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The Feasibility of the Re-introduction of the Karner Blue Butterfly to Ontario

Jennifer Bernard

Siobhan Dunets

Brittany Hammill

Elizabeth Hunter

Kelli McKay

Christopher Wagner

April 2, 2012

University of Guelph, 50 Stone Road East, Guelph, Ontario, N1G 2W1, Canada

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Executive Summary

The Karner Blue butterfly (Lycaeides melissa samuelis), named for its characteristic blue

colouration, is an extirpated species in Ontario and an endangered species in parts of the U.S.

(COSEWIC, 2000). In southern Ontario the butterfly occupied oak savanna habitat characterized

by wild lupine and mixed canopy cover as a result of fire disturbance (COSEWIC, 2000). Land

conversion and fire suppression, among other threats, have led to the loss of oak savannas which

has caused the extirpation of the Karner Blue butterfly and made re-introductions difficult to

accomplish (U.S. Fish and Wildlife Service, 2003). Previous Karner Blue butterfly re-

introduction efforts in Ohio, Indiana, and New Hampshire, as well as the re-introduction of the

Large Blue butterfly and Frosted Elfin demonstrate how re-introductions may be carried out,

often with successful results.

This literature review addressed the question: what is required to re-introduce the Karner Blue

butterfly and stabilize its population? Using information on the biology of the Karner Blue

butterfly and the findings of previous re-introduction projects, the list of requirements for re-

introduction were determined to be as follows:

● Dry sandy prairie or oak savanna habitat characteristics (Chan and Packer, 2006)

● Sands or well-drained soils that are frequently disturbed (Halpern, 2005; Pavlovic

and Grundel, 2009; Shi et al., 2005)

● Early successional sites (Chan and Packer, 2006)

● Wild lupine (1.50 stems/m2) (Chan and Packer, 2006)

● Nectar species (first brood 47.25 stems/m2 and second brood 47.85 stems/m2) (Chan

and Packer, 2006)

● Five tending ant species (showed to increase larvae survival) (Chan and Packer,

2006)

● Open/patchy canopy which is connected to facilitate movement (Huntzinger, 2003;

Halpern, 2005; Pavlovic and Grundel, 2009)

● Reduced cover/density of woody species (Halpern, 2005; Pavlovic and Grundel,

2009)

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● Increased light intensity (standard deviation of integrated light intensity of 16.82%)

(Chan and Packer, 2006)

● Minimum area of 800 ha (Smallidge and Leopold, 1997)

● Minimum population of second generation adults of 1000 individuals (Smallidge and

Leopold, 1997)

All information collected has been from peer-reviewed articles and reputable sources, such as

government websites. This literature review can be used by those involved in the re-introduction

of this species as an up to date collection of significant information, as well as providing

information for further research that may need to take place.

Key words: Karner Blue butterfly, re-introduction, extirpated, endangered, wild lupine biology,

description, mating, wild lupine, predators, disease, parasites, ants, threats, habitat requirements

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Table of Contents Executive Summary ..................................................................................................................................... 2  

Table of Contents......................................................................................................................................... 4  

1. Introduction.............................................................................................................................................. 5  

2. Biology .................................................................................................................................................... 6  

General Description ................................................................................................................................. 6  

Life Cycle ................................................................................................................................................ 7  

Mating...................................................................................................................................................... 9  

Food Source ............................................................................................................................................. 9  

Predators and Parasites .......................................................................................................................... 11  

Disease ................................................................................................................................................... 12  

3. Habitat ................................................................................................................................................... 13  

4. Relationships With Other Species.......................................................................................................... 16  

5. Wild Lupine ........................................................................................................................................... 19  

Habitat and the Importance of Fire ........................................................................................................ 21  

Reasons for the Decline of Wild Lupine................................................................................................ 21  

Re-introduction of Wild Lupine and Maintenance of Populations ........................................................ 21  

6. Re-introduction Projects for the Karner Blue Butterfly in the US ......................................................... 23  

Captive Rearing ..................................................................................................................................... 24  

Re-introduction Projects of Other Species ............................................................................................. 25  

7. Requirements for the Re-introduction of the Karner Blue Butterfly...................................................... 26  

Habitat Requirements ............................................................................................................................ 27  

Habitat Assessment Prior to Re-introduction......................................................................................... 28  

Restoration and Maintenance of Re-introduction Sites.......................................................................... 28  

Central Threats....................................................................................................................................... 30  

8. Methods and Materials........................................................................................................................... 31  

9. Conclusion ............................................................................................................................................. 31  

References ................................................................................................................................................. 34  

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1. Introduction

With the world’s growing and expanding human population, many species have become

endangered or extirpated because of poaching and loss of habitat, among other reasons. The

specific species of concern in this literature review is the Karner Blue butterfly (Lycaeides

melissa samuelis). Historically, several colonies of this species were present throughout southern

Ontario, including the most recently documented colonies at Port Franks and St. Williams (Fig.

1; COSEWIC, 2000). However, the Karner Blue butterfly has since been listed as extirpated in

Canada and has been declared endangered throughout the U.S. (Grundel et al., 1998). The

Karner Blue butterfly is in this situation because it has very specific requirements for continued

survival that are not currently being met (Forrester et al., 2005).

Figure 1. Map of historical locations of Karner Blue butterfly populations in southern Ontario. Provided

by COSEWIC (2000).

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This butterfly is important because it could be considered an “umbrella species”, that is, a

species whose own conservation and that of its habitat protects other species in a given area

(Chan and Laurence, 2006). The problem that is addressed in this review is the lack of

consolidation of all pertinent information to the Karner Blue butterfly and its re-introduction.

This literature review contains a comprehensive overview of relevant research concerning the

life cycle and habitat needs of this species. Using this information in combination with the

findings of previous butterfly re-introduction projects, general recommendations are made

regarding requirements for re-introduction.

2. Biology

General Description

The Karner Blue butterfly is a small (wingspan of 2.2-3.2cm) North American species of

butterfly named for its characteristic blue colouration (Fig. 2; COSEWIC, 2000). The adult male

and female differ in wing colouration, with the male exhibiting iridescent blue wings surrounded

with a black outline and white outer fringe (COSEWIC, 2000). The wings of the female display a

gradient from dull-blue in the center to purple-brown near the edges, and have a border of black

spots with orange crescents (COSEWIC, 2000). The underside of the male and female wings is

grey with black spots, but the females again exhibit orange crescents surrounding silver spots

that are outlined in black (COSEWIC, 2000).

Figure 2. Markings of the adult female (left), adult male (middle), and underside of the wings of both

sexes (right) of the Karner Blue butterfly. Provided by U.S. Fish and Wildlife Service (2003).

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Life Cycle

The life cycle of the Karner Blue butterfly has been extensively documented. The Karner Blue

butterfly undergoes four life stages, consisting of egg, larvae, pupae, and adult (COSEWIC,

2000). Each summer, two generations of eggs are hatched, one around mid-April and the other in

June, although exact timing varies depending on weather conditions (Fig. 3; Haack, 1993). The

first generation hatches from overwintered eggs laid by the second generation from the previous

summer (Cryan and Dirig, 1978). The first generation is 3-4 times smaller than the second,

possibly due to less food availability at that time of year (COSEWIC, 2000). The ratio between

sexes in each generation is near 1:1 (Packer, 1987).

Figure 3. Approximate timing of life stages of the first and second generations of Karner Blue butterflies.

Timing may vary based on climate and weather (ie. warm conditions will cause hatching and thus other

stages to occur earlier). Provided by Haack (1993).

The eggs of the Karner Blue butterfly are pale green, flattened and round in shape (Fig. 4) and

are deposited on the lower leaves and leaf petioles of the wild lupine plant (around the first leaf

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petiole) (COSEWIC, 2000; Packer, 1990). Eggs from the first generation hatch after

approximately 7 days, whereas eggs from the second generation fall to the ground with the wild

lupine leaves in winter and are preserved there before hatching next spring (COSEWIC, 2000).

Figure 4. Drawings (left and middle) and photograph (right) depicting the size, shape, and overall

appearance of the Karner Blue butterfly egg. Provided by U.S. Fish and Wildlife Service (2003).

Karner Blue butterfly larvae are dorsally flattened and pubescent, with green bodies that

effectively camouflage them on the wild lupine plant (Cryan and Dirig, 1978; Packer, 1987).

Savignano (1990) proposed there are four larval instar stages while Opler and Krizek (1984)

proposed five. The larval stage lasts from 18 to 21 days (Cryan and Dirig, 1978). Lane (1999)

found larvae have improved growth and survivorship on wild lupine growing in the shade.

Following the larval stage, the pupa stage lasts for 8 days (Cryan and Dirig, 1978). Starting out

pea green, pupae turn blackish-purple before adult immergence (Haack, 1993). Pupae can be

found attached to lupine plants or in underlying litter (Hess, 1993).

Adult Karner Blue butterflies have an average life span of 5 days in the wild (Cuthrell, 1990). In

Ontario, the flight season for the first generation takes place, on average, from May 25 to June 24

and the second generation from July 20 to August 18 with variation based on the weather during

the particular year (COSEWIC, 2000). Unlike larvae, adults prefer sunlit locations, which Lane

(1999) proposes is likely for the purpose of thermoregulation. Adults have weak flying

capabilities and as such tend to be sedentary, not travelling farther than one kilometer (Lawrence

and Cook, 1989). In warm temperatures butterflies are active all day (Lawrence and Cook,

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1989). However, overly hot or cold days and rain reduce and stop activity respectively (Haack,

1993; Lawrence and Cook, 1989). Temperatures below 24.6°C for females or 26.4°C for males

lead to greatly reduced flight activity, whereas heat stress tends to occur at temperatures above

35.6°C in females and 36.8°C in males (Lane, 1999). Temperature sensitivity in the Karner Blue

butterfly means that daily temperature affects which habitat areas are used by the butterflies

(Lane, 1999). Butterflies will tend to occupy areas with open canopies at lower temperatures and

seek shaded or partially shaded areas at high temperatures (Lane, 1999). Temperature influences

where oviposition occurs in the same way, with oviposition occurring in shaded areas when

ambient temperature is high (Lane, 1999).

Mating

Adult butterflies mate and lay eggs within their five day lifespan (COSEWIC, 2000). Grundel et

al. (1998) proposed that mating occurs mainly in large openings of canopy cover, as opposed to

more shaded areas. Males are only weakly territorial, and as a result disperse slightly more than

females (COSEWIC, 2000). Breeding sites historically varied in Ontario and coincided spatially

with historical colony locations (COSEWIC, 2000).

Food Source

The larval stage of the Karner Blue butterfly relies solely on wild lupine as a food source (Fig 5;

Haack, 1993). Younger larvae feed only on the leaves of the plant, but later stages may feed on

buds and flowers, although to a lesser extent (Haack, 1993). Packer (1987) and Cryan and Dirig

(1978) propose that the first instars feed within the leaf, and that subsequent stages consume the

underside and inner flesh. However, Schweitzer (1989) has suggested that the larvae instead feed

on the upper surface of the leaf.

Quality of the wild lupine plant can vary due to site conditions and this can affect the larva's

fitness (Grundel et al., 1998). Grundel et al. (1998) found that larvae which fed on shade-grown

wild lupine grew faster than those that fed upon lupine grown in the sun. This likely occurs as a

result of sun vs. shade exposure influencing plant nutrients, flowering state, plant health, and

levels of secondary plant compounds such as alkaloids (U.S. Fish and Wildlife Service, 2003).

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Non-flowering wild lupine appears to provide a better food source (increases larvae survival)

than flowering wild lupine (Maxwell, 1998).

Figure 5. (A) Line drawing depicting shape of wild lupine flower, leaf and seed pod. (B) Photograph of

wild lupine plant with blue flowers. Flower colour can range from pink to blue to white. Provided by

Natural Resources Conservation Service (2012).

Adult butterflies will consume nectar from almost any available plant (Table 1; Savignano and

Zaremba, 1993). Food source species vary between the two generations and are also based on the

site (COSEWIC, 2000). For example, Packer (1990) listed seven plant species used by the first

generation of butterflies of the Port Franks population in Ontario, and an additional five available

only to the second generation. More nectaring flowers are available to the second generation,

including Butterfly Weed (Asclepias tuberose) and New Jersey Tea (Ceanothus americana) at

the Port Franks site (Schweitzer, 1984). Lack of nectaring plants in an area can limit dispersal

despite the presence of wild lupine (Packer, 1987).

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Table 1. Scientific and common names of the most common food (nectar providing) plants of the first

and second generations of Karner Blue butterflies. Provided by COSEWIC (2000).

Common Nectaring Plants of the Karner Blue Butterfly in Ontario.

First Generation

Wild Lupine, Lupinus perennis

Blackberry, Rubus allegheniensis

Strawberry, Fragaria virginiana

Yellow hawkweed, Hieracium pilosella

Dewberry, Rubus flagellaris

Thyme-Leaved Sandwort, Arenaria serpyllifolia

Lyre-Leaved Rock Cress, Arabis lyrata

Juneberry, Amelanchier sp.

Wild Geranium, Geranium maculatum

Puccoon, Lithospermum caroliniense

Second Generation

Knapweed, Centaurea maculosa

New Jersey Tea, Ceanothus americanus

Butterfly Weed, Asclepias tuberosa

Flowering Spurge, Euphorbia corollata

Dwarf Blazing-star, Liatris cylindracea

Wild Bergamot, Monarda fistulosa

Black-Eyed Susan, Rudbeckia hirta

Predators and Parasites

Generalist predators including Polistes wasps (Polistes fuscatus, and P. metricus), pentatomid

stink bugs (Podisus maculiventris), Formica ants (F. schaufussi and F. incerta) and spiders have

all been noted to prey on Karner Blue butterfly larvae (Savignano, 1990; Packer, 1987).

Shellhorn et al. (2005) found that the seven spotted ladybug (Coccinella septempunctata), a

biocontrol species introduced for aphid control, tends to co-occur with the Karner Blue butterfly

and will feed on the butterfly larvae. Shellhorn et al. (2005) found that predation of the larvae by

C. septempunctata significantly increased with predator density. Therefore, these results would

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suggest that the proposed habitat for the re-introduction of the butterfly should be located a

minimum distance away from agricultural fields applying C. septempunctata (Shellhorn et al.,

2005). However, a recommended minimum distance has not yet been quantified, and as such

further research on C. septempunctata dispersal is needed (Shellhorn et al., 2005).

Predators of the adult butterfly include crab spiders, robber flies, ambush bugs, assassin bugs,

and dragonflies (Haack, 1993). Grazing by deer can also contribute directly to larvae mortality

(Schweitzer, 1994). It is currently unknown whether birds feed on Karner Blue butterflies,

although beak marks have occasionally been found on wings of the adult butterfly (U.S. Fish and

Wildlife Service, 2003).

Four parasites of Karner Blue butterfly larvae were reported by Savignano (1990): the tachinid

fly (Aplomya theclarum), the braconid wasp (Apanteles sp., possibly epinotiae), and two

ichneumonid wasps (Neotypus nobilitator nobilitator and Parania geniculate). However, key

predators and parasites, as well as the extent of predation and parasitism, have yet to be

investigated (Schellhorn et al., 2005). It is currently unknown how significant an impact

predation has, relative to resource availability, on the survival of various life stages of the Karner

Blue butterfly.

Disease

The impacts of disease on Karner Blue butterfly populations is still widely unknown and requires

further investigation (Haack, 1993). However, some significant research and documentation has

been conducted on a bacterial infection affecting the Karner Blue butterfly. Nice et al. (2009)

found widespread Wolbachia in Karner Blue butterfly populations west of Lake Michigan.

Wolbachia is a heritable endosymbiotic bacteria that is maternally inherited and alters its host’s

reproductive biology (Nice et al., 2009). While these infections can manifest in varying

detrimental phenotypes, this particular infection appears to have induced cytoplasmic

incompatibility in the western Karner Blue butterfly population (Nice et al., 2009). This means

sperm of infected males cannot fertilize eggs of uninfected females or females infected with a

different strain (Nice et al., 2009). The result is fewer viable embryos being produced, leading to

a decrease in population until the infection is fixed or disappears (Nice et al., 2009). Though

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temporary, this decrease in population size can put individual populations at risk (Nice et al.,

2009).

3. Habitat

The Karner Blue butterfly is typically found in frequently disturbed, early successional habitats

which in Ontario consisted of oak savannas, beach dunes, or woodlands prior to extirpation

(COSEWIC, 2000). These habitats are characterized by well-drained sandy soils and open

canopy sustained by fire disturbance, with canopy cover ranging from zero to between fifty and

eighty percent (U.S. Fish and Wildlife Service, 2003). Forbs and grasses are the main understory

vegetation and tend to colonize in canopy openings (U.S. Fish and Wildlife Service, 2003).

Habitat structure is maintained by conditions wet enough (determined by precipitation, soil

drainage, and topography) to permit tree growth while maintaining conditions dry enough to

allow for cyclic fire disturbance (U.S. Fish and Wildlife Service, 2003). Habitat temperature is

typical of that found in southern Ontario (COSEWIC, 2000). The Karner Blue butterfly thrives in

this highly specific habitat for a number of reasons, including the nectar plants and symbiotic ant

species the habitat supports (U.S. Fish and Wildlife Services, 2003). However, the main features

of oak savanna habitat conducive to Karner Blue butterfly survival is wild lupine presence and

mixed canopy cover (Grundel et al., 1998).

While the necessity of wild lupine for larvae survival has already been commented on, canopy

structure is also key to reproduction and adult and larval survival. Grundel et al. (1998) reported

the percentage of male and female Karner Blue butterfly habitat use as well as the percentage of

lupine cover is directed as a function of canopy cover.

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Figure 6. Male and female habitat use and percent lupine cover as a function of canopy cover. Provided

by Grundel et al. (1998).

It is evident that male and female Karner Blue butterflies show behavioural separation of habitat

use in relation to canopy cover (Fig. 6; Grundel et al., 1998). This is largely due to the fact that

male Karner Blue butterflies prefer larger canopy openings for mating and feeding on various

nectar sources, which generally decrease with increased canopy cover (Grundel et al., 1998).

Conversely, female Karner Blue butterflies prefer a canopy cover percentage of around 30-60%

for oviposition (Fig. 7; Grundel et al., 1998). It is apparent that the expected use and the actual

distribution of oviposition sites differs significantly (Grundel et al., 1998). As mentioned earlier,

Grundel et al. (1998) also showed that larvae preferentially fed on larger lupine plants that

tended to grow in shaded areas. Overall, these findings point at the importance of habitat with a

mixture of open canopy and shaded areas to ensure Karner Blue butterfly survival.

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Figure 7. Distribution of oviposition sites and mean lupine abundance score as a function of overhead

canopy cover. Striped bars represent the distribution of oviposition sites, empty bars the expected use, and

the line the mean lupine abundance score. Provided by Grundel et al. (1998).

Abundance of suitable Karner Blue butterfly habitat such as oak savannas has been rapidly

declining (U.S. Fish and Wildlife Service, 2003). Loss or degradation of habitat may occur as a

result of actual habitat destruction, fragmentation, or increase in canopy cover of native habitat.

Destruction or fragmentation occurs as a result of conversion of land to urban or agricultural uses

(U.S. Fish and Wildlife Service, 2003). Increase in canopy cover has occurred as a result of fire

suppression, in an effort to prevent property damage and also preserve other forest species, and

also likely as a result of extensive planting of pine trees in native habitat (Konecny, 1986; U.S.

Fish and Wildlife Service, 2003).

In areas where the Karner Blue butterfly is not yet extinct, habitat loss has often forced the

species into remnant early successional habitats such as power-line corridors, trails and road

systems of forests, airports, and military training areas (Forrester et al., 2005; Smith et al., 2002,

Warren et al., 2007). These new habitat areas all share structurally similar attributes such as: the

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amount of canopy cover, understory vegetation, soil types, and disturbance regimes (Forrester et

al., 2005).

4. Relationships With Other Species

Like many members of the Lycaenidae family, the Karner Blue butterfly larvae form a

mutualistic relationship with certain ant species (Shapiro, 1973). The ants benefit from the

Karner Blue butterfly through harvesting the larvae that provide a nectar-like solution from

special glands (Savignano, 1990; Savignano, 1994). Tending by ants has been shown to increase

survival of larvae in the field (Savignano, 1990; Savignano, 1994). Savignano (1990) showed

lower predation of larvae with ant tending, suggesting larval benefit is a result of protection from

predation and parasitism. A list of all the tending ant species found to be associated with the

Karner Blue butterfly was provided by the U.S. Fish and Wildlife Service (2003) (Table 2).

Table 2. Ant species tending Karner blue butterfly larvae and pupae. Ont. (Ontario), WI (Wisconsin), MN

(Minnesota), NY (New York), MI (Michigan).

Ant Species Tending Larvae Locality References

Aphaenogaster rudis Ont. Packer (1991)

Brachymyrmex debilis Emery MN, WI Lane (1999)

Camponotus americanus Mayr NY Savignano (1994)

Camponotus ferrugineus WI Bleser (1992)

Camponotus novaeboracensis Fitch NY Savignano (1994)

Camponotus pennsylvanicus Ont. Packer (1991)

Crematogaster ashmeadi WI Bleser (1992)

Crematogaster cerasi Fitch NY Savignano (1994)

Crematogaster lineolata (Say) MI Herms (1996)

Dolichonderus (Hypoclinea)plagiatus Mayr NY, WI Savignano (1994), Lane (1999)

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Dolichonderus mariae Forel MI, WI Herms (1996), Lane (1999)

Dolichonderus pustulatus Mayr MI Herms (1996)

Formica difficilis Emery NY Savignano (1994)

Formica exsectoides Ont. Packer (1991)

Formica fusca WI Bleser (1992)

Formica lasioides Emery NY Savignano (1994)

Formica Montana WI Bleser (1992)

Formica (Neoformica) incerta Emery NY, MN, WI Savignano (1994), Lane (1999)

Formica (Neoformica) nitidventris Emery NY Savignano (1994)

Formica (Neoformica) schaufussi Mayr NY, MI Savignano (1994), Herms (1996)

Formica neogatates Emery MI Herms (1996)

Formica obscuripes Forel WI, MI Herms (1996), Lane (1999)

Formica obscuriventris Mayr MI Herms (1996)

Formica querquetulana Wheeler NY Savignano (1994)

Formica schaufussi WI Bleser (1992)

Formica subnuda Emery WI Lane (1999)

Formica subsericea Say NY, MI, WI Savignano (1994), Herms (1996),

Lane (1999)

Lasius alienus Foerster NY, MN, WI Savignano (1994), Lane (1999)

Lasius neoniger Emery NY, MI Savignano (1994), Herms (1996)

Monomorium emarginatum DuBuois NY Savignano (1994)

Monomorium pharaonis (L.) MI Herms (1996)

Myrmica americana Weber NY, MI, MN,

WI Savignano (1994), Herms (1996),

Lane (1999)

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Myrmica emeryana Forel MN, WI Lane (1999)

Myrmica fracticornis Emery NY, MI Savignano (1994), Herms (1996)

Myrmica lobifrons MN, WI Lane (1999)

Myrmica punctiventris Ont. Packer (1991)

Myrmica sculptilis NY Savignano (1990)

Paratrechina parvula Mayr NY Savignano (1994)

Prenolepsis imparis (Mayr) MN Lane (1999)

Tapinoma sessile Say NY,WI,MN Bleser (1992), Savignano (1994),

Lane (1999)

Tetramorium caespitum WI Bleser (1992)

Ant Tending Pupae

Crematogaster lineolata (Say) WI Lane (1999)

Dolichonderus tashenbergi (Mayr) WI Lane (1999)

Formica obscuripes Forel WI Lane (1999)

Lasius alienus Foerster WI Lane (1999)

Lasius neoniger Emery WI Lane (1999)

Leptothorax sp. WI Lane (1999)

Myrmica emeryana Forel WI Lane (1999)

Tapinoma sessile Say WI Lane (1999)

In addition to this direct mutualism with ants, the Karner Blue butterfly could be considered to

share an important indirect relationship with numerous other species. A number of species co-

occur with the Karner Blue butterfly including plants such as the Prairie thistle (Cirsium hillii

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Fernald) and insects such as the Frosted Elfin (Callophrys irus Godart), among others (Chan and

Packer, 2006).

5. Wild Lupine

Wild lupine is essential to the survival of Karner Blue butterfly larvae and therefore a working

knowledge of its properties and habitat requirements is crucial to the butterfly’s re-introduction.

Wild lupine is a nitrogen fixing, herbaceous perennial plant that is found on sandy, nutrient poor,

slightly acidic soils (Pavlovic and Grundel, 2009; Bernhardt et al., 2008; Halpern, 2005). The

plant can grow to 20-60cm in height and appears to grow best in small forest clearings where it

experiences partial shade, as it suffers in habitat with too little or too much direct sunlight

(Pavlovic and Grundel, 2009). The habitat requirements of wild lupine overlap with those of the

Karner Blue butterfly in that it is typically found in oak savannas and clearings of disturbed areas

(USDA Forest Service, n.d.).

The life cycle of wild lupine generally extends from April to July with flowering taking place in

May or June in northern regions (USDA Forest Service, n.d.). The plant can begin flowering in

its second year at the earliest, depending on environmental conditions, and may not sprout every

year (USDA Forest Service, n.d.). Wild lupine can produce sexually or asexually, although

asexual reproduction results in less fruit and seed production (Shi et al., 2005; Bernhardt et al.,

2008). Pollination occurs via bumblebees (Bombus spp.), orchard bees (Osmia spp.), honeybees,

eastern carpenter bees, and butterflies (including the Karner Blue butterfly) (Bernhardt et al.,

2008; Halpern, 2005; USDA Forest Service, n.d.). Following pollination 3-5cm seed pods

containing on average four to nine seeds are formed (U.S. Fish and Wildlife Service, 2003).

Seeds are dispersed with the opening of the seed pod and can be transported a few feet from the

plant, meaning wild lupine dispersal or colonization is likely slow (U.S. Fish and Wildlife

Service, 2003). These seeds may then germinate the same summer or remain dormant for three

years (USDA Forest Service, n.d.). The number of seeds that germinate depends on water

availability, temperature, predation, and scarification (USDA Forest Service, n.d.).

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Historically, this plant had a range throughout eastern North America extending from central and

southern Ontario to Minnesota and south to Florida and Louisiana (Gleason and Cronquist,

1963). However, it is now becoming rare throughout its Ontario and U.S. range (COSEWIC,

2000). In Ontario, wild lupine is found primarily in discreet, localized colonies on the Norfolk

sand plain and on the dunes of the Pinery Provincial Park or Grand Bend area (Fig. 8;

COSEWIC, 2000). Although several other populations have been confirmed, no investigation

has been performed since the 1980’s (COSEWIC, 2000).

Figure 8. Distribution of purported wild lupine populations in southern Ontario. Provided by COSEWIC

(2000).

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Habitat and the Importance of Fire

The shade-intolerant wild lupine tends to grow in habitat maintained at early successional stages

by fire disturbance regimes (Bernhardt et al., 2008; Pavlovic and Grundel, 2009; Smallidge and

Leopold, 1997; Smallidge et al., 1996).

Wild lupine flourishes in fire-disturbed habitat as its nitrogen fixing capability provides it with

an advantage over potential competitors (due to the loss of nitrogen during fire events)

(Maxwell, 1998; Pfitsch and Williams, 2009). Regular fire disturbance regimes also serve to

prevent vegetation cover from becoming too dense, which would be detrimental to the shade

intolerant wild lupine (Pavlovic and Grundel, 2009). In a study by Grigore and Tramer (1996)

they compared burned and unburned populations of wild lupine based on germination, growth,

flowering, and seed set. They found that burning plots affected individuals by increasing plant

nitrogen content, increasing leaf cover, increasing plant biomass, and increasing seed set

(Grigore and Tramer, 1996). They also found that burning plots increased seed and seedling

mortality, but this effect could be reduced by staggering burns by a minimum of two years and

only performing burning before seedling emergence in the fall (Grigore and Tramer, 1996;

Smallidge and Leopold, 1997).

Reasons for the Decline of Wild Lupine

Wild lupine populations have declined or been extirpated due to decreasing habitat disturbances

in the form of fires (Smallidge and Leopold, 1997). Without disturbance former wild lupine

habitat can reach later stages of succession resulting in lowered light intensity and therefore

lowered reproductive ability of wild lupine (Smallidge and Leopold, 1997). The introduction of

coniferous plantations, in its original habitat, is also harmful to the shade intolerant plant

(COSEWIC, 2000; Pavlovic and Grundel, 2009). Urbanization is another major factor in the

decline of wild lupine since many savannas have been transformed into urban areas (U.S. Fish

and Wildlife Service, 2003).

Re-introduction of Wild Lupine and Maintenance of Populations

There are many factors that can affect the establishment of wild lupine in re-introduction areas.

In order to ensure the health and survival of wild lupine when growing it for re-introduction

 22  

purposes a few factors must be considered. Larger seed sizes of wild lupine increase the growth

of seedlings, survival rates, plant size, and reproductive ability (Halpern, 2005). Wild lupine

seeds have a seed coat that can reduce the germination of seeds (Mackay et al., 1996). In order to

remove the seed coat scarification needs to occur by placing the seeds in concentrated sulfuric

acid (36N) for 30 minutes which will increase germination significantly (Mackay et al., 1996).

Seeds of wild lupine germinate the fastest at temperatures between 24-29°C (Mackay et al.,

1996). Wild lupine does best when planted in the fall (Pavlovic and Grundel, 2009). This allows

the seeds to cold stratify during the winter (Pavlovic and Grundel, 2009). Wild lupine also

performs well in disturbed sandy substrate (Smallidge and Leopold, 1997).

Optimal site selection and maintenance must also be considered when re-introducing wild lupine

populations or maintaining current populations. As stated previously, wild lupine performs best

in partial shade and open areas, but does not perform well in dense shade (Halpern, 2005;

Pavlovic and Grundel, 2009). Wild lupine thrives in areas of light intensity over 69% that have

been recently disturbed either through natural fire regimes or management practices (Smallidge

et al., 1996). Low litter cover, moderate vegetative cover, and intermediate canopy cover is ideal

for wild lupine habitat (Halpern, 2005; Pavlovic and Grundel, 2009). Prescribed burnings can be

used to regulate litter cover, vegetative cover, and canopy cover, but must not reduce vegetative

cover too greatly (Pavlovic and Grundel, 2009). Mechanical treatments or herbicide treatments

can also effectively reduce woody species cover and increase the light intensity and therefore

increase and maintain the wild lupine population (Smallidge and Leopold, 1997).

Wild lupine experiences significant inbreeding depression and therefore pollination and

outcrossing are vital to long term population viability (Shi et al., 2005). Densely packed

populations of wild lupine that are greater than a few hundred individuals are more appealing to

pollinators because of increased foraging efficiency (Shi et al., 2005). Increased pollinator

presence within wild lupine populations will increase pollination rates and outcrossing and will

therefore allow the population to remain sustainable in the long term (Shi et al., 2005).

 23  

6. Re-introduction projects for the Karner Blue butterfly in the U.S.

In the United States, re-introductions of the Karner Blue butterfly have been attempted in Ohio,

Indiana, and New Hampshire (U.S. Fish and Wildlife Service, 2003; Suckling, 2006). The goal

of these re-introduction projects is to establish several viable metapopulations of adult butterflies

within the Karner Blue butterfly’s range, in order to ensure that populations become established

and begin reproducing successfully on their own (U.S. Fish and Wildlife Service, 2003). More

specifically, the goal is establishment of populations of approximately 3000 butterflies in

nineteen metapopulations and 6000 butterflies in eight large metapopulations (U.S. Fish and

Wildlife Service, 2003).

In 1988 the Karner Blue butterfly was considered extirpated from Ohio and in 1998 a re-

introduction program for this butterfly began (Ohio Karner blue butterfly Recovery Team:

OKBBRT, 2010). The goal of this re-introduction program was to have several viable

metapopulations of the Karner Blue butterfly present to ensure the establishment of successfully

reproducing populations (OKBBRT, 2010; U.S. Fish and Wildlife Service, 2003). This program

is currently considered to be partially successful, as minimum viable population sizes for the

butterfly have not yet been reached (Soorae, 2008; OKBBRT, 2010). Even though the population

goal has not been reached this re-introduction program has provided valuable information with

regards to key Karner Blue butterfly re-introduction requirements and learning experiences,

which will help ensure the success of future re-introduction programs (OKBBRT, 2010). The

main lesson that has been taken away from this project is that the following factors are important

to successful re-introduction: an abundant supply of wild lupine populations, appropriate canopy

cover to protect the butterflies from the elements (freeze events, severe storms), the appropriate

abundance of nectar plants, appropriate abundance of mutualistic ant species, considering the

impact of land use changes, and ensuring public awareness of the re-introduction program

(OKBBRT, 2010). Soorae (2008) summarizes the Ohio re-introduction program by highlighting

some of the difficulties associated with the program. The Ohio re-introduction program involved

Karner Blue butterflies bred in captivity and then released into the wild (Soorae, 2008). One of

the main challenges faced was the maintenance of wild lupine populations large enough in size

to ensure successful reproduction of the butterflies in the wild (Soorae, 2008). Wild lupine plants

 24  

experienced a lot of stress due to larvae feeding and usually were unable to recover (Soorae,

2008). Therefore new plants had to be grown and introduced every year to ensure a sufficient

supply for the Karner Blue butterflies (Soorae, 2008). Soorae (2008) also mentions that a widely

spaced oak canopy cover would help protect the butterflies from freeze events.

Re-introduction projects were also started in New Hampshire and Indiana in 2000 (U.S. Fish and

Wildlife Service, 2003; Suckling, 2006). The Nature Conservancy and several reports have

stated that re-introduction of the Karner Blue butterfly into the Indiana site have been successful

thus far but monitoring programs are still required (U.S. Fish and Wildlife Service, 2003;

Suckling, 2006). The New Hampshire re-introduction project has been successful and the

populations of the Karner Blue butterfly have started mating and reproducing in the wild on their

own since 2003 (Suckling, 2006). This re-introduction site is approximately 300 ha, located in

the Pine Barrens in Concord (New Hampshire Fish and Game Dept, 2012a). The wildlife action

plan for the Karner Blue butterfly profile taken from the New Hampshire Fish and Game website

highlights some of their re-introduction project details (New Hampshire Fish and Game Dept,

2012b). They find that suitable habitat sites must be at least 0.25ha. Habitat areas of 0.25-5ha

should have at least 500-800 lupine stems per 0.4ha, and habitat areas greater than 5ha should

have at least 0.1 lupine stem per m2 or 405 lupine stems per 0.4ha (New Hampshire Fish and

Game Dept., 2012b). Monitoring programs are still ongoing for this project as well, and both the

Indiana and New Hampshire projects will require habitat expansion and an increase in the wild

lupine and nectar source plants population numbers in order to further improve the establishment

of the Karner Blue butterfly (Suckling, 2006).

Captive Rearing

Captive rearing programs have been performed in the past for the Karner Blue butterfly and

could help to bolster populations with low numbers or be used to re-establish populations which

have disappeared (Herms et al., 1996).

Herms et al. (1996) provide a detailed account of how captive rearing may be successfully

carried out. First the Karner Blue butterflies were collected during their spring generation from

two areas in Michigan (Herms et al., 1996). Individuals were caught and placed in glassine

 25  

envelopes and were then placed in a bag kept in a cooler at 20°C (Herms et al., 1996). Healthy

wild lupine was collected from fields near the areas where the Karner Blue butterflies were

collected (Herms et al., 1996). The stems of the wild lupine were cut and then placed in a

container filled with water where they were re-cut underwater (Herms et al., 1996). Once the

wild lupine arrived at the laboratory they were placed in plastic bags and stored at 5°C until

needed (Herms et al., 1996). Wild lupine was harvested every five days in order to ensure an

ample amount of fresh plants were available (Herms et al., 1996).

The butterflies were kept in aluminium frame cages (61x61x61cm) in an environmental chamber

at 24-26°C with an 18:6 hour light:dark photoperiod and relative humidity of 57-68% for five

days (Herms et al., 1996). In each cage butterflies were provided with a water source (wet

sponge), partial shading (paper towels covering one part of the cage), a nectar source (petri dish

with 5% honey and 95% water), and a wild lupine stem (20-30cm tall) in a water filled flask

(Herms et al., 1996).

Eggs were removed from leaves and placed into individual small plastic cups (Herms et al.,

1996). Cups were then placed in a lidded plastic box with moist paper towel to provide a relative

humidity of 80-85% (Herms et al., 1996). When eggs were close to hatching wild lupine foliage

was added (Herms et al., 1996). Once eggs hatched fresh wild lupine was provided every two

days (Herms et al., 1996). Once larvae reached their third or fourth level instar they were reared

on petri dishes with an entire wild lupine leaf which was replaced every two days (Herms et al.,

1996). Eggs, larvae and pupae were kept in environmental chambers at 24°C with a 18:6 hour

light:dark photoperiod (Herms et al., 1996). After adults emerged they were placed in a

refrigerator for a day at 5°C prior to field release (Herms et al., 1996). This method, or

derivations of it, may be crucial to future re-introduction projects.

Re-introduction Projects of Other Species

The Large Blue butterfly (Maculinea arion) was considered extinct from Britain in 1979

(Cookson, 2009; Barnett and Warren, 1995). Through a re-introduction project, which started

over 25 years ago, the species has now been successfully re-introduced into the wild (Cookson,

2009; Barnett and Warren, 1995). A major reason for the decrease in population of the Large

 26  

Blue butterfly was the lack of knowledge about the driving factors of the decline (Cookson,

2009; Barnett and Warren, 1995). In the 1970’s it was determined that the driving factors for this

decline in population were the decline in Myrmica sabuleti populations (an ant species critical

for the growth and development of the Large Blue butterfly larvae) and loss of habitat (Thomas,

Simcox, and Bourn, 2011). Habitat for the Large Blue butterfly has been restored to over 50

areas and within those areas 33 are currently occupied by the Large Blue butterfly (Cookson,

2009). Due to the success of the approaches and models used in this re-introduction project they

are now being used for the re-introduction of other rare species of butterflies across Europe

(Cookson, 2009; Spitzer et al., 2009). The success of this study shows that with the right

implementation and information the re-introduction of species can be successful (Cookson, 2009;

Spitzer et al., 2009). This study also demonstrates that presence of mutualistic ant species can be

a critical factor in the survival of a specific butterfly population.

The Frosted Elfin butterfly (Callophrys irus (Godart), another re-introduction success story, also

demonstrates how crucial presence of a supporting organism can be to butterfly population

survival (Pfitsch and Williams, 2009). Frosted Elfin butterflies rely on wild lupine to survive and

reside in small isolated colonies in close proximity to their host plant throughout their life span

(Pfitsch and Williams, 2009). An experiment conducted in the Rome Sand Plains in central New

York has shown that by decreasing canopy cover more light reaches the ground providing

favourable habitat for wild lupine (Pfitsch and Williams, 2009). With an increase in flowering

stems of wild lupine there was an increase in Frosted Elfin butterfly reproduction (Pfitsch and

Williams, 2009). The Frosted Elfin butterflies responded to this increase in habitat within a year

of the removal of some trees (Pfitsch and Williams, 2009). This success in increasing Frosted

Elfin butterfly populations through the expansion of wild lupine habitat is important for wild

lupine specialist butterflies and could be implemented in other areas where these species are

declining (Pfitsch and Williams, 2009).

7. Requirements for the Re-introduction of the Karner Blue Butterfly

The preceding accounts of Karner Blue butterfly biology and past re-introduction projects point

towards potential requirements for future re-introduction projects. This final analysis will draw

 27  

from this information in combination with directly applicable new information to clearly outline

the major requirements Karner Blue butterfly re-introduction projects must fulfill to improve

probability of success.

Habitat requirements

The predominant reason for this species being extirpated can be attributed to the fact that it has

very specific habitat requirements, therefore putting it at a high risk of extinction (Forrester et

al., 2005). A potential re-introduction site must meet the following habitat requirements in order

for the introduced Karner Blue butterfly population to have potential to survive: possess sands or

well-drained soils that are frequently disturbed, be an early successional site with open/patchy

canopy cover, contain wild lupine and various nectar species, have sufficiently high light

intensity at the forest floor, and have sufficiently large area (Huntzinger, 2003; Smallidge et al.,

1996; Kleintjes et al., 2003).

In a study done by Chan and Packer (2006) minimum standards for the requirements needed to

re-introduce the Karner Blue butterfly were highlighted. They calculated these minimum

standards through assessing the minimum values of four variables in U.S. sites where the Karner

Blue butterfly is present (Chan and Packer, 2006). The minimum standards for re-introduction

are density of wild lupine of 1.50 stems/m2, standard deviation of integrated light intensity of

16.82%, five ant species which tend Karner Blue butterfly larvae, first brood nectar source plant

density of 47.25 stems/m2, and second brood nectar source plant density of 47.85 stems/m2

(Chan and Packer, 2006).

Although a host of habitat features necessary for re-introduction have been clearly identified,

throughout the literature several habitat features have been found to be more important to the

success of the Karner Blue butterfly than others, especially at different stages during its life

cycle. At the larval stage the most limiting factor for the butterfly’s success is wild lupine

(Forrester et al., 2005; Pfitsch and Williams, 2009). The first and second brood of Karner Blue

butterfly adults also require wild lupine but as a nectar source instead of a food source, and are

less dependent due to the fact that they nectar on various other plant species both native and

exotic (Haack, 1993).

 28  

Despite the heavy reliance of the Karner Blue butterfly larvae on wild lupine there is one habitat

feature that is even more crucial to the survival of Karner Blue butterfly and that is optimal

canopy cover (Grundel et al., 1998). As demonstrated previously, shade heterogeneity in a

potential re-introduction habitat is important to ensure that the population has optimal conditions

for mating, oviposition, and larval growth (Grundel et al., 1998). Heterogeneous canopy cover

will also allow for proliferation of wild lupine (Grundel et al., 1998). While studies such as

Smallidge et al. (1996) have generally shown that lupine thrives in open areas, Grundel et al.

(1998) showed that while lupine density was highest in open canopy, larger plants (preferred by

larvae) were found in partially shaded areas. Thus, a mixture of shade and open canopy is also

important for ensuring wild lupine growth optimal for Karner Blue Butterfly larva survival.

Although it is clear that wild lupine plays an important role at the larval stage of the Karner Blue

butterfly’s life cycle, ensuring optimal canopy cover is necessary for both Karner Blue butterfly

and wild lupine success (Smallidge et al., 1996).

Habitat Assessment Prior to Re-introduction

Packer and Chan (2006) outlined methods for assessing habitat suitability prior to re-introduction

of the Karner Blue butterfly. Sites were visited during first and second flights of brood adults

(Packer and Chan, 2006). The sites were surveyed using the transect quadrat method used to

count herbaceous cover (transects were placed where wild lupine numbers were highest) (Packer

and Chan, 2006). Number of individuals, stems, and percent cover for each plant species were

recorded and the data from two visits were pooled (Packer and Chan, 2006). Mean number of

lupine stems and nectar plants per m2 were calculated for each site (Packer and Chan, 2006). Ant

species were collected and preserved in ethanol for identification (Packer and Chan, 2006). Light

intensity was measured using portable data loggers (Packer and Chan, 2006). Detailed habitat

assessment using similar methods to these, and other analyses as needed, is crucial to ensuring an

optimal re-introduction site is selected.

Restoration and Maintenance of Re-introduction Sites

As a large proportion of oak savanna habitat has already been degraded, it is likely that potential

sites selected for re-introduction will be found to be sub-optimal. Measures can be taken to

restore these sites to more suitable conditions. In a study by Kleintjes et al. (2003) they restored

 29  

an area for use as suitable habitat for the Karner Blue butterfly. The area was a sand prairie and

pine barren (similar to oak savanna) that had become forested due to fire suppression (Kleintjes

et al., 2003). They found an effective mixture of seeding and transplanting of wild lupine helped

in the restoration (Kleintjes et al., 2003). Transplanting was done in early April or after wild

lupine senescence (Kleintjes et al., 2003). The seed mixture was 40% grasses, 50% forbs

(including nectar plants), and 10% wild lupine and was applied by hand to the site in the fall at a

rate of 22.6lbs seed/hectare with a ratio of 1 part seed mix: 8 parts damp saw dust (Kleintjes et

al., 2003). The seed mix was composed of seeds harvested from plants within 120km of the

restoration site (Kleintjes et al., 2003). They found that tree thinning instead of establishment of

fire regimes was effective at maintaining the essential habitat characteristics of this re-

introduction area (Kleintjes et al., 2003). The thinning (25% tree cover) provided a mix of

openings and shaded areas for the Karner Blue butterfly (Kleintjes et al., 2003).

As demonstrated by the later steps taken by Kleintjes et al. (2003), areas where the Karner Blue

butterfly is re-introduced will need to be intensively managed in order to maintain the butterfly’s

specific habitat requirements and in order to ensure that environmental stochasticity does not

extirpate the population (U.S. Fish and Wildlife Service, 2003). Weather variability can affect

the timing of wild lupine senescence which in turn can affect the larval stage of the Karner Blue

butterfly (U.S. Fish and Wildlife Service, 2003). If annual weather negatively affects wild lupine

populations then they may senesce later, which could result in starvation of Karner Blue butterfly

larvae (U.S. Fish and Wildlife Service, 2003). Management strategies (ie. weather monitoring)

will need to be created in order to prevent this scenario from resulting in an extirpated or lowered

population.

In addition to weather monitoring, areas that Karner Blue butterflies are re-introduced to will

also require maintenance in order to prevent them from succeeding the early successional stages,

which Karner Blue butterflies prefer, and to maintain the proper wild lupine populations. There

are a few options for maintaining litter, vegetation, and canopy cover of these areas (King,

2003). Prescribed burns in November and July are cooler and do not disrupt key life stages of the

Karner Blue butterfly (King, 2003). Mowing is another option and can be done in the fall (late

August) and will not disrupt any key life stages of the Karner Blue butterfly as long as the

 30  

mowing height is above approximately 20 cm, which will prevent the majority of Karner Blue

butterfly eggs from being damaged (King, 2003).

If canopy cover becomes too high then trees on re-introduction sites can be cut down (Pfitsch

and Williams, 2009). As well, the removal of some trees can impact amounts of litter cover and

help to lower it to levels preferred by wild lupine (Pfitsch and Williams, 2009). In a study by

Pfitsch and Williams (2009) they found that removal of trees on pine barrens helped to increase

wild lupine cover through reducing pH and increasing sunlight. They later go on to suggest that a

mix of herbicides, mechanical cutting, and prescribed fires are the best management options

(Pfitsch and Williams, 2009).

Central Threats

The narrow habitat requirements of the Karner Blue butterfly, along with its large dependency on

the abundance of wild lupine and percentage of canopy cover, leave this already endangered

species vulnerable to a number of major threats. In order for successful long-term re-

introductions, threats to the Karner Blue butterfly must be considered. These threats include

habitat loss, urbanization, fire suppression, unmanaged agricultural and forestry practices,

increases in woody vegetation, and reductions in wild lupine abundance (Kleintjes et al, 2003).

As mentioned previously, Nice et al. (2009) noted a possible emerging threat to the Karner Blue

butterfly, which is an infection by endosymbiotic bacteria called Wolbachia. Action has been

taken on a small scale as described by Forrester et al. (2005) by carefully managing power-line

corridors, trails, and road systems of forests, airports, and military training areas which are

critical habitat for the Karner Blue butterfly (Smith et al., 2002, Warren et al., 2007). As

previously mentioned, prescribed fires can also be considered to combat habitat loss on a small

scale (Hunzinger, 2003). However, larger conservation efforts need to be made in order to

increase the populations of the Karner Blue butterfly to a satisfactory level. These efforts will

need to coincide with increased funding and research to provide a concrete basis on which to

enact large-scale conservation efforts.

 31  

8. Methods and Materials

The main method used to complete this report was conducting a thorough literature review of the

requirements needed for the Karner Blue butterfly to be successfully re-introduced to Ontario.

This literature review was completed using many peer reviewed journal articles, found through

Google Scholar, Scholars Portal, JSTOR, Wiley and ProQuest, the University of Guelph’s

library search engine.

While compiling this research a number of key words were used repeatedly in database searches.

Key words: Karner Blue butterfly, re-introduction, review, extirpated, endangered, wild

lupine, biology, description, mating, predators, disease, propagation, parasites, ants, larvae,

life cycle, threats, habitat requirements, and distribution.

Along with the peer reviewed articles other reports and articles written by reputable sources such

as the Ontario Ministry of Natural Resources were reviewed for this report. The various peer-

reviewed articles accessed covered a range of topics including information on the Karner Blue

butterfly, wild lupine, and related re-introduction strategies from other sources in North America.

9. Conclusion

This literature review is a compilation of research and information on the Karner Blue butterfly’s

biology, habitat, threats, and essential re-introduction considerations. The literature on Karner

Blue butterfly biology and habitat requirements cements the importance of wild lupine and oak

savanna habitat characteristics to successful re-introductions. In addition, this background

research provides sufficient information to develop new conclusions regarding additional re-

introduction requirements. Successful past re-introduction and rehabilitation of some Karner

Blue butterfly populations as well as other species gives some ideas as to where to start and what

methods can be successful.

Drawing from this comprehensive collection of information, the habitat characteristics necessary

for the successful re-introduction of the Karner Blue butterfly can be summarized as follows:

 32  

● Sands or well-drained soils that are frequently disturbed (Halpern, 2005; Pavlovic

and Grundel, 2009; Shi et al., 2005)

● Early successional sites (Chan and Packer, 2006)

● Wild lupine (1.50 stems/m2) (Chan and Packer, 2006)

● Nectar species (first brood 47.25 stems/m2 and second brood 47.85 stems/m2) (Chan

and Packer, 2006)

● Five tending ant species (showed to increase larvae survival) (Chan and Packer,

2006)

● Open/patchy canopy which is connected to facilitate movement (Huntzinger, 2003;

Halpern, 2005; Pavlovic and Grundel, 2009)

● Reduced cover/density of woody species (Halpern, 2005; Pavlovic and Grundel,

2009)

● Increased light intensity (standard deviation of integrated light intensity of 16.82%)

(Chan and Packer, 2006)

Potential limitations and weaknesses to this review may include missed research and relevant

information, and these concerns have been addressed as best as possible by making sure

thorough searches have been completed and all possible gaps have been filled. Where gaps in

available information have been encountered, recommendations have been made as to where

further research should take place. Recommendations for filling current gaps in knowledge can

be summarized as follows:

● Further investigation of the key predators and parasites of the Karner Blue butterfly, as

well as the extent of their predation and parasitism (Schellhorn et al., 2005)

● Understanding how significant an impact, relative to resource availability, predation is on

the survival of various life stages of this species

● Further investigation of the impact of disease organisms on Karner Blue butterfly

populations, especially Wolbachia (Nice et al., 2009; Haack, 1993)

 33  

This comprehensive literature review has been completed on the subject of the Karner Blue

butterfly and its re-introduction to ensure that all pertinent information on this species is in one

document. The review was created in an effort to facilitate re-introduction programs by

eliminating the added time of searching and consolidating multiple sources of information. It can

continue to be updated as new studies and papers are completed.

 34  

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