• The specificity of orchids for their fungi can vary substantially, from highly specialist interactions to more generalist interactions, but little is known about the evolutionary history of the mycorrhizal specificity of orchids. • Here, we used a network analysis approach to investigate orchid mycorrhizal associations in 16 species of the genus Orchis sampled across 11 different regions in Europe. We first examined in detail the structure of the network of associations and then tested for a phylogenetic signal in mycorrhizal specificity and identified the fungi with which the orchids associated. • We found 20 different fungal lineages that associated with species of the genus Orchis, most of them being related to members of the Tulasnellaceae (84.33% of all identified associations) and a smaller proportion being related to members of the Ceratobasidiaceae (9.97%). Species associations formed a nested network that is built on asymmetric links among species. Evolution of mycorrhizal specificity in Orchis closely resembles a Brownian motion process, and the interaction between Orchis and Tulasnellaceae fungi is significantly influenced by the phylogenetic relationships between the Orchis species. • Our results provide evidence of the presence of phylogenetic conservatism in mycorrhizal specificity in orchids and demonstrate that evolutionary processes may be an important factor in generating patterns of mycorrhizal associations.

Summary: 1. The effects of habitat fragmentation on plant population genetic structure and diversity are relatively well studied. Yet, most of these studies used molecular tools focusing on neutral genetic markers, and much less is known about the potential evolutionary consequences of habitat fragmentation on ecologically relevant plant traits.2. It can be expected that the altered biotic and abiotic conditions and limited gene flow following habitat fragmentation may impose strong selection pressures on traits important for plant fitness. Responses to these selection pressures may, however, be hampered by reduced genetic diversity through genetic drift. Conversely, evolutionary changes in flower or dispersal traits may itself impact the strength of inbreeding and genetic drift.3. In this review, we highlight different reproductive plant traits that may be under selection following habitat fragmentation, and we examine studies that have shown indications for rapid evolutionary responses.4. There are still relatively few studies that have convincingly shown that habitat fragmentation generates rapid evolutionary responses. Separating genetic from plastic trait responses and quantifying the long‐term fitness consequences of evolutionary changes in plant traits also remain major challenges.5. Synthesis. Many plant traits may be subject to selection following habitat fragmentation, but, until now, studies that quantified evolutionary responses to habitat fragmentation are limited. To study the consequences of changing plant traits for population fitness, we advocate applying population models that link demographic vital rates, the responding plant traits and their heritability.

Summary: 1. In woodland herbs, the probability of flowering and costs associated with reproduction may strongly depend on environmental context (shade vs. light habitats) and on plant size. This may be particularly true for tuberous orchids that inhabit woodlands, as the amount of incoming radiation and total leaf area strongly determine photosynthetic capacity and hence the amount of carbohydrates that can be relocated to below‐ground storage organs that form next year’s rosette and flowering stalk. 2. To fully comprehend the impact of size‐dependent reproduction on population dynamics under varying light conditions, life cycle models should therefore include plant size in a continuous manner. In this study, annual changes in plant size and demographic behaviour of the tuberous perennial orchid Orchis purpurea were monitored during seven consecutive years (2003–2009) in open and shaded woodland. Integral projection models (IPMs) and life table response experiments (LTRE) were used to investigate the extent to which variation in plant size affected the overall population dynamics of this species and to decompose differences in population growth rates between populations of open and shaded woodland into contributions from growth, survival and reproduction. 3. Both plants in shaded and light environments needed to be a certain size to initiate flowering, but this threshold size was almost three times as large in shaded environments as in light environments. Plants in open woodlands flowered more frequently over the years, showed less size regression after flowering and produced significantly more fruits than plants in shaded environments, resulting in significantly larger population growth rates. 4. Our life cycle models revealed that costs of reproduction, measured at the population‐level, were small in the light environment, and more than buffered by the increase in survival of flowering plants compared to non‐flowering plants. In the shade environment, however, the costs of reproduction were significant and made the difference between a stable (current) and a growing (without reproduction costs) population. 5. Synthesis. Light penetration to the soil is a key variable determining population dynamics of woodland orchids. Our analyses show that differences in vital rates related to size‐dependent reproduction (flowering) and growth are essential drivers of changes in orchid population dynamics in different light environments. The combination of IPMs and LTREs thus proves to be very useful when studying the fitness consequences of size‐ and context‐dependent phenomena like flowering strategies and costs of reproduction.