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Evolution of Life History in Three High Elevation Puya (Bromeliaceae), Leah Veldhuisen and Dr

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Evolution of Life History in Three High Elevation Puya (Bromeliaceae), Leah Veldhuisen and Dr 2 Evolution of Life History in Three High Elevation Puya (Bromeliaceae), Leah Veldhuisen and Dr. Rachel Jabaily, Organismal Biology & Ecology, Colorado College The Andes are known as a hotspot for biodiversity and high species endemism for both plants and animals. Two important tropical, high-elevation ecosystems in the Andes are the puna in Peru, Bolivia and Chile between 7° and 27° South, and the páramo in Colombia, Venezuela and Ecuador between 11° North and 8° South; both are found at elevations above 3500 meters. The genus Puya (Bromeliaceae) is found throughout the puna and the páramo, and is relatively under-studied. Life history of most Puya species is largely unknown, with the notable exception of entirely semelparous Puya raimondii, which flowers once right before dying and does not reproduce clonally. Other species in the genus do reproduce clonally to varying degrees; their life history strategies have not been defined. Decreased cloning ability in Puya may be evolving convergently as in other plant groups endemic to high- elevation, tropical ecosystems. We studied three species of Puya (P. raimondii, P. cryptantha and P. goudotiana) across the two ecosystems in Bolivia and Colombia, and collected data on threshold size at flowering and clonal reproduction. Data were also analyzed in conjunction with life history theory to hypothesize each species’ life history strategy. All three species were found to have a consistent and predictable minimum size at flowering, while P. cryptantha was found to also have a minimum size for clonal reproduction. No such evidence was found for P. goudotiana. Our data supported that P. raimondii is fully semelparous, and indicated that P. goudotiana and P. cryptantha may be semi-semelparous. Keywords: Bromeliaceae, evolution, life history, Andes 3 Acknowledgments: I would like to thank all the generous donors who made this project possible: the Keller Family Venture Grants and matching funds, the OBE Department funds, and the JH Enderson Research Award from Dr. Sharon Smith. Additionally, Dr. Rachel Jabaily, Dr. Jim Ebersole, Dr. Julián Aguirre-Santorro, Dr. Mary Carolina Garcia Lino and her family, many Bolivian students and professors, and everyone on Team Colombia who made the field research possible. Also thank you to Kate McGinn for data analysis and R help. 4 Introduction Life history theory Semelparity is a life history strategy that is found in a variety of annual plant species, as well as in the animal kingdom, but is rare in long-lived perennial plants. It occurs when an organism has one, large sexual reproductive effort at the end of its life, which results in death (Young & Augspurger, 1991). The single reproductive effort of semelparity requires a higher one time energetic input than in each of the repeated reproduction efforts of iteroparity, and can result in one higher reproductive output with larger inflorescence, more flowers, fruits, and seeds (Augspurger, 1985; Huxman & Loik, 1997; Young & Augspurger, 1991). Semelparity has been well-studied in some species (Barry, Unwin, Malison, & Quinn, 2001; Lazenby-Cohen & Cockburn, 1988; Young, 1984, 1985), and its evolution has been studied in some high elevation rosette plants such as the Lobelia genus that evolved convergently with other tropical alpine species (Schaffer & Schaffer, 1977, 1979; Schaffer & Rosenzweig, 1977; Young, 1990; Young & Augspurger, 1991). These studies resulted in three potential models for the selective pressures that favor semelparity. One model for semelparity evolution defines a tradeoff between the energy devoted to the one sexual reproductive effort and the elimination of energy for future survival and reproduction; all the plant’s transferable resources are dedicated completely to reproduction to maximize reproductive success (Young, 1990; Young & Augspurger, 1991). In this reproductive effort model, the increase in reproductive success must be an ever-increasing function of reproductive effort (Young, 1990). One example of this model is seen with pollinator preference: delayed flowering in semelparous species allows for enough of an increase in rosette and inflorescence size 5 to attract significantly more pollinators and expand seed set, thus negating the potential benefits of flowering earlier (Inouye & Taylor, 1980). There are two other models for the evolution of semelparity: the bet-hedging model and the demographic model. The bet-hedging model predicts that semelparity will be selected for in highly variable and unpredictable environments, although some evidence also suggests the opposite (Schaffer & Rosenzweig, 1977; Young & Augspurger, 1991). Unpredictable environments may favor semelparity over iteroparity, as annual and biennial life history (short- lived semelparous taxa) evolves repeatedly in plant lineages in unpredictable environments (Young & Augspurger, 1991). Long-lived semelparous rosette plants, such as Lobelia telekii, are common in drought-prone areas that also have highly variable temperatures (Young, 1990; Young & Augspurger, 1991). The demographic model considers the demographic conditions, such as population growth rate and age-class survivorship, when the loss of all future reproduction is made up for by success of one semelparous reproductive output (Young, 1990; Young & Augspurger, 1991). All three models suggest that semelparity will be selected for as the likelihood of repeated reproduction decreases. Iteroparous species also delay reproduction to minimize reproductive cost, but the reproduction is not lethal (Miller, Williams, Jongejans, Brys, & Jacquemyn, 2012). Size threshold background Both iteroparous and semelparous plants proceed through life cycle phases when enough energy has been acquired and stored to move forward, particularly into energetically costly processes like sexual and asexual reproduction (Lacey, 1986). While size and age are often closely correlated, some species reproduce based on a size threshold, while others base their life cycle phases on age (Lacey, 1986). Size in particular is thought to be a good predictor of a wide variety of species, and a consistent minimum size at reproduction has been determined in some species 6 (Lacey, 1986; Miller, Williams, Jongejans, Brys, & Jacquemyn, 2012b; Mora, Chaparro, Bonilla, & Vargas, 2005; Young, 1984). In semelparous species in particular, increased vegetative size is an important element of reproduction timing, as it has been shown to correlate with reproductive output and associated pollinator preference for larger inflorescences (Huxman & Loik, 1997; Rocha, Valera, & Eguiarte, 2005). Fecundity increases with size, often leading to plants reproducing at their largest possible size to maximize fitness. This trend is not true, however, for plants where increased size leads to increased mortality, predation or significantly limited growth, such as Lobelia telekii (Bonser & Aarssen, 2009; Young, 1985). L. telekii faces more herbivory at larger sizes, despite the fact that it is relatively large compared to other iteroparous species in the genus (Young, 1985). Variability in threshold reproductive size within a species can evolve from inconsistency in the environment or resource availability; plants in resource-limited or disturbed habitats will reproduce at half the size of plants in undisturbed habitats with readily available resources (Bonser & Aarssen, 2009). Because all environments will be at least somewhat variable, plasticity in threshold reproductive size is quite prevalent in almost all plant populations (Bonser & Aarssen, 2009). Both semelparous Lobelia telekii and iteroparous Lobelia keniensis are high elevation rosette plants on Mount Kenya, and have a minimum size at sexual reproduction (Young, 1990). Additionally, other annual, biennial and iteroparous plant species also have consistent minimum reproductive size (Kuss, Rees, Ægisdóttir, Ellner, & Stöcklin, 2008; Werner, 1975; Young, 1984). Similarly, Puya hamata has a relatively uniform rosette diameter of the vegetative body at the time of flowering (Garcia Meneses & Ramsay, 2014). In resource-limited habitats, the minimum size at reproduction will take longer to reach, and result in a longer life span, like that of long-lived semelparous plants (Young, 1984, 1985). For many plants, both the likelihood of survival and 7 flowering increase as plant size also increases (Bonser & Aarssen, 2009; Metcalf, Rose, & Rees, 2003). This trend holds true specifically in Puya dasylirioides with the number of mature fruits growing exponentially as rosette radii increases (Augspurger, 1985). Additionally, plant size alone is a good predictor of life history evolution since growth rate, chance of survival and reproduction, and reproductive output all depend on size (Augspurger, 1985; Metcalf et al., 2003; Werner, 1975). There is significantly less evidence for a size threshold in clonal or asexual reproduction, and evidence that does exist varies widely among genera and species (Ashmun & Pitelka, 1985; Mora et al., 2005; Schmid, Bazzaz, & Weiner, 1995; Young, 1984). In addition to size, plant reproduction timing is influenced by several factors. Age, and especially the interaction between age and size, environmental nitrogen levels, and photoperiod, among other things, play a significant role in when plants reproduce, although size alone is the most influential factor in scaling of reproductive output (Schmid et al., 1995; Song, Smith, To, Millar, & Imaizumi, 2012). Additionally, vernalization in combination with photoperiods and size are
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