Coccinellinae) from Historic

ASSESSING GEOGRAPHIC DISTRIBUTION AND RELATIVE ABUNDANCE PATTERNS OF

NATIVE AND NON-NATIVE LADY BEETLES (COCCINELLINAE) FROM HISTORIC

OCCURRENCE DATA

A Thesis

Submitted to the Graduate Faculty

In Partial Fulfilment of the Requirements

For the Degree of Master of Science

Department of Biology

Faculty of Science

University of Prince Edward Island

Meghan Marriott

2012

Published Heritage Direction du 1+1 Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A0N4 Ottawa ON K1A 0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-94052-5

Our file Notre reference ISBN: 978-0-494-94052-5

NOTICE: AVIS: The author has granted a non L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distrbute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats.

The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission.

In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these.

While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. Canada CONDITIONS FOR THE USE OF THE THESIS

The author has agreed that the Library, University o f Prince Edward Island, may make this thesis freely available for inspection. Moreover, the author has agreed that permission for extensive copying of this thesis for scholarly purposes may be granted by the professor or professors who supervised the thesis work recorded herein or, in their absence, by the Chair of the Department or the Dean of the Faculty in which the thesis work was done. It is understood that due recognition will be given to the author of this thesis and to the University of Prince Edward Island in any use of the material in this thesis. Copying or publication or any other use of the thesis for financial gain without approval by the University of Prince Edward Island and the author's written permission is prohibited.

Requests for permission to copy or to make any other use o f material in this thesis in whole or in part should be addressed to:

Chair of the Department of Biology Faculty of Science University of Prince Edward Island 550 University Avenue, Charlottetown, PE Canada CIA 4P3 SIGNATURE PAGE

6 > >) 4 '(\\J )

REMOVED ABSTRACT

Native species of lady beetles are believed to be declining in distribution and/or relative abundance across North America, but an accurate broad scale picture has been difficult to obtain. One source of data that could be tapped to assess broad distributional questions is the information associated with specimens in natural history collections. Natural history collection data have proven to be effective for assessing various ecological questions relating to species diversity and abundance patterns, changes in distribution, and in documenting the spread of invasive species for a wide variety of taxa. However, specimen data in collections may contain temporal and spatial gaps which can cause interpretation biases that vary with the type of collection, so collection data must be critically reviewed to assess its usability and limitations. The first goal of this study was to assess potential data gaps (and corresponding strengths) associated with lady beetle collection records from Atlantic Canada to determine whether these data can be used to determine long term distribution and relative abundance patterns for lady beetles in eastern Canada. Twenty-seven Atlantic Canadian collections representing four types of collections (Government, Museum, University and

Private) were assessed for potential data gaps by comparing numbers of specimen records collected for different time periods, seasons, taxonomic groups (especially native vs. non-native species), and geographical areas for each type of collection using

Mann-Whitney U-tests or Kruskal-Wallis tests. Each analysis was run for each collection type, and again with data combined from all collections to determine whether observed spatial or temporal differences could be minimized by combining collection data for multiple collection types. Individual collection types did show significant differences among these variables, indicating that use of data from individual collections, even large ones, results in data gaps that could interfere with interpretation of lady beetle distributions. However, differences (especially temporal gaps) among these variables were minimized, or were no longer significant when data from all collections were combined, indicating that biases could be minimized by including data from multiple collections. Therefore, the second goal of this study was to use data from multiple collections to assess temporal patterns in distribution and relative abundance in a suite of five native and five non-native lady beetle species in eastern Canada. Relative abundances for the 10 species were calculated by decade and compared using a G-test

(likelihood ratio test) to assess changes in lady beetle populations overtime.

Geographic area occupied by each species was calculated for each tim e period though

'rubber banding' in GIS software. Collection data showed a clear decline in relative abundance and geographic extent in four of the five native lady beetles studied, though timing of declines varied with species. In contrast, the non-native species showed a clear increase in relative abundance from the time of their establishment at least to the end of the study period in 2009, though three of the five species experienced range contractions after reaching peaks in abundance and area. This study provides strong evidence for the decline in several native species of lady beetles in eastern Canada, and does so over a wide geographic area. ACKNOWLEDGEMENTS

Funding was provided for this project by a Natural Sciences and Engineering

Research Council (NSERC) Discovery Grant to D. Giberson and D. McCorquodale and a contract from Environment Canada (Committee on Endangered Species and Wildlife in

Canada, COSEWIC).

My sincere thanks to all the individuals who provided access to lady beetle collections throughout eastern Canada, as well as those who shared pre-existing databases (Table 3.1 in thesis), particularly Patrice Bouchard and the Canadian National

Collection of Insects, Arachnids and Nematodes (CNC), for providing me with data for more than half of the ~19,600 specimens used in this study.

Thanks to Dr. David McCorquodale, whose interest in lady beetles precipitated this study, for teaching me the finer points of insect collecting, specimen preservation and databasing; and Dr. Donna Giberson, for her seemingly endless patience and invaluable counsel. I would also like to thank Dr. Christian Lacroix and Dr. Christine

Noronha for taking the time to be a part of my committee.

A special thanks goes to my family and friends, who have given me the support I needed to persevere. I hope I have made them proud. TABLE OF CONTENTS

TITLE PAGE...... i CONDITIONS FOR THE USE OF THE THESIS...... ii PERMISSION TO USE POSTGRADUATE THESIS...... iii CERTIFICATION OF THESIS WORK...... iv ABSTRACT...... v ACKNOWLEDGEMENTS...... vii LIST OF TABLES...... x LIST OF FIGURES...... xii GLOSSARY...... xvi 1.0 INTRODUCTION AND LITERATURE REVIEW...... 1 1.1 INTRODUCTION...... 1 1.2 LITERATURE REVIEW...... 5 NATURAL HISTORY COLLECTIONS...... 5 COLLECTION FACILITIES, FUNDING AND ACCESS TO DATA...... 8 DIGITIZATION OF COLLECTION DATA TO USE IN DISTRIBUTION STUDIES...... 12 POTENTIAL BIASES THAT CAN AFFECT PATTERNS DERIVED FROM COLLECTION DATA ...... 17 APPLICATIONS OF COLLECTION DATA TO BIODIVERSITY STUDIES...... 21 LADY BEETLES, CONSERVATION CONCERNS AND COLLECTION DATA...... 23 NON-NATIVE SPECIES OF LADY BEETLES IN CANADA...... 27 NATIVE LADY BEETLES SPECIES OF CONSERVATION INTEREST...... 31 1.3 STUDY OBJECTIVES...... 37 1.4 LITERATURE CITED...... ,...... 39 DO SMALL COLLECTIONS ADD VALUE TO THE BIG PICTURE? AN EVALUATION OF ATLANTIC CANADIAN LADY BEETLE DATA RETRIEVED FROM NATURAL HISTORY COLLECTIONS THROUGHOUT EASTERN CANADA...... 50 ABSTRACT...... 51 2.1 INTRODUCTION...... 52 2.2 MATERIALS AND METHODS...... 55 2.2.1 SELECTION OF LADY BEETLE SPECIES FOR STUDY...... 55 2.2.3 DATASET PREPARATION...... 60 2.2.4 DATA MAPPING...... 61 2.2.5 DATA ANALYSES...... 61 2.3 RESULTS...... 64 2.3.1 TEMPORAL DIFFERENCES IN RECORDS AMONG COLLECTIONS...... 64 2.3.2 SEASONAL DIFFERENCES IN COLLECTION EFFORT...... 71 2.3.3 TAXONOMIC DIFFERENCES IN COLLECTING EFFORT...... 71 2.3.4 SPATIAL DIFFERENCES IN COLLECTING EFFORT...... 75 2.4 DISCUSSION ...... 80 2.5 LITERATURE CITED...... 86 USING OCCURRENCE DATA FROM COLLECTIONS TO EVALUATE RELATIVE ABUNDANCE AND GEOGRAPHIC RANGE OF LADY BEETLE SPECIES IN EASTERN CANADA...... 90 ABSTRACT...... 91 3.1 INTRODUCTION...... 92 3.2 MATERIALS AND METHODS...... 95 3.3 RESULTS...... 101 3.4 DISCUSSION...... 110 3.5 LITERATURE CITED...... 117 4.0 CONCLUSIONS...... 123 APPENDIX...... 125 LIST OF TABLES

Table 1.1 Native species and locations of lady beetles for which declines have been reported in Canada and the northern United States and the non-native species being implicated (based Table 2 from Marriott et al. 2009 and Table 2 from McCorquodale etal. 2011)...... 2

Table 2.1 Comparison of the numbers of Atlantic Canada records for 11 study species of lady beetles found in insect collections throughout eastern Canada. Refer to page 61 for definitions of collection categories ...... 56

Table 2.2 List of insect collections in eastern Canada which store Atlantic Canada specimens of the 11 study species of Coccinellinae (see Table 2.1). For each collection physically located in Atlantic Canada with >5 records, the radius that included 75% of records for these species is also given. Where available, collection abbreviations are from Insect and Spider Collections of the World http://hbs.bishopmuseum.org/codens/codens-inst.html ...... 58

Table 2.3 Changes in collecting effort by decade for four categories o f insect collections. a (*** p < 0.001; ** p < 0.01; * p < 0.05). Bolded observed values indicate those which are lower than expected based on the G-test ...... 66

Table 2.4 List of universities, colleges and museums with Atlantic Canadian lady beetle specimens, the year the institution was founded and the year the first Atlantic Canada specimen was collected ...... 67

Table 2.5 Summary results of statistical analyses for Mann-Whitney U- tests and Kruskal-Wallis and Dunn's Multiple Comparisons tests used to assess seasonal, temporal and taxonomic biases in four categories of collections ...... 70

Table 2.6 Kruskal-Wallis and Dunn's Multiple Comparisons test used to assess spatial differences in four categories of collections ...... 79

Table 3.1 Collections from which data for this study were assessed, including location, contact information of curator or collection manager (MM: Meghan Marriott; DBMcC: David McCorquodale; CM: Chris Majka; DD: Denis Doucet), and total number of specimens for Ontario, Quebec, New Brunswick, Nova Scotia and Newfoundland and Labrador for the 10 study species ...... 96

Table 3.2a Comparison of observed versus expected number of records for 10 species of lady beetles across six time periods based on the G-test.a ( * p < 0.001; ** p < 0.01; * p < 0.05). Bolded observed values indicate those which are lower than expected based on the G-test...... 106

x Table 3.2b Comparison of observed versus expected number of records for 10 species of lady beetles across six tim e periods based on the G-test (excluding Newfoundland and Labrador).a (*** p < 0.001; ** p < 0.01; * p < 0.05). Bolded observed values indicate those which are lower than expected based on the G- test ...... 107

Table 3.3 Absolute number of records (and percent) for each time period for ten lady beetle species in eastern Canada. Records are shown for all o f eastern Canada (Ontario - Newfoundland) and with Newfoundland and Labrador excluded in case of bias due to lower collecting effort there ...... 108 LIST OF FIGURES Figure 1.1 Lady beetle specimen storage, a) Storage for insect collection at Cape Breton University, b) A drawer of lady beetles from the collection at the University of Prince Edward Island, c) A lady beetle specimen with locality, determination and unique identifier labels (New Brunswick Museum) ...... 7

Figure 1.2 Insect labels associated with insect specimens in the UPEI collection and Canadian National Collection of Insects (CNC): a) coded locality/date label associated with a student insect survey of Prince Edward Island during 1983 (UPEI); b) coded locality/taxon label from a survey of tributaries along the Mackenzie River in NWT during 1972 from the Mackenzie Pipeline survey (CNC); and c) locality/date label from the Agriculture Canada farm site at Harrington, PEI in 1981 (UPEI); d) detailed locality/date label, including GPS coordinates, associated with a student lady beetle project at UPEI ...... 14

Figure 1.3 Coccinella undecimpunctata (sketch from Gordon 1985) ...... 27

Figure 1.4 Propylea quatuordecimpunctata (sketch from Gordon and Vandenberg 1991)...... 28

Figure 1.5 Coccinella septempunctata (sketch from Gordon 1985) ...... 28

Figure 1.6 Hippodamia variegata (sketch from Gordon and Vandenberg 1991) ...... 29

Figure 1.7 Harmonia axyridis (sketch from Gordon and Vandenberg 1991) ...... 30

Figure 1.8 Coleomegilla maculata lengi (sketch from Gordon 1985) ...... 31

Figure 1.9 Hippodamia parenthesis (sketch from Gordon 1985) ...... 32

Figure 1.10 Hippodamia tredecimpunctata tibialis (sketch from Gordon 1985) ...... 33

Figure 1.11 Adalia bipunctata (sketch from Gordon 1985) ...... 33

Figure 1.12 Coccinella novemnotata (sketch from Gordon 1985) ...... 34

Figure 1.13 Coccinella transversoguttata richardsoni (sketch from Gordon 1985) ...... 35

Figure 1.14 Coccinella trifasciata perplexa (sketch from Gordon 1985) ...... 35

Figure 1.15 Anatis mali (sketch from Gordon 1985) ...... 36

Figure 2.1 Total numbers of records for each decade and pattern of accumulation of records for the entire period for which records are available, separated by collection category and for all collections combined ...... 65

xii Figure 2.2 Differences in number of records reported from tw o time periods (before 1980 and after 1980) for four collection categories and combined collection data. Time periods which are significant at p < 0.05 are represented by 'a' (based on Mann-Whitney U-test, Statistica v.6). Refer to Table 2.3 for p values ...... 69

Figure 2.4 Differences in number of records of native versus non-native lady beetles for different collection categories and combined collection data. Records which are significant at p < 0.05 are represented by 'a' (based on Mann-Whitney U-test, Statistica v.6). Refer to Table 2.3 for p values ...... 73

Figure 2.5 Differences in number of records of native lady beetles reported from two time periods for different collection categories and combined collection data. Time periods which are significant at p < 0.05 are represented by 'a' (based on Mann-Whitney U-test, Statistica v.6). Refer to Table 2.3 for p values ...... 74

Figure 2.6 Differences in number of records of non-native lady beetles reported from two time periods for different collection categories and combined collection data Time periods which are significant at p < 0.05 are represented by 'a' (based on Mann-Whitney U-test, Statistica v.6). Refer to Table 2.3 for p values ...... 76

Figure 2.7 Spatial coverage for collection categories graphed by cumulatively adding number of counties represented in each category of collection in the following order: government, museum, university and private ...... 77

Figure 2.8 Differences in collecting area (Km2) which includes 75% o f records for each collection, grouped by collection category. Collection categories which are significantly different are represented by a, b or c (based on Kruskal-Wallis, Dunn's Multiple Comparisons Test, Statistica v.6). Refer to Table 2.3 for p values ...... 78

Figure 3.1a Relative abundance by time period for lady beetle species in eastern Canada 102

Figure 3.1b Relative abundance by time period for lady beetle species in eastern Canada (excluding NL & LB) ...... 103

Figure 3.2a Geographic range (km2) by time period for lady beetles in eastern Canada ...... 104 Figure 3.2b Geographic range (km2) lady beetles in eastern Canada (excluding NL & LB) ...... 105

Figure A -l Distribution of Adalia bipunctata (twospotted lady beetle) in eastern Canada from 1891 to 1959, and by decade from 1960 to 2009 ...... 126

Figure A-2 Distribution of Coleomegilla maculata lengi (spotted lady beetle) in eastern Canada from 1890 to 1959, and by decade from 1960 to 2009 ...... 127

Figure A-3 Distribution of Coccinella novemnotata (ninespotted lady beetle) in eastern Canada from 1881 to 1959, and by decade from 1960 to 2009 ...... 128

Figure A-4v Distribution of Coccinella transversoguttata richardsoni (transverse lady beetle) in eastern Canada from 1889 to 1959, and by decade from 1960 to 2009 ...... 129

Figure A-5 Distribution of Hippodamia tredecimpunctata tibialis (thirteenspotted lady beetle) in eastern Canada from 1885 to 1959, and by decade from 1960 to 2009 ...... 130

Figure A-6 Distribution of Coccinella septempunctata (sevenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 ...... 131

Figure A-7 Distribution of Coccinella undecimpunctata undecimpunctata (elevenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009...... 132

Figure A-8 Distribution of Harmonia axyridis (multicoloured Asian lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 ...... 133

Figure A-9 Distribution of Hippodamia variegata (variegated lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 ...... 134

Figure A-10 Distribution of Propylea quatuordecimpunctata (fourteenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 ...... 135 Figure A - ll Distribution of Adalia bipunctata (twospotted lady beetle) in eastern Canada from 1891 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador)...... 136 Figure A-12 Distribution of Coleomegilla maculata lengi (spotted lady beetle) in eastern Canada from 1890 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador) ...... 137 Figure A-13 Distribution of Coccinella novemnotata (ninespotted lady beetle) in eastern Canada from 1881 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador) ...... 138

xiv Figure A-14 Distribution of Coccinella transversoguttata richardsoni (transverse lady beetle) in eastern Canada from 1889 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador) ...... 139 Figure A-15 Distribution of Hippodamia tredecimpunctata tibialis (thirteenspotted lady beetle) in eastern Canada from 1885 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador) ...... 140

Figure A-16 Distribution of Coccinella septempunctata (sevenspotted lady beetle) in eastern Canada from 1885 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador) ...... 141

Figure A-17 Distribution of Coccinella undecimpunctata undecimpunctata (elevenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador) ...... 142

Figure A-18 Distribution of Harmonia axyridis (multicoloured Asian lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador) ...... 143

Figure A-19 Distribution of Hippodamia variegata (variegated lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador) ...... 144

Figure A-20 Distribution of Propylea quatuordecimpunctata (fourteenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador) ...... 145

XV GLOSSARY

INSECT MORPHOLOGY TERMS:

Elytra (singular elytron) - hardened forewings which act as a protective covering for the

flight wings in beetles.

Pronotum - the upper surface of the prothorax.

Prothorax - the anterior segment of the thorax (one of three); bearing the first pair of

legs.

Thorax - the part of the body between the head and the abdomen in an insect.

Tibia-th e fourth segment (from the body) of an insect leg.

MAPPING TERMS:

Georeferencing - the process of aligning geographic data to a known coordinate

system.

Holarctic - refers to the combined geographic regions which comprise the Palaearctic

and Nearctic.

N earctic- geographic region comprised of Greenland and North America north of the

tropic of cancer.

Palaearctic - geographic region comprised of Europe and northern regions of Asia and

Africa.

xvi 1.0 INTRODUCTION AND LITERATURE REVIEW

1.1 INTRODUCTION

Lady beetles are colourful and popular insects which are important ecologically as predators of other insects. Since the 1990s, lady beetles have prompted conservation concern with reports that some native species may be declining in distribution and/or relative abundance throughout North America (see examples and references in Table 1.1). Possible reasons for a decline include habitat alteration

(Harmon et al. 2007), parasites, pathogens, and parasitoids (Riddick e tal. 2009), competition from non-native lady beetles (e.g. Lucas etal. 2002, Michaud 2002, Cottrell and Shapiro-llan 2003, Yasuda e tal. 2004), and interspecific predation (Evans 1991).

However, before evaluating reasons for declines it is important to confirm that species are actually experiencing declines. Confirmation could be achieved through long-term surveys, but these data are rarely available for insects such as lady beetles, especially over broad spatial scales. Therefore, other methods must be employed. One of these that may well be suited to lady beetles is to compare historic ranges to present day ones using historical records.

However, determining historic ranges of species, especially insect species, is a continuing challenge. Many species are highly motile, with patchy distributions scattered across broad geographic ranges (Koch and Strange 2009). They can also be difficult to identify, so historic written records may not be reliable. Even where distribution maps do exist, they may be misleading, since traditional distribution maps

"connect the dots" between locations in which specimens have been collected, with the Table 1.1 Native species and locations of lady beetles for which declines have been reported in Canada and the northern United States and the non-native species being implicated (based Table 2 from Marriott et al. 2009 and Table 2 from McCorquodale et al. 2011).

Native species reported Province / State Non-native species being Sources to be declining ______implicated ______

Adalia bipunctata New Brunswick, Nova Coccinella septempunctata, Boiteau et al. 1999; Cormier et al. 2000; Scotia, South Dakota, Harmonia axyridis Elliott et al. 1996; Hesler et al. 2004; Michigan, Missouri Hesler and Kieckhefer 2008; Colunga- Garcia and Gage 1998; Fothergill and Tindall 2010 Coccinella novemnotata Ontario, Quebec, north Coccinella septempunctata Marshall 2008; Skinner and Domaine eastern United States, 2010; Wheeler and Hoebeke 1995; South Dakota, Missouri Harmon etal. 2007; Hesler and Kieckhefer 2008; Fothergill and Tindall 2010 Coccinella Manitoba, New Brunswick, Coccinella septempunctata, Turnock etal. 2003; Boiteau etal. 1999; transversoguttata Maine, South Dakota Harmonia axyridis, Boiteau 1983; Alyokhin and Sewell 2004; richardsoni Propylea Finlayson etal. 2008; Elliott etal. 1996; quatuordecimpunctata Hesler et al. 2004; Hesler and Kieckhefer 2008

Coccinella trifasciata Manitoba Coccinella septempunctata Turnock etal. 2003 perplexa

Continued...

2 Table 1.1 Continued, Native species reported to Province / State Non-native species being Sources be declining implicated Cycloneda munda Michigan Harmonia axyridis Colunga-Garcia and Gage 1998 Hippodamia convergens Manitoba, Michigan, Coccinella septempunctata, Turnock et al. 2003; Colunga-Garcia and Pennsylvania, Harmonia axyridis Gage 1998; Wheeler and Stoops 1996; Connecticut Ellis et al. 1999 Hippodamia parenthesis Manitoba Coccinella septempunctata Turnock etal. 2003 Hippodamia Quebec, New Coccinella septempunctata Lucas et al. 2007; Boiteau 1983; Boiteau tredecimpunctata tibialis Brunswick, Midwestern etal. 1999; Obrycki etal. 2000; Alyokhin United States, Maine and Sewell 2004; Finlayson et al. 2008

3 underlying assumption that the species' distribution is continuous between points

(McCorquodale et al. 2011). Most species have habitat associations or ecological niches which are not continuous, so such maps should be used with caution when looking for range changes over time.

More accurate distribution maps can be produced from locality data from labels on specimens in natural history collections within museums, government facilities, universities and even private homes. These specimen records represent many years of collecting effort from both amateur and professional collectors, and can date back hundreds of years, depending on the taxonomic group and the location of the museum

(Lovejoy et al. 2010). Historical collection data have provided the basis for most of what we know about biodiversity and geographic ranges of flora and fauna worldwide

(Johnson 2007). Collection data can also be used to infer how distribution patterns change over space and time (e.g. Newbold 2010, Elith et al. 2006, Guralnick and Van

Cleve 2005, McCarthy 1998). However, natural history collections can contain gaps in data which may result in biased interpretations of distribution patterns. These include gaps in what taxa are collected, when they are collected, and how they are collected

(Newbold 2010), and the potential for data gaps needs to be considered before these data should be used for in any sort of analysis.

Lady beetles are well represented in insect collections thanks to their showy colours and popularity among collectors, and Canadian collection records for lady beetles date back to the 19th Century (Gordon 1985). Therefore, they should be a good

4 taxon for which to use collection data to assess distribution patterns, as long as potential gaps in data do not cause collection biases.

The main objective of this study is to use historical collection data to determine whether lady beetles (both native and non-native) are changing in distribution and/or relative abundance in eastern Canada over the last ~100 years. Since collection data are known to have gaps that might bias interpretation of distribution patterns, a secondary objective is to investigate the biases in different types of collections, to assess the usefulness of the collections for evaluating distribution patterns in lady beetles.

1.2 LITERATURE REVIEW

NATURAL HISTORY COLLECTIONS

Determining historic ranges of species which are currently thought to be declining is a challenge for scientists worldwide (Shaffer et al. 1998), but especially for insect species, because of their high motility, patchy distributions, and difficulty in identification (Koch and Strange 2009). The pattern for historical distribution maps to just "connect the dots" between the locations in which specimens have been collected

(McCorquodale et al. 2011) can lead to errors in interpretation of range data because of missing data on habitat associations or ecological niches. Therefore, obtaining as much information as possible about species ecology and geographic ranges is an important, but difficult, first step in conservation planning (Ferrier 2002, Funk and Richardson 2002,

Rushton et al. 2004, Newbold 2010). In at least some cases, though, information

5 associated with specimens collected over broad areas and time periods, and stored in natural history collections can be mined to infer spatial and temporal patterns.

Natural history collections throughout the world contain >2.5 billion preserved biological specimens (Cotterill 1995) which have been retained from taxonomic, ecological, and biodiversity studies (Danks 1991, Cotterill 1995). These collections represent many years of collecting efforts from both amateur and professional collectors and date back as early as the beginning of the nineteenth century (Lovejoy et al. 2010) (Figure 1.1). Most of what we know about biodiversity and geographic ranges of flora and fauna worldwide comes from the studies represented by these collections

(Johnson 2007).

More than 120 biological collections exist in Canada, ranging in size from several hundred to 16.7 million specimens (Lovejoy et al. 2010). According to a recent survey conducted by the Government of Canada, the largest Canadian collections are maintained by the federal government or universities and include the Canadian National

Collection (CNC) of Insects, Arachnids and Nematodes in Ottawa (16.7 million specimens); the Canadian Museum of Nature, Ottawa (7.4 million specimens); the

University of Alberta, Edmonton (3.5 million specimens); McGill University, Montreal

(3.4 million specimens); the Royal Ontario Museum, Toronto (3.2 million specimens); the University of Guelph (2.5 million specimens); Universite de Montreal (2.5 million specimens); University of Manitoba, Winnipeg (2.1 million specimens); and the

University of British Columbia, Vancouver (2.1 million specimens). Collectively, these large collections house representative specimens of most of the flora and fauna found in c)

Figure 1.1 Lady beetle specimen storage, a) Storage for insect collection at Cape Breton University, b) A drawer of lady beetles from the collection at the University of Prince Edward Island, c) A lady beetle specimen with locality, determination and unique identifier labels (New Brunswick Museum).

7 Canada (Wiggins etal. 1991). However, small regional collections in universities,

research facilities, museums, non-government facilities, ecological preserves and private

homes (Wiggins etal. 1991, Chapman 2005) provide additional security by having specimens located in multiple locations in the event of a disaster (e.g. fire or flooding)

(Lovejoy et al. 2010). Access to the collections to study the specimens can vary among the collection types. The large collections are used mainly fo r research, but require considerable planning to obtain permission to visit. In contrast, small collections can be easier to enter and work in than large ones, but may be difficult to travel to, so are frequently used for teaching and local research purposes (Lovejoy et al. 2010). Small collections usually also have a regional focus, so may have specimens which represent flora and fauna at specific local scales more completely than the larger scale collections.

They therefore provide important information on baselines to document changes in populations (Lovejoy etal. 2010, Snow 2005, Danks 1991, Johnson 2007, Wiggins etal.

1991).

COLLECTION FACILITIES, FUNDING AND ACCESS TO DATA

Canadian collections are housed in a variety of institutions which can be grouped into four broad categories: Federal Government, Other Government, University, and

Other Collection, such as those in homes of private collectors (Lovejoy et al. 2010). A recent survey conducted by the Government of Canada (Lovejoy et al. 2010) found that

43% of Canadian natural history collections are housed in universities or colleges, while

8 22% of collections are held in federal government facilities, 17% in provincial government facilities, and 18% in other facilities.

The different types of collections focus on different taxa, habitats, and localities, depending on their mandate, sources of funding, and location. Federal government collections in Canada range from the large Canadian National Collection of Insects in

Ottawa (CNC) with a broad taxonomic and geographical focus, to smaller collections associated with the focus (e.g. agriculture, forestry, health) o f particular departments.

Federal government collections are usually also research-based collections, but they can vary in their management. For example, the CNC collection is maintained and accountable to the federal government through a board of trustees (Lovejoy et al.

2010). The CNC was established by the Department of Agriculture, with an original mandate to preserve specimens of Canadian crop and livestock pest insects, but it has expanded to house representative arthropod species from numerous government sectors (Lonsdale and Huber 2011). In contrast) the other federal government collections are usually smaller in scope and in size than the CNC. For example, the regional federal Agriculture and Forestry research labs are administered locally and focus on their specific research goals, and national museum collections (such as the

Canadian Museum of Nature) reflect the expertise of the staff. The Provinces and

Territories may also maintain collections in their museums, and where present, these collections are administrated by the respective provincial governments (Lovejoy etal.

2010). Provincial and Territorial collections are used for a wide range of purposes which include research, teaching and interfacing the scientific community with the public. University collections are primarily teaching and research-based collections of modest size, with the exception of a few large collections which have the added value of public exhibits and outreach programs (Lovejoy et al. 2010). University collections generally reflect the research and teaching interests of their faculty and staff (Lovejoy et al. 2010).

All of these collection types may receive donations of specimens from other collections, broadening their focus. In addition, universities benefit from broad-scale collecting by students, combining specialist graduate-level student collecting with that of untrained students who may focus on the larger and more conspicuous charismatic taxa

(McCorquodale et al. 2011). The final category of collections, "Other collections", includes private, non-profit and hospital (microbial) collections, and usually reflects collecting focus of a single collector or study goal (Lovejoy et al. 2010).

Curation and collection maintenance are on-going challenges for natural history collections, and the sources and levels of support for these operations also varies with collection type. Operational funding including salaries and infrastructure is generally provided by the institution which houses the collection. Consequently, only the largest collections tend to have paid curatorial staff, and small museums rely either on volunteers or researchers to curate material in the collection (Lovejoy et al. 2010).

Some provincial or territorial museums have operational budgets that cover cabinets and general maintenance whereas smaller institutions may rely on external funding and donations to remain operational. Private and non-profit collections receive the least amount of external support (Lovejoy et al. 2010).

10 The amount of funding and curatorial support a collection facility receives directly impacts the preservation and quality of the biological specimens available for study. Small regional collection facilities have limited storage space which restricts the number of specimens which can be preserved, forcing curators to make difficult decisions about which taxa are kept or maintained. Older facilities may also have antiquated cabinets which do not seal to keep out pests (e.g. dermestid beetles which feed on dead organic material in museums), leading to damaged or missing specimens

(Snow 2005, O'Connell et al. 2004). All collections have a backlog of unidentified material, but small collections may be dominated by unidentified material due to lack of staffing and/or expertise, outdated equipment and/or taxonomic literature, and maintenance issues (such as remounting specimens for identification) (Snow 2005,

Cotterill 1997). The financial issues can be critical, since many facilities have received significant cuts to funding in recent years (Gropp 2003, Saurez and Tsutsui 2003, Snow

2005, Gabel et al. 2007, Lovejoy et al. 2010), forcing them to make difficult decisions about their management. For example, funding constraints may force collections to narrow their focus to specific taxa or project goals, and small regional facilities may have to downsize or close completely (Snow 2005). When the small collections close, they may be able to transfer specimens to larger institutions, since most will accept donations of curated material. However, not all the material in small collections is curated, and Gabel et al. (2007) argue that removing smaller regional collections from their original geographic context considerably depreciates the value of the collections.

11 The value of the collection for assessing distribution patterns is also dependent on the accessibility of the data, and access is one of the main challenges associated with using collection data in biodiversity and population studies. According to Lovejoy et al.

(2010), only 61% of specimens held in Canadian collections are accessible. The remaining specimens are in variable stages of preservation, ranging from needing to be curated, needing to be catalogued and needing to be integrated into collections. Access to specimens may also be restricted for reasons such as the protection o f specimens representing rare and endangered species, or issues relating to ownership rights to the data (Williams et al. 2002). One way to improve access to the data is to digitize the information from the collection labels, providing both a digital record and a unique identifier that can be associated with particular specimens (Wheeler et al. 2001).

However, despite a steady increase in the number of collections that have been digitized, only 5-10% of data from specimens in collections worldwide have been made available electronically (Graham et al. 2004), mainly because facilities lack the resources to digitize their collection records (Snow 2005, O'Connell et al. 2004). Digital data also need to be checked carefully for errors, both in data entry and in specimen identification, before their use in distribution studies.

DIGITIZATION OF COLLECTION DATA TO USE IN DISTRIBUTION STUDIES

Specimen data are used to track species distribution patterns by translating the locality and date from the specimen labels to "dots on the map" for different time periods. In the past, mapping of specimen records was done manually, but computer

1 2 mapping programs can now be used to analyse patterns in distribution, area of occupancy, and other parameters. These methods require that the data be digitized to be useful. The potential for errors in digital records can be high, however, especially if the data entry technician is not familiar with the taxonomic group or collection localities. For example, specimens may be misidentified, so entering label data verbatim can result in incorrect species names becoming associated with a specimen in a database. Other errors can include errors in interpreting dates or localities reported on labels, and misfiling of specimens into incorrect taxonomic groupings (Graham et al.

2004, Newbold 2010). Misidentification of specimens due to human error and/or outdated taxonomy is a particular challenge because it can only be detected through the careful examination of individual specimens. The process of verifying identifications can be tedious and time consuming, but the ability to re-examine specimens to confirm or update the species information is one advantage associated with using collection data (Graham etal. 2004, Newbold 2010).

Even if label data have been digitized, considerable work can still be required to make the data useful for computer mapping and analysis. Errors or misinterpretation of dates or locality information on labels may occur due to transcription errors on the labels themselves (when data are transcribed from field note books), or simply from changes in label standards that result in different information being recorded (Wheeler et al. 2001). For example, early labels may give locality information based on project codes, road mileage, or certain landmarks or place names which can change overtim e

(e.g. Figure 1.2 a, b), whereas most recent labels will contain precise geographical

13 a)

d) CAN;NS:Kings Co. Grand Pr4 45.096984°^, -64.303564^ August 2009 Feeding on unripe mulberries L. Marriott

14 locations obtained from hand-held GPS units (though see below for discussion on potential errors from GPS readings) (Wieczorek et al. 2004). Locality/date labels should contain a full textual description of where and when the specimen was collected (e.g.

Figure 1.1 c; Wheeler et al. 2001) so that changes to place names do not result in inaccuracies when georeferencing and for mapping (Murphey et al. 2004). Date codes can also cause confusion if not used consistently, and especially in long-term studies,

incomplete dates (e.g. '01' for 1901) may be misinterpreted (e.g. confused with 2001).

Other labels, particularly those associated with ecological surveys and those in collections without a curation manager, may consist of a code number associated with a field note book, making it even more difficult to determine patterns in species distribution. Over time, the "code books" may be lost or discarded, along with the

information needed to make use of these specimens in population studies.

Examples of different types of coded or incomplete locality labels can be seen in

Figure 1.2. Figure 1.2a shows a reference code on a label in the University of Prince

Edward Island (UPEI) collection giving a 2-part code: the first part is the year ('83' =

1983) and the second part is a number that corresponds to a detailed record in a field note book from a PEI insect survey. In this case, the field note book was lost as a result of a building fire at UPEI in 1994 (D. Giberson, UPEI, personal communication), so considerable local research was needed to even determine that the locality was in

Prince Edward Island, and during that year. In contrast, collection notes associated with the specimen label pictured in Figure 1.2b were published in a government technical report (Brunskill et al. 1973), so are still available. However, decoding the label

15 information is necessary (and time consuming) before specimen data can be analysed.

In this case, the site (TR = Trail River, NWT) was given in one place in the report, so maps can be used to determine the locality, and the sampling dates were explained in another place in the report (the last six digits are a date code, corresponding to Nov. 23,1972).

The sampling methods were described elsewhere in the report and correspond to other parts of the coded label (AB = artificial basket; "oil" refers to sample collected in zone contaminated with oil). The "IS" on the label is not explained, so has not been decoded. Figure 1.2c shows a third problem: a label with an incomplete textual description of a collection site. Though the label provides a place name ("Harrington"), no province is indicated and the Geographical Names of Canada gazetteer

(http://www.nrcan.gc.ca/earth-sciences/search/search_e.php) reveals there are currently four places called Harrington in Canada (in Prince Edward Island, Quebec and

Ontario), and one formerly named Harrington in Nova Scotia. As long as the specimen remains associated with the UPEI collection, the PEI location can be inferred, but if specimens are transferred then this geographical information may be lost, or interpreted incorrectly. All these label types present challenges in deriving useful information from preserved specimens.

The greatest challenge in mapping and analyzing distribution data from such highly variable and imprecise locality labels occurs when allocating geographical coordinates or "georeferencing" (Peterson et al. 2004, Murphey et al. 2004, Graham et al. 2004). Georeferencing can be time-consuming and difficult, and has no standardized guidelines (Wieczorek et al. 2004, Guralnick et al. 2006, Newbold 2010). Imprecise

16 georeferencing can cause false interpretations when studying distributions, extinctions, species richness and community composition (Graham et al. 2008, Rowe, 2005, M iller et al. 2007, Tingsley and Beissinger 2009). Contemporary collectors recognize the value of having geographic coordinates on labels, and include them as common practice because devices to collect this information (handheld GPS or global positioning system devices) are inexpensive and easy-to-use (Wieczorek etal. 2004). Handheld GPS units have not eliminated georeferencing problems, though, since many collectors do not record necessary information such as map scale, coordinate system, datum, and accuracy while collecting coordinates, which can cause spatial errors, particularly when combining modern data with historical records (Rowe 2005).

In addition to the challenges discussed regarding the digitization of collection data, there may be temporal or spatial gaps in specimen records that can bias interpretation of distribution patterns from collection data alone (Chapman 2005,

Newbold 2010). Newbold (2010) states that when used appropriately, collection data can be an invaluable resource for evaluating spatio-temporal distribution patterns.

However, Chapman (2005) argues that it is important to establish what biases are present in the data before deciding on its usefulness.

POTENTIAL BIASES THAT CAN AFFECT PATTERNS DERIVED FROM COLLECTION DATA

Newbold (2010) lists four major interpretation biases that can be associated with gaps in records for specimen data in natural history collections: 1) Taxonomic bias, where a collection may focus on certain taxa; 2) Spatial bias, where a collection may

17 focus on particular locations; 3) Environmental bias, where the collection may focus on particular habitats or seasons to the exclusion of others; and 4) Temporal bias, where a collection may focus on certain tim e periods. These potential gaps in space or time covered by the collection can vary with time and with collection type, depending on financial resources, the background of collectors contributing to the collection, and the mandate of the collection.

Collections may show taxonomic bias when certain taxa are preferentially collected because they are of interest to the collector either personally or professionally

(Newbold 2010), because different sampling methods target different taxa, or due to the rarity of a species. A collection maintained by an agricultural research lab, for example, will be dominated by taxa from agricultural habitats. Similarly, collections with specialists on particular taxonomic groups should include good representation of those groups. Even general surveys can be biased towards certain taxa, since not all sampling methods sample all groups equally. For example, insects may be collected o using different types of bulk sampling methods that target different habitats (e.g. sweep netting for insects on vegetation, and leaf litter sampling for litter invertebrates), or behaviours (e.g. Malaise traps for flying insects, pitfall traps fo r walking insects) (Danks

1991), but no method targets taxa equally. This type of bias usually cannot be evaluated since specimen labels rarely include information about collection method, so absence of certain species from a collection or tim e period cannot be taken as proof of absence from the locality.

18 As with taxonomic bias, spatial biases depend heavily on the type of collection being considered. Collections with a particular mandate, such as agriculture or forestry, will focus on agricultural or forest localities, with relatively poor representation from other habitats. Even in more general collections, specimens are often collected opportunistically based on the collecting opportunities presented to the collectors

(Margules and Austin 1994, Williams et al. 2002, Graham et al. 2004, Elith et al. 2006).

This results in domination by specimens which have been collected from easily accessible sites such as close to the homes of the collectors, along roads and rivers, and near towns or research stations. Other specimens may have been collected for specific projects, for example from known biodiversity hotspots or ecological niches (Hijmans et al. 2000, Soberon et al. 2000, Reddy and Davalos 2003, Williams et al. 2002,

Randrianasolo et al. 2002, Graham et al. 2004, Pyke and Ehrlich 2010).

Collections may also show gaps in specimen records due to environmental bias, or sampling bias that results from neglecting certain inaccessible habitats or seasons, due to difficulty in sampling them. Even when sampling is systematic, it is a challenge to collect without environmental biases. For example, the geographic distribution of an organism may cover a wide range of environmental gradients in variables such as temperature, precipitation and elevation (Newbold 2010) but individuals living at the extremes of their distribution may be underrepresented (e.g. the uppermost extremes of an elevation or latitudinal gradient may be virtually inaccessible without expensive resources). Collection data may also be seasonally biased due to targeting certain life cycle stages which can be easily identified or collected (Peterson etal. 1998 IN Newbold 2010). For example, many insect collections are dominated by adult butterflies, which are obvious in summer when collectors are in the field (Newbold 2010).

It is also rare to find consistent collecting over long periods of tim e for species,

resulting in temporal gaps in specimen records. This results in a temporal bias, where different taxa may be collected or retained at different time periods, as collectors'

interests change or they retire, as the mandate of the collection changes, or as collection space fills up (Newbold 2010). Workers in well-curated collections may stop

retaining common specimens once they are well represented in their collection, subsequently focusing on rarer or new taxa. If specimen records for certain species suddenly decline or disappear in a collection, this trend might be confused with actual changes in range or abundance of a species when collection data are mapped (McCarthy

1998). It is nearly impossible to remove temporal biases from collection data, though

research into a collection's history can help to determine whether it exists. However, combining data from multiple collections can reduce errors caused by temporal bias by

increasing the number of specimens collected from the same localities over time

(Guralnick and Van Cleve 2005, Pyke and Ehrlich 2010). For example, university student collections may continue to acquire common taxa as students make course-based collections, but their collections may miss rare or obscure taxa. By combining broad and on-going types of collections with more focused ones, a more complete picture may appear.

These patterns indicate that although collection data can be valuable in looking at current and past patterns in species distribution and abundance, characteristics of

2 0 individual collections must be considered when using these for assessment of species distributions. Since a single collection is rarely representative of the abundance and distribution patterns in a location for each collecting period, it is important to determine the extent and types of gaps that exist for each type of collection, so that assumptions can be corrected to prevent or mitigate biases. Many of the concerns considered in this section are especially pertinent to insect collections due to the relatively large number of specimens, and the high diversity of the groups found in the collections.

APPLICATIONS OF COLLECTION DATA TO BIODIVERSITY STUDIES

Once label data have been deciphered, databased, and georeferenced, they can be analyzed to track the arrival of non-native species, monitor distributions of endangered species, or carry out studies of long term distribution patterns. The number of studies using collection data to address various ecological questions has grown considerably since the mid-1990s, and covers a wide variety of taxa. For example, herbarium records have been used to reconstruct the spread of invasive plant species

(Delisle et al. 2003, Lavoie et al. 2007), as well as to assess their impact on plant biodiversity in Quebec (Lavoie etal. 2003). Mammal collection records have been used to document changes in relative abundance and distribution of woodland rodents and opossums in the northern Great Lakes region (Myers etal. 2009), to infer conservation status of marsupials and monotremes in south-west Western Australia (McCarthy 1998), and to document the decline of prairie habitat in the Chicago region (Pergams and

Nyberg 2001). Amphibian species declines have been assessed in Great Central Valley,

2 1 California using collection records (Fisher and Shaffer 1996), and collection data were used to model habitat suitability of Erhard's wall lizard in Crete (Herkt 2007).

Insect collections provide useful data to evaluate patterns of distribution and species diversity, since they are usually composed o f large numbers of specimens collected over a long time period. For example, insect records were used by Meier and

Dikow (2003) to estimate species richness of robber-flies and McCorquodale et al.

(2007) used collection records to assess changes in species richness in long-horned wood boring beetles in Ontario. Collections have also provided data on rare and invasive bees in a series of studies aimed at confirming anecdotal impressions about declines, predicting sites to search for rare species, and documenting invasions of various bee species. Declining bumblebee species have been documented from collection data in Ireland by Fitzpatrick et al. (2007), in Ontario by Colla and Packer

(2008), and in British Columbia by Colla and Ratti (2010). Recently, Koch and Strange

(2009) used bumblebee records and climatic/habitat data from North America to generate probable historical range maps to aid monitoring programs for species at risk

(Koch and Strange 2009). Suitable habitat for several non-native species of bees has also been modelled based on collection records from the bees' native ranges to predict probable distributions should the bees become established in the novel region

(Hinojosa-Diaz et al. 2005, Hinojosa-Diaz etal. 2009). This approach was pioneered by

Sutherst etal. (1996) who used CLIMEX, a computer program for comparing climatic variables and species occurrence data within a geographical context, to examine the potential spread o f the cane toad in Australia. Samways et al. (1999) predicted

2 2 geographical ranges of lady beetles (Coccinellidae, Chilocorus spp.) worldwide using

CLIMEX, and though the findings of this study have been questioned by Sutherst (2003), it supports the idea that lady beetles are good candidates fo r these types of analyses.

The studies mentioned above highlight the emergence of new modeling and analytical techniques which are greatly expanding the scope and applications of collection-based occurrence data. For example, Tingley and Beissinger (2009) have created two occupancy models which allow for the unbiased comparison of historical and modern species occurrence data, and suggest methods for measuring changes in range. Elith et al. (2006) compared 16 different modeling programs fo r assessing > 200 species of plants, birds and mammals from six regions of the world to determine which modeling methods were the most effective when using collection (occurrence) data. As with the bumblebee study reported above (Koch and Strange 2009), these models combine occupancy data with other data to predict occurrence for un-sampled localities, to guide current sampling programs. They can also be used to predict species richness; Guralnick and Van Cleve (2005) used a modeling approach to compare collection data, survey data, and combined collection and survey data to predict species richness for birds in north-eastern Colorado.

LADY BEETLES, CONSERVATION CONCERNS AND COLLECTION DATA

Lady beetles, also commonly known as ladybugs, ladybirds and ladybird beetles, . are an example of an insect (like the bumblebees listed above) which has been reported to be experiencing declines in range and abundance (Table 1.1). Proposed reasons for

23 the decline include habitat alteration (Harmon e tal. 2007), parasites, pathogens, and parasitoids (Riddick et al. 2009), competition from non-native lady beetles (e.g. Lucas et al. 2002, Michaud 2002, Cottrell and Shapiro-llan 2003, Yasuda etal. 2004), and interspecific predation (Evans 1991). However, information on lady beetle declines remains largely anecdotal except in a few cases, so more data are needed to determine which, if any, species may be declining before focusing on causes. Since lady beetles are well represented in insect collections because of their bright colours and their presence in habitats frequented by insect collectors, natural history collections are good potential sources of material for investigation. Not only are lady beetles common in insect collections of all types, there is a long record of lady beetle collection in Canada.

Therefore, there is potential to use collection data to assess distribution patterns, if possible biases can be identified and accounted for by using specimen information in conjunction with taxonomic and ecological information (Graham etal. 2004, Newbold

2010).

Lady Beetle Biology

Lady beetles are in the family Coccinellidae, with the larger, brightly coloured lady beetles belonging in the subfamily Coccinellinae. Other subfamilies of

Coccinellidae, such as the Scymninae, are also present in Canada, but are less well known because they tend to be smaller, less colourful beetles than the coccinellines.

Overall, 481 species of Coccinellidae in 60 genera have been reported in North America with 166 species occurring in Canada. Fifty-five of these 166 Canadian species are in the

24 subfamily Coccinellinae, and represent 19 genera (Gordon 1985, Gordon and

Vandenberg 1991).

Lady beetles are predatory, both in the larval and in the adult stages. Most lady beetles feed on slow moving and plant sucking herbivorous insects such as aphids, adelgids, psyllids, mealy bugs, and scale insects (Insecta: Hemiptera). Aphids are often the preferred food source, since the aphids can occur in large numbers and are easy to capture and consume (Acorn 2007). Although lady beetles are generalist predators, they may specialize on residents of certain habitats such as open fields with herbaceous or annual flowering plants, forest zones, or agricultural fields (Honek 1985). This wide feeding habitat range means that lady beetle specimens may be found in a wide variety of collections, including agricultural and forestry research stations, as well as regional and university museums.

The success of lady beetles at preying on aphids has led to intentional releases

(Obrycki and Kring 1998) of many non-native lady beetle species as biological control agents in North America, dating back as far as the late 1800s (Gordon 1985). Some of the studies into lady beetle declines have, in fact, blamed competition with these non native "invaders" for the observed declines. However, few of these intentional introductions have led to established populations (Gordon 1985), and the five non native Coccinellinae that have become common and widespread in Canada (as least for some time periods) are the result of unintentional introductions. The earliest establishment of a non-native species in Canada is reported to have occurred in New

Brunswick and Prince Edward Island in 1939 (Brown 1940; Coccinella u.

25 undecimpunctata Linnaeus) and the most recent was in 1994 in Ontario (University of

Guelph natural history collection; Harmonia axyridis (Pallas)). Three of the five common and widespread non-native lady beetle species in North America have been associated, at least anecdotally, with the declines of some native species (Table 1.1).

M arriott etal. (2009) conducted a preliminary literature assessment of lady beetle species in Canada, and identified eight species of lady beetles native to eastern

Canada that show different patterns relative to presence of the most common non native species: a) those experiencing declines in relative abundance and/or geographic distribution, b) those experiencing increases in relative abundance and/or geographic distribution, or c) those that are thought to be unchanged (so may serve as a "control" for assessing other patterns). These have therefore been selected for study of long term patterns using insect collections, along with the five non-native species of the subfamily

Coccinellinae currently established in Canada. Specific information on each species can be found in the next sections (illustrations and descriptions of each species are from

Gordon 1985 and Gordon and Vandenberg 1991, and are used with permission from the

New York Entomological Society). Official common names of the Entomological Society of Canada (ESC) and Entomological Society of America (ESA) are used when available; alternative common names from Acorn (2007), Devries (2007), Eaton and Kaufman

(2007), Marshall (2008) and BugGuide (VanDyk 2009) are noted where applicable.

2 6 NON-NATIVE SPECIES OF LADY BEETLES IN CANADA

There are currently five species of non-native lady beetles established in Canada:

Coccinella undecimpunctata undecimpunctata Linnaeus, Propylea quatuordecimpunctata (Linnaeus), Coccinella septempunctata Linnaeus, Hippodamia variegata (Goeze), and Harmonia axyridis (Pallas).

Coccinella undecimpunctata undecimpunctata Linnaeus

Official common name: Elevenspotted lady beetle (ESC) Alternate common name: None Description: A 4.0 - 5.0 mm beetle with red-orange elytra bearing eleven black spots, five on each elytron and one central spot near the pronotum. Biogeographic region: Palaearctic Habitat: Agricultural areas, seashore grassy communities and Figure 1.3 Coccinella parklands (Belicek 1976). undecimpunctata Distribution in Canada: British Columbia and Ontario to (sketch from Gordon Newfoundland and Labrador. 1985). Colonization history: This was the first Palaearctic subspecies of lady beetle to become established in North America following an accidental introduction in Massachusetts in 1912 (Schaeffer 1912). It became widespread in North America, spreading along the Atlantic seaboard and the St. Lawrence River waterway from Ohio into the Maritime Provinces and Newfoundland (Watson 1979, Wheeler and Hoebeke 1981, Gordon 1985). In Canada, the elevenspotted lady beetle was first recorded from New Brunswick and Prince Edward Island in 1939 (Brown 1940), Nova Scotia in 1945 (Majka and McCorquodale 2006), Ontario in 1960 (Hicks 1971), and Quebec in 1968 (Wheeler and Hoebeke 1981). Brown (1940) reports an earlier record for Quebec; however, there is no date of collection associated with the specimen. A separate and unrelated introduction occurred on the west coast of North America in Washington and southern British Columbia in the 1970s (Watson 1979, Belicek 1976). Notes: There are no reports of this lady beetle directly affecting natives; however, since the 1980s, it has not been recorded in surveys throughout the eastern United States, which coincided with the arrival of Coccinella septempunctata (Harmon etal. 2007).

27 Propylea quatuordecimpunctata (Linnaeus)

Official common name: None Alternate common name: Fourteenspotted lady beetle (VanDyk 2009) Description: A 3.5-5.2 mm beetle with yellow elytra bearing variable black markings (somewhat resembling a checkerboard). Biogeographic region: Palaearctic Figure 1.4 Propylea Habitat: Meadows, bogs, grasslands and agricultural areas quatuordecimpunctata (Kuznetsov 1997). (sketch from Gordon Distribution in Canada: Ontario to Nova Scotia and Prince and Vandenberg 1991). Edward Island. Colonization history: The first established populations of this Palaearctic species were recorded in Quebec in 1968 (Chantel 1972), Maine in 1988 (Day etal. 1994), and from southern New York.in 1989 (Yanega 1996). It was first recorded in NS in 1990 (McCorquodale 1998), and by 1995 it was being collected from all three Maritjme Provinces (Hoebeke and Wheeler 1996). Range expansion for this lady beetle has been relatively slow compared to other non-native species (McCorquodale 1998, Obrycki et al. 2000). Notes: There are no reports of this lady beetle directly affecting natives; however, in Maine, a highly significant positive correlation was found between high densities of this species and two other non-native species ( Coccinella septempunctata and Harmonia axyridis) (Alyokhin and Sewell 2004).

Coccinella septempunctata Linnaeus

Official common name: Sevenspotted lady beetle (ESC, ESA) Alternative common name: Seven-spot ladybug (Acorn 2007) Description: A 6.5-7.8 mm beetle with orange-red elytra bearing seven black spots, three on each elytron and one central spot near the pronotum. Biogeographic region: Palaearctic Habitat: Grasslands, agricultural areas and occasionally in arboreal environments (Kuznetsov 1997). Figure 1.5 Coccinella septempunctata Distribution in Canada: British Columbia to Newfoundland and (sketch from Gordon Labrador. 1985). Colonization history: This Palaearctic species was introduced

28 multiple times into North America between 1955 and 1971 for biological control of aphids (Gordon 1985). It became established through an unintentional introduction in 1973 in New Jersey and likely others in the early 1970s. In Canada, it was first found in New Brunswick in 1972, therefore an independent, unintentional introduction (Majka and McCorquodale 2010), Quebec in 1973 (Larochelle 1979), Prince Edward Island in 1982 (Majka and McCorquodale 2006), Nova Scotia in 1982 (Majka and McCorquodale 2010), Manitoba in 1988 (Turnock e ta l. 2003) and British Columbia in 1990 (Humble 1991). Notes: There have been noticeable declines in the abundance of the natives Coccinella novemnotata, Coccinella transversoguttata richardsoni, Adalia bipunctata, Hippodamia convergens, Hippodamia parenthesis and Hippodamia tredecimpunctata tibialis since the arrival of the sevenspotted lady beetle (Wheeler and Hoebeke 1995, Alyokhin and Sewell 2004, Elliott et al. 1996, Ellis et al. 1999, Finlayson etal. 2008, Hesler and Kieckhefer 2008, Turnock etal. 2003, Brown and Miller 1998, Obrycki et al. 2000, Elliott et al. 1998, Boiteau etal. 1999). In contrast, numerous reports suggest that some native lady beetles such as Coleomegilla maculata lengi, continue to coexist with this non-native lady beetle (Elliott et al. 1998, Obrycki et al. 1998b, Lucas et al. 2007).

Hippodamia variegata (Goeze)

Official common name: None Alternative common names: Variegated lady beetle (VanDyk 2009); Adonis' ladybird (VanDyk 2009); white collared ladybird (VanDyk 2009); Russian wheat-aphid lady beetle (VanDyk 2009); spotted amber lady beetle (Marshall 2008) Description: A 4.4-5 mm beetle with orange elytra bearing variable black spots. It is frequently misidentified as the larger, similar looking, Hippodamia convergens Figure 1.6 Hippodamia (convergent lady beetle), which lacks the raised margin on variegata (sketch from Gordon and its pronotum. Vandenberg 1991). Biogeographic region: Palaearctic Habitat: Grasslands and some types of arboreal environments (Kuznetsov 1997). Distribution in Canada: Ontario to Nova Scotia and Prince Edward Island. Colonization history: This Palaearctic species was introduced into North America many times between 1957 and 1983 as a biological control agent for aphids, but it did not become established in Canada until an inadvertent introduction in 1984 in Quebec

29 (Gordon 1987, Gordon and Vandenberg 1991). Range expansion for this lady beetle has been relatively slow compared to other non-native species (McCorquodale 1998, Obrycki et al. 2000). It was first recorded in Ontario and New Brunswick in 1991 (based on specimens at the University of Guelph and University of New Brunswick; M arriott 2009), and Nova Scotia and Prince Edward Island in 1995 (McCorquodale 1998). Notes; There are no reports of this lady beetle directly affecting natives.

Harmonia axyridis (Pallas)

Official common name: Multicoloured Asian lady beetle (ESA) Alternative common name: Harlequin ladybird (Majerus et al. 2006), Halloween ladybug (Acorn 2007) Description: A 4.8-7.5 mm beetle with highly variable colour morphs. The most common colouration is orange-red elytra with highly variable black markings (zero to 19 spots) and a light pronotum with up to five black spots that form an M-shaped marking characteristic of this Figure 1.7 Harmonia axyridis (sketch from species. A less common colour morph has black elytra Gordon and with light coloured spots. Vandenberg 1991). Biogeographic region: Palaearctic Habitat: Forest, grassland and agricultural areas (Kuznetsov 1997). Distribution in Canada: British Columbia and Manitoba to Newfoundland. Colonization history: Since 1916, this lady beetle has been repeatedly released in North America as a biological control agent for aphids (Gordon 1985). In 1988, an unintentionally introduced population was found in Louisiana (Chapin and Brou 1991). It spread rapidly, becoming one of the most common lady beetles in eastern North America within 10 years. The first eastern Canada records are from Quebec (Coderre et al. 1995) and Ontario in 1994 (Marshall 2008), and the first western Canada records are from British Columbia (based on specimens in Royal British Columbia Museum; Marriott 2009) in 1995, indicating completely separate introductions. It was recorded in New Brunswick and Nova Scotia in 1995 (Wheeler and Hoebeke 1996, McCorquodale 1998), Prince Edward Island in 1998 (Majka and McCorquodale 2006), and Manitoba (Wise etal. 2001) and Newfoundland in 2000 (Hicks et al. 2010).

30 Notes: Following the establishment of this species, studies within agroecosystems in Maine, West Virginia, Michigan and Florida have reported declines in the abundance of native lady beetles (Alyokhin and Sewell 2004, Brown and Miller 1998, Colunga- Garcia and Gage 1998, Michaud 2002).

NATIVE LADY BEETLES SPECIES OF CONSERVATION INTEREST

The eight species identified in M arriott et al. (2009) as native to eastern Canada that show one of three patterns relative to presence of the most common non-native species: 1) clearly documented local declines ( Coccinella novemnotata Herbst and

Coccinella transversoguttata richardsoni Brown, Hippodamia parenthesis (Say)), 2) suspected declines ( Adalia bipunctata (Linnaeus) and Hippodamia tredecimpunctata tibialis (Say), Coccinella trifasciata perplexa Mulsant, Anatis mali (Say)), 3) reported to have remained unchanged (Coleomegilla maculata lengiTimberlake).

Coleomegilla maculata lengi Timberlake

Official common name: None Alternative common name: Spotted lady beetle (Marshall 2008, VanDyk 2009); pink spotted lady beetle (DeVries 2007) Description: A 4.2-6.6 mm beetle with pink to red elytra bearing black spots. This lady beetle is easily confused with another native species, Naemia seriata, making it Figure 1.8 Coleomegilla especially important to verify the species data in maculata lengi (sketch collections. It can be confirmed by the absence of the from Gordon 1985). toothed tarsal claw that is characteristic of genus Naemia. Biogeographic region: Nearctic Habitat: Agricultural areas. Distribution in Canada: Manitoba, southern Ontario and Quebec.

31 Notes: This lady beetle switches to pollen when aphid densities are low (Gordon 1985, Obrycki et al. 1998b), and has been reported to coexist with non-native lady beetles in many agroecosystems (Elliott et al. 1996, Obrycki et al. 1998a, Colunga-Garcia and Gage 1998, Day and Tatman 2006, Mignault et al. 2006, Lucas et al. 2007).

Hippodamia parenthesis (Say)

Official common name: None Alternative common name: Parenthesis ladybug (Acorn 2007) Description: A 3.7-5.6 mm beetle with orange to red elytra bearing black, parenthesis-shaped markings near the tip of the elytra. The name-sake parenthesis-shaped markings can vary, sometimes resembling two large blotches or four small spots, so this species can be confused in western Figure 1.9 Hippodamia parenthesis (sketch Canada with other taxon of Hippodamia such as from Gordon 1985). Hippodamia expurgata, Hippodamia lunatomaculata dobzhanski, Hippodamia sinuata sinuata and Hippodamia oregonensis (Gordon 1985). These species do not occur in the east, but in the west, dissection of male genitalia is required to confirm identification (Belicek 1976, Gordon 1985, Acorn 2007). Biogeographic region: Nearctic Habitat: Grasslands, parklands, agricultural areas and other areas with low-growing vegetation (Belicek 1976, Acorn 2007). Distribution in Canada: Yukon and Northwest Territories in the north, and from British Columbia to Nova Scotia. Notes: In Manitoba, the relative abundance of the parenthesis lady beetle declined following the establishment of the non-native Coccinella septempunctata (Turnock et al. 2003).

32 Hippodamia tredecimpunctata tibialis (Say)

Official common name: Thirteenspotted lady beetle (ESC, ESA) Alternative common name: Thirteen-spot ladybug (Acorn 2007) Description: A 4.5-6.4 mm beetle with orange elytra bearing thirteen black spots. Biogeographic region: Holarctic Habitat: Agricultural areas, bogs, grasslands and parklands, wet meadows, sedge marshes and near water (Belicek 1976, Figure 1.10 Hippodamia Kuznetsov 1997, Acorn 2007). tredecimpunctata tibialis Distribution in Canada: Yukon and Northwest Territories in (sketch from Gordon the north, and from British Columbia to Newfoundland 1985). and Labrador. Notes: There are reports of a decrease in the relative abundance of this lady beetle following the arrival of the non-native species Coccinella septempunctata, for example, in the midwestern United States (Obrycki etal. 2000), Maine (Alyokhin and Sewell 2004, Finlayson etal. 2008), eastern South Dakota (Elliott etal. 1996), Quebec (Lucas etal. 2007) and New Brunswick (Boiteau 1983, Boiteau etal. 1999). In contrast, in Manitoba and Ontario, it appears to coexist with non-native lady beetles (Turnock etal. 2003, Marshall 2008).

Adalia bipunctata (Linnaeus)

Official common name: Twospotted lady beetle (ESC, ESA) Alternative common name: Two-spot ladybug (Acorn 2007) Description: A 3.5-5.2 mm beetle with highly variable colour patterns and number of spots (particularly in western Canada). The elytra base colour ranges from red to black and m the spots range from black to a pale colour; though typically this species has a red base colour and a black spot on each Figure 1.11 Adalia elytron. bipunctata (sketch Biogeographic region: Holarctic from Gordon 1985). Habitat: Often arboreal, but is also found in agricultural areas and parkland (Acorn 2007, Belicek 1976). Distribution in Canada: Yukon and Northwest Territories in the north, and from British Columbia to Newfoundland and Labrador. Notes: Numerous studies have reported declining numbers of this species in agroecosystems and other disturbed habitats since the 1980s (Elliott et al. 1996,

33 Colunga-Garcia and Gage 1998, Boiteau e tal. 1999, Cormier etal. 2000, Rytwinski 2004, Hesler et al. 2004, Harmon et al. 2007, Hesler and Kieckhefer 2008). Non native lady beetles such as Coccinella septempunctata and Harmonia axyridis may be, in part, contributing to the apparent decline of the twospotted lady beetle (Obrycki etal. 2000, Colunga-Garcia and Gage 1998).

Coccinella novemnotata Herbst

Official common name: Ninespotted lady beetle (ESC) Alternative common name: Nine-spot ladybug (Acorn 2007) Description: A 4.7-7 mm beetle with pale orange to red- orange with nine black spots, four on each elytron and one central spot near the pronotum. A black line along the elytral suture distinguishes it from most other Coccinella species. Biogeographic region: Nearctic Figure 1.12 Coccinella novemnotata (sketch Habitat: Agricultural areas, parkland and sand dunes (Belicek from Gordon 1985). 1976, Acorn 2007). Distribution in Canada: Historically found from Northwest Territories in the north, and from British Columbia to Quebec. Notes: By the mid-1980s, it was rare or extirpated in southern Ontario and Quebec (Wheeler and Hoebeke 1995, Marshall 2008), as well as in South Dakota and the northeastern USA (Hesler and Kieckhefer 2008). A single adult was discovered in Arlington, Virginia in 2006 (Losey et al. 2007) and in Mont Saint-Hilaire, Quebec, this lady beetle was observed by M. Racine between 2006 and 2010 (Skinner and Domaine 2010). The disappearance of this species from much of its historical range, while continuing to persist in western regions of North America (Wheeler and Hoebeke 1995) such as Alberta and Saskatchewan (Acorn 2007) is currently under study. There is only anecdotal information linking its decline with the introduction of Coccinella septempunctata.

34 Coccinella transversoguttata richardsoni Brown

Official common name: Transverse lady beetle (ESC, ESA). Alternative common name: Suggested alternative (this study): Richardson's transverse lady beetle. Description: A 5-7.8 mm beetle with orange-red elytra bearing a black band across the front of its elytra (near the head) and two additional black blotches on each elytron. Biogeographic region: Holarctic Habitat: Agricultural areas, parklands and forests (Belicek Figure 1.13 Coccinella transversoguttata 1976, Acorn 2007). richardsoni (sketch Distribution in Canada: Yukon and Northwest Territories in from Gordon 1985). the north, and from British Columbia to Newfoundland and Labrador. Notes: There are many reports of a decrease in the relative abundance of this lady beetle following the arrival of non-native species. In South Dakota (Hesler et al. 2004, Hesler and Kieckhefer 2008), Manitoba (Turnock et ol. 2003) and New Brunswick (Boiteau 1983, Boiteau et al. 1999) it experienced a decline following the arrival of C. septempunctata in the 1980s. In Maine, it also declined in the 1980s following the arrival of C. septempunctata (Elliott et al. 1996), but declined further in the mid-1990s following the arrival of H. axyridis and P. quatuordecimpunctata (Alyokhin and Sewell 2004, Finlayson et al. 2008).

Coccinella trifasciata perplexa Mulsant

Official common name: None Alternative common name: Three-banded lady beetle (Marshall 2008, VanDyk 2009) or three-banded ladybug (Acorn 2007); suggested alternative (this study): perplexed three banded lady beetle for this subspecies. Description: A 4-5 mm beetle with orange-red elytra bearing a black band across the front of the elytra (near the head) Figure 1.14 Coccinella and two additional smaller, black bands on each elytron. trifasciata perplexa Habitat: Agricultural areas, parklands and boreal forests (sketch from Gordon (Belicek 1976). 1985). Biogeographic region: Holarctic Distribution in Canada: Yukon and Northwest Territories in the north, and from British Columbia to Newfoundland and Labrador.

35 Notes: In Manitoba this lady beetle decreased in relative abundance following the arrival of C. septempunctata in 1988 (Turnock etal. 2003), while in Michigan populations did not change following the arrival of Harmonia axyridis (Colunga- Garcia and Gage 1998).

Anatis mali (Say)

Official common name: Eyespotted lady beetle (ESC) Alternative common name: American eyespot ladybug (Acorn 2007) Description: A 7.3-10 mm beetle with yellow to brownish-red elytra bearing black spots, each with a pale halo surrounding it, arranged in three transverse lines. The fifteenspotted lady beetle (Anatis labiculata) has a similar number and orientation of spots, but w ithout the pale halo. Figure 1.15 Anatis mali (sketch from Biogeographic region: Nearctic Gordon 1985). Habitat: A variety of wooded habitats (Belicek 1976, Acorn 2007). Distribution in Canada: Yukon and Northwest Territories in the north, and from British Columbia to Newfoundland. Notes: In Pemiscot and Madrid Counties of Missouri this lady beetle has not been collected since 1966 (Fothergill and Tindall 2010).

36 1.3 STUDY OBJECTIVES

The overall goal of this study is to determine whether changes in lady beetle distributions and relative abundance have occurred in eastern Canada, and whether patterns can be determined from historical specimen data in natural history collections.

Specific objectives include:

1) To verify identifications and database specimen label data for eastern Canadian lady

beetles found in collections across eastern Canada, including georeferencing

location information as necessary;

2) To assess gaps in records that could lead to collection biases in different sorts of

collections, and determine whether collection data are complete enough to assess

distribution patterns over time for native and non-native species of conservation

interest;

3) To map species and compare range distributions and relative abundance patterns

over time for native and non-native species of conservation interest. Three distinct

patterns in the collection data are expected:

a. Declines in relative abundance and geographical extent are expected for two

native species with clearly documented local declines ( Coccinella

novemnotata and Coccinella transversoguttata richardsoni), and two native

species with suspected declines ( Adalia bipunctata and Hippodamia

tredecimpunctata tibialis);

37 b. Relatively constant relative abundance and geographical extent is expected

for one native species, Coleomegilla maculata lengi, which has been reported

to have remained unchanged. c. Clear evidence of range expansion and increases in relative abundance is

expected for the five non-native species following their establishment in a

region (Coccinella septempunctata, Coccinella u. undecimpunctata, Harmonia

axyridis, Hippodamia variegata and Propylea quatuordecimpunctata).

38 1.4 LITERATURE CITED

Acorn, J. 2007. Ladybugs of Alberta, finding and connecting the dots. The University of Alberta Press, Edmonton, Alberta. 169 pp.

Alyokhin, A. and Sewell, G. 2004. Changes in a lady beetle community following the establishment of three alien species. Biological Invasions. 6: 463-471.

Belicek, J. 1976. Coccinellidae of western Canada and Alaska with analyses of the transmontane zoogeographic relationships between the fauna of British Columbia and Alberta (Insecta: Coleoptera: Coccinellidae). Quaestiones Entomologicae. 12: 283-409.

Boiteau, G. 1983. The arthropod community of potato fields in New Brunswick. The Canadian Entomologist 115: 847-853.

Boiteau, G., Y. Bousquet and Osborn, W.P.L. 1999. Vertical and temporal distribution of Coccinellidae (Coleoptera) in flight over an agricultural landscape. The Canadian Entomologist 131: 269-277.

Brown, W.J. 1940. Notes on the American distribution of some species of Coleoptera common to the European and North American continents. The Canadian Entomologist. 72: 65-78.

Brown, M.W. and Miller, S.S. 1998. Coccinellidae (Coleoptera) in apple orchards of eastern West Virginja and the impact of invasion by Harmonia axyridis. Entomological News 109:143-151.

Brunskill, G.J., Rosenberg, D.M., Snow, N.B., Vascotto, G.L. and Wageman, R. 1973. Ecological studies of aquatic systems in the MacKenzie-Porcupine drainages in relation to proposed pipeline and highway developments. Task Force on Northern Oil Development (Canada), Environmental-Social Committee, Volumes I & II. Reports 73-40 (131 pp) & 73-71 (345 pp).

Chantel, C. 1972. Additions a la faune Coleopterique du Quebec. Le Naturaliste Canadien. 99: 243-244.

Chapin, J.B. and Brou, V.A. 1991. Harmonia axyridis (Pallas), the third species of the genus to be found in the United States (Coleoptera: Coccinellidae). Proceedings of the Entomological Society of Washington. 93: 630-635.

Chapman, A.D. 2005. Uses of primary species-occurrence data, version 1.0. Report for the Global Biodiversity Information Facility, Copenhagen. 106 pp.

Coderre, D., E. Lucas and Gagne, 1.1995. The occurrence of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in Canada. The Canadian Entomologist. 127: 609-611.

39 Colla, S.R. and Packer, L. 2008. Evidence for decline in eastern North American bumblebees (Hymenoptera: Apidae), with special focus on Bombus affinis Cresson. Biodiversity and Conservation. 17:1379-1391.

Colla, S.R. and Ratti, C.M. 2010. Evidence for the decline of the western bumble bee (Bombus occidentalis Greene) in British Columbia. The Pan-Pacific Entomologist. 86: 32-24.

Colunga-Garcia, M. and Gage, S.H. 1998. Arrival, establishment, and habitat use of the multicolored Asian lady beetle (Coleoptera: Coccinellidae) in a Michigan landscape. Environmental Entomology. 27:1574-1580.

Cormier, C.M., T.A. Forbes, T.A. Jones, R.D. Morrison and McCorquodale, D.B. 2000. Alien invasion: the status of non-native lady beetles (Coleoptera: Coccinellidae) in industrial Cape Breton, Nova Scotia. Northeastern Naturalist. 7: 241-247.

Cotterill, F.P. 1995. Systematics, biological knowledge and environmental conservation. Biodiversity and Conservation. 4:183-205.

Cotterill, F.P.D. 1997. The second Alexandrian tragedy, and the fundamental relationship between biological collections and scientific knowledge. In Nudds, J.R. and Pettitt, C.R. The values and valuation of natural science collections (Proceedings of Conference at the Manchester Museum, Manchester, UK, 19-21 April 1995). Geological Society, London, pp. 217-241.

Cottrell, T.E. and Shapiro-llan, D.l. 2003. Susceptibility of a native and an exotic lady beetle (Coleoptera: Coccinellidae) to Beauveria bassiana. Journal of Invertebrate Pathology. 84:137-144.

Danks, H.V. 1991. Museum collections: fundamental values and modern problems. Collection Forum. 7: 95-111.

Delisle, F., Lavoie, C., Jean, M., and Lachance, D. 2003. Reconstructing the spread of invasive plants: taking into account biases associated with herbarium specimens. Journal of Biogeography. 30:1033-1042.

Day, W.H., D.R. Prokrym, D.R. Ellis and Chianese, R.J. 1994. The known distribution of the predator Propylea quotuordecimpunctata (Coleoptera: Coccinellidae) in the United States, and thoughts on the origin of this species and five other exotic lady beetles in eastern North America. Entomological News. 105: 244-256.

Day, W.H. and Tatman, K.M. 2006. Changes in abundance of native and adventive Coccinellidae (Coleoptera) in alfalfa fields, in northern New Jersey (1993-2004) and Delaware (1999-2004), U.S.A. Entomological News. 117: 491-502.

40 DeVries, PJ. 2007. Coccinellidae of Wisconsin; ladybird beetles of Wisconsin. Retrieved July 4, 2011, from http://insects.entomology.wisc.edu/Coleoptera/Coccinellidae/index.html.

Eaton, E.R. and K. Kaufman. 2007. Kaufman field guide to insects of North America. Houghton Mifflin Company, Boston. 391 pp.

Elith, J., Graham, C.H., Anderson, R.P., Dudik, M., Ferrier, S., Guisan, A., Hijmans, R.J., Huettmann, F., Leathwick, J.R., Lehmann, A., Li, J., Lohmann, L.G., Loiselle, B.A., Manion, G., Moritz, C., Nakamura, M., Nakazawa, Y., Overton, J.McC., Peterson, A.T., Phillips, S.J., Richardson, K., Scachetti-Pereira, R., Schapire, R.E., Soberon, J., Williams, S., Wisz, M.S. and Zimmermann, N.E. 2006. Novel methods improve prediction of species' distributions from occurrence data. Ecography. 29:129- 151.

Elliott, N.C., Kieckhefer, R.W. and Kauffman, W.C. 1996. Effects of an invading coccinellid on native coccinellids in an agricultural landscape. Oecologia 105: 537-544.

Elliott, N.C., Kieckhefer, R.W., Lee, J.H. and French, B.W. 1998. Influence of within-field and landscape factors on aphid populations in wheat. Landscape Ecology. 14: 239-252.

Ellis, D.R., Prokrym, D.E. and Adams, R.G. 1999. Exotic lady beetle survey in northeastern United States: Hippodamia variegata and Propylea quatuordecimpunctata (Coleoptera: Coccinellidae). Entomological News. I l l : 73-84.

Evans, E.W. 1991. Intra versus interspecific interactions of lady beetles (Coleoptera: Coccinellidae) attacking aphids. Oecologica. 87: 401-408.

Finlayson, C.J., K.M. Landry and Alyokhin, A.V. 2008. Abundance of native and non native lady beetles (Coleoptera: Coccinellidae) in different habitats in Maine. Annals of the Entomological Society of America. 101:1078-1087.

Ferrier, S. 2002. Mapping spatial pattern in biodiversity for regional conservation planning: where to from here? Systematic Biology. 51: 331-363.

Fisher, R.N. and Schaffer, H.B. 1996. The decline of amphibians in California's Great Central Valley. Conservation Biology. 10:1387-1397.

Fitzpatrick, U., Murray, T.E., Paxton, R.J., Breen, J., Cotton, D., Santorum, V. and Brown, M.J.F. 2007. Rarity and decline in bumblebees - a test of causes and correlates in the Irish fauna. Biological Conservation. 136:185-194.

Fothergill, M-J. and Tindall, K.V. 2010. Lady beetle (Coleoptera: Coccinellidae: Coccinellinae) occurrences in southeastern Missouri agricultural systems: differences between 1966 and present. The Coleopterists Bulletin. 64: 379-382.

41 Funk, V. and Richardson. K. 2002. Systematic data in biodiversity studies: use it or lose it. Systematic Biology. 51: 303-316.

Gabel, M.L., Johnson, P.J., Ode, D.J., Downing, H.A., and Kostel, G.M. 2007. South Dakota's natural collections: an endangered teaching and research resource. Proceedings of the South Dakota Academy of Science. 86: 71-82.

Gordon, R.D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Entomological Society. 95:1-912.

Gordon, R.D. 1987. The first North American records of Hippodamia variegata (Goeze) (Coleoptera: Coccinellidae). Journal of the New York Entomological Society 95: 307-309.

Gordon, R.D. and Vandenberg, N. 1991. Field guide to recently introduced species of Coccinellidae (Coleoptera) in North America, with revised key to North America genera of Coccinellini. Proceedings of the Entomological Society of Washington. 93: 845-864.

Graham, C.H., Elith, J., Hijmans, R.J., Guisan, A., Peterson, A.T., and Loiselle, B.A. 2008. The influence of spatial errors in species occurrence data used in distribution models. Journal of Applied Ecology. 45: 239-247.

Graham, C.H., Ferrier, S., Huettman, F., Moritz, C., and Peterson, A.T. 2004. New developments in museum-based informatics and applications in biodiversity analysis. Trends in Ecology and Evolution. 19: 497-503.

Gropp, R.E. 2003. Are university natural science collections going extinct? BioScience. 53:550.

Guralnick, R.P., Wieczorek, J., Beaman, R., Hijmans, R.J., and the BioGeomancer working Group. 2006. BioGeomancer: Automated georeferencing to map the World's biodiversity data. PLOS Biology. 4:1908-1909.

Guralnick, R., and Van Cleve, J. 2005. Strengths and weaknesses of museum and national survey data sets for predicting regional species richness: comparative and combined approaches. Diversity and Distributions. 11: 349-359.

Harmon, J.P., Stephens, E. and Losey, J. 2007. The decline of native coccinellids (Coleoptera: Coccinellidae) in the United States and Canada. Journal o f Insect Conservation 11: 85-94.

Herkt, M. 2007. Modelling habitat suitability to predict the potential distribution of Erhard's wall lizard Podarcis erhardiil on Crete. Masters thesis, International Institute for Geo-information Science and Earth Observation (ITC), The Netherlands, pp 80.

42 Hesler, L.S., Kieckhefer. R.W. and Catangui, M.A. 2004. Surveys and field observations of Harmonia axyridis and other Coccinellidae (Coleoptera) in eastern and central South Dakota. Transactions of the American Entomological Society. 130:113- 133.

Hesler, L.S. and Kieckhefer, R.W. 2008. Status of exotic and previously common native coccinellids (Coleoptera) in South Dakota landscapes. Journal of the Kansas Entomological Society. 81: 29-49.

Hicks, B., Majka, C.G. and Pardy Moores, S. 2010. Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) found in Newfoundland. The Coleopterists Bulletin. 64: 50.

Hicks, S.D. 1971. Comments on certain Coleoptera from extreme southern Ontario and the Ottawa District. The Coleopterists Bulletin. 25:103-106.

Hijman, R.J., Schreuder, M., de la Cruz, J., and Guarino, L. 1999. Using GIS to check co ordinates of genebank accessions. Genetic Resources and Crop Evolution. 46: 291-296.

Hinojosa-Diaz, I.A., Yanez-Ordonez, O., Chen, G., Peterson, A.T. and Engel, M.S. 2005. The North American invasion of the giant resin bee (Hymenoptera: Megachilidae). Journal of Hymenoptera Research. 14: 69-77.

Hinojosa-Diaz, I.A., Feria-Arroyo, T.P. and Engel, M.S. 2009. Potential distribution of orchid bees outside their native range: the cases of Eulaema polychrome (Mocsary) and Euglossa viridissima Friese in the USA (Hymenoptera: Apidae). Diversity and Distributions. 15:421-428.

Hoebeke, E.R. and Wheeler, A.G. 1996. Adventive lady beetles (Coleoptera: Coccinellidae) in Canadian Maritime Provinces, with new eastern US records on Harmonia quadripunctata. Entomological News. 107: 281-290.

Honek, A. 1985. Habitat preferences of aphidophagous coccinellids (Coleoptera). Entomophaga. 30: 253-264.

Humble, L.M. 1991. Occurrence of Coccinella septempunctata (L) (Coleoptera, Coccinellidae) in central British Columbia. Pan-Pacific Entomologist. 67: 224-226.

Johnson, N.F. 2007. Biodiversity informatics. Annual Review of Entomology. 52:421-438.

Koch, J.B. and Strange, J.P. 2009. Constructing a species database and historic range maps for North American bumblebees (Bombus sensu stricto Latreille) to inform conservation decisions. Bee Science. 9:97-108.

43 Kuznetsov, V.N. 1997. Lady beetles of the Russian Far East. Center for Systematic Entomology, Gainesville. 248 pp.

Larochelle, A. 1979. Les Coleopteres Coccinellidae du Quebec. Cordulia (Supplement). 10:1-111.

Lavoie, C., Jean, M., Delisle, F. and Letourneau, G. 2003. Exotic plant species o f the St Lawrence River wetlands: a spatial and historical analysis. Journal of Biogeography. 30: 537-549.

Lavoie, C., Jodoin, Y. and Goursaud de Merlis, A. 2007. How did common ragweed (Ambrosia artemisiifolia L.) spread in Quebec? A historical analysis using herbarium records. Journal of Biogeography. 34:1751-1761.

Lonsdale, O and Huber, J.T. 2011. Insect collections of Canada series - Canadian National Collection of Insects, Arachnids & Nematodes, Ottawa. Newsletter of the Biological Survey of Canada, 30:15-40.

Losey, J.E., Perlman, J.E. and Hoebeke, E.R. 2007. Citizen scientist rediscovers rare ninespotted lady beetle, Coccinella novemnotata, in eastern North America. Journal of Insect Conservation. 11: 415-417.

Lovejoy, T.E., Brouillet, L., Doolittle, W.F., Gonzolas, A., Green, D.M., Hall, P., Herbert, P., Herrmann, T.M., Hyde, D., Lee, J., Maddison, W.P., Otto, S.P., Sperling, F. and Thompson, R.P. 2010. Canadian taxonomy: exploring biodiversity, creating opportunity. Report for the Government of Canada, Ottawa. 126 pp.

Lucas, E., Gagne, I and Coderre, D. 2002. Impact of the arrival of Harmonia axyridis on adults of Coccinella septempunctata and Coleomegilla maculata (Coleoptera: Coccinellidae). European Journal of Entomology. 99: 457-463.

Lucas, E,, Vincent, C., Labrie, G., Chouinard, G., Fournier, F., Pelletier, F., Bostanian, N.J., Coderre, D., Mignault, M.-P. and Lafountaine, P. 2007. The multicolored Asian ladybeetle Harmonia axyridis (Coleoptera : Coccinellidae) in Quebec agroecosystems ten years after its arrival. European Journal of Entomology 104: 737-743.

Majerus, M., Strawson, V. and Roy, H. 2006. The potential of the arrival of the harlequin ladybird, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), in Britain. Ecological Entomology. 31: 207-215.

Majka, C. G. and McCorquodale, D.B. 2006. The Coccinellidae (Coleoptera) of the Maritime Provinces of Canada: new records, biogeographical notes, and conservation concerns. Zootaxa. 1154: 49-68.

44 Majka, C. G. and McCorquodale, D.B. 2010. Ladybird beetles (Coleoptera: Coccinellidae) of the Atlantic Maritime Ecozone, Assessment o f Species Diversity in the Atlantic Maritime Ecozone (D.F. McAlpine, and I.M. Smith, Eds), NRC Research Press. Ottawa, Ontario, pp. 439-452.

Margules, C.R., and Austin, M.P. 1994. Biological models for monitoring species decline: the construction and use of data bases. Philosophical Transactions, Royal Society, London B. 344: 69-75.

Marriott, S.M., Giberson, DJ. and McCorquodale, D.B. 2009. Changes in the status and geographic ranges of Canadian lady beetles (Coccinellidae) and the selection of candidates for risk assessment. Part I Foundation Report. Report to COSEWIC Arthropods Species Specialist Committee. 53 pp.

Marshall, S. 2008. Lady beetles of Ontario. Retrieved December, 2008, from http://www.uoguelph.ca/debu/lady/lady-beetles.htm.

McCarthy, M.A. 1998. Identifying declining and threatened species with museum data. Biological Conservation. 83: 9-17.

McCorquodale, D.B. 1998. Adventive lady beetles (Coleoptera: Coccinellidae) in eastern Nova Scotia, Canada. Entomological News 109:15-20.

McCorquodale, D.B., Brown, J.M., and Marshall, S.A. 2007. A decline in the number of long-horned wood boring beetles (Coleoptera: Cerambycidae) species in Ontario during the 20th century? Journal of the Entomological Society of Ontario. 138: 105-133.

McCorquodale, D.B., Giberson, DJ. and M arriott, S.M. 2011. Changes in the status and geographic range of Canadian lady beetles (Coleoptera: Coccinellidae: Coccinellinae) and the selection of candidate species for risk assessment by the Committee on the Status of Endangered Wildlife in Canada - Part 3. Report to COSEWIC Arthropods Species Specialist Committee. 60 pp.

Meier, R., and Dikow, T. 2004. Significance of specimen databases from taxonomic revisions for estimating and mapping the global species diversity of invertebrates and repatriating reliable specimen data. Conservation Biology. 18: 478-488.

Michaud, J.P. 2002. Invasion of the Florida citrus ecosystem by Harmonia axyridis (Coleoptera: Coccinellidae) and asymmetric competition with a native species, Cycloneda sanguinea. Environmental Entomology. 31: 827-835.

Mignault, P.-M., Roy, M. and Brodeur, J. 2006. Soybean aphid predators in Quebec and the suitability of Aphis glycines as prey for three Coccinellidae. BioControl 51: 89- 109.

45 Miller, B.P., Enright, N.J., and Lamont, B.B. 2007. Record error and range contraction, real and imagined, in the restricted shrub Banksia hookeriana in south-western Australia. Diversity and Distributions. 13: 406-417.

Murphey, P.C., Guralnick, R.P., Glaubitz, R., Neufeld, D., and Ryan, J.A. 2004. Georeferencing of museum collections: a review of problems and automated tools, and the methodology developed by the Mountain and Plains Spatio- Temporal Database-lnformatics Initiative (Mapstedi). Phylolnformatics. 3:1-29.

Myers, P., Lundrigan, B.L., Hoffman, S.M., Haraminac, A.P., and Seto, S.H. 2009. Climate- induced changes in the small mammal communities of the Northern Great Lakes Region. Global Change Biology. 15:1434-1454.

Newbold, T. 2010. Applications and limitations of museum data for conservation and ecology, with particular attention to species distribution models. Progress in Physical Geography. 34: 3-22.

Obrycki, J.J. and Kring, T.J. 1998. Predaceous Coccinellidae in biological control. Annual Review of Entomology. 43: 295-321.

Obrycki, J.J., Giles, K.L. and Ormond, A.M. 1998a. Interactions between an introduced and indigenous coccinellid species at different prey densities. Oecologia. 117: 279-285.

Obrycki, J.J., Giles, K.L. and Ormond, A.M. 1998b. Experimental assessment of interactions between larval Coleomegilla maculata and Coccinella septempunctata (Coleoptera: Coccinellidae) in field-cages. Environmental Entomology. 27:1280-1288.

Obrycki, J.J., Elliott, N.C. and Giles, K.L. 2000. Coccinellid introductions: potential for and evaluation of non-target effects, Nontarget effects of biological control (P.A. Follet, and JJ. Duan, Eds.), Dordrecht, Netherlands; Kluwer: 127-145.

O'Connell Jr., A E., Gilbert, A.T., and Hatfield, J.S. 2004. Contribution of natural history collection data to biodiversity assessment in national parks. Conservation Biology. 18:1254-1261.

Pergams, O.R.W. and Nyberg, D. 2001. Museum collections of mammals corroborate the exceptional decline of prairie habitat in the Chicago region. Journal of Mammalogy. 82: 984-992.

Peterson, A.T., Navarro-Siguenza, A.G., and Pereira, R.S. 2004. Detecting errors in biodiversity data based on collectors' itineraries. Bulletin of the British Ornithology Club. 124:143-151.

46 Pyke, G.H., and Ehrlich, P.R. 2010. Biological collections and ecological/environmental research: a review, some observations and a look to the future. Biological Reviews. 85: 247-266.

Randrianasolo, A., Miller, J.S., and Consiglio, T.K. 2002. Application o f IUCN criteria and Red List categories to species of five Anacardiaceae genera in Madagascar. Biodiversity and Conservation. 11:1289-1300.

Reddy, S., and Davalos, L.M. 2003. Geographical sampling bias and its implications for conservation priorities in Africa. Journal of Biogeography. 30: 1719-1727.

Riddick, E.W., Cottrell, T.E. and Kidd, K.A. 2009. Natural enemies of the Coccinellidae: parasites, pathogens, and parasitoids. Biological Control. 51: 306-3.

Rowe, R.J. 2005. Elevation gradient analyses and the use of historical museum specimens: a cautionary tale. Journal of Biogeography. 32: 1883-1897.

Rushton, S.P., Ormerod, SJ. and Kerby, G. 2004. New paradigms for modeling species distributions? Journal of Applied Ecology. 41: 193-200.

Rytwinski, T. 2004. Plant use by native and non-native lady beetles (Coccinellidae) in Cape Breton Island, Nova Scotia. Honours thesis, University College of Cape Breton.

Samways, M.J., Osborn, R., Hastings, H. and Hattingh, V. 1999. Global climate change and the accuracy of predictions of species' geographical ranges: establishment success of introduced ladybirds (Coccinellidae, Chilocorus spp.) worldwide. Journal of Biogeography. 26: 795-812.

Schaeffer, C. 1912. Coccinella undecimpunctata Linn, in Massachusetts. Psyche 19:104- 105.

Shaffer, H.B., Fisher, R.N. and Davidson, C. 1998. The role of natural history collections in documenting species declines. Trends in Ecology and Evolution. 13: 27-30.

Skinner, B. and Domaine, E. 2010. Rapport sur la situation de la coccinelle a neuf points (Coccinella novemnotata) au Quebec. Ministere des Ressources naturelles et de la Faune du Quebec, Faune Quebec, pp 37.

Snow, N. 2005. Successfully curating smaller herberia and natural history collections in academic settings. BioScience. 55: 771-779.

Soberon, J.M., Llorente, J.B., and Onate, L. 2000. The use of specimen-label databases for conservation purposes: an example using Mexican Papilionid and Pierid butterflies. Biodiversity and Conservation. 9:1441-1466.

47 Suarez, A.V., and Tsutsui, N.D. 2004. The value of museum collections fo r research and society. BioScience. 54: 66-74.

Sutherst, R.W. 2003. Prediction of species geographical ranges. Journal of Biogeography. 30: 805-816.

Sutherst, R.W., Floyd, R.B. and Maywald, G.F. 1996. The potential geographical distribution of the cane toad, Bufo marinus L. in Australia. Conservation Biology. 10: 294-299.

Tingley, M.W., and Beissinger, S.R. 2009. Detecting range shifts from historical species occurrences: new perspectives on old data. Trends in Ecology and Evolution. 24: 625-633.

Turnock, W.J., Wise, I.L., and Matheson, F.O. 2003. Abundance of some native coccinellines (Coleoptera: Coccinellidae) before and after the appearance of Coccinella septempunctata. The Canadian Entomologist. 135: 391-404.

VanDyk, J. 2009. BugGuide. Retrieved February, 2009, from http://bugguide.net.

Watson, W.Y. 1979. North American distribution of Coccinella u. undecimpunctata L. (Coleoptera: Coccinellidae). The Coleopterists Bulletin. 33: 85-86.

Wheeler, Jr., A.G. and Hoebeke, E.R. 1981. A revised distribution of Coccinella undecimpunctata L. in eastern and western North America (Coleoptera: Coccinellidae). The Coleopterists Bulletin. 35: 213-216.

Wheeler, Jr., A.G. and Hoebeke, E.R. 1995. Coccinella novemnotata in northeastern North America: historical occurrence and current status (Coleoptera: Coccinellidae). Proceedings of the Entomological Society of Washington. 97: 701- 716.

Wheeler, Jr., A.G. and Stoops, C.A. 1996. Status and spread of the Palaearctic lady beetles Hippodamia variegata and Propylea quatuordecimpunctata (Coleoptera: Coccinellidae) in Pennsylvania, 1993-1995. Entomological News. 107: 291-298.

Wheeler, T.A., Huber, J.T. and Currie, D.C. 2001. Label data standards for terrestrial arthropods. Brief for the Biological Survey of Canada (Terrestrial Arthropods). Retrieved on July 20, 2011, from http://www.biology.ualberta.ca/bsc/pdf/labelbrief.pdf.

Wieczorek, J., Guo, Q., and Hijmans, RJ. 2004. The point-radius method for georeferencing locality descriptions and calculating associated uncertainty. International Journal of Geographical Information Science. 18: 745-767.

48 Wiggins, G.B., Marshall, S.A. and Downes, J.A. 1991. The importance of research collections of terrestrial arthropods. Brief for the Biological Survey of Canada (Terrestrial Arthropods). 23:1-16.

Williams, P.H., Margules, C.R., and Hilbert, D.W. 2002. Data requirements and data sources for biodiversity priority area selection. Journal of Bioscience. 27: 327- 338.

Wise, I.L., Turnock, W.J. and Roughley, R.E. 2001. New records of coccinellid species for the province of Manitoba. Proceedings of the Entomological Society of Manitoba. 57: 5-10.

Yanega, D. 1996. Records of Propylea quatuordecimpunctata (Coleoptera: Coccinellidae) from Long Island, New York: evidence for a naturalized population before 1991. Entomological News. 107: 36-38.

Yasuda, H., Evans, E.W., Kajita, Y., Urakawa, K. and Takizawa, T. 2004. Asymmetrical larval interactions between introduced and indigenous ladybirds in North America. Oecologia. 141:722-731.

49 CHAPTER 2

DO SMALL COLLECTIONS ADD VALUE TO THE BIG PICTURE? AN EVALUATION OF

ATLANTIC CANADIAN LADY BEETLE DATA RETRIEVED FROM NATURAL HISTORY

COLLECTIONS THROUGHOUT EASTERN CANADA

50 ABSTRACT

Collection data have been effective fo r assessing various ecological questions relating to species diversity, abundance and distribution patterns, and documenting the spread of invasive species for a wide variety of taxa. However, temporal and spatial gaps in collection data can bias interpretation of patterns, and the extent and nature of the gaps vary with the category of collection. The goal of this study is to assess potential gaps in Atlantic Canadian lady beetle collection records from a variety of natural history collection categories, to determine whether these data can be used to determine long term distribution and relative abundance patterns fo r lady beetles in eastern Canada.

Twenty-seven collections representing four categories of collections were assessed for potential biases by comparing numbers of specimen records through different time periods, different seasons, different geographic regions, and different taxa. Numbers of records for each grouping were compared using Mann-Whitney U-tests or Kruskal-Wallis tests to determine whether there were significant differences that would indicate gaps.

Analyses were also performed with data combined for all collections to determine whether addition of data from multiple collections reduced potential biases by filling in the gaps in the records. Individual collection categories did show gaps in record data that could bias interpretation of lady beetle distributions, but these gaps were minimized by combining data from all collections. Lady beetles are well represented in collections from Atlantic Canada and therefore there is potential to assess temporal changes in distribution from collection data, as long as the potential biases can be accounted for.

51 2.1 INTRODUCTION

There have been a growing number of studies since the mid-1990s which have used Natural History Collection data to address ecological questions. Collection data have been used to assess species richness or abundance patterns (e.g. Meier and Dikow

2003, McCorquodale et al. 2007, Myer et al. 2009), to assess changes in distribution

(e.g. Fitzpatrick etal. 2007, Colla and Ratti 2010, Colla and Packer 2009), and to document the spread of invasive species (e.g. Lavoie et al. 2007, Delisle et al. 2003) over a wide variety of taxa including plants, mammals, reptiles, and insects. Natural history collections worldwide contain >2.5 billion preserved biological specimens (Cotterill

1995) from taxonomic, ecological, and biodiversity studies (Danks 1991, Cotterill 1995) from the early 1800s to the present (Lovejoy et al. 2010). In Canada, >120 biological collections exist, ranging in size from several hundred to 16.7 million specimens (Lovejoy etal. 2010) and fall into four broad categories: 1) Federal government (museums and laboratories); 2) Other government (e.g. provincial museums); 3) University; 4) Other collections (e.g. private collections) (Lovejoy et al. 2010).

The different categories of collections have different strengths and limitations for biodiversity research depending on their focus and level of funding, resulting in potential taxonomic, spatial, environmental, or temporal gaps in records that may bias interpretation of the data. For example, taxonomic gaps can occur when there is preferential collecting by specialists associated with a collection (resulting in domination by certain taxa), or when there are inaccuracies due to misidentifications (Newbold

2010). Spatial gaps occur where the collection localities fo r taxa found in a collection

52 depend on the location of the collection (Newbold 2010). This may occur when sampling is opportunistic as opposed to systematic, targeting 'hotspots' or easily accessible sites such as close to the collectors' homes, along roads and rivers, and near towns or research stations (Soberon et al. 2000, Reddy and Davalos 2003, Williams et al.

2002, Randrianasolo et al. 2002, Graham et al. 2004, Pyke and Ehrlich 2010). A focus on certain habitats or seasons in a collection can result in environmental gaps (Newbojd

2010), and can occur when taxa have not been sampled across their entire environment gradient (e.g. no collection from high elevations), or when collecting is restricted to specific seasons due to the life cycle of a target organism or collector preferences

(Newbold 2010). Finally, temporal gaps can occur in collections because collecting efforts fluctuate over time as collectors' interests change or they retire, or as collection space fills up (Newbold 2010). The different collections vary with respect to these four potential gaps in data due to their different mandates; for example, there would be an agricultural or forestry focus for agricultural or forestry research labs, compared to a broader collecting focus from University collections with undergraduate entomology programs (Lovejoy et al. 2010). Since some collections may have temporal gaps and others may have spatial or taxonomic gaps, examination o f specimens from multiple collection categories may reduce potential biases that might occur from gaps in data from just one category of collection (Guralnick and Van Cleve 2005, Pyke and Ehrlich

2010).

Insects represent good taxa with which to test whether or not gaps exist in collections that could bias interpretations in ecological studies, and also whether or not

53 combining data from multiple collections can minimize such gaps. One insect group, the lady beetles, has been well represented in insect collections for centuries because of their bright colours and their presence in habitats frequented by insect collectors. Not only are they common in insect collections of all types, but there is also a long record of lady beetle collection in Canada, and an extensive body of literature documenting taxonomy and distribution of these beetles (e.g. Belicek (1976) for western Canada and

Alaska, Larochelle (1979) for Quebec, and Gordon (1985) and Gordon and Vandenberg

(1991) for North America (north of Mexico)).

The goal of this study is to compile records from Atlantic Canada fo r a suite of lady beetle species to assess 1) whether there are gaps in the records for specific categories of collections that could bias interpretation of distribution patterns, and 2) whether combined collection data can be used to determine long term distribution and relative abundance patterns in lady beetles in eastern Canada. I expect that collections with specific mandates (e.g. government research collections) will show taxonomic, spatial and environmental biases from targeted collecting in specific habitats (e.g. agricultural), as well as temporal gaps in records associated with lack of collecting between projects/studies. I expect provincial museum collections to be biased temporally due to a reduction in collecting or specimen retention for taxa which are well represented in collections since resources for storage and maintenance are often limiting factors for museum collections. I expect private collections to be biased both temporally and spatially as most private collectors in Atlantic Canada are contemporaries who focus on sampling close to home. Finally, I think it likely that the

54 broad-scale collecting by untrained students, combined with specialist graduate-level student collecting will result in university collections which are diverse taxonomically, spatially, environmentally and temporally.

2.2 MATERIALS AND METHODS

2.2.1 SELECTION OF LADY BEETLE SPECIES FOR STUDY

Of the 55 species of Coccinellinae (the subfamily which includes the relatively large, aphid-eating lady beetles) in Canada, only a few are common enough to assess broad scale collection patterns in eastern Canada. Six native lady beetle species were chosen for this study (Table 2.1) because they were present in all collection types, and a previous study (Marriott et al. 2009) indicated that they include species that are a) universally reported to be experiencing declines in relative abundance and/or geographic distribution (Hippodamia parenthesis, Adalia bipunctata, Coccinella transversoguttata richardsoni, Anatis mali), or b) reported to be experiencing declines in relative abundance and/or geographic distribution in some regions while remaining unchanged in others (Hippodamia tredecimpunctata tibialis, Coccinella trifasciata perplexa). In addition, the five non-native species now common in Canada (Table 2.1) were also included in this study as examples of species which might cause taxonomic and temporal biases in collections while novel to a region (e.g. during early colonization).

55 Table 2.1 Comparison of the numbers of Atlantic Canada records for 11 study species of lady beetles found in insect collections throughout eastern Canada. Refer to page 61 for definitions of collection categories.

Lady Beetle Collection Type Species* Government Museum University Private Total Native Twospot 143 68 195 10 416 Eyespot 124 23 100 12 259 Transverse 122 51 139 2 314 Three-banded 65 38 113 11 227 Parenthesis 7 3 21 1 32 Thirteenspot 95 24 83 6 208 Total 556 207 651 42 1456

Non-native Sevenspot 43 36 223 35 337 Elevenspot 44 13 22 1 80 Asian 21 25 51 36 133 Variegated 11 6 49 1 67 Fourteenspot 57 27 81 31 196 Total 176 . 107 426 104 813

*Twospot: Adalia bipunctata; Eyespot: Anatis mali; Transverse: Coccinella transversoguttata richardsoni; Three-banded: Coccinella trifasciata perplexa; Parenthesis: Hippodamia parenthesis; Thirteenspot: Hippodamia tredecimpunctata tibialis; Sevenspot: Coccinella septempunctata; Elevenspot: Coccinella undecimpunctata undecimpunctata; Asian: Harmonia axyridis; Variegated: Hippodamia variegata; Fourteenspot: Propylea quatourdecimpunctata

56 2.2.2 SPECIMEN EXAMINATION AND VERIFICATION

Twenty-seven collections were located that housed lady beetle specimens from

Atlantic Canada (Table 2.2), and arrangements were made to either visit or borrow the lady beetles from each collection for verification of identifications, databasing of label information where necessary and analysis. Other collections were examined (e.g. the

Canadian Museum of Nature) but if they did not include Atlantic Canadian lady beetle specimens, they were not included here. Pinned adult specimens of the eleven species of Coccinellinae (Coleoptera: Coccinellidae) (Table 2.1) were identified using Gordon

(1985), supplemented by Gordon and Vandenberg (1991). Label data, including verbatim information on location, date, collector, and habitat information (where available) were recorded for all specimens collected up to and including 2009. Pre existing identifications were verified and miscellaneous or misfiled lady beetle specimens were identified and consolidated into main collections during visits. If specimens were not already databased, they were given unique identifier labels (usually an abbreviation of the collection/institution name and a number) which corresponded to existing databases when possible, and a copy of the database was left with each institution. Where collections were already databased by a credible source, a copy of the database was obtained and corrections were made as necessary to specimen identifications. Approximately 3460 specimens corresponding to 2269 records were examined for this study (Table 2.2). A "record" was defined within each collection as all specimens of the same species, collected from the same location/place, on the same date. The total number of records fo r each species is given in Table 2.1. Table 2.2 List of insect collections in eastern Canada which store Atlantic Canada specimens o f the 11 study species o f Coccinellinae (see Table 2.1). For each collection physically located in Atlantic Canada w ith >5 records, the radius that included 75% of records for these species is also given. Where available, collection abbreviations are from Insect and Spider Collections of the World http://hbs.bishopmuseum.org/codens/codens-inst.html.

Collection/Institution Collection Total# of Total # of Radius (km) Name and Location Abbreviation Specimens Records distance of 75% circle Government Collections Agriculture & Agri-Food Canada, ACPE 48 46 47.4 Charlottetown, PE Agriculture & Agri-Food Canada, ACNS 100 73 18.6 Kentville, NS Agriculture & Agri-Food Canada, 250 96 118.4 St. John's, NL Atlantic Forestry Centre AFC '231 119 236.4 Fredericton, NB Atlantic Forestry Service NFNR 106 55 730.0 Corner Brook, NL Canadian National Collection of CNC 344 166 Insects, Ottawa, ON Fundy National Park FNP 2 2 Alma, NB Great Lakes Forestry Centre, GLFC 4 3 Sault Ste. Marie, ON Nova Scotia Department of NSNR 230 172 76.0 Natural Resources, Shubenacadie, NS

Museum Collections New Brunswick Museum, St. NBMC 105 100 173.6 John, NB Nova Scotia Museum of Natural NSMC 213 196 116.8 History, Halifax, NS Royal Ontario Museum, ROME 24 18 Toronto, ON Continued...

58 Table 2.2 Continued

Collection/Institution Collection Total# of Total # of Radius (km) Name and Location Abbreviation Specimens Records distance of 75% circle University Collections Acadia University, Wolfville, NS 66 57 27.7 Cape Breton University, Sydney, CBU 330 295 19.6 NS Memorial University of MUNC 154 91 210 Newfoundland, St. John's, NL Nova Scotia Agricultural College, NSAC 300 172 101.9 AD Pickett Collection, Bible Hill, NS St. Francis Xavier University, STFX 48 47 12.0 Antigonish, NS Universite de Moncton, UMNB 159 151 84.3 Moncton, NB University of New Brunswick, DFRU 170 162 190.0 Fredericton, NB University of Guelph DEBU 37 29 Guelph, ON University of Prince Edward UPEI 386 73 34.9 Island, Charlottetown, PE

Private Collections D. Doucet Collection, Pelerin, NB 31 31 21.3 R. Harding Collection, 70 67 31.3 Summerville, PE B. Hicks Collection, Carbonear, 9 8 44.8 NL C. Majka Collection, Halifax, NS CGMC 25 24 115.0 Newfoundland Insectarium, NINF 4 2 Deer Lake, NL R. Webster Collection, Charters RWC 14 14 84.9 Settlement, NB

59 2.2.3 DATASET PREPARATION

The main preparation required fo r use of the database was to verify that dates were correct and to georeference locality information. Specimens which lacked geographical coordinates (approximately 77% of the specimens examined) were located using the Natural Resources Canada guide to Geographical Names o f Canada

(http://www.nrcan.gc.ca/earth-sciences/search/search_e.php), supplemented by reference to Google Earth®. The geographical names of Canada database provides coordinates relative to the North American datum of 1983 (NAD83), and usually represents the centre of a geographical feature (e.g. a town, hamlet or community)

(Canadian Permanent Committee on Geographical Names 1992). Specimens w ithout sufficient information to assign a locality for georeferencing were omitted from this study (approximately 4% of specimens). In instances where specific location descriptions were given but georeferenced coordinates could not be found, coordinates for a more general place were used (e.g., a label with a specific location of

"Headquarters" in the Fundy National Park was assigned the coordinates of the general place, Fundy National Park, New Brunswick). When less precise geographic coordinates were assigned to specimen data in the spreadsheet, this fact was recorded in the notes field of the spreadsheet. Specimen records for which the date of collection could not be defined in terms of a decade and/or season were omitted from this study.

60 2.2.4 DATA MAPPING

Collection data were mapped using Maplnfo® v.7 by first converting the longitude and latitude into decimal degrees and importing longitude, latitude, year and month of collection, genus, species and name o f collection into the Maplnfo® program.

These data were projected onto a map of eastern Canada which was retrieved from

Natural Resources Canada - GeoGratis (http://www.geogratis.gc.ca).

2.2.5 DATA ANALYSES

Potential gaps in records within the different collections were assessed by comparing numbers of specimen records in each category of collections for different spatial scales, taxonomic groupings, time periods, and seasons. The collection categories were based on the four categories outlined by Lovejoy et al. (2010): Federal government, Other government, University and Other collections. The Federal government category ("Government" in Table 2.1 and 2.2) included government-run facilities such as the CNC, Agriculture & Agri-Food Canada laboratories and national parks. Other government collections ("Museum" in Table 2.1 and 2.2) included the provincial museums from New Brunswick and Nova Scotia, as well as the Royal Ontario

Museum. University collections included university and college facilities, and Other collections for Atlantic.Canadian specimens ("Private" in Table 2.1 and 2.2) only represented private collections. Numbers of records for each comparison group (i.e. temporal, spatial, taxonomic, and seasonal) were compared within each collection category to identify whether significant differences in the comparison groups occurred within specific collection types. Analyses were then repeated with data combined from all collections to determine whether potential gaps in collecting were removed by combining collection data.

Collections were assessed for temporal gaps using two different tim e periods: by decade, and for periods relating to the establishment of the four most common non native lady beetle species. In the first analysis, numbers of records were compiled for each decade from 1900 to 2009 and compared to "expected" values using a G-test (Zar

1999). For each interval, expected values were calculated from proportions that should have occurred if collection effort was similar throughout the period of record, based on the method given in Myers et al. (2009):

(total # coll. category A in all decades) * (total # all coll. category for one decade) (total # of all coll. categories in all decades)

In the second analysis, numbers of records were compared for the periods before and after the arrival of the four common non-native lady beetles (i.e. before and after 1980) using a Mann-Whitney U-test (Statistica v.6) to determine whether overall collecting effort increased after the arrival o f novel species in a region.

Seasonal patterns in collection records (whether certain collections were dominated by specimens collected on certain dates) were assessed by sorting collection dates into three seasons (spring = March 20-June 20, summer = June 21-August 31, fall =

September 1-December 21). Seasonal patterns were compared for the different types of collections using a Kruskal-Wallis test (Statistica v.6).

Potential taxonomic biases could have included misidentifications or differences in species collected because of collector preferences. Errors due to misidentification

62 were minimized by verifying species identities of the specimens used in this study. Gaps in collecting effort may have resulted from preferential collecting of novel species i* following introduction of non-native species. These were tested by determining whether records of native vs. non-native species differed in the different collection types for the two time periods (before and after 1980) for each collection type using a

Kruskal-Wallis test (Statistica v.6).

To determine whether different collections showed different spatial patterns, spatial coverage was evaluated first by determining the number of counties represented for each species in each collection, and then by quantifying the radius comprising the majority of the collection records. In the first analysis, spatial coverage for the different types of collections was graphed by cumulatively adding the number of counties represented in each type of collection for each of the eleven species of lady beetles in the following order: government, museum, university and private. In the second analysis, spatial differences in the different categories of collections were considered to have occurred if some collections appeared to have more records from localities close to the facilities/homes in which the collections/collectors were housed. Locations for each record were plotted individually for each of the 21 collections physically based in

Atlantic Canada, and the area around the location for each collection containing the closest 75% of records was calculated for each collection category. The value of 75% was chosen to reflect the majority of the records, but not be affected by outliers. The ellipse tool in Maplnfo® v.7) was then used to create circles centrally located over the geographical location of each collection, extending to encompass 75% of the records from that collection, and then the radius and area were calculated using Maplnfo®. The

"collection area" was then compared for each category of collection with a Kruskal-

Wallis test to determine whether there were significant differences in sampling area based on collection.

2.3 RESULTS

2.3.1 TEMPORAL DIFFERENCES IN RECORDS AMONG COLLECTIONS

Collecting Effort by Decade

Each collection category showed a different trend in timing of collecting effort

(Figure 2.1) that could be evaluated by comparing observed values by decade with an expected value generated from the overall proportions (Table 2.3). The Museum collections had fairly consistent numbers of records over time, with significantly fewer than expected records in only tw o decades (1970s and 1990s; G-test, p<0.05; Table 2.3).

This pattern reflects the long period of record since the establishment of natural history museums containing Atlantic Canadian material (Table 2.4). In contrast, the majority of

Private collection records were collected in the 2000s, with significantly fewer records than expected in periods before 2000 (G-test, p<0.05; Figure 2.1; Table 2.3). The

University collection records fluctuated over time, with spurts in collecting effort in the

1940s, 1970s and particularly in the 1990s (Figure 2.1). Significantly fewer records than expected were found in University collections from 1930-1960 (G-test, p<0.05;

Table2.3), whereas significantly more records than expected were found from 1970-

2009 (G- test, p<0.05; Table 2.3). The collection pattern generally did not correspond

64 Figure 2.1 Total numbers of records for each decade and pattern of accumulation of of accumulation of pattern and decade each for records of numbers Total 2.1 Figure Number of records Number of records Number of records 250 275H 300 325i 200 225 0 0 1 125- 150- 175- 25- 50- 75- 0 - - rr HrtrtHHrtHN H t r H H t r t r H rtrt H 0)0)O><7><7)Ch0)O)0>O ocnchoitfichGhocftChot-icsm?>nor>ooa> oooooooooo O m O O N f t l U ^ M N H oooooooooo collection category and for all collections combined. collections all for and category collection records for the entire period for which records are available, separated by by separated available, are records which for period entire the for records All Collections University Decade Museum Decade Decade

rl.O -0.3 -0.4 -0.5 -0.7 - -0.9 0.0 0.1 0.2 0.6 0.8 ■o o o o.n 0) c n * 65 ■n 125- S 250- -S Z 5 150-fc 175 o £ o I/I 3

200 225 300 325 100 275 Acmltd Proportion Accumulated — 50- 75- 25 Records/Decade ■ 0

o >O O l )O lO ) _ _ N t r t r t O r n H O H O H N t S r I H A t I r V H n N i oimoioioioioioioioio r O o o o o o o o o o o o =-r- o *r n o o m Government

§ Decade Private Decade . in « N N « in 0)0)0)0 0) 0)0)0)0 O O O O O T I T I

00 /

J 9 ) I

I IM o

rl.O -0.5 - -0.7 -0.9

0.0 0.1 0.2 0.3 0.4 0.6 0.8 cuuae pooto Acmltd proportion Accumulated proportion Accumulated Table 2.3 Changes in collecting effort by decade for four categories of insect collections. a (*** p < 0.001; ** p < 0.01; * p < 0.05). Bolded observed values indicate those which are lower than expected based on the 6-test. a) Observed records ______Collection Observed Type 1900s 1910s 1920s 1930s 1940s 1950s 1960s 1970s 1980s 1990s 2000s * ** ** ------... “ ii**"- *** *** University 3 15 6 6 112 89 32 142 123 309 222 * **♦ *** Private 0 4 3 0 0 0 3 0 2 17 125 *** *** Museum 20*** 0 0 16 47 27** 35 26** 39” 39*** 69* - *«* *** • * *•« Government 0 2 10* 27 102*’ * 131*** 140 87 66 141 36 *** Total 23 21 19 49 261 247 210 225 230 506 452 b) Expected records Collection Expected6 Type 1900s 1910s 1920s 1930s 1940s 1950s 1960s 1970s 1980s 1990s 2000s University 10.7 9.8 8.9 22.8 121.6 115.1 97.8 118.8 107.2 235.8 210.6 Private 1.6 1.4 1.3 3.3 17.7 16.7 14.2 17.3 15.6 34.3 30.6 Museum 3.2 2.9 2.7 6.9 36.5 34.6 29.4 35.7 32.1 70.8 63.2 Government 7.5 6.9 6.2 16.0 85.2 80.6 68.6 83.2 75.1 165.2 147.6 a Differences between Observed and Expected tested with G-test: G = 2*[Q[ observed In observed) - (£ observed In expected)] (Zar 1999) b For each interval, Expected values were calculated as: (total # coll. type A in all decadesHtotal # all coll. types for one decade) (based on Myers et al. 2009) (total # of all coll. types in all decades)

6 6 Table 2.4 List of universities, colleges and museums with Atlantic Canadian lady beetle specimens, the year the institution was founded and the year the first Atlantic Canada specimen was collected.

Institution Year of Establishment Year of First Specimen Universities and Colleges - Acadia University 1838 1922 Cape Breton University 1951 - as St. Francis Xavier University 1985 Sydney Campus 1974 - University College of Cape Breton 2005 - renamed Cape Breton University Memorial University of 1925 - Memorial University College 1950 Newfoundland 1949 - Memorial University of Newfoundland Nova Scotia Agricultural College 1905 1906 St. Francis Xavier University 1853 - St. Francis Xavier College 1996 1866 - St. Francis Xavier University Universite de Moncton 1963 1974 University of New Brunswick, 1785 1938 Fredericton Campus University of Guelph 1862 - Ontario Veterinary College* 1899 1874-Ontario Agricultural College* 1903- Macdonald Institute* 1964 - University of Guelph University of Prince Edward 1854- St. Dunstan's University* 1916 Island 1860 - Prince of Wales College* 1969 - University of Prince Edward Island Museums Royal Ontario Museum 1912 1881 Nova Scotia Museum of Natural 1868 - Nova Scotia Museum 1900 History 1947 - Nova Scotia Museum of Science 1 9 70 - Nova Scotia Museum of Natural History 1968 New Brunswick Museum 1929 1901 * Amalgamation of institutions which led to the establishment of current university.

67 with the timing of establishment of universities in the region (Table 2.4). Finally, collecting effort in Government collections peaked in the 1960s (Figure 2.1), though significantly more records than expected were collected for the period of time between

1920 and 1969 (G-test, p<0.05; Table 2.3) and significantly fewer specimens than expected after 1979 (G-test, p<0.05; Table 2.3).

Combining the collection data showed that collecting effort increased over the period of study, resulting in three distinct periods for lady beetle collecting (Figure 2.1).

Between 1900 and the 1930s few specimens were collected (19 to 49 records per decade), then an increase in collecting was seen between 1940 and 1989 (210 to 261 specimens per decade), and again in the 1990s and 2000s (506 and 452 records in the

1990s and 2000s, respectively).

Overall Collecting Effort Before and After Establishment of Non-natives

Collecting effort before and after establishment of non-native lady beetles also varied by collection type (Figure 2.2). Government collections had significantly more lady beetle records collected before 1980 than after 1980 (Table 2.5, Mann-Whitney U- test, p = 0.037), and Private collections had significantly more records collected after < 1980 (Table 2.5, Mann-Whitney U-test, p = 0.001). University and Museum collections did not have significantly different numbers of records collected before and after 1980.

When lady beetle records were combined for the four different categories of collections there were no significant differences in the total number of lady beetle records before and after 1980 (Table 2.5, Mann-Whitney U-test, p = 0.261).

6 8 Figure 2.2 Differences in number of records reported from tw o time periods (before (before periods time o tw from reported records of number in Differences 2.2 Figure Number of records Number of records Number of records -10 140 100 120 100 120 140 -20 -20 40 50 60 70 80 20 30 10 40 60 80 40 20 80 20 60 0 0 values. data. Time periods which are significant at p < 0.05 are represented by 'a' 'a' by represented are p0.05 < at significant are which periods Time data. (based on Mann-Whitney U-test, Statistica v.6). Refer to Table 2.3 for p for 2.3 Table to Refer v.6). Statistica U-test, Mann-Whitney on (based 1980 and after 1980) for four collection categories and combined collection collection combined and categories collection four for 1980) after and 1980 Combined Collections Combined nvriyCletosPrivateCollections UniversityCollections Museum Collections Government Collections Government Collections Museum Before 1980 After 1980 After 1980 Before Before 1980 After 1980 After 1980 Before Before 1980 Before After 1980 After 69 -Q S X -Q U— z “O 3 E o O in i_ 3 E

Collection Type Biases University Private Museum Government Combined Temporal** <1980 vs. >1980 p=0.133 p=0.001 p=0.931 p=0.037 p=0.261 Seasonal* 1, 2, 3 p=0.513 p=0.563 p=0.123 p=0.019 p=0.015 Taxonomic** Native vs. Non-native p=0.001 p=0.828 p=0.121 p<0.050 p<0.001 Supplementary** Native p=0.236 p=0.023 p=0.085 p<0.050 p=0.014 <1980 vs. >1980 Non-native p<0.001 p=0.002 p=0.016 p=0.012 p<0.001 <1980 vs. >1980

* Kruskal-Wallis - Dunn's Multiple Comparisons Test, significance at p < 0.05 l=spring (Mar. 20-Jun. 20), 2=summer (Jun. 21-Aug. 31), 3 =fall (Sep. 1-Dec. 20) <1980 = records collection before 1980, >1980 = records collected between 1980 and 2009. * * Mann-Whitney U-test, significance at p < 0.05

70 2.3.2 SEASONAL DIFFERENCES IN COLLECTION EFFORT

Differences in seasonal collecting effort were noted only in Government collections (Figure 2.3), which had significantly more records collected during the summer than the fall (Table 2.5, Kruskal-Wallis, Dunn's Multiple Comparisons Test, p =

0.019). University, Private and Museum collection types did not show seasonal differences in collecting effort (Table 2.5, Kruskal-Wallis, Dunn's M ultiple Comparisons

Test, p>0.05), and even after combining lady beetle records from the four different categories of collections, there was still a trend for higher collecting effort in summer than during other seasons (Table 2.5, Kruskal-Wallis, Dunn's Multiple Comparisons Test, p = 0.015). Therefore, species with adult stages present during June 21-Aug. 31 dominated the collections and species that occur earlier or later than that time period may have been underrepresented.

2.3.3 TAXONOMIC DIFFERENCES IN COLLECTING EFFORT

University and Government collections had significantly more records of native species than non-native species overall (Mann-Whitney U-test, p<0.05; Figure 2.4; Table

2.5), as did the entire dataset with all records combined (Mann-Whitney U-test, p<0.05;

Table 2.5). Government collections had significantly more records of native species collected before 1980 than after 1980 (Mann-Whitney U-test, p<0.001; Figure 2.5; Table

2.5), suggesting that workers in these collections focused on acquiring novel species after non-native species appeared in the area. In contrast, Private collections had significantly more records of native species collected after 1980 than before 1980

(Mann-Whitney U-test, p = 0.023, Figure 2.5; Table 2.5), reflecting an overall pattern for

71 University Collections Private Collections 140 40 120 i/i 100 Y 30 8 25 80 QJ £ 20 60 o V_ 40 0) -Q 10 E 20 3 z 0 -20 Spring Summer Fall Spring Summer Fall

Museum Collections Government Collections 80 120 70 V) 100 60 T3 O 80 50

Combined Collections 140 a 120 100 80 60 Key: 40 Box=upper and lower quartiles 20 Square inside box = median I n 1 0 Whiskers = range of data -20 Spring Summer Fall

Figure 2.3 Differences in number of records reported from three seasonal time periods for different collection categories and combined collection data. Seasons which are significant at p < 0.05 are represented by 'a' (based on the Kruskal- Wallis, Dunn's Multiple Comparisons Test, Statistica v.6). Refer to Table 2.3 for p values. Figure 2.4 Differences in number of records of native versus non-native lady beetles for for beetles lady non-native versus native of records of number in Differences 2.4 Figure Number of records Number of records Number of records 40 40 45 30 30 35 100 120 t 140 20 25 10 15 140 100 120 -20 -5 -20 40 40 60 60 80 20 0 5 40 80 20 60

0 0 i

test, Statistica v.6). Refer to Table 2.3 for p values. for 2.3 Table to Refer v.6). Statistica test, are significant at p < 0.05 are represented by 'a' (based on Mann-Whitney U- Mann-Whitney on (based 'a' which Records by data. represented are p<0.05 at collection significant combined are and categories collection different Combined Collections Combined nvriyCletosPrivateCollections UniversityCollections Museum Collections Museum Native aieNon-native Native Native Non-Native Non-native 73 JD -3 z I X T3 z 3 E O) o 14 3 16 5 . u 3 E (U

20 20 22 -10 12 18 30 30 40 20 i 50 10 o

Whiskers = range o f data f o =range Whiskers median = box inside Square Box=upperand lower quartiles quartiles lower Box=upperand Key: Government Collections Government Native Native Non-native Non-native Figure 2.5 Differences in number of records of native lady beetles reported from two from reported beetles lady native of records of number in Differences 2.5 Figure Number of records Number of records Number of records -10 -10 -10 0 ■ 40 30 40 20 30 40 20 50 20 30 50 50 10 10 10 0 0 0

Time periods which are significant at p < 0.05 are represented by 'a' (based (based 'a' by represented are -10 40 -1 20 30 50 10 0 2 3 4 6 1 5 0 Government Collections Government Whiskers = range of data of =range Whiskers median = box inside Square Box=upperand lower quartiles quartiles lower Box=upperand Key: Before 1980 After 1980 After 1980 Before Before 1980 After 1980 After 1980 Before Private Collections more recent collecting in the Private collections examined. More records of native lady

beetles were collected before 1980 than after 1980 even when all collections were combined (Mann-Whitney U-test, p = 0.014; Figure 2.5; Table 2.5). All collection categories had significantly more non-native species records after 1980 than before

1980 (Mann-Whitney U-test, p < 0.050; Figure 2.6; Table 2.5), since most common non

native lady beetles became established in Atlantic Canada around 1980.

2.3.4 SPATIAL DIFFERENCES IN COLLECTING EFFORT

The spatial coverage of the lady beetle data increased for all eleven species as the different collection categories were added (Figure 2.7). One species ( Anatis mali)

reached greatest spatial coverage with Government and Museum collection data. Three species ( Coccinella t. richardsoni, Hippodamia parenthesis and Coccinella septempunctata) reached greatest spatial coverage with Government, Museum and

University collection data. The remaining seven species reached greatest spatial coverage when all four collection categories were combined.

The different categories of collections showed dramatic differences in the extent of collecting area comprising 75% of the records (Figure 2.8). Specimens in Government collections were collected from a similar geographical extent as Museum collections, and both of these were more extensive geographically than University or Private collections (Table 2.6, Kruskal-Wallis and Dunn's Multiple Comparisons Test, p < 0.05).

75 Figure 2.6 Differences in number of records o f non-native lady beetles reported from from reported beetles lady non-native f o records of number in Differences 2.6 Figure Number of records Number of records Number of records 24 20 22 140 10 12 14 16 18 100 120 140 100 120 -2 -20 -20 0 4 40 80 40 80 6 8 20 60 20 60 2 0 0 values. two time periods for different collection categories and combined collection collection combined and categories collection different for periods time two data. Time periods which are significant at p < 0.05 are represented by 'a' 'a' by represented are p0.05 < at significant are which periods Time data. (based on Mann-Whitney U-test, Statistica v.6). Refer to Table 2.3 for p for 2.3 Table to Refer v.6). Statistica U-test, Mann-Whitney on (based Combined Collections Combined 1980 After 1980 Before nvriyCletosPrivate Collections University Collections Museum Collections Museum Before 1980 After 1980 After 1980 Before Before 1980 After 1980 After 1980 Before 76 .a T3 z -Q 73 3 E a to o o in 3 E i

20 22 10 12 14 16 18 -10 -2 20 30 40 50 10 0 4 8 2 6 0 Whiskers = range of data of range = Whiskers median = box inside Square Box=upper and lower quartiles quartiles lower and Box=upper Key: Government Collections Government Before 1980 After 1980 After 1980 Before □ 25%-75% □ Before 1980 After 1980 After 1980 Before X Min-Max X ° Median Median ° Figure 2.7 Spatial coverage for collection categories graphed by cumulatively adding bycumulatively graphed categories collection for coverage Spatial 2.7 Figure

Accumulated counties ^ Accumulated counties ^ Accumulated counties ^ Accumulated f j counties Coccinella septempunctata * z . z z 20 25 30 35-| 30- 351 i k i k £ Hlppodamla varlegata Hlppodamla W / / : Coccinella perplexa t.

<5- dlabpntt Anatismali bipunctata Adalia A*' ^ f s Z Z ntefloigodr oenet uem nvriyad private. and university museum, government, collection of order: category each in following the specimens inby represented counties of number

£ ■a i < 10 i 0 s 3 £ u u 3 E 3 Coccinella undeclmpunctata u. 25- g 1 / <

ZZ.Z Hlppodamla parenthesis Hlppodamla 20- 30- 35-i 25- 35- 35-1 10- 15 ^ r > V / quatuordecimpunctata 0- Z Z Z Z z z z z ■ III Propylea z z z 77 z

10 E z 35-1 35-1 351 Coccinella richardsoni t. y z Hlppodamla * Harmonla axyrldis Harmonla z " z t. t. tibialis

Figure 2.8 Differences in collecting area (km ) which includes 75% of records for each for records 75% of includes )which (km area collecting in Differences 2.8 Figure Area (millionsof square kilimetres) 0.8xl06 0.8xl06 0.2xl06 0.2xl06 0.4xl06 0.6xl06 1.2xl06 1.2xl06 - 1.4xl06 - 1.6xl06 1.8x10 .xO •l.OxlO6 significantly different are represented by a, b or c (based on Kruskal-Wallis, Kruskal-Wallis, on (based c bor a, by are which represented are categories Collection different significantly category. collection by grouped collection, Dunn's Multiple Comparisons Test, Statistica v.6). Statistica Test, Comparisons Multiple Dunn's 0 University Whiskers = range of data of range = Whiskers Square inside box = median median = box inside Square Box=upper and lower quartiles quartiles lower and Box=upper Key: rvt uemGovernment Museum Private 78 a,c Table 2.6 Kruskal-Wallis and Dunn's Multiple Comparisons test used to assess spatial differences in four categories of collections.

University Private Museum Government University p=0.011 p=0.089 p<0.001 Private p=0.011 p<0.001 p<0.001 Museum p=0.089 p<0.001 p>0.100

79 2.4 DISCUSSION

Lady beetle collection records from individual collection categories showed differences in collecting effort, resulting in temporal, seasonal, taxonomic and spatial gaps in data that could bias interpretations of distribution patterns. Records from the

Government collection category showed highest collecting effort before the approximate 1980 establishment of common non-native species (McCorquodale 1998,

Hoebeke and Wheeler 1996, Majka and McCorquodale 2006, Wise et al. 2001, Hicks et al. 2010). Despite a pattern for more collecting of native species prior to 1980 and more non-native species after 1980, as well as a bias for collecting in summer (versus the fall), these collections still showed high numbers of lady beetle records. In addition, although records from University and Private collection categories showed a spatial bias for collection close to the institutions, it was clear that when all collections were combined, there was good spatial coverage over the Atlantic Provinces. Private collections were also biased temporally, showing the highest amount of collecting since 1990, and especially since 2000. University collections were also biased taxonomically, showing higher numbers of native species than non-native species. By identifying gaps in the data for different collection categories prior to combining all data into one dataset, it was found that the combined dataset had fewer gaps which could bias interpretation of distribution patterns. This suggests it is possible to use combined collection data to evaluate temporal changes in lady beetles in Atlantic Canada.

80 The main temporal bias that has been identified in this study, and which should be considered when assessing lady beetle temporal patterns, is a gradual increase in the number of specimen records collected between 1900 and 2009. This increase in collecting effort should actually be beneficial to using lady beetle collection records to assess change in lady beetle populations, especially since these data also show that numbers of lady beetle species collected after 1980 continues to be high, despite the pattern in Government collections for more records from before 1980 than after 1980.

Early in the 20th century, the importance of maintaining natural history collections was not yet fully understood and there were few institutions available to provide the resources necessary for long-term retention of specimens (Fairweather and McAlpine

2011). The increase in sampling during the middle of the century was largely driven by extensive regional government surveys such as the Forest Insect and Disease Survey

(FIDS) which ran for four decades after its start in 1932 (Power 1988). The large jum p in overall collecting effort since 1990 can be attributed to a combination of greater emphasis on student collecting in regional universities, as well as the interest in lady beetles that was generated by the arrival of non-native species Coccinella septempunctata (Gordon 1985, Gordon and Vandenberg 1991), Propylea quatuordecimpunctata (Gordon 1985, Gordon and Vandenberg 1991), Hippodamia variegata (Gordon and Vandenberg 1991) and Harmonia axyridis (Gordon and

Vandenberg 1991). The concern that a reduction in specimen records could be confused with overall reductions (McCarthy 1998) was eliminated by considering a wide range of collection categories in this study; therefore, any reduction of a particular species during this period can be considered to be an actual reduction, and not an artefact of collecting effort.

The spatial bias that has been identified in this study, and which should be considered when assessing lady beetle distribution patterns, is the tendency to collect specimens in close proximity to institutions which house University and Private collections. The limited spatial coverage of these collection data was apparent when assessing collection categories individually; however, it was clear that as collection data were combined, spatial representation of the Atlantic Provinces began to improve. For seven of eleven species of lady beetles, data from all four categories of collections were required to reach the greatest spatial coverage; for three species, data from three categories of collections were required to optimize geographic area; and for one species only, the greatest spatial coverage of the Atlantic Provinces was reached with data from two categories of collections. The concern that specimen records may not accurately reflect the geographic extent of a lady beetle's distribution was minimized by considering a wide range of collection categories in this study; therefore, any range contraction or expansion experienced by a particular species during this period can be considered to be an actual pattern, and not a reflection of collecting effort.

A major concern in this study was potential for collections to have a bias towards collecting, or at least retaining, targeted species (e.g. novel species or species which inhabit agricultural areas). Since many collections have limited space and maintenance budgets (Lovejoy et al. 2010), a trend towards retaining only new material in a collection could give a misleading impression of a decline in certain groups concurrent with an increase in new ones (Acorn 2007). An important finding in this study was that both overall lady beetle and native lady beetle collecting were higher following 1980 (the period when non-native species began to become established in Atlantic Canada) than before 1980, indicating that collecting of native species continued, despite the establishment of the new arrivals. This pattern was driven mainly by the general surveys in government labs and by university students, confirming the importance of combining data from multiple collection categories before attempting to assess distribution patterns. Similarly, there was some concern that certain categories of collections might focus more on certain habitats (such as agricultural habitats from

Agriculture & Agri-Food Canada labs) (Margules and Austin 1994, Williams etal. 2002,

Graham et al. 2004, Elith et al. 2006). Habitat patterns were difficult to assess in this study, since habitat information was not listed on most collection labels. However, spatial biases were clearly addressed by combining collection data; thereby reaching the greatest spatial coverage of the data and reducing the likelihood of biased collecting from specific habitats. Noticeable differences in collecting effort for the different categories of collections were also expected due to factors such as timing of establishment of academic institutions and/or the arrival of entomologists to these institutions. The peaks in collecting effort, however, did not coincide with the establishment time of universities, so more details on when specific entomologists began working in the universities may be required to understand the University patterns. The apparent trend for recent collecting efforts in Private collections may be explained by the fact that many older private collections have been assimilated into other types of collections and are counted with those collections. Two examples of this phenomenon include an early private collection of Robert Mutch which was incorporated into the UPEI insect collection (D. Giberson, UPEI, personal communication), and the private collection of John and Bertha Carr (containing

~100,000 specimens) which was donated to the CNC in 2001 (Larson 2006).

Interestingly, Museum collections were expected to show temporal bias due to reduced collecting once specimens were already included in a collection (D. Giberson, UPEI and

D.B. McCorquodale, CBU, personal communication), but Museum collections in this study retained fairly consistent numbers of records over time. The three museums included in this study continue to acquire new material through on-going research programs (e.g. a project by C. Majka at the N.S. Museum to identify all Coleoptera in

Atlantic Canada) and donations.

The seasonal bias found in this study (collecting during the summer months), is not expected to affect interpretation of lady beetle distribution patterns. In regions of the Palaearctic where the ecoregion is comparable to Atlantic Canada (Olson et al.

2001), it has been reported that common lady beetle species (e.g. Hippodamia tredecimpunctata, Coccinella septempunctata and Propylea quatuordecimpunctata) start emerging from hibernation at the end of April, with new adults emerging from July through August regardless of the number of generations produced in one year

(Kuznetsov 1997). Therefore, it is reasonable to assume all lady beetle taxa which occur in Atlantic Canada are also likely to have adults present during the summer months, though they may differ in detectability at different times in the summer due to specific lifecycle timing for adult emergence or overwintering aggregations (Iperti 1999). Since sampling focused on the main summer activity period, however, this bias is not expected to affect interpretation of lady beetle distribution patterns.

Based on the results of this study, I conclude that individual natural history collection categories have gaps in collection data that could interfere with interpretation of lady beetle distributions, but these can be minimized by combining data from multiple collection categories. In particular, the high collecting effort and specimen retention of lady beetles since 1990 means that natural patterns should generally reflect patterns shown in the combined collections data set, and any spatial bias has been minimized by the broad geographic range of the collections investigated.

85 2.5 LITERATURE CITED

Acorn, J. 2007. Ladybugs of Alberta, finding and connecting the dots. The University of Alberta Press, Edmonton, Alberta. 169 pp.

Belicek, J. 1976. Coccinellidae o f western Canada and Alaska with analyses of the transmontane zoogeographic relationships between the fauna of British Columbia and Alberta (Insecta: Coleoptera: Coccinellidae). Quaestiones Entomologicae. 12: 283-409.

Canadian Permanent Committee on Geographical Names. 1992. Guide to the field collection of Native geographical names (provisional edition). Retrieved January 26, 2012, from http://www.nrcan.ec.ca/sites/www.nrcan.ec.ca.earth- sciences/files/pdf/native field euide e.pdf.

Colla, S.R. and Packer, L. 2008. Evidence for decline in eastern North American bumblebees (Hymenoptera: Apidae), with special focus on Bombus affinis Cresson. Biodiversity and Conservation. 17:1379-1391.

Colla, S.R. and Ratti, C.M. 2010. Evidence for the decline of the western bumble bee (Bombus occidentalis Greene) in British Columbia. The Pan-Pacific Entomologist. 86: 32-24.

Cotterill, F.P. 1995. Systematics, biological knowledge and environmental conservation. Biodiversity and Conservation. 4:183-205.

Danks, H.V. 1991. Museum collections: fundamental values and modern problems. Collection Forum. 7: 95-111.

Fairweather, A.D. and McAlpine, D.F. 2011. History and status of the Natural History Society of New Brunswick entomology collection: 1897-1931. Journal of the Acadian Entomological Society. 7:14-19.

8 6 Fitzpatrick, U., Murray, T.E., Paxton, R.J., Breen, J., Cotton, D., Santorum, V. and Brown, M.J.F. 2007. Rarity and decline in bumblebees - a test of causes and correlates in the Irish fauna. Biological Conservation. 136:185-194.

Gordon, R.D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Entomological Society. 95:1-912.

Hicks, B., Majka, C.G. and Pardy Moores, S. 2010. Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) found in Newfoundland. The Coleopterists Bulletin. 64: 50.

Iperti, G. 1999. Biodiversity of predaceous Coccinellidae in relation to bioindication and economic importance. Agriculture, Ecosystems and Environment. 74: 232-342.

Kuznetsov, V.N. 1997. Lady beetles o f the Russian Far East. Center for Systematic Entomology, Gainesville. 248 pp.

Larochelle, A. 1979. Les Coleopteres Coccinellidae du Quebec. Cordulia (Supplement). 10:1-111.

Larson, D. 2006. Biography of Bertha and John Carr. Entomological Society of Alberta Carr Biography. Retrieved August 22, 2011, from http://www.entsocalberta.ca/carrbio.htm.

87 Lovejoy, T.E., Brouillet, L., Doolittle, W.F., Gonzolas, A., Green, D.M., Hall, P., Herbert, P., Herrmann, T.M., Hyde, D., Lee, J., Maddison, W.P., Otto, S.P., Sperling, F. and Thompson, R.P. 2010. Canadian taxonomy: exploring biodiversity, creating opportunity. Report for the Government of Canada, Ottawa. 126 pp.

Majka, C. G. and McCorquodale, D.B. 2006. The Coccinellidae (Coleoptera) of the Maritime Provinces of Canada: new records, biogeographical notes, and ^ conservation concerns. Zootaxa. 1154: 49-68.

Margules, C.R., and Austin, M.P. 1994. Biological models fo r monitoring species decline: the construction and use of data bases. Philosophical Transactions, Royal Society, London B. 344: 69-75.

M arriott, S.M., Giberson, DJ. and McCorquodale, D.B. 2009. Changes in the status and geographic ranges of Canadian lady beetles (Coccinellidae) and the selection of candidates for risk assessment. Part I Foundation Report. Report to COSEWIC Arthropods Species Specialist Committee. 53 pp.

McCarthy, M.A. 1998. Identifying declining and threatened species with museum data. Biological Conservation. 83: 9-17. -

McCorquodale, D.B. 1998. Adventive lady beetles (Coleoptera: Coccinellidae) in eastern Nova Scotia, Canada. Entomological News 109:15-20.

Newbold, T. 2010. Applications and limitations of museum data for conservation and ecology, with particular attention to species distribution models. Progress in Physical Geography. 34: 3-22.

Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D'Amjco, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wettengel, W.W., Hedao, P., and Kassem, K.R. 2001. Terrestrial ecoregions of the world: a new map of life on earth. BioScience. 51: 933-938.

8 8 Power. J.M. 1988. Decision support system for the forest insect and disease survey and for pest management. Forestry Chronicle. 64:132-135.

Pyke, G.H., and Ehrlich, P.R. 2010. Biological collections and ecological/environmental research: a review, some observations and a look to the future. Biological Reviews. 85: 247-266.

Reddy, S., and Davalos, L.M. 2003. Geographical sampling bias and its implications for conservation priorities in Africa. Journal of Biogeography. 30:1719-1727.

Soberon, J.M., Llorente, J.B., and Onate, L. 2000. The use of specimen-label databases for conservation purposes: an example using Mexican papilionid and pierid butterflies. Biodiversity and Conservation. 9:1441-1466.

Williams, P.H., Margules, C.R., and Hilbert, D.W. 2002. Data requirements and data sources for biodiversity priority area selection. Journal of Bioscience. 27: 327- 338.

Wise, I.L., Turnock, W.J. and Roughley, R.E. 2001. New records of coccinellid species for the province of Manitoba. Proceedings of the Entomological Society of Manitoba. 57: 5-10.

Zar, J.H. 1999. Biostatistical Analysis. 4th Edition. Prentice Hall, New Jersey.

89 CHAPTER 3

USING OCCURRENCE DATA FROM COLLECTIONS TO EVALUATE RELATIVE ABUNDANCE / AND GEOGRAPHIC RANGE OF LADY BEETLE SPECIES IN EASTERN CANADA

90 ABSTRACT

Native species of lady beetles are believed to be declining in distribution and/or relative abundance across North America, but an accurate broad scale picture is difficult to obtain because, to date, only localized studies exist (summarized by Wheeler and

Heobeke 1995, Harmon et at. 2007). One source of data, information from specimens in natural history collections, has been effective for assessing changes in distribution for a wide variety of taxa, but has not been used on a broad scale for lady beetles in Canada.

Therefore, the goal of this study is to assess temporal patterns in distribution and relative abundance in a suite of five native and five non-native lady beetle species in eastern Canada using data from collections. Observed patterns in relative abundance were compared to expected patterns using a G-test to determine whether abundance was higher or lower than expected for six time periods (covering a period from 1881 to

2009). Spatial patterns were assessed by plotting locality data and then applying '

'rubber banding' in GIS software to calculate geographic area of occupancy for each time period. There was a clear decline in relative abundance and geographic extent for four native species ( Adalia bipunctata, Coccinella novemnotata, Coccinella transversoguttata richardsoni and Hippodamia tredecimpunctata tibialis), but not for

Coleomegilla maculata lengi. In contrast, four non-native species showed an increase in relative abundance from the time of establishment to the end of the study period in

2009, despite some minor range contractions in more recent decades. This study provides strong evidence for the decline over a wide geographic area in four native species of lady beetles in eastern Canada. 3.1 INTRODUCTION

An increasing number of studies document local or regional declines in native lady beetles throughout North America (Wheeler and Hoebeke 1995, Harmon et al.

2007 and reviewed in M arriott et al. 2009, and also see Table 1.1 pp. 2). Two species in particular that have been reported to be declining in eastern North America are

Coccinella novemnotata (Wheeler and Hoebeke 1995, Marshall 2008, Harmon et al.

2007, Skinner and Domaine 2010, Hesler and Kieckhefer 2008, Fothergill and Tindall

2010) and Coccinella transversoguttata richardsoni (Elliott et al. 1996; Acorn 2007;

Hesler et al. 2004; Turnock et al. 2003; Hesler and Kieckhefer 2008; Boiteau et al. 1999;

Boiteau 1983; Alyokhin and Sewell 2004; Finlayson et al. 2008). Two additional species,

Adalia bipunctata (Elliott etal. 1996, Colunga-Garcia and Gage 1998, Boiteau etal. 1999,

Cormier et al. 2000, Hesler et al. 2004, Hesler and Kieckhefer 2008, Fothergill and Tindall

2010) and Hippodamia tredecimpunctata tibialis (Boiteau 1983, Boiteau et al. 1999,

Alyokhin and Sewell 2004, Lucas e tal. 2007, Finlayson etal. 2008) have conflicting reports regarding geographic distribution and relative abundance. Possible reasons for declines include habitat alteration (Harmon etal. 2007), parasites, pathogens, and parasitoids (Riddick et al. 2009), competition from non-native lady beetles (e.g. Lucas et al. 2002, Michaud 2002, Cottrell and Shapiro-llan 2003, Yasuda et al. 2004) and interspecifc predation (Evans 1991). However, no studies have attempted to evaluate abundance or distribution patterns at a broad scale to determine the extent of declines or even if they occur at a wider scale.

92 Determining historic ranges and relative abundance of species which are currently thought to be declining is a recurring challenge (Shaffer et al. 1998). Insects are particularly difficult to assess because many species are highly motile, and have patchy distributions scattered across broad geographic ranges (Koch and Strange 2009).

Therefore, obtaining as much information as possible about species ecology, natural history and geographic ranges is a crucial first step in conservation planning (Ferrier

2002, Funk and Richardson 2002, Rushton e ta l. 2004, Newbold 2010), even though for many species such detailed information is not known or not readily available (Elith et al.

2006).

Natural history collections (or "collections") throughout the world contain >2.5

billion preserved biological specimens (Cotterill 1995) which have been retained from decades of taxonomic, ecological, and biodiversity studies (Danks 1991, Cotterill 1995,

Lovejoy et al. 2010). These collections represent most of what is known about

biodiversity and geographic ranges of flora and fauna worldwide (Johnson 2007).

Collection data can be used to evaluate spatial and temporal trends in relative abundance and geographic range (Chapman 2005, Newbold 2010) as long as potential limitations in using collection data (see Chapter 2) are considered and accounted for.

Lady beetles are well represented in natural history collections of all types so historical data from museums should work well to evaluate long term patterns. Their abundance in a wide variety of collection types means that potential temporal and spatial biases in collection data can be minimized or eliminated through examining multiple collections (Chapter 2). The goal o f this study is to assess temporal patterns in distribution and relative abundance in a suite of native and non-native lady beetle species commonly occurring in eastern Canada to determine whether there is broad- scale evidence in native species, and if these coincide with arrival and spread of non native species. Declines, if they are present, are expected to be seen in both the relative abundance of specimens in collections and in the geographical extent of the collection localities. Therefore, the species assessed in this study include the native species with clearly documented local declines ( Coccinella novemnotata and Coccinella transversoguttata richardsoni) and two species with suspected declines (Adalia bipunctata and Hippodamia tredecimpunctata tibialis). A fifth native species,

Coleomegilla maculata lengi, which has been reported to have remained unchanged in southern Ontario and Quebec and neighbouring US (Elliott etal. 1996, Obrycki etal.

1998a, Colunga-Garcia and Gage 1998, Day and Tatman 2006, Mignault etal. 2006,

Lucas et al. 2007) is included in this study to monitor collecting effort over time.

Distributions and relative abundance of the five established non-native species

(Coccinella septempunctata, Coccinella u. undecimpunctata, Harmonia axyridis,

Hippodamia variegata and Propylea quatuordecimpunctata) will be assessed concurrently with examination of the native species. Based on the literature (e.g. Brown

1940, Chantel 1972, Belicek 1976, Larochelle 1979, Wheeler and Hoebeke 1981, Gordon

1985, Gordon and Vandenberg 1991, Coderre et al, 1995, Hoebeke and Wheeler 1996,

McCorquodale 1998, Turnock et al. 2003, Majka and McCorquodale 2006), these species show clear evidence of range expansion and increases in relative abundance in collections following their establishment in a region. 3.2 MATERIALS AND METHODS

Approximately 18,000 specimens of the 10 study species were examined and identified or verified using Gordon (1985) and Gordon and Vandenberg (1991). The specimens represented all the specimens from localities in ON, QC, NB, NS, PE and NL that had been deposited in 32 collections located in eastern Canada (Table 3.1).

Approximately half of these specimens, mainly those in smaller regional collections, had not been previously databased. Specimens which lacked geographical coordinates

(approximately 92% of the specimens examined) were georeferenced relative to the

North American datum of 1983 (NAD83) using the Geographical Names o f Canada

(http://www.nrcan.gc.ca/earth-sciences/search/search_e.php) database, supplemented by reference to Google Earth®. If specific location descriptions were given but coordinates could not be found, coordinates fo r a more general place were used (e.g., a label with a specific location of "Headquarters" in the Fundy National Park was assigned the coordinates of the general place, Fundy National Park, New Brunswick). In some cases this will result in slight errors in plotting, since the Geographical Names of Canada database location usually represents the centre of a geographic feature (e.g. a town, hamlet or community) (Canadian Permanent Committee on Geographical Names 1992) whereas the actual species may have been collected a few kilometres from the centre.

When less precise geographic coordinates were assigned to specimen data in the spreadsheet, this fact was recorded in theliotes field of the spreadsheet. Specimens without sufficient information to assign a locality for georeferencing were omitted from this study (approximately 7% of specimens).

95 Table 3.1 Collections from which data for this study were assessed, including location, contact information of curator or collection manager (MM: Meghan Marriott; DBMcC: David McCorquodale; CM: Chris Majka; DD: Denis Doucet), and total number of specimens for Ontario, Quebec, New Brunswick, Nova Scotia and ______Newfoundland and Labrador for the 10 study species. ______Prov. Institution Contact Notes ON Canadian Museum Robert Anderson 323 specimens. of Nature [email protected] . Verified and databased by MM, November 2008 ON CFS Great Lakes Kathryn Nystrom 332 specimens. Forestry Centre knystrom(5)nrcan-mcan.gc.ca Verified and databased by MM, May 2009. ON Canadian National Patrice Bouchard 6641 specimens. Collection of Insects [email protected] Verified and databased mostly by CNC staff, and partially by Roughley and DBMcC (Manitoba west) and MM and DBMcC (Ontaro east - 3108 specimens, 2009). ON University of S.A. Marshall 3641 specimens. Guelph samarsha(S)uoguelph.ca Databased by MM, November 2008 and May 2009.

ON Royal Ontario Doug Currie 1099 specimens. Museum dcurrie(S)zoo.utoronto.ca Databased by MM, November Brad Hubley, Collection 2008. Manager [email protected]

QC Complexe Celine Piche 977 specimens. Scientifique du [email protected] Verified and databased by Quebec Michele Roy MAPAQwith assistance from [email protected] MM, April 2009.

QC Rene Martineau Jan Klimaczewski 213 specimens. Insectarium, [email protected] Verified and databased by MM, Laurentian Forestry April 2009. Centre QC Laval University Conrad Cloutier, 655 specimens. [email protected] Verified and databased by MM, April 2009. QC Lyman Terry Wheeler, 137 specimens. Entomological Stephanie Boucher Verified and databased by Museum, Stephanie. [email protected] DBMcC and MacDonald in 2006. McDonald Campus, McGill University ... Continued

96 Table 3.1 Continued Prov. Institution Contact Notes NB Atlantic Forestry Jon Sweeney 174 specimens. Centre isweenev(3)nrcan.gc.ca Verified and databased by MM, Ed Hurley August 2008. ehurlev(5)nrcan.ec.ca NB University of New Dan Quiring 324 specimens. Brunswick, Forestry auiring(S)unb.ca Verified and databased by DBMcC, 2007. NB New Brunswick Don McAlpine 149 specimens. Museum Donald. McAloine(S>nbm- Databased by museum and MM, mnb.ca verified by DBMcC, MM and DD, April 2009. NB Universite de Gaetan Moreau, Pauline Duerr 113 specimens. Moncton oauline.duerr(5)umoncton.ca Verified and databased by CM and DD, April 2009. NB Fundy National Park Obtained from ChrisMajka 2 specimens. c.maika(5)ns.svmoatico.ca Verified and databased by CM.

NB D. Doucet Denis Doucet 21 specimens. Collection ddodguv57(®gmail.com Verified and databased by DD.

NB R. Webster Reginald Webster 9 specimens. Collection Verified and databased by CM. NS St. Francis Xavier Randy Lauff 46 specimens. University rlauff(S)stfx.ca Verified and databased by CM.

NS Acadia University Soren Bondrup-Nielsen 59 specimens. soren.bondruo- Verified by MM and DBMcC, nielsen(S)acadiau.ca databased by MM, September 2009. NS Cape Breton David McCorquodale 306 specimens. University David McCorauodale(S)cbu. Databased by CBU students, ca verified by DBMcC. NS Nova Scotia Peter Hicklenton 73 specimens. Agriculture & Agri- Deter. hicklentonOagr.gc.ca Databased by AAFC. Food Canada - Kentville NS Nova Scotia Jeff Ogden 115 specimens. Department of ogdenib(S>gov.ns.ca Databased by J. Ogden, updated Natural Resources - by MM, April 2009. Shubenacadie ... Continued

97 Table 3.1 Continued Prov. Institution Contact Notes NS Nova Scotia Chris Cutler 230 specimens. Agricultural College [email protected] Verified and databased by MM, August 2009. NS Nova Scotia Andrew Hebda, Curator of 172 specimens. Museum of Natural Zoology Databased by NS Museum, History [email protected] verified by CM and DBMcC.

NS C. Majka Collection Chris Majka 31 specimens. c.maika^ns.svrnoatico.ca ~ Verified and databased by CM.

NS J. Ogden Collection Jeff Ogden 30 specimens. [email protected] Databased by J. Ogden, verified by MM, April 2009.

PE Agriculture & Agri- Christine Noronha 34 specimens. Food Canada - [email protected] Databased by AAFC. Charlottetown

PE University of Prince Donna Giberson 435 specimens. Edward Island, [email protected] Verified and databased by MM, Charlottetown September 2009.

NL Memorial David Langor 83 specimens. University of [email protected] Verified and databased by D. Newfoundland, St. Langor. John's

NL Newfoundland Lloyd Hollet 7 specimens. Insectarium [email protected] Verified and databased by MM, August 2009. NL Agriculture & Agri- Peggy Dixon 214 specimens. Food Canada - St [email protected] Databased by AAFC, updated by John's MM, August 2009.

NL Atlantic Forestry Lucie Royer 70 specimens. Service, Corner Lucie. Rover@NRCan- Verified and databased by MM, Brook Office RNCan.gc.ca August 2009.

NL B. Hicks Collection Barry Hicks 7 specimens. [email protected] Databased by MM, February 2010, indenfications verified by CM.

98 A "record" was defined as all specimens of the same species, collected from the same location/place, in the same year. If the decade of collection could not be discerned from the collection label, the specimen was omitted from the study. Records for each species were sorted into six time periods to assess temporal changes in distribution of the target species. Due to relatively low collecting effort prior to 1960

(see chapter 2), all records prior to 1960 were combined into one tim e period to create approximately equal number of records in each tim e period. Distribution patterns were then assessed by decade ("time period") for pre-1960,1960s, 1970s, 1980s, 1990s and

2000s (covering a period from 1881 to 2009). These six tim e periods encompass pre introduction (pre-1960s and 1960s), colonization (1970s and 80s) and subsequent spread (1990s and 2000s) of the four most common of the five non-native species which occur in Canada. For each species, relative abundance in collections and geographical distribution and extent were calculated for each tim e period. Potential geographic bias due to low collecting effort in Newfoundland and Labrador (NL) relative to other parts of

Canada was assessed by looking at patterns with and without the NL data. Data were plotted for the entire region, and for the region with Newfoundland and Labrador excluded, in case the low collecting effort in Newfoundland and Labrador influenced the area or abundance patterns.

Relative abundance was calculated by dividing the total number of records for each species by the total number of records for all ten species for the time period.

Expected values were calculated by determining the relative proportions of records for

99 each species and each time period that should have occurred had proportions remained constant over time (Myers et al. 2009):

(total # of species A in all time periods) * (total # o f all species for one time period) (1) (total # of all species in all tim e periods)

Differences between observed and expected records across time periods were tested using the G-test (Zar 1999):

G = 2*[(2 observed In observed) - (X observed In expected)] (2)

Geographical patterns were evaluated by determining the area encompassed by records for each species for each time period in eastern Canada. To plot the localities, data were given a North America Lambert Conformal Conic projection with a NAD83 datum. The Lambert Conformal Conic Projection visually represented shape more accurately than area; however, this projection's linear unit of metres allowed for the accurate measurement of area (distribution) for each species for each time period

(Environmental Systems Research Institute, Inc., 2007). A visual inspection of the data was carried out to check for erroneous records (e.g. points outside of geographic range shown in Gordon 1985), and corrections were made where necessary. For each species for each time period a polygon was created around the outermost records using the convex hull tool in Maplnfo® (RouteWare, http://www.routeware.dk/toolbox.php). This technique, also known as 'rubber banding', produced polygons which consisted of the least number of points necessary to ensure all records fell within or along the edge of the polygons with no interior angle greater than 180 degrees (Maplnfo Corporation,

1 0 0 2007). The resultant polygons were then clipped to the extent of the study area

(eastern Canada) to remove portions of polygons which extended into the ocean or the

United States, and area was determined from these new polygons. Geographical areas were then plotted for each species for each time period to assess changes overtim e.

3.3 RESULTS

Four of the five native lady beetles declined in relative abundance while most non-native lady beetles increased. This pattern was generally similar regardless of whether or not Newfoundland and Labrador was excluded from the analysis. Trends in relative abundance (Figure 3.1a and b) were paralleled by changes in geographic distribution, though reductions in area lagged behind the pattern for abundance for native species (Figures 3.2a and b, and Appendix Figures A -l to A-5 and A - ll to A-15).

The two native species with documented declines from previous studies also showed declines here. Coccinella novemnotata declined 100% in relative abundance and geographic distribution and represented a steady and significant decline in observed number of records across the entire study period (Figures 3.1 and 3.2;Table 3.2a, G-test, p<0.05; Table 3.3). Coccinella transversoguttata richardsoni declined 35.8% in relative abundance and 13.7% in geographic distribution, and experienced its main decline in observed number of records in the 1980s (Figures 3.1 and 3.2; Table 3.2a, G-test, p<0.05; Table 3.3). Those native species that were suspected of declining also showed declines here. Adalia bipunctata steadily and significantly declined in observed number

1 0 1 Adalla bipunctata Coleomegilla m. lengi Coccinella novemnotata

Relative abundance Relativeabundance Relative abundance r o o o i-» o k» ui o b o o o o p o o p o o o NRr. \ ° i s

Relativeabundance Relativeabundance oooo

Coccinella septempunctata Coccinella u. undecimpunctata Harmonia axyridis 0.41 0.4-1 0.4-i

poo poo »-» io w Relative abundance Relative Relativeabundance Relative abundance Relative o M M oooo iu o oooo I I r o' 5 I Relativeabundance Relativeabundance

Figure 3.1a Relative abundance by time period for lady beetle species in eastern Canada Adalia bipunctata Coleomegilla m. lengi Coccinella novemnotata

i*> b » n Relative abundance Relative Relative abundance Relative abundance Relative 0 0 0 0 0 o o o o o o i-» Relative abundance Relative Relative abundance Relative

Coccinella septempunctata Coccinella u. undecimpunctata Harmonia axyridis

___ 1 ___ 1 N» ___ 1 Relative abundance Relative abundance Relative Relative abundance Relative

o i-» k» w * 00000 o M Nl U ^ 00 00 0 O % < 9 ° V A \ s i i X | o' § I 3 c a S's. 3 Q 2 S'

Relative abundance Relativeabundance Z 0 0 0 0 0 o M ki lu ^ 0 0 0 0 0 o i-» ki lu ia. D 3 CO o n 0 V) r-h -I O QD QJ Q. CD C ro rt> 0) fP "O CTQ (excluding NL & LB). o u> Adalia bipunctata Coleomegilla m. lengi Coccinella novemnotata 2.0 - 2.0-1 2 .0-1

~~5 1.5 S 1.5- “ 1.5-~~

~~i 1.0 E 1.0- § l.o-~~

~~5 0.5 •= 0.5- 0.5-~~

~~A oS? tSf* aCP qSf5 rSf rSf»~~

~~Coccinella t. richardsoni Hippodamia t. tibialis 2 .0*1 2.0 -~~

~~E 1.0- E 1.0~~

~~Coccinella septempunctata Coccinella u. undecimpunctata Harmonia axyridis 2.0 - 2.0 - 2.0 -~~

~~3 1.5 3 1.5-1~~

~~J 1.0- E 1.0 tiJn | 0.5 H~~

~~! . i l l 1------1------1------r- J ✓ / y y w ///< // ////// Propylaea Hippodamia variegata quatuordecimpunctata 2.0 n 2.0-~~

~~S 1.5- 3 1.5H X IS § 1.0- | 1.0~~

~~Figure 3.2a Geographic range (km2) by tim e period for lady beetles in eastern Canada.~~

~~104 Adalia bipunctata Coieomegilla m. lengi Coccinella novemnotata 1.5 1.5-,~~

~~rlo Ho * 1.0 X l . o - x 1.0~~

~~2 0.5 3 0.5 3 0.5~~

~~ofo* <$#» ^ 0?*’^ ’ / # , < / / ^ V V V V V~~

~~Coccinella t. richardsoni Hippodamia t. tibialis 15-1~~

~~x 1.0 x 1.0-~~

~~2 0.5-~~

~~I ^ r T r T ^ / / / / / / ////// i> V V V V~~

Coccinella septempunctata Coccinella u. undecimpunctata Harmonia axyridis 1.5 l.S-i 1.5-1 to O o rlo x 1.0 X 1 .0 IN™"'ZL 10 J 1 2 0.5 s 0.5 s < llll I i I I a ////// ✓/WW //.w / Propyiaea Hippodamia variegata quatuordecimpunctata 1.5 1.5- tO 0 rto 1 10 Ji 1Q1 pS""* N je£ Jt£ *3 0.5 s 0.5 <

~~~i 1 r- 0-1—i----- 1 r ////// pf>~~

~~Figure 3.2b Geographic range (km2) lady beetles in eastern Canada (excluding NL & LB).~~

~~105 2009 2000- 14.0 109.8 71.0~~

~~16.4 Hippodamia Hippodamia~~

Hv: Hv: Htt: 110.3 58.5 50.0 156.6 83.0 30.9 245.9 130.3 111.5 76.1 40.3 34.5 243.3 128.9 110.3 Expected6 163.4 161.7 104.1 73.4 Harmonia axyridis; Ha: Ha: 1960- 1970- 1980- 1990- 65.8 161.0 242.2 128.4 66.1 28.8 8.4 20.5 102.8 30.0 168.2 49.0 119.9 180.4 95.6 81.8 145.9 42.5 24.9 7.3 17.8 26.7 14.2 12.1 229.1 66.8 225.8 Coccinella transversoguttata richardsoni; Coccinella *** *** *** Ctr: Ctr: *** *** ** *** 2009 1969 1979 1989 1999 151 70.9 20.7 50.6 140 173 57 2 17 37*** 226.7 45*** 4 106 *** *** *** ««* *♦* *** 140 105 56 1* 209*” 732 626 7*** 35 91 0 0 63.9 18.6 45.6 68.5 36.3 31.1 88 Coccinella undecimpunctata undecimpunctata; Coccinella Coccinella novemnotata; Coccinella *«* *** *** *** *** 16*** 130 ; Cn: Cn: ; 517*** 184 223 2 5 Cuu: Cuu: Observed *** • * • *** *** *** *** 1970- 1980- 1990- 2000- <1960 193 153*** 115 167 0 6*** 264 ***264 *** 145 38 45 48 *** *** *** 1969 1979 1989 1999 1960- 24 136*’ * 374 918 1381 2** 48*** 30 0 90 30 Coccinella septempunctata; Coccinella Coleomegilla maculataColeomegilla lengi Cs: Cs: *** *** *** *** *** *** Cml: Cml: <1960 15 191 235 6 2 ’ ** 368 100 0 0 0 0 0 0 377 Propylea quatourdecimpunctataPropylea 2009) (total # ofall species in all tim e periods) on the 0.05). < 0.01; 0.001;p G-test.a < < p p * ** (*** Bolded observed values indicate those which are lowerexpected than on based the G-test. Pq: Pq: etal. Adalia bipunctata; Speciesc (Myers tredecimpunctata tibialis; Hv variegata; Ha Pq Cuu Htt 3 betweenDifferences tested 3 with and Observed Expected = 2*[(£G-test: G - observed) (£expected)] In observed In observed (Zar 1999) were each interval, values as: calculated For Expected b (total # of species A in all tim e periodsHtotal Ab: c # of all species for one tim e period) Cn Cml Total 1287 Non-natives Cs 0 Ctr Natives Ab Table 3.2a Comparison ofobserved versus expected number of recordsfor of species 10 lady beetlestime six across periods based

~~106 2000- 102.0 79.8 14.0 50.2 34.8 71.5 9.5~~

~~1999 2009 1990- 124.4 106.4 16.4 11.1 119.2 36.7 31.4 93.2 58.7 Hippodamia Hippodamia~~

Hv: Hv: Htt: 157.7 83.6 245.421.0 130.1 111.3 Expectedb 152.6114.3 234.6 175.8 102.6 159.6 20.1 31.0 49.8 76.6 40.6 45.1 69.3 Harmonia axyridis; Ha: 60.6 28.6 72.0 110.7 63.4 58.1 146.3 224.9 8.0 19.3 5.4 13.7 215.4 225.3 ______Coccinella transversoguttata richardsoni; *** *** Ctr: Ctr: *** * 2009 1969 1979 1989 2000- <1960 1960- 1970- 1980- 139*** 101.7 170 150 70.4 19.8 15*** 161.5 45.4 1 57 28.4 31*** 206.5 0 63.6 17.9 45*** 144.8 40.7 (* (* 0.001;< p 0.01; < p 0.05). < p Bolded observed values ).3 107 *** *** *** *** *** 1999 103 140 203 35 7*** 3*** 55 88** 0 Coccinella Coccinella undecimpunctata undecimpunctata; Coccinella novemnotata; *** *** *** Cn: Cn: 144*** 168*** 83*** 514 5 0 6 Cuu: Cuu: Observed *** «** #** *** *** 189 147*** 115 223*** 255 38 8*** 38*** 876 1347 714 611 48 48*** 130**’ *** *** *** *** ** 1969 1979 1989 124 348 22 30*** 60 2 80 145 0 Coccinella septempunctata; Coleomegilla maculata lengi; *«* Cs: Cs: *** *** *** **« *** Cml: Cml: 1237 11 191 224 100 30*** 361 00 0 0 0 350 Propylea quatourdecimpunctata indicate those which are lower than expected on thebased G-test. on the G-test (excluding Newfoundland and Labrador 2009) (tota| # 0f a|| Specjes jn a|| tjme periods) Pq: Pq: et al. Adalia bipunctata; Species0 <1960 1960- 1970- 1980- 1990- (Myers variegata; Hv Pq Ha 0 0 0 2*** Htt tredecimpunctata tibialis; Cuu b For each interval, Expected values were calculated as: were values interval, each as: calculated For Expected b (total # of species A in all tim e periodsHtotal # of all species for one tim e period! Cs Cn Ctr a betweenDifferences tested a withand Expected 2*[(][Observed = G-test; G - (£observed) expected)] In In observed observed (Zar 1999) Total c Ab: Ab: c Non-natives Cml Natives Ab Table 3.2b Comparison of observed versus expected number of records for of species 10 lady beetlestime six across periods based

107 Table 3.3 Absolute number of records (and percent) for each time period for ten lady beetle species in eastern Canada. Records are shown for all of eastern Canada (Ontario - Newfoundland) and with Newfoundland and Labrador excluded in case of bias due to lower collecting effort there. a) All records Species3 <1960 1960s 1970s 1980s 1990s 2000s Total Natives Twospot 368(28.6) 90(24.1) 167(18.2) 184(13.3) 91(12.4) 37(5.9) 937 Spotted 100(7.8) 30(8.0) 115(12.5) . 223(16.1) 88(12.0) 45(7.2) 601 Ninespot 191(14.9) 30(8.0) 38(4.1) 5(0.4) 0 0 264 Transverse 377(29.3) 136(36.4) 264(28.8) 145(10.5) 7(1.0) 4(0.6) 933 Thirteenspot 235(18.3) 62(16.6) 193(21.0) 153(11.1) 35(4.8) 17(2.7) 695 Non-natives Sevenspot 0 0 48(5.2) 517(37.4) 209(28.6) 173(27.6) 947 Elevenspot 15(1.2) 24(6.4) 45(4.9) 16(1.2) 1(0.1) 2(0.3) 103 Asian 0 0 0 2(0.1) 140(19.1) 151(24.1) 293 Variegated 0 0 0 6(0.4) 56(7.7) 57(9.1) 119 Fourteenspot 0 2(0.5) 48(5.2) 130(9.4) 105(14.3) 140(22.4) 425 Total 1287 374 918 1381 732 626 5317 b) Eastern Canada excluding Newfoundland and Labrador Species3 <1960 1960s 1970s 1980s 1990s 2000s Total Natives Twospot 350(28.3) 80(23.0) 145(16.6) 168(12.5) 83(11.6) 31(5.1) 857 Spotted 100(8.1) 30(8.6) 115(13.1) 223(16.6) 88(12.3) 45(7.4) 601 Ninespot 191(15.4) 30(8.6) 38(4.3) 5(0.4) 0 0 264 Transverse 361(29.2) 124(35.6) 255(29.1) 144(10.7) 7(1.0) 3(0.5) 894 Thirteenspot 224(18.1) 60(17.2) 189(21.6) 147(10.9) 35(4.9) 15(2.5) 670 Non-natives Sevenspot 0 0 48(5.5) 514(38.2) 203(28.4) 170(28.7) 935 Elevenspot 11(0.9) 22(6.3) 38(4.3) 8(0.6) 0 1(0.2) 80 Asian 0 0 0 2(0.1) 140(19.6) 150(24.8) 292 Variegated 0 0 0 6(0.4) 55(7.7) 57(9.3) 118 Fourteenspot 0 2(0.6) 48(5.5) 130(9,7) 103(14.4) 139(22.7) 422 Total 1237 348 876 1347 714 611 5133 “Twospot: Adalia bipunctata; Spotted: Coleomegilla maculata lengi; Ninespot: Coccinella novemnotata; Transverse: Coccinella transversoguttata richardsoni; Thirteenspot: Hippodamia tredecimpunctata tibialis; Sevenspot: Coccinella septempunctata; Elevenspot: Coccinella undecimpunctata undecimpunctata; Asian: Harmonia axyridis; Variegated: Hippodamia variegata; Fourteenspot: Propylea quatourdecimpunctata.

~~108 of records over time, as well as in relative abundance by 22.7% and in geographic distribution by 95.7% (Figures 3.1 and 3.2;Table 3.2a, G-test, p<0.05; Table 3.3).~~

~~Hippodamia tredecimpunctata tibialis declined more recently in observed number of records, as well as in relative abundance by 18.3% and in geographic distribution by~~

~~40.6% (Figures 3.1 and 3.2;Table 3.2a, G-test, p<0.05; Table 3.3). In contrast,~~

~~Coleomegilla maculata lengi experienced minor changes throughout the entire study~~

~~period with relative abundance and geographic distribution only fluctuating by 8.9% and~~

~~26.4%, respectively, throughout the study period (Figures 3.1 and 3.2;Table 3.2a, G-test,~~

~~p<0.05; Table 3.3).~~

Three general trends in abundance were shown by native species. The first trend was a steady decline in relative abundance through successive tim e periods (C. novemnotata and A. bipunctata), which began prior to the arrival of four of the non

~~native species. The second trend was for minor fluctuations in relative abundance~~

~~before 1980, followed by a noticeable and continuing decline in relative abundance~~

~~beginning in the 1980s (C. transversoguttata richardsoni and H. tredecimpunctata~~

~~tibialis). The third trend was for generally similar abundance through the entire study~~

~~period, with a small peak in the period from 1970-1990 ( Coleomegilla maculata lengi).~~

~~In contrast to native species investigated, four of five non-native species~~

~~increased in relative abundance following their establishment to the end of the study~~

~~period in 2009 (Figure 3.1a, G-test, p<0.05) while range expansion showed a period of~~

~~rapid increase in geographic extent, peaking in the 1960s for C. u. undecimpunctata, and~~

~~the 1990s for C. septempunctata, H. variegata and P. quatuordecimpunctata, followed~~

~~109 by at least one decade of minor range contractions (Figures 3.2a and b, and Appendix~~

~~Figures A-6 to A-10 and A-16 to A-20). Relative abundance and geographic distribution o f C. u. undecimpunctata (the earliest non-native lady beetle to become established in~~

~~Canada) peaked in the 1960s (when excluding NL) at 6.3% and ~319,000 km2 respectively, with significantly fewer records observed than expected from the 1980s to~~

2009 (Table 3.2a and b, G- test, p<0.05). Coccinella septempunctata peaked in relative abundance and geographic distribution in the 1980s at 37.4% and ~1,253,000 km2 respectively, despite being one of the most commonly collected lady beetle species in the 2000s (Figure 3.1a). Hippodamia variegata and Propylea quatuordecimpunctata were still increasing in relative abundance at the end of the study period (9.1% and

~~22.4%) despite peaking in geographic distribution in the previous decade (approximately~~

~~315,000 km2 and ~412,000 km2). Harmonia axyridis (the most recent non-native lady beetle to become established in Canada) was still increasing in relative abundance~~

~~(24.1%) and geographic distribution (~786,000 km2) at the end of the study period.~~

~~3.4 DISCUSSION~~

~~There has been a clear decline in abundance and geographic extent for the native lady beetles ( Coccinella novemnotata, Coccinella transversoguttata richardsoni,~~

Adalia bipunctata and Hippodamia tredecimpunctata tibialis) based on specimen record data. The decline is most profound for Coccinella novemnotata and Coccinella transversoguttata richardsoni, which have virtually disappeared from eastern Canada, though C. novemnotata persists on M ont St. Hilaire, Quebec (Skinner and Domaine

~~1 1 0 2010). These are the two species that had been documented as declining most dramatically in regional studies (e.g. Wheeler and Hoebeke 1995, Elliott eto/. 1996,~~

~~Boiteau etal. 1999, Turnock etal. 2003, Alyokhin and Sewell 2004, Harmon etal. 2007,~~

~~Skinner and Domaihe 2010, Fothergill and Tindall 2010). Adalia bipunctata and~~

Hippodamia tredecimpunctata tibialis have also been reported to be declining in some areas, though not to the same extent as C. novemnotata and C. transversoguttata richardsoni (e.g. Elliott etal. 1996, Boiteau etal. 1999, Cormier etal. 2000, Alyokhin and

Sewell 2004, Lucas et al. 2007, Fothergill and Tindall 2010). Results of this study confirm significant declines in both geographic area and relative abundance in collections over the past several decades.

~~In contrast, Coleomegilla maculata lengi remains common despite the arrival of~~

~~non-native species of lady beetles (Elliott etal. 1996, Colunga-Garcia and Gage 1998,~~

~~Day and Tatum 2006, Mignault et al. 2006, Lucas et al. 2007), indicating that the~~

declines are not affecting all lady beetles equally. Collection records for Coleomegilla maculata lengi remained fairly constant in numbers and geographic extent, confirming that the species relative abundance and geographic distribution has not changed over time. The consistent pattern of abundance and distribution for this species, based on the combined records in the 32 collections, has shown that when a lady beetle is

~~present in a region, it has continued to be collected. Therefore, the observed declines in some species likely reflect actual trends and are not artefacts of inadequate collecting.~~

~~Marriott (Chapter 2) found that by combining natural history collection data from a wide variety of collection types, temporal and spatial biases present in individual collection~~

~~1 1 1 data can be minimized, making the data a viable option for assessing relative abundance and geographic distribution patterns of lady beetles.~~

The difference in abundance and distribution patterns for the different lady beetles may relate to responses to changes in food availability. Of the five native species of lady beetles looked at in this study, Coleomegilla maculata lengi is unique in that it switches to pollen when aphid densities are low (Gordon 1985, Obrycki et al.

1998b), and is still capable of normal larval development on a non-aphid diet (Michaud and Grant 2005). Therefore, any changes that could reduce prey (mainly aphid) densities or access to prey might have a greater effect on some lady beetles than others.

~~One of the most cited hypotheses for declines in native lady beetle species has been competition or interference from non-native species (Evans 2000, Cottrell 2005,~~

~~Cottrell and Yeargan 1998, Majerus etal. 2006, Obrycki etal. 2000, Obrycki etal. 1998a,~~

Obrycki et al. 1998b; Table 1.1, page 2). Collection records confirm that during the periods of declines for four native species, three of the common non-native species showed a clear increase in range and relative abundance. Coccinella septempunctata and Coccinella u. undecimpunctata both experienced periods of rapid expansion and population growth in the decades immediately after colonization, followed by a minor drop in relative abundance for Coccinella septempunctata and a more significant decline for Coccinella u. undecimpunctata. Interestingly, Coccinella u. undecimpunctata has virtually disappeared from eastern Canada since the 1990s, with current distributions restricted to coastal regions of Atlantic Canada. This trend allows for speculation that non-native species may increase rapidly following introduction, but then decline in the following years, confirming the pattern reported in the literature (Harmon etal, 2007).

Harmon (2007) suggested that a similar recent decline in Coccinella septempunctata in the US may relate to displacement by the most recently introduced non-native species of lady beetle, Harmonia axyridis.

The hypothesis that the arrival of a new species of lady beetle can displace previously established species (native or non-native) is not a new one. It has been difficult to establish definitively whether or not native species are experiencing declines as it has to determine whether non-native species are the cause. Long term studies in a single location give important information about declining native species numbers concurrent with increases in non-native species (e.g. Turnock et al. 2003), but these studies are rare. Therefore, many studies looking for changes in native lady beetle populations have compared the relative abundance of native and non-native species to see if the natives decline over time in the proportions captured (Alyokhin and Sewell

~~2004, Day and Tatum 2006, Elliott et al. 1996, Finlayson et al. 2008, Harmon et al. 2007).~~

However, in many cases, the high abundance of the non-native species after their initial colonization may obscure patterns in the native species (e.g. an increase in the number of non-natives present increases the total lady beetle density, resulting in a decrease in the proportion of natives). This decrease can be misinterpreted as a loss of native species of lady beetles concurrent with arrival of non-natives, though in fact, the overall numbers may not change (Harmon etal. 2007). It is also not uncommon for a lady beetle community to be dominated by two to four species which can represent >90% of individuals, or for the dominant species to change over time (Hodek and Honek 1996,

113 cited from Acorn 2007), further obscuring interpretations of population increases or declines. One factor that argues against a decline in native species relating to the arrival of non-natives is that three of the native taxa in this study are present, at least at species level, in the Holarctic ( Hippodamia tredecimpunctata, Adalia bipunctata and

Coccinella transversoguttata), and coexist in Eurasia with the same non-native species which are now common in North America. For example, Coccinella transversoguttata shares a variety of habitats throughout Eurasia with Coccinella septempunctata and

~~Harmonia axyridis (Kuznetsov 1997).~~

Due to the large geographical extent of this collection-based study it is possible to assess patterns in the native and non-native species which may not be evident from localized studies. Declines (both in relative abundance and geographic extent) for two species, Coccinella transversoguttata richardsoni and Hippodamia tredecimpunctata tibialis, did coincide with the establishment (1970s) and rapid spread of the non-native

Coccinella septempunctata. Both native species were widely distributed and frequently collected across eastern Canada before 1980, but much less frequently between 1980 and 2009, despite extensive collecting effort. Instead, Coccinella septempunctata was collected from the same locations and habitats where the two native species were previously found. Since there was continued collecting of other native species (e.g.

Coleomegilla maculata lengi) as well as high numbers of this non-native species, the pattern is not likely due to preferential collecting of novel species, and suggests instead that C. septempunctata had displaced the native species. In contrast, the observed decline in relative abundance and geographic extent of Coccinella novemnotata and

114 Adalia bipunctata does not relate temporally to the arrival of non-native species. These declines began prior to the establishment of the now common species of non-native lady beetles. This is particularly true fo r Coccinella septempunctata which had been implicated in the decline of both of these native species (see Table 1.1 pp. 2 for examples and references).

~~Several mechanisms have been advanced fo r the decline in native lady beetles.~~

Where arrival of non-native species has been implicated, the non-native lady beetles may negatively affect the native populations through competition, intraguild predation and the introduction of parasites and pathogens. For example, Harmonia axyridis is a more efficient predator than other species of lady beetles (native and non-native) (Lucas et al. 2002); and also feeds directly on larvae of other lady beetles (Yasuda e t al. 2004,

Michaud 2002). Harmonia axyridis is also highly resistant to a fungal parasite which commonly infects some species of native lady beetles, so this may give this non-native species a competitive advantage over natives (Cottrell and Shapiro-llan 2003). However, c it is more likely multiple stressors are affecting the native lady beetles, and that observed declines in some species are not as simple as interactions with non-native species. For example, habitat change has occurred coincidentally with the decline of native species and the arrival of non-natives (Harmon et al. 2007). McCorquodale et al.

(2011) explores the possibility that European settlement of eastern North America in the 1800s facilitated an increase in lady beetle populations through the creation of preferred habitats (e.g. open areas such as meadows, forest edges, and farmland). As farmland has subsequently been abandoned throughout large areas of eastern North

~~115 America and land has reverted to forest (e.g. Davis and Browne 1996, Loo and Ives~~

~~2003), lady beetle populations would also be expected to decline, a trend which could be misattributed to the arrival of non-native species of lady beetles.~~

This study provides strong evidence for the decline in several species of lady beetles in eastern Canada. Analysis of collection records does not support the hypothesis that declines are related to the arrival of non-native species, at least for

Coccinella novemnotata and Adalia bipunctata, though further study on this factor is warranted for Coccinella transversoguttata richardsoni and Hippodamia tredecimpunctata tibialis. Factors which may contribute to the decline of native lady beetles include: habitat alteration, parasites, pathogens, parasitoids, interspecific predation and competition from non-native lady beetles.

~~116 3.5 LITERATURE CITED~~

~~Acorn, J. 2007. Ladybugs of Alberta, finding and connecting the dots. The University of Alberta Press, Edmonton, Alberta. 169 pp.~~

~~Alyokhin, A. and Sewell, G. 2004. Changes in a lady beetle community following the establishment of three alien species. Biological Invasions. 6:463-471.~~

Belicek, J. 1976. Coccinellidae o f western Canada and Alaska with analyses o f the transmontane zoogeographic relationships between the fauna of British Columbia and Alberta (Insecta: Coleoptera: Coccinellidae). Quaestiones Entomologicae. 12: 283-409.

~~Boiteau, G. 1983. The arthropod community of potato fields in New Brunswick. The Canadian Entomologist 115: 847-853.~~

~~Boiteau, G., Y. Bousquet and Osborn, W.P.L. 1999. Vertical and temporal distribution of Coccinellidae (Coleoptera) in flight over an agricultural landscape. The Canadian Entomologist 131: 269-277.~~

~~Brown, W.J. 1940. Notes on the American distribution of some species of Coleoptera common to the European and North American continents. The Canadian Entomologist. 72: 65-78.~~

Canadian Permanent Committee on Geographical Names. 1992. Guide to the field collection of Native geographical names (provisional edition). Retrieved January 26, 2012, from http://www.nrcan.gc.ca/sites/www.nrcan.gc.ca.earth- sciences/files/pdf/native field guide e.pdf.

~~Chantel, C. 1972. Additions a la faune Coleopterique du Quebec. Le Naturaliste Canadien. 99: 243-244.~~

~~Chapman, A.D. 2005. Uses of primary species-occurrence data, version 1.0. Report for the Global Biodiversity Information Facility, Copenhagen. 106 pp.~~

~~Coderre, D., E. Lucas and Gagne, 1.1995. The occurrence of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in Canada. The Canadian Entomologist. 127: 609-611.~~

~~117 Cotterill, F.P. 1995. Systematics, biological knowledge and environmental conservation. Biodiversity and Conservation. 4:183-205.~~

~~Cottrell, T.E. 2005. Predation and cannibalism of lady beetle eggs by adult lady beetles. Biological Control. 34: 159-164.~~

~~Cottrell, T.E. and Shapiro-llan, D.l. 2003. Susceptibility of a native and an exotic lady beetle (Coleoptera: Coccinellidae) to Beauveria bassiana. Journal of Invertebrate Pathology. 84:137-144.~~

Cottrell, T.E. and Yeargan, K.V. 1998. Intraguild predation between an introduced lady beetle, Harmonia axyridis (Coleoptera: Coccinellidae), and a native lady beetle, Coleomegilla maculata (Coleoptera: Coccinellidae). Journal of the Kansas Entomological society. 71:159-163.

~~Danks, H.V. 1991. Museum collections: fundamental values and modern problems. Collection Forum. 7: 95-111.~~

~~Davis, D.S. and Browne, S. 1996. The natural history of Nova Scotia. Volume 1 topics and habitats. Nova Scotia Museum of Natural History and Nimbus Publishing, Halifax, NS. 518 pp.~~

~~Elliott, N.C., Kieckhefer, R.W. and Kauffman, W.C. 1996. Effects of an invading coccinellid on native coccinellids in an agricultural landscape. Oecologia 105: 537-544.~~

Environmental Systems Research Institute, Inc. 2007. Lambert conformal conic-ArcGIS 9.2 desktop help. Retrieved September 5, 2011, from http://webhelp.esri.eom/arcgisdesktop/9.2/index.cfm ?TopicName=Lambert_Co nformal_Conic.

~~Evans, E.W. 1991. Intra versus interspecific interactions of lady beetles (Coleoptera: Coccinellidae) attacking aphids. Oecologica. 87:401-408.~~

~~118 Evans, E.W. 2000. Habitat displacement of North American ladybirds by an introduced species. Ecology. 85: 637-647.~~

~~Ferrier, S. 2002. Mapping spatial pattern in biodiversity for regional conservation planning: where to from here? Systematic Biology. 51: 331-363.~~

Finlayson, C.J., K.M. Landry and Alyokhin, AA/. 2008. Abundance of native and non native lady beetles (Coleoptera: Coccinellidae) in different habitats in Maine. Annals of the Entomological Society of America. 101:1078-1087.

~~Funk, V. and Richardson. K. 2002. Systematic data in biodiversity studies: use it or lose it. Systematic Biology. 51: 303-316.~~

~~Gordon, R.D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. Journal of New York Entomological Society. 95:1-912.~~

~~Harmon, J.P., Stephens, E. and Losey, J. 2007. The decline of native coccinellids (Coleoptera: Coccinellidae) in the United States and Canada. Journal o f Insect Conservation 11: 85-94.~~

Hesler, L.S., Kieckhefer. R.W. and Catangui, M.A. 2004. Surveys and field observations of Harmonia axyridis and other Coccinellidae (Coleoptera) in eastern and central South Dakota. Transactions of the American Entomological Society. 130:113- 133.

~~Hesler, L.S. and Kieckhefer, R.W. 2008. Status of exotic and previously common native coccinellids (Coleoptera) in South Dakota landscapes. Journal of the Kansas Entomological Society. 81: 29-49.~~

Hodek, I. and Honek, A. (with contributions by P. Ceryngier and I. Kovar). 1996. Ecology of Coccinellidae. Series Entomologica, Vol. 54. Kluwer Academic Publishers, Dordrecht, the Netherlands. 464 pp. (cited from Acorn, J. 2007. Ladybugs of Alberta, finding and connecting the dots. The University of Alberta Press, Edmonton, Alberta. 169 pp.)

119 Hoebeke, E.R. and Wheeler, A.G. 1996. Adventive lady beetles (Coleoptera: Coccinellidae) in Canadian Maritime Provinces, with new eastern US records on Harmonia quadripunctata. Entomological News. 107: 281-290.

~~Johnson, N.F. 2007. Biodiversity informatics. Annual Review of Entomology. 52:421-438.~~

Koch, J.B. and Strange, J.P. 2009. Constructing a species database and historic range maps for North American bumblebees ( Bombus sensu stricto Latreille) to inform conservation decisions. Bee Science. 9: 97-108.

~~Kuznetsov, V.N. 1997. Lady beetles of the Russian Far East. Center for Systematic Entomology, Gainesville. 248 pp.~~

~~Larochelle, A. 1979. Les Coleopteres Coccinellidae du Quebec. Cordulia (Supplement). 10: 1- 111.~~

~~Loo, J and Ives, N. 2003. The Acadian forest: historical condition and human impacts. The Forestry Chronicle. 79: 462-474.~~

Lucas, E., Vincent, C., Labrie, G., Chouinard, G., Fournier, F., Pelletier, F., Bostanian, N.J., Coderre, D., Mignault, M.-P. and Lafountaine, P. 2007. The multicolored Asian ladybeetle Harmonia axyridis (Coleoptera : Coccinellidae) in Quebec agroecosystems ten years after its arrival. European Journal of Entomology 104: 737-743.

~~Majka, C. G. and McCorquodale, D.B. 2006. The Coccinellidae (Coleoptera) of the Maritime Provinces of Canada: new records, biogeographical notes, and conservation concerns. Zootaxa. 1154: 49-68.~~

~~Maplnfo Corporation. 2007. MapBasic Version 9 Reference Guide. Retrieved July 13, 2011, from http://reference.mapinfo.com/software/mapbasic/english/9_0/MB_Ref.pdf.~~

120 Marriott, S.M., Giberson, DJ. and McCorquodale, D.B. 2009. Changes in the status and geographic ranges of Canadian lady beetles (Coccinellidae) and the selection of candidates for risk assessment. Part I Foundation Report. Report to COSEWIC Arthropods Species Specialist Committee. 53 pp.

~~Marshall, S. 2008. Lady beetles of Ontario. Retrieved December, 2008, from http://www.uoguelph.ca/debu/lady/lady-beetles.htm.~~

~~McCarthy, M.A. 1998. Identifying declining and threatened species with museum data. Biological Conservation. 83: 9-17.~~

McCorquodale, D.B., Giberson, DJ. and Marriott, S.M. 2011. Changes in the status and geographic range of Canadian lady beetles (Coleoptera: Coccinellidae: Coccinellinae) and the selection of candidate species for risk assessment by the Committee on the Status of Endangered Wildlife in Canada - Part 3. Report to COSEWIC Arthropods Species Specialist Committee. 60 pp.

Michaud, J.P. and Grant, A.K. 2005. Suitability of pollen sources for the development and reproduction of Coleomegilla maculata (Coleoptera: Coccinellidae) under simulated drought conditions. Biological Control. 32: 363-370.

~~Mignault, P.-M., Roy, M. and Brodeur, J. 2006. Soybean aphid predators in Quebec and the suitability of Aphis glycines as prey for three Coccinellidae. BioControl 51: 89- 109.~~

~~Newbold, T. 2010. Applicationss and limitations of museum data for conservation and ecology, with particular attention to species distribution models. Progress in Physical Geography. 34: 3-22.~~

~~Obrycki, J.J., Giles, K.L. and Ormond, A.M. 1998a. Interactions between an introduced and indigenous coccinellid species at different prey densities. Oecologia. 117: 279-285.~~

121 Obrycki, J.J., Elliott, N.C. and Giles, K.L. 2000. Coccinellid introductions: potential for and evaluation of non-target effects, Nontarget effects o f biological control (P.A. Follet, and J.J. Duan, Eds.), Dordrecht, Netherlands; Kluwer: 127-145.

~~Riddick, E.W., Cottrell, T.E. and Kidd, K.A. 2009. Natural enemies of the Coccinellidae: parasites, pathogens, and parasitoids. Biological Control. 51: 306-3.~~

~~Rushton, S.P., Ormerod, SJ. and Kerby, G. 2004. New paradigms for modeling species distributions? Journal of Applied Ecology. 41:193-200.~~

~~Shaffer, H.B., Fisher, R.N. and C. Davidson. 1998. The role of natural history collections in documenting species declines. Trends in Ecology and Evolution. 13: 27-30.~~

~~Yasuda, H., Evans, E.W., Kajita, Y., Urakawa, K, and Takizawa, T. 2004. Asymmetrical larval interactions between introduced and indigenous ladybirds in North America. Oecologia. 141:722-731.~~

~~Zar, J.H. 1999. Biostatistical Analysis. 4th Edition. Prentice Hall, New Jersey.~~

~~122 4.0 CONCLUSIONS~~

Natural history collection data from four different types of natural history collections in Atlantic Canada have biases which could interfere with interpretation of lady beetle distributions. However, these biases can be minimized by combining data from all collections, making natural history collection data a viable option for assessing changes in lady beetle distributions and relative abundance when used with caution.

Historical and current specimen data pooled from 32 natural history collections across eastern Canada provides evidence of changes in lady beetle distributions and relative abundance. Collection data shows there is a clear decline in abundance and geographic extent in all of the native lady beetles studied, except one which continues to persist at relatively constant levels. At the same time, non-native species show a clear increase in relative abundance from the time of establishment to the end of the study period in 2009, despite experiencing minor range contractions in more recent decades for three of the species.

The broad geographic scope of this study makes it possible to assess patterns in the native and non-native species which may not be evident from localized studies. This study provides strong evidence that the decline of Coccinella transversoguttata richardsoni and Hippodamia tredecimpunctata tibialis does coincide with the increase in numbers and distribution of non-native species, while the decline of Coccinella novemnotata and Adalia bipunctata began prior to the establishment of the now common species of non-native lady beetles. Therefore, these data suggest that the

~~123 decline of some species of native lady beetles is actually happening, but is not solely due to the arrival of non-native species.~~

~~124 APPENDIX~~

~~Maps of eastern Canada showing modified polygons giving range of lady beetle species, based on collection records for the tim e periods are shown in this section.~~

~~Records are mapped showing all localities in eastern Canada with lady beetle records~~

(Figures A -l to A-10) and excluding records from Newfoundland and Labrador (in case of bias due to low collecting effort in those areas; Figures A - ll to A-20). For both sets of maps, distributions of native lady beetle species are given in red, and those for non natives are given in blue. In cases where a polygon could not be created (<3 records), records are shown as a coloured dot.

~~125 Adalia bipunctata~~

~~Before 1960 1960-1969~~

~~A=CT~~

~~1970-1979 1980-1989~~

~~' ' l —^ 1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-l Distribution of Adalia bipunctata (twospotted lady beetle) in eastern Canada from 1891 to 1959, and by decade from 1960 to 2009.~~

~~126 Coleomegilla maculata lengi~~

~~Before 1960 1960-1969~~

~~1980-19891970-1979~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-2 Distribution of Coleomegilla maculata lengi (spotted lady beetle) in eastern Canada from 1890 to 1959, and by decade from 1960 to 2009.~~

~~127 Coccinella novemnotata~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-3 Distribution of Coccinella novemnotata (ninespotted lady beetle) in eastern Canada from 1881 to 1959, and by decade from 1960 to 2009.~~

~~128 ~ \~~

~~Coccinella transversoguttata richardsoni~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-4 Distribution of Coccinella transversoguttata richardsoni (transverse lady beetle) in eastern Canada from 1889 to 1959, and by decade from 1960 to 2009.~~

~~129 Hippodamia tredecimpunctata tibialis~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-5 Distribution of Hippodamia tredecimpunctata tibialis (thirteenspotted lady beetle) in eastern Canada from 1885 to 1959, and by decade from 1960 to 2009.~~

~~130 Coccinella septempunctata~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hal! Polygon~~

~~Figure A-6 Distribution of Coccinella septempunctata (sevenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009.~~

~~131 Coccinella undecimpunctata undecimpunctata~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-7 Distribution of Coccinella undecimpunctata undecimpunctata (elevenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009.~~

~~132 Harmonia axyridis~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-8 Distribution of Harmonia axyridis (multicoloured Asian lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009.~~

~~133 Hippodamia variegata~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-9 Distribution of Hippodamia variegata (variegated lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009.~~

~~134 Propylea quatuordecimpunctata~~

~~Before 1960 1960-1969~~

~~tp>\~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-10 Distribution of Propylea quatuordecimpunctata (fourteenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009.~~

~~135 Adalia bipunctata~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~2000-20091990-1999~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A -ll Distribution of Adalia bipunctata (twospotted lady beetle) in eastern Canada from 1891 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador).~~

~~136 Coleomegilla maculata lengi~~

~~1990-1999 2000-2009~~

~~n -.---~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

Figure A-12 Distribution of Coleomegilla maculata lengi (spotted lady beetle) in eastern Canada from 1890 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador). Coccinella novemnotata

~~1970-1979 1980-1989~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-13 Distribution of Coccinella novemnotata (ninespotted lady beetle) in eastern Canada from 1881 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador).~~

~~138 Coccinella transversoguttata richardsoni~~

~~Before 1960 1960-1969~~

~~1980-19891970-1979~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

Figure A-14 Distribution of Coccinella transversoguttata richardsoni (transverse lady beetle) in eastern Canada from 1889 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador).

~~139 Hippodamia tredecimpunctata tibialis~~

~~Before 1960~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

Figure A-15 Distribution of Hippodamia tredecimpunctata tibialis (thirteenspotted lady beetle) in eastern Canada from 1885 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador).

~~140 Coccinella septempunctata~~

~~Before 1960 1960-1969~~

~~1980-19891970-1979~~

~~2000-20091990-1999~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-16 Distribution of Coccinella septempunctata (sevenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador).~~

~~141 Coccinella undecimpunctata undecimpunctata~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~142 Harmonia axyridis~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Canada Modified Convex Hall PolygonEastern~~

~~Figure A-18 Distribution of Harmonia axyridis (multicoloured Asian lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador).~~

~~143 Hippodamia variegata~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-19 Distribution of Hippodamia variegata (variegated lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador).~~

~~144 Propylea quatuordecimpunctata~~

~~Before 1960 1960-1969~~

~~1970-1979 1980-1989~~

~~1990-1999 2000-2009~~

~~United States Eastern Canada Modified Convex Hall Polygon~~

~~Figure A-20 Distribution of Propylea quatuordecimpunctata (fourteenspotted lady beetle) in eastern Canada from 1900 to 1959, and by decade from 1960 to 2009 (excluding Newfoundland and Labrador).~~

~~145~~