Environmental filters shaping angiosperm tree assembly along climatic and geographic gradients

Question

Global‐scale forest censuses provide an opportunity to understand diversification processes in woody plant communities. Based on the climatic or geographic filtering hypotheses associated with tropical niche conservatism and dispersal limitation, we analysed phylogenetic community structures across a wide range of biomes and evaluated to what extent region‐specific processes have influenced large‐scale diversity patterns of tree species communities across latitude or continent.

Location

Global.

Methods

We generated a data set of species abundances for 21,379 angiosperm woody plants in 843 plots worldwide. We calculated net relatedness index (NRI) for each plot, based on a single global species pool and regional species pools, and phylogenetic β‐diversity (PBD) between plots. Then, we explored the correlations of NRI with climatic and geographic variables, and clarified phylogenetic dissimilarity along geographic and climatic differences. We also compared these patterns for South America, Africa, the Indo‐Pacific, Australia, the Nearctic, Western Palearctic and Eastern Palearctic.

Results

NRI based on a global‐scale species pool was negatively associated with precipitation and positively associated with Quaternary temperature change. PBD was positively associated with geographic distance and precipitation difference between plots across tropical and extratropical biomes. Moreover, phylogenetic dissimilarity was smaller in extratropical regions than in regions including the tropics, although temperate forests of the Eastern Palearctic showed a greater dissimilarity within extratropical regions.

Conclusions

Our findings support predictions of the climatic and geographic filtering hypotheses. Climatic filtering (climatic harshness and paleoclimatic change) relative to tropical niche conservatism played a role in sorting species from the global species pool and shaped the large‐scale diversity patterns, such as the latitudinal gradient observed across continents. Geographic filtering associated with dispersal limitation substantially contributed to regional divergence of tropical/extratropical biomes among continents. Old, long‐standing geographic barriers and recent climatic events differently influenced evolutionary diversification of angiosperm tree communities in tropical and extratropical biomes.     

Pollination system of the Pilosocereus leucocephalus columnar cactus (tribe Cereeae) in eastern Mexico

It has been suggested that there is a geographic dichotomy in the pollination systems of chiropterophilous columnar cacti: in intra‐tropical areas they are pollinated almost exclusively by bats, whereas in extratropical areas they are pollinated by bats, birds and bees. However, currently the studies are clumped both taxonomically (mainly Pachycereeae species) and geographically (mainly in the Tehuacan Valley and the Sonoran Desert). This clumping limits the possibility of generalising the pattern to other regions or cactus tribes. Only four of the 36 chiropterophilous cacti in Pilosocereus have been studied. Despite the tropical distribution of two Pilosocereus species, bees account for 40–100% of their fruit set. We examined how specialised is the pollination system of P. leucocephalus in eastern Mexico. As we studied tropical populations, we expected a bat‐specialised pollination system. However, previous studies of Pilosocereus suggest that a generalised pollination system is also possible. We found that this cactus is mainly bat‐pollinated (bats account for 33–65% of fruit set); although to a lesser degree, diurnal visitors also caused some fruit set (7–15%). Diurnal visitors were more effective in populations containing honeybee hives. P. leucocephalus is partially self‐compatible (14–18% of fructification) but unable to set fruit without visitors. Despite the variation in pollination system, P. leucocephalus shows more affinity with other columnar cacti from tropical regions than with those from extratropical regions. Although we report here that a new species of tropical Pilosocereus is relatively bat‐specialised, this Cereeae genus is more flexible in its pollination system than the Pachycereeae genera.     

Breeding systems in Balsaminaceae in relation to pollen/ovule ratio,pollination syndromes,life history and climate zone

• Pollen/ovule (P/O) ratios are often used as proxy for breeding systems. Here, we investigate the relations between breeding systems and P/O ratios, pollination syndromes, life history and climate zone in Balsaminaceae.
• We conducted controlled breeding system experiments (autonomous and active self‐pollination and outcrossing tests) for 65 Balsaminaceae species, analysed pollen grain and ovule numbers and evaluated the results in combination with data on pollination syndrome, life history and climate zone on a phylogenetic basis.
• Based on fruit set, we assigned three breeding systems: autogamy, self‐compatibility and self‐incompatibility. Self‐pollination led to lower fruit set than outcrossing. We neither found significant P/O differences between breeding systems nor between pollination syndromes. However, the numbers of pollen grains and ovules per flower were significantly lower in autogamous species, but pollen grain and ovule numbers did not differ between most pollination syndromes. Finally, we found no relation between breeding system and climate zone, but a relation between climate zone and life history.
• In Balsaminaceae reproductive traits can change under resource or pollinator limitation, leading to the evolution of autogamy, but are evolutionary rather constant and not under strong selection pressure by pollinator guild and geographic range changes. Colonisation of temperate regions, however, is correlated with transitions towards annual life history. Pollen/ovule‐ratios, commonly accepted as good indicators of breeding system, have a low predictive value in Balsaminaceae. In the absence of experimental data on breeding system, additional floral traits (overall pollen grain and ovule number, traits of floral morphology) may be used as proxies.

     

Pollinator individual‐based networks reveal the specialized plant–pollinator mutualism in two biodiverse communities

Generalization of pollination systems is widely accepted by ecologists in the studies of plant–pollinator interaction networks at the community level, but the degree of generalization of pollination networks remains largely unknown at the individual pollinator level. Using potential legitimate pollinators that were constantly visiting flowers in two alpine meadow communities, we analyzed the differences in the pollination network structure between the pollinator individual level and species level. The results showed that compared to the pollinator species‐based networks, the linkage density, interaction diversity, interaction evenness, the average plant linkage level, and interaction diversity increased, but connectance, degree of nestedness, the average of pollinator linkage level, and interaction diversity decreased in the pollinator individual‐based networks, indicating that pollinator individuals had a narrower food niche than their counterpart species. Pollination networks at the pollinator individual level were more specialized at the network level (H′₂) and the plant species node level (d′) than at the pollinator species‐level networks, reducing the chance of underestimating levels of specialization in pollination systems. The results emphasize that research into pollinator individual‐based pollination networks will improve our understanding of the pollination networks at the pollinator species level and the coevolution of flowering plants and pollinators.     

Latitudinal gradients in biotic interactions: Are cacti pollination systems more specialized in the tropics?

• Biotic interactions are said to be more specialized in the tropics, and this was also proposed for the pollination systems of columnar cacti from North America. However, this has not yet been tested for a wider set of cactus species. Here, we use the available information about pollination in the Cactaceae to explore the geographic patterns of this mutualistic interaction, and test if there is a latitudinal gradient in its degree of specialization.
• We performed a bibliographic search of all publications on the pollination of cacti species and summarized the information to build a database. We used generalized linear models to evaluate if the degree of specialization in cacti pollination systems is affected by latitude, using two different measures: the number of pollinator guilds (functional specialization) and the number of pollinator species (ecological specialization).
• Our database contained information about the pollination of 148 species. The most frequent pollinator guilds were bees, birds, moths and bats. There was no apparent effect of latitude on the number of guilds that pollinate a cactus species. However, latitude had a small but significant effect on the number of pollinator species that service a given cactus species.
• Bees are found as pollinators of most cactus species, along a wide latitudinal gradient. Bat and bird pollination is more common in the tropics than in the extra-tropics. The available information suggests that cacti pollination systems are slightly more ecologically specialized in the tropics, but it does not support any trend with regard to functional specialization.

     

Characterizing ecological generalization in plant-pollination systems

Despite the development of diversity indices in community ecology that incorporate both richness and evenness, pollination biologists commonly use only pollinator richness to estimate generalization. Similarly, while pollination biologists have stressed the utility of pollinator importance, incorporating both pollinator abundance and effectiveness, importance values have not been included in estimates of generalization in pollination systems. In this study, we estimated pollinator generalization for 17 plant species using Simpson’s diversity index, which includes richness and evenness. We compared these estimates with estimates based on only pollinator richness, and compared diversity estimates calculated using importance data with those using only visitation data. We found that pollinator richness explains only 57–65% of the variation in diversity, and that, for most plant species, pollinator importance was determined primarily by differences in visitation rather than by differences in effectiveness. While simple richness may suffice for broad comparisons of pollinator generalization, measures that incorporate evenness will provide a much more accurate understanding of generalization. Although incorporating labor-intensive measurements of pollinator effectiveness are less necessary for broad surveys, effectiveness estimates will be important for detailed studies of some plant species. Unfortunately, at this point it is impossible to predict a priori which species these are.     

Pollination efficiency and foraging behaviour of honey bees and non‐Apis bees to sweet cherry

1. Crop pollination generally increases with pollinator diversity and wild pollinator visitation. To optimize crop pollination, it is necessary to investigate the pollination contribution of different pollinator species. In the present study, we examined this contribution of honey bees and non‐Apis bees (bumble bees, mason bees and other solitary bees) in sweet cherry.
2. We assessed the pollination efficiency (fruit set of flowers receiving only one visit) and foraging behaviour (flower visitation rate, probability of tree change, probability of row change and contact with the stigma) of honey bees and different types of non‐Apis bees.
3. Single visit pollination efficiency on sweet cherry was higher for both mason bees and solitary bees compared with bumble bees and honey bees. The different measures of foraging behaviour were variable among non‐Apis bees and honey bees. Adding to their high single visit efficiency, mason bees also visited significantly more flower per minute, and they had a high probability of tree change and a high probability to contact the stigma.
4. The results of the present study highlight the higher pollination performance of solitary bees and especially mason bees compared with bumble bees and honey bees. Management to support species with high pollination efficiency and effective foraging behaviour will promote crop pollination.

     

Pollination efficiency of wild bees and hoverflies provided to oilseed rape

• 1 Declining numbers in honeybees and various wild bee species pose a threat to global pollination services. The identification and quantification of the pollination service provided by different taxa within the pollinator guild is a prerequisite for the successful establishment of nature conservation and crop management regimes.
• 2 Wild bees and hoverflies are considered to be valuable pollinators in agricultural and natural systems. Although some information on pollination efficiency of individual pollinator species is available, comparative studies of both taxa at different densities are rare. In the present study, the efficiency of the solitary mason bee Osmia rufa and two hoverfly species (Eristalis tenax and Episyrphus balteatus) as pollinators of oilseed rape Brassica napus was examined in a standardized caged plant breeding regime. Honeybee Apis mellifera colonies were used as a reference pollinator taxon.
• 3 Yield parameters responded differently to pollinator density and identity. Fruit set and number of seeds per pod increased with increasing pollinator density, although these were stronger in the mason bee than the hoverfly treatment. Weight per 1000 seeds did not respond to any pollinator treatment, indicating that seed quality was not affected. Oilseed rape yield in the highest tested densities of both pollinator taxa resulted in yield values close to the efficiency of small honeybee colonies.
• 4 Hoverflies required approximately five‐fold densities of the red mason bees to reach a similar fruit set and yield. Thus, mason bees are more efficient in plant breeding and managed pollination systems. Both natural pollinator taxa, however, are of potential value in open and closed crop production systems.

     

Reproductive biology of a hummingbird‐pollinated Billbergia: light influence on pollinator behaviour and specificity in a Brazilian semi‐deciduous forest

• Ornithophily has evolved in parallel several times during evolution of angiosperms. Bird pollination is reported for 65 families, including Bromeliaceae. One of the most diverse bromeliad is Billbergia, which comprises species pollinated mainly by hummingbirds.
• Based on investigations on flowering phenology, morpho‐anatomy, volume and concentration of nectar, pollinators and breeding system, this paper explores the reproductive biology and pollinator specificity of B. distachia in a mesophytic semi‐deciduous forest of southeastern Brazil.
• The results have show that B. distachia is pollinated by a single species of hermit hummingbird, Phaethornis eurynome, which search for nectar produced by a septal nectary, where the secretory tissue is located above the placenta. The species is self‐incompatible. The combination of pollinator specificity, due to long corolla tubes that exclude visitation of short‐billed hummingbirds, complete self‐incompatibility and non‐territorial behaviour of pollinators, it is very important to reduce pollen loss and increase gene flow within population.
• Our results indicate that studies on pollination biology and reproduction are essential to understand the evolutionary history of pollination systems of plants since, at least in Billbergia, variation in the pollinator spectrum has been recorded for different habitats among Brazilian forests. Furthermore, according to our data, foraging of Phaethornis on flowers is independent of air temperature and humidity, while the main factor influencing hummingbird visitation is daylight. Considering current knowledge on climatic parameters influencing hummingbird foraging, pollination and reproductive biology of Neotropical flora and environment of the hermit hummingbird in tropical forests, new insights on plant–pollinator interaction are provided.

     

Reproductive biology of Oreocereus fossulatus (Cactaceae), a long‐lived columnar cactus endemic to the tropical Andes

The geographic dichotomy hypothesis suggests that columnar cacti inhabiting the tropics depend on flower visitors (birds and bats) for their pollination, showing highly specialized animal‐pollinated systems. This pattern has been demonstrated for the northern hemisphere; however, our knowledge of the species of columnar cacti growing in the southern tropics is still scarce. In this study, we studied the reproductive biology of Oreocereus fossulatus (Cactaceae, Tribe Trichocereae), an endemic, columnar cactus of the tropical Andes, to determine if its pollination system tends to be more generalized (mixed systems of autogamy and xenogamy) than specialized (xenogamy) as a consequence of the geographic position of where it lives. Observations of the frequency of visits showed that Patagona gigas (Trochillidae) is its main pollinator. It visits the flowers when they are open, coinciding with the periods of greater floral reward (dusk and dawn of the first day of anthesis). The treatments of autogamy, xenogamy and geitonogamy produced fruits, showing that O. fossulatus exhibits a generalized pollination system, in the same way as its congeneric species O. celsianus, which is distributed farther south in the Prepuna biogeographical region. Our results suggest that species partially specializing in pollination by hummingbirds and, besides, capable of autopollination, could be common in the tropical and subtropical Andes, probably as a response to the unpredictable environments present in their ecosystems.     

Specialization clines in the pollination systems of agaves (Agavaceae) and columnar cacti (Cactaceae): A phylogenetically controlled meta-analysis

The biogeography of plant-animal interactions is a novel topic on which many disciplines converge (e.g., reproductive biology, biogeography, and evolutionary biology). Narrative reviews have indicated that tropical columnar cacti and agaves have highly specialized pollination systems, while extratropical species have generalized systems. However, this dichotomy has never been quantitatively tested. We tested this hypothesis using traditional and phylogenetically informed meta-analysis. Three effect sizes were estimated from the literature: diurnal, nocturnal, and hand cross-pollination (an indicator of pollen limitation). Columnar cactus pollination systems ranged from purely bat-pollinated in the tropics to generalized pollination, with diurnal visitors as effective as nocturnal visitors in extratropical regions; even when phylogenetic relatedness among species is taken into account. Metaregressions identified a latitudinal increase in pollen limitation in columnar cacti, but this increase was not significant after correcting for phylogeny. The currently available data for agaves do not support any latitudinal trend. Nectar production of columnar cacti varied with latitude. Although this variation is positively correlated with pollination by diurnal visitors, it is influenced by phylogeny. The degree of specificity in the pollination systems of columnar cacti is heavily influenced by ecological factors and has a predictable geographic pattern.     

Geographic variation in cone volatiles and pollinators in the thermogenic African cycad Encephalartos ghellinckii Lem

• Heat and odour production can have profound effects on pollination in cycads. It is therefore expected that these traits would co‐vary geographically with pollinator assemblages. Such intraspecific variation, may lead to the evolution of pollination ecotypes, which can be an early stage of pollinator‐mediated speciation.
• We measured cone temperatures using miniature temperature data loggers and examined the composition of cone volatile odours using headspace sampling and analysis with gas chromatography‐mass spectrometry in four populations spanning the range of the African cycad Encephalartos ghellinckii. Pollinator assemblages were also investigated in three populations.
• Male and female cones were thermogenic at pollen shed and receptive stages, respectively, but patterns of thermogenesis did not vary among populations. Scent emissions from cones in populations in the Drakensberg Mountains were characterised by cis‐β‐ocimene, β‐myrcene and (3E)‐1,3‐octadiene, while camphene and α‐pinene were characteristic of scent emissions from cones in populations closer to the coast. These differences in volatile blends corresponded with differences in insect assemblages.
• These results confirm intraspecific variation in volatile emissions of E. ghellinckii and support the predictions that intraspecific variation in volatile emissions will be associated with shifts in pollinator assemblages. While further work needs to be done to test for local adaptation in this system, this preliminary evidence is consistent with the formation of pollination ecotypes in the E. ghellinckii species complex.

     

Patterns of speciation are similar across mountainous and lowland regions for a Neotropical plant radiation (Costaceae: Costus)

High species richness and endemism in tropical mountains are recognized as major contributors to the latitudinal diversity gradient. The processes underlying mountain speciation, however, are largely untested. The prevalence of steep ecogeographic gradients and the geographic isolation of populations by topographic features are predicted to promote speciation in mountains. We evaluate these processes in a species-rich Neotropical genus of understory herbs that range from the lowlands to montane forests and have higher species richness in topographically complex regions. We ask whether climatic niche divergence, geographic isolation, and pollination shifts differ between mountain-influenced and lowland Amazonian sister pairs inferred from a 756-gene phylogeny. Neotropical Costus ancestors diverged in Central America during a period of mountain formation in the last 3 million years with later colonization of Amazonia. Although climatic divergence, geographic isolation, and pollination shifts are prevalent in general, these factors do not differ between mountain-influenced and Amazonian sister pairs. Despite higher climatic niche and species diversity in the mountains, speciation modes in Costus appear similar across regions. Thus, greater species richness in tropical mountains may reflect differences in colonization history, diversification rates, or the prevalence of rapidly evolving plant life forms, rather than differences in speciation mode.     

Evolutionary consequences of ecological factors: pollinator reliability predicts mating‐system traits of a perennial plant

The reproductive‐assurance hypothesis predicts that mating‐system traits will evolve towards increased autonomous self‐pollination in plant populations experiencing unreliable pollinator service. We tested this long‐standing hypothesis by assessing geographic covariation among pollinator reliability, outcrossing rates, heterozygosity and relevant floral traits across populations of Dalechampia scandens in Costa Rica. Mean outcrossing rates ranged from 0.16 to 0.49 across four populations, and covaried with the average rates of pollen arrival on stigmas, a measure of pollinator reliability. Across populations, genetically based differences in herkogamy (anther–stigma distance) were associated with variation in stigmatic pollen loads, outcrossing rates and heterozygosity. These observations are consistent with the hypothesis that, when pollinators are unreliable, floral traits promoting autonomous selfing evolve as a mechanism of reproductive assurance. Extensive covariation between floral traits and mating system among closely related populations further suggests that floral traits influencing mating systems track variation in adaptive optima generated by variation in pollinator reliability.     

Variation in floral morphology and plant reproductive success in four Ipomoea species (Convolvulaceae) with contrasting breeding systems

• This study tested the hypothesis that self‐compatibility would be associated with floral traits that facilitate autonomous self‐pollination to ensure reproduction under low pollinator visitation. In a comparison of two pairs of Ipomoea species with contrasting breeding systems, we predicted that self‐compatible (SC) species would have smaller, less variable flowers, reduced herkogamy, lower pollinator visitation and higher reproductive success than their self‐incompatible (SI) congeners.
• We studied sympatric species pairs, I. hederacea (SC)– I. mitchellae (SI) and I. purpurea (SC)–I. indica (SI), in Mexico, over two years. We quantified variation in floral traits and nectar production, documented pollinator visitation, and determined natural fruit and seed set. Hand‐pollination and bagging experiments were conducted to determine potential for autonomous self‐pollination and apomixis.
• Self‐compatible Ipomoea species had smaller flowers and lower nectar production than SI species; however, floral variation and integration did not vary according to breeding system. Bees were primary pollinators of all species, but visitation rates were seven times lower in SC than SI species. SC species had a high capacity for autonomous self‐pollination due to reduced herkogamy at the highest anther levels. Self‐compatible species had two to six times higher fruit set than SI species.
• Results generally support the hypothesis that self‐compatibility and autonomous self‐pollination ensure reproduction under low pollinator visitation. However, high variation in morphological traits of SC Ipomoea species suggests they maintain variation through outcrossing. Furthermore, reduced herkogamy was associated with high potential for autonomous self‐pollination, providing a reproductive advantage that possibly underlies transitions to self‐compatibility in Ipomoea.

     

The least effective pollinator principle: specialized morphology despite generalized ecology

• The large body of work on the adaptation of plants to pollinators is still somewhat incomplete because most studies focus on one‐to‐one interactions. How will adaptation proceed in a multi‐pollinator environment? According to Stebbins' Most Effective Pollinator Principle, ‘the characteristics of the flower will be moulded by those pollinators that visit it most frequently and effectively.’
• To test this hypothesis, we studied the pollination biology of Pelargonium incrassatum (Geraniaceae) in the Namaqualand Region of Southern Africa. This species has a long floral tube and we expected its most important pollinator to have a long proboscis.
• Contrary to expectations, the most important pollinator was a short proboscid fly (a new species of Prosoeca), while Prosoeca peringueyi, which had a proboscis that matched the floral tube length, was a rare visitor. Consistent with the high degree of trait mismatching, we did not detect selection on tube length at most sites.
• The paradox of mismatching traits can be resolved by considering the strength of the trade‐off involved. Adaptation to the rare species can apparently occur without incurring the cost of reduced pollination by the abundant species. Generally, species may often evolve specialized morphology if they do not incur the cost of ecological specialization.

     

Evolutionary disruption in the pollination system of Vanilla (Orchidaceae)

• Pollination of the pantropical Vanilla has been linked to melittophily and food deception.
• Here we investigated the role of flower traits on the reproduction of Neotropical Vanilla. We also studied the evolution of pollination systems in order to understand the origin of production of flower resources and the diversification of pollinators in this orchid genus. Our study was founded on data of adaptations in flower morphology, production of resources, scent release, pollinators and breeding systems of Vanilla and presenting new data on reproductive biology of V. palmarum. Data on reproductive biology of Vanilla were mapped onto a phylogeny to address our queries on the evolution of pollination systems in this genus.
• Vanilla palmarum shows a mixed mating system, with its facultative autogamous flowers being pollinated by hummingbirds. Its yellow flowers are scentless and produces nectar. Mapping of the pollination system onto trees resulted in one origin for bird pollination and at least two origins for autogamy in Vanilla. Nectar secretion has a single origin in the Neotropical thick-leafed lineage.
• Bird pollination of Vanilla is shown for the first time. The origin of ornithophily within a bee-pollinated clade is supported by flower morphology. Floral transitions to ornithophily have been favoured by the occupation of a distinct niche from that of the other thick-leafed Vanilla species. Despite its specialized pollination, V. palmarum is autogamous. A mixed mating system can promote reproductive assurance in the case of a decline in pollinator populations, or in areas where pollinator services are irregular or absent.

     

Relationships between wild bees,hoverflies and pollination success in apple orchards with different landscape contexts

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