Plants with extrafloral nectaries share indirect defenses and shape the local arboreal ant community.

Associational resistance (AR) is a positive interaction in which a plant suffers less herbivore damage due to its association with a protective plant. Here, we evaluated whether plants with extra-floral nectaries (EFNs) can share indirect defenses with neighboring plants. We sampled 45 individuals of an EFN-bearing liana (Smilax polyantha) and recorded whether their support species had EFNs. In S. polyantha, we measured foliar herbivory and flower and fruit production. We examined the ant species composition and visitation of S. polyantha and whether they changed according to the supporting plant type (with or without EFNs). We experimentally determined whether S. polyantha supplemented with artificial nectaries could share indirect defenses with defenseless neighboring plants. Support plants with EFNs indirectly benefited S. polyantha by sharing mutualistic ant species. Smilax polyantha supported by plants with EFNs had a more specific ant species composition, a higher number of visiting ants and ant species richness, and exhibited nearly 3 times less foliar herbivory. However, we did not observe differences in fruit production between the two groups of S. polyantha. Finally, we observed that S. polyantha with artificial nectaries increased ant visitation on neighboring plants 2.5 times. We provide evidence that interspecific neighbors with EFNs can experience reciprocal benefits by sharing indirect defenses. Such local effects might escalate and affect the structure of plant communities.

The effects of ants on pest control: a meta-analysis.

Environmental impacts of conventional agriculture have generated interest in sustainable agriculture. Biological pest control is a fundamental tool, and ants are key players providing ecological services, as well as some disservices. We have used a meta-analytical approach to investigate the contribution of ants to biological control, considering their effects on pest and natural enemy abundance, plant damage and crop yield. We also evaluated whether the effects of ants are modulated by traits of ants, pests and other natural enemies, as well as by field size, crop system and experiment duration. Overall (considering all meta-analyses), from 52 studies on 17 different crops, we found that ants decrease the abundance of non-honeydew-producing pests, decrease plant damage and increase crop yield (services). In addition, ants decrease the abundance of natural enemies, mainly the generalist ones, and increase honeydew-producing pest abundance (disservices). We show that the pest control and plant protection provided by ants are boosted in shaded crops compared to monocultures. Furthermore, ants increase crop yield in shaded crops, and this effect increases with time. Finally, we bring new insights such as the importance of shaded crops to ant services, providing a good tool for farmers and stakeholders considering sustainable farming practices.

Haemosporidian parasites and incubation period influence plumage coloration in tanagers (Passeriformes: Thraupidae).

Birds are highly visually oriented and use plumage coloration as an important signalling trait in social communication. Hence, males and females may have different patterns of plumage coloration, a phenomenon known as sexual dichromatism. Because males tend to have more complex plumages, sexual dichromatism is usually attributed to female choice. However, plumage coloration is partly condition-dependent; therefore, other selective pressures affecting individuals' success may also drive the evolution of this trait. Here, we used tanagers as model organisms to study the relationships between dichromatism and plumage coloration complexity in tanagers with parasitism by haemosporidians, investment in reproduction and life-history traits. We screened blood samples from 2849 individual birds belonging to 52 tanager species to detect haemosporidian parasites. We used publicly available data for plumage coloration, bird phylogeny and life-history traits to run phylogenetic generalized least-square models of plumage dichromatism and complexity in male and female tanagers. We found that plumage dichromatism was more pronounced in bird species with a higher prevalence of haemosporidian parasites. Lastly, high plumage coloration complexity in female tanagers was associated with a longer incubation period. Our results indicate an association between haemosporidian parasites and plumage coloration suggesting that parasites impact mechanisms of sexual selection, increasing differences between the sexes, and social (non-sexual) selection, driving females to develop more complex coloration.

Host life-history traits predict haemosporidian parasite prevalence in tanagers (Aves: Thraupidae).

Vector-borne parasites are important ecological drivers influencing life-history evolution in birds by increasing host mortality or susceptibility to new diseases. Therefore, understanding why vulnerability to infection varies within a host clade is a crucial task for conservation biology and for understanding macroecological life-history patterns. Here, we studied the relationship of avian life-history traits and climate on the prevalence of Plasmodium and Parahaemoproteus parasites. We sampled 3569 individual birds belonging to 53 species of the family Thraupidae. Individuals were captured from 2007 to 2018 at 92 locations. We created 2 phylogenetic generalized least-squares models with Plasmodium and Parahaemoproteus prevalence as our response variables, and with the following predictor variables: climate PC1, climate PC2, body size, mixed-species flock participation, incubation period, migration, nest height, foraging height, forest cover, and diet. We found that Parahaemoproteus and Plasmodium prevalence was higher in species inhabiting open habitats. Tanager species with longer incubation periods had higher Parahaemoproteus prevalence as well, and we hypothesize that these longer incubation periods overlap with maximum vector abundances, resulting in a higher probability of infection among adult hosts during their incubation period and among chicks. Lastly, we found that Plasmodium prevalence was higher in species without migratory behaviour, with mixed-species flock participation, and with an omnivorous or animal-derived diet. We discuss the consequences of higher infection prevalence in relation to life-history traits in tanagers.

Spatiotemporal niche-based mechanisms support a stable coexistence of ants and spiders in an extrafloral nectary-bearing plant community.

Mechanisms promoting stable coexistence allow multiple species to persist in the same trophic level of a given network of species interactions. One of the most common stabilizing mechanisms of coexistence is niche differentiation, such as temporal and spatial patchiness. To understand the limits of coexistence between species we have to understand the limits of competitive interactions which translate in species exclusion or patterns of non-co-occurrence. We evaluated spatiotemporal niche-based mechanisms that could promote stable coexistence between ants and spiders which forage on extrafloral nectary (EFN)-bearing plants. We observed co-occurrence and overlapping patterns between ants and spiders in a temporal and spatial scale in nine different EFN-bearing plant species in a Neotropical savanna, using both community and species-level approach. Ants and spiders showed asynchrony of their abundances over the year with low temporal overlapping patterns between them (temporal niche specialization). Greater abundance of ants occurred between September and March, whereas greater abundance of spiders occurred between March and August, exactly at the time when the abundance of ants decreases on plants. However, there might also be some levels of temporal overlapping, but then individual ants and spiders occupy different branches (spatial segregation). Finally, we also observed a spatial negative effect of the abundance of ants on the presence of spiders. Our results suggest that spatiotemporal partitioning between ants and spiders may be one of the potential mechanisms behind a stable coexistence between these two groups of organisms that forage on EFN-bearing plants in the Brazilian savanna.

The study of biotic interactions in the Brazilian Cerrado as a path to the conservation of biodiversity.

As a focus for conservation efforts, biodiversity has received increased attention in the last fifty years. Searching for patterns in biodiversity, researchers have suggested studies including: ecological communities, cladistics classifications, hierarchical compositions of different levels of organization, and groups of taxonomically related species. Here, we propose that the study of the biodiversity of interactions may present a new perspective in the efforts to conserve biodiversity, especially in endangered ecosystems like the tropical savannas. We suggest that Cerrado, like other tropical savannas, is a particularly important ecosystem in which we can direct efforts to explain what determines the major part of variation in the outcomes of species interactions.

Mimicry and eavesdropping enable a new form of social parasitism in ants.

Social parasitism is defined by the exploitation of the social mechanisms of one society by another whole society. Here, we use quantitative ecological data and experiments to identify the components of a new form of social parasitism by the recently discovered "mirror turtle ant," Cephalotes specularis. We show that C. specularis workers visually mimic and actively avoid contact with foragers of the hyperaggressive host ant Crematogaster ampla, allowing them to move freely in the extensive and otherwise defended foraging networks of host colonies. Workers from parasite colonies have immediate access to these networks by nesting exclusively within host territories, and 89% of all potential host territories were parasitized. Inside the network, parasite workers eavesdrop on the host's trail pheromones to locate and exploit food resources that are defended by the host to the exclusion of all other ants. Experiments demonstrated the unprecedented capacity of the parasite for superior foraging performance on its host's pheromone trails than on trails of its own. Considered together, the apparent Batesian-Wallacian mimicry, pheromone-based interceptive eavesdropping, kleptoparasitism, and xenobiotic nesting ecology displayed by C. specularis within the territory and foraging network of a host ant represents a novel adaptive syndrome for social exploitation.

The role of tending ants in host plant selection and egg parasitism of two facultative myrmecophilous butterflies.

Ovipositing adult females of myrmecophilous lycaenids are expected to select plants based on ant presence in order to maximize the survivorship of immature stages. Usually, larvae feed ants with honey-like solutions and, in turn, ants ward off parasitoids. Nonetheless, a rarely investigated approach is whether ant partners can also extend their protective behavior towards lycaenids eggs. Here, we investigated the ant-related oviposition pattern of Allosmaitia strophius and Rekoa marius; then, we compared egg parasitism according to the presence of ants. Lycaenid oviposition and egg parasitism (in percent) were experimentally compared in ant-present and ant-excluded treatments. The study plant, Heteropterys byrsonimifolia, is an extrafloral nectaried shrub which supports several ant species. We sampled 280 eggs, of which 39.65 % belonged to A. strophius and 60.35 % to R. marius. Both lycaenids eggs were significantly more abundant on branches with ants, especially those with Camponotus crassus and Camponotus blandus, two ant species known to attend to lycaenids. A. strophius and R. marius parasitism was 4.5- and 2.4-fold higher, respectively, in ant-present treatments, but the results were not statistically significant. Our study shows that ant-mediated host plant selection in lycaenids might be much more widespread than previously thought, and not restricted to obligate myrmecophilous species. Tending ants may be inefficient bodyguards of lycaenid eggs, because unlike larvae which release sugared liquids, eggs do not offer obvious rewards to ants. Ants can ward off parasitoids of larvae, as observed elsewhere, but our findings show that positive ant-lycaenid interactions are conditional and depend on immature ontogeny.

Description of Cephalotes specularis n. sp. (Formicidae: Myrmicinae)-the mirror turtle ant.

We describe here Cephalotes specularis n. sp. (Hymenoptera: Formicidae: Myrmicinae: Cephalotini) based on minor and major workers, gynes and larvae from Uberlândia, Minas Gerais state, Brazil. On morphological and molecular grounds, this new species belongs to the C. fiebrigi + C. bruchi species complex, of which there are 11 previously described species (one in C. bruchi group and 10 in the C. fiebrigi group). All members of these groups are found in, or are limited to the South American "arid diagonal", comprised of the Argentinian Chaco, the Cerrados of central South America, and the Brazilian northeastern caatingas. Workers of C. specularis n. sp. have an extremely shiny gaster which is mirror-like, notwithstanding its sparse covering by minute hairs. This species engages in a form of resource-based social parasitism of the host ant Crematogaster ampla (Myrmicinae: Crematogastrini). Cephalotes specularis foragers move freely in the dense traffic of Crematogaster ampla foraging trails. They exhibit highly atypical body posturing for turtle ants, which makes them hard to distinguish from the Crematogaster foragers.

Emerging Trends in Ant-Pollinator Conflict in Extrafloral Nectary-Bearing Plants.

The net outcomes of mutualisms are mediated by the trade-offs between the costs and benefits provided by both partners. Our review proposes the existence of a trade-off in ant protection mutualisms between the benefits generated by the ants' protection against the attack of herbivores and the losses caused by the disruption of pollination processes, which are commonly not quantified. This trade-off has important implications for understanding the evolution of extrafloral nectaries (EFNs), an adaptation that has repeatedly evolved throughout the flowering plant clade. We propose that the outcome of this trade-off is contingent on the specific traits of the organisms involved. We provide evidence that the protective mutualisms between ants and plants mediated by EFNs have optimal protective ant partners, represented by the optimum point of the balance between positive effects on plant protection and negative effects on pollination process. Our review also provides important details about a potential synergism of EFN functionality; that is, these structures can attract ants to protect against herbivores and/or distract them from flowers so as not to disrupt pollination processes. Finally, we argue that generalizations regarding how ants impact plants should be made with caution since ants' effects on plants vary with the identity of the ant species in their overall net outcome.

Effect of post-fire resprouting on leaf fluctuating asymmetry, extrafloral nectar quality, and ant-plant-herbivore interactions.

Fires in the Cerrado savanna are a severe form of disturbance, but some species are capable of resprouting afterwards. It is unknown, however, how and whether post-fire resprouting represents a stressful condition to plants and how their rapid re-growth influences both the production of biochemical compounds, and interactions with mutualistic ants. In this study, we examined the influence of post-fire resprouting on biotic interactions (ant-plant-herbivore relationships) and on plant stress. The study was performed on two groups of the extrafloral nectaried shrub Banisteriopsis campestris (Malpighiaceae); one group was recovering from fire while the other acted as control. With respect to biotic interactions, we examined whether resprouting influenced extrafloral nectar concentration (milligrams per microliter), the abundance of the ant Camponotus crassus and leaf herbivory rates. Plant stress was assessed via fluctuating asymmetry (FA) analysis, which refers to deviations from perfect symmetry in bilaterally symmetrical traits (e.g., leaves) and indicates whether species are under stress. Results revealed that FA, sugar concentration, and ant abundance were 51.7 %, 35.7 % and 21.7 % higher in resprouting plants. Furthermore, C. crassus was significantly associated with low herbivory rates, but only in resprouting plants. This study showed that post-fire resprouting induced high levels of plant stress and influenced extrafloral nectar quality and ant-herbivore relationships in B. campestris. Therefore, despite being a stressful condition to the plant, post-fire resprouting individuals had concentrated extrafloral nectar and sustained more ants, thus strengthening the outcomes of ant-plant mutualism.

Extrafloral Nectary-Bearing Plants Recover Ant Association Benefits Faster and More Effectively after Frost-Fire Events Than Frost.

The Cerrado confronts threats such as fire and frost due to natural or human-induced factors. These disturbances trigger attribute changes that impact biodiversity. Given escalating climate extremes, understanding the effects of these phenomena on ecological relationships is crucial for biodiversity conservation. To understand how fire and frost affect interactions and influence biological communities in the Cerrado, our study aimed to comprehend the effects of these two disturbances on extrafloral nectar (EFN)-bearing plants (Ouratea spectabilis, Ochnaceae) and their interactions. Our main hypothesis was that plants affected by fire would grow again more quickly than those affected only by frost due to the better adaptation of Cerrado flora to fire. The results showed that fire accelerated the regrowth of O. spectabilis. Regrowth in plants with EFNs attracted ants that proved to be efficient in removing herbivores, significantly reducing foliar herbivory rates in this species, when compared to the species without EFNs, or when ant access was prevented through experimental manipulation. Post-disturbance ant and herbivore populations were low, with frost leading to greater reductions. Ant richness and diversity are higher where frost precedes fire, suggesting that fire restores Cerrado ecological interactions better than frost, with less impact on plants, ants, and herbivores.

Time course of inducibility of indirect responses in an ant-defended plant.

Plants have evolved inducible defenses that allow them to minimize costs associated with the production of constitutive defenses when herbivores are not present. However, as a consequence, some plants might experience a period of vulnerability between damage and the onset of defense and/or between the cessation of damage and relaxation of defense. Few studies have examined the time course in the inducible protective mutualism between ants and extrafloral nectary (EFN)-bearing plants. None has compared the inducibility of EFNs on vegetative versus reproductive parts or in response to different levels of herbivore damage. Here, we disentangle the inducibility process by evaluating extrafloral nectar production and ant attendance over time, the time course of inducibility on different plant parts, and the time course of inducibility in response to different levels of foliar damage in a Brazilian tree, Qualea multiflora (Vochysiaceae). Using simulated herbivory on leaves and flowers, we found that (a) the production of extrafloral nectar from foliar and floral EFNs, as well as ant attendance, exhibited a lag between the moment of damage and the peak of response, followed by a response peak (usually 24 h after damage) at which the defense remains at its maximum level, then declines to prestimulus levels; (b) the time course of inducibility and the peak activity did not differ between EFNs located in vegetative versus reproductive parts, except for sugar concentration, which was higher in EFNs on vegetative parts; and (c) the time course of inducibility of foliar EFNs depended on damage level. Although considered a cost-saving strategy, inducible defenses can be disadvantageous since they can leave plants vulnerable to attack for extended periods. Our results illuminate the dynamics of the induced response and the underlying mechanisms that might mediate it, ultimately providing new insights into defense strategies employed by plants.

Context-Dependent Ant-Pollinator Mutualism Impacts Fruit Set in a Hummingbird-Pollinated Plant.

Context-dependence in mutualisms is a fundamental aspect of ecological interactions. Within plant-ant mutualisms, particularly in terms of biotic protection and pollination, research has predominantly focused on elucidating the benefits while largely overlooking potential costs. This notable gap underscores the need for investigations into the drawbacks and trade-offs associated with such mutualistic relationships. Here, we evaluated the role of pericarpial nectaries (PNs) in shaping the dynamics of ant-pollinator mutualisms. Specifically, we investigated whether ants visiting the PN of Palicourea rigida (Rubiaceae) could deter hummingbirds and disrupt pollination, ultimately influencing fruit production. Our research involved manipulative experiments and observation of ant-pollinator interactions on P. rigida plants in the Brazilian savannah. We found that visiting ants can deter hummingbirds and/or disrupt pollination in P. rigida, directly influencing fruit set. However, these results are species-specific. The presence of very aggressive, large predatory ants, such as E. tuberculatum, had a negative impact on hummingbird behavior, whereas aggressive mid-sized ants, such as C. crassus, showed no effects. Our study illuminates the multifaceted aspects of ant-plant mutualisms and underscores the importance of evaluating costs and unexpected outcomes within these ecological relationships.

From species descriptions to diversity patterns: the validation of taxonomic data as a keystone for ant diversity studies reproducibility and accuracy.

Research findings in natural sciences need to be comparable and reproducible to effectively improve our understanding of ecological and behavioural patterns. In this sense, knowledge frontiers in biodiversity studies are directly tied to taxonomic research, especially in species-rich tropical regions. Here we analysed the taxonomic information available in 470 studies on Brazilian ant diversity published in the last 50 years. We aimed to quantify the proportion of studies that provide enough data to validate taxonomic identification, explore the frequency of studies that properly acknowledge their taxonomic background, and investigate the primary resources for ant identification in Brazil. We found that most studies on Brazilian ant diversity (73.6%) explicitly stated the methods used to identify their specimens. However, the proportion of papers that provide complete data for the repository institutions and vouchered specimens is vanishingly small (5.8%). Additionally, only 40.0% of the studies consistently presented taxon authorities and years of description, rarely referencing taxonomic publications correctly. In turn, the number of specialists and institutions consulted for ant identification in Brazil has increased in the last years, along with the number of studies that explicitly provide their taxonomic procedures for ant identification. Our findings highlight a shift between generations regarding the recognition of taxonomy as fundamental science, deepening our understanding of biodiversity.

Can morphological and behavioral traits predict the foraging and feeding dynamics of social arachnids?

Complex social insect species exhibit task specialization mediated by morphological and behavioral traits. However, evidence of such traits is scarce for other social arthropods. We investigated whether the social pseudoscorpion Paratemnoides nidificator exhibits morphologically and behaviorally specialized individuals in prey capture. We measured body and chela sizes of adult pseudoscorpions and analyzed predation processes. Larger individuals spent more time moving through the colony and foraging than smaller pseudoscorpions. Individuals that captured prey had increased body and absolute chelae sizes. Although larger individuals had relatively small chelae size, they showed a higher probability of prey capture. Larger individuals manipulated prey often, although they fed less than smaller pseudoscorpions. Individuals that initiated captures fed more frequently and for more time than the others. Natural selection might be favoring individuals specialized in foraging and colony protection, allowing smaller and less efficient adults to avoid contact with dangerous prey. To our knowledge, there is incipient information regarding specialized individuals in arachnids, and our results might indicate the emergence of a morphologically specialized group in this species.

Cotesia itororensis sp. nov. from Brazilian savanna: a new reared microgastrine wasp (Hymenoptera: Braconidae) described using an integrative taxonomic approach.

A new species of microgastrine wasp, Cotesia itororensis Sousa-Lopes Whitfield, sp. nov., is described from a Brazilian savanna area in Uberlândia, Minas Gerais. This species is a koinobiont endoparasitoid recorded from caterpillars of Oospila pallidaria (Schaus, 1897) (Lepidoptera: Geometridae), feeding on Mimosa setosa var. paludosa (Fabaceae: Mimosoideae). Morphological, molecular, biological, ecological and geographical data are used to describe the new species and distinguish it from others formally recorded for the Neotropical region.

Unmasking the architecture of ant-diaspore networks in the Brazilian Savanna.

Ant-diaspore interactions are directly related to fruit consumption, seed predation and dispersal, being determinant for the plant fitness. However, although abundant and diversified, these ecological interactions have been neglected in network studies. Understanding the structure of these networks is the first step in preserving these ecological functions. However, describing the network structure is not enough; we need to understand what mechanisms are behind the network patterns. In this study, for the first time, we describe the structure of the ant-diaspore network, considering only the interactions that can benefit plants, separating it into fruit consumption and diaspore removal networks in the Brazilian Savanna. We postulated that ant-diaspore interactions tend to be more specialized in the diaspore removal network compared to the fruit consumption network. Furthermore, we tested whether morphological features, such as size of mandibles of ants and diaspores, could modulate these ecological networks. Overall, we recorded 24 ant and 29 plant species interacting. We found that fruit consumption and diaspore removal networks exhibited similar patterns of interactions (i.e., non-modular), although only the diaspore removal network was nested. The diaspore removal network did not show a more specialized pattern than the fruit consumption network, since both networks consisted of opportunistic interactions. We found that ant mandible and diaspore size does not explain the structure of ecological networks, but in diaspore removal networks the relationship between these morphological traits may explain the pattern of interactions. Thus, we showed that mandible size of ants may have implications on seedling recruitment, suggesting that mandible size can predict possible effects on plant fitness within in diaspore removal networks. Overall, ant-diaspore networks maintain important ecological functions, such as fruit consumption and seed dispersal, which often implies an increase in reproductive success of the plants.

Females of two species of Argia from Chapada dos Guimarães National Park, Brazil (Odonata: Coenagrionidae).

The female of Argia tupi Calvert, 1909 (BRAZIL, Mato Grosso, Chapada dos Guimarães National Park, Cachoeira do Marimbondo (15.4330° S, 55.7198° W, 370 m), 01 xi 2015) is described, illustrated and diagnosed based on comparison with sympatric species of Argia Rambur, 1842. We also augmented the description of Argia bicellulata (Calvert, 1909) female (BRAZIL, Mato Grosso, Chapada dos Guimarães National Park, Rio Paciencia (15.3438° S, 55.8322° W, 280 m), 25 x 2015).