Modeling the Life History of Sessile Rotifers: Larval Substratum Selection Through Reproduction
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Modeling the life history of sessile rotifers: larval substratum selection through reproduction Andrea N. Young, Rick Hochberg, Elizabeth J. Walsh & Robert L. Wallace Hydrobiologia The International Journal of Aquatic Sciences ISSN 0018-8158 Hydrobiologia DOI 10.1007/s10750-018-3802-x 1 23 Your article is protected by copyright and all rights are held exclusively by Springer Nature Switzerland AG. This e-offprint is for personal use only and shall not be self- archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Hydrobiologia https://doi.org/10.1007/s10750-018-3802-x (0123456789().,-volV)(0123456789().,-volV) ROTIFERA XV Modeling the life history of sessile rotifers: larval substratum selection through reproduction Andrea N. Young . Rick Hochberg . Elizabeth J. Walsh . Robert L. Wallace Received: 14 July 2018 / Revised: 13 October 2018 / Accepted: 15 October 2018 Ó Springer Nature Switzerland AG 2018 Abstract Although the theoretical underpinnings of cohort of larvae. Our Basic Model assessed fitness as habitat selection by marine invertebrate larvae have simply settling on a substratum using only the larval been well studied, this theory has been neglected for elements and revealed statistically greatest fitness freshwater sessile rotifers. To study how substratum when swimming speed decreased with age and when selection affects larval fitness, we developed a substratum preference was constant or exhibited mid- dynamic model to examine influences of three age competence. The Reproductive Model assessed elements of larval life (survival, substratum accep- fitness (mean number of offspring adult-1)asa tance, substratum encounter probability) and substra- function of substratum. We compared reproduction tum-dependent reproductive success in adults. Monte on neutral substrata to substrata where quality varied Carlo simulation models were run using an initial and separately as a function of adult population density: coloniality (synergism) versus competition. The model showed fitness was statistically greatest Guest editors: Steven A. J. Declerck, Diego Fontaneto, Rick Hochberg & Terry W. Snell / Crossing Disciplinary when larval swimming speed decreased with age and Borders in Rotifer Research when coloniality increased reproduction. We also explored conditions where populations of planktonic Electronic supplementary material The online version of adults could survive. The model is applicable to sessile this article (https://doi.org/10.1007/s10750-018-3802-x) con- tains supplementary material, which is available to authorized organisms and may be modified to examine other life users. A. N. Young R. L. Wallace (&) Department of Mathematical Sciences, Ripon College, Department of Biology, Ripon College, Ripon, WI 54971, Ripon, WI 54971, USA USA e-mail: [email protected] R. Hochberg Department of Biological Sciences, University of Massachusetts at Lowell, Lowell, MA 01854, USA E. J. Walsh Department of Biological Sciences, University of Texas at El Paso, El Paso, TX 79968, USA 123 Author's personal copy Hydrobiologia history activities including selection of mates, prey, or et al., 2004; Kinlan et al., 2005; Toonen & Tyre, 2007; territory. Burgess et al., 2012). However, the life history characteristics of sessile rotifers (Monogononta: Gne- Keywords Behavior Á Dynamic modeling Á siotrocha: Atrochidae, Collothecidae, and Flosculari- Gnesiotrocha Á Planktonic Á Substratum-dependent idae) differ from those of marine invertebrates. For reproduction Á Rotifera example, marine invertebrates often release large numbers of relatively long-lived (days, weeks, months) larvae that may disperse over short Larvae … are indeed not helpless, for they are endowed with the power of choice, and with a period of time during which that choice (* 1 m) to long ([ 750 km) distances and have may be made. — Wilson (1952). extended development times (days to months) (Kempf, 1981; Hadfield, 1998; Shanks, 2009). They also exhibit extreme diversity in size (* 200 to [ 3000 lm) and form (Levin & Bridges, 1995), and as a group use varied energy sources: endosymbiotic Introduction autotrophy, lecithotrophy, osmotrophy, and plank- totrophy, as well as poecilogony (Vance, 1973; Understanding a species requires comprehending its McEdward, 1995; Chia et al., 1996; Allen & Pernet, entire life history, not just the morphological and 2007). Additionally, during their free-swimming ecological features attendant to the adult (Werner & phase, marine larvae usually experience an extensive, Gilliam, 1984; Werner, 1988). Undeniably many relatively uniform, open-water existence, where suit- studies have documented ontogenetic (developmental) able substrata are generally not encountered. As a shifts; these include shifts in diet (e.g., polychaetes: consequence many, but certainly not all, marine larvae Hentschel, 1998; harpacticoid copepods: Decho & are capable of postponing settlement in the absence of Fleeger, 1988; fishes: Daly et al., 2009; amphibians: suitable habitat (Pechenik, 1980). In contrast, sessile Schriever & Williams, 2013), behavior (Despland & rotifers release relatively few, short-lived (hours to Hamzeh, 2004), physiology (Woods & Wilson, 2013), days) larvae that exhibit little variation in size (ca. and habitat use (Richards, 1992) (see de Roos & 400 lm) or form (Wallace et al., 2015), and which Persson, 2013). appear to be lecithotrophic or possess limited ability to Interpreting the life history of sessile invertebrates feed (Wallace et al., 1998; Hochberg, 2014; Hochberg is no different; it requires an understanding of their et al. (in this volume). Moreover hatchlings immedi- distinct bipartite existence: larval life is transitory, ately encounter a mosaic of habitats dominated by ending either in death or with a cascade of searching diverse hydrophytes that provide a rich landscape for and settling behaviors culminating in attachment and settlement with variable morphologies and chemis- subsequent metamorphosis into an adult (Crisp & tries that often differ over short distances (Meksuwan Meadows, 1963; Wallace, 1980; Pawlik, 1992; et al., 2014). Burgess et al., 2009). Accordingly, a successful larva Free-swimming larvae may provide sessile rotifers must survive planktonic life, encounter at least one several advantages including (1) colonization of a substratum, attach, metamorphose to adulthood, and preferred substratum, including new hydrophyte reproduce. Clearly, if settlement is irreversible, choice growth that has not been weakened by herbivores, of a high-quality substratum becomes critical to pathogens, or age; (2) establishment of a colony or maximize fitness (Hodin et al., 2015). Thus, one may alternatively avoidance of competition; and (3) wider infer that sessile invertebrates must make evolutionary dispersal that may reduce inbreeding. In contrast, tradeoffs that are not intuitively obvious. disadvantages include (1) death in the plankton due to Undeniably there is a rich literature on the life starvation and/or predation; (2) delayed metamorpho- history of most marine invertebrate larvae (Chia & sis with subsequent inability to complete it; and (3) Rice, 1978; Young, 1990; McEdward, 1995; Hadfield, poor adult reproduction due to selection of an inferior 1998), including conceptualizing the process of sub- habitat (Pechenik, 1999). stratum selection (Doyle, 1975; Roughgarden et al., To better understand the life history of sessile 1985; Strathmann, 1985; Rumrill, 1990; Marechal rotifers, we developed a dynamic model with the 123 Author's personal copy Hydrobiologia capacity to vary environmental and sessile rotifer life with the potential for reproduction. Components of the history features including (a) relative availability of model are fully described in Supplemental Document. potential substrata, (b) cohort size, (c) planktonic In the basic model (BM), we defined fitness simply as survival, (d) substratum choice, and (e) adult survival settling and metamorphosing, with the assumption that and subsequent reproduction. Here we present our metamorphosis leads to equal reproduction on all model, explore some of its results, offer testable hy- substrata (Doyle, 1975). In the reproductive model potheses regarding sessile rotifer life histories based (RM), we defined reproductive fitness as metamor- on outcomes of model runs, and suggest areas where phosing with subsequent reproduction, which could additional research is needed. vary as a function of substratum or density of settled larvae. The RM followed the behavior of the original population and one generation of offspring. We also Materials and methods explored conditions where adults could survive in the plankton. Following, we provide a brief overview of Model development the environmental and life history functions (LHF) that we incorporated into the model (Table 1). Our model reflects ideas from several previously LHF 1 Larval survival in the plankton. Larval published models of sessile animals (Doyle, 1975; survival was set such that during each time