Mutualistic Partners Research Articles

In vitro interactions between the ectomycorrhizal Pisolithus tinctorius and the saprotroph Hypholoma fasciculare fungi: morphological aspects and volatile production

ABSTRACT Ectomycorrhizal fungi are crucial for forests sustainability. For Castanea sativa, ectomycorrhizal fungus Pisolithus tinctorius is an important mutualist partner. Saprotrophic fungi Hypholoma fasciculare, although used for biocontrol of Armillaria root disease, it negatively affected the interaction between the P. tinctorius and plant host roots, by compromise the formation of P. tinctorius-C. sativa mycorrhizae. In this work, fungal morphology during inhibition of H. fasciculare against P. tinctorius was elucidated. P. tinctorius growth was strongly affected by H. fasciculare, which was significantly reduced after six days of co-culture and become even more significant through time. During this period, P. tinctorius developed vesicles and calcium oxalate crystals, which were described as mechanisms to stress adaption by fungi. H. fasciculare produced different volatile organic compounds in co-cultures over time and differ between single or in dual-species. H. fasciculare highly produced sesquiterpenes (namely, α-muurolene) and nitrogen-containing compounds, which are recognised as having antimicrobial activity.

Positive interactions within and between populations decrease the likelihood of evolutionary rescue.

Positive interactions, including intraspecies cooperation and interspecies mutualisms, play crucial roles in shaping the structure and function of many ecosystems, ranging from plant communities to the human microbiome. While the evolutionary forces that form and maintain positive interactions have been investigated extensively, the influence of positive interactions on the ability of species to adapt to new environments is still poorly understood. Here, we use numerical simulations and theoretical analyses to study how positive interactions impact the likelihood that populations survive after an environment deteriorates, such that survival in the new environment requires quick adaptation via the rise of new mutants-a scenario known as evolutionary rescue. We find that the probability of evolutionary rescue in populations engaged in positive interactions is reduced significantly. In cooperating populations, this reduction is largely due to the fact that survival may require at least a minimal number of individuals, meaning that adapted mutants must arise and spread before the population declines below this threshold. In mutualistic populations, the rescue probability is decreased further due to two additional effects-the need for both mutualistic partners to adapt to the new environment, and competition between the two species. Finally, we show that the presence of cheaters reduces the likelihood of evolutionary rescue even further, making it extremely unlikely. These results indicate that while positive interactions may be beneficial in stable environments, they can hinder adaptation to changing environments and thereby elevate the risk of population collapse. Furthermore, these results may hint at the selective pressures that drove co-dependent unicellular species to form more adaptable organisms able to differentiate into multiple phenotypes, including multicellular life.

Chronic, low-level nitrogen deposition enhances abundances of ant-protected herbivores inhabiting an imperiled foundation species

Atmospheric nitrogen (N) deposition is driving changes in plant assemblages worldwide, but animal responses are not well understood, especially in relation to current widespread rates of low-level N deposition. Big sagebrush (Artemisia tridentata) is a foundation species in steppe ecosystems of western North America, where elevated, but low-level N deposition is prevalent. Sagebrush hosts numerous herbivorous and predatory arthropods, some of which form mutualistic (i.e. mutually beneficial) herbivore-predator associations that may be harmful for sagebrush. Here, I evaluated the response of a sagebrush arthropod assemblage to four and nine years of low-level N enrichment in Colorado, USA. Though sagebrush cover and arthropod abundances were not different in the fourth year, by the ninth year, N additions increased arthropod abundances due to greater numbers of sap-feeding insects that are mutualist partners of predaceous ants. Ant and non-mutualist herbivore abundances did not significantly respond to N treatments in the ninth year, but non-mutualist predators were only found in ambient N plots, suggesting ants provided greater anti-predator protection for mutualist herbivores under N enrichment. These responses occurred during drought conditions, which may have increased sagebrush nutritional quality for sap-feeding insects. Sagebrush is experiencing multiple stressors that are transforming steppe ecosystems; these results suggest chronic exposure to common rates of low-level N deposition may create further stress as herbivorous insects proliferate. Additional work is needed to understand and manage the repercussions of N deposition for plant-animal assemblages, particularly in contexts including species of conservation concern and both mutualistic and antagonistic species interactions.

Limitation by a shared mutualist promotes coexistence of multiple competing partners

Although mutualisms are often studied as simple pairwise interactions, they typically involve complex networks of interacting species. How multiple mutualistic partners that provide the same service and compete for resources are maintained in mutualistic networks is an open question. We use a model bacterial community in which multiple ‘partner strains’ of Escherichia coli compete for a carbon source and exchange resources with a ‘shared mutualist’ strain of Salmonella enterica. In laboratory experiments, competing E. coli strains readily coexist in the presence of S. enterica, despite differences in their competitive abilities. We use ecological modeling to demonstrate that a shared mutualist can create temporary resource niche partitioning by limiting growth rates, even if yield is set by a resource external to a mutualism. This mechanism can extend to maintain multiple competing partner species. Our results improve our understanding of complex mutualistic communities and aid efforts to design stable microbial communities.

EFFECT OF CLIMATE CHANGE ON BEE FARMING. A CRITICAL REVIEW OF LITERATURE

Purpose: Climate warming affects the phenology, local abundance and large‐scale distribution of bees. Despite this, there is still limited knowledge of how climate affect plant‐pollinator mutualisms and how changed availability of mutualistic partners influences the persistence of interacting species. This article reviews the evidence of climate warming effects on bee farming and discuss how their interactions may be affected by change in climate. Bees provide the majority of biotic pollination and are at risk from a multitude of factors; changes in land use, intensive agricultural practices, mono-cropping (growing a single crop year after year on the same land), and the use of pesticides have all contributed to large-scale losses, fragmentation and degradation of bee habitat. The general objective of the study was to establish the effect of effect of climate change on bee farming. 
 Methodology: The paper used a desk study review methodology where relevant empirical literature was reviewed to identify main themes and to extract knowledge gaps.
 Findings: The study found out Climate change is causing temperature shifts which are leaving bees unable to pollinate in time. Bees are severely vulnerable to extreme weather and climate change has caused flowers to emerge and bloom earlier. Changing temperatures have also reduced the size of their wild range by approximately five miles.
 Recommendations: The study recommends that the local community needs to be enlightened on the need to form self-help group. These will provide them a platform to access more incentives and be able to share more information in relation to honey yield and to put more emphasis on providing food and water to bees during dry season

Coevolved mutualists experience fluctuating costs and benefits over time

Understanding how mutualisms persist over time requires investigations of how mutualist species coevolve and adapt to the interaction. In particular, the key factors in the evolution of mutualisms are the costs and benefits mutualists experience during the interaction. Here, we used a yeast nutritional mutualism to test how mutualists coevolve and adapt in an obligate mutualism. We allowed two yeast mutualists to evolve together for 15 weeks (about 150 generations), and then we tested if the mutualists had coevolved using time-shift assays. We also examined two mutualistic traits associated with the costs and benefits: resource use efficiency and commodity production. We found that the mutualists quickly coevolved. Furthermore, the changes in benefits and costs were nonlinear and varied with evolutionary changes occurring in the mutualist partner. One mutualist initially evolved to reduce mutualistic commodity production and increase efficiency in mutualistic resource use; however, this negatively affected its mutualist partner that evolved reduced commodity production and resource use efficiency. As a result, the former increased commodity production, resulting in an increase in benefits for its partner. The quick, nonlinear, and asynchronous evolution of yeast mutualists closely resembles antagonistic coevolutionary patterns, supporting the view that mutualisms should be considered as reciprocal exploitation.

Metabolic Contributions of an Alphaproteobacterial Endosymbiont in the Apicomplexan Cardiosporidium cionae.

Apicomplexa is a diverse protistan phylum composed almost exclusively of metazoan-infecting parasites, including the causative agents of malaria, cryptosporidiosis, and toxoplasmosis. A single apicomplexan genus, Nephromyces, was described in 2010 as a mutualist partner to its tunicate host. Here we present genomic and transcriptomic data from the parasitic sister species to this mutualist, Cardiosporidium cionae, and its associated bacterial endosymbiont. Cardiosporidium cionae and Nephromyces both infect tunicate hosts, localize to similar organs within these hosts, and maintain bacterial endosymbionts. Though many other protists are known to harbor bacterial endosymbionts, these associations are completely unknown in Apicomplexa outside of the Nephromycidae clade. Our data indicate that a vertically transmitted α-proteobacteria has been retained in each lineage since Nephromyces and Cardiosporidium diverged. This α-proteobacterial endosymbiont has highly reduced metabolic capabilities, but contributes the essential amino acid lysine, and essential cofactor lipoic acid to C. cionae. This partnership likely reduces resource competition with the tunicate host. However, our data indicate that the contribution of the single α-proteobacterial endosymbiont in C. cionae is minimal compared to the three taxa of endosymbionts present in the Nephromyces system, and is a potential explanation for the virulence disparity between these lineages.

Stage-Specific Parasitism by a Mutualistic Partner Can Increase the Host Abundance

Mutualism is one of the major interaction types occurring in ecological communities, and has significant consequences for population dynamics and community structure. A major feature of mutualism is its context dependence; i.e., potentially mutualistic relationships may become parasitic or neutral depending on the abiotic conditions and presence of other species. Recently, stage structure has been recognized as an additional important factor underlying the context-dependence of mutualism, giving rise to the concept of stage-structured mutualism, which states that mutualistic partners at one life-history stage may become parasitic at other life-history stages. However, the ecological consequences of such a stage-structured interaction remain unknown. Here, we present a simplified one host–one partner model with a juvenile–adult stage structure of the host species, in which the partner species improves fecundity of the adult host but suppresses growth and maturation of the juvenile host. Numerical simulations identified two ecological conditions for stage-specific parasitism to counterintuitively increase host abundance: (i) the benefits derived by the adult host from the partner species are large and the costs imposed on the juvenile host by the partner species are small (the partner investment hypothesis), and (ii) host population growth is limited by adult reproduction instead of juvenile maturation (the ontogenetic asymmetry hypothesis). Although the partner investment hypothesis appears more intuitive, exhaustive parameter exploration revealed that the ontogenetic asymmetric hypothesis would be more likely to operate. This counterintuitive result implies that the dynamics of mutualistic interactions are more complex than previously thought. The present study provides novel insights into the ecological consequences of mutualism and illustrates the importance of the ontogenetic perspective of mutualism in community ecology. Further research is required in both empirical and theoretical approaches to better understand the mechanisms by which stage-structured mutualism mediates population dynamics and community structure in more complex systems.

The Evolution of Mutualistic Dependence

While the importance of mutualisms across the tree of life is recognized, it is not understood why some organisms evolve high levels of dependence on mutualistic partnerships, while other species remain autonomous or retain or regain minimal dependence on partners. We identify four main pathways leading to the evolution of mutualistic dependence. Then, we evaluate current evidence for three predictions: ( a) Mutualisms with different levels of dependence have distinct stabilizing mechanisms against exploitation and cheating, ( b) less dependent mutualists will return to autonomy more often than those that are highly dependent, and ( c) obligate mutualisms should be less context dependent than facultative ones. Although we find evidence supporting all three predictions, we stress that mutualistic partners follow diverse paths toward—and away from—dependence. We also highlight the need to better examine asymmetry in partner dependence. Recognizing how variation in dependence influences the stability, breakdown, and context dependence of mutualisms generates new hypotheses regarding how and why the benefits of mutualistic partnerships differ over time and space.

Evolution: The Legacy of Endosymbiosis in Ants

A symbiotic partnership with Blochmannia bacteria is thought to underpin the ecological success of carpenter ants. Disentangling the molecular interactions between the mutualistic partners supports an old hypothesis that many other ants also had similar symbioses and lost them.

Clade-Specific Sterol Metabolites in Dinoflagellate Endosymbionts Are Associated with Coral Bleaching in Response to Environmental Cues.

Cnidarians cannot synthesize sterols (which play essential roles in growth and development) de novo but often use sterols acquired from endosymbiotic dinoflagellates. While sterol availability can impact the mutualistic interaction between coral host and algal symbiont, the biosynthetic pathways (in the dinoflagellate endosymbionts) and functional roles of sterols in these symbioses are poorly understood. In this study, we found that itraconazole, which perturbs sterol metabolism by inhibiting the sterol 14-demethylase CYP51 in dinoflagellates, induces bleaching of the anemone Heteractis crispa and that bleaching perturbs sterol metabolism of the dinoflagellate. While Symbiodiniaceae have clade-specific sterol metabolites, they share features of the common sterol biosynthetic pathway but with distinct architecture and substrate specificity features of participating enzymes. Tracking sterol profiles and transcripts of enzymes involved in sterol biosynthesis across time in response to different environmental cues revealed similarities and idiosyncratic features of sterol synthesis in the endosymbiont Breviolum minutum Exposure of algal cultures to high levels of light, heat, and acidification led to alterations in sterol synthesis, including blocks through downregulation of squalene synthase transcript levels accompanied by marked growth reductions.IMPORTANCE These results indicate that sterol metabolites in Symbiodiniaceae are clade specific, that their biosynthetic pathways share architectural and substrate specificity features with those of animals and plants, and that environmental stress-specific perturbation of sterol biosynthesis in dinoflagellates can impair a key mutualistic partnership for healthy reefs.

Chemical convergence between a guild of facultative myrmecophilous caterpillars and host plants

Ants exert strong selective pressure on herbivorous insects, although some caterpillars can live in symbiosis with them using chemical defensive strategies. We investigated the adaptive resemblance of cuticular hydrocarbons (CHCs) in multitrophic systems involving a guild of facultative myrmecophilous caterpillar species (Lepidoptera: Lycaenidae), tending ants (Hymenoptera: Formicidae), and host plants from three families. We hypothesised that the CHCs of the caterpillars would resemble those of their host plants (chemical camouflage). We analysed CHCs using gas chromatography/mass spectrometry. Morisita's similarity index (SI) was used to compare CHC profiles of caterpillar species with different types of ant associations (commensal or mutualistic), ants, and host plants. We found strong convergence between caterpillars' CHCs and plants, especially for commensal species that do not provide secretion rewards for ants. Moreover, we found unexpected chemical convergence among mutualistic (trophobiotic) caterpillar species that offer caloric reward secretions to ants. These results show that the studied caterpillars acquire CHCs through their diet and that they vary according to host plant species and type of ant association (commensalism or mutualism). This ‘chemical camouflage’ of myrmecophilous caterpillars may have arisen as a defensive strategy allowing coexistence with ants on plants, whereas ‘chemical conspicuousness’ may have evolved in the context of honest signalling between mutualistic partners. We suggest the existence of chemical mimicry among myrmecophilous species, especially between mutualistic caterpillars. Cuticular chemical mixtures can play a key adaptive role in decreasing ant attacks and increasing caterpillar survival in multimodal sensory systems.

The entomophagous caterpillar fungus Ophiocordyceps sinensis is consumed by its lepidopteran host as a plant endophyte

Abstract Endophytic insect pathogenic fungi of the order Hypocreales reside in plant tissues as mutualistic partners that protect plants against insect herbivores. Caterpillar fungus Ophiocordyceps sinensis (order: Hypocreales, family: Ophiocordycipitaceae) is often described as an entomophagous fungus that parasitizes lepidopteran larvae of the ghost moth genus Thitarodes (family: Hepialidae), but recent evidence suggests involvement of a plant-endophytic stage in O. sinensis infection of insect larvae. In this study, we screen sample plant species in a caterpillar fungus collection site at Mt. Gongga, Sichuan, China, for endophytic O. sinensis and analyze the diet of Thitarodes larvae living in the soil of the same habitat. We show that the entomophagous caterpillar fungus O. sinensis is a prevalent plant endophyte (present in 52.6% of all plant genera and 66.7% of plant families) and that the presence of endophytic O. sinensis in plant leaf tissue is significantly correlated with its presence in plant root tissue. Our analysis of larval diet content reveals a high diversity generalist diet and confirms that the plant families harboring endophytic O. sinensis are consumed by Thitarodes larvae, although the composition and diversity of an individual's diet do not predict its level of fungal infection. These results suggest a reproductive strategy of O. sinensis involving a plant endophytic stage that facilitates infection of root-eating host larvae. Our study highlights the role of resident plant communities as a third participating partner in this complex symbiosis.

Urbanization drives an early spring for plants but not for pollinators

Urbanization is one of the major threats to wild plants and pollinators, and its global increase demands a better understanding of the mechanism driving its negative impact. Urban warming and altered local environmental conditions have the potential to affect the timing of flowering and of pollinator activity. While previous evidence has shown that plant phenology tends to advance in urban areas, little is known about its effects on pollinator phenology. In this study we simultaneously assessed the response of the timing of flowering of native plants and of the flight period of wild pollinators to increased urbanization. We collected data from 12 sites along an urbanization gradient in northern France, a region under strong anthropogenic pressure. Overall, we recorded more than 70 plant species, and we sampled more than 4300 wild bees and hoverflies belonging to 154 species. Plant flowering showed a strong response to urbanization at the community level with a striking advancement of the flowering peak in sites at high urbanization. On the contrary, pollinator communities did not show any clear shift of their flight phenology along the gradient, neither regarding abundance nor diversity. Our results indicate that phenologies of plant and pollinator communities can respond differently along the same urbanization gradient. These asymmetric responses can drive modifications in the structure of plant–pollinator networks, and potentially negatively affect the fitness of both mutualistic partners.

Parasitism within mutualist guilds explains the maintenance of diversity in multi-species mutualisms

We consider here mutualisms where there are multiple species sharing a resource supplied by the same partner. If, as commonly assumed, there is competition between the species, then only the superior competitor should persist. Nevertheless, coexistence of multiple species sharing the same mutualistic partner is a widespread phenomenon. Regulation of nutrient exchange, where each species receives resources from the partner in proportion to the strength of the mutualism between the two, has been proposed as the main mechanism for coexistence in multi-species mutualisms involving the transfer of nutrients. Significant arguments, however, challenge the importance of partner selection processes. We present a mathematical model, applied to the arbuscular mycorrhizal symbiosis, to propose an alternative explanation for this coexistence. We show that asymmetric resource exchange between the plant and its fungal guild can lead to indirect parasitic interactions between guild members. In our model, the amount of carbon supplied by the plant to the fungi depends on both plant and fungal biomass, while the amount of phosphorus supplied by the fungi to the plant depends on both plant and fungal biomass when the plant is small, and effectively on fungal biomass only when the plant is large. As a consequence of these functional responses, more beneficial mutualists increase resource availability, and are indirectly exploited by less beneficial species that consume the resource and grow larger than they would in the absence of the better mutualists. As guild mutualists are not competing, competitive exclusion does not occur. Hence, the interaction structure can explain the maintenance of diversity within guilds in the absence of spatial structure and niche-related processes.

Intermingling of gut microbiota with brain: Exploring the role of probiotics in battle against depressive disorders

Depression is a debilitating psychiatric ailment which exerts disastrous effects on one’s mental and physical health. Depression is accountable for augmentation of various life-threatening maladies such as neurodegenerative anomalies, cardiovascular diseases and diabetes. Depressive episodes are recurrent, pose a negative impact on life quality, decline life expectancy and enhance suicidal tendencies. Anti-depression chemotherapy displays marked adverse effects and frequent relapses. Thus, newer therapeutic interventions to prevent or combat depression are desperately required. Discovery of gut microbes as our mutualistic partner was made a long time ago and it is surprising that their functions still continue to expand and as of yet many are still to be uncovered. Experimental studies have revealed astonishing role of gut commensals in gut-brain signaling, immune homeostasis and hormonal regulation. Now, it is a well-established fact that gut microbes can alleviate stress or depression associated symptoms by modulating brain functions. Here in, we provide an overview of physiological alleyways involved in cross-talk between gut and brain, part played by probiotics in regulation of these pathways and use of probiotic bacteria as psychobiotics in various mental or depressive disorders.

Covert Cross-Feeding Revealed by Genome-Wide Analysis of Fitness Determinants in a Synthetic Bacterial Mutualism.

Microbial interactions abound in natural ecosystems and shape community structure and function. Substantial attention has been given to cataloging mechanisms by which microbes interact, but there is a limited understanding of the genetic landscapes that promote or hinder microbial interactions. We previously developed a mutualistic coculture pairing Escherichia coli and Rhodopseudomonas palustris, wherein E. coli provides carbon to R. palustris in the form of glucose fermentation products and R. palustris fixes N2 gas and provides nitrogen to E. coli in the form of NH4+ The stable coexistence and reproducible trends exhibited by this coculture make it ideal for interrogating the genetic underpinnings of a cross-feeding mutualism. Here, we used random barcode transposon sequencing (RB-TnSeq) to conduct a genome-wide search for E. coli genes that influence fitness during cooperative growth with R. palustris RB-TnSeq revealed hundreds of genes that increased or decreased E. coli fitness in a mutualism-dependent manner. Some identified genes were involved in nitrogen sensing and assimilation, as expected given the coculture design. The other identified genes were involved in diverse cellular processes, including energy production and cell wall and membrane biogenesis. In addition, we discovered unexpected purine cross-feeding from R. palustris to E. coli, with coculture rescuing growth of an E. coli purine auxotroph. Our data provide insight into the genes and gene networks that can influence a cross-feeding mutualism and underscore that microbial interactions are not necessarily predictable a prioriIMPORTANCE Microbial communities impact life on Earth in profound ways, including driving global nutrient cycles and influencing human health and disease. These community functions depend on the interactions that resident microbes have with the environment and each other. Thus, identifying genes that influence these interactions will aid the management of natural communities and the use of microbial consortia as biotechnology. Here, we identified genes that influenced Escherichia coli fitness during cooperative growth with a mutualistic partner, Rhodopseudomonas palustris Although this mutualism centers on the bidirectional exchange of essential carbon and nitrogen, E. coli fitness was positively and negatively affected by genes involved in diverse cellular processes. Furthermore, we discovered an unexpected purine cross-feeding interaction. These results contribute knowledge on the genetic foundation of a microbial cross-feeding interaction and highlight that unanticipated interactions can occur even within engineered microbial communities.

Interactions among interactions: The dynamical consequences of antagonism between mutualists

Species often interact with multiple mutualistic partners that provide functionally different benefits and/or that interact with different life-history stages. These functionally different partners, however, may also interact directly with one another in other ways, indirectly altering net outcomes and persistence of the mutualistic system as a whole. We present a population dynamical model of a three-species system involving antagonism between species sharing a mutualist partner species with two explicit life stages. We find that, regardless of whether the antagonism is predatory or non-consumptive, persistence of the shared mutualist is possible only under a restrictive set of conditions. As the rate of antagonism between the species sharing the mutualist increases, indirect rather than direct interactions increasingly determine species’ densities and sometimes result in complex, oscillatory dynamics for all species. Surprisingly, persistence of the mutualistic system is particularly dependent upon the degree to which each of the two mutualistic interactions is specialized. Our work investigates a novel mechanism by which changing ecological conditions can lead to extinction of mutualist partners and provides testable predictions regarding the interactive roles of mutualism and antagonism in net outcomes for species’ densities.

Land-Use and Climate Impacts on Plant–Pollinator Interactions and Pollination Services

Most flowering plants rely on animals for pollination and most animal pollinators rely on flowering plants for food resources. However, there is an ongoing concern that anthropogenic-induced global change threatens the mutualistic association between plants and pollinators. Two of the most important factors of global change are land-use and climate change. Land-use and climate change may affect species distributions and species phenologies, leading to spatial and temporal mismatches between mutualistic partners. Land-use and climate change may also influence species abundances, nesting habitats, floral resources and the behaviors of pollinators. Thus, mutualistic plant–pollinator interactions should be more susceptible to global change than simple measures of biodiversity, such as species richness and species composition. The potential negative impacts of land-use and climate change on plant–pollinator interactions may have large consequences for the conservation of threatened plants and pollinators and economically by diminishing crop productivity. Here I highlight ‘fruitful avenues’ for research into better understanding the influence of land-use and climate change on plant–pollinator interactions.

Dispersal patterns of large-seeded plants and the foraging behaviour of a frugivorous bat

AbstractMutualistic interactions are biologically important, diverse and poorly understood. Comprehending these interactions and the effectiveness of the mutualistic partners has been the central focus of ecological and evolutionary studies, as this task requires disentangling the pieces of mutualism under study. Here, we tested the hypothesis that feeding activity of Artibeus phaeotis influences density, diversity and distance effects of the seed rain of large-seeded plant species in a Neotropical dry forest of Costa Rica. We predicted that bats’ activity increases density and diversity metrics on dispersing sites and that bats behave as short-distance dispersers. Our data revealed that, by dispersing hundreds of seeds within their small foraging areas and mostly close to fruiting trees, the feeding activity of A. phaeotis increases the density and richness of seeds underneath night feeding roosts. The functional role of A. phaeotis as disperser may vary among plant species, as we also detected that some seeds are more dispersed than others. Further inquiries to elucidate mutualistic interactions between frugivorous bats and large-seeded plant species should focus on understanding demographic and fitness outcomes, as strong selective pressures may be shaping morphological and behavioural traits of both plants and animals, ultimately influencing the survival of each partner.