Regal Fritillary
Encyclopedia
The Regal Fritillary is a striking nymphalid
butterfly
found among some of the remaining tallgrass
and mixed-grass prairie
s in the east-central United States
. This prairie-specialist butterfly has a characteristic deep orange color and unmistakable dark hindwings with two bands of spots (Brock 2003). On the female, both bands of spots are white. However, on the male, the outer band of spots is orange in color. Females also tend to be slightly larger than males. The ventral surface of the hindwings is olive brown to black in color with bold silvery white spots (Selby 2007). The wingspan of S. idalia usually measures 68–105 mm (2.7–4.1 ) (Selby 2007). Flight is in the summertime from approximately June to September and adults tend to be swift in flight, coasting close to the ground (Brock 2003).
Regal fritillary larvae are approximately 0.08 inches long after they hatch and reach a length of approximately 1.75 inches when fully developed (Edwards 1879). The mature larvae have a black body with yellowish-orange bands and stripes. There are yellowish middorsal and lateral stripes and a number of dorsal, subdorsal, and lateral fleshy spines extending from the body. The head of the mature larvae is rounded and small, orangish-red on top and black underneath (Edwards 1879).
The larval food source for the regal fritillary and all members of the genus Speyeria
are violets (Viola spp.) (Selby 2007). The violets are an extremely important component of habitat sustainability for the regal fritillary and there is a correlation between the number of violets present and the number of butterflies found in a given area (Kelly and Debinski 1998). Violet species that the larvae feed on include Viola pedata
(bird’s-foot violet), V. pedatifida
(blue prairie violet), V. papilionacea (common blue violet), V. lanceolata (lance-leafed violet), V. nuttallii
(Nuttall’s Violet) (Kelly and Debinski 1998), V. sagittata (arrowleaf violet), and V. tricolor (Johnny Jumpup) (Selby 2007). These various violet species are associated with the different areas of the regal fritillary’s range. For example, the bird’s-foot violet and the prairie violet tend to be the preferred larval food source for the regal in the Midwest
and Great Plains
regions (Selby 2007).
The adult butterflies may feed on a variety of nectar plants and their availability throughout the summer flight time can be as important as the presence of larval food plants in determining whether an area can support populations of butterfly species (Selby 2007). Milkweeds
, thistle
s, coneflower
s, blazing-stars
, bergamots
, clover
s, goldenrod
s, and ironweeds
are some of the most important nectar sources for adult regal fritillaries. Milkweeds and thistles have been observed to be the preferred nectar source throughout the regal fritillary’s range (Selby 2007). These two types of plants provide a constant supply of nectar due to their staggered growth times. Common milkweed starts blooming when male regal fritillaries begin to emerge early in the summer and thistles tend to bloom later in the season which is crucial to females approaching oviposition
(Selby 2007).
, or a suspended period of development. The ovaries remain undeveloped during this time. Oogenesis does not initiate until late August when juvenile hormone sharply increases (Kopper et al. 2001). Once oogenesis takes place, the eggs are fertilized and soon after, oviposition occurs. The female lays over 1,000 eggs and possibly more than 2,000 (Vaughan and Shepherd, 2005). The small larvae hatch in late September and into October. Immediately after hatching, the tiny larvae seek protective covering in the leaf litter and overwinter there. At this stage the larvae delay development over the winter months and this is known as larval diapuase (Kopper et al. 2001). Once spring arrives, the larvae emerge and begin feeding on violets. They grow and mature through six instar stages until late May when they pupate (Selby 2007).
The life cycle
of the regal fritillary is quite unique and is thought to be an adaptation
to the phenology
, or seasonal timing and nature of their larval food plant, the violet (Kopper et al. 2001). These small perennial violets produce abundant foliage in the spring for the growing larvae. However, in most areas they senesce
in the heat of the summer and become unavailable to the larvae at that time. When this occurs, the regal fritillary is entering into its adult life and is no longer dependent on the violet. The fact that the violets remain unable to support larvae throughout the rest of the summer helps to explain the regal fritillary’s univoltine life cycle. It is through larval diapause coupled with the adult female reproductive diapuase that enables the larvae to maximize the benefits of fresh and abundant violet foliage when they are active in the spring (Kopper et al. 2001).
to Maine
. However, due to habitat loss and large-scale population declines, their range has been far reduced, especially in the east.
From the 1960s through the early 1990s, eastern populations had crashed so severely that only a few eastern populations remain (Powell et al. 2006). The regal fritillary was once present in 18 states east of Illinois. Today, only three populations remain in the eastern region. These populations have been located in Indiana, Pennsylvania, and Virginia. Populations in the Midwest and Great Plains are much more widespread; however, they remain extremely vulnerable as their numbers continue to decline. The regal fritillary is not federally listed as an endangered species; however, it has been assigned a NatureServe conservation status of G3 which is considered vulnerable (Selby 2007). S. idalia was a Category II species, or a possible candidate for listing under the Endangered Species Act
until 1996 when this category was eliminated by the federal government (Kelly and Debinski 1998).
(Powell et al. 2006). A number of factors are continually contributing to the loss, fragmentation, and degradation of the butterfly’s habitat. Row crop agriculture, urban developments such as housing and business construction, road construction, and gravel mining
, are all activities that contribute to the disappearance and degradation of the prairies that regal fritillaries depend on. Since regal fritillaries require relatively non-degraded native tallgrass and mixed-grass prairies, the alteration of these landscapes have pushed them into a highly vulnerable status (Selby 2007).
Largely due to the loss and degradation of the tallgrass prairie landscape, violet plant density tends to be limited or far reduced in certain areas. This has been shown to have a rather negative impact on regal fritillary populations. A study conducted by Kelly and Debinski (1998) looked at how larval food source limitations may be a factor in the declining regal fritillary populations. In the study, they correlated violet plant density to population size and weights of regal fritillary butterflies. It was found that prairies with significantly lower violet densities had smaller populations of S. idalia. Butterfly weights were also slightly lower in areas with low violet density (Kelly and Debinski 1998). As a result, a number of concerns have been raised regarding the health of the regal fritillary. Areas with few or no violets can be detrimental to female fecundity
because there are few suitable places for the eggs to be laid (Kelly and Debinski 1998). Also, smaller fragmented populations are susceptible to restricted gene flow
and reduced genetic variability
(Williams et al. 2003). This study also showed that habitat quality for the regal fritillary is just as important as the amount of habitat available. Increased violet density and nectar availability are essential to maintaining healthy populations (Kelly and Debinski 1998).
Habitat fragmentation
and isolation can have large scale genetic effects on high gene flow species such as the regal fritillary. There is an increased likelihood of population extirpation
among high gene flow species experiencing habitat fragmentation (Williams et al. 2003). Williams et al. (2003) conducted a study that compared levels of genetic differentiation and diversity among populations with a relatively continuous habitat to populations in isolated habitat areas. It was found that the isolated and highly fragmented populations had increased differentiation, or divergence from other populations, and decreased genetic diversity in comparison to non-fragmented populations. Restricted gene flow and population bottlenecks likely occur among populations in fragmented habitat areas, causing these phenomena (Williams et al. 2003). As habitat fragmentation continues to increase in much of the regal fritillary’s Midwestern range, genetic problems may become a real threat, disrupting gene flow and increasing the risk of disease.
Prescribed burning
is an attractive and widely used conservation tool among land managers
today. The historic role that fire played in the prairie landscape is important and can be highly beneficial to many plant species. Prescribed burns have also become a popular low-cost alternative for removing woody vegetation on rural and agricultural lands (Powell et al. 2006). However, there has been some evidence that heavy fire management used on prairie lands can negatively affect the regal fritillary. A study conducted by Powell et al. (2006) examined the effects of prescribed prairie burns by surveying a number of prairies in Kansas. Butterflies were observed on both recently burned and unburned sites to determine the effects of prescribed burning on the populations. Population density of the regal fritillary may tend to vary among sites but was generally quite a bit higher at sites that had not been burned in the past year (Powell et al. 2006). Prescribed burns are usually conducted in the early spring while the first instar larvae remain vulnerable, buried in the leaf litter. Extensive prairie burns kill the overwintering larvae and can have a drastic effect on their population in the following years. One recently burned prairie that was studied used minimal patch burning by burning only small portions of the entire area at a time. This site had by far the highest regal fritillary abundance of any burned site (Powell et al. 2006).
A serious potential threat to the regal fritillary was discovered in a captive breeding study. Wagner (1995) found that disease is a possible mortality factor in some S. idalia populations. In a captive group, nuclear polyhedrosis virus
(NPV) caused an 80% loss. The virus is transmitted from females to offspring in eggs or between individuals through excreta (Wagner 1995). NPV could potentially be damaging to populations in the wild (Mason 2007), thus for reintroduction purposes, culturing a virus-free line is critical.
The regal fritillary is highly vulnerable to environmental factors year-round. Extreme weather
conditions over a large geographical range can severely influence their populations. First instar
larvae are highly sensitive to extreme weather conditions as they overwinter in the leaf litter and as they begin their initial search for food plants in the spring. Hard frosts late in the spring, severe storms, and cool damp conditions have all been shown to negatively impact larvae survival (Selby 2007). Larval development rates tend to be proportional to the temperature. Therefore, unusually cool conditions in the spring can drastically slow larval growth rates, increasing their exposure to mortality factors (Selby 2007). Some environmental factors can limit adult regal fritillary activity as well. Prolonged periods of cooler temperatures, cloudy skies, and rain can encumber normal activities, perhaps limiting reproduction (Selby 2007).
The increased use of pesticide
s and herbicide
s can have profound negative effects on the regal fritillary as well. Heavy spraying of herbicides can eliminate nearby larval food plants and nectar sources that they are dependent on (Selby 2007). The indiscriminant use of pesticides also poses a big threat to regal fritillaries and other prairie-specialist butterflies. Pesticides such as the bacterial pathogen Bacillus thuringiensis
("Bt", the agent used in gypsy moth
control) is lethal to all Lepidoptera larvae. It is thought that the gypsy moth control programs used in the east along wooded grassland edges may have been a final factor leading to the loss of some populations (Selby 2007). Broadcast spraying of insecticides for pest control on adjacent crop land and rangeland continues to be a direct threat to the regal fritillary.
The Midwest landscape includes few prairie remnants which are embedded in an agricultural matrix (Davis et al. 2007). Thus, it is extremely important that the land surrounding prairie remnants be included in management decisions and practices. The quality of the matrix surrounding a particular habitat fragment often makes a difference on a species’ dispersal ability (Davis et al. 2007). Edge effects must be considered and managed for to improve the butterfly’s dispersal capabilities. Connectivity between regal fritillary habitats is also extremely important to consider in land management practices and would likely increase gene flow and genetic diversity in certain areas. This would help to reduce some of the negative effects associated with habitat fragmentation, increasing the overall health of regal fritillary populations.
Though it was found that extensive fire management can have direct negative impacts on regal fritillary populations, it is important to remember that prescribed burns can be beneficial to many plant species, including violets and nectar plants (Selby 2007). This could in turn provide some increased habitat benefits for the butterfly if prescribed burns are used appropriately. Fire management also helps to remove cool season exotics and woody vegetation that encroach on native prairie plants such as the violet. Thus it is important to understand both the positive and negative effects of fire management and the combined effects that it has on the long-term survival of the regal fritillary (Selby 2007). Management should account for this and it is recommended that only small portions, no more than 20% of the total butterfly habitat, be burned in a given year. It has also been suggested that 3 to 5 year burn rotations be used where a certain plot of land must remain unburned for at least 3 to 5 years before it can be burned again. These practices would likely minimize the negative effects that fire management tends to have on regal fritillary populations, while likely providing them with some increased benefits due to higher quality food and nectar resources. It has also been suggested that very light grazing is also beneficial to these prairie specialist butterflies and can effectively be used in combination with limited prescribed burning (Selby 2007).
A limited use of herbicides and pesticides is fine; however, it should be heavily monitored and carefully applied where the regal fritillary is concerned (Selby 2007). Increasing the awareness among surrounding agricultural areas regarding the use of herbicides and pesticides is critical. Decreasing the use of herbicides and pesticides on agricultural lands directly adjacent to prairie habitats may be quite beneficial to regal fritillary populations as well as to many other native prairie insects. It is best to avoid or limit the widespread and indiscriminant use of these control agents in these areas (Selby 2007). Prairies that manage for invasive plants and woody vegetation by applying herbicides should do so with extreme caution and moderation. The use of selective applications such as spot spraying is better as well as using non-persistent herbicides. Reseeding may also be necessary after some herbicide applications so that beneficial native species remain plentiful. However, the use of herbicide on prairie lands should be a last resort to removing unwanted vegetation in order to protect regal fritillary populations (Selby 2007).
Since there is limited knowledge about the exact distribution and abundance of many regal fritillary populations, it is crucial that inventory and monitoring practices are carried out. Through a better understanding of the regal fritillary, it is possible to further conservation efforts in the right direction. Accurately surveying populations is crucial to monitoring practices and provides a great deal of useful information about specific populations. Pollard transect surveys obtain results of relative abundance values which can be used to track trend in abundance over time (Selby 2007). This method is better for long term monitoring of populations and can be quite useful to tracking the status of the regal fritillary over time.
Nymphalidae
The Nymphalidae is a family of about 5,000 species of butterflies which are distributed throughout most of the world. These are usually medium sized to large butterflies. Most species have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called...
butterfly
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...
found among some of the remaining tallgrass
Tallgrass prairie
The tallgrass prairie is an ecosystem native to central North America, with fire as its primary periodic disturbance. In the past, tallgrass prairies covered a large portion of the American Midwest, just east of the Great Plains, and portions of the Canadian Prairies. They flourished in areas with...
and mixed-grass prairie
Prairie
Prairies are considered part of the temperate grasslands, savannas, and shrublands biome by ecologists, based on similar temperate climates, moderate rainfall, and grasses, herbs, and shrubs, rather than trees, as the dominant vegetation type...
s in the east-central United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...
. This prairie-specialist butterfly has a characteristic deep orange color and unmistakable dark hindwings with two bands of spots (Brock 2003). On the female, both bands of spots are white. However, on the male, the outer band of spots is orange in color. Females also tend to be slightly larger than males. The ventral surface of the hindwings is olive brown to black in color with bold silvery white spots (Selby 2007). The wingspan of S. idalia usually measures 68–105 mm (2.7–4.1 ) (Selby 2007). Flight is in the summertime from approximately June to September and adults tend to be swift in flight, coasting close to the ground (Brock 2003).
Regal fritillary larvae are approximately 0.08 inches long after they hatch and reach a length of approximately 1.75 inches when fully developed (Edwards 1879). The mature larvae have a black body with yellowish-orange bands and stripes. There are yellowish middorsal and lateral stripes and a number of dorsal, subdorsal, and lateral fleshy spines extending from the body. The head of the mature larvae is rounded and small, orangish-red on top and black underneath (Edwards 1879).
The larval food source for the regal fritillary and all members of the genus Speyeria
Speyeria
Speyeria, commonly known as greater fritillaries, is the genus of butterflies in the family Nymphalidae commonly found in North America and Asia...
are violets (Viola spp.) (Selby 2007). The violets are an extremely important component of habitat sustainability for the regal fritillary and there is a correlation between the number of violets present and the number of butterflies found in a given area (Kelly and Debinski 1998). Violet species that the larvae feed on include Viola pedata
Viola pedata
Viola pedata is an ornamental plant in the Violaceae family. Monophyletic in stature, this violet, also known as the "mountain pansy" is endemic to eastern North America, where it favors well drained, acidic soils in full to partial sun environments...
(bird’s-foot violet), V. pedatifida
Viola pedatifida
Viola pedatifida Viola pedatifida Viola pedatifida (Prairie violet, Crow-foot violet, larkspur violet, purple prairie violet, coastal violet; syn. Viola pedatifida subsp. brittoniana (Pollard) L. E. McKinney, Viola pedatifida G. Don subsp. pedatifida, Viola palmata L. var...
(blue prairie violet), V. papilionacea (common blue violet), V. lanceolata (lance-leafed violet), V. nuttallii
Viola nuttallii
Viola nuttallii is a perennial herbaceous plant in the Violet family , and is one of the few violet species with lanceolate leaves...
(Nuttall’s Violet) (Kelly and Debinski 1998), V. sagittata (arrowleaf violet), and V. tricolor (Johnny Jumpup) (Selby 2007). These various violet species are associated with the different areas of the regal fritillary’s range. For example, the bird’s-foot violet and the prairie violet tend to be the preferred larval food source for the regal in the Midwest
Midwestern United States
The Midwestern United States is one of the four U.S. geographic regions defined by the United States Census Bureau, providing an official definition of the American Midwest....
and Great Plains
Great Plains
The Great Plains are a broad expanse of flat land, much of it covered in prairie, steppe and grassland, which lies west of the Mississippi River and east of the Rocky Mountains in the United States and Canada. This area covers parts of the U.S...
regions (Selby 2007).
The adult butterflies may feed on a variety of nectar plants and their availability throughout the summer flight time can be as important as the presence of larval food plants in determining whether an area can support populations of butterfly species (Selby 2007). Milkweeds
Asclepias
Asclepias L. , the milkweeds, is a genus of herbaceous perennial, dicotyledonous plants that contains over 140 known species...
, thistle
Thistle
Thistle is the common name of a group of flowering plants characterised by leaves with sharp prickles on the margins, mostly in the family Asteraceae. Prickles often occur all over the plant – on surfaces such as those of the stem and flat parts of leaves. These are an adaptation that protects the...
s, coneflower
Coneflower
Coneflower is a common name of at least four genera of flowering plants in the family Asteraceae:*Dracopis*Echinacea*Rudbeckia*Ratibida...
s, blazing-stars
Liatris
Liatris is a genus of ornamental plants in the Asteraceae family, native to North America, Mexico, and the Bahamas. These plants are used as a popular summer flowers for bouquets.They are perennials, surviving the winter in the form of corms....
, bergamots
Monarda didyma
Monarda didyma is an aromatic herb in the family Lamiaceae, native to eastern North America from Maine west to Ohio and south to northern Georgia. Its name is derived from its odor, which is considered similar to that of the bergamot orange...
, clover
Clover
Clover , or trefoil, is a genus of about 300 species of plants in the leguminous pea family Fabaceae. The genus has a cosmopolitan distribution; the highest diversity is found in the temperate Northern Hemisphere, but many species also occur in South America and Africa, including at high altitudes...
s, goldenrod
Goldenrod
Solidago, commonly called goldenrods, is a genus of about 100 species of flowering plants in the family Asteraceae. Most are herbaceous perennial species found in the meadows and pastures, along roads, ditches and waste areas in North America. There are also a few species native to Mexico, South...
s, and ironweeds
Vernonia
Vernonia is a genus of about 1000 species of forbs and shrubs in the family Asteraceae. Some species are known as Ironweed. Some species are edible and of economic value. They are known for having intense purple flowers. The genus is named for English botanist William Vernon. There are numerous...
are some of the most important nectar sources for adult regal fritillaries. Milkweeds and thistles have been observed to be the preferred nectar source throughout the regal fritillary’s range (Selby 2007). These two types of plants provide a constant supply of nectar due to their staggered growth times. Common milkweed starts blooming when male regal fritillaries begin to emerge early in the summer and thistles tend to bloom later in the season which is crucial to females approaching oviposition
Oviposition
Oviposition is the process of laying eggs by oviparous animals.Some arthropods, for example, lay their eggs with an organ called the ovipositor.Fish , amphibians, reptiles, birds and monetremata also lay eggs....
(Selby 2007).
Reproduction and life cycle
The regal fritillary is univoltine, having a single generation per year (Selby 2007). Adult male butterflies emerge in early June along with the first milkweed plants. Adult females emerge shortly after and mating takes place in late June and early July. After mating, females enter a 6 to 8 week period of reproductive diapauseDiapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...
, or a suspended period of development. The ovaries remain undeveloped during this time. Oogenesis does not initiate until late August when juvenile hormone sharply increases (Kopper et al. 2001). Once oogenesis takes place, the eggs are fertilized and soon after, oviposition occurs. The female lays over 1,000 eggs and possibly more than 2,000 (Vaughan and Shepherd, 2005). The small larvae hatch in late September and into October. Immediately after hatching, the tiny larvae seek protective covering in the leaf litter and overwinter there. At this stage the larvae delay development over the winter months and this is known as larval diapuase (Kopper et al. 2001). Once spring arrives, the larvae emerge and begin feeding on violets. They grow and mature through six instar stages until late May when they pupate (Selby 2007).
The life cycle
Biological life cycle
A life cycle is a period involving all different generations of a species succeeding each other through means of reproduction, whether through asexual reproduction or sexual reproduction...
of the regal fritillary is quite unique and is thought to be an adaptation
Adaptation
An adaptation in biology is a trait with a current functional role in the life history of an organism that is maintained and evolved by means of natural selection. An adaptation refers to both the current state of being adapted and to the dynamic evolutionary process that leads to the adaptation....
to the phenology
Phenology
Phenology is the study of periodic plant and animal life cycle events and how these are influenced by seasonal and interannual variations in climate...
, or seasonal timing and nature of their larval food plant, the violet (Kopper et al. 2001). These small perennial violets produce abundant foliage in the spring for the growing larvae. However, in most areas they senesce
Senescence
Senescence or biological aging is the change in the biology of an organism as it ages after its maturity. Such changes range from those affecting its cells and their function to those affecting the whole organism...
in the heat of the summer and become unavailable to the larvae at that time. When this occurs, the regal fritillary is entering into its adult life and is no longer dependent on the violet. The fact that the violets remain unable to support larvae throughout the rest of the summer helps to explain the regal fritillary’s univoltine life cycle. It is through larval diapause coupled with the adult female reproductive diapuase that enables the larvae to maximize the benefits of fresh and abundant violet foliage when they are active in the spring (Kopper et al. 2001).
Declining populations
With a loss of more than 99% of the original native tallgrass prairie landcover today (Powell et al. 2006), decreased sustainable habitat area for the regal fritillary has become a real threat. Drastic declines in regal fritillary populations have led to much concern about the butterfly’s future (Kopper et al. 2001). Historically, the regal fritillary’s range extended from eastern ColoradoColorado
Colorado is a U.S. state that encompasses much of the Rocky Mountains as well as the northeastern portion of the Colorado Plateau and the western edge of the Great Plains...
to Maine
Maine
Maine is a state in the New England region of the northeastern United States, bordered by the Atlantic Ocean to the east and south, New Hampshire to the west, and the Canadian provinces of Quebec to the northwest and New Brunswick to the northeast. Maine is both the northernmost and easternmost...
. However, due to habitat loss and large-scale population declines, their range has been far reduced, especially in the east.
From the 1960s through the early 1990s, eastern populations had crashed so severely that only a few eastern populations remain (Powell et al. 2006). The regal fritillary was once present in 18 states east of Illinois. Today, only three populations remain in the eastern region. These populations have been located in Indiana, Pennsylvania, and Virginia. Populations in the Midwest and Great Plains are much more widespread; however, they remain extremely vulnerable as their numbers continue to decline. The regal fritillary is not federally listed as an endangered species; however, it has been assigned a NatureServe conservation status of G3 which is considered vulnerable (Selby 2007). S. idalia was a Category II species, or a possible candidate for listing under the Endangered Species Act
Endangered Species Act
The Endangered Species Act of 1973 is one of the dozens of United States environmental laws passed in the 1970s. Signed into law by President Richard Nixon on December 28, 1973, it was designed to protect critically imperiled species from extinction as a "consequence of economic growth and...
until 1996 when this category was eliminated by the federal government (Kelly and Debinski 1998).
Threats
The greatest threat the regal fritillary faces is habitat destructionHabitat destruction
Habitat destruction is the process in which natural habitat is rendered functionally unable to support the species present. In this process, the organisms that previously used the site are displaced or destroyed, reducing biodiversity. Habitat destruction by human activity mainly for the purpose of...
(Powell et al. 2006). A number of factors are continually contributing to the loss, fragmentation, and degradation of the butterfly’s habitat. Row crop agriculture, urban developments such as housing and business construction, road construction, and gravel mining
Gravel pit
Gravel pit is the term for an open cast working for extraction of gravel. Gravel pits often lie in river valleys where the water table is high, so they may fill naturally with water to form ponds or lakes. Old, abandoned gravel pits are normally used either as nature reserves, or as amenity areas...
, are all activities that contribute to the disappearance and degradation of the prairies that regal fritillaries depend on. Since regal fritillaries require relatively non-degraded native tallgrass and mixed-grass prairies, the alteration of these landscapes have pushed them into a highly vulnerable status (Selby 2007).
Largely due to the loss and degradation of the tallgrass prairie landscape, violet plant density tends to be limited or far reduced in certain areas. This has been shown to have a rather negative impact on regal fritillary populations. A study conducted by Kelly and Debinski (1998) looked at how larval food source limitations may be a factor in the declining regal fritillary populations. In the study, they correlated violet plant density to population size and weights of regal fritillary butterflies. It was found that prairies with significantly lower violet densities had smaller populations of S. idalia. Butterfly weights were also slightly lower in areas with low violet density (Kelly and Debinski 1998). As a result, a number of concerns have been raised regarding the health of the regal fritillary. Areas with few or no violets can be detrimental to female fecundity
Fecundity
Fecundity, derived from the word fecund, generally refers to the ability to reproduce. In demography, fecundity is the potential reproductive capacity of an individual or population. In biology, the definition is more equivalent to fertility, or the actual reproductive rate of an organism or...
because there are few suitable places for the eggs to be laid (Kelly and Debinski 1998). Also, smaller fragmented populations are susceptible to restricted gene flow
Gene flow
In population genetics, gene flow is the transfer of alleles of genes from one population to another.Migration into or out of a population may be responsible for a marked change in allele frequencies...
and reduced genetic variability
Genetic variability
Genetic variability is a measure of the tendency of individual genotypes in a population to vary from one another. Variability is different from genetic diversity, which is the amount of variation seen in a particular population. The variability of a trait describes how much that trait tends to...
(Williams et al. 2003). This study also showed that habitat quality for the regal fritillary is just as important as the amount of habitat available. Increased violet density and nectar availability are essential to maintaining healthy populations (Kelly and Debinski 1998).
Habitat fragmentation
Habitat fragmentation
Habitat fragmentation as the name implies, describes the emergence of discontinuities in an organism's preferred environment , causing population fragmentation...
and isolation can have large scale genetic effects on high gene flow species such as the regal fritillary. There is an increased likelihood of population extirpation
Local extinction
Local extinction, also known as extirpation, is the condition of a species which ceases to exist in the chosen geographic area of study, though it still exists elsewhere...
among high gene flow species experiencing habitat fragmentation (Williams et al. 2003). Williams et al. (2003) conducted a study that compared levels of genetic differentiation and diversity among populations with a relatively continuous habitat to populations in isolated habitat areas. It was found that the isolated and highly fragmented populations had increased differentiation, or divergence from other populations, and decreased genetic diversity in comparison to non-fragmented populations. Restricted gene flow and population bottlenecks likely occur among populations in fragmented habitat areas, causing these phenomena (Williams et al. 2003). As habitat fragmentation continues to increase in much of the regal fritillary’s Midwestern range, genetic problems may become a real threat, disrupting gene flow and increasing the risk of disease.
Prescribed burning
Controlled burn
Controlled or prescribed burning, also known as hazard reduction burning or Swailing is a technique sometimes used in forest management, farming, prairie restoration or greenhouse gas abatement. Fire is a natural part of both forest and grassland ecology and controlled fire can be a tool for...
is an attractive and widely used conservation tool among land managers
Land management
Land management is the process of managing the use and development of land resources. Land resources are used for a variety of purposes which may include organic agriculture, reforestation, water resource management and eco-tourism projects.-See also:*Sustainable land management*Acreage...
today. The historic role that fire played in the prairie landscape is important and can be highly beneficial to many plant species. Prescribed burns have also become a popular low-cost alternative for removing woody vegetation on rural and agricultural lands (Powell et al. 2006). However, there has been some evidence that heavy fire management used on prairie lands can negatively affect the regal fritillary. A study conducted by Powell et al. (2006) examined the effects of prescribed prairie burns by surveying a number of prairies in Kansas. Butterflies were observed on both recently burned and unburned sites to determine the effects of prescribed burning on the populations. Population density of the regal fritillary may tend to vary among sites but was generally quite a bit higher at sites that had not been burned in the past year (Powell et al. 2006). Prescribed burns are usually conducted in the early spring while the first instar larvae remain vulnerable, buried in the leaf litter. Extensive prairie burns kill the overwintering larvae and can have a drastic effect on their population in the following years. One recently burned prairie that was studied used minimal patch burning by burning only small portions of the entire area at a time. This site had by far the highest regal fritillary abundance of any burned site (Powell et al. 2006).
A serious potential threat to the regal fritillary was discovered in a captive breeding study. Wagner (1995) found that disease is a possible mortality factor in some S. idalia populations. In a captive group, nuclear polyhedrosis virus
Nuclear Polyhedrosis Virus
The nuclear polyhedrosis virus which belongs to the sub group Baculoviruses is a virus affecting insects, predominantly moths and butterflies. It has been used as a pesticide for crops infested by insects susceptible to contraction...
(NPV) caused an 80% loss. The virus is transmitted from females to offspring in eggs or between individuals through excreta (Wagner 1995). NPV could potentially be damaging to populations in the wild (Mason 2007), thus for reintroduction purposes, culturing a virus-free line is critical.
The regal fritillary is highly vulnerable to environmental factors year-round. Extreme weather
Extreme weather
Extreme weather includes weather phenomena that are at the extremes of the historical distribution, especially severe or unseasonal weather. The most commonly used definition of extreme weather is based on an event's climatological distribution. Extreme weather occurs only 5% or less of the time...
conditions over a large geographical range can severely influence their populations. First instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...
larvae are highly sensitive to extreme weather conditions as they overwinter in the leaf litter and as they begin their initial search for food plants in the spring. Hard frosts late in the spring, severe storms, and cool damp conditions have all been shown to negatively impact larvae survival (Selby 2007). Larval development rates tend to be proportional to the temperature. Therefore, unusually cool conditions in the spring can drastically slow larval growth rates, increasing their exposure to mortality factors (Selby 2007). Some environmental factors can limit adult regal fritillary activity as well. Prolonged periods of cooler temperatures, cloudy skies, and rain can encumber normal activities, perhaps limiting reproduction (Selby 2007).
The increased use of pesticide
Pesticide
Pesticides are substances or mixture of substances intended for preventing, destroying, repelling or mitigating any pest.A pesticide may be a chemical unicycle, biological agent , antimicrobial, disinfectant or device used against any pest...
s and herbicide
Herbicide
Herbicides, also commonly known as weedkillers, are pesticides used to kill unwanted plants. Selective herbicides kill specific targets while leaving the desired crop relatively unharmed. Some of these act by interfering with the growth of the weed and are often synthetic "imitations" of plant...
s can have profound negative effects on the regal fritillary as well. Heavy spraying of herbicides can eliminate nearby larval food plants and nectar sources that they are dependent on (Selby 2007). The indiscriminant use of pesticides also poses a big threat to regal fritillaries and other prairie-specialist butterflies. Pesticides such as the bacterial pathogen Bacillus thuringiensis
Bacillus thuringiensis
Bacillus thuringiensis is a Gram-positive, soil-dwelling bacterium, commonly used as a biological pesticide; alternatively, the Cry toxin may be extracted and used as a pesticide. B...
("Bt", the agent used in gypsy moth
Gypsy moth
The gypsy moth, Lymantria dispar, is a moth in the family Lymantriidae of Eurasian origin. Originally ranging from Europe to Asia, it was introduced to North America in the late 1860s and has been expanding its range ever since...
control) is lethal to all Lepidoptera larvae. It is thought that the gypsy moth control programs used in the east along wooded grassland edges may have been a final factor leading to the loss of some populations (Selby 2007). Broadcast spraying of insecticides for pest control on adjacent crop land and rangeland continues to be a direct threat to the regal fritillary.
Conservation
Future losses among regal fritillary populations can be prevented by identifying critical habitat areas and managing them to maintain and improve habitat size, quality, and connectivity (Selby 2007). It is essential that land is set aside and protected in this manner. Land management practices should focus on the maintenance of intact native prairie remnants and the vegetation that is crucial to the regal fritillary’s continued survival. Managing for abundant larval food plants and nectar resources is extremely important. The timing, intensity, level, and duration of management activities must be adapted and monitored (Selby 2007). It has been suggested that proper management practices could play a crucial role in slowing, and possibly even reversing, the current wave of regal fritillary extirpations (Swengel 2004).The Midwest landscape includes few prairie remnants which are embedded in an agricultural matrix (Davis et al. 2007). Thus, it is extremely important that the land surrounding prairie remnants be included in management decisions and practices. The quality of the matrix surrounding a particular habitat fragment often makes a difference on a species’ dispersal ability (Davis et al. 2007). Edge effects must be considered and managed for to improve the butterfly’s dispersal capabilities. Connectivity between regal fritillary habitats is also extremely important to consider in land management practices and would likely increase gene flow and genetic diversity in certain areas. This would help to reduce some of the negative effects associated with habitat fragmentation, increasing the overall health of regal fritillary populations.
Though it was found that extensive fire management can have direct negative impacts on regal fritillary populations, it is important to remember that prescribed burns can be beneficial to many plant species, including violets and nectar plants (Selby 2007). This could in turn provide some increased habitat benefits for the butterfly if prescribed burns are used appropriately. Fire management also helps to remove cool season exotics and woody vegetation that encroach on native prairie plants such as the violet. Thus it is important to understand both the positive and negative effects of fire management and the combined effects that it has on the long-term survival of the regal fritillary (Selby 2007). Management should account for this and it is recommended that only small portions, no more than 20% of the total butterfly habitat, be burned in a given year. It has also been suggested that 3 to 5 year burn rotations be used where a certain plot of land must remain unburned for at least 3 to 5 years before it can be burned again. These practices would likely minimize the negative effects that fire management tends to have on regal fritillary populations, while likely providing them with some increased benefits due to higher quality food and nectar resources. It has also been suggested that very light grazing is also beneficial to these prairie specialist butterflies and can effectively be used in combination with limited prescribed burning (Selby 2007).
A limited use of herbicides and pesticides is fine; however, it should be heavily monitored and carefully applied where the regal fritillary is concerned (Selby 2007). Increasing the awareness among surrounding agricultural areas regarding the use of herbicides and pesticides is critical. Decreasing the use of herbicides and pesticides on agricultural lands directly adjacent to prairie habitats may be quite beneficial to regal fritillary populations as well as to many other native prairie insects. It is best to avoid or limit the widespread and indiscriminant use of these control agents in these areas (Selby 2007). Prairies that manage for invasive plants and woody vegetation by applying herbicides should do so with extreme caution and moderation. The use of selective applications such as spot spraying is better as well as using non-persistent herbicides. Reseeding may also be necessary after some herbicide applications so that beneficial native species remain plentiful. However, the use of herbicide on prairie lands should be a last resort to removing unwanted vegetation in order to protect regal fritillary populations (Selby 2007).
Since there is limited knowledge about the exact distribution and abundance of many regal fritillary populations, it is crucial that inventory and monitoring practices are carried out. Through a better understanding of the regal fritillary, it is possible to further conservation efforts in the right direction. Accurately surveying populations is crucial to monitoring practices and provides a great deal of useful information about specific populations. Pollard transect surveys obtain results of relative abundance values which can be used to track trend in abundance over time (Selby 2007). This method is better for long term monitoring of populations and can be quite useful to tracking the status of the regal fritillary over time.