CSL LTVCA Project

Assessment of the Factors Affecting the Spread of Allaria petiolata (Garlic Mustard)

Submitted: April 5, 2012 Authors: Omar Champsi Jake Balkos, Maisa Fumagalli, Khoa Tran

Professor: Dr. Daria Koscinski

Contents
1.0 2.0 Background.. 2 Factors Affecting the Spread of A. petiolata...3
2.1 Secondary Composition....4 2.2 Lack of Natural Enemies...5 2.3 Life Cycle......5 2.4 Phenotypic Plasticity and Disturbances7 2.5 Satellite Communities.......7 2.6 Human Influence and Management Techniques...8

3.0 4.0 5.0

Summary..11 References12 Appendix..17

 1.0 Background Invasive species are those that have been introduced into an environment in which they are non-native; in this way their introduction usually leads to various forms of environmental damage (IUCN, 2012). In the current study, we assess the factors affecting the spread of the invasive species Alliaria petiolata, also known as garlic mustard. The spread of invasive species such as A. petiolata are believed to be the second largest threat to biodiversity amongst plants (Lomusico, 2011). Garlic mustard is a biennial plant of the mustard family, originating in Europe, western Asia, North Africa and Asia Minor, it first appeared in North America during the colonial times in the mid 19th century (NCC, 2007; Lomusico, 2011). At the time colonists used garlic mustard as a substitute for garlic as well as for various medicinal purposes such as clearing infections (Schooley, 2005). Since garlic mustard is a biennial plant, its appearance varies between stages (Figure 1). In its first year, it remains a low growing cluster of dark green, kidney shaped leaves (Figure 2; Saanich, 2011; Huffman, 2005). In its second year, the garlic mustard plant has the potential to grow to a height greater than one meter; during this time its leaves take a more narrow shape and form a tip at the end (Figure 3; Saanich, 2011). Additionally, during this period the plant begins to flower and we begin to see white petals forming, in the shape of a cross at the top of the plant (Figure 3; Huffman, 2005). Apart from its appearance, garlic mustard can also be identified via its smell, when crushed up it emits a garlic odour (Saanich, 2011). As previously mentioned, garlic mustard is an invasive species of plant which was introduced to the woodland forests of North America in the mid 19th century. Upon

 introduction into the North American forest system, it was found that this plant threatened the existence of many native plant species (NCC, 2007). Currently, garlic mustard spreads at a rate of approximately 6400 km per year (Lomusico, 20011). In doing so, it destroys the ability for the native plants to thrive and grow, thereby leading them to their demise. The features allowing this plant thrive and inflict such damage can be described as advantages it has over other native plants (NCC, 2007). These features can be classified into categories involving biotic, abiotic and human interactions. The goal of this paper is to assess the factors that may affect the spread of garlic mustard across these three categories in the Lower Thames Valley Conservation Area (LTVCA), specifically, in the Longwoods Conservation Area (LCA). The LCA is an area that boasts 63 hectares of Carolinian forests, wetland and meadows, additionally, the conservation area provides a grounds to perform various recreational activities such as hiking, cross country skiing, snowshoeing and camping (LTVCA, 2012). In addition to this, the LCA is also home to many different plant, animal and insect species (LTVCA, 2012). Due to the pinnacle role that the LCA plays in its community, it is imperative that invasive species such as garlic mustard are eradicated as their spread may lead to the death of many native plants in the conservation area, thereby destroying the original environment. This will in turn affect both the people who use LCA for recreational, educational and other purposes as well as, the various ecosystems within it. 2.0 Factors affecting the spread of A. petiolata The spread and invasive success of garlic mustard is not attributed to a single trait alone, instead, it is the result of a medley of different mechanisms acting in a collective

 fashion (Rodgers et al., 2008). These mechanisms include a combination of its slightly dissimilar biological composition (secondary compounds), life cycle and phenotypic plasticity from plants within its vicinity (Rodgers et al., 2008). Additionally, its spread is fostered via the lack of natural enemies, the growth of its satellite colonies and various forms of human intervention. In this section of the paper, we will review how each of these factors affect the spread of garlic mustard. 2.1 Secondary Compounds One of the primary factors associated with the garlic mustards endurance is the composition of its secondary compounds. These compounds include an assortment of flavonoids, glycosides, glucosinolates, as well as defensive proteins, thereby making its consumption unfavorable by herbivores and providing it an advantage over incumbent plants (Daxenbichler et al., 1991; Rodgers et al., 2008). One example of this advantage of secondary compounds can be seen via the cyanide production pathway in garlic mustard. In this case, cyanide, an inhibitor of the respiratory electron transport chain, is derived from glucosinolates via hydrolysis in garlic mustard (Fahey et al., 2001). This cyanide acts as a toxic deterrent for consumption to a wide range of organisms (Fahey et al., 2001). This characteristic is also one plausible reason for butterflies such as the West Virginia White butterfly (Pieris virginiensis) to oviposit on garlic mustard leaves over their usual preference of toothwort (Dentaria spp.) even though the fraction of eggs hatching on garlic mustard has been observed to be significantly lower (Porter, 1994). Furthermore, garlic mustard has intrinsic biofumigating properties it releases antifungal chemicals that inhibit mutualistic mycorrhizae relationships in the soil (Stinson et al., 2006). These relationships are are required for the optimal growth of many native

 species (Rodgers et al., 2008). In this way it sterilizes the soil and produces unfavorable conditions for the growth of plants aside from itself. Following biofumigation, garlic mustards secondary compounds take over to defend its existence in the soil, thus contributing to it sustainability and spread. 2.2 Lack of Natural Enemies Another factor observed in the success of invasive plants in general, is the lack of natural enemies (Keane and Crawley, 2002). As previously mentioned, garlic mustards secondary compounds make it an unfavorable meal for many herbivores (Keane and Crawley, 2002). Although there are at least 69 different insects known to consume garlic mustard, there are no species native or present in North America that seem to do this (Szentesi, 1991). Due to this, there is a lack of control mechanisms to limit the production of garlic mustard plants and seeds (Szentesi, 1991). An example of this can be seen through the feeding behavior of the white tailed deer (Odocoileus virginianus), these herbivores have chosen to consume plants other than garlic mustard (Williams and Ward, 2006). In doing so, they choose to consume competing native plants surrounding garlic mustard, this suppresses the competition of the native plant species and further allows the spread of garlic mustard (through the growth of the plant and the spreading of its seeds) to occur without a natural mechanism of control (Williams and Ward, 2006). The introduction of non-native insects is a possible solution to limit the spread of garlic mustard due to their effectiveness in eradicating large colonies as per Hinz and Gerber (2000) but further considerations required on host specificity. 2.3 Life Cycle The life cycle of garlic mustard is also a characteristic that differentiates itself

 from the native species. Many of the woodland herbs which are native competitors to this invasive species are perennials with limited seed production, growth which is limited by understory light availability and a strong influence of resource availability affecting its relative spatial distribution (Whigham, 2004). On the contrary, garlic mustard is an obligate biennial, a plant with a two-year life cycle in any particular environment which is divided into two stages, a rosette stage and an adult stage (Rodgers et al., 2008). The first stage begins when garlic mustard is present as basal rosettes (in the winter season), during early spring it germinates and produces dark green leaves, this tends to take place before most of the native species begin to develop and garlic mustard is generally fully-grown before most of its perennial native competitors (Lomuscio, 2010; Rodgers et al., 2008). The growth of its dark green leaves block sunlight to the soil beneath, thus preventing the growth of the native species, as many of them are shade intolerant, additionally this lack of light also affects certain microbe and soil activities (Lomuscio, 2010; Rodgers et al., 2008). In the subsequent year, the plant begins its adult stageduring early spring it begins to flower, allowing it to take advantage of the soils nutrients before the inactive competing plants get a chance (Rodgers et al., 2008). This further enhances the plants viability and endurance in its non-native environment and assist in its spread. Another part of garlic mustards life cycle that adds to its success as an aggressive invasive species is the mating system it employs, this system leads to a high seed production (Rodgers et al., 2008). Firstly, the flowers of garlic mustard are able to self-pollinate, therefore, if typical pollinators fail to arrive, a single plant can begin an entire invasive population on its own (Rodgers et al., 2008). Secondly, a single garlic

 mustard plant is capable of producing over 3500 seeds, (Susko and Lovett-Doust, 2000) and a population can reach 107,000 seeds per m2 (as observed in Ontario) (Cavers et al., 1979). This substantially outnumbers the native plant species seed production and is another reason for garlic mustards strong invasive effects (Cavers et al., 1979). 2.4 Phenotypic plasticity and disturbances Another reason why invasive species are so successful is that they are able to grow in almost any condition and are often R-strategists that thrive from disturbances (Yates et al., 2004). From a molecular perspective, this feature is known as phenotypic plasticity and allows garlic mustard to adjust to physiological and morphological changes in the environment. For example, in order to cope with variable environments consisting of fluctuating availability of water and nutrients, it has responded by adjusting its internal enzymatic activity and leaf size (Hillstrom and Cipollini, 2011). More generally however, garlic mustard has the ability to colonize areas that have been disturbed, therefore, frequently disturbed areas such as landscape boundaries or transitional zones, with large open areas are susceptible to colonization by Garlic mustard (Stinson et al., 2006). An example of this can be seen through the varying landscape elevations of the LCA. This variation in elevation makes LCA susceptible to flooding (a form of disturbance), these floods provide a means of seed dispersion, which may contribute to the plants dispersion through various regions of LCA. 2.5 Satellite communities Garlic mustard spreads out from a core through multiple small, satellite populations. The rate of spread is a factor of the rate of colonization of satellite populations (Nuzzo, 1999). The spread of the population is also dependent on natural

 (wind and flooding) and anthropogenic (recreation, fragmentation) disturbances (Stinson et al., 2006; Lomuscio, 2010). Whitman (2006), recommends focusing on preventing the spread of satellite colonies when resources are scarce as they can then become the main population from which new satellites stem from. Garlic mustard has an average growth rate of 1.9 cm/day, this growth is described by an advanceretreat pattern, where the final result is an increase in the population of garlic mustard via satellite populations (Rodgers et al., 2008; Nuzzo, 1999). This was evident during our visit to the LCA, as we saw large established garlic mustard populations separated from smaller, presumably satellite populations of garlic mustard by areas of no garlic mustard growth (Figure 4). 2.6 Human influence and management techniques In this section, we will discuss the ways in which various management techniques may ameliorate the spread of garlic mustard. More specifically, we will focus on the ways in which common management techniques are performed incorrectly, therein facilitating the spread of garlic mustard. Currently, it seems that the most effective way to reduce the spread of garlic mustard is through various forms of human intervention. This is because no natural occurring factors (ie. herbivores) exist in North America (Szentesi, 1991), more specifically the LCA to eradicate garlic mustard. Therefore, it is of utmost importance to understand the ways in which management may negatively impact the reduction of garlic mustard via the improper use of management techniques. Something to keep in mind is that management techniques require one, all or a combination of the following resources: man-power, time and funding. In certain cases, not enough of these resources are available; this limits the ability to eradicate this invasive species from our

 plant population. The first management solution is manual removal. This method is by far the cheapest, however, it demands a large time commitment and lots of support in the form of manpower. This method involves pruning the flower heads of the plant and severing its stems as well as hand-pulling the plant out of the ground (NCC, 2007). If this process is not completed correctly, it may in fact promote the growth of the garlic mustard plant and further its spread in two ways. Firstly, according to Czarapata (2005), improper removal of the plant will turn the soil and aid in the weeds recovery. Secondly, cutting the flower heads alone, without severing the stems of the plant will promote the growth of new flower heads, thereby promoting further spread (Whitman, 2006). In order to circumvent these issues and decrease the frequency and scale of spread of garlic mustard in LCA, the LTVCA may consider encouraging visitors to avoid pulling plants out of the ground unless they understand the appropriate removal and disposal techniques. They may be able to do this by putting up signs describing these techniques at the start of various trails to educate hikers as well as all other users of the grounds. Additionally, in order to ensure proper disposal, they may suggest that those pulling plants out of the ground place them in a garbage bag immediately after removal. This will ensure that once a plant has been uprooted, it is removed from the LCA site itself. This is especially important if flowering has occurred in the plant, as these plants will release their seeds and create an opportunity for more plants to grow (Herms et al., 2002). In doing so, they may be able to effectively mitigate the spread of garlic mustard to other sites within LCA. Another management technique, which has shown to increase the spread of garlic mustard when implemented incorrectly is, controlled burns. When performed correctly,

 these burns are effective in eradicating the existing garlic mustard plants from the soil and allow the sprouting of the native understory plants (Nuzzo et al., 1996). However, Whitman (2006) suggests that if this technique is performed incorrectly via a short, low intensity burn, this will actually increase the population of the garlic mustard plant. It is also important to consider the fact that the burning does not eliminate the existing seed bank, and therefore can only prevent the addition of new seeds to the seed bank for that year (NCC, 2007). Furthermore, in a site such as LCA, these burns, even when performed correctly, may prove to be quite dangerous. The reason for this is due to the fact that much of the garlic mustard spread in LCA is located in or around forested regions of the conservation area, therefore burning may cause a larger fire to the surrounding areas. In addition to this, the LCA is home to many species of insects and animals, these native species may not be able to handle the heat and intensity of the burning process and therefore it may be detrimental to the health of the LCA ecosystem. Other mechanisms of human influence in preventing the spread of garlic mustard include the use of Roundup, a glyphosate based chemical. Although this chemical has been shown to be successful in eliminating established plants of garlic mustard, it does not eliminate the seeds that the plants produce (Murphy et al., 2007). Furthermore, if only applied as a one-time use without annual follow-up, it has been shown to increase garlic mustard density relative to no treatment (Murphy et al., 2007). Additionally, a method such as this would require a constant funding, something which may or may not be available to the LTVCA. Apart from this, Roundup can also harm the growth of other native plants if applied incorrectly; thereby further destroying the native plant species population (Murphy et al., 2007).

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 Overall, the success of humans as a form of intervention to the spread of garlic mustard depends heavily on the application of the method chosen. In this way human intervention has the potential to, increase, decrease, or have no effect garlic mustard spread and population growth. 3.0 Summary To summarize, there are many factors affecting the spread of garlic mustard. In the case of LCA, it is possible that eradicating garlic mustard from the conservation grounds may be an unrealistic goal for the near future, however it may be possible in the long-term. In order to make this possible an appropriate management plan must be established. This plan should encompass management solutions for all the factors discussed above. Even if this plan is not able to completely eradicate the growth of garlic mustard, it should decrease the plants prevalence on LCA grounds. It is important to consider the fact that there may already be millions of seeds from the garlic mustard plant scattered in various strata of the soil, which remain viable for years. This is one of the primary reasons a project such as this is an ongoing process and will take years and years to complete. Finally, one possible consideration may be to introduce another plant native to North American soil with a similar life history strategy to that of garlic mustard (fast growing, shade-tolerant) to areas with already growing garlic mustard (NCC, 2007). The logic here is that, in doing so, you may be introducing a form of competition to the garlic mustard plant, thereby decreasing its prevalence and eventually eliminating it from LCA. However, due to the robust features associated with the spread of garlic mustard, even this may not be enough.

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 References:
Note: References were cited according to the guidelines provided by the Journal of Zoology

Baskin J.M. and Baskin C.C. 1992. Seed-germination biology of the weedy biennial Alliaria petiolata. Natural Areas Journal. 12: 191-197.

Carlson, A.M., Gorchov, D.L. 2004. Effects of herbicide on the invasive biennial Alliaria petiolata (garlic mustard) and initial responses of native plants in a southwestern Ohio forest. Restoration Ecology 12: 559-567

Cavers, P.B., Heagy, M.I., and Kokron R.F. 1979. The biology of Canadian weeds, 35: Alliaria petiolata (M. Bieb.) Cavara and Grande. Canadian Journal of Plant Sciences 59:217-229.

Czarapata, E. 2005. Invasive Plants of the Upper Midwest: an illustrated guide to their identification and control. The University of Wisconsin.

Daxenbichler, M.E., Spencer, G.F., Carlson, D.G., Rose, G.B., Brinker A.M., and Powell R.G. 1991. Glucosinolate composition of seeds from 297 of wild plants. Phytochemistry, 30:2623-2638.

Fahey, J.W., Zalcmann, A.T., and Talalay, P. 2001. The chemical diversity and distribution of glucosinolates and isothiocyanates among plants. Phytochemistry 56:5-51.

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 Herms, C.P., M.N. Frey, and J. Cardina. 2002. Novel approaches to halting seed production and accelerating seed death in the exotic forest herb, Alliaria

petiolata. The Ohio State University. ESA Annual Meeting: Poster Presentation.

Hillstrom, C. and Cipollini, D. 2011. Variation in Phenotypic Plasticity among Native and Invasive Populations of Alliaria petiolata. Journal of Plant Sciences 172:763-772.

Hinz, H. L. and E. Gerber. 2000. Investigations on potential biological control agents of garlic mustard, Alliaria petiolata (Bieb.) Cavara & Grande. Annual Report for 1999. CABI Bioscience, Delemont, Switzerland.

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hypothesis. Trends in Ecology and Evolution. 17:164-170.

Lomuscio III, M.F. 2010. Modeling the spread of Alliaria petiolata across different landscape distributions Western Carolina University, 33 pages.

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 Lower Thames Valley Conservation Authority. LTVCA conservation areas. Availiable from http://www.lowerthames-conservation.on.ca/ [accessed 1 April 2012].

Meekins, J.F. & McCarthy, B.C. 2002. Effect of population density on the demography of an invasive plant (Alliaria petiolata, Brassicaceae) population in a southeastern Ohio forest. American Midland Naturalist Journal. 147:256-278.

Murphy, SD. 2005. Concurrent management of an exotic species and initial restoration efforts in forests. Restoration Ecology 13:584-593.

Murphy, S.D., Flanagan, J., Noll, K., Wilson, D. & Duncan, B. 2007. How Incomplete Exotic Species Management Can Make Matters Worse. Journal of Restoration Ecology 25:85-93.

Nature Conservancy of Canada (NCC). 2007. Control Methods for the Invasive Plant Garlic Mustard (Alliaria petiolata) within Ontario Natural Areas. V1.0. NCC Southwestern Ontario Region, London, Ontario. 16 pp.

Nuzzo, V. 1999. Invasion Pattern of Herb Garlic Mustard (Alliaria petiolata) in High Quality Forests. Biological Invasions. Biological Invasions. 1:169-179.

Nuzzo, V., McClain, W. & Strole, T. (1996). Fire impact on groundlayer forest in a sand forest: 1990-1994. American Midland Naturalist 136:207-221.

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Porter, A. 1994. Implications of introduced garlic mustard (Alliaria petiolata) in the habitat of Pieris virginiensis (Pieridae). Journal of the Lepidopterists Society 48:171-172.

Department of Environmental Services. 2011. Invasive Plant Alert: Alleria petiolata.

Schooly, J. 2005. Problem weed of the month: garlic mustard [online]. Available from http://www.omafra.gov.on.ca/english/crops/hort/news/hortmatt/2005/10hrt05a4. htm [accessed 1 April 2012].

Slaughter, B, D. Gorchov. 2004. Reducing the invasive biennial Alliaria petiolata affects the plant community in deciduous forests in Ohio. Poster Session 7: ESA 2004 Annual Meeting. Portland, Oregon.

Susko, D.J., and Lovett-Doust, L. 1998. Variable patterns of seed maturation and abortion in Alliaria petiolata (Brassicaceae). Canadian Journal of Botony 70:1677-1686.

Szenthesi, A. 1991. Controversial components of plant apparency in Alliaria petiolata Cavara & Grande (Cruciferae). Symposium Biologis, Hungarica 39:237-244.

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Rodgers, V.L., Stinson, K.A., and Finzi, A.C. 2008. Ready or not, garlic mustard is moving in: Alliaria petiolata as a member of eastern North American forests. BioScience, 58:426-436.

Whigham, D.E. 2004. Ecology of the woodland herbs in temperate deciduous forests. Annual Review of Ecology, Evolution, and Systematics 35:583-621.

Whitman, M. 2006. Garlic Mustard: Odiferous Invader. Wild Ones Journal. http://www.for-wild.org/download/garlicmustard.pdf. Accessed [accessed 1 April 2012].

Williams, S.C. and Ward J.S. 2006. Exotic seed dispersal by white-tailed deer in southern Connecticut. Natural Areas Journal 26:383-390.

Winterer, J, Walsh, MC, Poddar, M, 2005. Spatial and temporal segregation of juvenile and mature garlic mustard plants (Alliaria petiolata) in a central Pennsylvania woodland. American Midland Naturalist 153:209-216.

Yates E.D., Levia Jr. D.F., and C.L. Williams. 2004. Recruitment of three non-native invasive plants into a fragmented forest in southern Illinois. Forest Ecology and Management. 190:119-130.

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 Appendix

Figure 1. Various life stages of A. Petiolata. (A) Rosette. (B) Plant just beginning to flower. (C) Top of older plant with many seedpods (Schooly, 2005).

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Figure 2. Low growing cluster of A. Petiolata (Schooly, 2005).

Figure 3. Flowering A. Petiolata with mature leaves (Schooly, 2005).

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Figure 4. From left to right, Maisa Fumagalli and Jake Balkos perform transects to count the size of a garlic mustard population at Longwoods Conservation Area. Note the space between garlic mustard populations A, B and C. Its possible that populations B and C are satellite populations of population A.

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