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The Scalesia Genus As an in Situ Model for Evolutionary Theory

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The Scalesia Genus As an in Situ Model for Evolutionary Theory The Scalesia Genus as an In Situ Model for Evolutionary Theory Title Image. A young Scalesia pedunculata plant at the Darwin Research Station on Santa Cruz (Birnbaum, 2018). Foster Birnbaum Evolution and Conservation in the Galapagos Professor Bill Durham 10/12/2018 Foster Birnbaum Evolution and Conservation in the Galapagos 10/12/2018 Abstract The towering Scalesia plants once formed the Galapagos’ most verdant environments: on the tops of otherwise arid islands rose lush Scalesia forests. These forests were even more unique given that Scalesia trees are, in fact, daisies. In total, fifteen Scalesia daisies grow throughout the archipelago, four of which are highland trees; the rest are arid- or transitional-zone shrubs. Recognizing the Scalesia genus’s high species count, researchers consider it a paragon of adaptive radiation (Stöcklin 2009: pp.33-48). This paper further uses the Scalesia genus to explore tenets of evolution. To do so, this paper examines three hypotheses: (1) geographic distance affects the distribution of variation in the bush Scalesia affinis; (2) different quantities of pollinators explain why only some Scalesia species have ray florets; and (3) severe El Niño events exert a selective pressure on the tree Scalesia pedunculata to grow and die in generational cohorts. Genetic data on seven S. affinis populations support the first hypothesis; experimental data on how ray florets affect reproductive success support the second hypothesis; and observational data on S. pedunculata groves demonstrate the insufficiency of the third hypothesis. In addition, this paper discusses how S. pedunculata’s generational lifecycle affects efforts to conserve the species. Introduction The Galapagos are most well-known as the birthplace of not a living organism but an idea: Darwin’s theory of speciation by natural selection, also known as Darwinian evolution. Recognizing the ongoing potential for the flora and fauna of the Galapagos to advance evolutionary principles, this paper proposes the Scalesia genus, an endemic genus in the 1 Foster Birnbaum Evolution and Conservation in the Galapagos 10/12/2018 Asteraceae (daisy) family, as a model for Darwinian conceptions of variation and natural selection. Genetic analysis of chloroplast DNA restriction sites suggests that Scalesia’s closest relative is the Pappobolus genus of the Andes Mountains, and the number of differences in the sites suggests that the genera diverged around 2 million to 6 million years ago (Schilling et al. 2004: pp.248-254). Since its arrival in the Galapagos, Scalesia has grown to include fifteen species. Eleven are shrubs that survive in the arid zone, and four are trees—two of which (S. pedunculata and S. cordata) stand over 10 meters—that grow in higher-elevation, moist environments. Regarding the distribution of the fifteen species, older, larger islands harbor more species (e.g., Santa Cruz and San Cristóbal have six and four, respectively, each including S. pedunculata) than do younger, smaller islands (e.g., Wolf and Pinta have one species of shrub each). Further, “all the species are nearly completely allopatric in distribution, with a wide distance between their individual ranges in the archipelago,” and few hybrid forms have been observed (Itow 1995: pp.17-30). In addition to differing in size, Scalesia species display marked differences in leaf appearance, as shown in Figure 1. These differences are at least somewhat related to habitat conditions (Stöcklin 2009: pp.33-48). The distribution of and variation in Scalesia species make the genus an excellent candidate for evolutionary study. 2 Foster Birnbaum Evolution and Conservation in the Galapagos 10/12/2018 A: Scalesia cordata; B,C: Scalesia microcephala var. cordifolia; D: Scalesia microcephala var. microcephala; E: Scalesia pedunculata; F–H: Scalesia aspera; I: Scalesia villosa; J: Scalesia stewartii; K–M: Scalesia atractyloides var. atractyloides; N–P: Scalesia divisa; R: Scalesia divisa; S,T,V: Scalesia baurii ssp. hopkinsii; U: Scalesia baurii ssp. baurii; W: Scalesia affinis ssp. gummifera; X: intermediate between Scalesia baurii and Scalesia crockeri; Y: Scalesia crockeri; Z: Scalesia affinis ssp. brachyloba; AA–CC: Scalesia helleri; Figure 1. Leaf variations among Scalesia species DD: Scalesia retroflexa. (Stöcklin 2009: pp.33-48). Accordingly, this paper will analyze two Scalesia species, S. affinis and S. pedunculata. Table 1 summarizes several characteristics of each species. Characteristic S. affinis S. pedunculata Height shrub – grows up to 3 m tall tree – grows about 12 m tall (Figure 2B) (Figure 2C) Zone(s) of Residence arid and transitional highland – forms dense forests Islands of Residence Fernandina, Isabela, Santa San Cristóbal, Santiago, Santa Cruz, and Floreana Cruz, and Floreana (Figure 2A) (Figure 2A) Leaf Type pubescent (i.e., hairy) and pubescent and alternate alternate Ray Florets1 present (Figure 2D) absent Seed Dispersal Mechanism possibly land iguanas probably tree finches2 3 Foster Birnbaum Evolution and Conservation in the Galapagos 10/12/2018 Lifecycle seeds germination after about germination and growth occurs 20 days; subsequent growth quickly: plants grow 7 m, half occurs quickly their adult height, in their first two years Table 1. Characteristics of S. affinis and S. pedunculata. (Blaschke and Sanders 2009: pp.177-191; Durham 2016; Itow 1995: pp.17-30; Nielsen et al. 2002: pp.139-153; Traveset et al. 2016: pp.207-213) Figure 2. A. Range of S. affinis (green) and of S. penduculata (blue) (Blaschke and Sanders 2009: pp.177-191). B. A juvenile S. affinis plant. C. A juvenile S. pedunculata plant. D. The head of an S. affinis flower with white ray florets. (Birnbaum, 2018) In explaining how these species support Darwinian evolution, this paper attempts the following: 4 Foster Birnbaum Evolution and Conservation in the Galapagos 10/12/2018 1. to describe the variation in S. affinis populations—the only species for which data are available—on different islands; 2. to investigate why S. affinis flowers have ray florets while those of S. pedunculata do not; and 3. to explain the evolutionary history of the S. pedunculata lifecycle, characterized by the growth and eventual mass die-off of one generational cohort and the resulting growth of a new cohort. Hypotheses 1. The variation in S. affinis is based on geography (i.e., organisms that live closer together are more similar than those that live farther apart); 2. S. affinis developed ray florets while S. pedunculata did not because fewer pollinators are present in the arid and transitional zones than in the highlands; and 3. S. pedunculata’s generational lifecycle resulted from periodic, major climate changes (i.e., El Niño Southern Oscillation events). In addition, this paper addresses how the S. pedunculata lifecycle affects efforts to conserve the species, ranked vulnerable by the International Union for Conservation of Nature (Tye and Loving 1998). Thus, this paper will encourage further exploration of the Scalesia genus as a model for in situ evolutionary study and will provide advice on how to best protect S. pedunculata, a critical member of many island ecosystems. Part 1: Intraspecies Variation in S. affinis Variation forms the backbone of Darwin’s theory of evolution in On the Origin of Species. While the Galapagos mockingbirds catalyzed his doubts concerning the immutability of 5 Foster Birnbaum Evolution and Conservation in the Galapagos 10/12/2018 species, his observations of domesticated animals—especially pigeons—led to his conclusion that variation exists not only between species but also within species. This concept, in turn, helped him show the arbitrariness of labelling any group of organisms that share similar features as an absolute, distinct species; further, he advanced the idea that any organisms considered as separate species are merely morphologically distinct variations of a common ancestor. In this way, one “species” with two isolated sub-populations can, given enough time, become two “species”: i.e., a “species” can originate. As L.R. Nielsen’s 2004 genetic research demonstrates, S. affinis displays the intra-species variation that Darwin discusses in On the Origin of Species. Nielsen (2004: pp.434-442) analyzed the DNA of seven S. affinis populations—four from Isabella, two from Floreana, and one from Santa Cruz—using biparental and maternal markers. (Biparental markers are located in autosomal chromosomes, meaning they are affected by both parents’ genetic makeup; maternal markers are located in maternally inherited mitochondrial DNA.) The biparental data show the overall variation in S. affinis. Of the 286 biparental markers, only 129 (45%) were monomorphic (i.e., their DNA sequence was identical in greater than 98% of all individuals) (Nielsen 2004: pp.434-442). By contrast, a similar biparental analysis of Helianthus annuus, the common sunflower and another Asteraceae species, found 72% monomorphism (Gedil et al. 2000: pp.213-221). (The increased variation in S. affinis compared to its more widespread relative may be due to the Galapagos’ remoteness: evidence suggests that isolation increases the survival of mutant forms of an organism (Nielsen 2004: pp.434-442).) Genetic analysis of S. affinis thus reveals significant intraspecies variation. Further, the biparental markers reveal that the variation in S. affinis populations corresponds to their geographic proximity to each other, supporting the paper’s first hypothesis 6 Foster Birnbaum Evolution and Conservation in the Galapagos
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