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Patterns of Nectar Robbing on Two Manzanita Species

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Patterns of Nectar Robbing on Two Manzanita Species Patterns of nectar robbing on two manzanita species Breana Dyste1, Avrodet Mourkus2, Kenny Ruiz1, Benjamin Vargas3 1 University of California, Davis; 2 University of California, Riverside; 3 University of California, Santa Barbara ABSTRACT Pollinators provide ecosystem services and facilitate plant reproduction. Plant-pollinator interactions often benefit both partners and are considered mutualisms. However, cheating can occur in these interactions when floral visitors take resources from plants without providing any benefit (Smithson 2013). One example of this kind of cheating is nectar robbing, which occurs when a pollinator collects nectar from a hole in the flower’s corolla and bypasses pollination. We compared the variation of floral nectar robbing between two species of manzanita, Arctostaphylos viscida (whiteleaf manzanita) and A. manzanita (common manzanita). We examined both buds and blooms to explore the relationship between age on the robbing behavior of floral visitors. We found no difference in the percent of nectar robbing between whiteleaf and common manzanita. There was no notable difference of nectar robbing between blooms and buds in common manzanita, however, there was nearly three times more nectar robbing on blooms than buds in whiteleaf manzanita. Studying nectar robbing behavior, as well as its impact on flowering plants, can help monitor successful and unsuccessful pollination. Keywords: mutualism, nectar robbing, common manzanita, whiteleaf manzanita, phenophase INTRODUCTION the coevolution of pollinators and plants (Chittka et. al 2006). Pollination aids in the sexual reproduction One aspect of coevolution between plants of plants and the evolution of floral and pollinators is mutualism, which is an morphology, which influences pollinator interaction that benefits both species preference (Kevan et. al 1990). Pollinator (Richardson and Bronstein 2012). An preferences can impact visitation rates to example of mutualism is between the Nile certain plant species and affect their crocodile and the Egyptian plover, where the reproductive output. For example, certain bird eats decaying meat from the crocodile’s species of bees prefer magenta-colored mouth, who in turn avoids oral infection. flowers to white flowers, causing white Another mutualism is between plants and flowers to set fewer fruits (Urbanowicz et. al pollinators. A plant’s flower gets pollinated, 2020). These selection pressures influence which increases its fitness, and in return changes in floral character, which leads to pollinators are rewarded with nectar. CEC Research | https://doi.org/10.21973/N3TH4F Winter 2021, Vol. 5 Issue 1 1/6 However, temporary changes in pollinator amount of nectar robbing seen on the behavior can give rise to exploitation, where sampled tree, as the pollinators have more they benefit themselves more than their options to choose from. mutualist partner. An example of this exploitation is nectar robbing, which occurs METHODS when a floral visitor collects nectar from a flower, through a hole made in the corolla, 2.1 Study System and Natural History and therefore neither acquires nor The study was conducted at the distributes pollen in the process (Richardson McLaughlin Natural Reserve and adjacent and Bronstein 2012). Nectar robbing impacts property owned by the Bureau of Land both the flower, through a reduction of Management in Lake County California from pollen donation, and other pollinators, by February 24–27 2021. A complex geological influencing their choices of flowers to visit. history of tectonic plate movement, and In this study, we examined Arctostaphylos volcanic and hydrothermal activity, formed a viscida (whiteleaf manzanita) and A. mosaic of serpentine and sedimentary soils. manzanita (common manzanita), which are These features support diverse plant targets of exploitation from pollinators communities including oak woodland, through nectar robbing. Whiteleaf chaparral, and grassland. Whiteleaf and manzanita is endemic to serpentine soil and common manzanita are associated with the is an obligate seeder, depending on a seed chaparral community, but isolated by soil bank for regeneration after a fire (Duren and type. Whiteleaf manzanita is endemic to Muir 2010). Conversely, common manzanita serpentine soil, while common manzanita is does not tolerate serpentine soil and is a found on non-serpentine soils (O’dell et al. facultative seeder, able to resprout and 2006; Grace et al. 2007). Both species of germinate post-fire (O’dell et. al 2006). On manzanita are evergreen, grow to a height of whiteleaf and common manzanita, we nearly 6 m, and share similar phenological focused on two phenophases or stages in a flowering periods (Richardson and Bronstein plant’s annual life cycle. We defined these 2012). In late winter, trees bloom with white phenophases as buds (unopened flowers) to pink, urn-shaped flowers arranged in and blooms (opened flowers). clusters (Eliyahu et al. 2015). Manzanitas Since whiteleaf manzanita is an obligate attract a wide variety of floral visitors seeder, it presumably invests more energy in including hummingbirds, butterflies, bees, nectar production to attract pollinators, so wasps, moths, and flies (Richardson and we hypothesized that it would experience Bronstein 2012). more robbing (Manetas 2000). We also hypothesized that buds in whiteleaf and 2.2 Flower Inspection for Evidence of Nectar common manzanita would have more nectar Robbing robbing holes because they would be more appealing to pollinators, as seen in Bombus Whiteleaf and common manzanita were spp. (bumblebee) who was found to favor identified flowering at four locations across young flowers (Willmer et. al 1994). We also the study area. Two sites, Devilhead and hypothesized that a higher number of Southbound, hosted whiteleaf manzanita on flowering neighbors would decrease the serpentine soil. Common manzanita was CEC Research | https://doi.org/10.21973/N3TH4F Winter 2021, Vol. 5 Issue 1 2/6 present at the Campground and Northbound RESULTS sites on non-serpentine soil. At each site, five trees of a given species were sampled with a We collected a total of 1,200 buds and minimum distance of 50 m between 1,200 blooms from whiteleaf and common individuals, to ensure independence. On manzanita. We found that nectar robbing each manzanita tree, we haphazardly did not differ between manzanita species (N collected 60 flowers of each phenophase, = 2400, F = 1.47, p = 0.349; Fig. 1). There was bud, characterized by closed petals, and no pronounced difference in nectar robbing bloom, characterized by open petals. Each between blooms and buds of common flower was inspected for nectar robbing. manzanita, however, there was nearly three Unrobbed flowers were defined by the times more nectar robbing on blooms than absence of holes, whereas robbed flowers buds of whiteleaf manzanita (N=2400, were categorized as having 1, 2, or 3 holes. If F=19.02; Fig. 1). Notably, blooms from flowers were eaten through such that there whiteleaf manzanita represented almost were no distinct holes, they were half of all nectar robbed flowers. categorized as 3+. The number of flowering Furthermore, we found a significant manzanita within a 10 m radius of the interaction of flower phenophase and sample tree was also recorded. manzanita species on nectar robbing, such that nectar robbing differs with flower 2.3 Insect Collection and Species Richness phenophase, but only in whiteleaf manzanita (N = 2400, F = 15.78, p = 0.0004; Fig. 1). On February 24, we collected insects present on common manzanita using a sweep net method. Individuals were captured and dispatched with acetone in a sealed jar. The specimens were later classified to the most detailed taxonomic rank as possible using iNaturalist (v3.1). The collection was preserved for future reference and study at McLaughlin Natural Reserve. Due to inclement weather, we were unable to adequately survey the species richness of Figure 1. Percent of nectar robbed flowers on common vs. white-leaf manzanita trees. Buds (N = insects on whiteleaf manzanita. 600) and blooms (N = 600) were sampled from 10 common and 10 whiteleaf manzanita trees. Buds 2.4 Statistical Analysis were identified as closed flowers and blooms as opened flowers. Nectar robbing is defined as one or Statistical analysis was conducted using more holes in the corolla, which was calculated as a JMP statistical software v14.1. We ran a two- percentage for each manzanita species. On common way mixed model ANOVA to determine the manzanita the percent of robbed blooms was similar to buds. However, on whiteleaf manzanita, a higher effects of manzanita species and floral percent of blooms was robbed than buds. Error bars phenophase, as well as their interaction, on represent ±1 S.E. the percent of nectar robbed flowers. In this model, site was considered a random effect. CEC Research | https://doi.org/10.21973/N3TH4F Winter 2021, Vol. 5 Issue 1 3/6 The Southbound Road site accounted for manzanita are obligate seeders that 54.2% of all robbed flowers, which included presumably invest more resources into buds and blooms (Fig. 2). In contrast, the nectar production. Therefore, the flowers of percent of nectar robbed flowers at the whiteleaf manzanita may be more attractive other three sites, including whiteleaf and to nectar robbers than those of common common manzanita, only ranged from 18.3% manzanita which are facultative seeders. to 20% (Fig. 2). According to our results, Whiteleaf manzanita at the Southbound 51.7% of the variance in
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