The Use of Fire Management to Restore Monarch Butterfly Populations - Va Vang Environmental Studies Department CSU, Sacramento ENVS 190 - Senior ...
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The Use of Fire Management to Restore Monarch Butterfly Populations Va Vang Environmental Studies Department CSU, Sacramento ENVS 190 – Senior Thesis May 17, 2018 1
Table of Contents Page # I. Abstract 3 II. Biology 3-5 III. Distribution & Generational Migratory Populations 5-8 a. Monarch Generations i. Generational Migration Routes IV. Milkweeds 8-12 a. Ethnobotany – Humans and Milkweeds b. Common Asclepias Species V. Status & Decline of Milkweed & Monarchs 12-15 a. Wildlife Friendly Agricultural Practices VI. Prescribe Fire Management 15-17 VII. Propagating Milkweeds 17-18 a. From Root Cuttings b. From Seeds VIII. Restoration: Fire on Milkweeds in Prairies 19-21 IX. Other Restoration Efforts/Ideas to Consider 21-23 a. Agricultural Landscapes – data required for restoration X. Conclusion/Discussion 23-24 XI. Figures & Tables 8, 25-32 XII. References 33-40 2
Abstract The interaction between monarch butterflies (Danaus plexippus) and its host plant, milkweed (Asclepias) species, are extraordinary during their migration route from Central Mexico, to United States, onto Canada and then back down to Mexico. The purpose of this thesis is to explore the root behind declining eastern monarch populations in relations to milkweed species native to those states. Prairie land conversions and herbicides from agricultural fields have been the two-main cause of milkweed habitat loss. Conservation research has shown the benefits of using prescribe fire on these ecosystems to restore milkweeds to support and increase monarch populations. Biology Monarch butterflies (Danaus plexippus) captivate many people because of their phenomenal migration from Mexico, to the United States, and onto Canada. They then migrate back to Mexico to overwinter at the end of the year. It is astonishing for an insect with such a small mass to fly up to 6,000 miles from their two-way migration (Oberhauser & Solensky, 2004). They also contribute as essential flower pollinators for plant ecosystems (Hopwood, 2013). As pollinators, monarchs play a significant role. Impacts on their population will signify similar impacts on plants that they are pollinators for. If the impacts negatively affect the plants that monarchs pollinate, then many seeds cannot be dispersed or reproduce. Monarch butterflies undergo four stages of metamorphosis: the egg, larva, pupa, and adult stages. They are known as the “milkweed (Asclepias species) butterfly” since they have a symbiotic relationship with this forb. Milkweeds serve as a significant host plant for monarch larvae during reproduction and breeding events during their migration. 3
Females oviposit eggs underneath milkweed leaves that are tender and healthy with only 1 egg per plant (Urquhart, 1960; Pitman et al., 2018). Eggs are about the size of the head of a pin and have a yellow base when laid. The color changes to grey as the embryo develops. The growth of the embryo differs with changes in temperature. The development rate occurs more rapidly during higher temperatures, compared to lower temperatures (Urquhart, 1960) Larva, or caterpillar, hatch once the egg becomes grey. This stage consists of 5 instar stages. Instars are the developmental phases of larva leading the final physical caterpillar form with high pigmentation of the black, white, and yellow color with fully grown antenna (Urquhart, 1960). Once the larva emerges, they consume their vacant shell and depend on milkweed leaves as food resources. This plant is their main and only source of nourishment up until the next phase. This process lasts about 2 weeks before the pupa stage (Borders et al., 2013). Larvae tends to remain on the underside of milkweed leaves even during feeding activities. They consume only tender parts of the leaves and known to finish one leaf in about four minutes, which lasts them for a maximum of 2 hours. Just like the growth of the embryo during the first stage, temperature also plays a crucial role in the rate of growth. If their host plant is expose to high temperatures, larvae will be more active and gradually grow at a rapid rate. There are very few larvae that are found in shaded areas. Monarch adults chooses wisely when laying their eggs onto milkweed leaves and aims for the plants that are unsheltered from sunlight (Urquhart, 1960). The pupa, or chrysalis, phase is the final formation of larvae into adult butterflies. Their physical appearance is orange with black vein markings once they are free from the chrysalis. Finding the perfect area for metamorphosis is significant. Larvae will leave behind its original 4
host plant when matured and travel to other milkweed plants (rare) or found on logs, tree limbs, window-sills, timbers, and fences to morph. They start out by spinning silk fibers to create a mat which will then cover the larva. The larva will hang upside down by its prolegs. This stage lasts between 10-14 days until the adult butterfly emerges (Urquhart, 1960). The lifespan of each monarch lasts between 2-6 weeks after morphing into a butterfly. Distribution & Generational Migratory Populations Monarch butterflies is dispersed in Central Mexico, United States, and Canada. Distribution of the species vary because of multi-generational populations and from breeding to non-breeding grounds (Webster et al., 2002). This proposed evaluation will focus on the eastern monarch population that are from Central Mexico migrating to the U.S The eastern U.S. population include the monarchs from Central Mexico traveling to the southern, central, and northeast of the U.S (figure 1). They breed east of the Rocky Mountains. This population is the bigger one of the two, western and eastern U.S. Concern is higher here since majority of the migratory populations reside in those states. Monarchs abandon their breeding territories during the end of summer and start migrating south towards Central Mexico to escape freezing winter because they prefer high temperatures (Urquhart, 1960). This begins their spectacular journey from Canada, 3,000 miles to Mexico. Butterfly’s migratory cycle is usually linked to the locations that they settle in during their journey. Interactions over spatial movement and time are factors that can show the influence of location (Webster et al., 2002; Flockhart et al., 2013). For instance, monarchs may come into contact with surrounding species in their selected location. These species can either be predators or rivals competing for resources. Monarch butterflies feed on nectar from a variety of plants. Pollinators, like bees, are also interested in nectar resources. Therefore, they both will compete 5
for plant species with nectar. However, monarchs rely heavily on monarch patches to breed. The best areas are the one with high abundance of milkweed and plant species with nectar. Monarch Generations There are four monarch generations each year contributing to the stages of this migration. Not every generation migrate from Mexico to Canada and then back down. Their lifespans are short and certain generations migrate only to certain states before dying. The population that travels back to overwintering sites in Central Mexico are the offspring (great-great-great grandchildren) of the monarchs that regularly overwinter there from the year before. They are the fourth-generation population from the previous year that will start the first-generation population for the new year in Southern U.S. They are referred to as parents of the first-generation monarchs. During the first part of migration, the parents from Mexico travels to Texas, Florida, Arkansas, Missouri, Oklahoma, and Virginia where eggs are laid (Flockhart et al., 2013; Inamine, 2016) and breeding begins for the first generation (figure 2). These areas are part of their migratory route migrating towards the Midwestern part of U.S. right when they morph into adults (figure 3). The start of this migration and breeding begins at the end of March until late April (Inamine, 2016). The adult monarch of this generation dies after their eggs are laid and the next generation begins from these southern U.S. states. The second-generation monarchs are known as the grandchildren of the overwintering population from Mexico (generation 4 from the year before). Their breeding process starts from April until June and larvae are found in the Midwest and eastern U.S (figure 4). Adults from this generation emerges in the middle of June to July and begin laying eggs right away on milkweeds. They can be found in similar areas where they reside during their larvae stage (figure 6
5). This generation moves north but does not migrate as far nor overwinter. Energy is used to maximize and produce their offspring. Their breeding grounds are in the Eastern part of the U.S. The next group is the third generation. Larvae from this group can be found in certain southern states but majority in the Midwest and eastern states similar to the 2nd generation (figure 6). Adult emerges during the middle to the end of summer, July to August. Generation 3 monarch eggs are distributed throughout the northern range in the east right after they become adults, which will become the 4th generation. Some from the 3rdgeneration continues the migration to Canada. Early monarchs that surfaces are able to produce other offspring for the next generation. The later monarch adults either start their migration to overwintering sites or they go into diapause, delayed maturity, because of the cold temperatures then also head south (figure 7). They remain in this state until the next spring and breed in the overwintering sites (University of Minnesota, 2018). The last generation’s larvae stages begin in July until October. Eggs from this generation are laid from June to August and larvae are found in the same regions as the 3rd generation (figure 6). Their maturity stage starts to appear from August to April. This generation are the population that overwinters and migrates south towards the overwintering colonies in Mexico (figure 7). This occurs during the autumn season and are the only monarch generation who overwinters. Majority of this population will undergo diapause. Migration to overwinter in Mexico begins in September until November and they reside there until spring. This generation will migrate back north in the spring and will lay eggs that begins the new generations for the new year. 7
Generational Migration Routes Generation Migration Overwinter? Larvae Stage Adult Stage 1 North No March – April April – June 2 None No May – July June – July 3 Some – Some July – August July – August North/South 4 South Yes July – October August – April Table 1: This table represents the migration routes of the eastern monarch population. Time when larvae and adult butterflies emerge are also shown. Milkweeds Milkweeds (Asclepias species) are perennial herbaceous herbs that grows from a deep rhizome (USDA, 2006). They can be grown throughout the year and can produce beautiful nectar which attracts pollinators (Gaertnar, 1979) and migrating monarch butterflies. They are known as stepping stones for monarchs to support and maintain their population. Milkweed species can be found in a broad range of habitat including prairies, wetlands, riparian areas, agricultural fields, roadsides, pastures, and forest margins (Borders et al., 2013; Bhowmik, 1994). Most of the Asclepias species are grassland species. There are 72 native milkweed species in the U.S. 30 species occurs in the eastern population region. Milkweeds are propagated both sexually and vegetatively from seeds with a height up to 2 meters. Milkweed seed pods split open when ripe, then will disclose the seeds inside. Seeds have fluffy pappus to be carried in the wind. The seeds come secured with silky, white hairs, and are distributed by wind pollination (Urquhart, 1960; Bhowmik, 1994). 8
The plants are called “milkweed” because of its stems and leaves. The milky sap, also known as latex. The latex consists of a chemical called cardenolides (Bhowmik & Bandeen, 1976; Borders et al., 2013). This complex chemical makes the plant poisonous and dangerous for many livestock species such as cattle and poultry (Urquhart, 1960) and vertebrates. However, milkweed stems and leaves are an obligate host for monarch larvae (Borders et al., 2013). The butterfly larvae consume the milkweed leaves, which does not affect them but are stored inside the insect larvae. The chemical stays with the butterflies, therefore, the color provides protection since it signals to predators that they are poisonous. Both caterpillars and butterflies are brightly colored, and indicate to predators that they are toxic and should be avoided. Ethnobotany – Humans and Milkweeds Milkweeds also serve as a beneficial plant for ethnobotanical uses by Native Americans, such as the Potawatomi and Meskwaki tribe. Ethnobotanical usage of milkweed are for fiber, food, and medicine. This desirable herbaceous plant is well known for being a cordage-fiber plant used by Native Americans. This forb is known for its ‘bast’ fibers. Once these fibers are extracted, they can be turn into cords and ropes which are utilized to make fishing nets, head bands, hair nets, belts, bow strings, animal nets, feather capes, and dance skirts (Anderson & Moratto, 1996; Stevens, 2001). Some of these items are made specifically for ceremonial regalia. Certain products made by these fibers require an immense amount of harvested stalks to be created. Burning helps maintain the forb to redevelop to obtain more stalks for their fiber. Milkweed stems are collected during the autumn to winter seasons when the stalks senesce (Anderson & Moratto, 1996; Stevens, 2000). This usually occurs after traditional fire management practices in the beginning of autumn. Fire burns senescent plant material to stimulate new growth in the spring. After burning events, milkweeds appear to have better 9
growth (Anderson, 1991; Anderson & Moratto, 1996). Milkweeds can also capture sunlight and recycle nutrients into the soil for development. The stalks are split to retrieve the fibers and may be use alone or mixed with other plant fibers. It is then rolled on the thigh and twisted together to create chordage. Housewives tend to use milkweed plants to substitute as other vegetables, like asparagus. Toxicity from this herbaceous plant should always be removed before consuming it as a replacement of food plants. The toxics are removed by a boiling process which lasts between 2-5 minutes before eating or cooking it with other ingredients (Stevens, 2000; 2001). Young shoots, fruits, flowers, stems, and roots are preferred for consumption by the eastern and mid-western Native Americans (Stevens, 2000). The plant was known as a pot herb and many people sold it in packages for two cents back in the 1960s (Gaertner, 1979). Milkweed latex can cause pain and discomfort without sufficient preparation. Some species can lead to death like Asclepias subverticillata. This toxic species is found in the southern U.S. and can be mistaken for Indian hemp (Apocynum canibinum) which grows in the same habitat. Certain species, like butterfly milkweed (Asclepias tuberosa) or the butterfly milkweed, are frequently used as medicine to treat diseases and injuries. Scrofula is a form of tuberculosis on the neck with symptoms such as chills, weight loss, fever, and swelling of the lymph nodes. Milkweeds are use as ointments to lower these symptoms. The forb provides generalized medicinal uses and serve as diarrhea and blindness medicine, use in as an ointment for sore throat, heal snakebites, apply the root for rashes, remove tapeworm, and treat colic. The plant can be made into a drink and women can produce more milk for feeding purposes (Stevens, 2000). The Omahas, Poncas, Dakotas, and Menominis tribes consumes the root of butterfly milkweed for bronchial and pulmonary issues. When chewed, the root can be placed on wounds 10
to cure it. It can also be applied as a remedy for sores and prevent nauseas feelings (Steinuaer, 2013). Milkweeds were acknowledged as one of the most effective and important medicine (Stevens, 2000; 2001). Common Asclepias Species 30 species of milkweeds can be found in the Eastern states. Many of these includes common, swamp, showy, prairie, butterfly, and whorled milkweed (table 2). These are some of the well-known species that monarchs breed in the southern, eastern, and Midwest U.S. throughout their migration routes and endangered ones that are critical for the butterflies (Borders et al., 2013). This analysis will focus on 2 milkweed species in the eastern U.S. that resides in agricultural fields and prairies. One of which is a common species that’s used extensively by monarchs and one that is also utilized but endangered and requires conservation attention. Species that will be evaluated includes common milkweed and mead’s milkweeds. Common milkweed (Asclepias syriaca) is one of the native species in eastern U.S. utilize by monarchs. It’s currently distributed across the Midwestern and northeastern states with climatic requirements of 18 and 32°C (figure 8; Bhowmik, 1994; Borders et al., 2013). This is within the range of the eastern migratory monarch route. The inflorescence of this species appears in the upper leaf axils with 20-130 flowers per inflorescence. Blooming occurs from May until August. Common milkweed grows in sandy, rocky, or clay soils along lakes, waterways, prairies, roadsides, ponds, and waste areas (Stevens, 2006). Roadsides are significant for oviposit and easy access (Laura et al., 2000; Oberhauser & Solensky, 2004). Common milkweed is most favored by monarchs and utilized in the eastern monarch’s breeding range with great potential for milkweed population colonization and reproduction (Bhowmik, 1997; Hartzler & Buhler, 2000; Borders et al., 2013). Approximately 92% of 11
monarchs that return to overwinter in Mexico has been found to feed on common milkweed during their larvae stages (Malcolm et al., 1993). Mead’s milkweed (Asclepias meadii) is a common species found in the eastern tall grass prairie ecosystems. This current distribution extends from Kansas, Iowa, Illinois, Missouri, and Indiana (figure 9; Bowles et al., 1998; USFWS, 2018). This plant appears late May to June. Once matured, the stem bears about 6 to 15 flowers. They prefer wet to moderately dry tallgrass prairie environments (USFWS, 2018). Pollination is by done by bee pollinators. Growing from seeds can be difficult and requires 4 or more years to fully grown from a germinating seed to a flowering plant. However, they can live up to decades (Betz, 1989). This is one of the downsides to this species, but still beneficial for monarch larvae when mature. Status & Decline of Milkweed & Monarchs Monarch butterfly’s decline involves many factors. Monarchs are at risk due primarily due to pesticides in agricultural and prairie fields and milkweed destruction for human utilization. Anthropogenic activities have been a threat for the species and this have affected their migratory populations from Mexico, North America, and Canada locations. The current status of monarch butterflies is still underway although it continues to decline. The U.S. Fish and Wildlife Service (2014) petitioned to list the monarch butterfly as an ‘endangered species.’ A Species Status Assessment and Status Review is currently underway until summer of 2019 to decide whether the species should be listed as endangered (USFWS, 2016). Monarchs benefit greatly from the milkweeds, which follows with growth and development and the need of continuous corridors along their entire migratory routes. Because of this commensalism relationship, if milkweeds start to diminish, so will monarch populations. 12
The eastern monarch population has declined dramatically since the 1990s until today (figure 10). Over the past decades, monarch butterflies have declined by 80% that points to extinction of the species. Listing them as ‘endangered’ will offer time, possible protection, and a roadmap for conservation efforts. An estimate showed that there is an 11-57% probability that the eastern monarch population will collapse within 20 years (Centers for Biological Diversity, 2016). Habitat destruction of milkweed patches has been the widespread cause of destruction for monarch communities (New et al., 1995) specifically land transformation to agricultural fields. Prairie regions are vulnerable to development and expansion of modern agriculture in monarch migration regions (Smith, 1981; New et al., 1995; Vogel et al., 2007). Monarchs that occupy this ecosystem are interrupted heavily along with milkweeds (Swengel et al., 2011). These lands are converted to yield crops for human consumption, which is an important requirement, but can destroy numerous species that has mutualistic relationships. Mead’s milkweed is a native in these conversion habitats. It has been listed under the U.S. Fish and Wildlife Service since 1988 with similar threats such as: habitat loss and hay mowing (Harrison, 1988; USFWS, 2018). Iowa and Wisconsin are states that have lost majority of their prairie habitats. The remaining fragmented prairies are less than 5 ha in each state (Rooney & Leach, 2010; Ries & Debinski, 200l). Exploitation of species’ environment for human use has ruined many of the native organisms in this ecosystem and can also affect migrating ones, like monarchs. Wildlife Friendly Agriculture Practices Hay mowing takes place in June to July in prairie fields which affects the milkweed during their flowering period. This procedure has the ability to clear away premature follicles from mead’s milkweeds and this leaves the plants incomplete preventing sexual reproduction and 13
seed dispersal (Betz, 1989; Bowles et al., 1998; USFWS, 2018). Without a complete life cycle, the plant cannot develop fully and serve its purpose as a host plant for monarch larvae. Larvae depend on fully grown leaves to receive adequate food consumption. They do not feed on immature leaves and stems. Mowing can create disturbances for prairies and total plant coverage will be reduce. This also produces different species composition base on timing of occurrence (Fynn et al., 2004; Van Dyke et al., 2004). The common milkweed species is grown in the Midwest agricultural landscapes and its existence on the same lands as agricultural crops has cause yields to lessen (Bhowmik, 1994). But the presence milkweeds attract pollinators. Farmers despises insects and larvae that are found on milkweeds and other plant species which could impact their crops. This concern has caused farmer’s need to get rid of plants that interact with insects. The creation of the herbicide glyphosate was to enhanced crop fields for further and continuation of human utilization without disturbances (Wyrill & Burnside, 1977). Glyphosate has been an effective herbicide for humans but the opposite for insects and weeds. This herbicide has also caused destructions in certain crop fields which led to the creation of BT crops. BT (Bacillus thingenienis) crops are genetically modified organisms (GMOs) that were established to avoid glyphosate. It is a bacterium that is harmful and toxic to many insects and pests but provide crops with protection from glyphosate and plant quality improvement. This process is done by identifying and extracting genetic material from a donor organism. Only genes that resists the herbicide are preferred and then modified into crops (Bessin, 2003). Although BT crops resists this herbicide, it does kill off many unwanted plants and insects that humans assumed could impact their crops. Many plant species are invasive and have huge abundance in certain fields. However, this does not apply that all of them impact the crops 14
negatively. But many, including milkweeds, have no choice but to die off in that community because of these situations. It was discovered that common milkweed declines in Iowa, one of the Midwest states, correlated to use of genetically modified BT corn and soybean crop fields. Genetically modified (GM) crops and glyphosate are the main contributors to the reduction of this milkweed species. Monarchs rely heavily on common milkweeds found in these two GM croplands. A study revealed that these croplands consist of 78 times more monarchs compared to other habitats. There was approximately a 58% decline of the weed in the Midwest and an 81% decline in the monarch population (Hartzler, 2010). The decrease of both the milkweed and monarch populations will affect the reproduction rates in Iowa and surrounding states within the same ecosystems (Oberhauser et al., 2001; Hartzler, 2010), usually the 3rd generation. After the creation of transgenic BT crops, pollen from these hybrids that expresses the toxin has impacted the common milkweed species. Glyphosate is the main herbicide that’s used in agricultural crops (Laura et al., 2000). Prescribe Fire Management Prescribed fire on vegetation and landscapes have influenced many of our ecosystems. Its purpose is to maintain the biodiversity of certain landscapes and significant for natural resource management (Huntzinger, 2003; Driscoll et al., 2010). Native plants and animals on prescribed fire landscapes are adapted to fire events. Understanding these species response to fire is necessary for conservation efforts (Driscoll et al., 2010). This will reveal how fire may impact species and if it continues. Considering the effects of time, scale, frequency, and conditions of specific landscapes are important for prescribed burns (Van Dyke et al., 2004). They may influence the abundance of native species in certain areas (Fynn et al., 2004). 15
Fire have been a tool used to manage ecosystems for utilitarian or religious purposes. Traditional fire management practices were passed down by many generations in indigenous tribes. These groups of people have traditional ecological knowledge (TEK) of fire usage to establish specific desired habits for usage. The main uses of fire are to encourage seed production or to clear agricultural fields. Burning can promote environmental stability, maintenance of ecotones, and increase diversity and resources (Raish et al., 2005). The Patch Mosaic Burning (PMB) concept is the influence of space and time of fire regimes. (Driscoll et al., 2010). PMB fire creates a mosaic of patches which can increase heterogeneity, this belief is also known as pyrodiversity (Parr & Brockett, 1999; Baum & Sharber, 2012). The burning of the patches is known to have a variety of habitats that fit the needs of certain landscapes (Parr & Anderson, 2006). Burn events will supply species on certain landscapes with their preferred resources. PMB arises from indigenous traditional burning techniques. These burning concepts are used universally by many indigenous people (Parr & Anderson, 2006). Mosaic burning has been used by one of the largest group of indigenous people, the Australian Aborigines. This management system has been used as technique to clear land for agriculture and hunting. As civilization developed and evolved over the years, fire management has been viewed as a destructive method to use (Old, 1969). But now it is being implemented back into management procedures and shown positive results for habitat and species conservation. PMB can be a complex concept that consists of two variables: visible and invisible mosaic (Parr & Andersen, 2006). Invisible mosaic is the term which describes the results of fire regimes over fuel age by years viewing the frequency, intensity, and lengths of past intervals (Bradstock et al., 2005; Foster et al., 2017). Visible mosaic refers to the time-since-fire, or post 16
fire, status of burned areas. It is easier to determine and see the visible mosaic patches since the burn in these patches are frequently burned. Invisible mosaics are often covered up since the burn may be from years ago (figure 11). All characteristics of these two variables explains the diversity of fire regimes (Parr & Andersen, 2006). Mosaic prescribe burning was used as a management system with the belief that it reduces the heavy fuels that are used on certain lands to reduce intense wildfires. It also generates the growth of vegetation leading to the increase of diversity of plants and animals (Burrows et al., 2004). It is crucial to understand the how species react to postburn areas and see the different spatial patterns from the burning (Driscoll et al., 2010). This determine locations where prescribe fire can be used at to improve biodiversity. Fire regimes vary across different locations. The size of prescribe fire and its impacts can either be devastating or beneficial. Large, uncontrollable prescribe fires can destroy species diversity and infrastructures if they are within parameters of the fire. Small prescribe fires may not be able to help germinate native species in certain locations. Finding a decent size prescribe fire will helpful for landscapes to gradually develop. Fire can alter landscapes but also heals the land. This does depend on education, knowledge, and management experiences. Propagating Milkweeds From Root Cuttings The Asclepias species conservation and propagation by cuttings of the tuberous rhizome are considered for conservation. This method can be easy and reliable to increase milkweed populations. Cuttings should be done when the plants are inactive and should be kept cool and moist. The cuttings are then placed into containers of sand or mix with peat moss. Regular misting and covering the container is necessary for humidity. The roots will develop shoots 17
within 6 weeks. They can then be planted back onto landscapes to increase milkweed populations (Luna & Dumroese, 2013). The rhizomes should be planted in conservation regions or gardens by autumn so they will be able to develop enough to survive the colder seasons (Stevens, 2006). Milkweeds are able to reproduce visible shoot buds and flowers frequently when their roots are cut then replanted. But this procedure shouldn’t be done often because the plants will have poor crown development and overwinter imperfectly (Luna & Dumroese, 2013). From Seeds Propagating milkweeds from seeds are easy to complete and follow. The following steps will help anyone grow their own milkweeds for any purpose (Stevens, 2006). 1. Collect milkweed seeds after their pods have matured before opening. Avoid losing the seeds. 2. Remove grasses or weeds in your chosen area to plant the seeds and cultivate the soil. 3. Seeds can be placed into the soil during autumn and sow the seed with fine sand at a rate of 1/8 oz per sq. yd. 4. In greenhouses, milkweeds should be cold-treated (low temperatures) for at least 3 months. 5. Milkweed seeds are very capable of surviving. Weed invasive species and water the seeds as needed. These two methods of propagating milkweeds usually do not mature and blossom until the second year. Some may occur earlier depending on the environment that it is placed in. seeds and cuttings are available in nurseries. They can thrive in semi-dry places and spread from there increasing the population (Kinscher, 1992; Stevens, 2006). 18
Restoration: Fire on Milkweeds in Prairies Monarch and milkweed management has shown that the decline of one species influences one another. Many conservation and restoration efforts have been established to enhance the milkweed species to conserve monarchs that reside in the Midwest, northeastern, and southern U.S. regions during their migration. Different states in these regions have developed milkweed conservation solutions to support the butterflies. Many of which includes prescribe fire/burning, grazing, timber harvest, and roadside milkweed control (USDA, 2015). Fire management has been a known system used by many farmers in prairie fields since the 1950s (Bowles et al., 1998). This procedure is the main source of management used in prairies for maintenance and conservation on milkweeds and other forbs (Kline, 1997; Bowles et al., 1998). Fire managed prairies in the Midwest has been incorporating it into their procedures to increase abundance and biodiversity. Periodic burning can increase monarch habitats when applied to milkweeds. Dormant, small scale fires reveal that it encourages the development of flowering (Betz, 1989). The burning will be able to germinate the seeds on landscapes by the heat or smoke it generates. This increases seeds and biomass productivity (Old, 1969; Rooney & Leach, 2010). Low intensity is preferred and creates faster yet cooler burning for milkweeds. Plant responses to prescribe fire varies since it is influenced by the season in which it is used. Autumn and spring is more common for prescribe burns but forb diversity tends to be reduced during these seasons. Autumn is the perfect season to use prescribe fire but summer burns has also shown that milkweed can reemerge for another growth cycle of the same year (figure 12; Borders et al., 2013; USFWS, 2018). The development during the summer procedure is best toward late flowering grasses and forbs as they reach the period of senescence (Howe, 19
1994a; 1994b; Baum & Sharber, 2012). Milkweed is one of the forbs that regenerates growth during summer burns. In the Midwest, rainfall is not limited which allows the soil content to have higher organic matter. Infiltration after post burns are less critical and has better growth compared to areas where rain is limiting (Old, 1969). In a study, Baum & Sharber (2012) used prescribed fire on tall grass prairie fields in Oklahoma. This burning was implemented during the spring and summer to create a mosaic of patches of milkweeds in 200x400 meter plots (total of 6). Data were recorded after patches were burned, typically in July for both spring and summer, for 3 years. From their results, burned areas showed higher density of milkweeds while significantly lower in unburned plots. The fire eliminated the plants and tender leaves resulting in new growth of milkweeds in burned plots. Availability of the new reproduced plants for monarch use are from August to October. Eggs and larvae were abundant in the newly produced milkweeds from monitoring. The climax of both eggs and larvae in the burned plots were at the end of August and early September. Without the use of fire management, the milkweeds eventually senescence by August. Monarchs who utilizes these milkweeds in Oklahoma are the 3rd and 4th generations that migrates south to overwinter in Mexico. They are usually pre-migrants that travels back early to breed. Summer prescribed fire had the capability of supplying milkweed corridors for the pre- migrants during a difficult time when the plant is generally unavailable. With new emergence of milkweeds, the butterflies will be able to reproduce more and increase their populations. The study concluded that space and time of fire during the summer are vital to help conserve and generate more milkweed patches therefore leading to the increase of monarch populations (Baum & Sharber, 2012). 20
Fire management has been a successful addition for a variety of monarch habitats. In the Midwest states, prescribe fire has been beneficial to regenerate milkweeds for late-season breeding monarchs that are migrating to overwinter in Mexico. This occurs during the summer season and in tallgrass prairie habitats (USDA, 2015). With the loss of this ecosystem, it can be detrimental for these last migrators since this is when hay mowing also occurs. Another solution is to use prescribe fire after mowing of the prairies. This method will help the butterflies breed when they migrate through prairie landscapes to overwinter. Other Restoration Efforts/Ideas to Consider Agricultural Landscapes – data required for restoration & suggestions Collecting data on the current abundance and population number of monarch and milkweeds can provide suggestions to take accurate actions. To compensate for the loss of monarch butterflies in agricultural fields, lost and current remaining population statistics are crucial data. In agricultural fields, procedures implemented into caring for soybean and corn fields can affect milkweed species and monarch butterflies. Common milkweed is the species that’s found on this landscape. John Pleasants conducted a study to calculate the magnitude of milkweed loss for possible restoration efforts. He did this by focusing on agricultural landscapes and grasslands in the Midwest (Pleasants et al., 2017). An estimation of the first usage of the herbicide, glyphosate, was used to compare the population density of milkweed species from 1999-2014. Iowa was the only area that was sampled for milkweed data but was used to estimate for other states. Data were found to be intertwined with the 3rd and 4th generation monarchs who overwinters (Pleasants & Oberhauser, 2013). Data has shown an 80% decline of milkweed populations from 1994-2014, monarch population decline follows (figure 13; Monarch Watch, 2015). Monarchs 21
tend to populate the milkweeds that are located near agricultural crops in the Midwest. Their conservation goal is estimated to be 6-ha overwintering habitats. Results showed that abundance of milkweed left in their sampled sites of corn and soybean fields decreased 96% from 1999-2014 and 99% from 2010-2014 in their sampled location, Iowa. This is a total of 862 million milkweed stems that has been destroyed during those years. The total remaining estimation of milkweeds in all the Midwest states is about 1.34 billion. Majority of the areas with higher population size is on conservation lands and roadsides which provides milkweed and monarchs safe environments to reside in. The number of milkweeds that needs to be restored is a 32% increase, approximately 4.15-ha, to be able to support the overwintering generations. This would equal to 425 million stems added. To reach the goal of 6-ha, it would be 1.6 billion milkweed stems (Pleasants et al., 2017). Pleasants and his team does not expect for further milkweed loss in agricultural fields because they have all been extirpated. They suggested that restoration efforts should be the main focus for what’s been destroyed. To compensate for all the loss of milkweeds in this ecosystem, it would require efforts and manpower. Because roadsides and Conservation Reserve Programs (CRP) are spread throughout monarch ranges in the Midwest, it would be more effective to start restoring milkweed stems to those areas (Pleasants et al., 2017). It will also benefit agricultural fields within CRP lands. Some CRP lands and roadsides still have milkweeds, while some do not. Starting out small with a restoration project will help plan for better, more effective projects after mitigation and observations that may show improvements. With this to consider, the geographic area for milkweeds are significant for conservation. Having visuals of milkweed’s 22
current distribution and loss can provide information for conservationists and scientists to fill in the gaps that were destroyed. Conclusion/Discussion Monarch populations are breathtakingly, beautiful sights to see. Their extreme flight from three countries leaves everyone in awe. The different generations interaction and linkage to their host plant, milkweeds, throughout the year showcases the astounding ecological relationship between two species. Unfortunately, the synergistic relationship between milkweeds and monarchs leaves them vulnerable. If one disappears, so does the other. Milkweed species have been declining due to many human impacts involving herbicides, prairie degradation, or even land exploitation for human utilization. Because the butterflies require milkweeds to develop, the reduction in population size of one species leads to the same results as other. This requires conservation and mitigation efforts to help manage both species. Restoring millions of milkweeds to the eastern part of the U.S. will require attention and participation by local communities. Some milkweed species, such as Mead’s milkweed, are also threatened and listed as endangered. Restoration are required for them because they will continue to influence the monarch populations’ downfall. The sectors that requires attention in the eastern part of the U.S. are prairies and agricultural fields. The concern has been over two decades (Thogmartin et al., 2017). The use of prescribe fire for conservation of the forb will be able to provide a manageable monarch population. Studies have reported the importance of fire usage and has shown benefits for common milkweed and mead’s milkweed. It can also provide habitats for other species, such as pollinators. Ecosystem services from these areas can benefit other organisms, therefore, are significant for our ecosystems. 23
Government agencies are efficient with this situation of monarch decline. The USFWS had restrict the use of pesticides in GM crops (Kurth, 2014; Thogmartin et al., 2017). Less exposure to herbicides will be one less issue to worry about. Destructions in prairie and agricultural fields have been reduced and milkweed species has increase along with monarch larvae. Although this can be an extremely immense task, it may be achievable if broken down into segments. Focusing on critically threatened areas will create simpler tasks and opportunities for this restoration journey. Restoration efforts should engage farmers and roadside managers to restoration projects while educating them about milkweeds and monarchs. Education is always key for everything. Informing them about the value of monarchs could decrease or even ban or reduce the use of herbicide for agricultural lands. This could also influence prairies because they will understand the activity of hay mowing and know when to and when not to mow (Pleasants et al., 2017). 24
Figures & Tables Figure 1: North America’s Monarch Populations, western (west of the Rocky Mountains) and eastern population (east of the Rocky Mountains, Monarch Watch, 2010). Figure 2: Monarch Generation 1 – larvae distribution in southern U.S. (University of Minnesota, 2018). 25
Figure 3: Monarch Generation 1 – adult migration northward and distribution (University of Minnesota, 2018). Figure 4: Monarch Generation 2 – larvae distribution in the Midwest and eastern U.S. (University of Minnesota, 2018). 26
Figure 5: Generation 2 – Adult migration northward and distribution (University of Minnesota, 2018). Figure 6: Generation 3 and 4 – larvae distribution in the Midwest, southern, and eastern U.S. (University of Minnesota, 2018). 27
Figure 7: Generation 3 and 4 – Adult migration towards north and south and distribution (University of Minnesota, 2018). Figure 8: Distribution of ‘common milkweed’ species in eastern U.S. (Bhowmik, 1994). 28
Table 2: This table depicts the 9 available milkweed species and 1 endangered species identified in the southern, eastern, and Midwest region of the U.S. for monarch butterflies. (Borders et al., 2013). Species – Scientific Name Common Name Habitat/Distribution Asclepias asperula capricornu Antelope horns Upland prairie pastures, grasslands, and road sides. Abundant in: TX, OK, KS, NE Asclepias hirtella Tall green milkweed Upland tallgrass prairie, lowland prairie, prairie hay meadows, glades, roadsides, and marshy areas. Abundant in: MO Asclepias incarnata incarnata Swamp milkweed Wet meadows/prairies, sloughs & roadside ditches, swamp borders, marshes, ponds, and lakes. Abundant in: TX, some eastern states Asclepias speciosa Showy milkweed Short grass prairies, roadsides, rivers, streams, slough, pond, and lakes. Abundant in: OK, KS, NE Asclepias sullivanti Prairie milkweed Lowland/upland prairies, wet meadows, creek banks, and river bottoms. Abundant in: KS, MO, IL, IA Asclepias syriaca Common milkweed Prairies, old fields, margins of woods, flood plains-lakes/ponds, creek banks, roadsides, and railways. Abundant in: KS, NE, IA, MO, IL, WI, IN, KY Asclepias tuberosa interior Butterfly milkweed Open woods, prairies, savannas, old fields, and roadsides. Abundant in: KS, OK, AR, TX, IL, MS, IN, WI, KY, TN Asclepias verticillata Whorled milkweed Prairies, glades, open woods, fields, flood plains, and hillsides. Abundant in: KS, NE, IA, MO, IL, WI, IN, KY, OK, AR, LA Asclepias viridis Green antelopehorn Prairies, glades, grasslands, roadsides, and pastures. Abundant in: KS, OK, AR, LA, MS, MO Asclepias meadii Mead’s milkweed – Tallgrass prairie. endangered Abundant/endangered in: IA, IL, MO, KS 29
Figure 9: Current distribution of mead’s milkweed in eastern U.S. (USFWS, 2018). Figure 10: Eastern monarch population that has dwindle since 1994 until 2018 (Center for Biological Diversity, 2018). 30
Figure 11: Image A showcases visible mosaic while image B shows the invisible mosaic (Parr & Andersen, 2006). 31
Figure 12: (a) represents burned plots in spring whereas (b) represents plots burned during the summer (Baum & Sharber, 2012). Figure 13: Monarch population decline, 1999-2013 (Borders et al., 2013; Monarch Watch, 2015). 32
References Anderson, M. K. (1991). California Indian horticulture: Management and use of redbud by the Southern Sierra Miwok. Journal of Ethnobiology, 11 (1), 145–57. Anderson, M. K. & Moratto, M. J. (1996). Native American land-use practices and ecological impacts. Aspen Bibliography. Paper 1815. doi: https://digitalcommons.usu.edu/ aspen_bib/1815 Baum, K., & Sharber, W. (2012). Fire creates host plant patches for monarch butterflies. Biology Letters, 8(6), 968-971. Bessin, R. (2003). Bt-Corn: What it is and how it works. University of Kentucky – College of Agriculture. Retrieved from https://entomology.ca.uky.edu/files/efpdf1/ef130.pdf Betz, R. F. (1989). Ecology of Mead's milkweed (Asclepias meadii Torrey). Pp. 187-191 in T. B. Bragg & J. Stub-bendieck (eds.). Proceedings of the Eleventh North American Prairie Conference: Prairie pioneers: Ecology, history and culture. Lincoln, Neb.: University of Nebraska. Bhowmik, P., & Bandeen, J. (1976). THE BIOLOGY OF CANADIAN WEEDS: 19. Asclepias syriaca L. Canadian Journal of Plant Science, 56(3), 579-589. Bhowmik, P.C. (1994). Biology and control of common milkweed (Asclepias syriaca). Reviews of Weed Science, 6, 227–250. Bhowmik, P.C. (1997). Weed Biology: Importance to Weed Management. Weed Science, 45(3), 349-356. Borders, B., Casey, A., King, R., Jacobs, A., Mader, E., Row, J. M., Taylor, C., Wynia, R. (2013). Pollinator Plants of the Central United States Native Milkweeds (Asclepias spp.). The Xerces Society for Invertebrate Conservation. Retrieved from 33
https://www.nrcs.usda.gov/Internet/FSE_PLANTMATERIALS/publications/ mopmcpu11905.pdf Bowles, M., McBride, J., & Betz, R. (1998). Management and Restoration Ecology of the Federal Threatened Mead's Milkweed, Asclepias meadii (Asclepiadaceae). Annals of the Missouri Botanical Garden, 85(1), 110-125. doi: 10.2307/2992001 Bradstock, R., Bedward, M., Gill, A., & Cohn, J. (2005). Which mosaic? A landscape ecological approach for evaluating interactions between fire regimes, habitat and animals. Wildlife Research, 32(5), 409. Burrows, N. D., Burbidge, A.A., & Fuller, P. J. (2004). Integrating indigenous knowledge of wildland fire and western technology to conserve biodiversity in an Australian desert. Proceedings of Millennium Assessment, bridging scales and epistemologies conference. Millennium ecosystem assessment. Retrieved from http://www.millenniumassessment.org /en/About.Meetings.Bridging.Proceedings.asx Centers for Biological Diversity. (2016). Agreement: Monarch Butterfly to Get Endangered Species Act Protection Decision by 2019. Retrieved from https://www.biologicaldiversity.org/news/press_releases/2016/monarch-butterfly-07-05- 2016.html Driscoll, D. A., Lindenmayer, D. B., Bennett, A. F., Bode, M., Bradstock, R. A., Cary, G. J. Clarke, M. F., Dexter, N., Fensham, R., Friend, G., Gill, M., James, S., Kay, G, Keith, D. A., MacGregor, C., Russell-Smith, J., Salt, D., Watson, J. E. M., & York, A. (2010). Fire management for biodiversity conservation: Key research questions and our capacity to answer them. Biological Conservation, 143(9), 1928-1939. doi: 10.1016/ j.biocon.2010.05.026 34
Flockhart, D. T. T., Wassenaar, L. I., Martin, T. G., Hobson, K. A., Wunder, M. B., & Norris, D. R. (2013). Tracking multi-generational colonization of the breeding grounds by monarch butterflies in eastern North America. Proc R Soc B, 280: 20131087. doi:10.1098/rspb.2013.1087 Foster, C. N, Barton, P. S., Robinson, N. M., MacGregor, C. I., & Lindenmayer, D. B. (2017). Effects of a large wildfire on vegetation structure in a variable fire mosaic. Ecological applications a publication of the Ecological Society of America, 27(8), 2369-2381. Fynn, R. W. S., Morris, C. D., & Edwards, T. J. (2004). Effect of Burning and Mowing on Grass and Forb Diversity in a Long-Term Grassland Experiment. Applied Vegetation Science, 7(1), 1. Gaertner, E. (1979). The history and use of milkweed (Asclepias Syriaca L.). Economic Botany, 33(2), 119-123. doi: 10.1007/BF02858278 Harrison, W.E. (1988). Endangered and threatened wildlife and plants: determination of threatened status for Asclepias meadii (Mead's milkweed). Federal Register, 53, 33992- 33995. Hartzler, R., & Buhler, D. (2000). Occurrence of common milkweed (Asclepias syriaca) in cropland and adjacent areas. Crop Protection, 19(5), 363-366. Hartzler, R. G. (2010). Reduction in common milkweed (Asclepias syriaca) occurrence in Iowa cropland from 1999 to 2009. Crop Protection, 29, 1542–1544. Hopwood, J. L. (2013). Roadsides as Habitat for Pollinators: Management to support Bees and Butterflies. The Xerces Society for Invertebrate Conservation. Retrieved from http://www.icoet.net/icoet_2013/documents/papers/ICOET2013_Paper403C Hopwood.pdf 35
Howe, H. (1994a). Response of Early- and Late-Flowering Plants to Fire Season in Experimental Prairies. Ecological Applications a Publication of the Ecological Society of America, 4(1), 121-133. Howe, H. (1994b). Managing Species Diversity in Tallgrass Prairie: Assumptions and Implications. Conservation Biology, 8(3), 691-704. doi: 10.1046/j.1523- 739.1994.08030691.x Huntzinger, M. (2003) Effects of fire management practices on butterfly diversity in the forested western United States. Biological Conservation, 113(1), 1–12. Inamine, H., Ellner, S. P., Springer, J. P., and Agrawal, A. A. (2016). Linking the continental migratory cycle of the monarch butterfly to understand its population decline. Oikos, 125(8), 1081-1091. doi: https://doi.org/10.1111/oik.03196 Kinscher, K. (1992). Medicinal wild plants of the prairie. An ethnobotanical guide. University Press of Kansas. Pp. 84-94. Kline, V. M. (Eds.). (1997). The tallgrass restoration handbook: for prairies, savannas, and woodlands. Planning a restoration, p. 31-46. Washington, DC: Island Press. Kurth, J. W. (2014). Use of agricultural practices in wildlife management in the National Wildlife Refuge system Memorandum (Washington, DC: US Fish and Wildlife Service). Laura, C., Hansen, J., & Obrycki, J. (2000). Field Deposition of Bt Transgenic Corn Pollen: Lethal Effects on the Monarch Butterfly. Oecologia, 125(2), 241-248. doi: http://www.jstor.org/stable/4222768 Luna, T. & Dumroese, R. K. (2013). Monarchs (Danaus plexippus) and milkweeds (Asclepias species): the current situation and methods for propagating milkweeds. Native Plants Journal, 14(1), 5–15. 36
Malcolm, S.B., Cockrell, B.J. & Brower, L.P. (1993). Spring recolonization of the eastern North America by the monarch butterfly: successive brood or single sweep migration? Biology and Conservation of the Monarch Butterfly (ed. by S.B. Malcolm and M.P. Zalucki), pp. 253–267. Science Series No. 38, Natural History Museum of Los Angeles County, Los Angeles, California. Monarch Watch. (2015). Monarch population status. Retrieved from http://monarchwatch.org/ blog/2016/02/26/monarch-population-status26/ New, T., Pyle, R., Thomas, J., & Hammond, P. (1995). Butterfly conservation management, Annual Review of Entomology 40, 57–83. Oberhauser, K. S., Prysby, M. D., Mattila, H. R., Stanley-Hom, D. E., Sears, M. K., Dively, G., Olson, E., Pleasants, J. M., Lam, W. F., Hellmich, R. L. (2001). Temporal and spatial overlap between monarch larvae and corn pollen. Proc. Nat. Acad. Sci. USA, 98, 11913- 11918. doi: https://doi.org/10.1073/pnas.211234298 Oberhauser, K. S. & Solensky, M. J. (2004). The Monarch Butterfly: Biology and Conservation. Ithaca, New York: Comstock Publishing Associates. Old, S. M. (1969). Microclimates, fire, and plant production in an Illinois prairie. Ecological Monographs, 39, 355-384. doi: 10.2307/1942353 Pleasants, J. M. & Oberhauser, K. S. (2013). Milkweed loss in agricultural fields because of herbicide use: effect on the monarch butterfly population. Insect Conservation and Diversity 6(2), 135-144. doi: 10.1111/j.1752-4598.2012.00196.x Pleasants, J., Leather, S. R., & Stewart, A. (2017). Milkweed restoration in the Midwest for monarch butterfly recovery: Estimates of milkweeds lost, milkweeds remaining and 37
milkweeds that must be added to increase the monarch population. Insect Conservation and Diversity, 10(1), 42-53. doi: 10.1111/icad.12198 Pitman, G. M., Flockhart, D., & Norris, D. (2018). Patterns and causes of oviposition in monarch butterflies: Implications for milkweed restoration. Biological Conservation, 217, 54-65. doi: 10.1016/j.biocon.2017.10.019 Parr, C. L., and Brockett, B.H. (1999). Patch-mosaic burning: a new paradigm for savanna fire management in protected areas? Koedoe, 42, 117-130. doi: 10.4102/koedoe.v42i2.237 Parr, C., & Andersen, A. (2006). Patch Mosaic Burning for Biodiversity Conservation: A Critique of the Pyrodiversity Paradigm. Conservation Biology, 20(6), 1610-1619. doi: 10.1111/j.1523-1739.2006.00492.x Raish, C., González-Cabán, A., & Condie, C. J. (2005). The importance of traditional fire use and management practices for contemporary land managers in the American Southwest. Global Environmental Change B: Environmental Hazards, 6(2), 115-122. Ries, L., and Debinski, D. (2001). Butterfly Responses to Habitat Edges in the Highly Fragmented Prairies of Central Iowa. The Journal of Animal Ecology, 70(5), 840-852. doi: 10.1046/j.0021-8790.2001.00546.x Rooney, R.P. & Leach, M.K. (2010). Replacing Hay-mowing with Prescribed Fire Restores Species Diversity and Conservation Value in a Tallgrass Prairie Sampled Thrice: A 59- Year Study. American Midland Naturalist, 164(2), 311-321. doi: https://doi.org/10.1674 /0003-0031-164.2.311 Smith, D. D. (1981) Iowa prairie-an endangered ecosystem. Proceedings of the Iowa Academy of Science, 88, 7-10. 38
Steinauer, G. (2013). Healing Plants of the Prairie. Nebraska Land. Retrieved from http://nativeplants.ku.edu/wp-content/uploads/2012/08/medicinal-plant-final-pdf.pdf Stevens, M. (2001). Plant guide for butterfly milkweed (Asclepias tuberosa). Greensboro. USDA Natural Resources Conservation Service, National Plant Data Center. Stevens, M. (2000). Plant guide for butterfly milkweed (Asclepias tuberosa). Greensboro. USDA Natural Resources Conservation Service, National Plant Data Center. Stevens, M. (2006). Plant guide for common milkweed (Asclepias syriaca). Greensboro. USDA. Natural Resources Conservation, National Plant Data Center. Swengel, S. R., Schlicht, D., Olsen, F., & Swengel, A. B. (2011). Declines of prairie butterflies in the midwestern USA. Journal of Insect Conservation, 15(1-2), 327-339. doi: 10.1007/s10841-010-9323-1 Thogmartin, W. E., López-Hoffman, L., Rohweder, J., Diffendorfer, J., Drum, R., Semmens, D., Black, S., Caldwell, I., Cotter, D., Drobney, P., Jackson, L. L., Gale, M., Helmers, D., Hilburger, S., Howard, E., Oberhauser, K., Pleasants, J., Semmens, B., Taylor, O., Ward, P., Weltzin, J. F., & Wiederholt, R. (2017). Restoring monarch butterfly habitat in the Midwestern US: ‘all hands on deck’. Environmental Research Letters, 12(7), 10. doi: 10.1088/1748-9326/aa7637 University of Minnesota. (2018). Annual Life Cycle. Retrieved from https://monarchlab.org/ biology-and-research/biology-and-natural-history/breeding-life-cycle/annual-life-cycle/ USDA. (2015). Conservation and Management of Monarch Butterflies: A Strategic Framework. Retrieved from https://wsfrprograms.fws.gov/subpages/Pollinators/USDA_ ConsMgmtMonarchsStrategicFramework_March2015%20 (1).pdf. 39
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