Convergent camouflage and the non‐monophyly of ‘seadragons’ (Syngnathidae: Teleostei): suggestions for a revised taxonomy of syngnathids

Wilson, N. G. & Rouse, G. W. (2010). Convergent camouflage and the non‐monophyly of ‘seadragons’ (Syngnathidae: Teleostei): suggestions for a revised taxonomy of syngnathids. —Zoologica Scripta, 39, 551–558. The phylogeny and classification of the charismatic Syngnathidae (e.g. pipefish, seahorses) has not been comprehensively examined to date. In particular, we assessed morphological hypotheses that previously suggested the three ‘seadragon’ genera (Phycodurus, Phyllopteryx, Haliichthys) do not form a monophyletic group. We used three mitochondrial markers to investigate evolutionary relationships within Syngnathidae, and demonstrated that Phycodurus + Phyllopteryx formed a clade that excluded Haliichthys, indicating the elaborate appendages used for camouflage have evolved independently. A time‐calibrated tree revealed the divergence of true seadragons as coincident with other kelp‐associated fauna. We found evidence for the resurrection of neglected subfamily names, and recovered Doryrhampinae, Nerophinae, Soleganthinae, Phyllopteryginae, Sygnathoidinae and Haliichthyinae as clades. Even after removing these groups from what is currently recognized as Syngnathinae, we showed that the remaining members of Syngnathinae are not monophyletic. In the light of this information, some conclusions about the diversity of reproductive strategies found within ‘Syngnathinae’ need to be re‐examined and further revision of syngnathid classification is needed.     

The good,the bad,and the ugly: which Australian terrestrial mammal species attract most research?

相似文献   

Ricochet pollination in Senna (Fabaceae) – petals deflect pollen jets and promote division of labour among flower structures

• Naturalists Fritz and Hermann Müller hypothesised that heteranthery often leads to a division of labour into ‘feeding’ and ‘pollinating’ stamens; the latter often being as long as the pistil so as to promote successful pollination on the bees’ back. In many buzz‐pollinated species of Senna, however, the so‐called pollinating stamens are short and not level with the stigma, raising the question of how pollen is shed on the bees’ back. Here we explore a mechanism called ‘ricochet pollination’. We test whether division of labour is achieved through the interaction between short lower stamens and strongly concave ‘deflector petals’.
• We studied the arrangement and morphology of the floral organs involved in the ricochet pollination, functioning of the flowers through artificial sonication and observed the interactions between bees and flowers in the field.
• The middle stamens are adapted to eject pollen downwards, which can be readily collected on the bee mid legs. Most of the pollen is ejected towards the deflector petal(s). Pollen from this set of stamens is more likely to contribute to pollination. The pollen grains seem to ricochet multiple times against the deflector petals to eventually reach the bee's back.
• The pollen ricochet mechanism promotes a division of labour by involving additional floral organs, such as petals, reinforcing the Müllers’ division‐of‐labour hypothesis. However, alternative, non‐multiexclusive hypotheses could be explored in genus Senna and other angiosperm species.

     

Predator–prey naïveté, antipredator behavior,and the ecology of predator invasions

We present a framework for explaining variation in predator invasion success and predator impacts on native prey that integrates information about predator–prey naïveté, predator and prey behavioral responses to each other, consumptive and non‐consumptive effects of predators on prey, and interacting effects of multiple species interactions. We begin with the ‘naïve prey’ hypothesis that posits that naïve, native prey that lack evolutionary history with non‐native predators suffer heavy predation because they exhibit ineffective antipredator responses to novel predators. Not all naïve prey, however, show ineffective antipredator responses to novel predators. To explain variation in prey response to novel predators, we focus on the interaction between prey use of general versus specific cues and responses, and the functional similarity of non‐native and native predators. Effective antipredator responses reduce predation rates (reduce consumptive effects of predators, CEs), but often also carry costs that result in non‐consumptive effects (NCEs) of predators. We contrast expected CEs versus NCEs for non‐native versus native predators, and discuss how differences in the relative magnitudes of CEs and NCEs might influence invasion dynamics. Going beyond the effects of naïve prey, we discuss how the ‘naïve prey’, ‘enemy release’ and ‘evolution of increased competitive ability’ (EICA) hypotheses are inter‐related, and how the importance of all three might be mediated by prey and predator naïveté. These ideas hinge on the notion that non‐native predators enjoy a ‘novelty advantage’ associated with the naïveté of native prey and top predators. However, non‐native predators could instead suffer from a novelty disadvantage because they are also naïve to their new prey and potential predators. We hypothesize that patterns of community similarity and evolution might explain the variation in novelty advantage that can underlie variation in invasion outcomes. Finally, we discuss management implications of our framework, including suggestions for managing invasive predators, predator reintroductions and biological control.     

Three new species of Pristionchus (Nematoda: Diplogastridae) show morphological divergence through evolutionary intermediates of a novel feeding‐structure polymorphism

Developmental plasticity is often correlated with diversity and has been proposed as a facilitator of phenotypic novelty. Yet how a dimorphism arises or how additional morphs are added is not understood, and few systems provide experimental insight into the evolution of polyphenisms. Because plasticity correlates with structural diversity in Pristionchus nematodes, studies in this group can test the role of plasticity in facilitating novelty. Here, we describe three new species, Pristionchus fukushimae sp. nov. , Pristionchus hoplostomus sp. nov. , and the hermaphroditic Pristionchus triformis sp. nov. , which are characterized by a novel polymorphism in their mouthparts. In addition to showing the canonical mouth dimorphism of diplogastrid nematodes, comprising a stenostomatous (‘narrow‐mouthed’) and a eurystomatous (‘wide‐mouthed’) form, the new species exhibit forms with six, 12, or intermediate numbers of cheilostomatal plates. Correlated with this polymorphism is another trait that varies among species: whereas divisions between plates are complete in P. triformis sp. nov. , which is biased towards a novel ‘megastomatous’ form comprising 12 complete plates, the homologous divisions in the other new species are partial and of variable length. In a reconstruction of character evolution, a phylogeny inferred from 26 ribosomal protein genes and a partial small subunit rRNA gene supported the megastomatous form of P. triformis sp. nov. as the derived end of a series of split‐plate forms. Although split‐plate forms were normally only observed in eurystomatous nematodes, a single 12‐plated stenostomatous individual of P. hoplostomus sp. nov. was also observed, suggesting independence of the two types of mouth plasticity. By introducing these new species to the Pristionchus model system, this study provides further insight into the evolution of polymorphisms and their evolutionary intermediates. © 2013 The Linnean Society of London     

Publication bias in ecology and evolution: an empirical assessment using the 'trim and fill' method

Recent reviews of specific topics, such as the relationship between male attractiveness to females and fluctuating asymmetry or attractiveness and the expression of secondary sexual characters, suggest that publication bias might be a problem in ecology and evolution. In these cases, there is a significant negative correlation between the sample size of published studies and the magnitude or strength of the research findings (formally the ‘effect size’). If all studies that are conducted are equally likely to be published, irrespective of their findings, there should not be a directional relationship between effect size and sample size; only a decrease in the variance in effect size as sample size increases due to a reduction in sampling error. One interpretation of these reports of negative correlations is that studies with small sample sizes and weaker findings (smaller effect sizes) are less likely to be published. If the biological literature is systematically biased this could undermine the attempts of reviewers to summarise actual biology relationships by inflating estimates of average effect sizes. But how common is this problem? And does it really effect the general conclusions of literature reviews? Here, we examine data sets of effect sizes extracted from 40 peer‐reviewed, published meta‐analyses. We estimate how many studies are missing using the newly developed ‘trim and fill’ method. This method uses asymmetry in plots of effect size against sample size (‘funnel plots’) to detect ‘missing’ studies. For random‐effect models of meta‐analysis 38% (15/40) of data sets had a significant number of ‘missing’ studies. After correcting for potential publication bias, 21% (8/38) of weighted mean effects were no longer significantly greater than zero, and 15% (5/34) were no longer statistically robust when we used random‐effects models in a weighted meta‐analysis. The mean correlation between sample size and the magnitude of standardised effect size was also significantly negative (r_s=‐0.20, P < 0‐0001). Individual correlations were significantly negative (P < 0.10) in 35% (14/40) of cases. Publication bias may therefore effect the main conclusions of at least 15–21% of meta‐analyses. We suggest that future literature reviews assess the robustness of their main conclusions by correcting for potential publication bias using the ‘trim and fill’ method.     

The Relationship Between Foraging,Learning Abilities and Neophobia in Two Species of Darwin’s Finches

The ability to unlearn a previously established association is an important component of behavioural flexibility and may vary according to species ecology. Previously, two closely related sympatric Darwin’s finches were found to differ in their learning abilities. Small tree finches (Camarhynchus parvulus) outperformed woodpecker finches (Cactospiza pallida) in reversal learning but performed worse in an operant task. We attributed this difference to the habit of woodpecker finches to engage in long bouts of energetic pecking during extractive foraging. Persistently repeating one action without reward could favour performance in operant tasks but also limit behavioural flexibility. Here, we tested whether perseverance is the reason for woodpecker finches’ depressed reversal learning performance. Two new reversal conditions allowed the disentanglement of two sources of error in reversal learning: perseverant choice of the previously rewarded stimulus and failure to respond to the previously non‐rewarded stimulus. For the within‐species comparison, we predicted that woodpecker finches should find it more difficult to learn to avoid the previously rewarded stimulus than learning to choose the previously non‐rewarded stimulus. For the species comparison, we predicted the woodpecker finches should make more errors of perseverance than small tree finches. As performance could also be influenced by reaction to novelty, we compared neophobic responses between species and related them to reversal learning proficiency. We found no significant difference in reversal learning in the predicted direction, but found a negative correlation between neophobia and reversal learning at the inter‐ and the intraspecific level, which points towards a general relationship between reaction to novelty and flexibility.     

Length–weight relationships of 40 freshwater fish species from two decades of monitoring in Flanders (Belgium)

Nearly 263 000 individual length–weight (L/W) data, collected during 2839 fish stock assessments between 1992 and 2009, were used to calculate L/W relationships (W = aL^b) of 40 freshwater fish species from Flanders (Belgium). Roach Rutilus rutilus, perch Perca fluviatilis, three‐spine stickleback Gasterosteus aculeatus, gudgeon Gobio gobio and European eel Anguilla anguilla were the most abundant species in our surveys. Mean a and b values were in accordance with those available in FishBase. Mean b across all examined species is 3.094 (95% CL = 3.041 to 3.148) indicating a tendency towards slightly positive allometric growth in most fishes (thirty‐three species with b > 3). On the basis of a form factor equation, we characterized 2 species as ‘eel‐like’, 7 as ‘elongated’, 22 as ‘fusiform’ and 9 species as ‘short and deep’. This paper adds a substantial dataset to the yet scarcely available a and b values of European populations of freshwater fish species.     

Phylogenetic position and composition of Zygiellinae and Caerostris,with new insight into orb‐web evolution and gigantism

Orb‐weaving spiders are good objects for evolutionary research, but phylogenetic relationships among and within orb‐weaving lineages are poorly understood. Here we present the first species‐level molecular phylogeny that includes the enigmatic orb weavers ‘Zygiellidae’ and Caerostris. Zygiellidae is interesting for the evolution of the sector web, and Caerostris is noteworthy for web gigantism and extraordinary silk biomechanics. We assembled a molecular data set using mitochondrial (COI, 16S) and nuclear (H3, 18S, 28S, ITS2) gene fragments for 112 orbicularian exemplars, focusing on taxa with diverse web architecture and size. We show that ‘Zygiellidae’ contains the Holarctic Zygiella genus group (Leviellus, Parazygiella, Stroemiellus, and Zygiella) and the Australasian Phonognatha and Deliochus. As this clade is placed with Araneidae in all analyses we treat it as a subfamily, Zygiellinae. Using the new phylogeny, we show that the sector web evolved eight times, and coevolved with the silk tube retreat, but that these features are not zygielline synapomorphies. Zygiellinae, Caerostris, and some other araneids form a basal grade of araneids that differ from ‘classical’ araneids in web‐building and preying behaviour. We also confirm that Caerostris represents the most striking case of spider‐web gigantism. © 2015 The Linnean Society of London     

Statistical correlation between enterovirus genome copy numbers and infectious viral particles in wastewater samples

Aims: Classic virological tests are time consuming and labour‐intensive; real‐time RT‐PCR has proven to be a fast method to detect and quantify enterovirus genomes in clinical and environmental samples. This method is unable to discriminate between infective and noninfective enterovirus particles; few clinical studies have compared real‐time RT‐PCR and viral culture. We wondered if the enterovirus genome quantification could be correlated to the infectivity. Methods and Results: We used the statistical approach to verify our hypotheses to correlate data, obtained by the standard method (most probable number of cytopathic units—MPNCU) and molecular test (real‐time RT‐PCR), on wastewater treatment plant samples. Chi‐squared test was used, considering several cut‐off values (‘50’‐‘100’‐‘200’ genome copy numbers), to determine statistical significance in comparison of the two methods. Chi‐square value was not significant when cut‐off of 50 (P = 0·103) and 100 (P = 0·178) was assumed but was significant with cut‐off of 200 (P = 0·044). Conclusion: This limit, 200 genome copy, could be used as cut‐off value to indicate enterovirus survival in environmental monitoring. Significant and Impact of the Study: To introduce a fast procedure that is able to compensate for disadvantages of cell culture method for viral environmental analyses.     

Bottom‐up regulation of malaria population dynamics in mice co‐infected with lung‐migratory nematodes

When and how populations are regulated by bottom up vs. top down processes, and how those processes are affected by co‐occurring species, are poorly characterised across much of ecology. We are especially interested in the community ecology of parasites that must share a host. Here, we quantify how resources and immunity affect parasite propagation in experiments in near‐replicate ‘mesocosms’’ – i.e. mice infected with malaria (Plasmodium chabaudi) and nematodes (Nippostrongylus brasiliensis). Nematodes suppressed immune responses against malaria, and yet malaria populations were smaller in co‐infected hosts. Further analyses of within‐host epidemiology revealed that nematode co‐infection altered malaria propagation by suppressing target cell availability. This is the first demonstration that bottom‐up resource regulation may have earlier and stronger effects than top‐down immune mechanisms on within‐host community dynamics. Our findings demonstrate the potential power of experimental ecology to disentangle mechanisms of population regulation in complex communities.     

Additive genetic variance in polyandry enables its evolution,but polyandry is unlikely to evolve through sexy or good sperm processes

Polyandry is widespread despite its costs. The sexually selected sperm hypotheses (‘sexy’ and ‘good’ sperm) posit that sperm competition plays a role in the evolution of polyandry. Two poorly studied assumptions of these hypotheses are the presence of additive genetic variance in polyandry and sperm competitiveness. Using a quantitative genetic breeding design in a natural population of Drosophila melanogaster, we first established the potential for polyandry to respond to selection. We then investigated whether polyandry can evolve through sexually selected sperm processes. We measured lifetime polyandry and offensive sperm competitiveness (P₂) while controlling for sampling variance due to male × male × female interactions. We also measured additive genetic variance in egg‐to‐adult viability and controlled for its effect on P₂ estimates. Female lifetime polyandry showed significant and substantial additive genetic variance and evolvability. In contrast, we found little genetic variance or evolvability in P₂ or egg‐to‐adult viability. Additive genetic variance in polyandry highlights its potential to respond to selection. However, the low levels of genetic variance in sperm competitiveness suggest that the evolution of polyandry may not be driven by sexy sperm or good sperm processes.     

Flexible dispersal strategies in native and non‐native ranges: environmental quality and the ‘good–stay,bad–disperse’ rule

Dispersal strategies are one of the most important determinants of range dynamics and a surrogate for invasiveness. We tested three inter‐related hypotheses derived from demographic and ecological models: (H₁) short‐distance dispersal strategies arise at native range margins due to their demographic advantage; (H₂) in non‐native areas a high diffusion rate is favoured at the advancing range front for niche filling; (H₃) environmental deterioration can increase dispersal and lead to a ‘good–stay, bad–disperse’ strategy. Spatially and temporally explicit rates of spread and dispersal kernels of the European starling Sturnus vulgaris were generated for its native range (Britain) using ringing records from 1909 to 2008, and for a non‐native area (South Africa) using ringing data and distributional records since its introduction in 1897. There was a marked spatial and temporal variation in the rate of spread within both native and non‐native ranges. In the native range the rate of spread declined with increasing distance from the species’ European distribution (contradicting H₁). In the non‐native range the rate of spread increased with distance from the introduction locality (supporting H₂). The annual rate of spread in the native range also increased significantly when environmental conditions were deteriorating as indicated by marked population declines and relatively low abundance (H₃), providing clear evidence for flexible dispersal strategies based on a ‘good–stay, bad–disperse’ rule. Starlings’ dispersal kernel followed an inverse power law and showed strong anisotropy and significant divergence between native and invasive populations, suggesting a flexible strategy comprising a dynamic response to spatial and temporal environmental variation with implications for predicting dispersal and range dynamics arising from environmental change.     

A single‐stranded conformational polymorphism (SSCP)‐derived quantitative variable to monitor the virulence of a Barley yellow dwarf virus‐PAV (BYDV‐PAV) isolate during adaptation to the TC14 resistant wheat line

A standardized single‐stranded conformational polymorphism (SSCP) procedure is proposed as an alternative to the time‐consuming biological characterization of Barley yellow dwarf virus‐PAV (BYDV‐PAV) isolates. Using this procedure, six of 21 overlapping regions used to scan the viral genome gave patterns specific to ‘4E’ (avirulent) or ‘4T’ (‘4E’‐derived virulent) isolates. The calibration of samples and integration of SSCP patterns corresponding to the nucleotide region 1482–2023 allowed the estimation of P_T values that reflect the proportions of a ‘4T’‐specific band. Analysis of the biological (area under the pathogen progress curve) and molecular (P_T) data suggested a positive linear relation between these variables. Moreover, sequence analysis of the nucleotide region 1482–2023 highlighted the presence of a nucleotide polymorphism (C/A₁₈₃₅) which can be considered as a candidate for virus–host interactions linked to the monitored virulence. According to these parameters, P_T values associated with ‘4E’‐ and ‘4T’‐derived populations show that: (i) long‐term infection of a BYDV‐PAV isolate on the ‘TC14’ resistant host leads to the fixation of virulent individuals in viral populations; and (ii) the introduction of susceptible hosts in successive ‘TC14’ infections results in the maintenance of low virulence of the populations. Thus, the presented study demonstrates that SSCP is a useful tool for monitoring viral populations during the host adaptation process. The described impact of host alternation provides new opportunities for the use of the ‘TC14’ resistance source in BYDV‐resistant breeding programmes. This study is part of the global effort made by the scientific community to propose sustainable alternatives to the chemical control of this viral disease.     

Dispersal behaviour of individuals in metapopulations of two British butterflies

Dispersal patterns are important in metapopulation ecology because they affect the dynamics and survival of populations. However, because little empirical information exists on dispersal behaviour of individuals, theoretical models usually assume random dispersal. Recent empirical evidence, by contrast, suggests that the butterfly Maniola jurtina uses a non‐random, systematic dispersal strategy, can detect and orient towards habitat from distances of 100–150 m, and prefers a familiar habitat patch over a non‐familiar one (‘homing behaviour’). The present study (1) investigated whether these results generalise to another butterfly species, Pyronia tithonus; and (2) examined the cause of the observed ‘homing behaviour’ in M. jurtina. P. tithonus used a similar non‐random, systematic dispersal strategy to M. jurtina, had a similar perceptual range for habitat detection and preferred a familiar habitat patch over a non‐familiar one. The ‘homing behaviour’ of M. jurtina was found to be context‐dependent: individual M. jurtina translocated within habitat did not return towards their capture point, whereas individuals translocated similar distances out of habitat did return to their ‘home’ patch. We conclude that butterfly ‘homing behaviour’ is not based on an inherent preference for a familiar location, but that familiarity with an area facilitates the recognition of suitable habitat, towards which individuals orient if they find themselves in unsuitable habitat. Contrary to conventional wisdom, we suggest that frequent, short ‘excursions’ over habitat patch boundaries are evolutionarily advantageous to individuals, because increased familiarity with the surrounding environment is likely to increase the ability of a straying animal to return to its natural habitat, and to reduce the rate of mortality experienced by individuals attempting to disperse between habitat patches. We discuss the implications of the non‐random dispersal for existing metapopulation models, including models of the evolution of dispersal rates.     

Process,correlation and parameter fitting in species distribution models: a response to Kriticos et al

In a recent article (Dormann et al., 2012, Journal of Biogeography, 39, 2119–2131), we compared different approaches to species distribution modelling and depicted modelling approaches along an axis from purely ‘correlative’ to ‘forward process‐based’ models. In their correspondence, Kriticos et al. (2013, Journal of Biogeography, doi: 10.1111/j.1365‐2699.2012.02791.x ) challenge this view, claiming that our continuum representation neglects differences among models and does not consider the ability of fitted process‐based models to combine the advantages of both process‐based and correlative modelling approaches. Here we clarify that the continuum view resulted from recognition of the manifold differences between models. We also reinforce the point that the current trend towards combining different modelling approaches may lead not only to the desired combination of the advantages but also to the accumulation of the disadvantages of those approaches. This point has not been made sufficiently clear previously.     

Increases in Both Acute and Chronic Temperature Potentiate Tocotrienol Concentrations in Wild Barley at ‘Evolution Canyon’

Biosynthesis of tocols (vitamin E isoforms) is linked to response to temperature in plants. ‘Evolution Canyon’, an ecogeographical microcosm extending over an average of 200 meters (range 100–400) wide area in the Carmel Mountains of northern Israel, has been suggested as a model for studying global warming. Both domestic (Hordeum vulgare) and wild (Hordeum spontaneum) barley compared with wheat, oat, corn, rice, and rye show high tocotrienol/tocopherol ratios. Therefore, we hypothesized that tocol distribution might change in response to global warming. α‐, β‐, γ‐, and δ‐tocopherol, and α‐, β‐, γ‐, and δ‐tocotrienol concentrations were measured in wild barley (H. spontaneum) seeds harvested from the xeric (African) and mesic (European) slopes of Evolution Canyon over a six‐year period from 2005–2011. Additionally, we examined seeds from areas contiguous to and distant from the part of the Canyon severely burned during the Carmel Fire of December 2010. Increased α‐tocotrienol (p<0.01) was correlated with 1) temperature increases, 2) to the hotter ‘African’ slope in contrast to the cooler ‘European’ slope, and 3) to propinquity to the fire. The study illustrates the role of α‐tocotrienol in both chronic and acute temperature adaptation in wild barley and suggests future research into thermoregulatory mechanisms in plants.     

Importance of Sox2 in maintenance of cell proliferation and multipotency of mesenchymal stem cells in low-density culture

Objectives: This study has aimed to repopulate ‘primitive’ cells from late‐passage mesenchymal stem cells (MSCs) of poor multipotentiality and low cell proliferation rate, by simply altering plating density. Materials and methods: Effects of low density culture compared t high density culture on late‐passage bone marrow (BM)‐derived MSCs and pluripotency markers of multipotentiality were investigated. Cell proliferation, gene expression, RNA interference and differentiation potential were assayed. Results and conclusions: We repopulated ‘primitive’ cells by replating late‐passage MSCs at low density (17 cells/cm²) regardless of donor age. Repopulated MSCs from low‐density culture were smaller cells with spindle shaped morphology compared to MSCs from high‐density culture. The latter had enhanced colony‐forming ability, proliferation rate, and adipogenic and chondrogenic potential. Strong expression of osteogenic‐related genes (Cbfa1, Dlx5, alkaline phosphatase and type Ι collagen) in late‐passage MSCs was reduced by replating at low density, whereas expression of three pluripotency markers (Sox2, Nanog and Oct‐4), Osterix and Msx2 reverted to levels of early‐passage MSCs. Knockdown of Sox2 and Msx2 but not Nanog, using RNA interference, showed significant decrease in colony‐forming ability. Specifically, knockdown of Sox2 significantly inhibited multipotentiality and cell proliferation. Our data suggest that plating density should be considered to be a critical factor for enrichment of ‘primitive’ cells from heterogeneous BM and that replicative senescence and multipotentiality of MSCs during in vitro expansion may be predominantly regulated through Sox2.     

Fruit misshapen in strawberry cultivars (Fragaria×ananassa) is related to achenes functionality

Fruit deformation is a phenomenon commonly observed in commercial strawberry cultivars (Fragaria×ananassa) with a negative impact on the economic benefit of crop production. To better understand this physiopathy, we evaluated fruit size and the relative amount of ‘small’ (<0.4 mm) and ‘big’ achenes (>0.6 mm) in misshapen (MF) and well‐formed (WF) fruits from ‘Camarosa’, ‘Ventana’ and ‘Medina’ strawberry cultivars growing in the field. In ‘Camarosa’, size‐dependent achenes functionality was assessed by analysing achenes germinability and differences in time to ripening between MF and WF. We found that the occurrence of fruit deformation was not only strongly dependent on the cultivar (i.e. genetic factor) but also was promoted by low temperatures (i.e. environmental factor). Regardless of the cultivar, MF showed higher percentage of ‘small’ non‐functional achenes than WF, indicating a failure in achene development, which could be related to the functional integrity of reproductive structures. To test this hypothesis, we performed an experiment by pollinating strawberry flowers with pollen developed under low and mild temperatures. Low temperatures reduced pollen viability, and flowers pollinated with low quality pollen led to a higher amount of MF. However, fruit deformation was not completely explained by differences in pollen quality, neither by differences in pistil maturity and receptivity nor ovule viability, suggesting that non‐functional achenes might result from a disruption in postfertilisation processes such as embryo abortion as a consequence of low temperatures.     

Bird–flower interactions in the Macaronesian islands

Aims Several bird‐pollinated or ornithophilous flowers are present on the Macaronesian archipelagos, the Canary Islands and Madeira, but absent from nearby NW Africa and Europe. In Macaronesia, no specialist nectar‐feeding birds are found, but several generalist passerine bird species visit flowers for nectar. Two hypotheses attempt to explain the origin and evolution of ornithophily in the Macaronesian flora. According to ‘the island de novo hypothesis’, bird‐flowers evolved from mainland insect‐pollinated ancestors after island colonization. Alternatively, ancestors of the ornithophilous Macaronesian plant species evolved bird‐flowers before reaching the islands (‘the relict hypothesis’). In this study we first compile information of Macaronesian bird–flower interactions from the literature and our own field observations. Secondly, we discuss the two hypotheses of origin of ornithophily in the light of evidence from recent molecular plant phylogenies, palaeontology, historical biogeography of the African avifauna and flora, and present‐day ecological patterns. Location Madeira and Canary Islands. Results At least eleven endemic Macaronesian plant species from six genera have typical ornithophilous floral traits. These genera are: Canarina and Musschia (Campanulaceae), Isoplexis (Scrophulariaceae), Echium (Boraginaceae), Lotus (Fabaceae) and Lavatera (Malvaceae). These lineages have clear affinities to the Mediterranean region, except for Canarina whose closest relatives grow in East African mountains. Six generalist passerine bird species of Sylvia, Phylloscopus (Sylviidae), Serinus (Fringillidae) and Parus (Paridae) visit this flora for nectar. Main conclusion We suggest that the origin and evolution of ornithophilous traits in these plant species took place mostly in mainland areas prior to island colonization. In Canarina and Lavatera, it is well supported that ornithophily is a relict condition, which originated in mainland areas possibly in association with specialist nectar‐feeding birds. For the remaining plant species except Echium wildpretii bird floral traits probably also are a relict condition. These species may be derived from ancestors, which were visited by specialist nectar‐feeding birds during geological periods when the Mediterranean and the Ethiopian vegetation were intermingled in mainland Africa. Probably, these mainland ancestors went extinct due to severe climatic fluctuations, while their Macaronesian descendants survived in ‘refuge’ on the islands. Finally, the island de novo hypothesis may explain the evolution of a mixed bird/insect‐pollination system in the neo‐endemic red‐flowered Echium wildpretii.