 PP RSC&SV 13 &PP X 10 RSC&SV 1 10 RSC&PP 1 PP RSC&SV 165 &PP X 2 RSC 3 RSC&SV X 2 RSC&SV X 1 PP 1 SV 1 RSC X 1 SV X 3 RSC&SV 7 RSC&SV&PP RSC&SV 6 &PP X 1 RSC 13 SV X

X X X X

X X

X X

X X

X X X X X

X X

Native Pinelands Bees 14 Megachilidae Megachilidae Megachilidae Osmia (Osmia) taurus Osmia virga Stelis (Stelis) laterallis 1 RSC 2 RSC&SV 1 SV X X X

Graph 1

Bee Species Richness Compared to Month

Number of Species 40 30 20 10 0

Month

Native Pinelands Bees 15

Graph 2

Species Richness Compared to Sites

Number of Species 50 40 30 20 10 0 S. Vienna Ave Richard Stockton Franklin Parker College of New Preserve Jersey Site

Native Pinelands Bees 16

Graph 3

Bee Species Richness Compared to Family

Number of Species Found 40 30 20 10 0

Family

For more details about these figures refer to the database. Analysis and Discussion Patterns arise from analyzing figure one as well as the database. It is interesting to look at the patters of bee families related to the different months that were sampled. Andrenidae, the largest family of bees in North America, consists of mainly many mining bees and are most common in spring. One common genus, Andrena, is known to have banded abdomens similar to those of sweat bees in Halictus (Nature Search, 2008). This family is also known to nest in the ground, sometimes relatively close to one another, in similar borrows to that of halictids (Triplehorn & Johnson, 2005). In Andrenidae, all Andrena were collected in May, two species in Vienna Ave, one species at Stockton, and one species at Vienna Ave and Stockton. Andrena are known to mainly be in the northern hemisphere and most occur in the spring but some are also

Native Pinelands Bees 17

know to occur in the summer, autumn, and even in the winter (Bug Guide, 2012). Calliopsis was collected June through July at Vienna Ave, Stockton, and Parker Preserve. For Andrenidae, May had the most species richness and diversity, while the majority of the species were found at Vienna Ave. Calliopsis are known to occur from Canada to Chile and Aregentina and tend to be specialists where they collect pollen from one or few species generally from one of the following families of plants: Leguminosae, Euphorbiaceae, Compositae, and Verbenaceae (Bug Guide, 2012). Apidae, or long-tongued bees (along with Megachilidae), have some species with particular behaviors such as males sleeping in aggregations on vegetation or gathering flower oils instead of the pollen. This family includes digger bees that nest in the ground, bumble bees that are the only native social bee to North America, carpenter bees where some have overlapping nests in dead wood so they are considered semisocial, cuckoo bees which act as cleptoparasites on other bees, and the invasive honey bee (Nature Search, 2008). Digger bees generally nest in borrows in the ground and may act solitary or nest colonially with a thin waxlike substance lining their cells (Triplehorn & Johnson, 2005). In Apidae, the invasive, social honey bee Apis mellifera was collected throughout the entire sampling time from May to October as well as all three sites. This shows that invasive species tend to spread rapidly from lack their native predators and outcompeting the native pollinators (Thomson, 2004). Bombus was not collected as often, yet seemed to be most significant in June. A couple of Bombus were also collected in July and August. Three of the species of Bombus were found at all three sites, while one species of Bombus was only found at Vienna Ave and Stockton. Bombus are found around the world but lacking in Africa and Australia and are mimicked by a few fly species (Bug Guide, 2012). Ceratina were collected in

Native Pinelands Bees 18

June, August, and October, but had more species richness in October. Both species of Ceratina were found at Stockton and one was also found at Vienna Ave. Ceratine, or small carpenter bees, occur worldwide and nest in the wood (Bug Guide, 2012). Peponapis was collected in July along with Ptilothrix, which was also collected in August. These were both found at Vienna Ave, while Ptilothrix was also found Stockton. Peponapis are found throughout the US and Canada, south to Argentina and associated with the plant family Cucurbitaceae which generally occur in dry, sandy, deserts. Ptilothrix occur from the US to Argentina. (Bug Guide, 2012). Xylocopa was collected in June and July as well and at Vienna Ave and Stockton. Xylocopa, or carpenter bees, look similar to bumble bees but have naked abdomens and occur worldwide, generally on flowers or nesting sites near woody plants (Bug Guide, 2012). These nesting sites may be in wood or the stems of other plants and maybe even excavate galleries in solid wood (Triplehorn & Johnson, 2005). Throughout Apidae, June appeared to have the most species richness and diversity, followed closely by July. Vienna Ave also had the most species richness, followed close by Stockton. Halictidae, or sweat bees, generally are the most colorful bees from black to blue to green with behaviors that range from solitary to semisocial and consist of over 500 species in North America (Nature Search, 2008). In Halictidae, there is more of a wide range of bees collected in the Pinelands. Agapostemon has species current throughout the entire season from May to October, with all species present in July, September, and October. These species were found at two Vienna Ave and Stockton or Vienna Ave and Parker Preserve. Agapostemon occur from Canada to Argentina, where A. splendens and a. texanus are mentioned to only occur in Florida (Bug Guide, 2012), yet they were collected within this Pinelands survey. Halictus is similar, for this group has species in every month sampled so far, yet the most species were collected in

Native Pinelands Bees 19

May. These species were only found at Vienna Ave, Stockton, or a combination of both. Halictus occur in many areas yet are more numerous in sandy areas and are believed to occur from early spring to late fall, but year-round in Florida and Texas (Bug Guide, 2012). Augochlora has one species from July to August found at Stockton and Parker Preserve which occur worldwide, Augochlorella has two species from May to July and September where one continues onto October found at all three sites or Vienna Ave and Parker Preserve which occur from Canada to Argentina in May to October, and Augochloropsis has one species present from May to July found at Stockton which occur from Canada to Argentina (Bug Guide, 2012). Lasioglossum has fifteen different species that occur variously through May to October, yet the highest species richness and diversity was in June. L. Dialictus were all found at Stockton or Stockton and a combination of the other two sites. L (Evylarus) were found at Parker Preserve or all three sites. Other Lasioglossum were found at combinations of all three sites. The lowest species diversity occurred in May. This shows that, although others were most common in May, Lasioglossum is not in this class and may need the warmer temperatures to hatch, later to be able to stay longer with the cold weather in October. Lasioglossum are generalists that occur worldwide, yet some are cleptoparasites, some are nocturnal, and some are oligolectic with behaviors from solitary to semisocial to eusocial (Bug Guide, 2012). Two Sphecodes species were collected in May and found at Vienna Ave and Stockton, another in August found at Vienna Ave, and another in October found at Parker Preserve. This could be because these bees have very similar niches, so they occupy different sites in the Pinelands at different times as that correspond to their hosts, for they are cleptoparasites, usually of other Halictinae, and occur worldwide (Bug Guide, 2012).

Native Pinelands Bees 20

Megachilidae consist of leaf-cutter, which cut circles from leaves to construct their nests, and mason bees, which nest in preexisting tunnels in wood, and others such as Coelioxys, which are cleptoparasites of Megachile. Females collect pollen with a brush on the underside of their abdomens and are considered long-tongued with over 600 species on North America (Nature Search, 2008). In Megachilidae, there are ten species that were found within the Pinelands. One species of Heriades found at Stockton and one species of Hoplitis found at Vienna Ave were only found in May. Heriades occur worldwide except for South America and Australia while Hoplitis is noted to occur in South Africa in Holarctic and Ethiopia, yet it was also found in the New Jersey Pinelands (Bug Guide, 2012). The two species of Megachile (L.) were very similar and both collected in July found at Vienna Ave and Stockton or all three sites, yet one continued and the other was not collected in August but in September and October. When occurrences like this happen, it leads to think that these bees fill multiple niches of the same sort, especially to similar plants within the same genera of bee. Another Megachile (X.) was only found in May and June but was collected at all three sites. This reassures the idea of fulfilling similar niches, therefore causing competition, which would result in patchiness of similar bees, where one species is around during a period of time and the other is around during another period of time. Megachile is known to occur in many areas such as the US in May through October but may start flying in earlier times in warmer areas and feed on a variety of pollen (Bug Guide, 2012). One species of Osmia was only collected in May at Stockton, two species were only collected in June at Vienna Ave and Stockton or just Stockton, and another species was collected in May and June at Vienna Ave. When species are only found in May and/or June, it leads researchers to think that this species either flourished for a longer period of time before the study site was sampled, or to think that the species has a very short life span. Osmia are known to occur from early to late

Native Pinelands Bees 21

spring with few around in the summer and to visit species in Vaccinium such as blueberries with a variety of nesting areas such as in the soil or hollowed stems (Bug Guide, 2012). Stelis was only found in May as well at Stockton, which may be because of one of the above reasons. Stelis occurs throughout most of the world except Australasia and equatorial Africa and are known to be cleptoparasites of other leaf cutter bees, mainly Megachile (Bug Guide, 2012). For Megachilidae, Stockton and Vienna Ave also were the most productive sites. About 23.66% of the bees different species were collected in June, followed closes by July with 20.61% of the species. May collected 19.1% of the bee species. September collected the least amount of species richness, the least amount of different species, with a significantly lower 10.69%. Right before and after September, both August and October, caught the same at 12.98% bee species richness. Figures 2 and 3 show this quite well in graphs. Along with these findings, 39% of the species were found at Vienna Ave, 42% of the species were found at Stockton College, and 19% of species were found at Parker Preserve. One would think that Parker Preserve would have the most species richness because of the general lack of people, but as of the research so far, Stockton College has the highest species richness. Vienna Ave is following Stockton close behind, so it will be interesting to see how this changes once all of the bees collected are put into the overall data. For the different families of bees found there were the following statistics: Andrenidae were 8.77% of the species, Apidae were 17.54% of the species, Halictidae were 56.14% of the species and Megachilidae were 17.54% of the species. These results suggest that most bees were found at Stockton in June in which most of them were in the family Halictidae. This being said, the data collected does have some error within it. First, all of the specimens were not identified yet, so the species spoken about in this essay are about only half of

Native Pinelands Bees 22

the total species, maybe a bit more than half. Because of this, although the data is showing that the greatest species richness is found in the month of June as well as at Stockton College, this may change once the other specimens are collected, databased, and analyzed. While all of these sites were within the Pinelands, it would be useful to take a closer look into these areas and distinguish their differences because many studies cause some error due to ignoring with-in site variability (Barker, 1994). There must also be some error calculated into the results due to things such as small sample size, weather on collection day, amount of time the bee bowls were left out during collection, how well the bee bowls were visible at each site to the bees, possible calculation errors, possible misspelling errors, and other possible human errors. This study will be more valid when it is more complete once all of the specimens are identified, databased, and studied. Further Research and Implications This research was designed with the idea that another Stockton professor and student would work together to research these study sites again. This data was surveyed for baseline data so that future studies may compare their results to this as well as a survey of basic comparison of months to species richness. In very general terms, species diversity was found to be greatest at Stockton, in July, and within the family Halictidae. This information will be shared and hopefully used in the future as baseline data for comparison. Since these data were recorded in the Pinelands, it is important to look at factors there that may be influencing the bee community. Frohnapple (2010) conducted research involving open-forest areas and how much habitat alteration bees could survive within this setting. The results showed that bees abundance was positively related to fire and negatively related to canopy cover. Since all of the study sites within the Pinelands research were in proximity to the

Native Pinelands Bees 23

forest, this shows that areas with less trees benefited bees over areas with more trees. Bees forage and travel in different radius of habitat due to their species and size, so it is safe to say most probably did not forage within the relatively close forest, but found other sources of food, nesting areas, etc. The infertility of the Pinelands soil (Robichaud, 1980) appeared to also benefit the bees by increasing nesting availability (Frohnapple, 2010). Another study, conducted by Ascher et al (2007), suggests that bees in the Pinelands and other forests benefit from some habitat change; the bee community was positively affected with 60% tree removal in the forests for this increased dead wood for nesting and allowed more sunlight to reach their soil nests. Griswold, Kremen, & Winfree, (2007) also studied New Jersey Pinelands in 1600 m radius transects to see how altered habitat could be compared to unaltered forests. Their results were consist with Frohnapples and Ascher et als in that the altered areas such as agricultural fields, suburban developments, and urban areas had greater species richness and diversity than extensive forests. GIS also applied to this research. While these results show that forests are not ideal habitat for bees, there was a note that not all land-use change was beneficial to bees, for most had a general negative effect on bees (Ascher et al, 2007). As Bartomeus, Cariveau, & Winfree (2011) discuss from reviewing over 200 articles on pollinators (not just bees), they have found that the data supports that generally land-use change has a ratio of 3:1 negative to positive effects. While land-use change may help somewhat in forested areas or land close to forests like the study sties chosen in the Pinelands, it is more harmful than beneficial in other areas, so one must consider this while studying the bee community, for human impact is only going to increase as the population increases. Their studies also show that the response of the pollinators is largely dependent on the change of floral resources more than the specific land-use change and that the major negative effects were from

Native Pinelands Bees 24

extremely modified habitat. When the land-use is moderate, there tends to be positive or negative responses depending on what was actually changed about the habitat. Other studies that looked at habitat types instead of simply looking at a gradient of change showed that land-use change had many positive effects. Mainly it was found that pollinator abundance and richness decreased where there was an increase in human land-use change in surrounding landscape, but the pollinator species and abundance increased when natural habitats were altered to anthropogenic areas. This is believed to be due to the methods of studying the pollinators and a smaller spatial scale of change in some habitats. Some believe this is due to the fact that land-use change generally creates early successional habitat that creates more open areas for bees. There is not enough research to understand all of the aspects yet, but it appears that land-use change is generally more negative toward bee communities, where specialists and social populations are more negatively affected than the others. There is not much research like this in agricultural areas either (Bartomeus, Cariveau, & Winfree, 2011). While this information is important, there still needs to me studies about how land-use change affects other factors involving a pollinators life such as nesting grounds. Budny et al (2005) conducted another survey within the Pinelands of New Jersey that tested the proximity of habitats and how it related to bees movement. This study found that the movement was generally based on foraging and that indices were an extremely difficult calculation within GIS software. Dafni et al (2003) researched linking bees to floral resources and came to the conclusion that much of the bee community was related to floral diversity, nectar diversity, available nesting sites, geography, and post-fire age the area if possible. This information would be interesting to analyze once all of the data identification were completed with this research and the habitats were examined more thoroughly.

Native Pinelands Bees 25

Since the results are not finished, this article will be revised once the results come and other technologies such as GIS will be applied to the data. When another research decides to study these sites, collection dates should begin earlier so that early spring species are not left out as well. Once these data are completely identified and databased, the research will hopefully be published. Conclusions These data collected from this survey will be shared with organizations such as the Pinelands Preserve Alliance and different scientists such as Sam Droege and Dr. John Ascher. These data will fit in the larger scheme of things to help other current researchers connect certain aspects found in these data to their own data. Perhaps this research will help them confirm where a specific species may occur or give new light to a species that was not found in this region before. These data will be used to in analysis to understand the entire bee community better to research questions such as why a specific species occurs where or why the bee community is structured the way it is and many other questions, which would require professional analysis, that need answering to help native bees. While the importance of this research may be overlooked by some who do not understand the pollinator-plant relationship or know anything about bees, this research is far from trivial or irrelevant. Pollinators, which largely consist of bees, provide food as security for the human race as well as other animals and are essential bio-indicators of an ecosystems functioning and diversity (Abrol, 2012). The significance of bees cannot be stressed enough. Native bees are known to pollinate native flora and many are considered to be specialists, which means that they pollinate particular species of flowers (Aigner, 2001). Because of this, many bees are at risk, especially the specialists, for they are vulnerable to change. Native bees are currently declining,

Native Pinelands Bees 26

yet there is not enough research conducted to understand exactly why or how to find a solution to help stop this decline. Some reasons are believed to be excessive use of pesticides, habitat destruction due to certain types of land-use change (Batra, 1992), and competition from invasive species like Apis mellifera (Thomson, 2004). More research is encouraged to find a gradient of change that bee communities can handle instead of simply focusing on one species. Not only this, but a focus should be on pollinator species composition and abundance instead of simply focusing on species richness (Bartomeus, Cariveau, & Winfree, 2011), so, once the results are completed, this research should attempt to analyze the bee community composition instead of simply noting what species were found. I plan to continue research such as this in graduate school to broader the known information about native bees and help preserve ecosystems diversity and food by conserving native bee populations. Acknowledgements I want to thank Sam Droege for initially identifying bees. I want to thank Dr. John Ascher for taking the time out of his day to identify many of the bees collected from the survey as well as giving us information about more reliable methods of databasing. I also want to thank Dr. William J. Cromartie for collecting the specimens, making the labels for the specimens, driving me to New York to Dr. Ascher to ID the specimens, and having much care about his students to help me make this project highly interesting and useful. Special thanks also to Franklin Parker Preserve for letting us sample there and Stockton College of letting us sample there as well. Another thanks to organizations working toward the conservation of native bees such as The Xerces Society for Invertebrate Conservations efforts and the Pinelands Preservation Alliance for letting us share our information with you, making the research hold even more importance.

Native Pinelands Bees 27

With research such as this, we may be able to begin finding answers to why native bees are declining and start proposing solutions to help them.

References Abrol, D. R. (2012). Pollination Biology: Biodiversity Conservation and Agricultural Production. Springer. London and NY. Pp 785. Aigner, P. (2001). Optimality modeling and fitness trade-offs: when should plants become specialists? Oikos. 95. 177-184. Albert, J., Gibbs, J., & Packer, L. (2011, Nov.). Dual origins of social parasitism in North American Dialictus (Hymenoptera: Halictidae) confirmed using phylogenetic approach. Cladistics, 28(2), 195-207. Ascher, J. et al. (2007). Impacts of logging on midsummer diversity of native bees (Apoidea) in a northern hardwood forest. Journal of the Kansas Entomological Society, 80(4), 327-338.

Ascher, J. & Hall, G. Surveys of bees (Hymenoptera: Apoidea: Anthophila) in natural areas of Alachua County in north-central Florida. The Florida Entomologist, 93(4), 609-629.

Azuma, H. et al. (2000, November). New perspective on the pollination biology of basal angiosperms. International Journal of Plant Sciences, 161(S6), S225-S235.

Barker, R. J. et al. (1994). With-in site variability in surveys of wildlife populations. Ecology, 75(4). 1097-1108.

Native Pinelands Bees 28

Bartomeus, I., Cariveau, D., & Winfree, R. (2011). Native pollinators in anthropogenic habitats. Annual Review of Ecology, Evolution, and Systematics, 42, 1-22. Bidlack, J. E. & Jansky, S. H. (2011). Sterns Introductory Plant Biology: Edition Twelve. New York: World Color Press Inc. Pp 432-451. Boyd, H. (1991). A field guide to the Pine Barrens of New Jersey. Its Flora, Fauna, Ecology and Historic Sites. Plexus Publishing. USA. Pp 423.

Brady, S. et al. (2006). The history of early bee diversification based on five genus plus morphology. Proceedings of the National Academy of Sciences of the Untied States of America. 103. 15118-15123.\

Batra, S. (1992). Bees and pollination. Pp 15-17 in Adams, J., Arnett, R., and Thomas, M., eds. Insect potppurri: adventures in entomology. The Sandhill Crane Press. Gainseville. 15-17.

Brownie, C. et al. Limitations to fruit and seed production be Lysimachia Asperulifolia Poir. (primulaceae), a rare plant species of the Carolinas. The Journal of the Torrey Botanical Society, 133(3) 403-411. Budny, R. V. et al. (2005, March). Testing simple indices of habitat proximity. The American Naturalist, 165(6). Bug Guide. (2012). Anthophila (Apoidea) Bees. Retrieved September 16th, 2012, from http://bugguide.net/node/view/8267. Cane, J. et al. (1985). Pollination ecology of Vaccinium stamineum (Ericaceae: Vaccinioideae). American Journal of Botany, 72(1), 135-142.

Native Pinelands Bees 29

Coelho, M. (2004, July). A review of the bee genus Augochlorella (Hymenoptera: Halictidea: Augochlorini). Systematic Entomology, 29(3), 282-323. Cromartie, W. J. (2012) Campus Plant List. Excel Spread Sheet. Retrieved December 9th, 2012, from Jamie.Cromartie@stockton.edu. Dafni, A. et al. (2003, October). Linking bees and flowers: How do floral communities structure pollinator communities? Ecology, 84(10), 2628-2642. Davis, C. et al. (2009). The lost micro-deserts of the Patuxent River: Using landscape history, insect and plant specimens, and field work to detect and define a unique community. Entomological Society Washington, 111(1), 132-144. Discover Life. (2012). Apoidea Bees; Apoid Wasps. Retrieved September 16th, 2012, from http://www.discoverlife.org/. Donavan, T. (2011, January 3). Bumble Bees in U.S. Suffer Sharp Decline, Joining Countless Other Species Disappearing Worldwide. The Internet Newspaper: Huff Post Green. Retrieved September 9th, 2012, from http://www.huffingtonpost.com/2011/01/03/ bumble-bee-decline_n_803896.html. Donovall, L. & vanEngelsdorp, D. (2010). A checklist of the bees (Hymenoptera: Apoidea) of Pennsylvania. Journal of Kansas Entomological Society, 83(1), 7-24. Droege, S. 2012. Tips on how to Use Bee Bowls to Collect Bees. Received September 16, 2012, from http://online.sfsu.edu/~beeplot/pdfs/bee%20bowl%20%20tip%20sheet1.pdf. Droege, S. 2011. Presentations. Retrieved 13 May 2011 from http://www.slideshare.net/ sdroege/presentations.

Native Pinelands Bees 30

Frohnapple, K. et al. (2010). Floral and nesting resources, habitat structure, and fire influence bee distribution across an open-forest gradient. Ecological Applications, 20(6), Abstract. Gibbs, J. (2011). Revision of the metallic Lasioglossum (Dialictus) if eastern North America (Hymenoptera: Halictidae: Halictini). Zootaxa, 3073. 1-216. Goff, C. (1937). Importance of bees in the production of watermelons. The Florida Entomologist, 20(2), 30-31. Griswold, T., Kremen, C., & Winfree, R. (2007, February). Effect of human disturbance on bee communities in a forested ecosystem. Conservation Biology, 21(1), 213-223. Hanula, J & Horn, S. (2011, Jan.). Removing an invasive shrub (Chinese privet) increases native bee diversity and abundance in riparian forests of southeastern United States. Insect Conservation and Diversity, 4(4), 275-283. Kearns, C. A., Inouye, D. W, & Waser, N. M. (1998). Endangered mutualisms: The conservation of plant-pollinator interaction. Annual Review of Ecology and Systematics, 29, 83-112. McCormick, J. (1970, December). The Pine Barrens: A Preliminary Ecological Inventory. New Jersey State Museum. Trenton, NJ. Pp 85. Michener, C. (1947). Bees of a limited area in southern Mississippi (Hymenoptera; Apoidea). American Midland Naturalist, 30(2), 443-455. Nature Search. (2008). Bees. Retrieved November 24, 2012 from http://www.fnanaturesearch.org/index.php?option=com_naturesearch&task=viewcat&ci =138.

Native Pinelands Bees 31

Neal, P., Pascarella, J., & Waddington, K. (1999). The bee fauna (hymenoptera: Apoidea) of Evergaldes National Park, Floridaand adjacent areas: Distribution, phenology, and biogepgraphy. Journal of the Kansas Entomological Society, 72(1), 32-45. Robichaud, B. (1980, March). A conceptual framework for pinelands decision-making. New Jersey Pinelands Commissions, 79(9), 1-78. Stevens, A. P. (2011). Discovering native bees. Green Teacher, 93, 1192-1285. Thomson, D. (2004). Competitive interactions between the invasive European honey bee and native bumble bees. Ecology, 85(2), 458-470. Triplehorn, C. & Johnson, N. (2005). Borror & Delong's Intro to Study of Insects, 7th Edition. Thomson. Brooks/Cole: United States. P. 864. Xerces Society for Invertebrate Conservation. (n.d.). Bumble Bee Conservation Initiative. Retrieved September 9th, 2012, from http://www.xerces.org/bumblebees/.