title: Desiccation plasticity in the embryonic life histories of non-annual rivulid species.
abstract: BACKGROUND
  Diapause is a developmental arrest present in annual killifish, whose eggs are able to survive long periods of desiccation when the temporary ponds they inhabit dry up. Diapause can occur in three different developmental stages. These differ, within and between species, in their responsiveness to different environmental cues. A role of developmental plasticity and genetic assimilation in diapause evolution has been previously suggested but not experimentally explored. We investigated whether plastic developmental delays or arrests provoked by an unusual and extreme environment could be the ancestral condition for diapause. This would be in agreement with plasticity evolution playing a role in the emergence of diapause in this group. We have used a comparative experimental approach and exposed embryos of non-annual killifish belonging to five different species from the former genus Rivulus to brief periods of desiccation. We have estimated effects on developmental and mortality rates during and after the desiccation treatment.
  
  
  RESULTS
  Embryos of these non-annual rivulids decreased their developmental rates in early stages of development in response to desiccation and this effect persisted after the treatment. Two pairs of two different species had sufficient sample sizes to investigate rates of development in later stages well. In one of these, we found cohorts of embryos in the latest stages of development that did not hatch over a period of more than 1 month without mortality. Several properties of this arrest are also used to characterize diapause III in annual killifish. Such a cohort is present in control conditions and increases in frequency in the desiccation treatment.
  
  
  CONCLUSIONS
  The presence of plasticity for developmental timing and a prolonged developmental arrest in non-annual rivulids, suggest that a plastic developmental delay or diapause might have been present in the shared ancestor of annual and non-annual South American killifish and that the evolution of plasticity could have played a role in the emergence of the diapauses. Further comparative experimental studies and field research are needed to better understand how diapause and its plasticity evolved in this group.
authors.affiliation: Naturalis Biodiversity Center, Darwinweg 2, Leiden 2333 CR, The Netherlands ; Current address: Instituto Gulbenkian de Ciência, Rua da Quinta Grande 6, 2780-156 Oeiras, Portugal.
authors.forename: Irma
authors.initials: I
authors.lastname: Varela-Lasheras
authors.affiliation: Naturalis Biodiversity Center, Darwinweg 2, Leiden 2333 CR, The Netherlands ; CNRS/UPMC/UPEC/UPD/IRD/INRA-UMR 7618 Institute of Ecology and Environmental Sciences Paris (iEES), Université Pierre et Marie Curie, Case 237, 7 Quai St Bernard, 75005 Paris, France.
authors.forename: Tom Jm
authors.initials: TJ
authors.lastname: Van Dooren
keywords: Diapause; Genetic assimilation; Heterokairy; Killifish; Lagged responses; Multi-state modelling; Plasticity evolution; Quiescence; Survival analysis