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Reproductive Delays in Mammals: an Unexplored Avenue for Post-Copulatory Sexual Selection

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Reproductive Delays in Mammals: an Unexplored Avenue for Post-Copulatory Sexual Selection Biol. Rev. (2014), pp. 000–000. 1 doi: 10.1111/brv.12085 Reproductive delays in mammals: an unexplored avenue for post-copulatory sexual selection Teri J. Orr1,∗,† and Marlene Zuk2 1Department of Biology, University of California, Riverside, CA 92521, U.S.A. 2Ecology, Evolution and Behavior, University of Minnesota, 1987 Upper Buford Circle, Saint Paul, MN 55108, U.S.A. ABSTRACT Numerous mammalian taxa exhibit reproductive delays, pauses in reproduction that occur between mating and fertilization, between fertilization and implantation of the embryo, or after an embryo has implanted. Of the 27 mammalian orders, 9 are known to exhibit reproductive delays, including Diptrotodontia, Dasyuromorphia, Eulipotyphyta, Cingulata, Carnivora, Rodentia, Chiroptera, Lagomorpha and Cetartiodactyla. Most researchers interested in delays have focused on their evolutionary origins. However, the consequences of these delays have not been considered fully. Given the lengthening of the period over which reproduction occurs, it is possible that this unique aspect of reproduction facilitates post-copulatory sexual selection. When considered in the context of sexual selection, delays may allow sperm competition and female manipulation of fertilization (cryptic female choice) as well as other post-copulatory processes. We investigate the potential for reproductive delays to facilitate post-copulatory sexual selection and suggest avenues for research that may further our knowledge of sexual selection. We also provide a general review of reproductive delays in mammals. Key words: Carnivora, chiroptera, cryptic female choice, delayed fertilization, delayed implantation, embryonic diapause, mammalia, post-copulatory sexual selection, reproductive delays, sperm competition. CONTENTS I. Introduction ................................................................................................ 2 II. The distribution of reproductive delays ..................................................................... 3 III. Species with facultative delays .............................................................................. 3 IV. The evolution of delays ..................................................................................... 6 V. Sexual selection and delays ................................................................................. 8 (1) Delays between copulation and fertilization ............................................................ 9 (a) Predictions and experiments: delayed fertilization .................................................. 10 (2) Delays between fertilization and implantation ......................................................... 14 (a) Cryptic female choice via selective implantation, zygote comparison, and re-absorption ........... 14 (b) Predictions and experiments: delayed implantation ................................................. 14 (3) Delays between implantation and birth (delayed development) ........................................ 14 (a) Predictions and experiments: delayed development ................................................. 14 VI. Additional consequences of delays .......................................................................... 17 (1) Life history consequences of delays .................................................................... 17 (2) Post-copulatory sexual selection and the evolution of delays ........................................... 18 (a) Predictions and experiments: post-copulatory sexual selection and the potential evolution of delays 18 VII. Conclusions ................................................................................................ 19 VIII. Acknowledgements ......................................................................................... 19 IX. References .................................................................................................. 19 * Author for correspondence (Tel: 951-723-7647; E-mail: [email protected]). † Present address: Departments of Psychology and Biology, University of Massachusetts, Amherst, MA 01003, U.S.A. Biological Reviews (2014) 000–000 © 2014 The Authors. Biological Reviews © 2014 Cambridge Philosophical Society 2 Teri J. Orr and Marlene Zuk I. INTRODUCTION Macdonald, 2004), which has led to some understanding of the advantages of delays (see Section IV below) and The study of vertebrate reproductive patterns has been of their taxonomic distribution (Mead, 1993). However, the interest to biologists for centuries. Historically, researchers consequences of such delays, once they have evolved, remain have approached the investigation of reproduction from to be explored fully. Because reproductive delays lengthen an anthropocentric point of view, whereby they assumed the duration of a reproductive event from copulation to the timing of events from fertilization to birth occurred birth, they may provide additional time for post-copulatory in the same manner as in humans. Despite this common sexual selection to operate (Table 1). For example, delayed assumption, the diversity of mammalian reproductive fertilization increases the time frame during which sperm physiology has also been recognized for some time (Hamlett, and ejaculates from various males might interact (sperm 1935). Indeed, even before the publication of Darwin’s On the competition; Parker, 1970; Birkhead & Møller, 1993) or origin of species by means of natural selection in 1859, an unusual when females could manipulate and select sperm (cryptic reproductive physiology, reproductive delays, had already female choice; Thornhill, 1983; Eberhard, 1996). During been observed in mammals (Ziegler, 1843). delayed implantation, post-copulatory processes could select Extended periods of inactivity during a reproductive event among zygotes, allowing some to implant and others to be are not restricted to mammals. Other animals and even plants rejected. Finally, delayed development extends the period of are known to experience periods of reproductive diapause time over which an embryo is in the female’s reproductive between fertilization and offspring sprouting/hatching or tract (Bernard & Cumming, 1997), potentially facilitating its birth. For example, plants may have seeds that remain re-absorption or rejection (abortion). dormant (Bewley, 1997), female insects often store sperm In this review we first describe the distribution of (Pitnick, Markow & Spicer, 1999), and birds lay clutches of reproductive delays among mammals, briefly review the eggs asynchronously that often hatch synchronously, so that a literature on the evolution of delays and define the types given egg may undergo a delay in development (Ewert, 1992). of reproductive delays. Delays are then discussed in greater Delays are less common in viviparous animals. However, in detail in the order in which they might occur in a typical addition to mammals, some viviparous lizards (e.g. Sceloporus pregnancy: delayed fertilization, implantation and then jarrovii) exhibit slow embryonic development similar to the development. For each type of delay we ask what modes reproductive delays of some mammals (Ballinger, 1979). of sexual selection might be operating. After evaluating the The first documentation of this type of physiology in likelihood of sexual selection during each delay type (and mammals was of the roe deer (Capreolus capreolus) which corresponding time frame) we suggest experiments to test were known in 1843 to delay implantation for 4–5 months the hypothesis that delays facilitate post-copulatory sexual after fertilization (Hayssen, van Tienhoven & van Tienhoven, selection, and offer potential avenues for future research. 1993). Similar types of delays in fertilization, implantation or Finally, for each delay type, we review the data consistent development have since been described in many additional with these expectations (Table 2). Although we focus here on species of mammals. Well over 100 mammalian species, mammals, the predictions outlined in this review would be including members of the Diptrotodontia, Dasyuromorphia, relevant to any animal or plant with delay-like reproductive Eulipotyphyta, Cingulata, Carnivora, Rodentia, Chiroptera, asynchrony. Lagomorpha and Cetartiodactyla have some form of The existence of reproductive delays presents an ideal reproductive delay (Hamlett, 1935; Mead, 1993), and opportunity for asking questions about female versus this type of reproductive physiology may be even more male control of reproduction. Because specific stages of widespread but still undocumented in additional species. reproduction are lengthened in species with delays, these Delaying fertilization, implantation, or pausing development stages can be scrutinized for what may be evidence of (delayed development) may in turn alter the timing of sexual selection. For example, fertilization may primarily subsequent events including parturition. For example, in the serve as an arena for male competitive strategies, while long-tailed weasel (Mustela frenata), implantation is delayed delayed development may provide opportunities for females for 7–9 months while active gestation lasts an additional to manipulate events (and the embryos themselves), 9.5 months (Wright, 1942, 1963). The California leaf-nosed and implantation might be a period of extreme sexual bat (Macrotus californicus) has delayed development for an conflict. We believe delays are an under-appreciated average of 4.5 months, with active gestation lasting an source of natural variation in reproductive physiology, the
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