Summary1. Herbarium collections contain long-term data for a wide range of taxa and provide unique opportunities to evaluate the importance of life-history components in driving species-specific responses to climate change. In this paper, we analyse the relationships between change in flowering dates and life-history traits within a phylogenetic framework. The study is based on an extensive data set of herbarium specimens of orchids collected in Hungary between 1837 and 2009, supplemented by recent field observations . 2. Of the 39 taxa investigated, 31 (79%) showed apparent advancement in mean flowering time. Among these, advancement was statistically significant in nine taxa. The rest (eight taxa) showed non-significant delays in flowering. Averaging across all taxa, flowering time advanced by 3 days (3.8% of flowering period) during the last 50 years compared with the period before 1960. In taxa showing significant advancement, flowering times advanced by 7.7 days (8.6% of the flowering period). The most extreme advancement was 13.9 days. 3. Multivariate models were used to evaluate ways in which life history may affect phenological responses to climate change. Pollination mode (i.e. deceptive vs. rewarding vs. autogamous), life span (i.e. short-lived vs. long-lived), biogeographical distribution type (i.e. Mediterranean vs. nonMediterranean) and flowering time (i.e. mean date of blooming) emerged as important factors that influence changes in flowering through time. Phylogenetic relatedness did not predict phenological response. The strongest response was observed in orchids that flower relatively early in spring, exhibit an autogamous or deceptive pollination mechanism, have a long life span and possess a Mediterranean centre of distribution. 4. Synthesis. Our investigation demonstrates that the majority of Hungarian orchids have shifted their yearly mean flowering to earlier dates during the past 50 years. Certain life-history traits, but not phylogenetic relatedness, were found to be important in predicting climatic responsiveness in European terrestrial orchids.

The red swamp crayfish (Procambarus clarkii), originally from North America, is one of the world's worst aquatic invaders. It is a favoured prey item for waterbirds, but the influence of this novel predator–prey relationship on dispersal of other organisms has not previously been considered. We investigated the potential for dispersal of plants and invertebrates by migratory waterbirds feeding on alien P. clarkii in European ricefields at harvest time. In November–December of 2014–2015, we collected propagules from the outside of 13 crayfish captured as they moved out of ricefields during harvest in Doñana, south‐west Spain. We also collected excreta (N = 76 faeces, 14 pellets) of lesser‐black backed gull (Larus fuscus). We recorded diaspores from at least 11 plant species (161 seeds from 10 angiosperm taxa, and 14 charophyte oogonia) on the outside of crayfish, together with 54 eggs from eight aquatic invertebrate taxa. Adults and juveniles of at least nine microcrustaceans, including the alien ostracods Hemicypris reticulata and Ankylocythere sinuosa, were also recovered from crayfish. No intact propagules were present in the digestive system of the crayfish. Contents of regurgitated pellets confirmed P. clarkii as the main food item for gulls. Diaspores from at least 12 plant species (154 seeds from 11 angiosperm taxa, and 17 charophyte oogonia) were recovered from gull excreta, together with 129 eggs of 12 aquatic invertebrate taxa. A statoblast of the alien bryozoan Plumatella vaihiriae was found in gull faeces. Seven of the plant species are important agricultural weeds, and two are alien to Spain. Diaspores from six plant taxa were germinated, confirming viability. These propagules were from a similar set of plants and invertebrates to those found on the outside of crayfish, suggesting that propagules in gull excreta were ingested inadvertently with their crayfish prey. Ricefields constitute a major artificial aquatic habitat covering an increasing proportion of the world's land surface and typically support native or alien crayfish. Crayfish invasion can lead to novel secondary dispersal pathways for plants and invertebrates through interactions with their predators, promoting the expansion of alien and native species (including weeds) through long‐distance dispersal via migratory waterbirds and increasing connectivity of organisms between artificial and natural ecosystems. This represents a previously overlooked impact of crayfish invasion on ecosystem services.

Modern literature on plant dispersal by birds focuses mainly on the importance of frugivory and scatter‐hoarding, yet recent studies show that endozoochory by migratory waterbirds is an important mechanism of long‐distance dispersal for a broad range of plants. Nevertheless, there is a lack of empirical field studies that identify the plants dispersed by waterbirds in a comprehensive manner. In particular, so far there are no detailed studies of the level of spatial variation in the plant taxa dispersed by a waterbird vector, and no clear demonstration that aquatic ferns can be dispersed by endozoochory. Consequently, we remain ignorant of the networks of dispersal interactions between granivorous waterbirds and plants. Five sets of faecal samples (total n = 215) were collected from mallards Anas platyrhynchos on autumn migration in the Hevesi‐holm and Balaton regions in Hungary, central Europe. Intact diaspores were extracted, identified and their germinability assessed under standard conditions. The plant communities recorded at different sites were compared with PERMANOVA and other multivariate methods. Macrospores of the floating watermoss Salvinia natans were recorded in 32 samples, and a total of 16 macrospores germinated, providing the first field demonstration of endozoochory of ferns by birds. Of 21 angiosperm taxa recorded (of which eight germinated), 13 were terrestrial species, although the most abundant taxa were aquatic species such as the alkali bulrush Bolboschoenus maritimus and the sago pondweed Potamogeton pectinatus. Two naturalized alien species, the common fig Ficus carica and the hackberry Celtis occidentalis were also recorded. Only four of the taxa had an endozoochory syndrome. The plant taxa dispersed varied at two different spatial scales, with minor but significant differences between samples from sites separated by less than 1 km, and major differences between the two regions separated by c. 220 km. Synthesis. This is a unique study of the spatial variation in plants dispersed by endozoochory by a migratory waterfowl species, with a high taxonomic resolution and the first demonstration of avian endozoochory of ferns. We recorded eight taxa not previously reported as dispersed by mallards, showing how more empirical studies are essential, so we can understand which plants are dispersed by migratory birds. We found evidence that networks of interactions between granivorous waterbird vectors and dispersed plants vary spatially.

Field studies have shown that waterbirds, especially members of the Anatidae family, are major vectors of dispersal by endozoochory for a broad range of plants lacking a fleshy fruit, yet whose propagules can survive gut passage. Widely adopted dispersal syndromes ignore this dispersal mechanism, and we currently have little understanding of what traits determine the potential of angiosperms for endozoochory by waterbirds. Results from previous experimental studies have been inconsistent as to how seed traits affect seed survival and retention time in the gut and have failed to control for the influence of plant phylogeny. Using 13 angiosperm species from aquatic and terrestrial habitats representing nine families, we examined the effects of seed size, shape, and hardness on the proportion of seeds surviving gut passage through mallards (Anas platyrhynchos) and their retention time within the gut. We compiled a molecular phylogeny for these species and controlled for the nonindependence of taxa due to common descent in our analyses. Intact seeds from all 13 species were egested, but seed survival was strongly determined by phylogeny and by partial effects of seed mass and hardness (wet load): species with seeds harder than expected from their size, and smaller than expected from their loading, had greater survival. Once phylogeny was controlled for, a positive partial effect of seed roundness on seed survival was also revealed. Species with seeds harder than expected from their size had a longer mean retention time, a result retained after controlling for phylogeny. Our study is the first to demonstrate that seed shape and phylogeny are important predictors of seed survival in the avian gut. Our results demonstrate that the importance of controlling simultaneously for multiple traits and relating single traits (e.g., seed size) alone to seed survival or retention time is not a reliable way to detect important patterns, especially when phylogenetic effects are ignored.

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Background and AimsThe charismatic Himantoglossum s.l. clade of Eurasian orchids contains an unusually large proportion of taxa that are of controversial circumscriptions and considerable conservation concern. Whereas our previously published study addressed the molecular phylogenetics and phylogeography of every named taxon within the clade, here we use detailed morphometric data obtained from the same populations to compare genotypes with associated phenotypes, in order to better explore taxonomic circumscription and character evolution within the clade.MethodsBetween one and 12 plants found in 25 populations that encompassed the entire distribution of the Himantoglossum s.l. clade were measured in situ for 51 morphological characters. Results for 45 of those characters were subjected to detailed multivariate and univariate analyses.Key ResultsMultivariate analyses readily separate subgenus Barlia and subgenus Comperia from subgenus Himantoglossum, and also the early-divergent H. formosum from the less divergent remainder of subgenus Himantoglossum. The sequence of divergence of these four lineages is confidently resolved. Our experimental approach to morphometric character analysis demonstrates clearly that phenotypic evolution within Himantoglossum is unusually multi-dimensional.ConclusionsDegrees of divergence between taxa shown by morphological analyses approximate those previously shown using molecular analyses. Himantoglossum s.l. is readily divisible into three subgenera. The three sections of subgenus Himantoglossum—hircinum, caprinum and formosum—are arrayed from west to east with only limited geographical overlap. At this taxonomic level, their juxtaposition combines with conflict between contrasting datasets to complicate attempts to distinguish between clinal variation and the discontinuities that by definition separate bona fide species. All taxa achieve allogamy via food deceit and have only weak pollinator specificity. Artificial crossing demonstrates that intrinsic sterility barriers are weak. Although we have found evidence of gene flow among and within the three sections of subgenus Himantoglossum, reports of natural hybrids are surprisingly rare, probably because putative parents are sufficiently similar to questionably warrant the status of species. Phenological separation and increased xeromorphy characterise the origin of subgenus Barlia. Several individual morphological characters show evidence of parallel acquisition, and loss of features is especially frequent in floral markings among members of section caprinum. Detailed patterns of gain and loss demonstrate that several different categories of flower markings are inherited independently. Along with the dimensions of labellar lobes, these pigmentation characters have been over-emphasised in previous taxonomic treatments. Increased plant vigour was a crucial element of the origin of the genus, but vegetative characters underwent remarkably little subsequent evolution. Attempts to reconstruct hypothetical ancestors at internal nodes of t...

Summary 1 1. Floral deception is widespread in orchids, with more than one third of the species being 2 pollinated this way. The evolutionary success of deceptive orchids is puzzling, as species 3 employing this strategy are thought to have low reproductive success (less flowers yielding 4 fruits) because of low pollination rates. However, direct measurements of seed production in 5 orchids are scarce due to the extremely small size of their seeds. 6 2. Here, we quantified seed numbers in 1,015 fruits belonging to 48 orchid species from the 7 Pannonian ecoregion (central Europe) and obtained fruit-set and thousand-seed weight data 8 for these species from the literature. We used phylogenetic comparative methods to test the 9 hypothesis that deceptive species should compensate for their lower fruit-set by having either 10 larger seeds or more seeds in a fruit. 11 3. Similarly to previous studies, we found that deceptive orchids have substantially lower 12 fruits-set than nectar-rewarding ones. Also, we found that deceptive species have more seeds 13 in a fruit but not larger seeds compared to nectar-rewarding ones. Based on our results, 14 deceptive species compensate for their lower fruit-set by having higher seed numbers per 15 fruit, thus their seed numbers per shoot do not differ from that of nectar-rewarding ones. 16 4. Together with other benefits of deceptive pollination (e.g. lower energy expenditure due to 17 the lack of nectar production and higher genetic variability due to decreased probability of 18 geitonogamous pollination), our results can explain why deceptive strategies are so 19 widespread in the orchid family. 20 5. Synthesis. Our study provides new seed number data for 48 terrestrial orchid species. 21Using these data we have tested the hypothesis that deceptive species should compensate for 22 their lower fruit-set by having either larger seeds or more seeds in a fruit than nectar-23 rewarding ones. Our results suggest that deceptive species have more seeds in a fruit but not 24 larger seeds compared to nectar-rewarding ones. As a consequence, there are no significant 25 3 differences in seed numbers per shoot between different pollination types. 1 2