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2018  Divergent strategies in pre- and postzygotic reproductive isolation between two closely related Dianthus species — Cahenzli F, Bonetti C & Erhardt A — Evolution 72: 1851–1862   doi> 10.1111/evo.13556 

Quantifying the relative contribution of multiple isolation barriers to gene flow between recently diverged species is essential for understanding speciation processes. In parapatric populations, local adaptation is thought to be a major contributor to the evolution of reproductive isolation. However, extrinsic postzygotic barriers assessed in reciprocal transplant experiments are often neglected in empirical assessments of multiple isolation barriers. We analyzed multiple isolation barriers between two closely related species of the plant genus Dianthus, a genus characterized by the most rapid species diversification in plants reported so far. Although D. carthusianorum L. and D. sylvestris Wulf. can easily be hybridized in crossing experiments, natural hybrids are rare. We found that in parapatry, pollinator mediated prezygotic reproductive isolation barriers are important for both D. carthusianorum (0.761) and D. sylvestris (0.468). In contrast to D. carthusianorum, high hybrid viability in D. sylvestris (-0.491) was counteracted by strong extrinsic postzygotic isolation (0.900). Our study highlights the importance of including reciprocal transplant experiments for documenting extrinsic postzygotic isolation and demonstrates clearly divergent strategies and hence asymmetric pre- and postzygotic reproductive isolation between closely related species. It also suggests that pollinator-mediated and ecological isolation could have interacted in synergistic ways, further stimulating rapid speciation in Dianthus.

Keywords: Dianthus carthusianorum - Dianthus sylvestris - extrinsic postzygotic isolation - reciprocal transplant experiments - reproductive isolation barriers - speciation


2015  Female butterflies adapt and allocate their progeny to the host-plant quality of their own larval experience — Cahenzli F, Wenk BA & Erhardt A — Ecology 96: 1966–1973   doi> 10.1890/14-1275.1 

Recent studies with diverse taxa have shown that parents can utilize their experience of the environment to adapt their offspring's phenotype to the same environmental conditions. Thus, offspring would then perform best under environmental conditions experienced by their parents due to transgenerational phenotypic plasticity. Such an effect has been dubbed transgenerational acclimatization. However, evidence that parents can subsequently ensure the appropriate environmental conditions in order that offspring benefit from transgenerational acclimatization has never been demonstrated. We reared Pieris rapae larvae in the parental generation on high-nitrogen and low-nitrogen host plants, and reared the offspring (F1) of both treatments again on high- and low-nitrogen plants. Furthermore, we tested if females prefer to oviposit on high- or low-nitrogen host plants in two-way choice tests. We here show not only that females adapt their offspring's phenotype to the host-plant quality that they themselves experienced, but that females also mainly oviposit on the host quality to which they adapt their offspring. Moreover, effects of larval host plant on oviposition preference of females increased across two generations in F1-females acclimatized to low-nitrogen host plants, showing an adaptive host shift from one generation to the next. These findings may have profound implications for host-race formation and sympatric speciation.

Keywords: butterfly reproduction - Hopkins' host selection principle - host shift - maternal effects - oviposition - phenotypic plasticity - Pieris rapae - transgenerational acclimatization


2013  Nectar amino acids enhance reproduction in male butterflies — Cahenzli F & Erhardt A — Oecologia 171: 197–205   doi> 10.1007/s00442-012-2395-8 

After over 30 years of research, it was recently shown that nectar amino acids increase female butterfly fecundity. However, little attention has been paid to the effect of nectar amino acids on male butterfly reproduction. Here, we show that larval food conditions (nitrogen-rich vs. nitrogen-poor host plants) and adult diet quality (nectar with or without amino acids) affected the amount of consumed nectar in Coenonympha pamphilus males. Furthermore, amino acids in the nectar diet of males increased progeny's larval hatching mass, irrespective of paternal larval reserves. Our study takes the whole reproductive cycle of male butterflies into account, and also considers the role of females in passing male nutrients to offspring, as males' realized reproduction was examined indirectly via nuptial gifts, by female performance. With this comprehensive approach, we demonstrate for the first time that nectar amino acids can improve male butterfly reproduction, supporting the old postulate that nectar amino acids generally enhance butterfly fitness.

Keywords: butterfly reproduction - larval feeding - Lepidoptera - pollination


2013  Transgenerational acclimatization in an herbivore-host plant relationship — Cahenzli F & Erhardt A — Proceedings of the Royal Society of London B - Biological Sciences 280: 20122856   doi> 10.1098/rspb.2012.2856 

Twenty years ago, scientists began to recognize that parental effects are one of the most important influences on progeny phenotype. Consequently, it was postulated that herbivorous insects could produce progeny that are acclimatized to the host plant experienced by the parents to improve progeny fitness, because host plants vary greatly in quality and quantity, and can thus provide important cues about the resources encountered by the next generation. However, despite the possible profound implications for our understanding of host-use evolution of herbivores, host-race formation and sympatric speciation, intense research has been unable to verify transgenerational acclimatization in herbivore–host plant relationships. We reared Coenonympha pamphilus larvae in the parental generation (P) on high- and low-quality host plants, and reared the offspring (F1) of both treatments again on high- and low-quality plants. We tested not only for maternal effects, as most previous studies, but also for paternal effects. Our results show that parents experiencing predictive cues on their host plant can indeed adjust progeny's phenotype to anticipated host plant quality. Maternal effects affected female and male offspring, whereas paternal effects affected only male progeny. We here verify, for the first time to our knowledge, the long postulated transgenerational acclimatization in an herbivore–host plant interaction.

Keywords: butterfly reproduction - larval feeding - Lepidoptera - maternal effects - paternal effects - phenotypic plasticity


2012  Enhancing offspring quality or quantity? Different ways for using nectar amino acids in female butterflies — Cahenzli F & Erhardt A — Oecologia 169: 1005–1014   doi> 10.1007/s00442-012-2254-7 

Butterfly-pollinated flowers offer nectar with higher amino acid concentrations than most flowers pollinated by other animals, and female butterflies of some species prefer to consume amino acid-rich nectar. However, for over 30 years, there has been an ongoing discussion about whether nectar amino acids benefit butterfly fitness. A clear positive effect was only shown for the nectar-feeding Araschnia levana, and females of the fruit-feeding Bicyclus anynana also increased offspring quality when they were fed amino acids as adults. Thus, severe doubts remain about the general significance of these single positive results. We therefore tested a further species from a phylogenetically different butterfly subfamily, the small heath (Coenonympha pamphilus L., Satyrinae), taking into account feeding conditions over the whole life cycle of this species. C. pamphilus females receiving nectar amino acids as adults, irrespective of larval food quality, produced heavier larvae and also increased the hatching success of their eggs offer the oviposition period. Furthermore, females raised under nitrogen-poor larval conditions tended to use nectar amino acids to increase the number of eggs laid. Thus, C. pamphilus females used nectar amino acids primarily to increase their offspring quality, and secondly tended to increase offspring quantity, if larval resources were scarce, showing a resource allocation pattern differing from both B. anynana and A. levana. Our study supports the old postulate that nectar amino acids generally enhance butterfly fitness.

Keywords: butterfly reproduction - Coenonympha pamphilus - larval feeding - lepidoptera pollination


2012  Nectar sugars enhance fitness in male Coenonympha pamphilus butterflies by increasing longevity or realized reproduction — Cahenzli F & Erhardt A — Oikos 121: 1417–1423   doi> 10.1111/j.1600-0706.2012.20190.x 

The principal components of floral nectar are water and the sugars sucrose, fructose and glucose. Several studies have shown the importance of nectar sugars for female butterfly fecundity, whereas to date little attention has been paid to the effect of nectar sugars on male butterfly reproduction. Clear evidence for an effect of nectar sugars on male realized reproductive success is still missing.
In this study, we fed male Coenonympha pamphilus butterflies nectar mimics with low (5%), medium (20%) or high (30%) total sugar concentrations with a sucrose:glucose:fructose ratio of 2.7:1.1:1. Sugar solutions were made mimicking Knautia arvensis, an essential nectar plant for C. pamphilus and many other European butterflies. Realized male reproductive success for each treatment was measured indirectly via nuptial gifts, by recording reproductive parameters and by characterizing time patterns over the oviposition period of their female partner.
Male butterflies fed high-concentrated nectar sugars had a longer lifespan than males fed low-concentrated nectar sugars. In contrast, offspring of males fed medium-concentrated nectar sugars had a higher hatching mass than progeny of males fed low-concentrated nectar sugars, indicating a tradeoff between somatic maintenance and reproduction in the use of nectar sugars. Thus, allocation patterns of nectar sugars differed according to sugar concentrations in adult food.
The method used in this experiment took into account the indispensable role of female butterflies in passing male nutrients to offspring. With this comprehensive approach, we can show the general importance of nectar sugars for male butterfly fitness and support previous findings suggesting a coevolutionary process between butterflies and flowers dependent on butterfly pollination.


2012  Host plant defence in the larval stage affects feeding behaviour in adult butterflies — Cahenzli F & Erhardt A — Animal Behaviour 84: 995–1000   doi> 10.1016/j.anbehav.2012.07.025 

Nitrogen is a key nutrient for fitness in insects, but host plant defence can deter herbivores from acquiring it. Therefore, coping with host plant defence is a predominant issue for herbivores. Butterflies have the ability to compensate for a nitrogen-poor diet both as herbivorous larvae and as nectar-feeding adults. We examined whether silica (S), the main anti herbivore defence in grasses (in both fertilized nitrogen-rich (F+) and nitrogen-poor (F-) Festuca rubra host plants) affects larval development and accordingly adult feeding behaviour in the small heath butterfly, Coenonympha pamphilus. High silica levels in nitrogen-poor hostplants (F-/S+) negatively affected larval performance, and, as a consequence, female and even male butterflies preferred to consume amino acid-rich nectar. Our findings show for the first time that plant defence in larval host plants affects feeding behaviour in adult butterflies, and that even male butterflies, which have so far been thought to be indifferent to nectar amino acids, preferred to consume amino acid-rich nectar. Hence larval food quality can influence plant–pollinator dynamics.

Keywords: amino acid - Coenonympha pamphilus - Festuca rubra - larval feeding - Lepidoptera - nectar - preference - Satyrinae - silica