Warning Patterns Research Articles

The visual memorability of natural warning patterns: insights from humans and machines

The visual memorability of natural warning patterns: insights from humans and machines

Pattern Mining-Based Warning Prioritization by Refining Abstract Syntax Tree

Static code analysis tools (SATs) are widely used to detect potential defects in software projects. However, the usability of SATs is seriously hindered by a large number of unactionable warnings. Currently, many warning prioritization approaches are proposed to improve the usability of SATs. These approaches mainly extract different warning features to capture the statistical or historical information of warnings, thereby ranking actionable warnings in front of unactionable warnings. Such features are extracted by extremely relying on domain knowledge. However, the precise domain knowledge is difficult to be acquired. Also, the domain knowledge obtained in a project cannot be directly applied to other projects due to different application scenarios among different projects. To address the above problem, we propose a pattern mining-based warning prioritization approach based on the warning-related Abstract Syntax Tree (AST). To automatically mine actionable warning patterns, our approach leverages an advanced technique to collect actionable warnings, designs an algorithm to extract the warning-related AST, and mines patterns from ASTs of all actionable warnings. To prioritize the newly reported warnings, our approach combines exact and fuzzing matching techniques to calculate the similarity score between patterns of the newly reported warnings and the mined actionable warning patterns. We compare our approach with four typical baselines on five open-source and large-scale Java projects. The results show that our approach outperforms four baselines and achieves the maximum MAP (0.76) and MRR (2.19). Besides, a case study on Defect4J dataset demonstrates that our approach can discover 83% of true defects in the top 10 warnings.

Adaptive introgression of a visual preference gene.

Visual preferences are important drivers of mate choice and sexual selection, but little is known of how they evolve at the genetic level. In this study, we took advantage of the diversity of bright warning patterns displayed by Heliconius butterflies, which are also used during mate choice. Combining behavioral, population genomic, and expression analyses, we show that two Heliconius species have evolved the same preferences for red patterns by exchanging genetic material through hybridization. Neural expression of regucalcin1 correlates with visual preference across populations, and disruption of regucalcin1 with CRISPR-Cas9 impairs courtship toward conspecific females, providing a direct link between gene and behavior. Our results support a role for hybridization during behavioral evolution and show how visually guided behaviors contributing to adaptation and speciation are encoded within the genome.

Divergent warning patterns influence male and female mating behaviours in a tropical butterfly.

Traits under divergent ecological selection that also function during mating can be important in maintaining species boundaries. Few studies have considered mutual mate choice, where both males and females base mating decisions on the same trait. Wing colouration in Heliconius butterflies evolved as a warning signal but also functions as a mating cue. We investigated the contribution of visual preference to assortative mating in an aposematic butterfly Heliconius cydno in the context of reproductive isolation with its sympatric, visually distinct relative Heliconius melpomene. Heliconius cydno have conspicuous white bands on their forewings, whereas those of H. melpomene are red in colour. We predicted that both sexes of H. cydno contributed to assortative mating by exhibiting visual preference towards conspecific wing colouration. We analysed published and new data from preference experiments, in which males were presented with conspecific and H. melpomene females. We also recorded female responses and mating outcomes in choice experiments, involving conspecific males with either the original white or artificially painted red forewing bands. Both sexes of H. cydno responded more positively towards the conspecific colouration, and males strongly preferred females of its own colours. In contrast, male colouration did not predict mating outcomes in female choice experiments. As courtships are initiated by males in butterflies, our findings suggest that female visual preference might be of secondary importance in H. cydno. Our data also suggest that the contribution of visual preference to reproductive isolation might be unequal between H. cydno and its sympatric relative H. melpomene.

Conspicuous stripes on prey capture attention and reduce attacks by foraging jumping spiders.

Many animals avoid predation using aposematic displays that pair toxic/dangerous defences with conspicuous achromatic warning patterns, such as high-contrast stripes. To understand how these prey defences work, we need to understand the decision-making of visual predators. Here we gave two species of jumping spiders (Phidippus regius and Habronattus trimaculatus) choice tests using live termites that had their back patterns manipulated using paper capes (solid white, solid black, striped). For P. regius, black and striped termites were quicker to capture attention. Yet despite this increased attention, striped termites were attacked at lower rates than either white or black. This suggests that the termite's contrast with the background elicits attention, but the internal striped body patterning reduces attacks. Results from tests with H. trimaculatus were qualitatively similar but did not meet the threshold for statistical significance. Additional exploratory analyses suggest that attention to and aversion to stripes is at least partially innate and provide further insight into how decision-making played out during trials. Because of their rich diversity (over 6500 species) that includes variation in natural history, toxin susceptibility and degree of colour vision, jumping spiders are well suited to test broad generalizations about how and why aposematic displays work.

Wearable Prophylaxis Tool for AI-Driven Identification of Early Warning Patterns of Pressure Ulcers.

As today's society ages, age-related diseases become more frequent. One very common but yet preventable disease is the development of pressure ulcers (PUs). PUs can occur if tissue is exposed to a long-lasting pressure load, e.g., lying on tissue without turning. The cure of PUs requires intensive care, especially for the elderly or people with preexisting conditions whose tissue needs longer healing times. The consequences are heavy suffering for the patient and extreme costs for the health care system. To avoid these consequences, our objective is to develop a pressure ulcer prophylaxis device. For that, we built a new sensor system able to monitor the pressure load and tissue vital signs in immediate local proximity at patient's predilection sites. In the clinical study, we found several indicators showing correlations between tissue perfusion and the risk of PU development, including strongly reduced SpO2 levels in body tissue prior to a diagnosed PU. Finally, we propose a prophylaxis system that allows for the prediction of PU developments in early stages before they become visible. This work is the first step in generating an effective system to warn patients or caregivers about developing PUs and taking appropriate preventative measures. Widespread application could reduce patient suffering and lead to substantial cost savings.

Does sexual conflict contribute to the evolution of novel warning patterns?

Why warning patterns are so diverse is an enduring evolutionary puzzle. Because predators associate particular patterns with unpleasant experiences, an individual's predation risk should decrease as the local density of its warning pattern increases, promoting pattern monomorphism. Distasteful Heliconius butterflies are known for their diversity of warning patterns. Here, we explore whether interlocus sexual conflict can contribute to their diversification. Male Heliconius use warning patterns as mating cues, but mated females may suffer costs if this leads to disturbance, favouring novel patterns. Using simulations, we show that under our model conditions drift alone is unlikely to cause pattern diversification, but that sexual conflict can assist such a process. We also find that genetic architecture influences the evolution of male preferences, which track changes in warning pattern due to sexual selection. When male attraction imposes costs on females, this affects the speed at which novel pattern alleles increase. In two experiments, females laid fewer eggs with males present. However, although males in one experiment showed less interest in females with manipulated patterns, we found no evidence that female colouration mitigates sex-specific costs. Overall, male attraction to conspecific warning patterns may impose an unrecognized cost on Heliconius females, but further work is required to determine this experimentally.

Evaluation of spatio-temporal electrocutaneous warning signals

Abstract Acoustic or visual warning signals for workers in hazardous situations might fail under loud and/or lowvisibility work situations. A warning system that uses electrocutaneous stimulation can overcome this problem. The aim of this pilot study was to find spatio-temporal stimulation patterns for appropriate electrical warning. Eight electrode pairs were attached to the upper right arm of 16 participants. The stimulation was conducted with bi-phasic rectangular pulses of 150 μs and an amplitude of up to 25 mA. Pulse intervals that generate a single pulse, pulsating, vibrating, and continuous perception as well as varying spatial patterns (e.g. alternating between electrode pairs or circumferentially around the arm) were investigated and evaluated with regard to alertness, discomfort, and urgency. The pilot study revealed that a stimulation signal that generates a vibrating perception and is applied as a circumferential signal around the arm showed the highest values of alertness and is therefore considered a potential warning pattern for future studies with larger study groups.

Balanced polymorphisms and their divergence in a Heliconius butterfly.

The evolution of mimicry in similarly defended prey is well described by the Müllerian mimicry theory, which predicts the convergence of warning patterns in order to gain the most protection from predators. However, despite this prediction, we can find great diversity of color patterns among Müllerian mimics such as Heliconius butterflies in the neotropics. Furthermore, some species have evolved the ability to maintain multiple distinct warning patterns in single populations, a phenomenon known as polymorphic mimicry. The adaptive benefit of these polymorphisms is questionable since variation from the most common warning patterns is expected to be disadvantageous as novel signals are punished by predators naive to them. In this study, we use artificial butterfly models throughout Central and South America to characterize the selective pressures maintaining polymorphic mimicry in Heliconius doris. Our results highlight the complexity of positive frequency‐dependent selection, the principal selective pressure driving convergence among Müllerian mimics, and its impacts on interspecific variation of mimetic warning coloration. We further show how this selection regime can both limit and facilitate the diversification of mimetic traits.

A revision of the minor species group in the millipede genus Nannaria Chamberlin, 1918 (Diplopoda, Polydesmida, Xystodesmidae).

Millipedes in the family Xystodesmidae (Polydesmida) are often referred to as “colorful, flat-backed millipedes” for their bright aposematic coloration and tendency to form Müllerian mimicry rings in the Appalachian region. However, there are many species of Xystodesmidae that do not display colorful warning patterns, and instead have more cryptic appearances. Perhaps for this reason, groups such as the genus Nannaria have remained understudied, despite containing a large number of undescribed species. Before his death in 2012, R. L. Hoffman worked on a revision of the genus Nannaria, and synthesized material and drawings since 1949. Here the work is continued, inferring a molecular phylogeny of the Nannariini (Nannaria + Oenomaeapulchella), and revealing two clades within the genus. One clade is named the minor species group, and the second is the wilsoni species group. This revision, using a molecular phylogenetic framework, is the basis for descriptions of 35 new species in the minor species group. A multi-gene molecular phylogeny is used to make taxonomic changes in the taxon. Eleven putative species of Nannaria are also illustrated and discussed. Additionally, detailed collection, natural history and habitat notes, distribution maps, and a key to species of the Nannariaminor species group are provided. These items are synthesized as a basis for a revision of the genus, which hopefully will aid conservation and evolutionary investigations of this cryptic and understudied group.

Light environment influences mating behaviours during the early stages of divergence in tropical butterflies.

Speciation is facilitated when traits under divergent selection also act as mating cues. Fluctuations in sensory conditions can alter signal perception independently of adaptation to the broader sensory environment, but how this fine-scale variation may constrain or promote behavioural isolation has received little attention. The warning patterns of Heliconius butterflies are under selection for aposematism and act as mating cues. Using computer vision, we extracted behavioural data from 1481 h of video footage, for 387 individuals. We show that the putative hybrid species H. heurippa and its close relative H. timareta linaresi differ in their response to divergent warning patterns, but that these differences are strengthened with increased local illuminance. Trials with live individuals reveal low-level assortative mating that is sufficiently explained by differences in visual attraction. Finally, results from hybrid butterflies are consistent with linkage between a major warning pattern gene and the corresponding behaviour, though the differences in behaviour we observe are unlikely to cause rapid reproductive isolation as predicted under a model of hybrid trait speciation. Overall, our results reveal that the contribution of ecological mating cues to reproductive isolation may depend on the immediate sensory conditions during which they are displayed to conspecifics.

Why has transparency evolved in aposematic butterflies? Insights from the largest radiation of aposematic butterflies, the Ithomiini.

Defended species are often conspicuous and this is thought to be an honest signal of defences, i.e. more toxic prey are more conspicuous. Neotropical butterflies of the large Ithomiini tribe numerically dominate communities of chemically defended butterflies and may thus drive the evolution of mimetic warning patterns. Although many species are brightly coloured, most are transparent to some degree. The evolution of transparency from a warning-coloured ancestor is puzzling as it is generally assumed to be involved in concealment. Here, we show that transparent Ithomiini species are indeed less detectable by avian predators (i.e. concealment). Surprisingly, transparent species are not any less unpalatable, and may in fact be more unpalatable than opaque species, the latter spanning a larger range of unpalatability. We put forth various hypotheses to explain the evolution of weak aposematic signals in these butterflies and other cryptic defended prey. Our study is an important step in determining the selective pressures and constraints that regulate the interaction between conspicuousness and unpalatability.

The fitness effects of a pale mutant in the aposematic seed bug Lygaeus simulans indicate pleiotropy between warning coloration and life history.

Conspicuous warning colors that signal chemical or other defenses are common in the natural world. For instance, such aposematic warning patterns of red‐and‐black or yellow‐and‐black are common among insect taxa, particularly in the order Hemiptera, often forming the basis of Batesian and/or Müllerian mimicry rings. In addition, it has been repeatedly noted that color polymorphisms or mutants that influence pigmentation can show pleiotropy with other behavioral, physiological, or life‐history traits. Here, we describe a pale mutant of the seed bug Lygaeus simulans that appeared in our laboratory population in 2012, which differs in color to the wild‐type bugs. Through multigenerational experimental crosses between wild‐type and pale mutant L. simulans, we first show that the pale phenotype segregates as a single Mendelian locus, with the pale allele being recessive to the wild type. Next, we show (a) that there is a large heterozygous advantage in terms of fecundity, (b) that pale females suffer reduced longevity, and (c) that pale males have increased body length compared to wild‐type homozygotes. Our data therefore suggest that the color locus is pleiotropic with a number of life‐history traits, opening the door for a more complete genetic analysis of aposematic coloration in this species. In addition, this phenotype will be useful as a visible genetic marker, providing a tool for investigating sperm competition and other post‐copulatory drivers of sexual selection in this species.

High warning colour polymorphism inHeliconiushybrid zone roosts

1. Communal roosting behaviour has been documented among a wide range of taxa, particularly among groups of butterflies that display warning colourations. These aggregations of conspecifics and/or other species that share mimetic warning colour patterns can have a large impact on predator learning, and thus the survival of an aposematic form. Yet there has been limited investigation of communal roosting within areas where new and diverse warning colour forms are generated, such as hybrid zones. 2. Here, roosting behaviour was examined in a Heliconius erato hybrid zone in French Guiana between races with divergent warning colourations on their wings. In this hybrid zone, native individuals with nine distinct warning colourations, as well as individuals with altered forms that are not native in the French Guiana population, were marked and observed to determine if divergently coloured individuals participated in communal roosting, and if the proportions of colour pattern forms at roosts differed from the proportions that are found in the population. 3. The results demonstrated that divergently coloured individuals of the same species, including altered, non-native forms, will readily and repeatedly participate in nocturnal communal roosting, often with extreme fidelity to specific perch locations. 4. These findings suggest that roosts composed of polymorphic warning patterns may be common in phenotypic transition zones, which could have major implications on predator training and selection dynamics in hybrid zones.

Seismic Sequence Structure and Earthquakes Triggering Patterns

Within a time distribution of magnitude values, before any mainshock some earthquake triggering patterns with several features develop, under tectonic processes’ influence, through which it is possible to early identify the preparation phase of big earthquakes. The purpose of this article was to identify and classify the warning patterns that develop before a big earthquake by considering space-time seismicity variations. The methodological approach adopted was of graphical type, based on procedures of technical analysis currently used to estimate the financial markets. In the initial phase of the study we have analyzed the seismic sequences types described in the bibliography (type 1: foreshocks-mainshock-aftershocks, type 2: mainshock-aftershock; type 3: swarm) and the main structure of the seismic cycle, within which maximum and minimum magnitude values characterize the pattern that it develops until the main event changes. Then, we assessed the position of foreshocks, mainshock and aftershocks within the seismic cycle in order to identify the warning pattern that characterized the exact time when the energy emission occurs. As to the evolution normally shown over time, we have grouped the warning patterns in 2 categories: 1) progressive earthquake pattern; 2) flash earthquake pattern. Finally, we have made a classification of the warning pattern related to the fluctuations of maximum and minimum magnitude values, compared its form with the mainshock’s focal mechanism and suggested some graphic procedures in order to estimate the mainshock magnitude value associated with each warning pattern. The results we obtained unquestionably allow a better comprehension of preparation process of a large earthquake, improving the earthquakes forecasting probability in the next future.

Designing Fatigue Warning Systems: The perspective of professional drivers

Professional drivers have been characterized as experiencing heavy fatigue resulting from long driving time in their daily work. This study aimed to explore the potential demand of Fatigue Warning Systems (FWSs) among professional drivers as a means of reducing the danger of fatigue driving and to examine their opinions regarding the design of FWSs. Six focus groups with 35 participants and a questionnaire survey with 600 respondents were conducted among Chinese truck and taxi drivers to collect qualitative and quantitative data concerning the current situation of fatigue driving and opinions regarding the design of FWSs. The results revealed that both truck and taxi drivers had a positive attitude toward FWSs, and they hoped this system could not only monitor and warn them regarding their fatigue but also somewhat relieve their fatigue before they could stop and rest. As for warning signals, participants preferred auditory warnings, as opposed to visual, vibrotactile or electric stimuli. Interestingly, it was proposed that verbal warnings involving the information regarding consequences of fatigue driving or the wishes of drivers' family members would be more effective. Additionally, different warning patterns, including graded, single and continuous warnings, were discussed in the focus group. Finally, the participants proposed many other suggestions, as well as their concerns regarding FWSs, which will provide valuable information for companies who wish to develop FWSs for professional drivers.

Discovery of a glowing millipede in California and the gradual evolution of bioluminescence in Diplopoda

The rediscovery of the Californian millipede Xystocheir bistipita surprisingly reveals that the species is bioluminescent. Using molecular phylogenetics, we show that X. bistipita is the evolutionary sister group of Motyxia, the only genus of New World bioluminescent millipedes. We demonstrate that bioluminescence originated in the group's most recent common ancestor and evolved by gradual, directional change through diversification. Because bioluminescence in Motyxia has been experimentally demonstrated to be aposematic, forewarning of the animal's cyanide-based toxins, these results are contrary to aposematic theory and empirical evidence that a warning pattern cannot evolve gradually in unpalatable prey. However, gradual evolution of a warning pattern is plausible if faint light emission served another function and was co-opted as an aposematic signal later in the diversification of the genus. Luminescence in Motyxia stem-group taxa may have initially evolved to cope with reactive oxygen stress triggered by a hot, dry environment and was repurposed for aposematism by high-elevation crown-group taxa colonizing new habitats with varying levels of predation. The discovery of bioluminescence in X. bistipita and its pivotal phylogenetic location provides insight into the independent and repeated evolution of bioluminescence across the tree of life.

Body size matters for aposematic prey during predator aversion learning

Aposematic prey advertise their toxicity to predators using conspicuous warning signals, which predators learn to use to reduce their intake of toxic prey. Like other types of prey, aposematic prey often differ in body size, both within and between species. Increasing body size can increase signal size, which make larger aposematic prey more detectable but also gives them a more effective and salient deterrent. However, increasing body size also increases the nutritional value of prey, and larger aposematic prey may make a more profitable meal to predators that are trading off the costs of eating toxins with the benefits of ingesting nutrients. We tested if body size, independent of signal size, affected predation of toxic prey as predators learn to reduce their attacks on them. European starlings (Sturnus vulgaris) learned to discriminate between defended (quinine-injected) and undefended (water-injected) mealworm prey (Tenebrio molitor) using visual signals. During this process, we found that birds attacked and ate more defended prey the larger they were. Body size does affect the probability that toxic prey are attacked and eaten, which has implications for the evolutionary dynamics of aposematism and mimicry (where species share the same warning pattern).

Issue Information

Front Cover: The bright warning patterns of the sister species H. himera (top) and H. erato (bottom) are also used as mating cues. Original images by Riccardo Papa. Photo reproduced by permission of Richard M. Merrill, Department of Zoology, University of Cambridge, Cambridge, U.K.

Divergent warning patterns contribute to assortative mating between incipient Heliconius species.

Theoretical models suggest that traits under divergent ecological selection, which also contribute to assortative mating, will facilitate speciation with gene flow. Evidence for these so-called “magic traits” now exists across a range of taxa. However, their importance during speciation will depend on the extent to which they contribute to reproductive isolation. Addressing this requires experiments to determine the exact cues involved as well as estimates of assortative mating in the wild. Heliconius butterflies are well known for their diversity of bright warning color patterns, and their amenability to experimental manipulation has provided an excellent opportunity to test their role in reproductive isolation. Here, we reveal that divergent color patterns contribute to mate recognition between the incipient species Heliconius himera and H. erato, a taxon pair for which assortative mating by color pattern has been demonstrated among wild individuals: First, we demonstrate that males are more likely to attempt to mate conspecific females; second, we show that males are more likely to approach pinned females that share their own warning pattern. These data are valuable as these taxa likely represent the early stages of speciation, but unusually also allow comparisons with rates of interbreeding between divergent ecologically relevant phenotypes measured in the wild.