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Canopy Effects on Abundance and Leaf Traits of a Spring Ephemeral: Erythronium Americanum

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Canopy Effects on Abundance and Leaf Traits of a Spring Ephemeral: Erythronium Americanum Botany Canopy effects on abundance and leaf traits of a spring ephemeral: Erythronium americanum Journal: Botany Manuscript ID cjb-2019-0083.R2 Manuscript Type: Article Date Submitted by the 28-Aug-2019 Author: Complete List of Authors: Greco, Danielle; Algoma University , Department of Biology Schamp, Brandon; Algoma University , Department of Biology Mercer, Kirstin; Algoma University , Department of Biology leaf-out, hard canopy, hemispherical photographs, specific leaf area, Keyword: Draft structural equation modeling Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? : https://mc06.manuscriptcentral.com/botany-pubs Page 1 of 25 Botany Canopy effects on abundance and leaf traits of a spring ephemeral: Erythronium americanum Danielle A. Greco, Brandon S. Schamp* & Kirstin A. Mercer Dept. of Biology, Algoma University, Sault Ste.Draft Marie, Ontario, Canada *Corresponding Author Email: [email protected] Phone: 705.949.2301 ext. 4358 Fax: 705.949.6583 1 https://mc06.manuscriptcentral.com/botany-pubs Botany Page 2 of 25 Abstract Spring ephemerals take advantage of high light available in the spring by completing the aboveground portion of their lifecycle before the canopy develops and while few other understory plant species are growing. The spring is marked by high resource availability, yet spring ephemerals are variably abundant throughout forests. Research indicates that canopy conditions can influence spring ephemeral growth; consequently, we tested whether variation in canopy conditions predicted variation in the abundance of Erythronium americanum across 50 forest plots. We also tested whether the specific leaf area (SLA) of E. americanum in plots was predicted by variation in plot-level Draftcanopy conditions, reflecting E. americanum’s ability to adapt to different canopy conditions. E. americanum abundance was significantly lower in plots with greater hard canopy closure (i.e., permanent cover: tree architecture + evergreen leaf cover) and significantly higher under canopies that reached full development earlier. Canopies with greater hard canopy cover at the start of the growing season were associated with significantly higher SLA, quantifying local adaptation by E. americanum to variable canopy conditions. E. americanum takes advantage of high light available in the spring. It is unclear at this time why higher E. americanum abundance is associated with earlier closing canopies. Keywords: leaf-out; hard canopy; hemispherical photographs; specific leaf area; structural equation modeling 2 https://mc06.manuscriptcentral.com/botany-pubs Page 3 of 25 Botany Introduction Spring ephemerals complete the aboveground portion of their lifecycle within spring, prior to leaf-out in the forest canopy. These plants are common in temperate forests where the winter- to-spring transition results in unique conditions of high soil and light resources. Snow melt results in high levels of soil moisture (Likens et al. 1977), soil nutrients, as organic matter from the previous autumn breaks down, and as low temperatures delay uptake by trees (Chapin 1980; Muller 2003). Furthermore, because summer-growing plants, which are more tolerant of shade, are not yet developed, spring ephemeral species experience relatively limited competition for many resources (Muller and Bormann 1976). Throughout much of the growingDraft season, light reaching the forest understory is limited, conducive only to the growth of shade-tolerant understory species. Light levels reach their highest point in the spring for a short period when the solar angle is high and the canopy is leafless (Hutchison and Matt 1976). This is the only period when the understory environment supports shade-intolerant spring ephemerals. As spring continues and temperatures warm, trees undergo leaf-out until canopy development peaks and light availability at the forest floor is greatly reduced. There are also early growing species that flower prior to leaf-out but continue to photosynthesize later into the summer; however, here we focus on a true spring ephemeral that senesces as the canopy develops and shade increases. Although Erythronium americanum Ker Gawl. grows during a period of relative high light, plentiful soil resources, it is not consistently abundant throughout the forest. Indeed, local abundance of this species varies widely within and among forests. It is currently unknown what drives this variation. Several factors may limit E. americanum’s abundance in different 3 https://mc06.manuscriptcentral.com/botany-pubs Botany Page 4 of 25 environments. For example, herbivores, dispersers, and pollinators may play a role. However, abundance varies strongly even within relatively small forests where the distribution of dispersers and pollinators are unlikely to be variable enough to account for relatively large observed differences in abundance. Additionally, our chosen vegetation plots did not have signs of herbivory. There is little evidence that deer browsing has an important influence on this species (Collard et al. 2010), although it may play a role. However, canopy conditions that contribute to different schedules of canopy closure appear to vary considerably within even small forests because of compositional differences. Canopies vary in the presence of evergreen tree species, which determines baseline shade to the forest floor, as well as in the presence and abundance of deciduous tree species thatDraft vary in leaf-out phenology (Dion et al. 2017). Consequently, we hypothesize that variation in the timing of leaf-out (start and end) limit the success of spring ephemerals that cannot acclimate to increased shade (Eickmeier and Schussler 1993; Dion et al. 2017). While the role of canopy variation in limiting E. americanum abundance has not been studied, evidence does suggest that this species is sensitive to canopy conditions. Marchand and Houle (2006) found that E. americanum patch sizes were greater in the interior of an Acer saccharum stand in southern Quebec, where canopy gaps were generally larger and provided less cover than along the edge. Taylor and Pearcy (1976) have noted that shaded E. americanum plants senesce earlier than those with prolonged gap exposure. These two studies support the influence of canopy variation on E. americanum. Collins and Pickett (1982) found no significant difference in E. americanum cover (i.e. abundance) across sites of varying canopy cover in the Kane Experimental Forest of western Pennsylvania. In situations where the canopy 4 https://mc06.manuscriptcentral.com/botany-pubs Page 5 of 25 Botany has been removed or canopy closure is delayed, senescence of E. americanum occurred as normal (Muller 1978) or only 5-7 days later than normal (Hughes 1992), indicating that factors beyond the canopy also contribute to senescence. Experimental work has demonstrated that early shade treatments can result in earlier senescence in a congener, E. japonicum (Kim et al. 2015). Finally, evidence suggests that other spring ephemeral species grow and/or reproduce better in response to delays in canopy closure (Trillium erectum L.: Routhier and Lapointe 2002; Allium tricoccum Ait.: Dion et al. 2017; Gagea lutea (L.) Ker Gawl: Sunmonu and Kudo 2014). Overall, evidence indicates that light conditions can influence the growth and success (i.e., local abundance) of spring ephemerals in general, and E. americanum specifically, under natural conditions. However, the above-mentionedDraft studies have not investigated whether local abundance is influenced by within-forest canopy variation. If canopies experience delayed leaf- out, or close earlier and shorten the window that ephemeral species have for growth and resource accumulation, variation in canopy structure and closure timing may account for observed spatial variation in spring ephemeral population density. Specific leaf area (SLA, in mm2/mg ), a functional trait that represents the leaf area per unit of dry mass investment by a plant , captures important information related to how plants deal with environmental variation related to light and moisture (Milla et al. 2008; Cornwell and Ackerly 2009). Research indicates, for example, that SLA, both within and among species, changes along light and moisture gradients (e.g., Cornwell and Ackerly 2009; Long et al. 2011). At least one study has found that E. japonicum plants have lower SLA under conditions of earlier canopy closure (Kim et al. 2015). Consequently, understanding how light impacts the 5 https://mc06.manuscriptcentral.com/botany-pubs Botany Page 6 of 25 abundance of a species requires that we also account for that species’ ability to acclimate to variation in these conditions via intraspecific variation in SLA. A dynamic canopy is a defining characteristic of the spring forest environment. It supports the establishment of spring ephemerals, but in spite of generally high resource availability in the spring, these species vary significantly in abundance throughout temperate forests. In this paper, we test whether variation in the abundance and SLA of the spring ephemeral E. americanum is influenced by local variation in spring canopy characteristics that alter the timeframe over which this species can effectively harvest light energy. Materials and methods Draft Study sites A total of 50 circular plots were established in April 2015 within the Hiawatha Highlands Conservation Area in Sault Ste. Marie, Ontario, Canada, which encloses 892 hectares of mixed coniferous-deciduous forest. Plots
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