: | ISSN 1713-7845

JOURNAL ¥ . oe

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ENTOMOLOGICAL SOCIETY

me : : ; .

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Volume One Hundred and Forty One =

2010 ¥

THE ENTOMOLOGICAL SOCIETY OF ONTARIO

OFFICERS AND GOVERNORS

2009-2010 .

President: G. UMPHREY ESO Regional Rep to ESC: H. DOUGLAS

Department of Mathematics and Statistics Canadian Food Inspection Agency :

University of Guelph, Guelph, ON N1G 2W1 960 Carling Ave., Ottawa ON K1A 06C

umphrey@uoguelph.ca douglash@inspection.ge.ca

Librarian: J. BRETT

President-Elect: H. FRASER

Ontario Ministry of Agriculture, Food and Rural Affairs

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Vineland, ON LOR 2E0

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Past President: C. SCOTT-DUPREE

Dept. of Environmental Biology

University of Guelph, Guelph, ON NIG 2W1

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Secretary: N. MCKENZIE

Vista Centre, 1830 Bank Street, P.O. Box 83025

Ottawa, ON K1V 1A3

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Treasurer: S. LI

Natural Resources Canada, Pest Management Centre

Building 57, 960 Carling Ave.

Ottawa, ON K1A 0C6

shiyou@nrcan.ge.ca

Directors:

H. DOUGLAS

Canadian Food Inspection Agency

960 Carling Ave., Ottawa ON K1A 06C

hume.douglas@inspection.ge.ca

S. KULLIK

Department of Environmental Biology

(2008-2010)

(2009-2011)

University of Guelph, Guelph, ON N1G 2W1

kullik.sigrun@gmail.com

S. LACHANCE

(2010-2012)

University de Guelph - Campus D’ Alfred

31 St. Paul Street, Alfred, ON KOB 1A0

slachance@alfredc.uoguelph.ca

K. RYALL (2009-2011)

Canadian Forest Service, Great Lakes Forestry Centre, Natural

Resources Canada, 1219 Queen Street East,

Sault Ste. Marie, ON P6A 2E5

kryall@nrcan.gc.ca

K. RYAN

Faculty of Forestry, University of Toronto

Toronto, ON MS5S 3B3

kathleen.ryan@utoronto.ca

1. SCOTT

Agriculture and Agri-Food Canada

1391 Stanford Street, London, ON NSV 4T3

ian.scott@agr.gc.ca

(2008-2010)

(2010-2012)

FELLOWS OF THE ENTOMOLOGICAL SOCIETY OF ONTARIO

W. W. BILL JUDD

C. RON HARRIS

GLENN WIGGINS

BERNARD PHILOGENE

FREEMAN MCEWEN

Library, University of Guelph

Guelph, ON NIG 2W1

jimbrett@uoguelph.ca

Newsletter Editor: J. ALLEN

Ontario Ministry of Agriculture, Food and Rural Affairs

1 Stone Road West, Guelph, ON NIG 4Y2

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Student Representative: J. GIBSON

Department of Biology, Carleton University

1125 Colonel By Drive, Ottawa, ON K1S 5B6

jgibsonS@connect.carleton.ca

Website: M. JACKSON

School of Environmental Science

University of Guelph, Guelph, ON NIG 2W1

jackson@uoguelph.ca

JESO Editor: M. RICHARDS

Dept. of Biological Sciences, Brock University

St. Catharines, ON L2S 3A1

miriam.richards@brocku.ca

Technical Editor: J. VICKRUCK

Dept. of Biological Sciences, Brock University

St. Catharines, ON L2S 3A1

jess.vickruck@brocku.ca

Associate Editors:

A. BENNETT

Agriculture and Agri-Food Canada

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N. CARTER

Engage Agro Corporation

1030 Gordon St., Guelph, ON, N1G 4X5

neilcarter@engageagro.com

C. SCOTT-DUPREE

Dept. of Environmental Biology

University of Guelph, Guelph, Ontario NIG 2W1

cscottdu@uoguelph.ca

J. SKEVINGTON

Agriculture and Agri-Food Canada

Eastern Cereal and Oilseed Research Centre

960 Carling Ave., Ottawa, ON K1A 0C6

2002

2003

2006

2010

MCZ

JESO Volume 14 lAR Y

JOURNAL \AN -

of the

ENTOMOLOGICAL SOCIETY OF ONTARIO) v="

VOLUME 141 2010

In my previous Editor’s message, I commented that in an age where entomologists

are becoming a rare species, an adaptive response is to broaden our perspective on

entomology to include all science focused on insects. When I first joined the ESO and

attended my first ESO conference, Yves Prevost, the previous JESO Editor, asked me in a

very friendly manner, whether I had just moved to Ontario. I replied that no, I had been

at Brock for five or six years. Yves asked why I had not been at previous meetings, and |

replied that I wasn’t really sure that I was an entomologist. Yves looked really puzzled, and

inquired if I studied insects. When I told him that I studied bees, he responded ‘Well, then,

you are an entomologist!” I have always treasured that conversation, because it represented

to me such a broad and welcoming perspective in defining entomology. This perspective

has also guided me as Editor of JESO — all sorts of insect science are welcome at ESO and

JESO.

[have been Editor of JESO for about 5 years, starting with Volume 135 (2005) which

I co-edited with Yves. I am very pleased that we have managed to accomplish so much in

that time. The Journal is back to a regular, annual publication cycle and is now distributed

electronically from the ESO website, as well as being published in the more classical paper

format. An exciting development is a plan to include JESO in a large electronic journal

listing service which will lead to a quantum leap in journal visibility. Although final details

are not yet settled, I anticipate that this listing will begin with the current volume. Another

exciting development is a plan to have all back issues made available electronically in the

near future.

With these developments, JESO will have successfully accomplished the mission

that I set out when I first became Editor. I sincerely thank Yves, all the Associate Editors,

and the ESO Board for entrusting JESO to me, in the process convincing me, that yes, I

am an entomologist. I am very proud of our accomplishments and very gratified that I

can pass the baton to a new Editor, knowing that JESO is thriving. I will be co-editing

Volume 142 (2011) with a new Editor, who I am sure will guide the Journal through its next

developmental phase.

Happy reading!

Miriam H. Richards

Editor

Aerial foraging in burying beetles JESO Volume 141, 2010

AERIAL FORAGING AND SEXUAL DIMORPHISM IN BURYING

BEETLES (SILPHIDAE: COLEOPTERA) IN A CENTRAL

ONTARIO FOREST

D. L. LEGROS AND D. V. BERESFORD!

Department of Biology, Trent University, 1600 West Bank Drive,

Peterborough, ON, Canada K9J 7B8

email: davidberesford@trentu.ca

Abstract J. ent. Soc. Ont. 141: 3-10

Burying beetles (Coleoptera: Silphidae) are commonly sampled on the

ground using pitfall traps. Recent work has shown that these beetles also

respond to aerial traps baited with carrion. In this study, we sampled the

Silphidae of Algonquin Park using traps baited with mouse and bird carrion,

and set at 2, 4, and 6 m heights. The most abundant species caught was

Nicrophorus tomentosus, followed by (in order) N. defodiens, N. sayi,

Oiceoptoma noveboracense, N. pustulatus, Necrophila americana, and

Necrodes surinamensis. Only N. tomentosus showed bait preference, with

higher than expected catches at traps baited with mice. Catches differed

based on trap height for two species, with most N. defodiens being caught

in the lower traps (2 m), and all N. pustulatus caught at the high traps (6

m). Nicrophorus tomentosus males caught in the 6 m traps were significantly

larger than males caught in the lower traps, and females caught at all heights.

Possible reasons are discussed.

Published November 2010

Introduction

The carrion beetles (Family Silphidae) have long been a favourite group with

collectors and naturalists in North America. There are 30 species of these in North America,

and individual species specialize in carrion types based on size and source, ranging from

large carcasses such as black bears (Watson and Carlton 2003) (e.g. subfamily Silphinae),

to small rodent sized carcasses such as mice and song birds (subfamily Nicrophorinae, the

burying beetles; Anderson and Peck 1985). Burying beetles use olfactory cues to locate

carcasses, and are able to locate fresh squirrel carcasses several metres distant in a few

minutes (Dethier 1947).

' Author to whom all correspondence should be addressed.

LeGros and Beresford a el ‘JESO Volume 141, 2010

In spite of the attention that this group has attracted in the past, basic life history

and niche information is still lacking. For example Nicrophorus pustulatus Herchel was

only recently determined to be a parasitoid on snake eggs (Blouin-Demers and Weatherhead

2000; Keller and Heske 2001). Regional surveys of extant Nicrophorus species do report

catching N. pustulatus in low numbers (e.g., Anderson 1982; Shubeck et al. 1981), and

outside of the range of the black rat snake (Smith et al. 2007). It appears that N. pustulatus

must utilize some other unknown source of carrion.

What this is may be consistent with the recently observed vertical distribution of N.

pustulatus in Georgia, where N. pustulatus were almost exclusively caught several metres

above ground (Ulyshen and Hanula 2007a) in the forest canopy. The canopy habitat includes

tree cavities, and these could offer a specialized niche of carrion such as dead nestlings, e.g.

squirrels, birds, or bats. Any species differences in how this habitat is exploited should be

reflected in catches obtained using carrion baited traps placed at various heights.

In this study, we report on the species composition of Silphidae caught in carrion

baited traps placed at three different heights above the ground in Algonquin Park, a large

forested region of central Ontario. We also compared trap catches at traps with avian and

mammalian baits.

For burying beetles, there are reproductive advantages to size, with larger species

and individuals winning fights over carcasses (Otronen 1988; Trumbo 1990). Body size is

also related to flight capability. During flight, beetles lose heat due to convection (Merrick

and Smith 2004), so that conserving heat would enable longer flights, for example, flight

associated with searching or localized dispersal. Such flights would take place at higher

levels (Taylor 1974), so we would expect elevated traps to catch these individuals. For the

most numerous species caught, NV. tomentosus, we tested for differences in the size of males

and females at each height, reasoning that such size differences might indicate sex biased

flight capability associated with dispersal from natal sites.

Materials and Methods

Silphid beetles were collected during August 2008, in Algonquin Provincial Park,

Ontario, Canada. The study site was located along Highway 60, at kilometre 20, Found

Lake, Peck Township on the Canadian Shield. The forest was composed of mature trees:

maple (predominantly Acer saccharum Marsh.), beech (Fagus grandifolia Ehrh.), hemlock

(Tsuga canadensis (L.) Carr.), and birch (Betula papyrifera Marsh and Betula allegheniensis

Britton). The understory was shaded, was easily accessible by foot, and composed mainly

of bracken (Pteridium aquilinum (L.) Kuhn) with some low density shrubs.

Twenty-four traps were deployed for seven day periods beginning 1, 7, and 14

August 2008, for a total of 72 samples. During sampling, many traps were destroyed by

wildlife (bears and raccoons) and wind. If damage occurred early in the sampling week,

traps were re-installed for later collection, so that the length of a trapping period varied for

some traps. Fourteen ruined traps could not be replaced. For each sampling week, traps

were positioned in a grid or block comprised of three rows of eight traps per row. The rows

Aerial foraging in burying beetles JESO Volume 141, 2010

were spaced 15 m apart, with traps in each row spaced 10 m. Traps were placed at 2, 4, and

6 m above ground, with 6 m traps placed in the upper trunk zones but just below the canopy

level. Trap height and bait type were randomized over the 24 possible locations in each

block (there were eight possible X and three possible Y coordinates), so that there were

four traps for each bait type at each height in the block.

Traps were constructed from 2 L plastic soda pop bottles with a 6 cm diameter

hole cut into the middle of one side to allow beetles to enter the traps. Bait was wrapped in

cheese cloth and hung in each bottle by a piece of wire pushed through the lid. The bottom

of each trap had a 1.5 cm deep layer of killing solution consisting of 5% dish soap and water

(Larsen 2005). A sling shot was used to send a weighted rope over a tree branch to raise

and lower traps.

Half the traps were baited with chicken wings and the other half with mice. Frozen

domestic mice (Mus musculus) were obtained from a reptile feed supply store. The mean

weight per mouse was ~ 25 g. Frozen chicken wings were obtained in bulk from the grocery

store (mean weight ~ 30 g). Baits were frozen until placed in traps without any prior aging

or ripening.

Trapped beetles were preserved in 70% isopropyl alcohol. A sub-sample of 320

Nicrophorus tomentosus, about 100 specimens from each height, were sexed and measured

for total body length.

Because of the missing data, ANOVA could not be used to test the effects of bait

type and trap height on trap catches. Instead, we used a chi-square test (Sokal and Rohlf

1995) to compare the observed to expected total catch frequencies of each species for

bait type and trap height. The expected frequencies were determined from the number of

trapping days for each category. For example, if 140 individuals of a species were caught

over 50 trapping days at traps baited with mice, and 220 were caught over 100 trapping days

at traps baited with birds, then the expected frequencies would be (140+220) x (50/150)

=120 for mouse traps and (140+220) x (100/150) = 240 for bird traps. Because several

comparisons were performed for each test, the critical values of chi-square at a significance

level of a = 0.05 were adjusted using the Bonferroni method by which a’ = a/k (Sokal and

Rohlf 1995). For the bait tests and the trap height tests we used k = 7 (number of species

tested). Our adjusted critical values for chi-square were 9.94 for 2 degrees of freedom

(three trap heights), and 7.33 for 1 degree of freedom (two bait types).

We tested whether there were differences in body length of N. tomentosus

between males and females at each height using a 2-way ANOVA with trap height and sex

as treatments. Statistical tests were done using an Excel spreadsheet and STATISTICA 7

(Statsoft Inc. 2004).

Results

We caught 2388 Silphidae from 7 species in the traps (Table 1). The most abundant

species was Nicrophorus tomentosus, accounting for 81% of the total catch. The rarest

species was Necrodes surinamensis, of which only a single individual was caught, this

at 4 m. Members of each species were more abundant in the 2 m traps except for N.

LeGros and Beresford JESO Volume 141, 2010

TABLE 1. Number of each species of Silphidae captured in carrion baited aerial traps at

three different heights in Algonquin Park, Ontario, in August 2008.

Trap height

Species . 2m 4m 6m Total

Nicrophorus tomentosus 863 497 585 1945

Nicrophorus defodiens 19] 76 42 309

Nicrophorus sayi 43 22 25 90

Oiceoptoma noveboracense 21 7 5 33

Nicrophorus pustulatus 0 0 6 6

Necrophila americana * 0 0 -

Necrodes surinamensis 0 l 0 l

Total 1122 603 663 2388

pustulatus, which was only found in 6 m traps (Table 1). MN. tomentosus was the only

species for which trap catches differed by bait types, with slightly more caught at mouse

baited traps than expected (observed = 856, expected = 791.9; Table 2).

More N. defodiens were caught in the lowest traps and more N. pustulatus in the

higher traps (Table 2). For the other species, observed trap catches at the three heights did

not differ from the expected proportions.

The biggest male N. tomentosus were caught in the 6 m traps; these were

significantly larger than females caught at all heights (Duncan’s Multiple Range post-hoc

test, 2 metres: p=0.003; 4 metres: p=0.024; 6 metres, p=0.025) and males at 2 metres (2

metres: p=0.012; 4 metres, p=0.07) (Table 3, Fig. 1).

female

male

a=_,

rs)

number per size class

—)

2 metres 4 metres 6 metres

length mm

FIGURE 1. Frequency distribution of body lengths of female and male N. tomentosus at 2,

4, and 6 metre traps.

JESO Volume 141, 2010

Aerial foraging in burying beetles

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Discussion

Nicrophorus tomentosus and N. defodiens were the most common species captured

in this study. Work done in North Carolina has shown that these species are frequently

encountered late in the warm season (Trumbo 1990). In southern Ontario, N. tomentosus

breeds from mid to late summer (Anderson and Peck 1985), with trap catches peaking in

late August (Anderson 1982); N. defodiens is active from May to early September and has

been trapped at similar levels during this period (Anderson 1982). Nicrophorus sayi, a

spring breeder in Ontario (Anderson 1982) was the third most common species captured,

consistent with a previously reported peak in early and late August of teneral adults

(Anderson 1982).

Only N. tomentosus showed any bait preference, with a minor preference for traps

with mice. For the Nicrophorus spp., small carrion would include dead fledglings as well

as dead rodents. In terms of possible tree hole exploitation, both types of carrion would be

present. Generally, Silphinae are attracted to larger carrion (Anderson and Peck 1985).

Height Preference

Nicrophorus were caught at all heights except N. pustulatus and N. defodiens.

Nicrophorus defodiens is a small species, and it compensates for this competitive

disadvantage by being able to locate carcasses quicker than its larger competitors, such as

N. orbicollis Say (Trumbo and Bloch 2002). That it was caught less often in the higher traps

may reflect a possible decreased flight height due to its smaller size rather than its inability

to find the bait. |

Our results support our initial reasoning that carrion in the forest canopy might be a

suitable specialized niche for exploitation for N. pustulatus. Our low catch of 6 individuals

is similar to low catches reported in previous studies (Anderson 1982; Robertson 1992).

Nicrophorus pustulatus does not appear to respond to traps baited with fresh carcasses

(Trumbo 1990). However, it has been caught in pitfall traps baited with well-rotted carrion

(Trumbo 1990; Anderson 1982).

Catching N. pustulatus at elevated traps is consistent with earlier work (Ulyshen

and Hanula 2007b). In Ontario, Robertson (1992) caught N. pustulatus at 1m to 2m. Low

catches of N. pustulatus at ground level could be due to this species specializing in canopy

or nest-cavity habitats rather than specific carrion sources; N. pustulatus will breed on dead

rodents in lab settings (Robertson 1992).

Nevertheless, black rat snake eggs are the only known wild breeding medium

(Smith et al. 2007; Blouin-Demers and Weatherhead 2000), which may explain the generally

low numbers caught using mammalian or avian baits. Because of this, the presence of N.

pustulatus is relevant to the conservation of regionally rare or endangered snake species

(Smith et al. 2007) e.g. the black rat snake Elaphe obsoleta (Say) in Ontario, a species not

found in Algonquin Park (Logier 1970). Our results suggest that surveying for NV. pustulatus

should include aerial traps.

Sex and Size Dimorphism in Nicrophorus tomentosus

For N. tomentosus, because males and females both remain with their brood after

Aerial foraging in burying beetles JESO Volume 141, 2010

covering a carcass, it is unlikely that there is a size advantage for one sex or the other in

terms of contests over carcasses. However, there may be a size advantage in terms of flight

capability. During flight, beetles lose heat due to convection (Merrick and Smith 2004), so

that conserving heat would enable longer flights, for example flight associated with local

dispersal. Nicrophorus tomentosus is a bumble bee mimic (Fisher and Tuckerman 1986),

and is covered in yellow hairs (Milne and Milne 1944), which can contribute to conserving

heat and regulating body temperature in Nicrophorus spp. (Merrick and Smith 2004).

The female Nicrophorus tomentosus we caught were the same size at all trap

heights, whereas males caught on 6 m traps were significantly larger than both females and

males from lower traps (Table 3). Trumbo (1990), found no difference in the size of male

and female Nicrophorus tomentosus, nor did we in our lower traps. The larger males were

largely present in 6 m traps only (Fig. 1). Sampling at the higher traps would over-represent

this group.

Acknowledgments

The authors thank Dr. James Sutcliffe of Trent University, Algonquin Provincial

Park biologist B. Steinberg, as well as Peter Mills, Nathan G. Miller, and Natalie Earl for

their assistance.

References

Anderson, R. S. 1982. Resource partitioning in the carrion beetle (Coleoptera: Silphidae)

fauna of southern Ontario: Ecological and evolutionary considerations. Canadian

Journal of Zoology 60: 1314-1325.

Anderson, R. S. and S. B. Peck. 1985. The Carrion Beetles of Canada and Alaska. Coleoptera:

Silphidae and Agyrtidae. The Insects and Arachnids of Canada, Part 13. 121 pp.

Blouin-Demers, G. and P. J. Weatherhead. 2000. A novel association between a beetle and

a snake: parasitism of Elaphe obsoleta by Nicrophorus pustulatus. Ecoscience 7:

395-456.

Dethier, V. G. 1947. The role of the antennae in the orientation of carrion beetles to odors.

Journal of the New York Entomological Society 55: 285-293.

Fisher, R. M. and R. D. Tuckerman. 1986. Mimicry of bumble bees and cuckoo bumble bees

by carrion beetles (Coleoptera: Sylphidae). Journal of the Kansas Entomological

Society 59: 20-25.

Keller, W. L. and E. J. Heske. 2001. An observation of parasitism of black rat snake (Elaphe

obsoleta) eggs by a beetle (Nicrophorus pustulatus) in Illinois. Transactions of the

Illinois State Academy of Science 94: 167-169.

Larsen, T. 2005.:Trap spacing and transect design for dung beetle biodiversity studies.

Biotropica 37: 322-325.

Logier, E. B. S. 1970. The snakes of Ontario. University of Toronto Press, Toronto. 94 pp.

LeGros and Beresford . JESO Volume 141, 2010

Merrick, M. J.and R .J. Smith. 2004. Temperature regulation in burying beetles (Nicrophorus

ssp. Coleoptera: Silphidae): effects of body size, morphology and environmental

temperature. The Journal of Experimental Biology 207: 723-733.

Milne, L. J. and M. J. Milne. 1944. Notes on the behavior of burying beetles (Nicrophorus

spp.). Journal ofthe New York Entomological Society 52: 311-327.

Otronen, M. 1988. The effect of body size on the outcome of fights in burying beetles

(Nicrophorus). Annales Zoologici Fennici. 25:191-201.

Robertson, I. C. 1992. Relative abundance of Nicrophorus pustulatus (Coleoptera: Silphidae)

in a burying beetle community, with notes on its reproductive behavior. Psyche 99:

189-198.

Shubeck, P. P., N. M. Downie, R. L. Wenzel and S. B. Peck. 1981. Species composition and

seasonal abundance of carrion beetles in an oak-beech forest in the Great Swamp

National Wildlife Refuge (N.J.). Entomological News 92: 7-16.

Smith, G., S. T. Trumbo, D. S. Sikes, M. P. Scott and R. L. Smith. 2007. Host shift by

the burying beetle, Nicrophorus pustulatus, a parasitoid of snake eggs. Journal of

Evolutionary Biology 20: 2389-2399.

Sokal, R. R. and F. J. Rohlf. 1995. Biometry. W. H. Freeman and Company, New York.

StatSoft Inc. 2004. STATISTICA (data analysis software system), version 7. http://www.

statsoft.com.

Taylor, L. R. 1974. Insect migration, flight periodicity and the boundary layer. Journal of

Animal Ecology 43: 225-238.

Trumbo, S. 1990. Reproductive success, phenology and biogeography of burying beetles

(Silphidae, Nicrophorus). The American Midland Naturalist 124: 1-11.

Trumbo, S. and P. L. Bloch. 2002. Competition between Nicrophorus orbicollis and N.

defodiens: resource locating efficiency and temporal partitioning. Northeastern

Naturalist 9: 13-26.

Ulyshen, M. D. and J. L. Hanula. 2007a. Burying beetles (Coleoptera: Silphidae) in the

canopy: The unusual case of Nicrophorus pustulatus Herschel. The Coleopterists

Bulletin 61: 121-132.

Ulyshen, M. D. and J. L. Hanula. 2007b. A comparison of the beetle (Coleoptera) fauna

captured at two heights above the ground in a North American temperate deciduous

forest. American Midland Naturalist 158: 260-278.

Watson, E. J. and C. E. Carlton. 2003. Spring succession of necrophilous insects on wildlife

carcasses in Louisiana. Journal of Medical Entomology 40: 338-347.

Natural enemies of Ceratina in the Niagara Region JESO Volume 141, 2010

: NATURAL ENEMIES OF THE BEE GENUS CERATINA

! (HYMENOPTERA: APIDAE) IN THE NIAGARA REGION,

: ONTARIO, CANADA

J. L. VICKRUCK’, J.T. HUBER? AND M. H. RICHARDS

Department of Biological Sciences, Brock University,

St. Catharines, ON, Canada L2S 3A1

email: jess.vickruck@brocku.ca

Abstract J. ent. Soc. Ont. 141: 11-26

Ceratina dupla and C. calcarata (Hymenoptera: Apidae) are abundant bees

in southern Ontario, commonly nesting in staghorn sumac (Rhus typhina),

wild raspberry (Rubus strigosus) and teasel (Dipsacus fullonum). Ceratina

nests were collected from April-September 2008 and parasitized individuals

were reared to adulthood in the laboratory. Pyemotes sp. (Pyemotidae) and

Baryscapus americana (Eulophidae) were the most common natural enemies,

followed by Baryscapus sp., Axima zabriskiei, and Hoplocryptus zoesmairi.

Eupelmus vesicularis, Coelopencyrtus sp. and Eurytoma sp. near apiculae

were rarely collected. This is the first record of E. vesicularis (Eupelmidae)

as a primary host on any member of the family Apidae. New host records are

also reported for H. zoesmairi (Ichneumonidae), two different Baryscapus

spp. (Eulophidae), Eurytoma sp. near apiculae (Eurytomidae) and Pyemotes

sp. (Pyemotidae) on C. calcarata and C. dupla. Detailed descriptions of

immature development of the parasitoids, and their preferences for host bee

species and host plant species are provided.

Published November 2010

Introduction

Parasite—host relationships have been studied for numerous species in a laboratory

setting (Harri et al. 2008; Jervis et al. 2008; Traynor and Mayhew 2005). These studies are

vital to help understand the dynamics of host—parasite interactions, however they often only

involve the most common one or two parasitoids associated with the host under study. Ina

' Author to whom all correspondence should be addressed

* Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, ON,

K1A 0C6

Vickruck et al. . JESO Volume 141, 2010

natural setting, hosts may be parasitized by a number of species af varying frequencies, each

using different parasitism and developmental strategies at different times. By describing

the life history, development and preferences of numerous parasite species attacking one

host, a more complete understanding of these interactions is gained.

Bees of the genus Ceratina Latreille (often referred to as dwarf carpenter bees),

are cosmopolitan, with the subgenus Zadontomerus Ashmead being found exclusively in

the Western Hemisphere (Michener 2007). The life history of Ceratina offers an excellent

opportunity to study the development and interactions of parasites with their hosts. All

offspring from eggs laid by a single female can be collected together in a nest, thus allowing

for observation of how the parasites interact with an individual host, as well as how nest

substrate, position in the nest, and interactions with other parasites and the foundress bee

occur.

The Niagara Region, Ontario, Canada, is home to two common species of Ceratina

(Zadontomerus): C. dupla Say and C. calcarata Robertson. Their nests are commonly

collected from staghorn sumac (Rhus typhina L.), wild raspberry (Rubus strigosus Michaux)

and teasel (Dipsacus fullonum L.; J. Vickruck, unp. data). Both sumac and wild raspberry

are native to the region whereas teasel is an obsolete crop plant introduced from Europe,

whose flower heads (when the seeds are mature) were once used to raise the knap on

wool (Rector et al. 2006). Sumac and raspberry are both perennial plants found at wood

margins, differing from teasel which is a biennial weed found in open, generally abandoned

agricultural fields.

Ceratina in the Niagara region are solitary and-univoltine, producing one brood

per year and overwintering as newly emerged, unmated adults (J. Vickruck, unpublished

data). Emergence and mating typically take place in mid-April, and new nests are founded

in May. Nests are not reused from year to year and can only be initiated in twigs with

exposed pith. After digging a linear tunnel females begin to forage, forming pollen and

nectar provisions into rounded masses upon which a single egg is laid (Grothaus 1962;

Johnson 1988; Kislow 1976). Each provision mass and egg is separated from its neighbours

by a cell septum formed by the foundress. Once finished provisioning, females sit and

guard the nest entrance until the eclosion of their offspring. The newly eclosed adults can

either overwinter in their natal nest or disperse to found new hibernacula for the winter

(Grothaus 1962; Kislow 1976).

Ceratina immatures can be classified into 18 developmental stages which were

originally described by Daly (1966) for C. dallatoreana. The first eight stages rank the

pupa in relation to the size of the pollen ball, after which the immature passes through a

pre—pupal stage followed by metamorphosis. The eyes of the pupa then pass from white

through to black (five stages), followed by darkening of the body (four stages). In the final

stage the black bodied pupa emerges as an adult with milky wings.

Natural enemies of Ceratina in the Niagara region include predators, parasites,

and parasitoids. Predators consume more than one prey individual in order to complete

development. Parasites feed on the host contents but do not consume the entire host

before completing development (Godfray 1994). Parasitoids consume a single host in

order to complete development (Godfray 1994). Parasitoids were classified as idiobionts

or koinobionts, endoparasitoids or ectoparasitoids, and gregarious or solitary. Idiobionts

prevent the larva from developing further after initial parasitisation (Askew and Shaw 1986).

Natural enemies of Ceratina in the Niagara Region JESO Volume 141, 2010

Koinobionts do not kill the host until it has reached a certain point in its development,

as the parasitoid benefits from the continued life of the host (Askew and Shaw 1986).

Ectoparasitoids develop outside the host (although they are often attached to it), while

endoparasitoids consume the host internally. In solitary species the parasitoid to host ratio

is 1:1, whereas in gregarious parasites multiple individuals develop in one host.

The objectives of this study were to identify and describe the development of

the natural enemies of Ceratina in the Niagara Region as well as quantify their host and

substrate preferences.

Materials and Methods

Host nest collections

All parasites were reared from a total of 107 nests of C. calcarata and C. dupla

collected from 14 April to 30 September 2008. Each week at least 15 Ceratina nests were

collected so that sampling effort was consistent over the season. Supplementary nest

collections also took place in June 2009 to aid with final parasite identifications only. The

2009 data are not included in the statistical analysis. All collections took place at the Brock

University campus (43.1197, —79.2492), the Glenridge Quarry Naturalization Site (43.1223,

—79.2375) and an abandoned old field site near the Welland Canal (43.1479, —79.1811).

Nests were collected from sumac, raspberry, and teasel and brought back to the laboratory

in early morning to ensure that all occupants were present inside. After being chilled, twigs

were carefully split open longitudinally to identify nest contents. Bee species, plant nest

substrate, position of any parasitized cells in the nest, and developmental stages of bees

and parasites were recorded on the day of collection. Dissected nests were then inserted in

transparent PVC tubing slightly larger than the diameter of the nest (ranging from '2—1 inch

depending on twig diameter) for protection and to allow for easy visual observation of nest

contents. This also allowed for behavioural observations of host—parasite interactions in

the laboratory.

Ceratina species were identified using the key of Rehan and Richards (2008).

Parasite identifications were made by JTH and Dr. Gary Gibson at the Canadian National

Collection of Insects, Arachnids and Nematodes (CNC), Dr. Michael Gates at the National

Musuem of Natural History (NMNH) in Washington, DC, as well as JV. Dr. Andrew

Bennett (CNC) verified identity and nomenclature of Hoplocryptus zoesmairi Dalla Torre.

Voucher specimens of Baryscapus sp. and americana, Eupelmus vesicularis (Retzius),

Coelopencyrtus sp., Axima zabriskiei Howard and Eurytoma sp. near apiculae, were

deposited in the CNC. Baryscapus sp., Coelopencyrtus sp., Eupelmus vesicularis and

Eurytoma sp. near apiculae are labelled as CNC Ident. lot # 2008-341, and Baryscapus sp.,

Axima zabriskiei and H. zoesmairi as 2009-188.

Parasite development and classification

Hosts were observed on a daily basis to detect parasitoid presence. Position in

the nest, stage parasitized, and parasitoid species were recorded as soon as they became

apparent. Developmental milestones such as defecation, pupation, pigmentation of the

exoskeleton as well as emergence dates were recorded for parasites whenever possible.

Vickruck et al. JESO Volume 141, 2010

Once parasitoids had pupated they were transferred to their own individual 0.2 mL

microcentrofuge tubes prior to eclosion. Upon emergence parasitoids were placed in 70%

ethanol for later identification.

Data Analysis

All data were analyzed using SAS 9.1.. Parasite prevalence is defined as the

number of individuals affected by a particular parasite species divided by the number

of hosts examined (Margolis et al. 1982). Parasite frequency is defined as whether that

particular species is present in the nest, regardless how many individuals in the nest were

parasitized. G-tests for goodness of fit were used when sample sizes were large, Fisher’s

exact tests were used when expected values were small (<5). Hoplocryptus zoesmairi has

not been included in statistics as it is a predator.

Results

Host parasitism

Eight species of arthropod parasites representing two classes (Insecta, Arachnida),

two orders (Hymenoptera and Trombidiformes), and seven families were reared from a

total of 107 C. dupla and C. calcarata nests containing 850 brood cells. Characteristics

of these eight species are compared in Table 1. Of the 107 nests collected, 64 were

teasel, 36 raspberry, and 7 sumac. Twenty-nine percent (243/850) of all brood cells were

parasitized, and 68% (73/107) of nests contained at least one parasite. Ceratina calcarata

had a significantly higher proportion of cells parasitized than C. dup/a but the proportion

of nests parasitized between host species did not differ significantly (Table 2). Parasitism

for each Ceratina species also varied by substrate, with significantly higher proportion of

cells and nests parasitized in raspberry compared to teasel (Table 2). Ceratina calcarata

was parasitized more often than C. dupla when nesting in raspberry (G=20.05, d.f.=1,

P<0.0001), however no difference was seen between species in teasel (G=0.04, d.f=1, n.s.)

Sumac nests were not included in substrate comparisons due to small sample size. Ceratina

dupla nesting in raspberry was the least parasitized with 15% of available cells affected

(Table 2, Fig. 1). Only seven sumac nests were found, all C. calcarata, in which 16/33

(48%) of immatures had been parasitized (Fig. 1).

Parasite and predator development

The frequency and prevalence, i.e., proportion of hosts affected, of all parasites

and predators in Ceratina nests is presented in Table 3 for affected cells and Table 4 for

affected nests. Detailed observations for each species are given below.

Predators

Hoplocryptus zoesmairi (Dalla Torre) (Hymenoptera: Ichneumonidae)

This species has previously been associated with C. dupla (as Habrocryptus

graenicheri Viereck; Viereck 1904). Hoplocryptus Thomson has until recently been

considered a junior synonym of Aritranis (Férster) (Yu and Horstmann 1997), but Yu et

JESO Volume 141, 2010

Natural enemies of Ceratina in the Niagara Region

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Vickruck et al. JESO Volume 141, 2010

TABLE 2. Prevalence of natural enemies associated with Ceratina dupla and C. calcarata

in each substrate. Due to low sample sizes sumac was excluded from statistical analysis of

parasitism amongst substrates.

Prevalence (“%)

6 t

se ea lyfe Brood cells Nests

C. dupla Teasel 107/426 27/49

Raspberry 15/97 9/10

Total 122/523 36/59

C. calcarata Teasel 25/96 8/15

Raspberry 80/198 25/26

Sumac 16/33 4/7

Total 121/327 37/48

Grand Total 243/850 73/107

C. dupla vs. C. calcarata G=18.42, d.f.=1, P<0.0001 G=3.21, d.f.=1, P=0.07

Teasel vs. Raspberry G=4.44, d.f=1, P=0.04 G=20.21, d.f.=1, P<0.0001

0.60

no]

N 0.50

@ 0.40

rm OBs.1

S = 8 Col

: 0.20 MEV.

= VILLETTE: Eyt.

a 0.10 - m@ Axm.

a gm Pym.

0.00 -

C. dupla C. dupla C. calcarata C. calcarata C. calcarata

Raspberry Teasel Teasel Raspberry Sumac

Host species and substrate

FIGURE 1. The proportion of available cells parasitized for Ceratina dupla and C. calcarata

in each substrate. Values associated with each bar indicate the number of available cells

for each species in each substrate. Abbreviations: Bs. 1.=Baryscapus americana, Bs.

2=Baryscapus sp., Col.= Coelopencyrtus sp., E.v.= Eupelmus vesicularis, Eyt.= Eurytoma

sp. near apiculae, Axm.=Axima zabriskiei, Pym.= Pyemotes sp.

Natural enemies of Ceratina in the Niagara Region

JESO Volume 141, 2010

TABLE 3. Prevalence of parasites on each Ceratina host by affected brood cells. Prevalence

is the proportion of brood parasitized in each host species in each nesting substrate.

Significance testing was conducted using G tests unless indicated by * where Fisher’s Exact

tests were used.

Parasite

Baryscapus americana

(Eulophidae)

Baryscapus sp.

(Eulophidae)

Coelopencyrtus sp.

(Encyrtidae)

Eupelmus vesicularis

(Eupelmidae)

Eurytoma sp. near apiculae

(Eurytomidae)

Axima zabriskiei

(Eurytomidae)

Pyemotes sp.

(Pyemotidae)

Host

C. dupla

C. calcarata

C. dupla

C. calcarata

C. dupla

C. calcarata

C. dupla

C. calcarata

Substrate

Teasel

Raspberry

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Total

Teasel

Raspberry

Sumac

Total

Prevalence cells

available

55/426

2/97

57/523

9/96

0/198

0/33

9/327

0/96

51/198

13/33

64/327

0/96

0/198

1/33

1/327

1/426)

0/97

1/523

1/96

0/198

0/33

1/327

0/426

8/97

8/523

0/96

14/198

2/33

16/327

51/426

5/97

56/523

15/96

15/198

0/33

30/327

Statistics

Species:

G= 21.42, d.f.=1,

P<0.0001

Substrate:

G=46.68 ,d.f=1,

P<0.0001

Substrate

G= 2.49, d.f.=1,

N.S.

Species:

G= 8.02, d.f.=1,

P=0.005

Substrate:

*X7= 0.08, d.f.=1,

n.s.

Species:

G=0.53, d.f.=1, n.s.

Substrate:

G= 7.56 d.f.=1,

P=0.006

al. (2005) consider Hoplocryptus a valid genus. This is the first time this species has been

recorded with C. calcarata.

This natural enemy is considered a predator as it always consumed multiple

Vickruck et al.

JESO Volume 141, 2010

TABLE 4. Infection rate of nests of Ceratina spp. by substrate type and species of natural

enemy. Nests were scored as infected if at least one individual of the eight natural enemies

were present in a nest. Significance testing was conducted using G tests unless indicated

by * where Fisher’s Exact tests were used.

Natural Enemy

Baryscapus americana

(Eulophidae)

Baryscapus sp.

(Eulophidae)

Coelopencyrtus sp.

(Encyrtidae)

Eupelmus vesicularis

(Eupelmidae)

Eurytoma sp. near apiculae

(Eurytomidae)

Axima zabriskiei

(Eurytomidae)

Pyemotes sp.

(Pyemotidae)

Host

C. dupla

C. calcarata

C. dupla

C. calcarata

C. dupla

C. calcarata

C. dupla

C. calcarata

Substrate

Teasel

Raspberry

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Total

Teasel

Raspberry

Sumac

Total

Teasel

Raspberry

Total

Teasel

Raspberry

Sumac

Total

ests wit

at least one

natural enemy/

Total nests

13/49

2/10

15/59

2/15

0/26

0/7

2/48

0/15

16/26

2/7

18/48

0/15

0/26

1/7

1/ 48

1/49 |

0/10

1/59

1/15

0/26

0/7

1/48

0/49

4/10

4/59

0/15

7/26

1/7

8/48

13/49

3/10

16/59

5/15

2/26

0/7

7/48

Statistics

Species:

G= 10.16, d.f.=1,

P=0.001

Substrate:

G= 6.03, d.f.=1, P=0.01

Substrate:

*X*= 2.45, d.f.=1, nus.

Species:

G= 2.60, d.f=1, n.s.

Substrate:

*X?= 0.87, d.f.=1, n.s.

Species:

G= 2.53, d.f.=1, n.s.

Substrate:

G= 2.80, d.f.=1, n.s.

Ceratina immatures before completing development. The predator egg was always laid

in the innermost cell of the nest. After the egg hatched, the larva attached to the small

Ceratina larva, but did not kill it immediately. Rather, the H. zoesmairi larva waited until the

Natural enemies of Ceratina in the Niagara Region JESO Volume 141, 2010

Ceratina larva was at least half as large as its pollen mass, at which point it consumed

the immature Ceratina and the remainder of its provisions. Once the entire contents of

the cell had been consumed the larva broke down the cell septum and consumed the next

larva and its pollen mass. This process was repeated, with individual H. zoesmairi larvae

devouring anywhere from two to five Ceratina immatures and pollen masses, then spinning

silken cocoons. Each H. zoesmairi larva then defecated and pupated inside its cocoon before

emerging as an adult. Development from time of hatching to adulthood took 27-48 days,

with emergence dates ranging from 28 July to 14 August 2008. There were four occurrences

of this predator, two in C. dupla nests (one in teasel and one in raspberry), one in a C.

calcarata nest (raspberry), and one in a Ceratina nest that contained no adult female and no

surviving offspring.

Parasitoids

Baryscapus americana (Ashmead) (Hymenoptera: Eulophidae)

Baryscapus americana was previously known to parasitize C. calcarata in

Georgia (Kislow 1976) and Missouri (Rau 1928). The species was transferred from the

genus Aprostocetus Westwood by Lasalle (1994). This is the first record of any member of

the genus Baryscapus Forster parasitizing C. dupla.

Baryscapus americana is a gregarious, koinobiont endoparasitoid of Ceratina

immatures. Their presence was undetectable until they began to consume their hosts (Fig.

2a), but the larvae grew to approximately half the length of their Ceratina host by the time its

contents had been entirely consumed. At this point the parasitoids migrated to the anterior

or posterior ends of the pre-pupal skin (Fig. 2b). The parasitoids then emerged in three

ways: either all individuals in the Ceratina larval skin pupated and emerged that summer

(Fig. 2c), or all of the individuals remained as prepupae to overwinter together and emerge

the following spring, or several individuals occupying a single host would pupate while

the rest would overwinter. The aforementioned strategies were also observed by Kislow

(1976). Of the 66 immature Ceratina parasitized, 20 (30%) showed total emergence, 35

(53%) overwintered as a group together, and 11 (17%) showed partial emergence, with

some individuals emerging that summer and some overwintering as prepupae. Average

development time was 21.6 + 2.3 days (range 11—37) once B. americana larvae had begun

to consume Ceratina immatures. Emergence was highly synchronized for non-diapausing

larvae, with all newly eclosed adults emerging from the host within 24 hours.

Baryscapus americana was the most common parasitoid species observed,

infecting 8% (66/850) of all cells, and 16% (17/107) of all nests. This parasitoid was

found in nests collected from 14 July through 1 August 2008. They were most often found

parasitizing nests in teasel, with low levels of infection in raspberry, and none in sumac

(Tables 3, 4). On average they infected 39% of available brood in an affected nest, ranging

from one immature to the entire nest. This parasitoid predominantly affected the prepupal

stage (8/9 C. calcarata and 53/56 C. dupla) and occasionally white eyed pupae. Individuals

of Baryscapus sp. 1 were often found in nests with other associated species (7/17, 41%),

including Eurytoma sp. near apiculae, Axima zabriskiei, Eupelmus vesicularis and Pyemotes

sp.

Vickruck et al. JESO Volume 141, 2010

FIGURE 2. Development of Baryscapus americana a) Parasitoid larvae consume the

contents of the Ceratina immature, leaving the larval skin intact. b) Full grown larvae move

to the anterior and posterior ends of the host. c) Thereafter, pupation and development

continue to eclosion or individuals overwinter as prepupae.

Baryscapus sp. (Eulophidae)

This parasitoid which is morphologically very similar to B. americana, but

mummifies its host, overwintered as prepupae in the larval or pupal skin of Ceratina

calcarata only. All individuals of this gregarious, koinobiont endoparasitoid that emerged

as adults were male. It infected 20% (64/327) of the total C. calcarata cells available and

17% (18/48) of all C. calcarata nests. It was found most commonly in raspberry (51 of

64 cells), occasionally in sumac (13 of 64 cells), and never in teasel. On average 3.8 + 0.6

cells per affected nest were parasitized, representing 51% of infected C. calcarata nests on

average. Other associated species were present in 8 of the 17 infected nests (47%); these

were always Pyemotes or Axima. Prepupae were the most commonly affected host stage

(43/64), but white-eyed pupae (21/64) were also susceptible to parasitism.

Parasitism went unnoticed until these internal parasitoids began to consume the

host. Infection became evident when the larval skin of the C. calcarata changed dramatically

in colour and consistency. The larval skin of living Ceratina is somewhat transparent and

the gut is often visible. Parasitism caused the larval skin of the Ceratina to become a rusty

red-brown colour; it also became much more brittle with the consistency of paper mache.

The parasitoids overwintered as full grown larvae in the host, and the tough pupal casing

of the larval or pupal skin may provide protection to the diapausing larvae (Legrand et al.

2004). Only males of this species emerged as adults from Ceratina immatures, in constrast

with Baryscapus americana where both sexes emerged. These parasites were collected

from Ceratina nests from 22 July through 21 August 2008.

Coelopencyrtus sp. (Hymenoptera: Encyrtidae)

A single C. calcarata larva in a sumac nest was affected by this gregarious,

Natural enemies of Ceratina in the Niagara Region JESO Volume 141, 2010

endoparasitic koinobiont. The only other observation of C. calcarata being attacked

by Coelopencyrtus Timberlake is by R.W. Matthews (reported by Daly et al. 1967),

who reported C. hylaei Burks parasitism on six consecutive cells in a nest collected in

Connecticut. Coelopencyrtus have also been reported to parasitize members of the twig-

nesting, bee genus Hylaeus Fabricius (Burks 1958).

The C. calcarata nest was collected on 7 July 2008 and parasitism became evident

on 10 July 2008 when more than 20 Coelopencyrtus larvae could be seen consuming the

bee larva, which was in the second innermost cell in a nest with six other immatures. Once

the entire contents of the Ceratina larva had been consumed, development of the parasitoids

continued inside the transparent larval skin. Eyes of the parasitoids began to darken on 4

August with their exoskeletons gaining pigmentation by 7 August. Synchronized emergence

took place on 13 August, when all of the new Coelopencyrtus adults emerged, except for

one individual that had died during development.

Eupelmus vesicularis Retzius (Hymenoptera: Eupelmidae)

One Eupelmus vesicularis specimen was reared from a C. dupla nest in teasel.

While this is the first host record of E. vesicularis parasitizing C. dupla, members of the

genus Eupelmus are well known for parasitizing a large number of different hosts (Burks

1979a; Gibson 1990). Eupelmus vesicularis has a Holarctic distribution, but may have been

introduced to North America from Europe in straw (Burks 1979a). Its first record in North

America was from Pennsylvania in 1915 (Burks 1979a).

Usually a primary parasitoid, E. vesicularis has been occasionally reported as a

secondary parasitoid (Burks 1979a). The wasp collected here had actually parasitized a

white-eyed bee pupa that had also been parasitized by Baryscapus americana. The E.

vesicularis egg had already been laid when the nest was collected on 15 July 2008. The

parasitoid hatched and began feeding externally on the bee larva on 20 July 2008. A day later

it became apparent that the bee larva had also been parasitized internally by B. americana.

Eupelmus vesicularis consumed the bee larva, followed by the B. americana parasitoids,

and pupated on | August. Body sclerotization was quite rapid, beginning 4 August and

finishing 2 days later. The adult E. vesicularis emerged on 8 August 2008, 19 days after

first hatching. This is the first record of the family Eupelmidae associated with a parasitoid

developing in bees.

Eurytoma sp. near apiculae Bugbee (Hymenoptera: Eurytomidae)

This is the first record of Eurytoma Illiger parasitizing C. calcarata. Eurytoma

apiculae and E. nodularis Boheman have been reported as parasitoids on C. callosa

Fabricius, C. dallatoreana Friese, C. nanua Cockerell, and C. punctigena Cockerell (Bugbee

1966; Burks 1979b; Daly 1966), and an unknown Eurytoma species has been observed as a

parasitoid of C. australensis Perkins in Queensland, Australia (S. Rehan, pers. comm.).

An external parasitoid of C. calcarata, only one E. sp. near apiculae individual

was collected which was parasitizing a larva that had almost finished eating its pollen ball

in a nest constructed in teasel. The E. sp. near apiculae egg was laid in the innermost brood

cell and by 16 July, 2008, had begun to feed on the host Ceratina larva. Over the course of

the next week the parasitoid finished consuming the host, after which it defecated and then

Vickruck et al. JESO Volume 141, 2010

pupated. The eyes of the E. sp. near apiculae began to darken on 27 July and the integument

was fully pigmented by | August. The teneral adult emerged on 3 August, 2008.

Axima zabriskiei Howard (Eurytomidae)

Axima zabriskiei has been reported as a parasitoid of both C. dupla and C.

calcarata (Kislow 1976; Krombein 1960; Rau 1928). An ectoparasitic idiobiont, 1—7

Axima individuals could be seen consuming a single Ceratina immature, always a pre-pupa

or white-eyed pupa, most often attached between the head and thorax and/or near the wing

buds of white eyed pupae (Fig 3b). The parasitoids consumed the hosts’ contents rapidly

(usually in 24 — 48 hours), leaving the skin intact. It was at this point that most lab-reared

parasitoids died, but two did pupate in the laboratory in 2008 (Fig. 3c). None of these

chalcid parasitoids were successfully reared to adulthood in the lab in 2008 but one was

reared to adulthood during 2009 collections.

Axima zabriskiei parasitoids infected 3% (24/850) of all available cells and 11%

(12/107) of available nests. Twenty-two of the infected cells were found in raspberry (11

nests) and two cells were in sumac (one nest), for an average of 1.9 + 0.3 cells per infected

nest, with a maximum of four infected Ceratina immatures but never representing more

than 50% of the total brood in a nest. Axima zabriskiei was found with other parasites

FIGURE 3. Axima zabriskiei wasp development. a) Newly hatched larvae pierce the soft

exoskeleton of the pupa and rapidly ingest the contents (a, circle), usually within 24-48

hours. Often multiple parasitoids will attack a single Ceratina immature (b). Once finished

feeding larvae pupate (c) before emerging as an adults (d).

Natural enemies of Ceratina in the Niagara Region JESO Volume 141, 2010

in 7/12 (58%) affected nests, most often in conjunction with Baryscapus sp.. The first

Ceratina nest containing Axima zabriskiei was collected on 14 July, with the last parasitized

nest collected on 21 August 2008.

Parasites (other than parasitoids)

Pyemotes sp. (Acari: Actinedida: Pyemotidae)

Pyemotes sp. were the most common natural enemy found on Ceratina immatures,

infecting 10% (86/850) of all available brood cells and 21% (23/107) of all available nests.

This is the first record of Pyemotes mites infecting C. dupla and C. calcarata, although

they have been reported on C. dallatorreana (Friese) in California (Daly 1966). They were

more common in teasel nests (66 of 86 infected brood) than in raspberry (20 of 86 infected

brood), and were not found in sumac (Table 3). On average Pyemotes sp. affected 3.7 + 0.7

immatures per nest, representing 28% of the total brood in affected nests. Pyemotes sp. was

present in nest collections from 25 June to 25 July 2008.

This external parasite was found to infect all immature stages, from small larvae

to fully pigmented pupae. Multiple individuals often infected a single larva or pupa, but a

single mite was effective in paralyzing and killing the host. Pyemotes seemed to monopolize

parasitism in a nest, being found with other parasitoids only 22% of the time (5/23 nests).

Pyemotes mites were also observed feeding on two A. zabriskiei larvae which subsequently

died. Other members of the genus Pyemotes have been known to decimate nests of the

bee Melipona colimana Ayala and the stem-nesting wasp, Psenulus interstitalis Cameron

(Macias-Macias and Otero-Colina 2004; Matthews 2000).

Discussion

Many parasitoids were more prevalent in one substrate than in another. Baryscapus

americana for example, was collected significantly more often from teasel nests, with only

two cells parasitized in raspberry (Table 3). While not statistically significant, Axima

zabriskiei was collected more in raspberry than sumac (Table 3). Pyemotes mites did not

parasitize one species more than another, but were significantly more common in teasel nests

then they were in raspberry (Tables 3 and 4). While Baryscapus sp. was found parasitizing

64 individuals in 18 nests, it was only ever a parasite of C. calcarata in raspberry and

sumac, never in teasel.

The parasitoid preferences seen for specific host substrates may be due to a number

of factors such as the structure or biology of the host plant itself. The following discussion

pertains mainly to raspberry and teasel, because so few sumac nests were collected. One

possible reason for higher parasitism rates in raspberry than in teasel may be the structure of

the plant species used for Ceratina nests. Teasel nests can only contain one nest per plant, in

the straight stalk that grows perpendicular to the ground. Shrubs like raspberry (and sumac)

have multiple branches in each plant and thus multiple possible nest substrates. These

shrubs also tend to grow in aggregations, with multiple plants in very close proximity to one

another. This can lead to higher nest densities in raspberry than in teasel. As Ceratina dupla

and C. calcarata females guard only their own nests, the high density of nests in shrubs may

Vickruck et al. JESO Volume 141, 2010

lead to increased rates of parasitism, as an individual parasitoid may be able to efficiently

locate and infect several nests in close proximity. When comparing parasitism rates for

a number of non-social hymenopteran species that nest solitarily and in aggregations,

Rosenheim (1990) found that aggregated nests had higher parasitism rates in most cases. —

Higher parasitism rates in raspberry may also relate to the habitat and biology of

the plants in which Ceratina nest. Teasel is an invasive plant found in large open fields,

almost always in full sunlight, while. raspberry and sumac are both native plants located

in shaded wood margins. In other words, Ceratina are nesting in different microclimates,

in substrates with different biology, and with different possible chemical signatures.

Numerous experiments have shown that many parasitoids are attracted to chemical cues

of the flora where their host species are commonly found (Drost et al. 1986; Elzen et al.

1986; Godfray 1994; Vet 1983). If parasitoids use the microclimate and/or chemical cues

emitted by the native substrates, then this might explain why the nests in native shrubs

had higher parasitisation. Members of the genus Eupelmus parasitize a very wide range

of host species (Gibson 1990). Gibson (1990) hypothesized that Eupelmus searched for

hosts in specific microclimates, with the microclimate being of more importance than the

host species. Searching for hosts by their preferred substrate may also be more effective

in temperate regions due to the relatively short and synchronized phenology of foraging

insects and nest substrates (Wcislo 1987).

Acknowledgements

The authors wish to thank Dr. Gary Gibson and Dr. Andrew Bennett at the Canadian

National Collection of Insects and Arachnids, and Dr. Michael Gates at the United States

National Museum for identification and confirmation of natural enemies. The comments of

Dr. Jeff Skevington as well as three anonymous reviewers also greatly helped to improve the

manuscript. This research was funded by a Brock University Deans Graduate Fellowship to

JLV and an NSERC discovery grant to MHR.

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Biodiversity in burned and unburned alvar woodland JESO Volume 141, 2010

ORTHOPTERANS (ORTHOPTERA), GROUND BEETLES

(COLEOPTERA: CARABIDAE), AND SPIDERS (ARANEAE)

IN BURNED AND UNBURNED ALVAR WOODLANDS — THE

IMPORTANCE OF POSTFIRE SUCCESSION TO INSECT

DIVERSITY

P. M. CATLING', H. GOULET’, R. BENNETT? AND B. KOSTIUK?

Agriculture and Agri-Food Canada, Environmental Health, Biodiversity,

Wm. Saunders Building, Central Experimental Farm, Ottawa, ON, Canada K1A 0C6

email: paul.catling@agr.gc.ca

Abstract J. ent. Soc Ont. 141: 27-37

To contribute to understanding the importance of successional habitat to insect

diversity and assist biodiversity management in globally imperilled alvar

ecosystems, we surveyed three groups of arthropods in an Ottawa Valley,

Ontario, alvar landscape. Using pitfall traps and sweeping, we compared

grasshopper (Orthoptera), ground beetle (Coleoptera: Carabidae), and

spider (Araneae) diversity in two sites on the same successional gradient: an

unburned climax alvar woodland and a corresponding burned woodland that

had developed into alvar shrubland nine years after fire. Between-site species

similarity was 47.4% for orthopterans (9 species), 6.9% for ground beetles (2

species), and 40.9% for spiders (10 species). Both sites included regionally

rare orthopterans and ground beetles. For all three groups species richness and

density was higher on the burned site. The value of Brillouin’s biodiversity

index was higher for both orthopterans and ground beetles in the burned site

but higher for spiders in the unburned alvar woodland. These results provide

evidence for: (1) the importance of successional habitat to insect diversity, (2)

the value of alvar shrublands to overall alvar landscape biodiversity, and (3)

the potential value of fire in maintaining alvar biodiversity.

Published November 2010

‘Author to whom all correspondence should be addressed

?Agriculture and Agri-Food Canada, Environmental Health, Biodiversity, K.W. Neatby

Building., Central Experimental Farm, Ottawa, ON, Canada K1A 0C6

email: henri.goulet@agr.gc.ca, brenda.kostiuk@agr.ge.ca

British Columbia Ministry of Forests, Saanichton, BC, email: rbennett@esc-sec.ca

Catling et al. JESO Volume 141, 2010

Introduction

Fire was a major ecological factor in much of southeastern Canada during pre-

settlement times (Day 1953; Wein & MacLaren 1983), maintaining a variety of seral stages

across the landscape. In the absence of fire, much of the landscape is now dominated by

woodlands. Since biodiversity is considered to be higher during middle and later stages

of vegetational succession in general (Bormann and Likens 1979; Brown 1984) and

specifically for insects (Brown 1984; Strong et al. 1984; Kayna & Giirkan 2007), the loss

of middle and late stages may be contributing to the decline in insect diversity. We chose

an imperilled alvar ecosystem to investigate this hypothesis and to improve understanding

of the importance of successional habitat to insect diversity. Specifically, the objective of

our work was to clarify the importance of fire and subsequent succession to insect diversity

through a comparative study. This involved three groups of arthropods, orthopterans

(Orthoptera), ground beetles (Coleoptera: Carabidae), and spiders (Araneae) in two different

habitats representing different temporal positions on the same successional gradient: an

early successional alvar shrubland (“burned woodland”, nine years post-fire) and a nearby

climax alvar woodland (“unburned woodland”).

Materials and Methods

The study area

The Burnt Lands, approximately 40 km SW of Ottawa in the Ottawa Valley of

eastern Ontario, is one of the richest (in terms of species number and rare species) and

most extensive alvar landscapes in the Great Lakes region. Alvars are globally imperilled

ecosystems with a fragmented distribution in North America (Catling and Brownell 1995,

1999; Reschke et al. 1999; Brownell and Riley 2000). Evidence of past fire is common

in alvars (Catling and Brownell 1998; Jones and Reschke 2005) and, considering the high

plant diversity in successional alvar habitats, large scale biomass removal may significantly

contribute to biodiversity protection (e.g. Catling et al. 2001, 2002; Catling and Sinclair

2002). The insect fauna of alvars is distinctive and significant (Bouchard 1997a, b, 1998;

Bouchard et al. 1998, 2001, 2005), making information on the importance of alvar succession

to insects particularly relevant.

The climax woodland study site included four hectares centered at 45.2569,

—76.1437. The burned woodland study site, also 4 hectares in size, is centered at 45.2507,

—76.1437, 0.5 km southeast of the climax site. Both sites are located in Burnt Lands

Provincial Park, Lanark County, Ontario, and are part of the Burnt Lands Alvar landscape

(Brunton 1986). Based on personal observations (annually 1985 to 1995) and examination

of aerial photographs obtained from the National Air Photo Library of Natural Resources

Canada (e.g., number A 16525-105, 29 May 1959), both of these study sites were semi-

open, mixed boreal forest until 23 June 1999 when a fire burned 152 hectares, including

the burned woodlands study site. The forest (Figure 1a) was dominated by Abies balsamea

(L.) P. Mill., Picea glauca (Moench) Voss, Pinus strobus L., Thuja occidentalis L., and

Populus tremuloides Michx. with an understory of mosses including Pleurozium schreberi

(Brid.) Mitt., and Dicranum polysetum Sw., and occasional depauperate shrubs including

Biodiversity in burned and unburned alvar woodland JESO Volume 141, 2010

FIGURE 1. Burned (a) and unburned (b) alvar woodland on the Burnt Land Alvar west of

Ottawa. a)Fallen and dead standing trees are Abies balsamea, Picea glauca, Pinus strobus,

and Thuja occidentalis, regrowth on upper left is Populus tremuloides and Arctostaphylos

uva-ursi can be seen flowering in the foreground. b)The forest is dominated by Abies

balsamea, Picea glauca, Pinus banksiana, Pinus strobus, Populus tremuloides and Thuja

occidentalis. Shrubs of Juniperus communis are present in the foreground. (a) Area burned

on 23 June 1999 was taken at 45.2507, —76.1437. The right photo was taken at 45.2569,

—76.1437 in late May. Both photograps by P. M. Catling.

he = Nee .

ayo. - te \ a.)

“\ . ; x a

~ Be r - xX o

Lad — ~~ >

— ’ a _§

> _ = “M4 : a =

> Ue Qoaime 3 F- 3 } Fai

> <4 . e

; { 24

= -

Juniperus communis L. var. depressa Pursh. Nine years after the fire, the burned area had

developed into a species-rich, open grassy shrubland dominated by graminoid plants such

as Danthonia spicata (L.) Beauv. ex Roemer & J. A. Schultes and Carex richardsonii R.

Br., herbs such as Packera (Senecio) paupercula (Michx.) A. & D. Love, and Solidago

nemoralis Ait. var. nemoralis, and shrubs such as Amelanchier alnifolia (Nutt.) Nutt. ex M.

Roemer var. compacta (Nielsen) McKay and Prunus virginiana L. Additional information

on the vegetation of the two study sites is in Catling (in press).

The Burnt Lands area is subject to fire because of its shallow soils and location on

an elevated plateau of porous limestone rock. It was named by settlers in 1870 following

an extensive fire. Additional background information on the Burnt Lands Alvar is in White

(1979), Brunton (1986), and Catling et al. (2001, 2002).

Collection and identification of insects

(1) Traps: Ten pitfall traps (15 cm x 10 cm x 5 cm deep) were buried 10 m apart

in east-west transects in each of the burned and unburned alvar woodlands. The traps

were buried so that the tops were flush with the ground surface and there were no nearby

obstructions. Each trap was filled with antifreeze and a drop of soap to half depth. Five days

Catling et al. | JESO Volume 141, 2010

after setting, the traps were checked. The survey continued from spring through summer to

fall with several gaps. The dates of checking the traps were 16 May; 1, 9 June; 1, 5, 12 July;

16, 21, 26 August; and 8, 13 September 2008. These traps, set out and checked by PMC

and BK, provided the entire basis for a comparison of Coleoptera and Araneae and most of

the information on Orthoptera.

(2) Sweeping: To additionally sample Orthoptera, at each site on each visit, 15

minutes was spent sweeping vegetation less than 1.5 m tall. Sweeping was always done on

vegetation selected randomly along the same 100 m transect.

Orthopterans were identified by PMC using Vickery and Kevan (1985). Ground

beetles were identified by HG (1961, 1963, 1966, 1968, 1969a, 1969b), Bousquet and

Larochelle (1993), and Goulet & Bousquet (2004). Spiders were identified by RB using

primary taxonomic literature and available regional keys (e.g., Dondale and Redner 1978,

1982, 1990; Paquin and Dupérré 2003; Platnick and Dondale 1992). Ninety-one juvenile

spiders and 52 juvenile orthopterans, unidentifiable to species, were excluded from the

analysis. Voucher specimens were deposited in the Canadian National Collection of Insects

at Agriculture and Agri-Food Canada (CNCI) in Ottawa.

Comparing biodiversity

Biodiversity was compared with respect to: (1) total number of species and

numbers of individuals within’species, (2) the presence of regionally rare species (confined

to alvars or known from less than four locations in the Ottawa valley), (3) the extent of

distinctive composition, and (4) by applying Brillouin’s Index which includes consideration

of heterogeneity (species richness and evenness), is relatively sensitive to the abundance of

rare species, and assumes a finite sample and collection of that sample without replacement

of individuals (Krebs 1999, 2008). Replacement in this case is expected to have been the

same in both habitats, would not likely have been from substantial distances (1.e., over 100

m) for these ground dwelling species, and would not have involved second generations

since most of these species have a single generation in a year.

Results

Overall, 19 species of orthopterans, 29 species of ground beetles, and 42 species of

spiders were recorded (Tables 1—3). Species composition differed between the burned and

unburned sites. The sites shared 9, 2, and 10 grasshopper, ground beetle, and spider species,

respectively (Tables 1—3), or 47.4 %, 6.9 %, and 40.9 % of all collected grasshopper, ground

beetle, and spider species. For species in common, the number of individuals was generally

highest in the burned woodland for orthopterans and ground beetles but not for spiders. At

each site certain unique species were common, e.g., the grasshopper Jefrix ornata (Say),

the spider Schizocosa avida Walckenaer in the burned site, and the ground beetle Synuchus

impunctatus (Say) in the unburned site.

Four regionally rare grasshopper species were present at each site (Table 1).

Of these, Melanoplus keeleri luridus (Dodge), Spharagemon bolli bolli Scudder, and

Encoptolophus sordidus (Burmeister) were found only in the burned site and Melanoplus

Biodiversity in burned and unburned alvar woodland JESO Volume 141, 2010

TABLE 1. Species and number of individuals of orthopterans (Orthoptera) recorded in

burned and unburned woodland on the Burnt Lands. Species marked with an asterisk (*)

are regionally rare.

Species Burned Unburned

Acrididae

Chloealtis conspersa Harris*

Chortophaga viridifasciata (De Geer)

Encoptolophus sordidus (Burmeister) *

Melanoplus bivitattus (Say)

M. dawsoni (Scudder) *

M. fasciatus (F. Walker)*

M. keeleri luridus (Dodge)*

M. punctulatus (Scudder) *

M. sanguinipes (Fabricius)

Spharagemon bolli bolli Scudder*

Gryllidae

Allonemobius fasciatus (De Geer) 19 2

Gryllus pennsylvanicus Burmeister 32 8

G. veletis (Alexander & Bigelow) 22 6

Oecanthidae

Oecanthus quadripunctatus (Beutenmiiller) 3 2

Phaneropteridae

Scudderia curvicauda (De Geer) | 1

S. furcata furcata Brunner von Wattenwy1 l |

Rhaphidophoridae

Ceuthophilus cf. maculata (Harris) - 7

Tetrigidae

Nomotettix cristatus cristatus (Scudder) 3 ]

Tetrix ornata ornata (Say) 23 -

Totals 178 44

—

eS)

ONWi Wes WOw ,

—

dawsoni (Scudder), M. punctulatus (Scudder), and Chloealtis conspersa Harris only in the

unburned site. Similarly, of the 6 regionally rare ground beetle species collected during the

study, Harpalus indigens Casey, Selenophorus gagatinus Dejean, S. opalinus (LeConte),

and the southern Calathus opaculus LeConte were found only in the burned woodland,

while the boreal species Harpalus fulvilabris Mannerheim and the alvar-restricted species

Pterostichus novus Straneo were found only in the unburned woodland. All collected

spiders were members of wide-ranging and relatively common species.

Species richness was greater in the burned woodland for all three groups. With

respect to orthopterans, the burned woodland had 15 species and unburned woodland had

13 species (Table 1, Figure 2). For spiders, the difference was greater with 31 species

in the burned compared to 22 species in the unburned site (Table 3, Figure 2). Ground

beetles provided the greatest contrast in species richness, with 21 species in the burned area

compared to 9 species in the unburned area (Table 2, Figure 2).

In all three groups there were more individuals in the burned area (Table 3). This

was most pronounced in the case of orthopterans with 3.55 times as many in the burned

woodland, and least for spiders with 1.69 times as many in the burned woodland.

Catling et al. | JESO Volume 141, 2010

TABLE 2. Number of various species of ground beetles (Coleoptera: Carabidae) in 10 pitfall

traps in each of a burned and unburned alvar woodland. Species marked with an asterisk (*)

are regionally rare.

Species Burned Unburned

Agonum cupripenne (Say)

Amara pennsylvanica Hayward

Anisodactylus rusticus (Say)

Calathus gregarius (Say)

C. opaculus LeConte*

Calosoma calidum Fabricius

Carabus nemoralis O.F. Miller 3

Chlaenius emarginatus Say

Cicindela punctulata Olivier

C. purpurea Olivier

C. sexguttata Fabricius

Diplocheila obtusa (LeConte)

Harpalus faunus Say 3

H. fulvilabris Mannerheim*

H. indigens Casey*

H. laevipes Zetterstedt

H. laticeps LeConte °

H. opacipennis (Haldeman)

H. pensylvanicus (De Geer)

Notiophilus aeneus (Herbst)

Poecilus lucublandus (Say)

Pterostichus novus Straneo*

P. mutus (Say)

P. pensylvanicus LeConte

Selenophorus gagatinus Dejean*

S. opalinus (LeConte)*

Sphaeroderus stenostomus lecontei Dejean

Synuchus impunctatus (Say) ‘ 34

Totals 124 66

The value of Brillouin’s biodiversity index was higher for both orthopterans and

ground beetles in the burned site but higher for spiders in the unburned woodland (Figure

2).

ORF OR W, BRR WRK MRS MN NUN NY

1 ns —OO— 1

i — i —s

Discussion

In some cases the much greater presence of a species in either habitat is to be

expected on the basis of known ecological associations of a general or specific nature. For

example, the orthopterans found only in woodland (Chloealtis conspersa and Melanoplus

punctulatus) are known to require wood for oviposition, and the latter species is found

mostly on tree trunks (Vickery and Kevan 1985). The ground beetles present are mostly

normal to one site or the other (Lindroth 1961). The large number of Carabus nemoralis

in the open areas may be a result of greater abundance of earthworms in the more mineral

substrates of the open area as compared to the more acidic needle litter of the woodland.

Harpalus faunus, which collects small seeds, may be more abundant in the burned area as a

Biodiversity in burned and unburned alvar woodland JESO Volume 141, 2010

TABLE 3. Numbers of various species of spiders (Araneae) in 10 pitfall traps in each of a

burned and unburned alvar woodland.

Species Burned Unburned

Agelenidae

Agelenopsis potteri (Blackwall) ) 6

Agelenopsis utahana (Chamberlin & Ivie) l -

Clubionidae

Clubiona mixta Emerton 1 -

Elaver excepta (L. Koch) - Zz

Corinnidae

Castianeira longipalpa (Hentz) 3 -

Gnaphosidae

Callilepis pluto Banks l

Drassodes neglectus (Keyserling)

Drassylus depressus (Emerton) ]

D. niger (Banks) -

D. socius Chamberlin -

Haplodrassus bicornis (Emerton) -

H. signifer (C.L. Koch) 8

Herpyllus ecclesiasticus Hentz -

Gnaphosa muscorum (L. Koch) 24

Micaria laticeps Emerton 1

—

oS)

Z. fratris Chamberlin

Z. hentzi Barrows

Hahniidae

Neoantistea magna (Keyserling)

Liocranidae

Agroeca ornata Banks - 4

A. pratensis Emerton 1 6

Lycosidae

Alopecosa aculeata (Clerck) 2

Hogna frondicola (Emerton) -

P. distincta (Blackwall) 9 16

P. moesta Banks -

Schizocosa avida (Walckenaer) 76 -

S. crassipalpata Roewer 10 -

S. saltatrix (Hentz) 14 6

Trochosa ruricola (De Geer) 3 -

T. terricola Thorell 3 23

Oxyopidae

Oxyopes scalaris Hentz - ]

Philodromidae

Thanatus formicinus (Clerck) z l

Salticidae

Evarcha hoyi (Peckham & Peckham)

Habronattus viridipes (Hentz)

Phidippus purpuratus Keyserling

Thomisidae

Xysticus alboniger Turnbull et al.

X. ampullatus Turnbull et al.

X. canadensis Gertsch

X. elegans Keyserling

X. luctuosus (Blackwall)

X. pellax O. P.-Cambridge

X. punctatus Keyserling

X. triguttatus Keyserling -

Totals Zoe 140

yar a ee,

— ae

SR i NR it NNR

NCO 1

—

NOR ne WHOM

Catling et al. JESO Volume 141, 2010

Fal Unburned Woodland

(EG Burned Woodland

te)

Number of Species

Number of Individuals

Brillouin’s Index Value

& o

& S

‘ $

O O

FIGURE 2. Number of species, number of individuals, and the value of Brillouin’s Index

for three arthropod groups in burned and unburned alvar woodland.

result of greater numbers of herbs with small seeds in that area. The abundance of Synuchus

impunctatus in the unburned woodland and its absence from the burned area is anticipated

on the basis of its association with deep litter (Lindroth 1961), which was removed by fire

from the burn. Likewise with respect to the spiders, some of the major differences were

anticipated based on known species and habitat associations, such as the relationship of

many Schizocosa and Xysticus species with dry, open habitats.

Our results suggest that the relatively long lasting successional shrubland that

follows burning of alvar woodland differs from the original woodland in species composition,

higher species richness and density, and higher biodiversity index values for some arthropod

groups. The fact that the Brilluoin Index has a slightly lower value for spiders in the burned

area than in the unburned is a consequence of less evenness in the number of individuals

of different species in that site, despite higher diversity and species number. Other studies

similarly found highest insect diversity in middle successional stages (e.g., Brown 1984;

Kayna & Giirkan 2007) and qualitative differences between burned and unburned sites

(e.g., Burger et al. 2005). The importance of succession and differing species compositions

in different seral stages is not surprising considering that opposite ends of the sere are

characterized by organisms with different life history strategies (Brown 1984).

Biodiversity in burned and unburned alvar woodland JESO Volume 141, 2010

The tendency for insect diversity to track plant diversity (e.g. Knops et al. 1999)

may help to explain the generally higher insect diversity in the burned woodland, which had

higher vascular plant diversity than the unburned alvar woodland (Catling in press). With

regard to herbivorous insect diversity, which is tracked by predator and parasite diversity

(e.g., Knops et al. 1999), other explanatory factors associated with high plant diversity may

include increased insect richness per plant host and higher average plant host specificity

(Lewinsohn et al. 2005), as well as increased structural diversity of plant hosts (Southwood

et al. 1979).

The evidence presented here for the importance of succession to ground-dwelling

insects on alvars is based on a single location and a rather limited sampling procedure.

However, the relationship seems to hold for other alvar sites in the Ottawa valley based

on general comparative surveys (Catling, unpublished data). Although management of

alvar vegetation with succession-initiating fire seems appropriate, it may also have negative

impacts (Siemann et al. 1996), especially if it does not allow survival of some species in

unburned patches that serve as refugia. Fire or any form of biomass removal should be

part of a broad, long-term landscape management plan that takes many species and species

groups into account.

Acknowledgements

The CanaColl Foundation (http://www.canacoll.org) provided financial support

for identification and processing of the spider collection. Useful reviews were provided by

P. Bouchard and J. Miskelly.

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Bombus of southern Ontario JESO Volume 141, 2010

THE BUMBLE BEES OF SOUTHERN ONTARIO: NOTES ON

NATURAL HISTORY AND DISTRIBUTION

S. R. COLLA' AND S. DUMESH?

Department of Biology, York University,

4700 Keele Street, Toronto, ON, Canada M3J 1P3

email: scolla@yorku.ca

Abstract J. ent. Soc. Ont. 141: 39-68

Although North American bumble bees are common and easily recognizable

insects in many habitats, details of their natural history are poorly known.

This study presents basic ecological information based on a literature review,

databased insect collections and recent survey work performed throughout

southern Ontario. As a result of this review, phenology, food plants,

distribution and habitat associations are summarized for each species of this

important group of pollinators.

Published November 2010

Introduction

The plight of native pollinators has recently gained the interest of scientists and

naturalists alike. Globally, bees have been documented as declining throughout their native

ranges (Berenbaum et al. 2007; Biesmeijer et al. 2006). Conservation plans are highly

dependent upon basic natural information, which is lacking for almost all North American

bee species. Southern Ontario is an important region to consider as it has relatively high

biodiversity and many potential threats to these native organisms (Allen et al. 1990). Habitat

loss and fragmentation due to high population density and agricultural production are major

threats to wild pollinators in this region (eg. Taki et al. 2007; Findlay and Houlahan 1997).

Understanding basic natural history aspects of native bees in southern Ontario is important

to conserve these pollinators and their associated native flowering plants.

Bumble bees are eusocial organisms with an annual colony cycle. Queens emerge

in the spring, find nest sites, build honey pots and lay eggs. Hatched workers then take over

foraging and nest tending duties, while the queen focuses on reproduction. Towards the end

of the colony cycle, males and new queens are produced. These reproductive individuals

leave the nest and mate. Newly mated queens overwinter in rotting logs, mulch or dirt

while the rest of the colony perishes with the onset of cold weather. Cuckoo bumble bees

(subgenus Psithyrus) are exceptions, as they do not produce a worker caste, but usurp the

colonies of other species and the host workers collect resources for the cuckoo’s offspring

‘Author to whom all correspondence should be addressed

7 email: sdumesh@yorku.ca

Colla and Dumesh | JESO Volume 141, 2010

instead. The timing of queen emergence, worker production and colony decline for each

species varies with latitude, elevation and weather variation from year to year (Benton 2006).

Here, using historical and'recent records of bumble bees in southern Ontario, information is

provided on distribution, phenology, habitat and forage for each of the 18 species found in

this region.

Materials and Methods

Phenology graphs were produced using bumble bee records from collections and

recent surveys. Bumble bee specimens from southern Ontario insect collections (University

of Guelph, Royal Ontario Museum, Canadian Museum of Nature, PCYU York University,

Algonquin Provincial Park) were databased with species identifications being determined

or verified by SR Colla. Surveys were made throughout southern Ontario during the

summers of 2005-2009 (Colla and Packer 2008; Colla.et al. 2006; Colla, unpublished).

Bees were collected using hand nets and were either identified and released or collected

for identification. Plant records were accumulated from specimen labels, surveys, the

literature (Macfarlane 1974; Robertson 1929). In Robertson (1929), plant records were not

determined in southern Ontario but were included if the taxon occurs naturally in the region.

Habitat notes were collected from Macfarlane (1974) and field surveys.

Results

A total of 9052 bumble bee records representing 18 species spanning the years

1876-2009 were accumulated from insect collections (n=6506) and field surveys (n=2546).

Laverty and Harder (1988) note the presence of B. frigidus Smith in southern Ontario, but

specimens for this species were not found during this study. Ecological information is

presented below for each species. Introduced plant species are marked with an asterisk (*).

Figures presenting the phenology and collection locations for each species are presented at

the end of the paper.

Bombus impatiens Cresson: the common eastern bumble bee

Phenology and Distribution: This species exhibits early spring emergence (earliest

record April 23) with a long colony cycle extending into autumn and has a widespread

distribution (Fig. 1).

Floral Records: Robertson (1929): Actinomeris, Agastache, Amphicarpaea, Arabis,

Asclepias, Blephilia, Camassia, Campanula, Caulophyllum, Cercis, Cicuta, Claytonia,

Clematis, Collinsia, Coreopsis, Crataegus, Delphinium, Dentaria, Desmodium,

Dasistoma, Dicentra, Fragaria, Gaura, Geranium, Gerardia, Gleditzia, Gymnocladus,

Helenium, Helianthus, Helianthus divaricatus, Heliopsis, Hydrophyllum, Impatiens,

Lactuca, Liatris, Lithospermum, Lobelia, Lonicera, Lycopus, Lythrum, Monarda,

Nelumbo, Osmorhiza, Penstemon, Petalostemum, Phlox, Podophyllum, Polemonium,

Prenanthes, Prunella, Prunus, Pycnanthemum, Pyrus, Ribes, Rudbeckia, Salix,

Bombus of southern Ontario JESO Volume 141, 2010

Scrophularia, Scutellaria, Sicyos, Silphium, Sium, Smilax, Stachys, Staphylea,

Symphoricarpos, Symphyotrichum, Teucrium, Tradescantia, Verbena, Viola, Zizia.

Macfarlane (1974): Acer ginnala*, Aesculus hippocataneum*, Althaea rosea*, Antirrhinum

majus*, Arctium minus*, Berberis thunbergii*, Caragana arborescens*, Carduus nutans *,

Cotoneaster adpressa*, Daucus carota*, Dipsacus sylvestris*, Echium vulgare*, Epilodium

angustifolium, Euthamia graminifolia, Helianthus annuus*, Hesperis matronalis*,

Hydrophyllum virginianum, Hypericum perforatum*, Impatiens capensis, Kalmia latifolia,

Kolkwitzia amabilis*, Ligustrum vulgare*, Lonicera caerulea, Lonicera periclymenum*,

Lonicera tatarica*, Medicago sativa*, Melilotus alba*, Nepeta cataria*, Oenthera biennis,

Phaseolus coccineus*, Philadelphus coronarius*, Potentilla, Prunus cerasus*, Prunus

tomentosa*, Pyrus malus*, Rhododendron, Ribes grossularia*, Ribes nigrum*, Ribes

rubrum*, Robinia fertilis*, Rubus, Salix, Salvia sylvestris*, Silene vulgaris*, Solanum

dulcamara*, Solidago canadensis, Solidago flexicaulis, Solidago rugosa, Stachys palustris *,

Symphyotrichum (Aster) ericoides, Symphyotrichum (Aster) lateriflorum, Symphyotrichum

novae-angliae, Symphytum officinale*, Syringa vulgaris*, Taraxacum officinale*, Tilia

platyphyllos*, Trifolium pratense*, Trifolium repens*, Vicia cracca*, Weigelia florida*.

Field and Museum Records: Amelanchier alnifolia, Ceanothus americanus, Centaurea

macrocephala, Cephalanthus, Cephalanthus occidentalis, Chelone glabra, Clinopodium

vulgare, Echinacea, Erythronium, Eupatorium fistulosum, Eupatorium maculatum, Justicia,

Lespedeza intermedia, Linum, Lupinus, Mentha, Onopordum acanthium*, Pediomelum,

Pontederia cordata, Prunus virginiana, Rhexia virginica, Rhus, Rubus idaeus, Rubus

occidentalis, Rudbeckia hirta, Solidago altissima, Solidago bicolor, Solidago caesia,

Solidago sempervirens, Spiraea, Spiraea alba, Uvularia, Vaccinium angustifolium,

Vaccinium vacillans.

Habitats: Close to or within wooded areas, open fields, urban parks and gardens,

wetlands.

Bombus bimaculatus Cresson: the two-spotted bumble bee

Phenology and Distribution: This species exhibits early spring emergence (earliest record

April 13) and has a widespread distribution (Fig. 2).

Floral Records: Robertson (1929): Amelanchier, Blephilia, Collinsia, Cephalanthus,

Dentaria, Dicentra, Geranium, Helianthus, Hydrophyllum, Mertensia, Monarda, Phlox,

Polemonium, Ribes, Triosteum, Uvularia, Verbena.

Macfarlane (1974): Ajuga genevensis*, Ajuga reptans*, Aesculus hippocataneum*,

Camassia scilloides, Carduus nutans*, Cotoneaster adpressa*, Cynoglossum officinale*,

Deutzia gracilis*, Dipsacus sylvestris*, Echium vulgare*, Hydrophyllum virginianum,

Hypericum perforatum*, Lamium amplexicaule*, Leonurus cardiaca*, Linaria vulgaris *,

Lonicera caerulea, Lonicera periclymenum*, Lonicera tatarica*, Mahonia aquifolium,

Melilotus alba*, Medicago sativa*, Monarda didyma, Onopordum acanthium*, Prunella

vulgaris*, Prunus tomentosa*, Pyrus malus*, Ribes grossularia*, Ribes nigrum*, Robinia

fertilis*, Salix, Solanum dulcamara*, Solidago canadensis, Solidago flexicaulis, Solidago

graminifolia, Symphytum officinale*, Syringa vulgaris*, Taraxacum officinale*, Trifolium

pratense*, Trifolium repens*, Vicia cracca*, Weigelia florida*.

Field and Museum Records: Arabis, Asclepias, Campanula, Campanula rotundifolia,

Caulophyllum, Ceanothus americanus, Centaureamacrocephala, Cephalanthus occidentalis,

Colla and Dumesh | JESO Volume 141, 2010

Claytonia, Clinopodium vulgare, Dasistoma, Desmodium, Dicentra cucullaria, Echinacea,

Erythronium albidum, Euthamia graminifolia, Gaylussacia, Hypericum prolificum, Kalmia

polifolia, Lonicera, Lotus corniculatus*, Lupinus, Lythrum alatum, Mentha, Pontederia

cordata, Rhexia virginica, Rhus glabra, Rubus odoratus, Spiraea, Spiraea alba, Tilia

americana, Vaccinium angustifolium, Vaccinium corymbosum, Vaccinium myrtilloides,

Vaccinium vacillans, Viola.

Habitats: Close to or within wooded areas, urban parks and gardens.

Bombus terricola Kirby: the yellow-banded bumble bee

Phenology and Distribution: This species exhibits early spring emergence (earliest record

April 24) and has historically a widespread distribution (except extreme southwestern

Ontario) with fewer recent records (Fig. 3).

Floral Records: Macfarlane (1974): Berberis thunbergii*, Caragana arborescens*,

Carduus nutans*, Centaurea jacea*, Cirsium arvense*, Cotoneaster adpressa*, Echium

vulgare*, Hydrophyllum virginianum, Hypericum perforatum*, Impatiens capensis, Lactuca

canadensis, Lonicera caerulea, Lonicera tatarica*, Melilotus alba*, Medicago sativa*,

Philadelphus coronarius*, Prunus cerasus*, Prunus tomentosa*, Pyrus malus*, Rhus

typhina, Ribes grossularia*, Ribes nigrum*, Robinia fertilis*, Salix, Solanum dulcamara’*,

Solidago canadensis, Solidago flexicaulis, Solidago hispida, Solidago juncea, Sonchus

oleraceus*, Sorbus americana, Spiraea, Symphyotrichum (Aster) ericoides, Symphyotrichum

(Aster) lateriflorum, Symphyotrichum novae-anglia, Symphytum officinale*, Syringa

vulgaris*, Taraxacum officinale*, Tilia platyphyllos*, Trifolium pratense*, Trifolium

repens *, Vicia cracca*, Weigelia florida*.

Field and Museum Records: Anaphalis margaritacea, Aquilegia canadensis, Aralia,

Arctostaphylos uva-ursi, Asclepias, Asclepias incarnata, Asclepias syriaca, Astragalus,

Baptisia tinctoria, Diervilla lonicera, Epigaea repens, Erigeron philadelphicus,

Epilodium angustifolium, Eupatorium fistulosum, Eupatorium maculatum, Eurybia

macrophylla, Euthamia graminifolia, Gaylussacia, Heracleum lanatum, Kalmia, Kalmia

augustifolia, Ledum groenlandicum, Linaria vulgaris*, Lupinus, Mertensia, Monarda

fistulosa, Onopordum acanthium*, Pontederia cordata, Prunus, Prunus pensylvanica,

Rhexia virginica, Rhus, Senecio, Solidago, Spiraea latifolia, Thalictrum pubescens, Tilia

americana, Trifolium hybridum*, Urticularia cornuta, Vaccinium angustifolium, Vaccinium

corymbosum.

Habitats: Close to or within wooded areas.

Bombus vagans Smith: the half-black bumble bee

Phenology and Distribution: This species exhibits early spring emergence (earliest record

April 21) with a long colony cycle extending into the fall and has a widespread distribution

(Fig. 4).

Floral Records: Robertson (1929): Blephilia, Cercis, Claytonia, Clematis, Delphinium,

Dicentra, Ellisia, Erigenia, Geranium, Hydrophyllum, Hypericum, Ipomoea, Liatris,

Lobelia, Mertensia, Mimulus, Monarda, Phlox, Physostegia, Polygonatum, Polemonium,

Prunella, Ribes, Scrophularia, Scutellaria, Stachys, Staphylea, Symphoricarpos, Teucrium,

Tradescantia, Triosteum, Verbascum, Verbena, Veronica, Viola, Zizia.

Macfarlane (1974): Aesculus hippocataneum*, Althaea rosea*, Arctium minus*, Berberis

Bombus of southern Ontario JESO Volume 141, 2010

thunbergii*, Carduus nutans*, Centaurea jacea*, Cichorium intybus*, Cirsium arvense*,

Crataegus, Dipsacus sylvestris*, Daucus carota*, Echium vulgare*, Erica cinerea*,

Helianthus annuus*, Hesperis matronalis*, Hydrophyllum_ virginianum, Hypericum

perforatum*, Impatiens capensis, Kolkwitzia amabilis*, Leonurus cardiaca*, Linaria

vulgaris*, Lonicera periclymenum*, Lonicera tatarica*, Lotus corniculatus*, Melilotus

alba*, Medicago sativa*, Nepeta cataria*, Onobrychis viciifolia*, Prunella vulgaris*,

Prunus cerasus*, Pyrus malus*, Ribes nigrum*, Rubus, Silene vulgaris*, Solanum

dulcamara*, Sonchus oleraceus*, Symphyotrichum (Aster) lateriflorum, Symphyotrichum

novae-angliae, Symphytum officinale*, Syringa vulgaris*, Tamarix gallica*, Taraxacum

officinale*, Trifolium pratense*, Trifolium repens*, Verbena hastata, Vicia cracca*,

Weigelia florida*.

Field and Museum Records: Amphicarpaea, Anaphalis margaritacea, Apocynum, Aquilegia

canadensis, Aralia, Aralia hispida, Asclepias incarnata, Asclepias syriaca, Astragalus

canadensis, Chelone glabra, Clinopodium vulgare, Collinsonia canadensis, Cornus,

Dasistoma, Decodon, Desmodium, Diervilla lonicera, Erythronium albidum, Eupatorium

fistulosum, Eupatorium maculatum, Eurybia macrophylla, Euthamia graminifolia, Gerardia

pedicularia, Hypericum prolificum, Lactuca canadensis, Ledum groenlandicum, Lonicera,

Lupinus, Mentha, Monarda fistulosa, Nymphaea odorata, Onopordum acanthium*,

Pontederia cordata, Prunus, Pyrus, Rhexia virginica, Rhododendron, Rubus idaeus, Rubus

odoratus, Sarracenia purpurea, Scutellaria lateriflora, Solidago altissima, Solidago

canadensis, Solidago flexicaulis, Solidago hispida, Solidago juncea, Solidago sempervirens,

Spiraea alba, Tilia americana, Vaccinium angustifolium.

Habitats: Close to or within wooded areas, urban parks and gardens.

Bombus perplexus Cresson: the confusing bumble bee

Phenology and Distribution: This species exhibits early spring emergence (earliest record

April 16) and has a widespread distribution (Fig. 5).

Floral Records: Macfarlane (1974): Althaea rosea*, Asclepias incarnata, Asclepias syriaca,

Berberis thunbergii*, Campanula rapunculoides*, Carduus nutans*, Cirsium arvense*,

Cotoneaster adpressa*, Dipsacus sylvestris*, Echium vulgare*, Helianthus annuus*,

Hesperis matronalis*, Hydrophyllum virginianum, Lonicera caerulea, Lonicera tatarica*,

Onopordum acanthium*, Philadelphus coronarius*, Prunus americana, Prunus cerasus*,

Pyrus malus*, Ribes grossularia*, Ribes nigrum*, Robinia fertilis*, Salix, Solidago,

Solidago altissima, Solidago canadensis, Solidago flexicaulis, Symphytum officinale*,

Taraxacum officinale*, Tilia americana, Tilia platyphyllos*, Trifolium pratense*, Weigelia

florida*.

Field and Museum Records: Aralia, Arctostaphylos uva-ursi, Astragalus canadensis,

Campanula rotundifolia, Cephalanthus, Collinsonia canadensis, Desmodium, Echinacea,

Erythronium albidum, Eupatorium, Eupatorium fistulosum, Eupatorium maculatum,

Euthamia graminifolia, Fragaria, Helianthus, Hypericum perforatum*, Hypericum

prolificum, Kalmia latifolia, Lonicera, Lotus corniculatus*, Lysimachia ciliata, Medicago

sativa*, Mentha, Monarda, Nymphaea odorata, Penstemon, Pontederia cordata, Prunus,

Rhexia virginica, Rubus odoratus, Rudbeckia hirta, Spiranthes lacera, Spiranthes

romanzoffiana, Spiraea, Spiraea alba, Spiraea latifolia, Symphyotrichum, Syringa

vulgaris*, Tamarix gallica*, Vaccinium angustifolium, Vaccinium corymbosum, Vaccinium

Colla and Dumesh JESO Volume 141, 2010

myrtilloides, Vaccinium vacillans, Viola.

Habitats: Close to or within wooded areas, urban parks and gardens, wetlands.

Bombus griseocollis DeGeer: the brown-belted bumble bee

Phenology and Distribution: This species exhibits late spring emergence (earliest record

May 11) and has a widespread distribution (Fig. 6).

Floral Records: Robertson (1929): Actinomeris, Agastache, Amorpha, Arabis, Astragalus,

Baptisia, Bidens, Blephilia, Camassia, Campanula, Caulophyllum, Cephalanthus,

Cercis, Collinsia, Cornus, Crataegus, Delphinium, Dentaria, Desmodium, Dicentra,

Eupatorium, Frasera, Gerardia, Geranium, Gleditzia, Hibiscus, Houstonia, Hydrophyllum,

Ipomoea, Iris, Justicia, Krigia, Liatris, Lobelia, Lythrum, Mertensia, Monarda, Nelumbo,

Penstemon, Petalostemon, Phlox, Podophyllum, Polemonium, Prunella, Prunus,

Pediomelum, Pycnanthemum, Pyrus, Rudbeckia, Salix, Scutellaria, Sisyrinchium, Spiraea,

Symphoricarpos, Symphyotrichum, Teucrium, Tradescantia, Triodanis, Verbena, Vernonia,

Viburnum, Viola, Vitis, Uvularia, Zizia.

Macfarlane (1974): Aesculus hippocataneum*, Carduus nutans*, Echium vulgare*,

Helianthus annuus*, Hesperis matronalis*, Hydrophyllum_ virginianum, Hypericum

perforatum*, Linaria vulgaris*, Lonicera tatarica*, Melilotus alba*, Medicago sativa*,

Pyrus malus*, Robinia fertilis*, Solanum dulcamara*, Solidago canadensis, Solidago

sempervirens, Symphyotrichum novae-angliae, Symphytum officinale*, Syringa vulgaris*,

Taraxacum officinale*, Trifolium pratense*, Vicia cracca*.

Field and Museum Records: Aesculus glabra, Asclepias incarnata, Asclepias syriaca,

Asclepias tuberosa, Clinopodium vulgare, Echinacea, Eupatorium dubium, Eupatorium

perfoliatum, Heliopsis helianthoides, Hypericum prolificum, Lactuca canadensis,

Onopordum acanthium*, Pontederia cordata, Rudbeckia hirta, Vaccinium angustifolium,

Vaccinium corymbosum, Vaccinium myrtilloides, Urticularia vulgaris.

Habitats: Open farmland and fields, urban parks and gardens, wetlands.

Bombus fervidus Fabricius: the yellow bumble bee

Phenology and Distribution: This species exhibits mostly late spring emergence with a

long colony cycle extending into autumn and has a widespread distribution (Fig. 7). The

accumulated database contained only one single queen record in April with an unknown

date. The majority of queen records were from May and June.

Floral Records: Macfarlane (1974): Ajuga genevensis*, Ajuga reptans*, Aesculus

hippocataneum*, Caragana arborescens*, Carduus nutans*, Centaurea jacea*, Cichorium

intybus*, Cirsium arvense*, Delphinium, Dipsacus sylvestris*, Echium vulgare*,

Gleditzia, Hydrophyllum virginianum, Iris, Kalmia latifolia, Kolkwitzia amabilis*, Lactuca

canadensis, Lathyrus latifolius*, Linaria vulgaris*, Lonicera caerulea, Lonicera tatarica*,

Lotus corniculatus*, Malus, Melilotus alba*, Medicago sativa*, Monarda didyma, Nepeta

cataria*, Onobrychis viciifolia*, Onopordum acanthium*, Potentilla, Prunella vulgaris*,

Pyrus malus*, Ribes odoratum, Robinia fertilis*, Salix, Solanum dulcamara*, Solidago

canadensis, Solidago graminifolia, Symphyotrichum novae-angliae, Symphytum officinale*,

Syringa vulgaris*, Taraxacum officinale*, Trifolium pratense*, Tulipa*, Vicia cracca*,

Weigelia florida*.

Field and Museum Records: Centaurea maculosa*, Erigeron, Euthamia graminifolia, Inula

Bombus of southern Ontario JESO Volume 141, 2010

helenium*, Lobelia cardinalis, Lonicera, Lythrum salicaria*, Melilotus alba*, Penstemon

digitalis, Pontederia cordata, Securigera varia*, Solidago, Sonchus oleraceus*, Spiranthes,

Urticularia vulgaris.

Habitats: Open farmland and fields.

Bombus ternarius Say: the tri-coloured bumble bee

Phenology and Distribution: This species exhibits early spring emergence (earliest record

April 15) and has a widespread distribution except in extreme southwestern Ontario (Fig.

8).

Floral Records: Macfarlane (1974): Salix discolor.

Field and Museum Records: Aralia, Aralia hispida, Aralianudicaulis, Astragalus canadensis,

Chamaedaphne_ calyculata, Cirsium vulgare*, Claytonia caroliniana, Cynoglossum

officinale*, Echium vulgare*, Epilodium angustifolium, Erythronium albidum, Eupatorium

fistulosum, Eupatorium maculatum, Eurybia macrophylla, Euthamia_ graminifolia,

Hypericum perforatum*, Impatiens capensis, Ledum groenlandicum, Medicago sativa*,

Melilotus alba*, Mentha, Monarda, Pilosella aurantiaca*, Prunus pensylvanica, Rhexia

virginica, Securigera varia*, Solidago altissima, Solidago canadensis, Solidago juncea,

Spiraea, Spiraea alba, Symphyotrichum novae-angliae, Symphyotrichum puniceum,

Syringa vulgaris*, Tanacetum vulgare*, Taraxacum officinale*, Tilia americana, Trifolium

hybridum*, Trifolium pratense*, Trillium grandiflorum, Urticularia vulgaris, Vaccinium

angustifolium, Verbascum thapsus*, Verbena hastata, Vicia cracca*.

Habitats: Close to or within wooded areas.

Bombus rufocinctus Cresson: the red-belted bumble bee

Phenology and Distribution: This species exhibits late spring emergence (earliest record

May 28) and has a widespread distribution with more records in the Greater Golden

Horseshoe region (Fig. 9).

Floral Records: Macfarlane (1974): Lactuca canadensis, Potentilla, Symphyotrichum

(Aster) lateriflorum.Field and Museum Records: Apocynum androsaemifolium, Arctium

minus*, Campanula, Centaurea maculosa*, Cichorium intybus*, Cirsium arvense*,

Daucus carota*, Echium vulgare*, Eupatorium, Eupatorium fistulosum, Eupatorium

maculatum, Geranium, Helianthus, Heracleum lanatum, Hypericum prolificum, Inula

helenium*, Lupinus, Medicago sativa*, Melilotus alba*, Mentha, Monarda /fistulosa,

Prunella vulgaris*, Rubus, Securigera varia*, Solidago, Solidago altissima, Solidago

bicolor, Syringa vulgaris*, Trifolium pratense*, Trifolium repens*, Verbena hastata, Vicia

cracca*.

Habitats: Close to or within wooded areas, urban parks and gardens.

Bombus affinis Cresson: the rusty-patched bumble bee

Phenology and Distribution: This species exhibits early spring emergence (earliest record

April 20) with a long colony cycle extending into the autumn and has a southern distribution

with fewer recent records (Fig. 10).

Floral Records: Macfarlane (1974): Arctium minus*, Asclepias syriaca, Berberis

thunbergii*, Carduus nutans*, Centaurea cyanus*, Cotoneaster adpressa*, Crataegus,

Cucumis melo*, Deutzia gracilis*, Echium vulgare*, Hydrophyllum_ virginianum,

Colla and Dumesh JESO Volume 141, 2010

Hypericum perforatum*, Impatiens campensis, Kalmia latifolia, Linaria vulgaris*,

Lonicera caerulea, Lonicera periclymenum*, Lonicera tatarica*, Lotus corniculatus*,

Melilotus alba*, Medicago sativa*, Nepeta cataria*, Prunus americana, Prunus cerasus*,

Prunus tomentosa*, Pyrus malus*, Rhododendron, Ribes grossularia*, Ribes nigrum*,

Ribes rubrum*, Robinia fertilis*, Rubus, Salix, Silene dichotoma*, Solanum dulcamara*,

Solidago canadensis, Solidago fiexicaulis, Sonchus oleraceus*, Stachys palustris*,

Symphyotrichum (Aster) ericoides, Symphyotrichum (Aster) lateriflorum, Symphyotrichum

novae-angliae, Symphytum officinale*, Syringa vulgaris*, Taraxacum officinale*, Trifolium

pratense*, Trifolium repens*, Vicia cracca*, Vinca minor*, Weigelia florida*.

Field and Museum Records: Eupatorium maculatum, Eupatorium perfoliatum, Eupatorium

rugosum, Euthamia graminifolia, Helianthus decapetalus, Helianthus divaricatus, Kalmia,

Onopordum acanthium*, Rhexia virginica, Rhus, Spiraea, Vaccinium angustifolium,

Vaccinium vacillans.

Habitats: Close to or within wooded areas, open fields, urban parks and gardens.

Bombus citrinus Smith: the lemon cuckoo bumble bee

Phenology and Distribution: Males and females of this species are found from early spring

until late autumn and this species has a widespread distribution (Fig. 11). The earliest spring

record is a female collected on April 26.

Floral Records: Robertson (1929): Blephilia, Verbena.

Field and Museum Records: Centaurea jacea*, Cephalanthus occidentalis, Cirsium

vulgare*, Daucus carota*, Epilobium, Eupatorium fistulosum, Eupatorium maculatum,

Eupatorium perfoliatum, Euthamia graminifolia, Helianthus, Melilotus alba*, Prunella

vulgaris*, Solidago, Solidago altissima, Solidago bicolor, Symphyotrichum (Aster)

ericoides, Trifolium pratense*, Verbena hastata, Vicia cracca™.

Habitats: Close to or within wooded areas.

Bombus ashtoni Cresson: Ashton’s cuckoo bumble bee

Phenology and Distribution: Males and females of this species are found from early spring

until late autumn (Fig. 12). Recent occurrences of this species in Ontario are scarce. The

earliest spring record is a female collected on April 21.

Floral Records: Field and Museum Records: Allium, Aralia, Cephalanthus, Eupatorium,

Inula helenium*, Melilotus alba*, Penstemon, Pilosella aurantiaca*, Rubus, Solidago,

Solidago canadensis, Symphyotrichum novae-angliae, Syringa vulgaris*, Taraxacum

officinale*, Trifolium hybridum*, Trifolium pratense*, Vaccinium angustifolium, Vaccinium

corymbosum.

Habitats: Close to wooded areas.

Bombus borealis Kirby: the northern amber bumble bee

Phenology and Distribution: This species exhibits late spring emergence (earliest record

May 14) and is sparsely distributed across southern Ontario (Fig. 13).

Floral Records: Field and Museum Records: Astragalus canadensis, Cirsium vulgare*,

Echium vulgare*, Eupatorium fistulosum, Inula helenium*, Medicago sativa*, Melilotus

alba*, Onopordum acanthium*, Rubus, Solidago, Symphyotrichum novae-angliae, Trifolium

pratense*, Vicia cracca*.

Bombus of southern Ontario JESO Volume 141, 2010

Habitats: Close to or within wooded areas.

Bombus pensylvanicus DeGeer: the American bumble bee

Phenology and Distribution: This species exhibits late spring emergence (earliest record

May 15) and is mostly distributed in southernmost regions where it is at the northern edge

of its range (Fig. 14).

Floral Records: Robertson (1929): Amelanchier, Amorpha, Antennaria, Asclepias,

Astragalus, Baptisia, Blephilia, Campanula, Cephalanthus, Cerastium, Cercis, Circaea,

Claytonia, Clematis, Collinsia, Coreopsis, Cornus, Crataegus, Delphinium, Desmodium,

Dentaria, Dicentra, Frasera, Gaura, Gentiana, Geranium, Gerardia, Gymnocladus,

Helenium, Helianthus, Heliopsis, Heuchera, Hibiscus, Hydrophyllum, Impatiens, Ipomoea,

Tris, Krigia, Lespedeza, Lithospermum, Lobelia, Ludwigia, Lycopus, Mertensia, Mimulus,

Monarda, Nelumbo, Oenothera, Orobanche, Oxalis, Petalostemum, Penstemon, Phlox,

Physostegia, Plantago, Podophyllum, Polemonium, Polygonatum, Polygonum, Polytaenia,

Potentilla, Prenanthes, Prunella, Prunus, Pycnanthemum, Rhamnus, Ribes, Rubus,

Rudbeckia, Ruellia, Sagittaria, Salix, Scutellaria, Sida, Silene, Silphium, Sium, Solidago,

Stachys, Staphylea, Strophostyles, Symphoricarpos, Teucrium, Tilia, Tradescantia,

Triosteum, Uvularia, Verbascum, Verbesina, Verbena, Vernonia, Veronica, Viburnum, Viola,

Zizia.

Macfarlane (1974): Althaea rosea*, Carduus nutans*, Dipsacus sylvestris*, Echium

vulgare*, Erigeron philadelphicus, Symphyotrichum novae-angliae, Hesperis matronalis*,

Hydrophyllum virginianum, Kalmia latifolia, Lonicera tatarica*, Medicago sativa*, Pyrus

malus*, Robinia fertilis*, Solidago flexicaulis, Solidago graminifolia, Sonchus oleraceus*,

Symphytum officinale*, Syringa vulgaris*, Trifolium pratense*, Vaccinium angustifolium,

Vicia cracca*, Weigelia florida*.

Field and Museum Records: Astragalus canadensis, Campsis, Decodon, Dasistoma,

Echinacea, Euthamia graminifolia, Justicia, Lupinus, Pontederia cordata, Pediomelum,

Rubus idaeus, Symphyotrichum, Triodanis.

Habitats: Open farmland and fields.

Bombus insularis Smith: indiscriminate cuckoo bumble bee

Phenology and Distribution: Males and females of this species are found from late spring

until early autumn (Fig. 15). Records of this species are across southern Ontario but are

scarce. The earliest spring record is for a female collected on May 30.

Floral Records Field and Museum Records: Centaurea maculosa*, Eupatorium maculatum,

Melilotus alba*, Rubus, Solidago, Trifolium pratense*, Trifolium repens*, Vaccinium

angustifolium, Vicia cracca*.

Habitats: Unknown.

Bombus auricomus Robertson: the black and gold bumble bee

Phenology and Distribution: This species exhibits mid-spring emergence (earliest record

May 5) and has mostly a southwestern distribution (Fig. 16).

Floral Records Field and Museum Records: Dipsacus fullonum*, Eupatorium perfoliatum,

Hypericum, Malus, Monarda, Penstemon, Rubus occidentalis, Solanum dulcamara*,

Trifolium pratense*.

Colla and Dumesh JESO Volume 141, 2010

Habitats: Open farmland and fields.

Bombus sandersoni Franklin: Sanderson’s bumble bee

Phenology and Distribution: Few records exist for this species, consequently little is

known on its distribution and phenology (Fig. 17). The earliest spring record for this species

is June 2 in southern Ontario, indicating it may be a later emerging species.

Floral Records: Field and Museum Records: Epilobium, Gaylussacia, Kalmia angustifolia,

Kalmia polifolia, Lonicera, Malus, Monarda, Penstemon, Rhododendron, Rubus, Salix,

Sarracenia purpurea, Scutellaria lateriflora

Habitats: Unknown.

Bombus fernaldae Franklin: Fernald’s cuckoo bumble bee

Phenology and Distribution: Very few records exist for this species in the region,

consequently little is known on its distribution and phenology (Fig. 18). The earliest spring

record for this species is May 27 in southern Ontario.

Floral Records: Field and Museum Records: Asclepias, Aster, Helianthus, Pilosella

aurantiaca*, Potentilla, Rubus, Solidago, Solidago hispida, Taraxacum officinale*,

Trifolium repens*, Vaccinium angustifolium.

Habitats: Unknown.

Discussion

The bumble bees of southern Ontario differ in phenology, food plant choice,

abundance and habitat selection. Unlike some invertebrate pollinators, bumble bees in

southern Ontario are food generalists, foraging on a variety of native plant genera. For

example, B. impatiens, the most common species in the region (Fig. 1), has been found

visiting over 100 native plant genera throughout its range. This is likely due to their long

colony life-cycles which span the flowering cycles of more than one plant species. Broad

diet tolerances allows for behavioural flexibility in highly competitive situations where

nectar is often a limiting resource (Fontaine et al. 2008). Differences in food choice are a

major factor in niche partitioning among bumble bees in Ontario (Harder 1985).

This review indicates species in southern Ontario are often associated with particular

habitat types and associated phenologies. Interestingly, the earliest emerging species (i.e.

B. bimaculatus, B. ternarius, B. perplexus, B. affinis, B. vagans, B. impatiens, B terricola)

are all associated with wooded habitats (Figs. 1-5, 8, 10). These species likely have co-

evolutionary relationships with woodland spring ephemerals. Species associated with open

fields tend to be later emerging species (i.e. B. auricomus, B. griseocollis, B. pensylvanicus,

and B.fervidus) and are likely more reliant on later blooming field flowers (Figs. 6, 7, 14,

16). It has been suggested that later emerging species may be more vulnerable to stressors

such as habitat loss (Williams et al. 2009).

Although this study focuses on natural history, some comments can also be made

on species abundances. These data presented are consistent with previous findings that B.

impatiens and B. bimaculatus are the most abundant species in southern Ontario (Colla

and Packer 2008). Whether this was historically the situation or the consequence of their

Bombus of southern Ontario JESO Volume 141, 2010

tolerance of urbanized habitats remains to be determined. In contrast, many species have

few records and are both historically and recently uncommon. For some rarer taxa this

may be because they are at the northern edge of their native ranges in southern Ontario

(e.g. B. auricomus and B. pensylvanicus; Figs. 14, 16). Additionally, very few records

exist for most of the socially parasitic species (i.e. B. ashtoni, B. fernaldae, B. insularis),

which have likely been always rare and dependent on host abundances. Basic information

on distribution, ecological requirements and phenology are still required for the rare and

uncommon species to aid in the conservation of these potentially at-risk species (COSEWIC

2010). Additionally, to better understand this important group of bees, further study is

needed on nesting requirements, queen overwintering requirements, mating behaviours and

dietary breadth for all Bombus species in southern Ontario. Additional studies on changes

in distribution and abundance over time will also better our understanding of the ecological

needs of these native pollinators.

Acknowledgements

This study would not have been possible without the use of valuable historical

specimens from the following collections: Canadian Museum of Nature, University of

Guelph Insect Collection, Royal Ontario Museum and Algonquin Provincial Park. Thanks

to Laurence Packer, Paul Williams, Paul Catling, James Thomson and Cory Sheffield for

valuable discussion. This is contribution No. 8 from the Canadian Pollination Initiative

(NSERC-CANPOLIN)

References

Allen, G. M.., P. F. J. Eagles and S.D. Price. 1990. Conserving Carolinian Canada. University

of Waterloo Press, Waterloo, ON. 346 pp.

Benton, T. 2006. Bumblebees. Collins New Naturalist Series, London, UK. 592 pp.

Berenbaum, M., P. Bernhardt, S. Buchmann, N. W. Calderone, P. Goldstein, D. W. Inouye,

P. G. Kevan, C. Kremen, R. A. Medellin, T. Ricketts, G.E. Robinson, A. A. Snow,

S. M. Swinton, L. B. Thien and F. C. Thompson. 2007. Status of pollinators in

North America. The National Academies Press, Washington, DC. 312 pp.

Biesmeijer, J. C., S. P. Roberts, M. Reemer, R. Ohlemueller, M. Edwards, T. Peeters, A.

Schaffers, S. G. Potts, R. Kleukers, C. D. Thomas, J. Settele, and W. E Kunin.

2006. Parallel declines in pollinators and insect-pollinated plants in Britain and the

Netherlands. Science 313: 351-354.

Colla, S. R., M. C. Otterstatter, R. J. Gegear, and J. D. Thomson. 2006. Plight of the

bumble bee: Pathogen spillover from commercial to wild populations. Biological

Conservation 129: 461-467.

Colla, S. R. and L. Packer. 2008. Evidence for decline in eastern North American bumblebees

(Hymenoptera: Apidae), with special focus on Bombus affinis Cresson. Biodiversity

and Conservation 17: 1379-1391.

COSEWIC. 2010. Assessment and status report on the rusty-patched bumble bee (Bombus

Colla and Dumesh JESO Volume 141, 2010

affinis). 40 pp.

Findlay, C. S. and J. Houlahan. 1997. Anthropogenic correlates of species richness in

southeastern Ontario wetlands. Conservation Biology 11: 1000—1009.

Fontaine, C., C. L. Collin and I. Dajoz. 2008. Generalist foraging of pollinators: diet

expansion at high density. Journal of Ecology 96: 1002-1010.

Harder, L. D. 1985. Morphology as a predictor of flower choice by bumble bees. Ecology

66: 198-210.

Laverty, T. M. and L. D. Harder. 1988. The Bumble Bees of Eastern Canada. The Canadian

Entomologist 120: 965-987.

Macfarlane, R. P. 1974. Ecology of Bombinae (tbymenopterst Apidae) of Southern Ontario,

with emphasis on their natural enemies and relationships with flowers. PhD Thesis.

University of Guelph, Guelph, ON. 210 pp.

Mitchell, T. B. 1962. Bees of the eastern United States. II. Technical bulletin (North Carolina

Agricultural Experiment Station)*152: 1-557.

Robertson, C. 1929. Flowers and insects. Lists of visitors to four hundred and fifty-three

flowers. Science Press Printing Company, Lancaster, PA. 221 pp.

Taki, H., P. G. Kevan and J. S. Ascher. 2007. Landscape effects of forest loss in a pollination

system. Landscape Ecology 22: 1575-1587.

Williams, P., S. R. Colla and Z. Xie. 2009. Bumblebee vulnerability: common correlates of

winners and losers across three continents. Conservation Biology 23: 931-940.

Bombus of southern Ontario JESO Volume 141, 2010

—e— Queens

i Workers

Frequency

—t— Males

APR MAY JUNE JULY AUG SEPT OCT

Legend

a Before 1950

a 1950-1979

FIGURE 1. Phenology and distribution for Bombus impatiens in southern Ontario

(n=3017).

Colla and Dumesh JESO Volume 141, 2010

—e— Queens

—&— Workers

Frequency

—#— Males

APR MAY JUNE JULY AUG SEPT OCT

FIGURE 2. Phenology and distribution for Bombus bimaculatus in southern Ontario

(n=1316).

Bombus of southern Ontario JESO Volume 141, 2010

—e— Queens

—@— Workers

Frequency

—t— Males

JUNE JULY AUG SEPT OCT

FIGURE 3. Phenology and distribution for Bombus terricola in southern Ontario (n=996).

Colla and Dumesh JESO Volume 141, 2010

—e— Queens

= Workers

Frequency

—t— Males

APR MAY JUNE JULY AUG SEPT OCT

FIGURE 4. Phenology and distribution for Bombus vagans in southern Ontario (n=914).

Bombus of southern Ontario JESO Volume 141, 2010

—e— Queens

fi Workers

Frequency

—#te— Males

APR MAY JUNE JULY AUG SEPT OCT

Legend

A Before 1950

a 1950-1979

e 1980-2010

[] mca

FIGURE 5. Phenology and distribution for Bombus perplexus in southern Ontario

(n=646).

Colla and Dumesh JESO Volume 141, 2010

—e Queens

—®— Workers

Frequency

—*— Males

APR MAY JUNE JULY AUG SEPT OCT

FIGURE 6. Phenology and distribution for Bombus griseocollis in southern Ontario

(n=611).

Bombus of southern Ontario JESO Volume 141, 2010

—e— Queens

—f— Workers

Frequency

—*— Males

APR MAY JUNE JULY AUG SEPT OCT

FIGURE 7. Phenology and distribution for Bombus fervidus in southern Ontario (n=585).

Colla and Dumesh JESO Volume 141, 2010

Frequency

APR MAY JUNE JULY AUG SEPT OCT

Legend

a Before 1950

1950-1979

FIGURE 8. Phenology and distribution for Bombus ternarius in southern Ontario (n=534).

Bombus of southern Ontario JESO Volume 141, 2010

—@— Queens

—i— Workers

Frequency

—k— Males

MAY JUNE JULY AUG SEPT OCT

FIGURE 9. Phenology and distribution for Bombus rufocinctus in southern Ontario

(n=527).

Colla and Dumesh JESO Volume 141, 2010

Frequency

APR MAY JUNE JULY AUG SEPT

FIGURE 10. Phenology and distribution for Bombus affinis in southern Ontario (n=310).

Bombus of southern Ontario JESO Volume 141, 2010

—e— Queens

(ii Workers

Frequency

—t— Males

APR MAY JUNE JULY AUG SEPT

FIGURE 11. Phenology and distribution for Bombus citrinus in southern Ontario (n=285).

Colla and Dumesh JESO Volume 141, 2010

—e— Queens

—i— Workers

Frequency

—t— Males

APR MAY JUNE JULY AUG SEPT OCT

FIGURE 12. Phenology and distribution for Bombus ashtoni in southern Ontario

(n= 275).

Bombus of southern Ontario JESO Volume 141, 2010

—e— Queens

ii Workers

Frequency

—t— Males

APR MAY JUNE JULY AUG SEPT OCT

FIGURE 13. Phenology and distribution for Bombus borealis in southern Ontario

(n= 191).

Colla and Dumesh JESO Volume 141, 2010

Frequency

MAY JUNE JULY AUG SEPT OCT

FIGURE 14. Phenology and distribution for Bombus pensylvanicus in southern Ontario

(n=135).

Bombus of southern Ontario JESO Volume 141, 2010

—e— Queens

=i Workers

Frequency

—t— Males

APR MAY JUNE JULY AUG SEPT OCT

FIGURE 15. Phenology and distribution for Bombus insularis in southern Ontario (n=54).

Colla and Dumesh -JESO Volume 141, 2010

—e— Queens

—@— Workers

Frequency

—t— Males

APR MAY JUNE JULY AUG SEPT

FIGURE 16. Phenology and distribution for Bombus auricomus in southern Ontario

(n=43).

Bombus of southern Ontario JESO Volume 141, 2010

—e— Queens

i Workers

Frequency

—t— Males

MAY JUNE JULY AUG SEPT OCT

FIGURE 17. Phenology and distribution for Bombus sandersoni in southern Ontario

(n=40).

Colla and Dumesh JESO Volume 141, 2010

—e— Queens

—&— Workers

Frequency

—*— Males

MAY JUNE JULY AUG SEPT OCT

FIGURE 18. Phenology and distribution for Bombus fernaldae in southern Ontario

(n= 19).

Weevil communities in hardwood forests JESO Volume 141, 2010

GROUND-DWELLING WEEVIL

(COLEOPTERA: CURCULIONIDAE) COMMUNITIES IN

FRAGMENTED AND CONTINUOUS HARDWOOD FORESTS

IN SOUTH-CENTRAL ONTARIO

E. PROCTOR’, R. S. ANDERSON’, E. NOL’,

J. M. GIRARD* AND S. RICHMOND*

Department of Environmental and Life Sciences, Trent University

1600 West Bank Drive, Peterborough, ON, Canada K9J 7B8

email: eproctor@trentu.ca

Abstract J. ent. Soc Ont. 141: 69- 83

Weevils (Coleoptera: Curculionidae) are the largest family in the animal

kingdom and can be found in any habitat where plants grow. Many species

not native to North America have invaded both anthropogenic and natural

habitats, and the aim of this paper is to determine whether forest landscape

continuity has discouraged introduced species. We compared the ground-

dwelling weevil communities of hardwood forest fragments to those in

hardwood stands in a continuously forested landscape, with the prediction

that the fragments would have more introduced species. Pitfall traps

caught 5090 individuals from 26 species. Both landscapes were dominated

by introduced weevils (96% of all individuals), but forest fragments were

dominated by Barypeithes pellucidus (Boheman), while Sciaphilus asperatus

(Bonsdorff) represented 74% of all weevils caught in the continuous forest.

Sixty-four percent of the introduced species were parthenogenetic, and all

parthenogenetic species were polyphagous and flightless. Fifteen native

species were captured but they accounted for only 4% of total individuals, and

the only numerous native species, Hormorus undulatus (Uhler), was absent

from the continuous forest. Seven native species were each represented by

a single individual, one of which, Sirocalodes sericans (LeConte) is the first

record for Ontario. Ground-dwelling weevil communities in central Ontario’s

forests are composed largely of non-native species, and relatively intact forests

do not provide conservation protection for this group of invertebrates.

Published November 2010

' Author to whom all correspondence should be addressed

? Canadian Museum of Nature, email: randerson@mus-nature.ca

> Biology Department, Trent University, email: enol@trentu.ca

+ Biology Department, Carleton University, email: jphilli8@connect.carleton.ca

> Faculty of Forestry, University of Toronto email: sonya.richmond@utoronto.ca

Proctor et al. JESO Volume 141, 2010

Introduction

The family Curculionidae (Coleoptera), hereafter weevils, is the largest family in

the animal kingdom and contains about 51000 species worldwide, in 4600 genera (Anderson

1997; Oberprieler et al. 2007). As larvae and adults, almost all weevils are phytophagous,

although a few species are saprophagous. While most of the phytophagous species are

associated with angiosperms, weevils can be found in association with almost any terrestrial

or freshwater plant species, and any plant part (Anderson 2002; Oberprieler et al. 2007).

The largest subfamily, Entiminae, have larvae that mostly live in the soil and feed on roots.

Both adult and larval entimines are largely polyphagous, feeding on many species of host

plants. The other 17 subfamilies contain species whose larvae tend to live and feed inside

plant parts including stems, leaves, roots and reproductive structures. These subfamilies

also tend to have a more restricted range of host plants (monophagous or oligophagous),

some limited to a single family, genus or species (Anderson 2002).

There are about 600 described weevil species in Canada, many of which are

introduced (McNamara 1991). Most of the introduced species are of European origin

(Langor et al. 2009) and most introductions are attributed to the importation of ornamental

plants and other products, or to dry ballast (rock, soil and sand) dumped by British ships at

North American seaports at the turn of the nineteenth century (Anderson 2002; Majka et al.

2007). Introduced weevils are prevalent in forested habitats. Pinski et al. (2005a) found

66.4% of all adult Curculionidae caught in northern hardwood forests in the Great Lakes

region to be an introduced species (Phyllobius oblongus L.) and Coyle et al. (2008) reported

a suite of nine invasive root-feeding weevils from this same region. Maerz et al. (2005)

found one introduced weevil, Barypeithes pellucidus (Boheman), to be more abundant

than all other beetle taxa combined in mature forest stands in New York and Pennsylvania.

These past studies show the prevalence of introduced weevils in forests, but it is not known

whether any large forest stands in Ontario provide refuge for native species.

Niemela and Mattson (1996) argue that European phytophagous insects are better

at invading North American forests than their North American counterparts are at invading

European forests, especially in disturbed or fragmented landscapes where the plant

communities are partially European in origin (Burke and Nol 1998). Periods of glaciation,

more severe in Europe due to its topography than elsewhere in the world (Huntley 1993),

and the past 6000 years of human habitation and exploitation (Ledig 1992) resulted in a

long history of expansion and contraction of European forests. Through this volatility,

traits having high survival-value in patchy landscapes were selected over those adapted

to expansive forests. Armed with such traits as phenotypic plasticity, high reproductive

potential, and stress-tolerance mechanisms, many introduced species have become the

dominant phytophagous insects in their invaded niches (Niemela and Mattson 1996).

Our objective is to compare the ground-dwelling weevil assemblages among forest

fragments in an agricultural landscape and continuous forest in central Ontario, Canada.

We predicted that forest fragments would have a greater proportion and higher abundances

of introduced weevil species than stands found in a continuously forested landscape. We

discuss the life history attributes of the weevils in light of their ability to become established

in continuously forested landscapes.

Weevil communities in hardwood forests JESO Volume 141, 2010

Materials and Methods

Study Sites

Forest Fragments —Twenty-two mature deciduous woodlots in the southern portion

of Peterborough County (44° 17’ N, 78° 20’ W) in central Ontario, Canada were selected

as part of a broader study on forest-breeding birds. The region is part of the Great Lakes-

St. Lawrence lowlands, at the southern edge of the Canadian Shield, in the Mixedwood

Plains Ecozone of the Ecological Land Classification of Canada (AAFC 2010). The region

experiences an average of 124 frost-free days a year, and 84 cm of precipitation (Environment

Canada 2008). Elevation ranges from 75 to 408 m with an average of 305 m (Wickware

and Rubec 1989). Though the soils in this region are variable in composition, all woodlots

were located in well-drained loam or sandy-loam soils with layers of organic material 8-10

cm thick (Webber et al. 1946; Gillespie and Acton 1981). The landscape is composed

of agriculture and forest fragments, interspersed with farms, housing developments, and

wetlands (Phillips et al. 2005). On average, intensive agriculture (row-crops) accounted

for 28% of the landscape within 2 km of the woodlots, less-intensive agriculture (hay, old

fields, and pastures) accounted for 30%, and forest cover accounted for 24% (Richmond

2006). All 22 woodlots were dominated by Acer saccharum (Aceraceae), with Fraxinus

americana (Oleaceae), Ostrya virginiana (Betulaceae), Thuja occidentalis (Cupressaceae),

Fagus grandifolia (Fagaceae) and Tilia americana (Tiliaceae) all occurring as frequent

canopy species. The woodlots ranged from 6.7 to 280.4 ha, with a mean of 37.1 ha. None

of the woodlots had been recently exposed to grazing, logging, fire or construction (Phillips

et al. 2005, Richmond 2006). The understory plant communities at the sampling points in

forest fragments were largely native (S. Richmond, pers. obs.).

Continuous Forest — Nine mature hardwood stands in the southern portion of

Algonquin Provincial Park (45° 35’ N, 78° 29’ W) in central Ontario, Canada were selected

as part of a larger study on sustainable forest management. Algonquin Park lies in a

transition zone between the boreal forest to the north and the Great Lakes-St. Lawrence

lowlands to the south (Rowe 1972), in the Boreal Shield Ecozone of the Ecological Land

Classification of Canada (AAFC 2010). The park covers an area of 7700km/’, the western

two-thirds of which (about 4600km7) consists of tolerant hardwood forests over rugged

terrain, interspersed with numerous lakes (Quinn 2004). Mean elevation is 396m, with

the western portion of the park experiencing an average of 84 frost-free days a year, and

100cm of precipitation (Strickland 2006). Soils characteristic of the hardwood forests are

fresh to moist, medium to coarse loams, with an average organic matter layer of 10cm

(Chambers et al. 1997). All nine stands were characterized by canopies dominated by Acer

saccharum, with lesser amounts of Fagus grandifolia, Betula alleghaniensis (Betulaceae),

Prunus serotina (Rosaceae), and Tsuga canadensis (Pinaceae). Sites ranged from 14.4 to

54.8ha, with a mean of 35.8ha. Few non-native plant species occurred in these stands (E.

Proctor, pers. obs.).

Insect Sampling

Ground-dwelling invertebrates were sampled in all sites using pitfall traps, which

consisted of 500ml containers dug into the ground so that the lips of the containers were

Proctor et al. JESO Volume 141, 2010

flush with the soil. The containers were filled halfway with water, a pinch of salt (as a

preservative) and a few drops of dish soap(to disrupt surface tension). Each trap was covered

with a wire grate to reduce small mammal and amphibian by-catch, and to discourage larger

mammals from disturbing them.

Sampling in Peterborough County took place in the spring and summer of 2001,

2003 and 2004, and the number-and layout of pitfall traps varied from year to year. In

2001, one trap was placed every 20m along a 100m transect that extended from the edge of

the woodlot into the interior, for a total of six pitfalls per site. Traps were active from mid

May until mid August, for a total of approximately 8800 trap-days. In 2003 and 2004, the

transect was 50m long, with one trap placed every 5m from 0 to 20m, and then every 10m

from 20 to 50m, for a total of eight traps per site. In 2003, sampling went continuously

from late-May to late-July, for a total of approximately 8500 trap-days. In 2004, trapping

began in June and continued until August, and traps were left inactive for six to seven days

following each collection. This resulted in approximately 3600 trap-days. Thus, in the

fragmented forest we had a sampling effort of 20,900 trap-days.

Sampling in Algonquin Provincial Park took place in the spring and summer of

2006 and 2007. Each of the nine stands had 12 pitfalls, placed in a three-by-four grid, with

10m between traps. Pitfalls were located in the centre of each stand and were active for

two seven-day periods: once in late May and again in mid June, for a total of approximately

1500 trap-days per year for a total of 3000.

All invertebrates were rinsed and preserved in 70% ethanol upon collection.

Weevils were later separated from the rest of the samples and identified to species. Voucher

specimens were deposited at Trent University and the Canadian Museum of Nature. We

characterized dominant species as those that represented 5% or more of the total number of

specimens for a region.

Analysis

To determine whether parthenogenesis or flightlessness were more prevalent in the

introduced weevil species than in the native species caught in this study, we used Fisher’s

exact tests. When comparing communities, problems arise if sample sizes differ because

larger samples are expected to contain a greater number of species. The Peterborough

County sites were sampled more intensively than those in Algonquin Park, with sampling

taking place throughout the growing season as compared to a week each in late-May and

mid-June. We used rarefaction to standardize all samples to a common sample size by

estimating the number of species expected in a random sample of individuals taken from

a collection (Krebs 1999). We used EcoSim (Gotelli and Enstminger 2009) to estimate

the number of species expected in Peterborough County had we caught 1000 weevils

there during the same time periods as those in Algonquin and to compare species richness

between the two regions.

Results

In total, 5090 weevil specimens were collected from 10 subfamilies, 21 genera, and

26 species. The fragmented forest sites yielded 24 species, 11 of which were introduced,

Weevil communities in hardwood forests JESO Volume 141, 2010

and the continuous forest sites yielded five species, 3 of which were introduced.

More than 4000 individuals were captured in the Peterborough County pitfall traps

(Table 1). Most individuals collected were of introduced species (95.1%) even though

fewer than half the species were introduced (11 out of 24). The three dominant species were

Barypeithes pellucidus Boheman (63.7% of total), Otiorhynchus raucus Fabricius (17.7%),

and Phyllobius oblongus L. (6.1%). Of the 13 native species, Hormorus undulatus Uhler

was the most abundant (3.2%). All other native species represented 0.6% of the specimens,

or less. Five natives (Anametis granulata Say, Rhyncolus brunneus Mannerheim, Grypus

equiseti Fabricius, Listronotus sparsus Say, and Listronotus oregonensis LeConte) were

each represented by a single individual.

One thousand weevils from five species were caught in the Algonquin Park pitfall

traps (Table 1). The three dominant species were all introduced and accounted for 99.8%

of the total: Sciaphilus asperatus Bonsdorff (74%), P. oblongus (20%), and Otiorhynchus

ovatus L. (5.8%). The two native species, Nemocestes horni Van Dyke and Sirocalodes

sericans LeConte, were each represented by a single individual, and were not caught in

the forest fragments. Otiorhynchus ovatus, P. oblongus, and S. asperatus were found in

both forest types, and were collected every year. Five species (H. undulatus, O. raucus, B.

pellucidus, Polydrusus sericeus Schaller, and Trachyphloeus bifoveolatus Beck) were only

caught in the forest fragments, but were caught in all three years.

More than half of the species (57.7%) and 98.7% of the individuals caught in

this study were from the subfamily Entiminae. Of these, 13 were flightless (86.7%) and

nine were parthenogenetic (60%). All but one of the introduced species was from this

subfamily. Seven out of 11 introduced entimines (63.6%) were parthenogenetic and

flightless, three were bisexual and capable of flight (27.3%), and one was bisexual and

flightless. The proportion of parthenogenetic introduced species was significantly greater

than the proportion of parthenogenetic native species [7 out of 11 (63.6%) parthenogenetic

introduced; 2 out of 15 (13.3%) parthenogenetic native; Fisher’s exact test, P= 0.01]. The

proportion of flightless introduced species was not significantly greater than the proportion

of flightless native species [8 out of 11 (72.7%) flightless introduced; 6 out of 15 (40%)

flightless native; Fisher’s exact test, P = 0.10].

Peterborough County sites caught 2016 weevils from 18 species during the

weeklong periods in late-May and mid-June that corresponded to Algonquin’s sampling.

Using the rarefaction method, in a random sample of 1000 weevils from this subsample of

Peterborough County, we would expect to see between 13 and 18 species (95% confidence).

The Peterborough County sites therefore have higher species richness than the Algonquin

sites (1000 individuals from five species).

Discussion

Contrary to our prediction, introduced weevils overwhelmingly dominated both

the fragmented and continuously forested landscapes of central Ontario but the assemblages

of the two regions were distinct and species richness was higher in the fragmented sites.

Most of the weevils in both regions were flightless, but introduced species were more likely

to be parthenogenetic.

Proctor et al. . JESO Volume 141, 2010

TABLE 1. Subfamilies, status in North.America (native or introduced), flight capability,

mode of reproduction in North America, and known host plants of weevils (Coleoptera:

Curculionidae) captured in pitfall traps in 22 hardwood forest fragments in Peterborough

County, Ontario and 9 hardwood stands in Algonquin Provincial Park, Ontario.

Peterborough Algonquin

sehaoaly Species 2001 2003 2004 2006 2007. /°t!

Dryophthorinae Sphenophorus minimus Hart 19 5 24

S. parvulus Gyllenhal 10 ; 10

Erirhininae Grypus equiseti Fabricius l ]

Curculioninae Tychius picirostris Germar l l 2

Baridinae Stethobaris ovata LeConte 3 ] 4

Ceutorhynchinae Sirocalodes sericans LeConte l ]

Cossoninae Rhyncolus brunneus Mannerheim l ]

Cryptorhynchinae Acalles carinatus LeConte 15 o 18

Cyclominae Listronotus oregonensis LeConte l l

L. sparsus Say l ]

Entiminae Hormorus undulatus Uhler 55 51 25 131

Otiorhynchus ovatus L. 12 38 14 5 53 122

O. raucus Fabricius FO? 256" YSIS 724

O. rugosostriatus Goeze l l

O. singularis L. 2 2

Nemocestes horni Van Dyke l l

Phyllobius oblongus L. 90 = 124 =F ela ET 67 45]

Polydrusus sericeus Schaller 4 l 4 9

Barypeithes pellucidus Boheman 295 1799 “312 2606

Sciaphilus asperatus Bonsdorff 39 71 67 118 622 917

Sitona lineelus Bonsdorff 2 i

Cathormiocerus aristatus Gyllenhal* 30 16 46

Trachyphloeus bifoveolatus Beck 3 3 p 8

Anametis granulata Say l ]

Phyxelis rigidus Say I 3 4

Molytinae Conotrachelus posticatus Boheman 2 2

TOTAL 594 2430 1066 256 744 5090

Weevil communities in hardwood forests JESO Volume 141, 2010

TABLE 1. continued...

Known Host

Species Status’ Flight Reproduction Plants Sources

S. minimus Native Yes Sexual Poaceae, — Vaurie 1951

Cyperaceae

S. parvulus Native Yes Sexual Poaceae, —_- Vaurie 1951

Cyperaceae

G. equiseti Native Yes Sexual Equisetaceae Cawthra 1957; Anderson 2002

T. picirostris Introduced Yes Sexual Fabaceae Anderson and Howden 1994

(Trifolium)

S. ovata Native Yes Sexual Orchidaceae _ Blatchley and Leng 1916; Howden 1988

S. sericans Native Yes Sexual Papaveraceae, Anderson 2002; Korotyaev 2008

Fumariaceae

R. brunneus Native Yes Sexual Downed woody Anderson 1997; Anderson 2002

debris

A. carinatus Native No Sexual Downed woody Anderson 2002; LaChowska et al. 2009

debris

L. oregonensis Native Yes Sexual Apiaceae Campbell et al. 1989

L. sparsus Native Yes Sexual Asteraceae, Boivin 1999; Anderson 2002

Chenopodiaceae

H. undulatus Native No Sexual Liliaceae — Blatchley and Leng 1916; Champlain

and Null 1921

O. ovatus Introduced No Asexual Polyphagous Takenouchi 1965; Anderson 2002

O. raucus Introduced No Asexual Polyphagous Anderson 2002

O. rugosostriatus Introduced No Asexual Polyphagous Wheeler 1999; Anderson 2002

O. singularis Introduced No Asexual Polyphagous Campbell et al. 1989

N. horni Native No Asexual Polyphagous Anderson 2002

P. oblongus Introduced Yes Sexual Polyphagous _ Pinski et al. 2005a

P. sericeus Introduced Yes Sexual Polyphagous _ Pinski et al. 2005a

B. pellucidus Introduced No Sexual Polyphagous Takenouchi 1965; Galford 1987

S. asperatus Introduced No Asexual Polyphagous Pinski et al. 2005a

S. lineelus Native No Sexual Polyphagous Loan 1963

C. aristatus® Introduced No Asexual Polyphagous Piper et al. 2001

T. bifoveolatus Introduced No Asexual Polyphagous Brown 1965; Piper et al. 2001

A. granulata Native No Sexual Polyphagous Campbell et al. 1989; McLain 1998

P. rigidus Native No Asexual’ —_ Brassicaceae, Levesque and Levesque 1994; Shellhorn

Rosaceae and Sork 1997

C. posticatus Native Yes Sexual Fagaceae = Anderson 2002

(Quercus)

TOTAL

a) Subfamily classification based on Anderson 2002

b) Sources: McNamara 1991; Anderson 1997

c) Formerly Trachyphloeus

d) No males in Canadian Museum of Nature or Royal Ontario Museum collections

Proctor et al. | JESO Volume 141, 2010

Barypeithes pellucidus dominated the forest fragments in Peterborough County

while S. asperatus dominated the continuously forested sites in Algonquin Park. Both of

these entimine weevils are flightless and polyphagous as larvae and adults (Witter and Fields

1977; Galford 1987; Anderson 2002) but B. pellucidus reproduces sexually (Takenouchi

1965), while S. asperatus reproduces through apomictic parthenogenesis (Suomalainen et

al. 1987). Many parthenogenetic organisms are successful colonizers due to their abilities

to continually propagate even at low population numbers, and to rapidly adapt because

of more frequent random mutations (Ledig 1992; Langor et al. 2009). With asexual

reproduction and a preference for Acer saccharum and other deciduous trees (Witter and

Fields 1977), S. asperatus has been able to colonize not only the continuous hardwood

forests of Algonquin Park, but also those in Michigan and Wisconsin (Werner and Raffa

2000; Pinski et al. 2005a; Coyle et al. 2008). It is a widespread species (McNamara 1991;

Bright and Bouchard 2008) and has been found as far north as Iroquois Falls, Ontario (48°

45’ N, 80° 41° W), and Edmonton, Alberta (53° 32’ N, 113° 29’ W; Bright and Bouchard

2008).

Barypeithes pellucidus, with a preferred diet that includes Quercus rubra

(Fagaceae), Aster spp. (Asteraceae), Medicago spp. (Fabaceae), Trifolium spp. (Fabaceae),

and weedy herbaceous plants (Galford 1987; Campbell et al. 1989) is the dominant species

in the forest fragments in this study and in New York and Pennsylvania (Maerz et al. 2005).

High numbers have been found in agricultural sites such as vineyards (Bouchard et al.

2005), berry plantations (Bomford and Vernon 2005), residential areas (Balsbaugh 1988)

and continuous forests of Wisconsin and Michigan (Werner and Raffa 2000).

We suggest several explanations for why this adaptable colonizing species was

absent from the Algonquin Park samples. Weevil abundance can vary considerably

seasonally, from year to year, and from place to place (Balsbaugh 1988; Bouchard et al. 2005)

and it is possible that B. pe/lucidus is established in the sites in Algonquin Park but the small

sampling effort failed to detect them. In Quebec, Bouchard et al. (2005) caught over 1000

individuals of B. pellucidus in one vineyard but caught only 19 in another vineyard 30km

away. They hypothesized that the high clay-content soils of the depauperate vineyard were

less favourable to this species’ pupation, but none of the soils in our study contained much

clay (Webber et al. 1946; Gillespie and Acton 1981; Chambers et al. 1997). Barypeithes

pellucidus are univoltine (Campbell et al. 1989). Adults usually emerge early in spring and

disappear by mid-summer (Galford 1987; Maerz et al. 2005), and have only been found in

the milder parts of Canada, such as southern British Columbia, around the Great Lakes, and

in the Maritimes (Bright and Bouchard 2008). Specimens have been collected from as far

north as Sault Sainte Marie, Ontario (46° 30’ N, 84° 20’ W; Takenouchi 1965) and Montreal,

Quebec (45° 32’ N, 73° 38’ W; Bright and Bouchard 2008) but the frost-free periods in

both these locations (120d and 140d, respectively), and in Peterborough County (124d), are

much longer than in Algonquin (84d; Marsan 1990; Strickland 2006; Environment Canada

2008; MRCC 2009). The short frost-free period in Algonquin may be insufficient for one

generation to find mates, lay eggs, and for the larvae of the next generation to hatch and

grow to a sufficient size to survive the winter. If B. pe//ucidus is present in Algonquin Park,

the combination of its flightlessness, sexual reproduction, and the short frost-free period, all

likely contribute to limiting its numbers there.

Otiorhynchus raucus, like B. pellucidus, was abundant in Peterborough County,

Weevil communities in hardwood forests JESO Volume 141, 2010

but absent from the Algonquin Park samples. This species is flightless, polyphagous, and

parthenogenetic in North America (Mazur 1992; Bright and Bouchard 2008), and has been

found as far north as Calgary, Alberta (51° 07’ N, 114° 19’ W; Bright and Bouchard 2008).

It was first reported in North America in 1936 at a nursery in Fonthill, Ontario (Hicks 1947),

and the larvae are serious pests of garden vegetables, while adults feed on the foliage and

shoots of fruit trees (Campbell et al. 1989). According to Mazur (1992), in Europe O.

raucus is acommon component in anthropogenic habitats such as urban parks, gardens, and

roadsides, and it is probably absent from Algonquin Park due to the lack of cultivated plants

on which it prefers to feed in both its native and introduced range.

Phyllobius oblongus was a dominant species in both landscapes. Unlike the

aforementioned species, P. oblongus is capable of flight, and thus is a good disperser. It is

established in continuous hardwood forests in the Great Lakes Region (Pinski et al. 2005a;

Coyle et al. 2008) and in Nova Scotia (McCorquodale et al. 2005). Host plants include

a wide variety of trees and shrubs, especially Acer saccharum (Witter and Fields 1977)

and Ostrya virginiana (Pinski et al. 2005b). These tree species were common in both

landscapes.

Otiorhynchus ovatus was present in both regions and in all five years, and was

a dominant species in Algonquin. Commonly known as the strawberry root weevil, it is

abundant, widely distributed, and can be found wherever plants occur (Bright and Bouchard

2008). It is flightless, parthenogenetic, extremely fecund, and has a very broad range of

hosts, including conifers (Warner and Negley 1976; Campbell et al. 1989). This broad

niche and the capability of overwintering as adults or larvae (Campbell et al. 1989), have

likely facilitated O. ovatus’ ability to colonize as far north as Fairbanks, Alaska (64° 50° N,

147° 38’ W; Bright and Bouchard 2008). Why it was collected less frequently in the forest

fragments than its congener O. raucus is unknown. Otiorhynchus raucus (5.5 to 7.5 mm) is

larger than O. ovatus (4 to 5.5 mm; Bright and Bouchard 2008) and since pitfalls select for

larger, more active individuals (Baars 1979), the high catches of O. raucus may not reflect

the true proportions of these two species.

The only introduced weevil species caught in this study not in the subfamily

Entiminae was Tychius picirostris Fabricius (subfamily Curculioninae). Commonly known

as the clover-seed weevil, its larvae feed inside the reproductive structures of naturalized and

cultivated clovers (Trifolium spp.; Anderson and Howden 1994; Anderson 2002). Though

there are a few species of clover native to Ontario (e.g. T: reflexum), most are introduced

from Europe (e.g. 7. repens and T. pratens), and have been spread throughout the continent

for their use as forage and in crop rotation (Taylor 1985; Voss 1985). Majka et al. (2007)

suggest 7. picirostris was introduced to North America in dry ballast, and with the ability to

fly and the widespread distribution of its host plants (i.e. introduced clovers), it has become

established throughout the continent (Anderson and Howden 1994).

Hormorus undulatus was the only abundant native weevil species captured in this

study, and it was only caught in the Peterborough sites. Little is known of this entimine

species beyond that it reproduces sexually, is flightless, and has been found on members of the

Liliaceae family (Convallaria, Maianthemum and Polygonatum; Blatchley and Leng 1916:

Champlain and Knull 1921). In Canada, it has been recorded as far north as Wawa, Ontario

(47° 59’ N, 84° 46’ W; Bright and Bouchard 2008), and three were collected from raspberry

(Rubus idaeus) in a Quebec plantation (Levesque and Levesque 1994). A few individuals

Proctor et al. JESO Volume 141, 2010

were collected in continuous hardwood forests in the Great Lakes Region compared to

the thousands of introduced specimens (Coyle et al. 2008). Though sampling was not as

intensive in Algonquin Park as it was in Peterborough County, the absence of H. undulatus

from the Park’s samples indicates that it is not an abundant species there. If it ever had been

a significant component of the weevil community in this forest and those elsewhere, it is

possible that the competitive abilities of the invasive European species already mentioned

have displaced them. The short growing season may also have prevented H. undulatus from

establishing in Algonquin Park and may explain the substantially lower species richness of

the weevil communities of the park as compared to the more southerly forest fragments.

All parthenogenetic species caught in this study were flightless and polyphagous.

With parthenogenetic reproduction and a polyphagous diet, these weevils do not need to

find mates nor travel far to find food. The high proportion of introduced species that were

parthenogenetic (63.6%) emphasizes the superior colonizing ability of these flightless

weevils over flightless sexual species like B. pellucidus. The proportion of flightless

introduced species (72.7%), however, was not significantly greater than the proportion of

flightless native species (40%), which suggests that flightlessness alone does not affect

colonizing ability. Therefore the polyphagous diets of entimines, in combination with

either parthenogenetic reproduction or the ability to fly, have made these introduced weevils

dominate the forested sites of this study.

Species richness was lower in the continuous forest sites than in the forest

fragments. We found only five species in 1000 individuals in Algonquin, whereas we would

expect 13 to 18 species in a same-sized sample from Peterborough County for the same

time period. With three common species in Peterborough County (B. pellucidus, O. raucus,

and H. undulatus) all seeming to be absent from Algonquin Park, it is likely that the shorter

frost-free period, as well as the lack of agricultural and anthropogenic habitats are limiting

others species as well.

This is the first report of Sirocalodes sericans for Ontario. Though it is generally

distributed in the western and southern regions of the United States, in Canada it has only

been documented in Manitoba (McNamara 1991; Anderson 2002). Other Sirocalodes

species are associated with Papaveraceae and Fumariaceae, with larvae mining the stems

or crowns of the host plants (Anderson 2002). Dicentra cucullaria (Fumariaceae) is a

common spring ephemeral in northern hardwood forests (Walton and Hufford 1994), and

is a possible host for this weevil. Further targeted sampling around this plant might help to

elucidate more of this species’ biology.

Pitfall traps are useful in assessing relative abundance of invertebrates active at

the ground level, and are the most efficient method to assess ground-dwelling invertebrate

communities (Prasifka et al. 2007). Further research on the weevil communities in these

areas would benefit from additional sampling techniques such as flight-intercept traps,

emergence traps, and sweep-netting.

The life histories and effects of invasive weevils are thoroughly studied in

agricultural systems (e.g. Otiorhynchus sulcatus Fabricius; Moorhouse et al. 1992) because

of the economic damage they can cause, but forest-invaders are poorly understood. We

lack information on native weevil assemblages in forests prior to the invasions (Pinski et

al. 2005a) and studying the tolerances and below-ground herbivory of introduced larval

entimines is difficult (Coyle et al. 2008). There is ample scope for further study of the

Weevil communities in hardwood forests JESO Volume 141, 2010

functional role of these adaptable insects in our forested ecosystems.

Acknowledgements

We thank the many field and lab assistants, D. R. Coyle, D. Beresford, and two

anonymous reviewers for their helpful comments. Funding for this work was provided by

the National Science and Engineering Research Council, the Forestry Futures Trust Grant,

the Ontario Ministry of Natural Resources, Environment Canada and Trent University.

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Megachile ericetorum in Ontario, Canada JESO Volume 141, 2010

DISCOVERY OF THE WESTERN PALEARCTIC BEE,

MEGACHILE (PSEUDOMEGACHILE) ERICETORUM

(HYMENOPTERA: MEGACHILIDAE), INONTARIO, CANADA

C. S. SHEFFIELD', T. GRISWOLD? AND M. H. RICHARDS?

Department of Biology, York University,

4700 Keele St., Toronto, ON, Canada M3J 1P3

email: corys@yorku.ca

Scientific Note J. ent. Soc. Ont. 141: 85-92

The indigenous bee fauna of North America north of Mexico contains approximately

3500 described species (Ascher and Pickering 2010), but nearly 30 exotic species of Old

World origin are published as established (Cane 2003; Committee on the Status of Pollinators

in North America 2007). A few of these introduced species have a long history in North

America. The first was the honey bee, Apis mellifera L., brought with European settlers in

the 1620’s (Crane 1999; Horn 2005) for honey and wax production, roles now overshadowed

in importance by pollination services (Free 1993; Delaplane and Mayer 2000). Other, albeit

very few, bee species have been purposely introduced into North America for evaluation as

crop pollinators (e.g., Torchio and Asensio 1985; Batra 2003), though none of these species

are used commercially. Most bee introductions have been accidental. Brown (1950) and

Lindroth (1957) theorized that many introduced insect species may have arrived in the

New World through the importation of dry ballast (e.g., rock, sand, soil). The earliest bee

introductions likely included the ground-nesting species Andrena wilkella (Kirby), which

has been in eastern North America since the 1800’s (Malloch 1918), and Lasioglossum

leucozonium (Schrank). Lasioglossum leucozonium was only recently determined to be an

introduced species (Giles and Ascher 2006; Zayed et al. 2007), and not naturally Holarctic

in distribution as previously assumed (McGinley 1986). The North American population(s)

may have established from a single mated female (Zayed et al. 2007). These and the few

other introduced ground-nesting bee species may have been introduced via ballast from

ships in eastern North America (Giles and Ascher 2006).

Most successful introductions of bees (ca. 80%) have involved cavity-nesting

species (Cane 2003; Committee on the Status of Pollinators in North America 2007), those

nesting in pre-existing or easily excavated cavities, such as in hollow or pithy plant stems or

beetle burrows in wood (Michener 2007). The alfalfa leafcutter bee, Megachile rotundata

Published November 2010

' Author to whom all correspondence should be addressed.

2 USDA-ARS Bee Biology and Systematics Laboratory, 5310 Old Main Hill, Logan, UT,

USA 84322-5310

3 Department of Biological Sciences, Brock University, St. Catharines, ON, Canada

L2S 3Al

Sheffield et al. JESO Volume 141, 2010

(F.), has been in North America since at least the 1930’s (Cane 2003) and is now widespread

across the United States and southern Canada, as far east as Nova Scotia (Sheffield et al.

2008), and two additional species of the subgenus Eutricharaea Thomson are also widely

established. Three additional cavity-nesting megachilid bees, Megachile sculpturalis

Smith, Anthidium manicatum L.,, and A. oblongatum (Illiger) are rapidly spreading in North

America (Paiero and Buck 2004; Hinojosa-Diaz 2008; Zavortink and Shanks 2008; Gibbs

and Sheffield 2009; Tonietto and Ascher 2009). Several species only recently detected in

North America, such as M. scu/pturalis (Magnum and Brooks 1997) and Hylaeus hyalinatus

Smith (Ascher 2001), are now widely distributed and locally abundant members of the North

American fauna. Many introduced cavity-nesting species do very well in urban settings

(Matteson et al. 2008). For example, Chelostoma campanularum (Kirby), a recent arrival

in Canada (Buck et al. 2006), is now relatively common in Ontario in the cities of Guelph,

St. Catharines, and Toronto. Urban settings can support introduced bee species due to the

presence of introduced plant species including floral hosts also visited in their native ranges

(Hanley and Goulson 2003; Matteson et al. 2008; Gibbs and Sheffield 2009).

Invasive species are one of the biggest threats to regional biodiversity (Wilson

1999: Chivian and Bernstein 2008). Although introduced bee species account for less

than 1% of the species in North America, they often constitute much larger proportions

of surveyed faunas (calculations exclude Apis): 15% in Grixti and Packer (2006); 29% in

Sheffield (2006); 27% in Matteson et al. (2008); and 8% in Tuell et al. (2009). Additionally,

a recent survey in Guelph, ON found 12.5% ofall bees captured in pan traps to be introduced

species (M. Horn, unpublished data).

Considering that some introduced bee species are thought to disrupt local

indigenous bee populations, and potentially pollination, through competition for floral (e.g.

Paini 2004; Paini and Roberts 2005) and/or nesting resources (Barthell et al. 1998), it is

especially important to note their presence and monitor their establishment (Cane 2003).

It is also important to establish patterns of floral use, since many introduced species share

floral resources with native species, especially in urban settings (Matteson et al. 2008) and

agricultural settings that may have limited native floral resources.

The purpose of this note is to report the discovery in southern Ontario of Megachile

ericetorum Lepeletier, a Western Palearctic bee species new to the Western Hemisphere. Its

biology and diagnostic characters are briefly summarized, and methods for monitoring its

potential establishment are discussed.

Megachile (Pseudomegachile) ericetorum Lepeletier, 1841

Megachile ericetorum is wide-ranging in the Old World, occurring throughout

most of Europe (excluding western Scandinavia), Asia Minor, the Caucasus, Central Asia,

western North A frica, and Syria (Westrich 1989; Ozbek and van der Zanden 1994; Banaszak

and Romasenko 1998). Like many members of the genus Megachile Latreille, this species

nests in pre-existing cavities in canes or wood (Westrich 1989; Banaszak and Romasenko

1998). Females lack beveled cutting edges in the interspaces of the mandibular teeth (Fig.

1), as Pseudomegachile Friese and other members of Chalicodoma sensu lato (Megachile

Group 2, as per Michener 2007) do not cut leaf sections for nest construction but instead use

other materials such as plant resins, sand, and pebbles to construct nest partitions (Mitchell

1980; Westrich 1989; Snelling 1990; Banaszak and Romasenko 1998; Michener 2007).

Megachile ericetorum in Ontario, Canada JESO Volume 141, 2010

FIGURE 1. Face of specimen of female Megachile ericetorum Lepeletier collected in St.

Catharines, ON, Canada, showing 4-dentate mandibles without cutting edges.

Westrich (1989) indicates that this species is oligolectic on Fabaceae, mainly Lotus and

Lathyrus; males have been collected on Stachys.

Although the subgenus Pseudomegachile is indigenous to the Old World, another

introduced species, M. (Pseudomegachile) lanata (F.), is commonly collected in Florida

(Leavenwood and Serrano 2005) and the West Indies (Genaro 1997). Sheffield et al. (in

press) provide keys and full descriptions to distinguish M. ericetorum from other leafcutter

bees in Canada, although at this time, evidence is lacking as to whether this species has

established successfully. The female of M ericetorum can be distinguished from most

Megachile in Canada by the lack of cutting edges between the mandibular teeth (Fig. 1),

excluding M. sculpturalis which is much larger (> 20mm) with much orange pubescence,

and M. angelarum Cockerell and M. campanulae (Robertson), which are both slightly

smaller than M. ericetorum and lack the single median apical tubercle on the clypeus of M.

ericetorum (Fig. 1). Although the male has not yet been observed in Canada, males of M.

ericetorum are distinguishable from all other Megachile in Canada, except M. coguilletti

Cockerell, by the unmodified yellowish front tarsomeres; it differs from that species by

lacking a lower triangular process on the mandible (Sheffield et al. in press).

Megachile ericetorum is currently only known in North America from a single

female specimen (Fig. 2) collected on the Niagara Escarpment in St. Catharines, Ontario in

2003 (14.vii.2003; coll. Amy Rutgers) in a former farm field east of, and contiguous with,

the Glenridge Quarry Naturalization Site (43.124, —79.237; elev. 170m), and bordered on

the west by Highway 406. The Naturalization Site was formerly a limestone quarry and

then a landfill, which was closed in 2001 and completely replanted by 2003, whereas the

field in which the specimen was found belongs to Brock University and has remained more

or less undisturbed for almost 50 years. Westrich (1989) indicated similar habitat use (.e.,

calcareous grasslands) by this species in Europe. The habitat in which the specimen was

Sheffield etal. ~ JESO Volume 141, 2010

FIGURE 2. Female Megachile ericetorum Lepeletier, A) lateral and B) dorsal view.

Specimen collected in St. Catharines, ON, Canada.

found and the method by which it was collected (pan-trapping) suggest that this species may

have had the opportunity to establish in the area because the site is directly adjacent to a

former landfill. This bee species could have been transported to the area in discarded lumber,

for instance in wooden skids commonly used in international shipping. St. Catharines is

part of the major shipping route for international materials arriving into Canada and the

United States, and is < 20 km north of the United States border. Major seaports offer many

opportunities for introduced species (Majka and LeSage 2006), and this region has a long

history of commercial sea traffic. Several cavity-nesting species have been intercepted at

Megachile ericetorum in Ontario, Canada JESO Volume 141, 2010

such ports of entry (Cane 2003), although they may not always have the opportunity to

establish.

Since M. ericetorum has been in Canada at least since 2003, monitoring its

establishment and spread should be done (Cane 2003). Ultimately, modeling its potential

range in North America based on habitat suitability (e.g., Hinojosa-Diaz et al. 2005) may

indicate if it has the ability to become widespread across many ecozones, as in its native

range. Trap-nest surveys (e.g., Fye 1965; Krombein 1967; Sheffield et al. 2008) would

provide a means of monitoring the establishment of this species in North America, as well as

the possible displacement of native species (Barthell et al. 1998). Males of M. ericetorum,

like those of the introduced Anthidium manicatum (Severinghaus et al. 1981; Wirtz et al.

1988), aggressively defend territories (Hass 1960). This behaviour should make the species

stand out among the native Megachile species, potentially assisting in documenting its

spread.

Acknowledgements

We appreciate the assistance of John Ascher (American Museum of Natural History,

New York, NY) who provided the initial identification of this specimen, as well as many

useful comments on the manuscript, Matthias Buck (Royal Alberta Museum, Edmonton,

AB) for reviewing the manuscript and providing additional useful comments, as well as an

anonymous reviewer.

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JESO Volume 141, 2010

ERRATUM

L. Timms. 2009. Growing pains: How the birth of the Entomological Society of Canada

affected the identity of the Entomological Society of Ontario. The Journal of the

Entomological Society of Ontario 140: 49-56.

The page numbers published for this manuscript in JESO 140 were incorrect. The correct

reference is:

L. Timms. 2009. Growing pains: How the birth of the Entomological Society of Canada

affected the identity of the Entomological Society of Ontario. The Journal of the

Entomological Society of Ontario 140: 46-53.

Below is how the title should appear. The editor apologizes for this error.

GROWING PAINS: HOW THE BIRTH OF THE

ENTOMOLOGICAL SOCIETY OF CANADA AFFECTED THE

IDENTITY OF THE ENTOMOLOGICAL SOCIETY OF ONTARIO

L. TIMMS

Faculty of Forestry, University of Toronto

33 Willcocks Street, Toronto, Ontario, Canada M5S 3B3

email: laura.timms@utoronto.ca

Special Contribution J. ent. Soc. Ont. 140: 46-53

JESO Volume 141, 2010

THE ENTOMOLOGICAL SOCIETY OF ONTARIO

OFFICERS AND GOVERNORS

2010-2011 ©

President: H. FRASER

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ENTOMOLOGICAL SOCIETY OF ONTARIO

The Society founded in 1863, is the second oldest Entomological Society in North America

and among the nine oldest, existing entomological societies in the world. It serves as an

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CONTENTS

T. FROM THE EDUT OR oceceosioseseininscsovccseintoninniansescasttebaaalaias ane sistas

I. ARTICLES

D. L. LEGROS and D. V. BERESFORD — Aerial foraging and sexual dimorphism in bu

beetles (Silphidae: Coleoptera) in a central Ontario forest............csscscsssssssscssssssessesssssseeeees 3-10

J. L. VICKRUCK, J. T. HUBER and M. H. RICHARDS — Natural enemies of the bee gen 1s”

Ceratina (Hymenoptera: Apidae) in the Niagara Region, Ontario, Canada..............s0+0-+.1 1-26

P.M. CATLING, H. GOULET, R. BENNETT and B. KOSTIUK — Orthopterans (Orthoptera),

ground beetles (Coleoptera: Carabidae), and spiders (Araneae) in burned and unburned alvar

woodlands — the importance of postfire succession to insect iversity...........ssssescssseseees 27-37

S. C. COLLA and S. DUMESH — The bumble bees of southern Ontario: Notes on natu al ;

history and distribution...............:cesccessvoshesenecsiibietaamitishiianemnmnapeanaaan 5

E. PROCTER, R. S. ANDERSON, E. NOL, J. M. GIRARD and S. RICHMOND — Groun d-

dwelling weevil (Coleoptera: Curculionidae) communities in fragmented and continuous

hardwood forests in south-central Ontari0............scecssseesesserseseesesseeesseseeseeseeesneenenseneenenees 69-83,

;

Ill. NOTE

C. S. SHEFFIELD, T. GRISWOLD and M. H. RICHARDS — Discovery of the Weste a

Palearctic bee Megachile (Pseudomegachile) ericetorum (Hymenoptera: Megachilidae), in

Ontario, Canada.... SOSOSSSSSSSSSSHESSSSOSEOSOSSEESSESSE EEO E OEE ESOS OS EEEEEEEEEEEEEEEEESEEEEESESSSESEESESESSEESEEEEEEEESS

IV. ERRATUM

L. TIMMS. 2009. Growing pains: How the birth of the Entomological Society of Canada

affected the identity of the Entomological Society of Ontario. The Journal of the Entomologiell

Society of Ontario 140: 49-56.......cccc.ccssccesescscessesssoesseersesssnasenstesaccnsessensnsnnienansieiessintinasedsnataaE a

V. ESO OFFICERS AND GOVERNORS 2010-201 1........scssssssssssssnsossssssssessnssssesssnscssnscenseeensess 000.94

VI. ESO OFFICERS AND GOVERNORS 2009-2010........ssssesssssessessesnesneeses ..inside front cover

VIL. FELLOWS OF THE ESO .o...conc:ccssccscecsansssensciasspennioserentetacmomnensatasnnanettd inside front cover

VIII. APPLICATION FOR MEMBERSHIP.............0cccescccsescossssenssssossetenesnsaseees inside back cover

IX. NOTICE TO CONTRIBUTORS i-csnsccsehecccenc shee inside back cover