Long-term observation of a pollination network: fluctuation in species and interactions, relative invariance of network structure and implications for estimates of specialization

We analysed the dynamics of a plant-pollinator interaction network of a scrub community surveyed over four consecutive years. Species composition within the annual networks showed high temporal variation. Temporal dynamics were also evident in the topology of the network, as interactions among plants and pollinators did not remain constant through time. This change involved both the number and the identity of interacting partners. Strikingly, few species and interactions were consistently present in all four annual plant-pollinator networks (53% of the plant species, 21% of the pollinator species and 4.9% of the interactions). The high turnover in species-to-species interactions was mainly the effect of species turnover (c. 70% in pairwise comparisons among years), and less the effect of species flexibility to interact with new partners (c. 30%). We conclude that specialization in plant-pollinator interactions might be highly overestimated when measured over short periods of time. This is because many plant or pollinator species appear as specialists in 1 year, but tend to be generalists or to interact with different partner species when observed in other years. The high temporal plasticity in species composition and interaction identity coupled with the low variation in network structure properties (e.g. degree centralization, connectance, nestedness, average distance and network diameter) imply (i) that tight and specialized coevolution might not be as important as previously suggested and (ii) that plant-pollinator interaction networks might be less prone to detrimental effects of disturbance than previously thought. We suggest that this may be due to the opportunistic nature of plant and animal species regarding the available partner resources they depend upon at any particular time.     

Long-term temporal variation in the organization of an ant–plant network

Background and Aims

Functional groups of species interact and coevolve in space and time, forming complex networks of interacting species. A long-term study of temporal variation of an ant–plant network is presented with the aims of: (1) depicting its structural changes over a 20-year period; (2) detailing temporal variation in network topology, as revealed by nestedness and modularity analysis and other parameters (i.e. connectance, niche overlap); and (3) identifying long-term turnover in taxonomic structure (i.e. switches in ant resource use or plant visitor assemblages according to taxa).

Methods

Fieldwork was carried out at La Mancha, Mexico, and ant–plant interactions were observed between 1989 and 1991, between 1998 and 2000, and between May 2010 and 2011. Occurrences of ants on extrafloral nectaries (EFNs) were recorded. The resulting ant–plant networks were constructed from qualitative presence–absence data determined by a species–species matrix defined by the frequency of occurrence of each pairwise ant–plant interaction.

Key Results

Network variation across time was stable and a persistent nested structure may have contributed to the maintenance of resilient and species-rich communities. Modularity was lower than expected, especially in the most recent networks, indicating that the community exhibited high overlap among interacting species (e.g. few species were hubs in the more recent network, being partly responsible for the nested pattern). Structurally, the connections created among modules by super-generalists gave cohesion to subsets of species that otherwise would remain unconnected. This may have allowed an increasing cascade-effect of evolutionary events among modules. Mutualistic ant–plant interactions were structured 20 years ago mainly by the subdominant nectarivorous ant species Camponotus planatus and Crematogaster brevispinosa, which monopolized the best extrafloral nectar resources and out-competed other species with broader feeding habits. Through time, these ants, which are still present, lost their position as network hubs and diminished in their importance in structuring the network; simultaneously, plants gained in importance.

Conclusions

The long-term network analysis reveals a decrease in attended plant species richness, a notable increase in plant species participation from 1990 to 2010 (sustained by less plant taxonomic similarity in the older 1990 network), an increase in the number of ant species and a diminishing dominance of super-generalist ants. The structure of the community has remained highly nested and connected with low modularity, suggesting overall a more participative, homogeneous, cohesive interaction network. Although previous studies have suggested that interactions between ants and EFN-bearing plants are susceptible to seasonality, abiotic factors and perturbation, this cohesive structure appears to be the key for biodiversity and community maintenance.     

Invasive plant integration into native plant–pollinator networks across Europe

The structure of plant–pollinator networks has been claimed to be resilient to changes in species composition due to the weak degree of dependence among mutualistic partners. However, detailed empirical investigations of the consequences of introducing an alien plant species into mutualistic networks are lacking. We present the first cross-European analysis by using a standardized protocol to assess the degree to which a particular alien plant species (i.e. Carpobrotus affine acinaciformis, Impatiens glandulifera, Opuntia stricta, Rhododendron ponticum and Solanum elaeagnifolium) becomes integrated into existing native plant–pollinator networks, and how this translates to changes in network structure.Alien species were visited by almost half of the pollinator species present, accounting on average for 42 per cent of the visits and 24 per cent of the network interactions. Furthermore, in general, pollinators depended upon alien plants more than on native plants. However, despite the fact that invaded communities received more visits than uninvaded communities, the dominant role of alien species over natives did not translate into overall changes in network connectance, plant linkage level and nestedness. Our results imply that although supergeneralist alien plants can play a central role in the networks, the structure of the networks appears to be very permeable and robust to the introduction of invasive alien species into the network.     

Global patterns of intraspecific leaf trait responses to elevation

Elevational gradients are often used to quantify how traits of plant species respond to abiotic and biotic environmental variations. Yet, such analyses are frequently restricted spatially and applied along single slopes or mountain ranges. Since we know little on the response of intraspecific leaf traits to elevation across the globe, we here perform a global meta‐analysis of leaf traits in 109 plant species located in 4 continents and reported in 71 studies published between 1983 and 2018. We quantified the intraspecific change in seven morpho‐ecophysiological leaf traits along global elevational gradients: specific leaf area (SLA), leaf mass per area (LMA), leaf area (LA), nitrogen concentration per unit of area (Narea), nitrogen concentration per unit mass (Nmass), phosphorous concentration per unit mass (Pmass) and carbon isotope composition (δ¹³C). We found LMA, Narea, Nmass and δ¹³C to significantly increase and SLA to decrease with increasing elevation. Conversely, LA and Pmass showed no significant pattern with elevation worldwide. We found significantly larger increase in Narea, Nmass, Pmass and δ¹³C with elevation in warmer regions. Larger responses to increasing elevation were apparent for SLA of herbaceous compared to woody species, but not for the other traits. Finally, we also detected evidences of covariation across morphological and physiological traits within the same elevational gradient. In sum, we demonstrate that there are common cross‐species patterns of intraspecific leaf trait variation across elevational gradients worldwide. Irrespective of whether such variation is genetically determined via local adaptation or attributed to phenotypic plasticity, the leaf trait patterns quantified here suggest that plant species are adapted to live on a range of temperature conditions. Since the distribution of mountain biota is predominantly shifting upslope in response to changes in environmental conditions, our results are important to further our understanding of how plants species of mountain ecosystems adapt to global environmental change.     

Plant richness and blooming cover affect abundance of flower visitors and network structure in Colombian orchards

1. Pollination is essential to fruit production. How plant diversity and blooming events in and around orchards affect the pollinator community and the plant-flower-visitor network in neotropical systems remains largely unknown.
2. We surveyed the flower visitors in deciduous fruit trees and alternative blooming resources (other crops, hedgerows and weeds) in Colombia across 6 orchards over 12 months. We evaluated whether plant species richness and blooming cover influenced abundance and richness of flower visitors, as well as network-level connectance and specialization. We also assessed the role of alternative blooming resources for the flower visitors of deciduous fruit trees.
3. Overall, we found 66 taxa of flower visitors, 35 of which visited deciduous fruit trees. There was a greater abundance of flower visitors when there was higher richness of weedy species and greater blooming cover of deciduous fruit trees. Networks were less connected when there was lower crop and weedy species richness. Finally, flower visitor abundance and specialization increased when there were multiple hedgerow species in bloom with a high blooming cover.
4. We highlight the importance of maintaining alternative blooming resources in and around the orchards to support deciduous fruit tree pollinators and diversity in the plant flower-visitor network.

     

Individual variation in resource use by opossums leading to nested fruit consumption

Despite recent findings on the ecological relevance of within population diet variation far less attention has been devoted to the role diet variation for ecological services. Seed dispersal is a key ecological service, affecting plant fitness and regeneration based on foraging by fruit‐eating vertebrates. Here we used a network approach, widely used to understand how seed‐dispersal is organized at the species level, to gain insights into the patterns that emerge at the individual‐level. We studied the individual fruit consumption behavior of a South American didelphid Didelphis albiventris, during the cool–dry and warm–wet seasons. In species–species networks the heterogeneity in specialization levels generates patterns such as nestedness and asymmetry. Because generalist populations may be comprised of specialized individuals, we hypo thesized that network structural properties, such as nestedness, should also emerge at the individual level. We detected variation in fruit consumption that was not related to resource availability, ontogenetic or sexual factors or sampling biases. Such variation resulted in the structural patterns often found in species–species seed‐dispersal networks: low connectance, a high degree of nestedness and the absence of modules. Moreover structure varied between the warm–wet and cool–dry seasons, presumably as a consequence of seasonal fluctuation in fruit availability. Our findings suggest individuals may differ in selectivity causing asymmetries in seed dispersal efficiency within the population. In this sense the realized dispersal would differ from the expected dispersal estimated from their average dispersal potential. Additionally the results suggest possible frequency‐dependent effects on seed dispersal that might affect individual plant performance and plant community composition.     

The distribution of vascular plant species and guilds in space and time along a desert gradient

Abstract. We studied distribution patterns of vascular plant species and environmental variables for three years along a permanent transect traversing a closed-drainage watershed in the northern Chihuahuan Desert of south-central New Mexico, USA. The transect extended for 2.7 km from a basin floor playa (1310 m elevation, fine-textured soil), across a piedmont slope, and onto the base of a granitic mountain (1410 m elevation, coarse-textured soil). The gradients in elevation and soils across our transect, along with variable seasonal rainfall, downslope redistribution of water and organic matter, and soil texture-related variation in infiltration, water holding capacity, and moisture release characteristics, interact to generate a complex spatial and temporal gradient of available soil water and nitrogen. We grouped plant species into guilds according to growth form and photo synthetic pathway type. These guilds are spatially and temporally differentiated along the transect such that particular groups utilize particular seasonal phases or spatial regions of the gradient. We identified six distinct plant communities along the transect. C₄ perennial grasses dominated the mesic/high nitrogen portion of the gradient, which occurred at the highest (upper piedmont grassland, dominated by Bouteloua eriopoda) and lowest (playa, dominated by Panicum obtusum) elevations along the transect. C₃ shrubs were dominant in the xeric/low nitrogen portion of the gradient located near the middle of the transect (bajada shrubland, dominated by Larrea tridentata). C₃ shrubs also dominated a narrow zone of vegetation located adjacent to the playa (playa fringe, dominated by Prosopisglandulosa). C₄ perennial grasses, C₃ subshrubs, and C₃ and C₄ perennial forbs and annuals were co-dominant in the intermediate locations along the gradient, which occurred below (mixed basin slopes) and above (lower piedmont grassland) the bajada shrubland. Life-form distribution patterns at the small scale of our study reflect some of the patterning that occurs at larger scales in response to climate gradients. The distributions of some species and guilds along the transect are apparently modified by competitive interactions.     

传粉网络非对称特化的地理模式

植物与传粉者间相互作用,构成了复杂的传粉网络。非对称特化是共生互作网络中的有趣现象和基本特点,也被认为是植物-传粉者互作网络的结构特征之一。根据文献总结分析了植物-传粉者互作网络非对称特化的重要名词术语,并采用线性回归法深入分析了植物-传粉者互作网络的地理变异模式,以及植物生活型和网络大小等传粉网络特征对非对称程度的影响。结果表明:传粉网络大小与网络的交互作用间呈线性正相关关系,并随总物种丰度呈指数增长。25个传粉网络的线性回归斜率(Lβ)变异范围在0.002至0.031间,且斜率值随植物丰度(P)、传粉者丰度(A)、总物种丰度(R)、交互作用(I)及网络大小(M)上升而降低。海拔高度对传粉网络非对称性有一定影响效果,而纬度的变化并不显著影响传粉网络非对称性。草本植物、灌木及乔木植物与其传粉者之间的相关系数分别为-0.197,-0.026和0.200,表明草本物种比乔木物种非对称性更强。     

Spatio-temporal variation in the structure of pollination networks

Pollination networks are representations of all interactions between co-existing plants and their flower visiting animals at a given site. Although the study of networks has become a distinct sub-discipline in pollination biology, few studies have attempted to quantify spatio-temporal variation in species composition and structure of networks. We here investigate patterns of year-to-year change in pollination networks from six different sites spanning a large latitudinal gradient. We quantified level of species persistence and interactions among years, and examined year-to-year variation of network structural parameters in relation to latitude and sampling effort. In addition, we tested for correlations between annual variation in network parameters and short and long-term climate change variables. Numbers of plant and animal species and interactions were roughly constant from one year to another at all sites. However, composition of species and interactions changed from one year to another. Turnover was particularly high for flower visitors and interactions. On the other hand, network structural parameters (connectance, nestedness, modularity and centralization) remained remarkably constant between years, regardless of network size and latitude. Inter-annual variation of network parameters was not related to short or long term variation in climate variables (mean annual temperature and annual precipitation). We thus conclude that pollination networks are highly dynamic and variable in composition of species and interactions among years. However, general patterns of network structure remain constant, indicating that species may be replaced by topologically similar species. These results suggest that pollination networks are to some extent robust against factors affecting species occurrences.     

Effect of temporal data aggregation on the perceived structure of a quantitative plant–floral visitor network

Seasonal turnover in plant and floral visitor communities changes the structure of the network of interactions they are involved in. Despite the dynamic nature of plant–visitor networks, a usual procedure is to pool year‐round interaction data into a single network which may result in a biased depiction of the real structure of the interaction network. The annual temporal dynamics and the effect of merging monthly data have previously been described for qualitative data (i.e. describing the occurrence of interactions) alone, while its quantitative aspect (i.e. the actual frequency with which interactions occur) remain little explored. For this, we built a set of 12 monthly networks describing year‐round plant–floral visitor interactions in a 30‐hectare planted forest and its adjacent agricultural landscape at Bahauddin Zakariya University Multan, Pakistan. A total of 80 plant and 162 insect species, which engaged in 1573 unique interactions, were recorded. Most network properties (particularly the number of plants, visitors and unique interactions) varied markedly during the year. Data aggregation showed that while animal species, plant species, unique interaction, weighted nestedness, interaction diversity and robustness increased, connectance and specialization decreased. The only metric which seemed relatively unaffected by data pooling was interaction evenness. In general, quantitative metrics were relatively less affected by temporal data aggregation than qualitative ones. Avoiding data aggregation not only gives a more realistic depiction of the dynamic nature of plant–visitor community networks, but also avoids biasing network metrics and, consequently, their expected response to disturbances such as the loss of species.     

Population sinks resulting from degraded habitats of an obligate life-history pathway

In networks of plant–animal mutualisms, different animal groups interact preferentially with different plants, thus forming distinct modules responsible for different parts of the service. However, what we currently know about seed dispersal networks is based only on birds. Therefore, we wished to fill this gap by studying bat–fruit networks and testing how they differ from bird–fruit networks. As dietary overlap of Neotropical bats and birds is low, they should form distinct mutualistic modules within local networks. Furthermore, since frugivory evolved only once among Neotropical bats, but several times independently among Neotropical birds, greater dietary overlap is expected among bats, and thus connectance and nestedness should be higher in bat–fruit networks. If bat–fruit networks have higher nestedness and connectance, they should be more robust to extinctions. We analyzed 1 mixed network of both bats and birds and 20 networks that consisted exclusively of either bats (11) or birds (9). As expected, the structure of the mixed network was both modular (M = 0.45) and nested (NODF = 0.31); one module contained only birds and two only bats. In 20 datasets with only one disperser group, bat–fruit networks (NODF = 0.53 ± 0.09, C = 0.30 ± 0.11) were more nested and had a higher connectance than bird–fruit networks (NODF = 0.42 ± 0.07, C = 0.22 ± 0.09). Unexpectedly, robustness to extinction of animal species was higher in bird–fruit networks (R = 0.60 ± 0.13) than in bat–fruit networks (R = 0.54 ± 0.09), and differences were explained mainly by species richness. These findings suggest that a modular structure also occurs in seed dispersal networks, similar to pollination networks. The higher nestedness and connectance observed in bat–fruit networks compared with bird–fruit networks may be explained by the monophyletic evolution of frugivory in Neotropical bats, among which the diets of specialists seem to have evolved from the pool of fruits consumed by generalists.     

Contrasting structures of plant–mite networks compounded by phytophagous and predatory mite species

Differences in the feeding habits between phytophagous and predatory species can determine distinct ecological interactions between mites and their host plants. Herein, plant–mite networks were constructed using available literature on plant-dwelling mites from Brazilian natural vegetation in order to contrast phytophagous and predatory mite networks. The structural patterns of plant–mite networks were described through network specialization (connectance) and modularity. A total of 187 mite species, 65 host plant species and 646 interactions were recorded in 14 plant–mite networks. Phytophagous networks included 96 mite species, 61 host plants and 277 interactions, whereas predatory networks contained 91 mite species, 54 host plants and 369 interactions. No differences in the species richness of mites and host plants were observed between phytophagous and predatory networks. However, plant–mite networks composed of phytophagous mites showed lower connectance and higher modularity when compared to the predatory mite networks. The present results corroborate the hypothesis that trophic networks are more specialized than commensalistic networks, given that the phytophagous species must deal with plant defenses, in contrast to predatory mites which only inhabit and forage for resources on plants.     

Insect-Flower Interaction Network Structure Is Resilient to a Temporary Pulse of Floral Resources from Invasive Rhododendron ponticum

Invasive alien plants can compete with native plants for resources, and may ultimately decrease native plant diversity and/or abundance in invaded sites. This could have consequences for native mutualistic interactions, such as pollination. Although invasive plants often become highly connected in plant-pollinator interaction networks, in temperate climates they usually only flower for part of the season. Unless sufficient alternative plants flower outside this period, whole-season floral resources may be reduced by invasion. We hypothesized that the cessation of flowering of a dominant invasive plant would lead to dramatic, seasonal compositional changes in plant-pollinator communities, and subsequent changes in network structure. We investigated variation in floral resources, flower-visiting insect communities, and interaction networks during and after the flowering of invasive Rhododendron ponticum in four invaded Irish woodland sites. Floral resources decreased significantly after R. ponticum flowering, but the magnitude of the decrease varied among sites. Neither insect abundance nor richness varied between the two periods (during and after R. ponticum flowering), yet insect community composition was distinct, mostly due to a significant reduction in Bombus abundance after flowering. During flowering R. ponticum was frequently visited by Bombus; after flowering, these highly mobile pollinators presumably left to find alternative floral resources. Despite compositional changes, however, network structural properties remained stable after R. ponticum flowering ceased: generality increased, but quantitative connectance, interaction evenness, vulnerability, H’₂ and network size did not change. This is likely because after R. ponticum flowering, two to three alternative plant species became prominent in networks and insects increased their diet breadth, as indicated by the increase in network-level generality. We conclude that network structure is robust to seasonal changes in floral abundance at sites invaded by alien, mass-flowering plant species, as long as alternative floral resources remain throughout the season to support the flower-visiting community.     

A year‐long plant‐pollinator network

Abstract In this work we analyse the pollination community in a South American forest known as ‘talar’. This is a vegetal woody community that inhabits fossil coastal banks characterized by seasonal temperate weather and calcareous soil, at the coast of the Río de la Plata, in the province of Buenos Aires, Argentina. We obtained data of the interactions between anthophylous insects and entomophylous flowering plants over an extensive period of time. We showed that pollination system parameters, such as partners’ identity, system size, and connectance, fluctuated among months, when sampled year‐long. Maximal network size occurred in early spring and early autumn, when both the number of mutualistic species and the number of interactions peaked, and this was also when network asymmetry was higher than average. Monthly connectance of the plant‐flower visitor matrix decreased to its lowest values at these peaks. Available data suggest that cumulative traditional connectance (i.e. the connectance calculated as the whole number of interactions registered in the community divided by the full size system) underestimates actual connectance values by a factor of c. 3 ×. Monthly values of connectance decreased exponentially as system size increased, and the distribution of interactions per species followed power‐law regimes for animals, and truncated power‐law regimes for plants, in accordance with patterns previously deduced from among‐network cumulative communities studies. We think that either within or and among pollination networks, systems that are organized as power‐law regimes may be a basic property of these webs, and provide examples of the fact. Both seasonal changes and interactions between mutualists like competition, and some degree of facilitation, may be very important to understand the performance of the system as a whole, and the role and importance of different species in the community. We suggest that communities of plant – pollinators that exhibit extended activity, such as temperate or tropical seasonal ones, should be studied through consecutive plant‐pollinator webs rather than cumulative ones. The partition of the system into smaller serial parts allows us to obtain outstanding information of every short period. This information is flattened by the average effect when we considered the combined analysis of the whole data.     

Native and Non-Native Supergeneralist Bee Species Have Different Effects on Plant-Bee Networks

Supergeneralists, defined as species that interact with multiple groups of species in ecological networks, can act as important connectors of otherwise disconnected species subsets. In Brazil, there are two supergeneralist bees: the honeybee Apis mellifera, a non-native species, and Trigona spinipes, a native stingless bee. We compared the role of both species and the effect of geographic and local factors on networks by addressing three questions: 1) Do both species have similar abundance and interaction patterns (degree and strength) in plant-bee networks? 2) Are both species equally influential to the network structure (nestedness, connectance, and plant and bee niche overlap)? 3) How are these species affected by geographic (altitude, temperature, precipitation) and local (natural vs. disturbed habitat) factors? We analyzed 21 plant-bee weighted interaction networks, encompassing most of the main biomes in Brazil. We found no significant difference between both species in abundance, in the number of plant species with which each bee species interacts (degree), and in the sum of their dependencies (strength). Structural equation models revealed the effect of A. mellifera and T. spinipes, respectively, on the interaction network pattern (nestedness) and in the similarity in bee’s interactive partners (bee niche overlap). It is most likely that the recent invasion of A. mellifera resulted in its rapid settlement inside the core of species that retain the largest number of interactions, resulting in a strong influence on nestedness. However, the long-term interaction between native T. spinipes and other bees most likely has a more direct effect on their interactive behavior. Moreover, temperature negatively affected A. mellifera bees, whereas disturbed habitats positively affected T. spinipes. Conversely, precipitation showed no effect. Being positively (T. spinipes) or indifferently (A. mellifera) affected by disturbed habitats makes these species prone to pollinate plant species in these areas, which are potentially poor in pollinators.     

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.     

Mammalian seed dispersal in Cantabrian woodland pastures: Network structure and response to forest loss

We evaluated the role of wild large mammals as dispersers of fleshy-fruited woody plants in woodland pastures of the Cantabrian range (N Spain). By searching for seeds in mammal scats across four localities, we addressed how extensive seed dispersal was in relation to the fleshy-fruited plant community, and applied a network approach to identify the relative role of mammal species in the seed dispersal process. We also tested the response of mammalian dispersers to forest availability at increasing spatial scales. Five carnivores and three ungulates dispersed seeds of eight fleshy-fruited trees and shrubs. Mammalian seed dispersal did not mirror community-wide fruit availability, as abundant fruiting trees were scarce whereas thorny shrubs were over-represented among dispersed species. The dispersal network was dominated by bramble (Rubus ulmifolius/fruticosus), the remaining plants being rarer and showing more restricted disperser coteries. Fox (Vulpes vulpes), badger (Meles meles), and wild boar (Sus scrofa) dispersed mostly bramble, whereas martens (Martes sp.) dispersed mostly wild rose (Rosa sp.). Ungulates occasionally dispersed holly (Ilex aquifolium) and hawthorn (Crataegus monogyna). The empirical network reflected a skewed distribution of interactions and some functional complementarity (as judged from the low levels of connectance and nestedness), but also some degree of specialization. Mammals overused uncovered microsites for seed deposition, and increased their disperser activity in those landscape sectors devoid of forest. Combined with previous findings on avian seed dispersal, this study suggest a strong functional complementarity coming from the low overlap in the main plant types that mammals and birds disperse – thorny shrubs and trees, respectively – and the differential patterns of seed deposition, with mammals mostly dispersing into deforested areas, and birds into forest-rich landscapes.     

Characterizing topology of ecological networks along gradients: The limits of metrics’ standardization

Species interact in nature to form complex ecological networks. There has been a rising interest in recent years to characterize the topology of such networks along various gradients (e.g. successional, climatic, elevational) to better understand how they assemble in space and time. However, to compare structure of networks that vary in size, shape and connectance, topological metrics need to be standardized (as most metrics covary with such network attributes). Traditionally, this has been done by transforming network metrics into z-scores prior comparisons. Here, I show that such standardized metrics are not independent of basic network properties such as connectance. Instead, I found that there was a consistent tendency for z-scores to approach 0 when connectance progressively decreased and approached its minimal value. This is probably due to the reduced null space available for null models to randomize interactions at such low connectance. I discuss ways to circumvent the problem in future studies.     

Comparative biogeography of the congener lilies Lilium distichum and Lilium tsingtauense in Korea

The main Korean mountain range (the so-called “Baekdudaegan”), which stretches from north to south across most of the Peninsula, has been thought to harbor glacial refugia for boreal and temperate plant species, where they likely found relatively stable habitats and maintained large population sizes. Under this scenario, high levels of genetic variation for plant populations can be anticipated. To test this hypothesis, we examined levels of allozyme diversity in the boreal herb Lilium distichum, which in Korea occurs largely along the Baekdudaegan and in its closely related congener L. tsingtauense, a temperate species that can occur from the coastal islands to high-altitude mountains in the Baekdudaegan. As expected, L. distichum harbored high levels of genetic variation within populations, and we found significant correlations between percentage of polymorphic loci (and allelic richness) and elevation. For L. tsingtauense we found moderate genetic variation, with its populations showing a significantly positive correlation between intra-population expected heterozygosity and elevation. Our results suggest that populations/species that occur within or near the main ridge of the Baekdudaegan probably endured the glacial periods in macrorefugia, whereas those populations/species located at low elevations (far away from mid- and high-elevation mountains) did it in microrefugia. Palaeodistribution modelling is in agreement with genetic data, indicating that the Baekdudaegan was suitable habitat for both species during the glacial (LGM) and interglacial periods (like the present). This study as well as the data from other studies compiled here provides strong evidence that the Baekdudaegan was a major refugial area for the East Asian flora throughout the Quaternary and, thus, meriting high priority for conservation biogeography.