Correction: A novel power-amplified jumping behavior in larval beetles (Coleoptera: Laemophloeidae).

[This corrects the article DOI: 10.1371/journal.pone.0256509.].

A morphofunctional study of the jumping apparatus in globular springtails.

Springtails are notable for their jumping apparatus and latch-mediated spring mechanism. The challenge, in the light of the tiny size and rapid movement of these organisms, has been to understand the morphological intricacies of this spring system. This study takes an approach that integrates SEM, MicroCT, cLSM and high-speed video recordings to understand the composition and functionality of the jumping apparatus in Megalothorax minimus (Neelipleona), Dicyrtomina ornata and Dicyrtomina minuta (Symphypleona). We focus on reconstructing, describing, and understanding the functioning of structures such as basal plates, musculature and furca. The dimensions of the jumping apparatus in Dicyrtomina and Megalothorax differ significantly from those in elongated springtails. A hypothesis of functional coherence between taxa, based on muscle connections and basal plates, is postulated. High-speed video recordings provide information on: 1) furca release timing and function during jumping and self-righting; 2) performance properties of manubrium, dens and mucro in interaction with the ground and in take-off; 3) possible pre-release furca moves. The study underscores the need for further research employing a variety of visualization methods in order to explore additional aspects such as retinaculum unlatching and furca flexion/extension muscles.

A novel power-amplified jumping behavior in larval beetles (Coleoptera: Laemophloeidae).

Larval insects use many methods for locomotion. Here we describe a previously unknown jumping behavior in a group of beetle larvae (Coleoptera: Laemophloeidae). We analyze and describe this behavior in Laemophloeus biguttatus and provide information on similar observations for another laemophloeid species, Placonotus testaceus. Laemophloeus biguttatus larvae precede jumps by arching their body while gripping the substrate with their legs over a period of 0.22 ± 0.17s. This is followed by a rapid ventral curling of the body after the larvae releases its grip that launches them into the air. Larvae reached takeoff velocities of 0.47 ± 0.15 m s-1 and traveled 11.2 ± 2.8 mm (1.98 ± 0.8 body lengths) horizontally and 7.9 ± 4.3 mm (1.5 ± 0.9 body lengths) vertically during their jumps. Conservative estimates of power output revealed that some but not all jumps can be explained by direct muscle power alone, suggesting Laemophloeus biguttatus may use a latch-mediated spring actuation mechanism (LaMSA) in which interaction between the larvae's legs and the substrate serves as the latch. MicroCT scans and SEM imaging of larvae did not reveal any notable modifications that would aid in jumping. Although more in-depth experiments could not be performed to test hypotheses on the function of these jumps, we posit that this behavior is used for rapid locomotion which is energetically more efficient than crawling the same distance to disperse from their ephemeral habitat. We also summarize and discuss jumping behaviors among insect larvae for additional context of this behavior in laemophloeid beetles.

Testing how different narrative perspectives achieve communication objectives and goals in online natural science videos.

Communication of science through online media has become a primary means of disseminating and connecting science with a public audience. However, online media can come in many forms and stories of scientific discovery can be told by many individuals. We tested whether the relationship of a spokesperson to the science story being told (i.e., the narrative perspective) influences how people react and respond to online science media. We created five video stimuli that fell into three treatments: a scientist presenting their own research (male or female), a third-party summarizing research (male or female), and an infographic-like video with no on-screen presenter. Each of these videos presented the same fabricated science story about the discovery of a new ant species (Formicidae). We used Qualtrics to administer and obtain survey responses from 515 participants (~100 per video). Participants were randomly assigned to one of the videos and after viewing the stimulus answered questions assessing their perceptions of the video (trustworthiness and enjoyment), the spokesperson (trustworthiness and competence), scientists in general (competence and warmth), and attitudes towards the research topic and funding. Participants were also asked to recall what they had seen and heard. We determined that when participants watched a video in which a scientist presented their own research, participants perceived the spokesperson as having more expertise than a third-party presenter, and as more trustworthy and having more expertise than the no-spokesperson stimuli. Viewing a scientist presenting their own work also humanized the research, with participants more often including a person in their answer to the recall question. Overall, manipulating the narrative perspective of the source of a single online video communication effort is effective at impacting immediate objective outcomes related to spokesperson perceptions, but whether those objectives can positively influence long-term goals requires more investigation.

Bacterial exposure leads to variable mortality but not a measurable increase in surface antimicrobials across ant species.

Social insects have co-existed with microbial species for millions of years and have evolved a diversity of collective defenses, including the use of antimicrobials. While many studies have revealed strategies that ants use against microbial entomopathogens, and several have shown ant-produced compounds inhibit environmental bacterial growth, few studies have tested whether exposure to environmental bacteria represents a health threat to ants. We compare four ant species' responses to exposure to Escherichia coli and Staphylococcus epidermidis bacteria in order to broaden our understanding of microbial health-threats to ants and their ability to defend against them. In a first experiment, we measure worker mortality of Solenopsis invicta, Brachymyrmex chinensis, Aphaenogaster rudis, and Dorymyrmex bureni in response to exposure to E. coli and S. epidermidis. We found that exposure to E. coli was lethal for S. invicta and D. bureni, while all other effects of exposure were not different from experimental controls. In a second experiment, we compared the antimicrobial ability of surface extracts from bacteria-exposed and non-exposed S. invicta and B. chinensis worker ants, to see if exposure to E. coli or S. epidermidis led to an increase in antimicrobial compounds. We found no difference in the inhibitory effects from either treatment group in either species. Our results demonstrate the susceptibility to bacteria is varied across ant species. This variation may correlate with an ant species' use of surface antimicrobials, as we found significant mortality effects in species which also were producing antimicrobials. Further exploration of a wide range of both bacteria and ant species is likely to reveal unique and nuanced antimicrobial strategies and deepen our understanding of how ant societies respond to microbial health threats.

External immunity in ant societies: sociality and colony size do not predict investment in antimicrobials.

Social insects live in dense groups with a high probability of disease transmission and have therefore faced strong pressures to develop defences against pathogens. For this reason, social insects have been hypothesized to invest in antimicrobial secretions as a mechanism of external immunity to prevent the spread of disease. However, empirical studies linking the evolution of sociality with increased investment in antimicrobials have been relatively few. Here we quantify the strength of antimicrobial secretions among 20 ant species that cover a broad spectrum of ant diversity and colony sizes. We extracted external compounds from ant workers to test whether they inhibited the growth of the bacterium Staphylococcus epidermidis. Because all ant species are highly social, we predicted that all species would exhibit some antimicrobial activity and that species that form the largest colonies would exhibit the strongest antimicrobial response. Our comparative approach revealed that strong surface antimicrobials are common to particular ant clades, but 40% of species exhibited no antimicrobial activity at all. We also found no correlation between antimicrobial activity and colony size. Rather than relying on antimicrobial secretions as external immunity to control pathogen spread, many ant species have probably developed alternative strategies to defend against disease pressure.

Snap-jaw morphology is specialized for high-speed power amplification in the Dracula ant, Mystrium camillae.

What is the limit of animal speed and what mechanisms produce the fastest movements? More than natural history trivia, the answer provides key insight into the form-function relationship of musculoskeletal movement and can determine the outcome of predator-prey interactions. The fastest known animal movements belong to arthropods, including trap-jaw ants, mantis shrimp and froghoppers, that have incorporated latches and springs into their appendage systems to overcome the limits of muscle power. In contrast to these examples of power amplification, where separate structures act as latch and spring to accelerate an appendage, some animals use a 'snap-jaw' mechanism that incorporates the latch and spring on the accelerating appendage itself. We examined the kinematics and functional morphology of the Dracula ant, Mystrium camillae, who use a snap-jaw mechanism to quickly slide their mandibles across each other similar to a finger snap. Kinematic analysis of high-speed video revealed that snap-jaw ant mandibles complete their strike in as little as 23 µsec and reach peak velocities of 90 m s-1, making them the fastest known animal appendage. Finite-element analysis demonstrated that snap-jaw mandibles were less stiff than biting non-power-amplified mandibles, consistent with their use as a flexible spring. These results extend our understanding of animal speed and demonstrate how small changes in morphology can result in dramatic differences in performance.

The evolution of cuticular fertility signals in eusocial insects.

A reproductive division of labor is a definitive characteristic of eusocial insect societies and it requires a means through which colony members can assess the presence and productivity of reproductive individuals. Cuticular hydrocarbons are the primary means of doing so across eusocial hymenopterans. However, recent experimental work presents conflicting views on how these chemical signals function, are interpreted by workers, and evolve. These recent advances include demonstrations of hydrocarbons as evolutionarily conserved 'queen pheromones' and as species-divergent 'fertility signals' used by both queens and workers. In this review, we synthesize conflicting studies into an evolutionary framework suggesting a transition of reproductive communication from cue-like signature mixtures, to learned fertility signals, to innate queen pheromones that evolved across eusocial insects.

Comparative analysis of fertility signals and sex-specific cuticular chemical profiles of Odontomachus trap-jaw ants.

The lipid mixture that coats the insect cuticle contains a number of chemical signals. Mate choice in solitary insects is mediated by sexually dimorphic cuticular chemistry, whereas in eusocial insects, these profiles provide information through which colony members are identified and the fertility status of individuals is assessed. Profiles of queens and workers have been described for a number of eusocial species, but there have been few comparisons of fertility signals among closely related species. Additionally, sexual dimorphism in cuticular lipid profiles has only been reported in two species of ants. This study describes the cuticular chemical profiles of queens, workers and males of three species of Odontomachus trap-jaw ants: O. ruginodis, O. relictus and O. haematodus. These are compared with fertility signals and sexually dimorphic profiles already described from O. brunneus. We report that fertility signals are not conserved within this genus: chemical compounds that distinguish queens from workers vary in number and type among the species. Furthermore, the compounds that were most abundant in cuticular extracts of O. ruginodis queens relative to workers were novel 2,5-dialkyltetrahydrofurans. Bioassays of extracts of O. ruginodis queens indicate that the dialkyltetrahydrofuran and hydrocarbon fractions of the profile are likely to work synergistically in eliciting behavioral responses from workers. In contrast, cuticular lipids that distinguish males from females are more conserved across species, with isomeric and relative abundance variations comprising the main differences among species. Our results provide new insights into how these contact chemical signals may have arisen and evolved within eusocial insects.

A social insect fertility signal is dependent on chemical context.

Identifying group members and individuals' status within a group are fundamental tasks in animal societies. For ants, this information is coded in the cuticular hydrocarbon profile. We manipulated profiles of the ant Odontomachus brunneus to examine whether the releaser and primer effects of fertility signals are dependent on chemical context. Fertility status is signalled through increased abundance of (Z)-9-nonacosene (Z9 : C29). Across the ant's distribution, populations have distinct hydrocarbon profiles but the fertility signal is conserved. Foreign queens and fertility-signal-treated workers from the same population, sharing a similar chemical background, elicited releaser effects from workers, whereas queens and fertility-signal-treated workers from different populations did not. Z9 : C29 presented without chemical background did not elicit releaser effects. A primer-effect experiment found that Z9 : C29, presented without a chemical background, did not inhibit worker reproduction. Our results demonstrate that a familiar chemical background is necessary for appropriate responses to fertility signals.

A conserved fertility signal despite population variation in the cuticular chemical profile of the trap-jaw ant Odontomachus brunneus.

Contact pheromones in the form of cuticular hydrocarbons are widespread among insects. Eusocial insects present a special challenge for understanding the evolution of the cuticular hydrocarbon profile because this blend is responsible for multiple distinct roles such as nestmate recognition and signalling fertility status. This study investigates these two signalling roles of the hydrocarbon profile in the trap-jaw ant Odontomachus brunneus. We demonstrate that the cuticular hydrocarbon profile is highly variable across populations and provide evidence that these differences are used for nestmate discrimination. Through manipulative experiments we also show that (Z)-9-nonacosene (Z9:C29) is used as a fertility signal and its role is conserved across populations. Our data demonstrate that both fertility and nestmate signalling influence the cuticular hydrocarbon profile and specifically the relative abundance of Z9:C29 on the cuticle of O. brunneus. Our study suggests that natural selection works on the cuticular chemical profile through multiple regulatory pathways, diversifying nestmate signals while conserving fertility signals.

The role of anchor-tipped larval hairs in the organization of ant colonies.

The spatial organization within a social insect colony is a key component of colony life. It influences individual interaction rates, resource distribution, and division of labor within the nest. Yet studies of social insect behavior are most often carried out in artificial constructions, which may change worker behavior and colony organization. We observed how workers of the ant Pheidole rhea organized brood in nests with deep chambers and textured walls that were designed to mimic their natural constructions more closely. Instead of clumping larvae into piles on the chamber floor, workers suspended fourth-instar larvae from the vertical walls and ceiling of each chamber while young larvae and pupae were clumped at the base. Fourth-instar larvae possess five rows of anchor-tipped hairs on their dorsal side, and we predicted that these hairs functioned to attach larvae to the nest walls. We gave larvae "haircuts," where only the anchor-tipped hairs were removed, and then tested their ability to adhere to a textured surface raised to an angle of 90° and then 120° with respect to the horizontal plane. Larvae whose hairs had been clipped came unattached in almost all trials, while larvae whose hairs remained intact stayed attached. This confirmed that anchor-tipped hairs functioned to attach larvae to the walls of the nest. The presence of anchor-tipped hairs is widespread and has been documented in at least 22 genera from the ant subfamily Myrmicinae, including species that occur in a variety of environments and represent a broad range of nesting habits. Based on our results, it is likely that many species exhibit this larval hanging behavior, and this could impact colony characteristics such as spatial organization and the care of developing larvae by nurse workers.

Reclaiming the crown: queen to worker conflict over reproduction in Aphaenogaster cockerelli.

In many social taxa, reproductively dominant individuals sometimes use aggression to secure and maintain reproductive status. In the social insects, queen aggression towards subordinate individuals or workers has been documented and is predicted to occur only in species with a small colony size and a low level of queen-worker dimorphism. We report queen aggression towards reproductive workers in the ant species Aphaenogaster cockerelli, a species with a relatively large colony size and a high level of reproductive dimorphism. Through analysis of cuticular hydrocarbon profiles, we show that queens are aggressive only to reproductively active workers. Non-reproductive workers treated with a hydrocarbon typical for reproductives are attacked by workers but not by queens, which suggests different ways of recognition. We provide possible explanations of why queen aggression is observed in this species.

Social insects inspire human design.

The international conference 'Social Biomimicry: Insect Societies and Human Design', hosted by Arizona State University, USA, 18-20 February 2010, explored how the collective behaviour and nest architecture of social insects can inspire innovative and effective solutions to human design challenges. It brought together biologists, designers, engineers, computer scientists, architects and businesspeople, with the dual aims of enriching biology and advancing biomimetic design.

Cuticular hydrocarbons reliably identify cheaters and allow enforcement of altruism in a social insect.

Cheaters are a threat to every society and therefore societies have established rules to punish these individuals in order to stabilize their social system. Recent models and observations suggest that enforcement of reproductive altruism (policing) in hymenopteran insect societies is a major force in maintaining high levels of cooperation. In order to be able to enforce altruism, reproductive cheaters need to be reliably identified. Strong correlational evidence indicates that cuticular hydrocarbons are the means of identifying cheaters, but direct proof is still missing. In the ant Aphaenogaster cockerelli, we mimicked reproductive cheaters by applying a synthetic compound typical of fertile individuals on nonreproductive workers. This treatment induced nestmate aggression in colonies where a queen was present. As expected, it failed to do so in colonies without a queen where workers had begun to reproduce. This provides the first direct evidence that cuticular hydrocarbons are the informational basis of policing behaviors, serving a major function in the regulation of reproduction in social insects. We suggest that even though cheaters would gain from suppressing these profiles, they are prevented from doing so through the mechanisms of hydrocarbon biosynthesis and its relation to reproductive physiology. Cheaters are identified through information that is inherently reliable.

Hydrocarbon signals explain the pattern of worker and egg policing in the ant Aphaenogaster cockerelli.

In ant societies, worker reproduction is regulated through policing behaviors, such as physical aggression or egg eating. The information used by policing individuals is thought to be in blends of hydrocarbons present on the cuticle and the surface of eggs. These fertility signals have been studied in numerous genera. However, signaling patterns that emerge across distinct subfamilies of ants have yet to be explained. We investigated policing behavior and the chemical signaling upon which policing behaviors are informed in the ant Aphaenogaster cockerelli. We found that worker-produced eggs are not policed, and we showed that there is a lack of chemical signaling for effective egg policing to occur in this species. Furthermore, we identified the available signals that demarcate workers to be policed physically. We showed that in A. cockerelli, a species with derived social organization, workers produce fertility signals identical to the queen. This queen-like signaling may be due to workers maintaining a high level of ovarian activity, linked to trophic egg production, in the presence of the queen.

Army ants as research and collection tools.

Ants that fall prey to the raids of army ants commonly respond by evacuating their nests. This documented behavior has been underexploited by researchers as an efficient research tool. This study focuses on the evacuation response of the southwestern desert ant Aphaenogaster cockerelli André (Hymenoptera: Formicidae) to the army ant Newamyrmex nigrescens Cresson. It is shown that army ants can be used to collect mature colonies of ants. The applicability of this tool to ecologically meaningful areas of research is discussed.