Beetle or roach: systematic position of the enigmatic Umenocoleidae based on new material from Zhonggou Formation in Jiuquan,Northwest China,and a morphocladistic analysis

The Umenocoleidae is among the most perplexing fossil insect taxa, widespread at lower and middle latitudes during the Early Cretaceous and probably becoming extinct after the mid-Cretaceous. This problematic taxon was first described from the Lower Cretaceous of China and has been variously assigned to stem-group Coleoptera, Protelytroptera, Blattaria, stem-group Dictyoptera, or Mantodea. The systematic position of the enigmatic family — especially its type species Umenocoleus sinuatus Chen et T’an, 1973 — has remained highly controversial owing to the lack of preserved phylogenetically informative or diagnostic characters. Here, we describe a new specimen of the type species from the type horizon of the type locality (Lower Cretaceous Zhonggou Formation at Jiuquan, Yumen City, China), and additionally two forewings of Ponopterix. We found that the Umenocoleidae is similar to stem-group Coleoptera in its forewing venation but differs in its multi-segmented antennae, orthognathous orientation of the head, subdivision of the pronotum by a transverse furrow, ultrastructure of the forewing punctures, and unfolded hindwings. Our phylogenetic analysis, using an extended matrix of 72 characters and 37 taxa, indicates that the Umenocoleidae is likely a specialized taxon of Dictyoptera, sister to Alienoptera. The beetle-like appearance is interpreted to be a result of convergent evolution.     

Myrmica ants host highly diverse parasitic communities: from social parasites to microbes

Myrmica ants have been model species for studies in a variety of disciplines, including insect physiology, chemical communication, ant social dynamics, ant population, community ecology, and ant interactions with other organisms. Species belonging to the genus Myrmica can be found in virtually every habitat within the temperate regions of the northern hemisphere and their biology and systematics have been thoroughly studied. These ants serve as hosts to highly diverse parasitic organisms from socially parasitic butterfly caterpillars to microbes, and many Myrmica species even evolved into parasitizing species of their own genus. These parasites have various impacts both on the individuals and on the social structure of their hosts, ranging from morphological malformations to reduction in colony fitness. A comprehensive review of the parasitic organisms supported by Myrmica and the effects of these organisms on individuals and on whole ant colonies has not yet been compiled. Here, we provide a review of the interactions of these organisms with Myrmica ants by discussing host and parasite functional, behavioral or physiological adaptations. In addition, for all “symbiont groups” of Myrmica ants described in this paper, we examine the present limitations of the knowledge at present of their impact on individuals and host colony fitness. In conclusion, we argue that Myrmica ants serve as remarkable resource for the evolution of a wide variety of associated organisms.     

Reduced ant defenses in <Emphasis Type="Italic">Macaranga</Emphasis> myrmecophytes (Euphorbiaceae) infested with a winged phasmid <Emphasis Type="Italic">Orthomeria cuprinus</Emphasis>

Macaranga is a tree genus that includes many species of myrmecophytes, which are plants that harbor ant colonies within hollow structures known as domatia. The symbiotic ants (plant–ants) protect their host plants against herbivores; this defense mechanism is called ‘ant defense’. A Bornean phasmid species Orthomeria cuprinus feeds on two myrmecophytic Macaranga species, Macaranga beccariana and Macaranga hypoleuca, which are obligately associated with Crematogaster ant species. The phasmids elude the ant defense using specialized behavior. However, the mechanisms used by the phasmid to overcome ant defenses have been insufficiently elucidated. We hypothesized that O. cuprinus only feeds on individual plants with weakened ant defenses. To test the hypothesis, we compared the ant defense intensity in phasmid-infested and non-infested M. beccariana trees. The number of plant–ants on the plant surface, the ratio of plant–ant biomass to tree biomass, and the aggressiveness of plant–ants towards experimentally introduced herbivores were significantly lower on the phasmid-infested trees than on the non-infested trees. The phasmid nymphs experimentally introduced into non-infested trees, compared with those experimentally introduced into phasmid-infested trees, were more active on the plant surface, avoiding the plant–ants. These results support the hypothesis and suggest that ant defenses on non-infested trees effectively prevent the phasmids from remaining on the plants. Thus, we suggest that O. cuprinus feeds only on the individual M. beccariana trees having decreased ant defenses, although the factors that reduce the intensity of the ant defenses remain unclear.     

Rainfall facilitates the spread, and time alters the impact, of the invasive Argentine ant

Climate change may exacerbate invasions by making conditions more favorable to introduced species relative to native species. Here we used data obtained during a long-term biannual survey of the distribution of ant species in a 481-ha preserve in northern California to assess the influence of interannual variation in rainfall on the spread of invasive Argentine ants, Linepithema humile, and the displacement of native ant species. Since the survey began in 1993, Argentine ants have expanded their range into 74 new hectares. Many invaded hectares were later abandoned, so the range of Argentine ants increased in some years and decreased in others. Rainfall predicted both range expansion and interannual changes in the distribution of Argentine ants: high rainfall, particularly in summer months, promoted their spread in the summer. This suggests that an increase in rainfall will promote a wider distribution of Argentine ants and increase their spread into new areas in California. Surprisingly, the distribution of two native ant species also increased following high rainfall, but only in areas of the preserve that were invaded by L. humile. Rainfall did not have a negative impact on total native ant species richness in invaded areas. Instead, native ant species richness in invaded areas increased significantly over the 13 years of observation. This suggests that the impact of Argentine ants on naïve ant communities may be most severe early in the invasion process.     

From species to individuals: does the variation in ant–plant networks scale result in structural and functional changes?

Predicting the outcomes of any mutualistic interaction between ants and plants can be a very difficult task, since these outcomes are often determined by the ecological context in which the interacting species are embedded. Network theory has been an important tool to improve our understanding about the organizational patterns of animal–plant interactions. Nevertheless, traditionally, network studies have focused mainly on species-based differences and ignoring the importance of individual differences within populations. In this study, we evaluated if downscaling an ant–plant network from species to the individual level results in structural and functional changes in a network involving different-sized plant individuals. For this, we studied the extrafloral-nectar producing-tree Caryocar brasiliense (Caryocaraceae) and their associated ants in a Neotropical savanna. We observed 254 interactions involving 43 individuals of C. brasiliense and 47 ant species. The individual-based ant–plant network exhibited a nested pattern of interactions, with all developmental stages contributing equally to structuring this non-random pattern. We also found that plants with greater centrality within the network were better protected by their ant partners. However, plants with higher levels of individual specialization were not necessarily better protected by ants. Overall, we presented empirical evidence that intra-population variations are important for shaping ant–plant networks, since they can change the level of protection against herbivores conferred by the ants. These results highlight the importance of individual-based analyses of ecological networks, opening new research venues in the eco-evolutionary dynamics of ant–plant interactions.     

Invasive ants carry novel viruses in their new range and form reservoirs for a honeybee pathogen

When exotic animal species invade new environments they also bring an often unknown microbial diversity, including pathogens. We describe a novel and widely distributed virus in one of the most globally widespread, abundant and damaging invasive ants (Argentine ants, Linepithema humile). The Linepithema humile virus 1 is a dicistrovirus, a viral family including species known to cause widespread arthropod disease. It was detected in samples from Argentina, Australia and New Zealand. Argentine ants in New Zealand were also infected with a strain of Deformed wing virus common to local hymenopteran species, which is a major pathogen widely associated with honeybee mortality. Evidence for active replication of viral RNA was apparent for both viruses. Our results suggest co-introduction and exchange of pathogens within local hymenopteran communities. These viral species may contribute to the collapse of Argentine ant populations and offer new options for the control of a globally widespread invader.     

Ant interactions with native and exotic seeds in the Patagonian steppe: Influence of seed traits,disturbance levels and ant assemblage

Invasive plants may establish strong interactions with species in their new range which could limit or enhance their establishment and spread. These interactions depend upon traits of the invader and the recipient community, and may alter interactions among native species. In the Patagonian steppe we studied interactions of native ant assemblages with seeds of native and exotic plants, and asked whether ant–seed interactions differ with seed types and disturbance levels and whether the amount and type of ant–seed interactions can be predicted if both ant and seed traits are known. To characterize and quantify ant–seed interactions, we offered baits with large seeds of Pappostipa speciosa (native) and medium-sized elaiosome-bearing seeds of Carduus thoermeri (exotic), near and far from a road (high vs. low disturbed areas), and compared ant abundance and composition between areas. Interaction frequency was the highest for C. thoermeri seeds far from the road. Composition of ants interacting with C. thoermeri in these areas differed from that near the road and from that interacting with native seeds. Ant composition and abundance were similar between areas, but some species interacted more with exotic seeds in low disturbed areas. Ant foraging type predicted ant–seed interactions since the abundance of seed harvesters was positively correlated to interactions with P. speciosa, and that of generalists and predators, with interactions with C. thoermeri. The high interaction of ants with exotic seeds in low invaded areas suggests that ant activity could influence plant invasion, either by predating or dispersing seeds of invasive plants.     

Macroevolutionary assembly of ant/plant symbioses: Pseudomyrmex ants and their ant-housing plants in the Neotropics

Symbioses include some of the clearest cases of coevolution, but their origin, loss or reassembly with different partners can rarely be inferred. Here we use ant/plant symbioses involving three plant clades to investigate the evolution of symbioses. We generated phylogenies for the big-eyed arboreal ants (Pseudomyrmecinae), including 72% of their 286 species, as well as for five of their plant host groups, in each case sampling more than 61% of the species. We show that the ant-housing Vachellia (Mimosoideae) clade and its ants co-diversified for the past 5 Ma, with some species additionally colonized by younger plant-nesting ant species, some parasitic. An apparent co-radiation of ants and Tachigali (Caesalpinioideae) was followed by waves of colonization by the same ant clade, and subsequent occupation by a younger ant group. Wide crown and stem age differences between the ant-housing genus Triplaris (Polygonaceae) and its obligate ant inhabitants, and stochastic trait mapping, indicate that its domatium evolved earlier than the ants now occupying it, suggesting previous symbioses that dissolved. Parasitic ant species evolved from generalists, not from mutualists, and are younger than the mutualistic systems they parasitize. Our study illuminates the macroevolutionary assembly of ant/plant symbioses, which has been highly dynamic, even in very specialized systems.     

Booms,busts and population collapses in invasive ants

The abundance of many invasive species can vary substantially over time, with dramatic population declines and local extinctions frequently observed in a wide range of taxa. We highlight population crashes of invasive ants, which are some of the most widespread and damaging invasive animals. Population collapse or substantial declines have been observed in nearly all of the major invasive ant species including the yellow crazy ant (Anoplolepis gracilipes), Argentine ants (Linepithema humile), big-headed or coastal brown ant (Pheidole megacephala), the tropical fire ant (Solenopsis geminata), red imported fire ants (Solenopsis invicta), and the little fire ant or electric ant (Wasmannia auropunctata). These declines frequently attract little attention, especially compared with their initial invasion phase. Suggested mechanisms for population collapse include pathogens or parasites, changes in the food availability, or even long-term effects of the reproductive biology of invasive ants. A critical component of the collapses may be a reduction in the densities of the invasive ant species, which are often competitively weak in low abundance. We propose that mechanisms causing a reduction in invasive ant abundance may initiate a local extinction vortex. Declines in abundance likely reduce the invasive ant’s competitive ability, resource acquisition and defense capability. These reductions could further reduce the abundance of an invasive ant species, and so on. Management of invasive ants through the use of pesticides is expensive, potentially ecologically harmful, and can be ineffective. We argue that pesticide use may even have the potential to forestall natural population declines and collapses. We propose that in order to better manage these invasive ants, we need to understand and capitalize on features of their population dynamics that promote population collapse.     

Differential Recruitment of Camponotus femoratus (Fabricius) Ants in Response to Ant Garden Herbivory

Although several studies have shown that ants can recognize chemical cues from their host plants in ant-plant systems, it is poorly demonstrated in ant gardens (AGs). In this interaction, ant species constantly interact with various epiphyte species. Therefore, it is possible to expect a convergence of chemical signals released by plants that could be acting to ensure that ants are able to recognize and defend epiphyte species frequently associated with AGs. In this study, it was hypothesized that ants recognize and differentiate among chemical stimuli released by AG epiphytes and non-AG epiphytes. We experimentally simulated leaf herbivore damage on three epiphyte species restricted to AGs and a locally abundant understory herb, Piper hispidum, in order to quantify the number of recruited Camponotus femoratus (Fabricius) defenders. When exposed to the AG epiphytes Peperomia macrostachya and Codonanthe uleana leaves, it was observed that the recruitment of C. femoratus workers was, on average, respectively 556% and 246% higher than control. However, the number of ants recruited by the AG epiphyte Markea longiflora or by the non-AG plant did not differ from paper pieces. This indicated that ants could discern between chemicals released by different plants, suggesting that ants can select better plants. These results can be explained by evolutionary process acting on both ants’ capability in discerning plants’ chemical compounds (innate attraction) or by ants’ learning based on the epiphyte frequency in AGs (individual experience). To disentangle an innate behavior, a product of classical coevolutionary process, from an ant’s learned behavior, is a complicated but important subject to understand in the evolution of ant-plant mutualisms.     

The Effect of Diet and Opponent Size on Aggressive Interactions Involving Caribbean Crazy Ants (Nylanderia fulva)

Biotic interactions are often important in the establishment and spread of invasive species. In particular, competition between introduced and native species can strongly influence the distribution and spread of exotic species and in some cases competition among introduced species can be important. The Caribbean crazy ant, Nylanderia fulva, was recently introduced to the Gulf Coast of Texas, and appears to be spreading inland. It has been hypothesized that competition with the red imported fire ant, Solenopsis invicta, may be an important factor in the spread of crazy ants. We investigated the potential of interspecific competition among these two introduced ants by measuring interspecific aggression between Caribbean crazy ant workers and workers of Solenopsis invicta. Specifically, we examined the effect of body size and diet on individual-level aggressive interactions among crazy ant workers and fire ants. We found that differences in diet did not alter interactions between crazy ant workers from different nests, but carbohydrate level did play an important role in antagonistic interactions with fire ants: crazy ants on low sugar diets were more aggressive and less likely to be killed in aggressive encounters with fire ants. We found that large fire ants engaged in fewer fights with crazy ants than small fire ants, but fire ant size affected neither fire ant nor crazy ant mortality. Overall, crazy ants experienced higher mortality than fire ants after aggressive encounters. Our findings suggest that fire ant workers might outcompete crazy ant workers on an individual level, providing some biotic resistance to crazy ant range expansion. However, this resistance may be overcome by crazy ants that have a restricted sugar intake, which may occur when crazy ants are excluded from resources by fire ants.     

Interspecific competition between the red imported fire ant,Solenopsis invicta Buren and ghost ant,Tapinoma melanocephalum Fabricius for honeydew resources produced by an invasive mealybug,Phenacoccus solenopsis Tinsiley

In natural as in agricultural ecosystems, interactions between ants and honeydew-producing hemipterans are commonly observed. Mutualisms between invasive ants and hemipterans have been extensively studied in recent years. However, native ant species can equally exploit the honeydew excreted by hemipterans, and establish close relationships with them. Up till present, little is known about the competition between exotic ants (such as Solenopsis invicta) and its co-occurring species (e.g., Tapinoma melanocephalum) for this food resource. In this study, we compared the competitive ability of the invasive ant S. invicta and its co-occurring species T. melanocephalum in the laboratory. We also determined whether the two ant species could coexist and share honeydew resource. Our results indicate that the foraging activity of T. melanocephalum was restrained by S. invicta. Mortality of S. invicta and T. melanocephalum was significantly higher in T. melanocephalum colony case than that in other cases. The invasive ability between the two ant species was significantly different. These results suggest that S. invicta suppresses exploitation of honeydew-producing hemipterans by native ants and occupies most of honeydew resource. S. invicta could not completely drive T. melanocephalum out of honeydew competition, with small numbers of T. melanocephalum workers coexisting and sharing the honeydew with S. invicta. This finding permits a better understanding of the invasion success of S. invicta, and its ability to occupy new habitats.     

Arboreal thorn-dwelling ants coexisting on the savannah ant-plant, Vachellia erioloba, use domatia morphology to select nest sites

Nest site selection in arboreal, domatia-dwelling ants, particularly those coexisting on a single host plant, is little understood. To examine this phenomenon we studied the African savannah tree Vachellia erioloba, which hosts ants in swollen-thorn domatia. We found four ant species from different genera (Cataulacus intrudens, Tapinoma subtile, Tetraponera ambigua and an unidentified Crematogaster species). In contrast to other African ant plants, many V. erioloba trees (41 % in our survey) were simultaneously co-occupied by more than one ant species. Our study provides quantitative field data describing: (1) aspects of tree and domatia morphology relevant to supporting a community of mutualist ants, (2) how ant species occupancy varies with domatia morphology and (3) how ant colony size varies with domatia size and species. We found that Crematogaster sp. occupy the largest thorns, followed by C. intrudens, with T. subtile in the smallest thorns. Thorn age, as well as nest entrance hole size correlated closely with ant species occupant. These differing occupancy patterns may help to explain the unusual coexistence of three ant species on individual myrmecophytic trees. In all three common ant species, colony size, as measured by total number of ants, increased with domatia size. Additionally, domatia volume and species identity interact to predict ant numbers, suggesting differing responses between species to increased availability of nesting space. The proportion of total ants in nests that were immatures varied with thorn volume and species, highlighting the importance of domatia morphology in influencing colony structure.     

Trail traffic flow prediction by contact frequency among individual ants

Ants build a trail that leads to a new location when they move their colony. The trail’s traffic flows smoothly, regardless of the density on the trail. To the best of our knowledge, such a phenomenon has been reported only for ant species. The trail’s capacity is known as trail traffic flow. In this paper, we propose a probabilistic model of trail traffic flow, which overcomes some inadequacies of the kinetic model previously proposed in the literature. Our model answers a question unsolved by the previous model, namely, how many worker ants form such a density-independent trail. We focus on ants’ responses to mutual contacts that involve individuals in trail formation. We propose a model in which contact frequency predicts the number of worker ants that form a trail. We verify that our model’s estimates match the empirical data that ant experts reported in the literature. In modeling and evaluation, we discuss an intelligent ant species, the house-hunting ant Temnothorax albipennis, which is popular among the ant experts.     

Asymmetric dispersal and colonization success of Amazonian plant-ants queens

Background

The dispersal ability of queens is central to understanding ant life-history evolution, and plays a fundamental role in ant population and community dynamics, the maintenance of genetic diversity, and the spread of invasive ants. In tropical ecosystems, species from over 40 genera of ants establish colonies in the stems, hollow thorns, or leaf pouches of specialized plants. However, little is known about the relative dispersal ability of queens competing for access to the same host plants.

Methodology/Principal Findings

We used empirical data and inverse modeling—a technique developed by plant ecologists to model seed dispersal—to quantify and compare the dispersal kernels of queens from three Amazonian ant species that compete for access to host-plants. We found that the modal colonization distance of queens varied 8-fold, with the generalist ant species (Crematogaster laevis) having a greater modal distance than two specialists (Pheidole minutula, Azteca sp.) that use the same host-plants. However, our results also suggest that queens of Azteca sp. have maximal distances that are four-sixteen times greater than those of its competitors.

Conclusions/Significance

We found large differences between ant species in both the modal and maximal distance ant queens disperse to find vacant seedlings used to found new colonies. These differences could result from interspecific differences in queen body size, and hence wing musculature, or because queens differ in their ability to identify potential host plants while in flight. Our results provide support for one of the necessary conditions underlying several of the hypothesized mechanisms promoting coexistence in tropical plant-ants. They also suggest that for some ant species limited dispersal capability could pose a significant barrier to the rescue of populations in isolated forest fragments. Finally, we demonstrate that inverse models parameterized with field data are an excellent means of quantifying the dispersal of ant queens.     

Leaf beetles are ant-nest beetles: the curious life of the juvenile stages of case-bearers (Coleoptera,Chrysomelidae, Cryptocephalinae)

Although some species of Cryptocephalinae (Coleoptera: Chrysomelidae) have been documented with ants (Hymenoptera: Formicidae) for almost 200 years, information on this association is fragmentary. This contribution synthesizes extant literature and analysizes the data for biological patterns. Myrmecophily is more common in the tribe Clytrini than in Cryptocephalini, but not documented for Fulcidacini or the closely-related Lamprosomatinae. Myrmecophilous cryptocephalines (34 species in 14 genera) primarily live among formicine and myrmecines ants as hosts. These two ant lineages are putative sister-groups, with their root-node dated to between 77–90 mya. In the New World tropics, the relatively recent radiation of ants from moist forests to more xeric ecosystems might have propelled the association of cryptocephalines and ant nests. Literature records suggest that the defensive behavioral profile or chemical profile (or both) of these ants has been exploited by cryptocephalines. Another pattern appears to be that specialized natural enemies, especially parasitoid Hymenoptera, exploit cryptocephaline beetles inside the ant nests. With the extant data at hand, based on the minimum age of a fossil larva dated to 45 mya, we can infer that the origin of cryptocephaline myrmecophily could have arisen within the Upper Cretaceous or later. It remains unknown how many times myrmecophily has appeared, or how old is the behavior. This uncertainty is compounded by incongruent hypotheses about the origins of Chrysomelidae and angiosperm-associated lineages of cryptocephalines. Living with ants offers multiple advantages that might have aided the colonization of xeric environments by some cryptocephaline species.     

Thorn‐dwelling ants provide antiherbivore defence for camelthorn trees,Vachellia erioloba,in Namibia

Ants are widely employed by plants as an antiherbivore defence. A single host plant can associate with multiple, symbiotic ant species, although usually only a single ant species at a time. Different plant‐ant species may vary in the degree to which they defend their host plant. In Kenya, ant–acacia interactions are well studied, but less is known about systems elsewhere in Africa. A southern African species, Vachellia erioloba, is occupied by thorn‐dwelling ants from three different genera. Unusually, multiple colonies of all these ants simultaneously and stably inhabit trees. We investigated if the ants on V. erioloba (i) deter insect herbivores; (ii) differ in their effectiveness depending on the identity of the herbivore; and (iii) protect the tree against an important herbivore, the larvae of the lepidopteran Gonometa postica. We show that experimental exclusion of ants leads to greater levels of herbivory on trees. The ants inhabiting V. erioloba are an effective deterrent against hemipteran and coleopteran, but not lepidopteran herbivores. Defensive services do not vary among ant species, but only Crematogaster ants exhibit aggression towards G. postica. This highlights the potential of the V. erioloba–ant mutualism for studying ant–plant interactions that involve multiple, simultaneously resident thorn‐dwelling ant species.     

Interactions between ant species increase arrival rates of an ant parasitoid

Behaviours or traits associated with aggression and communication may increase an animal's conspicuousness to predators or parasitoids. Most examples of this come from instances of aggression or communication within a species. We tested whether interspecific encounters between ants enhance the host location success of a parasitoid (Diptera: Phoridae) that attacks ants in the genus Linepithema. At food resources recruited to by Linepithema, parasitoid discovery rates were lower when Linepithema was alone than when other ant species were present. In experimentally controlled encounters, parasitoid discovery rates were elevated when Linepithema confronted an ant species that elicited use of chemicals, but not when it confronted an ant species that primarily elicited physical aggression. These results indicate that phorid parasitoids of Linepithema use the ant's chemicals as host location cues. Because Linepithema is known to abandon food resources in the presence of its phorid parasitoids, its use of chemicals during interspecific encounters may diminish its competitive success when phorids are nearby. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.