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.     

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。     

Population dynamics of plant and pollinator communities: stability reconsidered

Plant-pollinator networks are systems of outstanding ecological and economic importance. A particularly intriguing aspect of these systems is their high diversity. However, earlier studies have concluded that the specific mechanisms of plant-pollinator interactions are destabilizing and should lead to a loss of diversity. Here we present a mechanistic model of plant and pollinator population dynamics with the ability to represent a broad spectrum of interaction structures. Using this model, we examined the influence of pollinators on the stability of a plant community and the relationship between pollinator specialization and stability. In accordance with earlier work, our results show that plant-pollinator interactions may severely destabilize plant coexistence, regardless of the degree of pollinator specialization. However, if plant niche differentiation, a classical stabilizing mechanism, is sufficiently strong to overcome the minority disadvantage with respect to pollination, interactions with pollinators may even increase the stability of a plant community. In addition to plant niche differentiation, the relationship between specialization and stability depends on a number of parameters that affect pollinator growth rates. Our results highlight the complex effects of this particular type of mutualism on community stability and call for further investigations of the mechanisms of diversity maintenance in plant-pollinator systems.     

60 million years of co-divergence in the fig-wasp symbiosis

Figs (Ficus; ca 750 species) and fig wasps (Agaoninae) are obligate mutualists: all figs are pollinated by agaonines that feed exclusively on figs. This extraordinary symbiosis is the most extreme example of specialization in a plant-pollinator interaction and has fuelled much speculation about co-divergence. The hypothesis that pollinator specialization led to the parallel diversification of fig and pollinator lineages (co-divergence) has so far not been tested due to the lack of robust and comprehensive phylogenetic hypotheses for both partners. We produced and combined the most comprehensive molecular phylogenetic trees to date with fossil data to generate independent age estimates for fig and pollinator lineages, using both non-parametric rate smoothing and penalized likelihood dating methods. Molecular dating of ten pairs of interacting lineages provides an unparalleled example of plant-insect co-divergence over a geological time frame spanning at least 60 million years.     

Burial disturbance leads to facilitation among coastal dune plants

There is growing evidence that interactions among plants can be facilitative as well as competitive, but knowledge of how disturbances influence these interactions and how they vary with species diversity is lacking. We manipulated plant density, species diversity (richness), and a burial disturbance in a controlled, complete factorial experiment to test theories about the relationships among species interactions, disturbance, and richness. The hypotheses tested were 1) burial disturbance reduces plant performance at all levels of density and richness, 2) burial disturbance can cause net plant interactions to become more facilitative, and 3) facilitation increases with species richness. Burial decreased plant survival by 60% and biomass by 50%, supporting the hypothesis that burial reduces plant performance. In the control (unburied) treatment, there was no difference in proportion survival or per plant biomass between low and high density plots, meaning that neither competition nor facilitation was detected. In the buried treatment, however, high density plots had significantly greater survival and greater per plant biomass than the low density plots, indicating net facilitative interactions. Thus facilitation occurred in the buried treatment and not in the unburied control plots, supporting the hypothesis that facilitation increases with increasing disturbance severity. The hypothesis that facilitation increases with increasing species richness was not supported. Richness did not affect survival or biomass, and there was no richness by burial treatment interaction, indicating that richness did not influence the response of the community to burial. The influence of the disturbance on plant interactions was thus consistent across levels of richness, increasing the generality of the relationship between disturbance and facilitation.     

Refuges from fire maintain pollinator–plant interaction networks

Fire is a major disturbance factor in many terrestrial ecosystems, leading to landscape transformation in fire‐prone areas. Species in mutualistic interactions are often highly sensitive to disturbances like fire events, but the degree and complexity of their responses are unclear. We use bipartite insect–flower interaction networks across a recently burned landscape to explore how plant–pollinator interaction networks respond to a recent major fire event at the landscape level, and where fire refuges were present. We also investigate the effectiveness of these refuges at different elevations (valley to hilltop) for the conservation of displaced flower‐visiting insects during fire events. Then, we explore how the degree of specialization of flower‐visiting insects changes across habitats with different levels of fire impact. We did this in natural areas in the Greater Cape Floristic Region (GCFR) biodiversity hotspot, which is species rich in plants and pollinators. Bees and beetles were the most frequent pollinators in interactions, followed by wasps and flies. Highest interaction activity was in the fire refuges and least in burned areas. Interactions also tracked flower abundance, which was highest in fire refuges in the valley and lowest in burned areas. Interactions consisted mostly of specialized flower visitors, especially in refuge areas. The interaction network and species specialization were lowest in burned areas. However, species common to at least two fire classes showed no significant difference in species specialization. We conclude that flower‐rich fire refuges sustain plant–pollinator interactions, especially those involving specialized species, in fire‐disturbed landscape. This may be an important shelter for specialized pollinator species at the time that the burned landscape goes through regrowth and succession as part of ecosystem recovery process after a major fire event.     

Does asymmetric specialization differ between mutualistic and trophic networks?

Recently, plant–pollinator networks have been found to be highly structured in a nested pattern in which specialists interact with generalist species. This structure is often assumed to be particular to mutualistic interactions in opposition to the compartmentalized pattern expected for antagonistic networks. We investigated the presence of asymmetric specialization in a data set assembled from the literature of 20 highly resolved plant–insect herbivore networks and compared them with 24 plant–pollinator networks. Our results indicate that these two types of networks differ, but not in the way it is generally assumed. Asymmetric specialization is present in plant–herbivore networks even if it appears less frequently than in plant–pollinator networks. Indeed, mean and median percentages of species showing asymmetric specialisation in herbivory webs are 33% and 14% respectively, compared to 57% and 60% in pollination webs. Furthermore, the amount of asymmetry is linked with species diversity and not to connectance in plant-pollinator networks whereas the opposite pattern is found in plant–herbivore networks. Our results offer promising perspectives for understanding both the mechanisms that structure ecological communities and their impact on community dynamics depending on the type of interaction.     

Responses of 'resistant' vertebrates to habitat loss and fragmentation: the importance of niche breadth and range boundaries

Abstract. An ability to predict species' sensitivities to habitat loss and fragmentation has important conservation implications, and numerous hypotheses have been proposed to explain interspecific differences observed in human-dominated landscapes. We used occupancy data collected on 32 species of vertebrates (16 mammals and 16 amphibians) in an agricultural landscape of Indiana, USA, to compare hypotheses that focus on different causal mechanisms underlying interspecific variation in responses to habitat alteration: (1) body size; (2) morphology and development; (3) behaviour; (4) niche breadth; (5) proximity to range boundary; and multiple-process models combining main effects and interactions of hypotheses (1)–(2) and (4)–(5). The majority of habitat alteration occurred over a century ago and coincided with extinction of several species; thus, our study dealt only with variation in responses of extant species that often are considered 'resistant' to human modifications of native habitat. Corrected Akaike scores and Akaike weights provided strongest support for models incorporating niche breadth and proximity to range boundary. Measures of dietary and habitat breadth obtained from the literature were negatively correlated with sensitivity to habitat alteration. Additionally, greater sensitivity was observed for species occurring at the periphery of their geographical ranges, especially at northern or western margins. Body size, morphological, developmental and behavioural traits were inferior predictors of tolerance to fragmentation for the species and landscape we examined. Our findings reinforce the importance of niche breadth as a predictor of species' responses to habitat alteration. They also highlight the importance of viewing the effects of habitat loss and fragmentation in a landscape within a biogeographical context that considers a species' level of adaptation to local environmental conditions.     

Asymmetric specialization and extinction risk in plant–flower visitor webs: a matter of morphology or abundance?

A recently discovered feature of plant–flower visitor webs is the asymmetric specialization of the interaction partners: specialized plants interact mainly with generalized flower visitors and specialized flower visitors mainly with generalized plants. Little is known about the factors leading to this asymmetry and their consequences for the extinction risk of species. Previous studies have proposed random interactions proportional to species abundance as an explanation. However, the simulation models used in these studies did not include potential biological constraints. In the present study, we tested the potential role of both morphological constraints and species abundance in promoting asymmetric specialization. We compared actual field data of a Mediterranean plant–flower visitor web with predictions of Monte Carlo simulations including different combinations of the potential factors structuring the web. Our simulations showed that both nectar-holder depth and abundance were able to produce asymmetry; but that the expected degree of asymmetry was stronger if based on both. Both factors can predict the number of interaction partners, but only nectar-holder depth was able to predict the degree of asymmetry of a certain species. What is more, without the size threshold the influence of abundance would disappear over time. Thus, asymmetric specialization seems to be the result of a size threshold and, only among the allowed interactions above this size threshold, a result of random interactions proportional to abundance. The simulations also showed that asymmetric specialization could not be the reason that the extinction risk of specialists and generalists is equalized, as suggested in the literature. In asymmetric webs specialists clearly had higher short-term extinction risks. In fact, primarily generalist visitors seem to profit from asymmetric specialization. In our web, specialists were less abundant than generalists. Therefore, including abundance in the simulation models increased the difference between specialists and generalists even more. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Testes asymmetry,condition and sexual selection in birds: an experimental test

The functional significance of the marked directional asymmetry in testes size observed in many bird species is obscure. Møller suggested that (i) the smaller of the two testes serves a compensatory role and increases in size (and hence reduces asymmetry) when the larger one is defective in some way, and (ii) as a consequence, the degree of directional asymmetry in testes size reflects male quality and covaries positively with the expression of secondary sexual traits. We conducted an experimental test of these two hypotheses in the zebra finch, Taeniopygia guttata. Neither hypothesis was supported. First, there was no significant relationship between the size of the left testis and relative testes asymmetry. Second, we obtained no support for the hypothesis that the degree of directional asymmetry in testes mass covaried with condition. On the contrary, directional asymmetry in testes mass was significantly greater in birds whose condition was experimentally reduced, compared with control birds. Moreover, we found no significant relationships between testes asymmetry and secondary sexual traits. We conclude that directional asymmetry in testes size does not reflect male condition in the zebra finch.     

Specialization in plant-hummingbird networks is associated with species richness, contemporary precipitation and quaternary climate-change velocity

Large-scale geographical patterns of biotic specialization and the underlying drivers are poorly understood, but it is widely believed that climate plays an important role in determining specialization. As climate-driven range dynamics should diminish local adaptations and favor generalization, one hypothesis is that contemporary biotic specialization is determined by the degree of past climatic instability, primarily Quaternary climate-change velocity. Other prominent hypotheses predict that either contemporary climate or species richness affect biotic specialization. To gain insight into geographical patterns of contemporary biotic specialization and its drivers, we use network analysis to determine the degree of specialization in plant-hummingbird mutualistic networks sampled at 31 localities, spanning a wide range of climate regimes across the Americas. We found greater biotic specialization at lower latitudes, with latitude explaining 20-22% of the spatial variation in plant-hummingbird specialization. Potential drivers of specialization--contemporary climate, Quaternary climate-change velocity, and species richness--had superior explanatory power, together explaining 53-64% of the variation in specialization. Notably, our data provides empirical evidence for the hypothesized roles of species richness, contemporary precipitation and Quaternary climate-change velocity as key predictors of biotic specialization, whereas contemporary temperature and seasonality seem unimportant in determining specialization. These results suggest that both ecological and evolutionary processes at Quaternary time scales can be important in driving large-scale geographical patterns of contemporary biotic specialization, at least for co-evolved systems such as plant-hummingbird networks.     

Can plant-pollinator network metrics indicate environmental quality?

Plant-pollinator interaction networks may be more informative than the diversity of species in the evaluation of the effects of environmental change. Considering that networks vary with the integrity of ecosystems, their changes may help to predict the consequences of anthropogenic impacts on biodiversity and ecological processes. This characteristic highlights its use as environmental quality indicator. However, to employ interaction networks as ecological indicators it is necessary to identify the most sensitive metrics and understand how and why they vary with environmental changes. This review aimed to identify, in empirical studies, which network metrics have been evidenced as being more sensitive to changes in environmental quality. We analyzed published empirical studies, that applied the network approach on environmental quality gradients. In addition to the network metric behavior, we studied the interactions between them and possible causes of their variation. The available empirical data indicated that degree, nestedness and connectance did not have a simple, linear or unidirectional response to habitat degradation. Conversely, the metrics interaction asymmetry, d' (reciprocal specialization index of the species) showed the most consistent responses to environmental change. The role of the species changed, ranging between generalists and specialists under different conditions. In addition, specialist species with morphological and behavioral constraints were lost in worse environmental quality situations. The identity of interacting species and their role in the network, with a further specification of groups and interactions most affected, are the properties with greater potential to indicate changes in environmental quality. Most of the available studies focused on metrics at the network level, but several studies and this review indicate that the patterns at the network level can be better understood in the light of metrics analyzed at the species level. Our results provide information that enrich the network analysis, highlighting the need to consider important features that are often neglected. Discussions and information compiled here are important for deciding how to look at empirical data and what to look for, as well as to indicate some caveats when interpreting data on plant-pollinator interactions with a complex network approach. Network metrics can be good indicators of environmental quality if the underlying ecological causes of the numerical changes are carefully analyzed.     

Why do some,but not all,tropical birds migrate? A comparative study of diet breadth and fruit preference

Annual migrations of birds profoundly influence terrestrial communities. However, few empirical studies examine why birds migrate, in part due to the difficulty of testing causal hypotheses in long-distance migration systems. Short-distance altitudinal migrations provide relatively tractable systems in which to test explanations for migration. Many past studies explain tropical altitudinal migration as a response to spatial and temporal variation in fruit availability. Yet this hypothesis fails to explain why some coexisting, closely-related frugivorous birds remain resident year-round. We take a mechanistic approach by proposing and evaluating two hypotheses (one based on competitive exclusion and the other based on differences in dietary specialization) to explain why some, but not all, tropical frugivores migrate. We tested predictions of these hypotheses by comparing diets, fruit preferences, and the relationships between diet and preference in closely-related pairs of migrant and resident species. Fecal samples and experimental choice trials revealed that sympatric migrants and residents differed in both their diets and fruit preferences. Migrants consumed a greater diversity of fruits and fewer arthropods than did their resident counterparts. Migrants also tended to have slightly stronger fruit preferences than residents. Most critically, diets of migrants more closely matched their preferences than did the diets of residents. These results suggest that migrants may be competitively superior foragers for fruit compared to residents (rather than vice versa), implying that current competitive interactions are unlikely to explain variation in migratory behavior among coexisting frugivores. We found some support for the dietary specialization hypothesis, propose refinements to the mechanism underlying this hypothesis, and discuss how dietary specialization might ultimately reflect past interspecific competition. We recommend that future studies quantify variation in nutritional content of tropical fruits, and determine whether frugivory is a consequence or a cause of migratory behaviour.     

Resource heterogeneity, diet shifts and intra-cohort competition: effects on size divergence in YOY fish

The pollination syndrome hypothesis has provided a major conceptual framework for how plants and pollinators interact. However, the assumption of specialization in pollination systems and the reliability of floral traits in predicting the main pollinators have been questioned recently. In addition, the relationship between ecological and evolutionary specialization in pollination interactions is still poorly understood. We used data of 62 plant species from three communities across southern Norway to test: (1) the relationships between floral traits and the identity of pollinators, (2) the association between floral traits (evolutionary specialization) and ecological generalization, and (3) the consistency of both relationships across communities. Floral traits significantly affected the identity of pollinators in the three communities in a way consistent with the predictions derived from the pollination syndrome concept. However, hover flies and butterflies visited flowers with different shapes in different communities, which we mainly attribute to among-community variation in pollinator assemblages. Interestingly, ecological generalization depended more on the community-context (i.e. the plant and pollinator assemblages in the communities) than on specific floral traits. While open yellow and white flowers were the most generalist in two communities, they were the most specialist in the alpine community. Our results warn against the use of single measures of ecological generalization to question the pollination syndrome concept, and highlight the importance of community comparisons to assess the pollination syndromes, and to understand the relationships between ecological and evolutionary specialization in plant-pollinator interactions.     

Individual-Area Relationship Best Explains Goose Species Density in Wetlands

Explaining and predicting animal distributions is one of the fundamental objectives in ecology and conservation biology. Animal habitat selection can be regulated by top-down and bottom-up processes, and is mediated by species interactions. Species varying in body size respond differently to top-down and bottom-up determinants, and hence understanding these allometric responses to those determinants is important for conservation. In this study, using two differently sized goose species wintering in the Yangtze floodplain, we tested the predictions derived from three different hypotheses (individual-area relationship, food resource and disturbance hypothesis) to explain the spatial and temporal variation in densities of two goose species. Using Generalized Linear Mixed Models with a Markov Chain Monte Carlo technique, we demonstrated that goose density was positive correlated with patch area size, suggesting that the individual area-relationship best predicts differences in goose densities. Moreover, the other predictions, related to food availability and disturbance, were not significant. Buffalo grazing probably facilitated greater white-fronted geese, as the number of buffalos was positively correlated to the density of this species. We concluded that patch area size is the most important factor determining the density of goose species in our study area. Patch area size is directly determined by water levels in the Yangtze floodplain, and hence modifying the hydrological regimes can enlarge the capacity of these wetlands for migratory birds.     

Quantitative measures of organization for multiagent systems

A set of "information theoretic" measures has been developed to quantify the degree of constraint inherent in the organization of a multiagent system. Separate measures can be provided to quantify spatial organization, trophic organization and, more generally, the overall structure of interactions. The additive character of these quantities allows them to be distributed in various fashions among species and places in a way that allows one to assign an "Importance Index" to those taxa and places. In addition, a measure to gauge the degree of adaptation of a species to a particular environment is proffered. The proposed measures allow one to formulate the following hypotheses in quantitative fashion: (1). that any disturbance of an ecosystem at a location associated with a high spatial Importance Index will exert a greater impact on the population dynamics than will a similar disturbance aimed at a place where the values of these indexes are lower; (2). that any disturbance in an ecosystem affecting a particular species with high individual Importance Indexes will cause a greater impact on the overall population dynamics than will a disturbance aimed at a species with a lower values of these indexes; (3). that the ascendancy of evolving system has a propensity to increase. The precise quantitative formulation of these hypothesis would permit them to be tested via multiagent simulation. Estimating the probablities pertaining to these hypotheses presents a number of problems that merit discussion.     

The future of plant-pollinator diversity: understanding interaction networks across time, space, and global change

Structural analysis of plant-pollinator networks has revealed remarkably high species and interaction diversity and highlighted the species important for pollination services. Although techniques to analyze plant-pollinator networks began to emerge a decade ago, the characterization of spatiotemporal variation of interactions is still in its infancy. Understanding the ecological and evolutionary causes and consequences of spatial and temporal variation in plant-pollinator interactions is important for both basic and applied questions in community structure and function, the evolution of floral traits, and the development of optimal conservation strategies. Here we review observational, theoretical, and experimental studies of temporal and spatial variation in plant-pollinator interaction networks to establish a foundation for future studies to incorporate perspectives in spatiotemporal variation. Such perspectives are crucial given the rapid environmental changes associated with habitat loss, climate change, and biological invasions, which we discuss in this context. The inherent plasticity of plant-pollinator interactions and network structure suggests that many species should be able to persist by responding to environmental changes quickly, even though the identity of their mutualistic partners may change.     

Asymmetries in specialization in ant-plant mutualistic networks

Mutualistic networks involving plants and their pollinators or frugivores have been shown recently to exhibit a particular asymmetrical organization of interactions among species called nestedness: a core of reciprocal generalists accompanied by specialist species that interact almost exclusively with generalists. This structure contrasts with compartmentalized assemblage structures that have been verified in antagonistic food webs. Here we evaluated whether nestedness is a property of another type of mutualism-the interactions between ants and extrafloral nectary-bearing plants--and whether species richness may lead to differences in degree of nestedness among biological communities. We investigated network structure in four communities in Mexico. Nested patterns in ant-plant networks were very similar to those previously reported for pollination and frugivore systems, indicating that this form of asymmetry in specialization is a common feature of mutualisms between free-living species, but not always present in species-poor systems. Other ecological factors also appeared to contribute to the nested asymmetry in specialization, because some assemblages showed more extreme asymmetry than others even when species richness was held constant. Our results support a promising approach for the development of multispecies coevolutionary theory, leading to the idea that specialization may coevolve in different but simple ways in antagonistic and mutualistic assemblages.     

Intraspecific variation in fruit–frugivore interactions: effects of fruiting neighborhood and consequences for seed dispersal

The extent of specialization/generalization continuum in fruit–frugivore interactions at the individual level remains poorly explored. Here, we investigated the interactions between the Neotropical treelet Miconia irwinii (Melastomataceae) and its avian seed dispersers in Brazilian campo rupestre. We built an individual-based network to derive plant degree of interaction specialization regarding disperser species. Then, we explored how intraspecific variation in interaction niche breadth relates to fruit availability on individual plants in varying densities of fruiting conspecific neighbors, and how these factors affect the quantity of viable seeds dispersed. We predicted broader interaction niche breadths for individuals with larger fruit crops in denser fruiting neighborhoods. The downscaled network included nine bird species and 15 plants, which varied nearly five-fold in their degree of interaction specialization. We found positive effects of crop size on visitation and fruit removal rates, but not on degree of interaction specialization. Conversely, we found that an increase in the density of conspecific fruiting neighbors both increased visitation rate and reduced plant degree of interaction specialization. We suggest that tracking fruit-rich patches by avian frugivore species is the main driver of density-dependent intraspecific variation in plants’ interaction niche breadth. Our study shed some light on the overlooked fitness consequences of intraspecific variation in interaction niches by showing that individuals along the specialization/generalization continuum may have their seed dispersed with similar effectiveness. Our study exemplifies how individual-based networks linking plants to frugivore species that differ in their seed dispersal effectiveness can advance our understanding of intraspecific variation in the outcomes of fruit–frugivore interactions.

     

Species abundance and asymmetric interaction strength in ecological networks

The strength of interactions among species in a network tends to be highly asymmetric. We evaluate the hypothesis that this asymmetry results from the distribution of abundance among species, so that species interactions occur randomly among individuals. We used a database on mutualistic and antagonistic bipartite quantitative interaction networks. We show that across all types of networks asymmetry was correlated with abundance, so that rare species were asymmetrically affected by their abundant partners, while pairs of interacting abundant species tended to exhibit more symmetric, reciprocally strong effects. A null model shows that abundance provides a sufficient explanation of the asymmetry structure in some networks, but suggests the role of additional factors in others. Although not universal, our hypothesis holds for a substantial fraction of networks analyzed here, and should be considered as a null model in all studies aimed at evaluating the ecological and evolutionary consequences of species interactions.