Abstract

Hatching timing is under strong selection and environmentally cued in many species. Embryos use multiple sensory modalities to inform hatching timing and many have spontaneous hatching patterns adaptively synchronized to natural cycles. Embryos can also adaptively shift their hatching timing in response to environmental cues indicating immediate threats or opportunities. Such cued shifts in hatching are widespread among amphibians; however, we know little about what, if anything, regulates their spontaneous hatching. Moreover, in addition to selection on hatching timing, embryos may experience benefits or suffer costs due to the spatial orientation of hatching. Amphibian eggs generally lack internal constraints on hatching direction but embryos might, nonetheless, use external cues to inform hatching orientation. The terrestrial embryos of red-eyed treefrogs, Agalychnis callidryas, hatch rapidly and prematurely in response to vibrational cues in egg-predator attacks and hypoxia if flooded. Here we examined A. callidryas’ use of light cues in hatching timing and orientation. To assess patterns of spontaneous hatching and the role of light cues in their diel timing, we recorded hatching times for siblings distributed across three light environments: continuous light, continuous dark, and a 12L:12D photoperiod. Under a natural photoperiod, embryos showed a clear diel pattern of synchronous hatching shortly after nightfall. Hatching was desynchronized in both continuous light and continuous darkness. It was also delayed by continuous light, but not accelerated by continuous dark, suggesting the onset of dark serves as a hatching cue. We examined hatching orientation and light as a potential directional cue for flooded embryos. Embryos flooded in their clutches almost always hatched toward open water, whereas individual eggs flooded in glass cups often failed to do so, suggesting the natural context provides a directional cue. To test if flooded embryos orient hatching toward light, we placed individual eggs in tubes with one end illuminated and the other dark, then flooded them and recorded hatching direction. Most embryos hatched toward the light, suggesting they use light as a directional cue. Our results support that A. callidryas embryos use light cues to inform both when and where to hatch. Both the spatial orientation of hatching and the timing of spontaneous hatching may affect fitness and be informed by cues in a broader range of species than is currently appreciated.

Highlights

  • 32 33 Maternally provided structures such as egg capsules and shells provide embryos with protection, yet limit their interaction with the surrounding environment

  • Hatching was desynchronized in both continuous light and continuous 19 darkness. It was delayed by continuous light, but not accelerated by continuous dark, 20 suggesting the onset of dark serves as a hatching cue

  • One common and widespread environmental factor which entrains endogenous hatching rhythms are light-dark cycles (Asoh & Yoshikawa 2002; Brännäs 1987; Brüning et al 2011; DeCoursey 1983; McAlary & McFarland 1993; Salmon et al 1986). 65 Environmentally cued shifts in hatching timing are widespread in amphibians (Warkentin 66 2011b)

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Summary

Introduction

32 33 Maternally provided structures such as egg capsules and shells provide embryos with protection, yet limit their interaction with the surrounding environment. Many tropical brachyuran crabs and reef fishes release their larvae during high-amplitude, nocturnal high tides, facilitating larval transport away from high predation locations (Christy 2003; Johannes 1978; Morgan & Christy 1995; Robertson 1991) This regulation of hatching timing in response to abiotic cycles in fish and crustacean embryos is hypothesized to be an adaptive strategy which serves as an antipredator defense mechanism (Christy 2003; McAlary & McFarland 1993; Morgan & Christy 1995). Because we know A. callidryas embryos hatch primarily at night (Gomez-Mestre & Warkentin 2007; Warkentin 1995), we hypothesized that embryos in continuous darkness would hatch earlier and those in continuous light would hatch later than those in other treatments

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