Effects of environmental changes on local butterfly populations

in Ottawa, Ontario

Alyssa Caravaggio

Jeremy T. Kerr

Monday, April 29, 2013

1
Abstract

Diversity and abundance of butterfly populations are influenced by local land use and weather

patterns. Using landscape and climate data for the Ottawa area, I examined butterfly surveys

taken at the same sites in 2004 and 2012 to see what effects these factors had on the local

butterfly population over time. The defined patterns of species distribution and diversity in 2004

were not replicated in 2012. Instead, species were more widespread and evenly distributed,

regardless of land use. Generalist species, having not been affected by warmer and drier weather,

were most abundant and evenly distributed in 2012. Due to this I propose that agricultural land

use impacts came secondary to effects of abnormal weather patterns in 2012. This is an example

of climate change induced weather fluctuations affecting local butterfly populations, which can

cause difficulties in focusing future conservation efforts.

2
Introduction

Butterflies are important species in the natural environment, not only as pollinators but as

bioindicators of the state of an ecosystem. Butterflies have a holometabolous life cycle, going

through complete metamorphosis between juvenile and adult stages. This creates a divide

between the resource needs of juvenile (caterpillars) and adult butterflies, each group filling

separate ecological niches. Many species of butterflies are also migratory, which leads to a large

dispersal of these species. Both factors make these insects very attuned to the quality of host

plants, habitats, and the local climate. Drivers of species richness have been shown to vary based

on geographic location and the ecology of the given butterfly species (Stefanescu et al., 2011).

Temperature and precipitation are both environmental factors that affect the habitats of

lepidopteran species. For example, cold weather can affect insect development, food

requirements, and can lead to population extinctions in periods of severe or unseasonably low

temperatures (Gilbert and Singer, 1975). High temperatures causing a drought may increase

butterfly abundances but also is damaging to plant life (Woods et al. 2008). With respect to

precipitation levels, heavy rain can have a negative impact on larvae, but may also have positive

impact on plant growth in the following year (Woods et al. 2008). Stefanescu et al. (2011) found

a positive correlation between increasing yearly rainfall and species richness. It is clear that there

is a delicate balance needed between temperature and precipitation to support stable butterfly

populations within a specific area.

Each butterfly species requires different environments and food sources for its larval,

pupal, and adult stages. Species can generally be categorized as habitat generalists or specialists,

depending on their specifications for survival. Habitat generalists do not require a specific

environment or host plant to thrive, which may allow them to survive in environments with

3
higher anthropogenic inputs. They have been shown to benefit from varying resources, are able

to move among suitable habitat patches, and deal with environmental changes more favorably

than specialist species (Dapporto and Dennis, 2012). Habitat specialists are often found in

natural environments, with little or no human impacts. This is because habitat specialists are

dependent on the diversity of resources and environments available (Menndez et al., 2007).

There is a large range of interactions of butterflies with their environment. There have

been many studies done throughout the world that have found that butterfly species are affected

by landscape changes (Benton et al., 2003, Flick et al., 2012, Menndez et al., 2007, Oliver et al.,

2010, Weibull et al., 2000). Often this is due to the implementation of agriculture or construction

of urban areas in what were once natural landscapes (Flick et al., 2012). Habitat heterogeneity is

cited as a factor that decreases butterfly abundance and diversity in agricultural areas, due to the

monotonous groups of crops that are planted. Benton et al. (2003) explained that increasing

habitat heterogeneity in agricultural areas leads to increases in biodiversity. Urban areas tend to

pose a problem for butterfly habitats because they reduce the amount of natural area and food

plants available.

A warming climate is hypothesized to have negative impacts on both generalist and

specialist groups (Stefanescu et al., 2011). The impacts of environmental changes including

fluctuations in the climate are different for each species in the local population and will differ in

relation to the species needs for survival (Menndez et al., 2007). It is this delicate system of

interactions between butterfly species and their environment that makes it so difficult to predict

the future trends of a population.

Populations of butterflies including both native and migrant species are observed in the

Ottawa area beginning every spring. With the expansion of agriculture and urbanization, changes

4
in the local butterfly population are to be expected. By analyzing data collected at sites in the

Ottawa area in 2004 and comparing it with data taken from the same sites in 2012, I observed

changes in the composition of the local butterfly population. The original data collected in 2004

looked for links between agricultural intensity and butterfly abundance and species richness.

This data showed localized patterns in species distribution for habitat generalists and specialists

and reduced butterfly species richness due to agricultural intensification (Feswick, 2005). My

research examines the changes that took place in the butterfly population between 2004 and 2012

in hopes of understanding the causes of these shifts. I looked at both the landscape and climate

factors affecting the local environment in relation to the butterfly species at the survey sites.

Levels of greenness at each site were used as a measure of land use and plant productivity.

Temperatures and precipitation levels were used as climatic variables representative of the

weather for each year. There were two main questions that I hoped to answer through my

research:

1) How do butterfly species richness and abundance patterns change through time

due to landscape change?

2) Did weather patterns have an effect on the butterfly population?

I expected that decreases in greenness due to intensifying agriculture at survey sites would

decrease the abundance and diversity of butterflies observed in 2012. I predicted that a greater

concentration of generalist butterfly species would be recorded in sites in 2012 due to the

landscape changes. I also hypothesized that changes in local weather patterns due to climate

change would affect the butterfly population. I made these hypotheses based on the trends that

had been previously observed in the data from 2004 and the conclusions of the several other

studies cited previously. This study provides the chance to understand the dynamics of the local

5
butterfly population as a whole, relative to changes in the local environment over eight years. It

may also have the potential to help in identifying trends in butterfly populations elsewhere and to

predict the impacts of future environmental change on local butterflies.

Materials and Methods

The data analyzed was collected in 2004 and 2012 from May to August at the same sites

in the Ottawa area (Figure 1). The total of 22 sites were carefully selected previous to data

collection in 2004. Factors taken into account when choosing sites were those that could affect

species richness and abundance, including site area, shape index, and plant species richness

(Feswick, 2005). The selected sites were divided into groups of control or treatment. Control

landscapes were not within 100 m of any active agriculture and were comprised of natural

meadows or old fields. The two treatment landscapes were varying in agricultural intensity based

on the type of crop grown. Forage crop agricultural areas were defined as medium intensity

impact, whereas cash crop sites were labeled as high intensity impact.

When analyzing butterfly counts, data from one medium intensity site was excluded from

both sets due to construction occurring during the collection period in 2012. Total abundance of

each species was calculated and those species with abundance counts of only one individual

siting were removed from further analysis. This was done in order to disregard species that had a

very low probability of being seen in collections the opposite year. The sampling effort was

shown to be equal for both years through a resampling test calculated with the statistics program

Estimate S. A Coleman curve was used as a resampling method rather than a rarefaction formula

to prove the evenness in the two sampling efforts (Figure 2). Removing the singletons, as noted

before, also improved the shape of the curve. With the data compiled, abundance and species

6
richness for each site was totaled. Species were determined to be habitat generalists or specialists

based on information in Layberry, Hall, and Lafontaines The Butterflies of Canada.

When extracting landscape variables, GIS system ArcMap 10 was used to plot the

latitude and longitude coordinates of each site. The points were layered over satellite landscape

images in order to extract values of greenness for each site. Buffer areas of 100, 200, and 500 km

around each site were calculated. The landscape variables extracted for each point were NDVI

(Normalized Difference Vegetation Index) and DTCG (Difference in Tasseled Cap Greenness

annually). Averages of NDVI and TCG were also calculated for each buffer area around each of

the sites.

Climate data for 2004 and 2012 came from the National Climate Data and Information

Archive records of Environment Canada. Mean monthly temperatures were calculated as an

average of the mean daily temperatures.

Results

Butterfly Population Diversity and Abundance

When listing the species that were recorded from the surveys in both years, it was noted

that there were several species present in 2004 that were not found in 2012 (Table 1). Several of

the species missing in 2012 are habitat specialists. Likewise, there are many new species

recorded in the observations for 2012, with many of these being extreme habitat generalists.

When comparing species richness for the sites between the two years, the trends for 2004 are not

mirrored in the data from 2012. In 2004, the greatest species richness was found among the

control sites. In 2012, there were large and nearly equal number of species found in all sites.

Jaccards index of similarity was used to compare the species composition of butterflies at

7
control, medium, and high sites between 2004 and 2012 (Figure 3). Control, medium, and high

sites all had a level of under 50% similarity between the two years. Comparing the evenness

index of distribution of species throughout the sites (Figure 4), there was a decreasing trend

among sites going from control to high intensity in 2004. In 2012, this trend is not seen, but

rather the evenness of species among sites is higher and the values are nearly equal.

Environmental changes: Landscape

When reviewing landscape data such as NDVI and TCG, there was very little change

when the difference between 2004 and 2012 was measured. Many sites measured 0 for the

difference in NDVI and TCG. A Pearson product-moment correlation between change in NDVI

for sites over time and 2012 species richness was calculated using R. Given r-value was -0.296

with p-value 0.9125. When looking at the average greenness in buffer areas surrounding the

sites, no significant changes are present that could have accounted for the differences in the

butterfly population.

Environmental changes: Climate

There were clear changes in the climate, including temperature and precipitation levels,

between 2004 and 2012. On average, the mean monthly temperature for 2012 was 1.87C

warmer than 2004 (Figure 5). From May to August when the surveys were taking place, the

mean monthly temperature was 2.275C warmer in 2012 than in 2004. The maximum

temperatures recorded in each month were also higher in 2012 (Figure 6). March 2012 showed

the most significant increase with a maximum temperature reaching 27.4C. This is more than

10C warmer than the highest temperature in March 2004 and is also higher than any maximum

8
temperature for the month recorded in any year since the last survey. When looking at the survey

months, the maximum temperatures are all higher than those months in 2004. Total amounts of

precipitation (in mm) per month in 2012 were generally lower than those in 2004 (Figure 7).

May to August 2012 all had lower total precipitation than those months in 2004. July 2012

specifically has 49.8 mm less precipitation than July 2004. Total yearly precipitation in mm was

much lower in 2012 than 2004. Total precipitation in the summer months was 338.4 mm in 2004

and 230 mm in 2012.

Discussion

The original aim of this research was to discover links between landscape changes, such

as land conversion to agriculturally intensive areas, and changes in the local butterfly population.

The minimal change in greenness and vegetation indices is what has lead me to believe that

landscape changes have not been intensifying over the past 8 years. Therefore there must be

some other cause for shifts that were observed in the butterfly population. The other form of

environmental disturbance that could be attributed to compositional change is the unusual

weather patterns in 2012. For one, the extreme maximum recorded temperature for March 2012

is enormously different from the maximum temperature recorded 8 years earlier. This early and

very warm start to the growing season could have caused harm to butterfly pupae that are

seasonally triggered by warmer temperatures to emerge from cocoons. The higher mean monthly

temperature and the lower total precipitation could have also had an effect on the survival of

many species. This caused a warmer and more arid environment throughout the survey sites.

This can be damaging to host plant abundance and moisture levels in butterfly habitats. Overall

9
this change caused large differences in the abundances and distributions of many butterflies,

some species even becoming completely absent from all sites.

Looking at the species recorded in each survey year there is a clear shift in the population

composition. The newly observed species could be present due to expansions of their ranges over

the years. Species that were not recorded in 2012 that had been seen in 2004 suggest that the

fluctuations in weather proved to be unfavorable for their survival. Due to the lack of knowledge

of the time period in between survey years, it is unknown whether these species had been

steadily decreasing in numbers or if they disappeared all at once. The Jaccards index of

similarity shows that only approximately half the same species were found in the sites between

2004 to 2012. This is also indicative of the shift in species composition. Results drawn from a

comparison of the evenness of the sites between 2004 and 2012 were not similar to the

expectations. With the way the sites are distributed, a high evenness in the distribution of species

would be expected across control sites since they are meant to provide natural habitat with even

dispersal. This would be followed by a slightly lower evenness for medium impact sites and

lower still for high impact sites, since it would be more likely for species to be randomly

disbursed in sites with agricultural impact. This downward trend is shown in the data of 2004,

but not in 2012. In 2012, all of the evenness values are nearly equal, with the high impact sites

having the same evenness as the control sites. This represents a largely even dispersal of species

among both control and treatment sites in 2012. The trends in the population that were visible in

2004 are not repeated in the data from 2012.

Specialist species such as Acadian hairstreak (Satyrium acadicum), bronze copper

(Lycaena hyllus), Harris's checkerspot (Chlosyne harrissi), and silvery checkerspot (Chlosyne

nycteis), were not present in 2012 surveys after having been seen in 2004. This could be due to

10
decline in abundance of their food plants and preferred moist habitat areas when the weather

patterns changed in 2012. These species contributed to the species richness of the sites in 2004,

so their absence is a loss to species composition, especially in the control sites where they

thrived. New species in 2012, such as painted lady (Vanessa cardui), were highly abundant and

distributed across both control and treatment sites. This species was the most abundant in 2012

and is also a migrant, so it is valid to assume that the weather at their original location was

favorable and therefore they were able to appear in the area in large numbers despite the

abnormal weather patterns. Several of the other species that were new in 2012, such as question

mark (Polygonia interrogationis), red admiral (Vanessa atalanta), and mourning cloak

(Nymphalis antiopa), are also extreme habitat generalists. Other generalist species, including

common ringlet (Coenonympha tullia) and cabbage white (Pieris rapae), were among the most

highly abundant species in 2012. The fact that all of these abundant and widely spread species

are habitat generalists explains their success at the sites in 2012. Variations in climate are less

likely to affect generalist species and they are able to thrive in nearly any habitat available. These

generalist species could also be spreading their habitat ranges due to the newly vacated niches

caused by the absence of several specialist species in 2012. The widespread nature of these

generalist species is what casts a shadow over the overall landscape patterns and the less

numerous specialist species groups.

Despite the strong expectation for landscape changes to influence the butterfly

population, there was no detectable change in habitat since 2004 that could have affected

community structure. The decrease in diversity and abundance of butterflies that I had

hypothesized was not seen. Species richness in 2012 was similar across all landscapes. This can

be accounted for by the large number of generalist species, such as common ringlet, painted lady,

11
and cabbage white, that were present in all habitats. Conversely, specialist species were found in

much lower abundances. As predicted, weather patterns did effect the butterfly population. It is

possible that the extreme weather observed in 2012 caused many butterflies to be present across

all landscapes, regardless of agricultural intensity or availability of habitat. This effectively

obscures the otherwise significant links between landscape factors and butterfly population

variables such as distribution and species diversity. The change in climate may be able to affect

how butterfly species distribute themselves within the local environment. If extreme climatic

events can cause landscape ecological patterns to rapidly disappear or fluctuate, it could present

a difficulty in prioritizing conservation strategies. Future work looking at Ottawa area butterfly

populations can be done in order to verify the impacts of climatic conditions on species

composition, abundance, and distribution.

12
References

Benton, Tim G., Juliet A. Vickery, and Jeremy D. Wilson. 2003. Farmland biodiversity: is

habitat heterogeneity the key? Trends in Ecology and Evolution 18.4:182-88.

Dapporto, Leonardo, and Roger L. H. Dennis. 2013. The generalist-specialist continuum: testing

predictions for distribution and trends in British butterflies. Biological Conservation

157:229-236.

Feswick, April. 2005. Impacts of agricultural intensity on local butterfly populations. Masters

thesis.

Flick, Tatyana, Sean Feagan, and Lenore Fahrig. 2012. Effects of landscape structure on

butterfly species richness and abundance in agricultural landscapes in Eastern Ontario,

Canada. Agriculture, Ecosystems and Environment 156:123-33.

Gilbert, Lawrence E., and Michael C. Singer. 1975. Butterfly ecology. Annual Review of

Ecology and Systematics 6.1:365-395.

Layberry, Ross A., Peter W. Hall, and J. Donald Lafontaine. 1998. The Butterflies of Canada.

University of Toronto Press Incorporated, Toronto, ON.

Menndez, Rosa, Adela Gonzlez-Megas, Yvonne Collingham, Richard Fox, David B. Roy,

Ralf Ohlemller, and Chris D. Thomas. 2007. Direct and indirect effects of climate and

habitat factors on butterfly diversity. Ecology 88.3:605-11.

Stefanescu, Constanti, Jofre Carnicer, and Josep Peuelas. 2011. Determinants of species

richness in generalist and specialist Mediterranean butterflies: the negative synergistic

forces of climate and habitat change. Ecography 34:353-363.

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Weibull, Ann-Christin, Jan Bengtsson, and Eva Nohlgren. 2000. Diversity of butterflies in the

agricultural landscape: the role of farming system and landscape heterogeneity.

Ecography 23.6:743-50.

Woods, Jennifer N., John Wilson, and James R. Runkle. 2008. Influence of climate on butterfly

community and population dynamics in Western Ohio. Community and Ecosystem

Ecology 37.3:696-706.

14
Figures and Tables

Figure 1: Map of sites in the Ottawa area where surveys were conducted. Map created by April

Feswick.

Figure 2: Coleman curves representative of the sampling effort for both 2004 and 2012

(including standard deviation for all points). Species counts were calculated with singletons

removed from all sites. Curve and standard deviation values were calculated using Estimate S.

Figure 3: A measure of Jaccards index of similarity between control, medium, and high

intensity sites for 2004 and 2012 species richness.

Figure 4: The evenness index of species distribution throughout control, medium, and high

intensity sites in 2004 versus 2012.

Figure 5: Mean monthly temperatures in degrees Celsius for 2004 and 2012. Data collected at

Environment Canada weather station at Macdonald-Cartier International Airport in Ottawa (45

19' N, 75 40' W).

Figure 6: Maximum recorded temperature in degrees Celsius for each month in 2004 and 2012.

Data collected at Environment Canada weather station at Macdonald-Cartier International

Airport in Ottawa (45 19' N, 75 40' W).

15
Figure 7: Total monthly rainfall (in mm) for 2004 and 2012. Data collected at Environment

Canada weather station at Macdonald-Cartier International Airport in Ottawa (45 19' N, 75 40'

W).

Table 1: Lists of species missing in 2012 and new in 2012. * beside a species indicates that they

are a habitat specialist species. ** beside a species indicates that they are extreme habitat

generalists.

16
Figure 1

17
Figure 2

45

40

35

30

25
2004
20 2012

15

10

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

18
Figure 3

0.6

0.5

0.4
Index of Similarity

CONTROL
0.3
MEDIUM
HIGH
0.2

0.1

0
SITE

19
Figure 4

0.9

0.8

0.7

0.6
Evenness index

0.5

2004
0.4
2012

0.3

0.2

0.1

0
Control Medium High
Site

20
Figure 5

25

20

15
Mean Monthly Temperature (C)

10

5
2004
2012
0
Jan Feb Mar Apr May Jn Jl Aug Sept Oct Nov Dec

-5

-10

-15

-20

21
Figure 6

40

35
Maximum recorded temperature (C)

30

25

20 2004
2012
15

10

0
Jan Feb Mar Apr May Jn Jl Aug Sept Oct Nov Dec

22
Figure 7

160

140

120
Total monthly precipitation (mm)

100

80 2004
2012

60

40

20

0
Jan Feb Mar Apr May Jn Jl Aug Sept Oct Nov Dec

23
Table 1
Missing in 2012
Acadian Hairstreak (Satyrium acadicum)*
Bronze Copper (Lycaena hyllus)*
Columbine Duskywing (Erynnis lucilius)
Coral Hairstreak (Satyrium titus)
Crossline Skipper (Polites origines)
Grey Comma (Polygonia progne)
Harris's Checkerspot (Chlosyne harrissi)*
Longdash Skipper (Polites mystic)
Peck's Skipper (Polites peckius)
Silver-Bordered Fritillary (Boloria selene)
Silvery Checkerspot (Chlosyne nycteis)*
Striped Hairstreak (Satyrium liparops)
Tawny-Edged Skipper (Polites themistocles)
New in 2012
American Lady (Vanessa virginiensis)**
Brown Elfin (Callophrys augustinus)**
Dreamy Duskywing (Erynnis icelus)
Eastern Comma (Polygonia comma)
Giant Swallowtail (Papilio cresphontes)
Henry's Elfin (Callophyrys henrici)
Mourning Cloak (Nymphalis antiopa)**
Northern Broken-Dash (Wallengrenia
egeremet)
Northern Cloudywing (Staphylus hayhurstii)
Orange Sulphur (Colias eurytheme)
Painted Lady (Vanessa cardui)**
Pink-Edged Sulphur (Colias interior)
Question Mark (Polygonia
interrogationis)**
Red Admiral (Vanessa atalanta)**
24