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Thesis Draft Rough Wesleyan University The Honors College Plant-pollinator interactions across California grassland and coastal scrub vegetation types on San Bruno Mountain, San Mateo County by Miles Gordon Brooks Class of 2020 A thesis submitted to the faculty of Wesleyan University in partial fulfillment of the requirements for the Degree of Bachelor of Arts with Departmental Honors from the College of the Environment Middletown, Connecticut April, 2020 1 2 Abstract Animal pollination of plants is a crucial ecosystem service for maintaining biodiversity and ecosystem function, worldwide. High pollinator abundance and diversity can likewise improve the reproductive success of the plant community. Plant-pollinator interaction networks have the potential to identify dominant, specialist, and generalist pollinator species within a system, and their host plant counterparts. Understanding these relationships is paramount for buffering natural systems from biodiversity loss in a world where pollinator abundance continues to decline rapidly. San Bruno Mountain (SBM) in San Mateo County, California, is one of the last natural, open spaces in the urban landscape in the northern San Francisco Peninsula. I conducted a series of timed meanders and vegetation surveys at eight sample sites within SBM (four grassland and four coastal scrub sites) to identify plant species prevalence and pollinator species visitation of flowering plants. I employed a multivariate approach for investigating plant and pollinator species richness, plant and pollinator community composition, and trophic-level interactions across the SBM landscape, and I evaluated differences in these relationships between grassland and coastal scrub habitats. A total of 59 pollinator species and 135 plant species were inventoried over the course of the study. While species richness did not vary significantly between vegetation types, the nonmetric multidimensional scaling results revealed significant differences in species composition and key indicator species between vegetation types. The bipartite analyses identified Bombus vosnesenskii, Eriophyllum stachaedifolium, Grindelia hirsutula and others as generalist pollinator and plant species that are 3 important for the long-term biodiversity conservation of SBM due to their interactions with a diverse array of other plant and pollinator taxa. In the future, adaptive restoration activities could be used at SBM and other similar habitats to bolster the abundance of herbaceous flowering plants for pollinators to conserve biodiversity and promote ecosystem health in a world that continues to experience declines in pollinator abundance. 4 Introduction Bees and other wild pollinators provide an important ecosystem service to agriculture crops and natural environments (Kearns et al. 1998, Aguilar et al. 2006, Klein et al. 2007, Kremen et al. 2007, Tuell et al. 2008, Potts et al. 2010). Pollination is the act of transferring pollen grains from a plant’s male reproductive parts, the anthers, to a plant’s female parts, the stigma or pistil (Meeuse 2018). Pollination is often facilitated by animals, who feed on the nectar and pollen of flowering plants. Approximately 35% of crops and 80% of native plants rely on animal pollination for reproduction (Klein et al. 2007, Potts et al. 2010). As such, pollination maintains biodiversity, facilitates the plant community’s reproductive health, and provides ecosystem services to ecosystems, worldwide (Waser et al. 1996, Kearns et al. 1998, Aguilar et al. 2006, Greenleaf and Kremen 2006, Klein et al. 2007, Kremen et al. 2007, Black et al. 2009, Potts et al. 2010, Elle et al. 2012, Smith DiCarlo et al. 2019). High pollinator diversity facilitates plant community diversity and ecosystem function (Aguilar et al. 2006, Fontaine et al. 2006, Kremen et al. 2007, Tuell et al. 2008, Blüthgen and Klein 2011). Many types of pollinators exist, but bees make up a significant proportion of pollinators around the world (Potts et al. 2003). They are one of the most diverse groups of insects, with an estimated 20,000 species, globally (Michener 2007, Smith DiCarlo et al. 2019). Other types of pollinators include butterflies, wasps, birds, and flies. These pollinator species have different plant preferences that allow them to fill complementary niches to bees (Fontaine et al. 2006, Blüthgen and Klein 2011, Venjakob et al. 2016, 5 Iwasaki et al. 2018). Greater pollinator diversity benefits plant community productivity because high pollinator diversity increases chance of plant pollination (Fontaine et al. 2006, Greenleaf and Kremen 2006, Tuell et al. 2008, Potts et al. 2010). Higher plant species richness provides a feedback for pollinator species richness because it boosts resource availability, which attracts a wider range of pollinators taxa to an area (Potts et al. 2003, Ebeling et al. 2008, Carman and Jenkins 2016, Maia et al. 2019). Greater pollinator diversity increases plant visitation frequency and the pollinator population’s resilience to perturbation, such as extinction of a plant species (Memmot et al. 2004, Fontaine et al. 2006, Ebeling et al. 2008, Tylianakis et al. 2010, Elle et al. 2012). Plant-pollinator interactions and richness are therefore crucial ecosystem health and integrity metrics. Recent global declines in wild pollinator abundances threaten community health, on scales spanning from local to global (Kearns et al. 1998, Aguilar et al. 2006, Klein et al. 2007, Kremen et al. 2007, Williams et al. 2009, Winfree et al. 2009, Potts et al. 2010, vanEngelsdorp et al. 2017). The loss of pollinator diversity has stimulated new interest in understanding the diversity and distribution of pollinator species across a range of habitats and land cover types (Grundel et al. 2010, Colteaux et al. 2013, Meiners 2016, Campell et al. 2018, Luong et al. 2019, Smith DiCarlo et al. 2019). Climate change, pesticide use, and the increased prevalence of pests are responsible for bee population decline (Kearns et al. 1998, Black et al. 2009, Potts et al. 2010, vanEngelsdorp et al. 2017). Habitat loss and fragmentation comprise other anthropogenic stressors on 6 wild pollinator populations (Aguilar et al. 2006, Fortuna and Bascompte 2006, McFrederick and LeBuhn 2006, Kremen et al. 2007, Goulson et al. 2008, Black et al. 2009, Winfree et al. 2009, Carman and Jenkins 2016). Bee communities depend on the ecosystem integrity (Kremen et al. 2007, Grundel et al. 2010, Ullmann et al. 2010), and the plant community depends on the presence of pollinators (Aguilar et al. 2006, Fontaine et al. 2006, Greenleaf and Kremen 2006, Klein et al. 2007). Further pollinator population decline could lead to further decline in biodiversity in the vegetation community (Memmot et al. 2004, Klein et al. 2007, Lundgren et al. 2016, Venjakob et al. 2016). While a myriad of negative effects from pollinator decline are predicted for the future, understanding contemporary plant-pollinator relationships in natural areas that span different habitats and plant cover types represents a key step for predicting the effects of future global change on pollinator population dynamics. Bipartite networks, networks that consist of interactions between two trophic levels, are useful ways to assess and conserve the mutualistic relationship between plants and pollinators (Blüthgen et al. 2008, Dormann et al. 2009, Bascompte 2010, Tylianakis et al. 2010, Elle et al. 2012, Carman and Jenkins 2016, Petanidou et al. 2018, Maia et al. 2019). Unlike traditional measures of conservation that only measure the abundance of a key species or community species richness, plant-pollinator networks capture interactions and their frequency between the two trophic levels (Tylianakis et al. 2010, Elle et al. 2012). In plant-pollinator networks there tend to be a few well-connected, generalist species that promote diversity by interacting lots of species (Waser et al. 1996, 7 Lopezaraiza-Mikel et al. 2007, Maia et al. 2019). Identifying key generalist plants and pollinators through bipartite networks allows conservation managers to determine important species for conservation to maintain ecosystem function and promote resilience to extinction (Memmot et al. 2004, Maia et al. 2019). Understanding the mutualistic interactions between plant and pollinators is paramount for buffering natural systems from biodiversity loss in a world where pollinator abundance continues to decline rapidly. San Bruno Mountain (SBM) in San Mateo County, California is an important site for conservation because it is an ecological ‘island’ surrounded by an urban matrix (Figure 1). The mountain has been protected by the San Bruno Mountain Habitat Conservation Plan since 1982 (Ormshaw 2018), and has been called one of the most important and threatened biodiversity sites in the world (Wilson 1999). SBM hosts both northern coastal scrub and grassland plant cover types, and a rich pollinator community. A lack of disturbance from wildfires, which were common on the mountain before European colonization, has resulted in widespread scrub encroachment and a subsequent decline of grassland community cover in the absence of fire (from 793 hectares of grassland in 1932 to 478 in 2014) (Weiss et al. 2015). Fire frequency is a well-known influence on the abundance and distribution of both woody and herbaceous vegetation in coastal habitats, where a lack of fire often results in shrub dominance at the expense of graminoid-dominated communities (Moyes et al. 2005, Zavaleta and Kettley 2006, Knapp et al. 2007, NPS 2007, Eviner 2016, Carlsen
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