Sequestered defensive toxins in tetrapod vertebrates: principles, patterns, and prospects for future studies

Chemical defenses are widespread among animals, and the compounds involved may be either synthesized from nontoxic precursors or sequestered from an environmental source. Defensive sequestration has been studied extensively among invertebrates, but relatively few examples have been documented among vertebrates. Nonetheless, the number of described cases of defensive sequestration in tetrapod vertebrates has increased recently and includes diverse lineages of amphibians and reptiles (including birds). The best-known examples involve poison frogs, but other examples include natricine snakes that sequester toxins from amphibians and two genera of insectivorous birds. Commonalities among these diverse taxa include the combination of consuming toxic prey and exhibiting some form of passive defense, such as aposematism, mimicry, or presumptive death-feigning. Some species exhibit passive sequestration, in which dietary toxins simply require an extended period of time to clear from the tissues, whereas other taxa exhibit morphological or physiological specializations that enhance the uptake, storage, and/or delivery of exogenous toxins. It remains uncertain whether any sequestered toxins of tetrapods bioaccumulate across multiple trophic levels, but multitrophic accumulation seems especially likely in cases involving consumption of phytophagous or mycophagous invertebrates and perhaps consumption of poison frogs by snakes. We predict that additional examples of defensive toxin sequestration in amphibians and reptiles will be revealed by collaborations between field biologists and natural product chemists. Candidates for future investigation include specialized predators on mites, social insects, slugs, and toxic amphibians. Comprehensive studies of the ecological, evolutionary, behavioral, and regulatory aspects of sequestration will require teams of ecologists, systematists, ethologists, physiologists, molecular biologists, and chemists. The widespread occurrence of sequestered defenses has important implications for the ecology, evolution, and conservation of amphibians and reptiles.     

Don’t get the blues: conspicuous nuptial colouration of male moor frogs (Rana arvalis) supports visual mate recognition during scramble competition in large breeding aggregations

Conspicuous male colouration is expected to have evolved primarily through selection by female choice. In what way conspicuous colours could be advantageous to males scrambling for mates remains largely unknown. The moor frog (Rana arvalis) belongs to the so-called explosive breeders in which spawning period is short; intrasexual competition is strong, and males actively search and scramble for females. During breeding, male body colouration changes from a dull brown (similar to females) to a conspicuous blue, and we wanted to test if male blueness influences mating success or facilitates male mate recognition. To do so, we first measured the colour of mated and non-mated males using a spectrophotometer. In an experiment, we then analysed interactions of actual male moor frogs in natural spawning aggregations with a brown (resembling a female or a non-breeding male) and a blue model frog. Mated and non-mated males did not differ in colouration, suggesting that female choice based on colour traits was unlikely. In our behavioural experiment, male moor frogs spent significantly more time in contact and in amplexus with the brown model than with the blue model. Our results suggest that the nuptial colouration in moor frogs can act as a new type of visual signal in anurans evolved to promote instantaneous mate recognition allowing males to quickly move between rivals while scrambling for females.     

Geographic variation in a predator-induced defense and its genetic basis

Predator-induced morphological defenses are a well-known form of phenotypic plasticity, but we continue to have a limited understanding of geographic variation in these responses and its genetic basis. Here we examine genetic variation and geographic differentiation in the inducible defenses of tadpoles (Rana pirica) in response to predatory salamander larvae (Hynobius retardatus). To do so, we crossed male and female frogs from a "mainland" Japanese island having predaceous salamanders and a more isolated island not having predaceous salamanders and raised resulting offspring in the presence and absence of H. retardatus. Mainland tadpoles exhibited a higher capacity to express the inducible morphology (a more bulgy body) than those from the predator-free island, and expression of the bulgy morph in mainland-island hybrids produced phenotypes that were intermediate to those produced by pure crosses. In addition, parental sex had no effect on expression of the bulgy morph. Our results support the hypothesis that geographic variation in inducible defenses is linked to the additive effects of autosomal alleles that are shaped by differences in historical exposure to the inducing predator.     

Conspicuous displays in cryptic males of a polytypic poison-dart frog

The evolution of aposematism is linked to increased opportunities for conspicuous sexual displays since detection by potential predators is no longer disadvantageous. Therefore, phenotypic divergence in aposematic species leading to relatively cryptic forms is expected to constrain such opportunities, by restoring the trade-off between natural and sexual selection on the boldness of sexual displays. We asked if and how a derived phenotype of the poison-dart frog Oophaga granulifera that appears relatively cryptic to potential predators exhibits conspicuous sexual displays for potential mates. We used visual modeling of frog contrasts against their natural backgrounds to test if for conspecifics green frogs appear less conspicuous than red frogs as they do for birds. We conducted behavioral observations of focal red and green males to determine if green frogs adjust their display behavior to the availability of potential mates. Dorsal brightness is known to influence female preferences in at least one poison frog species. We found that, despite being less visible under some measures, green frogs may appear as bright as red frogs for conspecifics but not birds, when viewed on dark backgrounds. Additionally, green males called more intermittently than red males when advertising to distant females, but they exhibited a dramatic increase in calling activity in proximity of a female and were as active as red males in this context. Together, our results suggest that green frogs retain context-dependent conspicuousness to conspecifics despite the evolution of relative crypsis to potential predators.     

The poison gland of the antMyrmicaria eumenoides and its role in recruitment communication

Summary The antMyrmicaria eumenoides is a significant arthropod predator. For rapid attraction of large numbers of nestmates to newly discovered food sources the ants use an efficient recruitment communication system based on the poison gland secretion. Workers exhibit age-based division of labour. Young workers perform brood-care; their poison gland reservoir develops and reaches its final size of 0.5 µl at an age of 6 weeks, when they become foragers. The secretion deposited during combat with enemies or prey is composed of equal amounts of both a high volatile and a low volatile fraction. Within the high volatile fraction (+)—limonene is the main component (97%) and is the only olfactory trigger to alert ants in the vicinity and to recruit them to places of combat, where they assemble. Ants respond to synthetic (+)—limonene in exactly the same way as to the poison gland secretion when applied at the same airborne concentrations. Further components of the high volatile fraction are four additional monoterpene hydrocarbons and hexanoic nitrile. The high volatile and the low volatile fraction of the poison gland secretion each have dual functions: The low volatiles, of which the main component is an alkaloid, serve as a fixative and extend the effective period of the limonene signal by modifying its evaporation kinetics. On the other hand the high volatile recruitment signal (+)—limonene is also the solvent for the alkaloid and enhances its spreading on the surface of the cuticle of arthropod enemies or prey.     

Introduced trout sever trophic connections in watersheds: consequences for a declining amphibian

Trophic linkages between terrestrial and aquatic ecosystems are increasingly recognized as important yet poorly known features of food webs. Here we describe research to understand the dynamics of lake food webs in relation to a native riparian amphibian and its interaction with introduced trout. The mountain yellow-legged frog Rana muscosa is endemic to alpine watersheds of the Sierra Nevada Mountains and the Transverse Ranges of California, but it has declined to a small fraction of its historical distribution and abundance. Although remaining frogs and introduced trout feed in different habitats of alpine lakes, our stable-isotope analyses clearly show that the same resource base of benthic invertebrates sustains their growth. During one period, insect emergence from naturally fishless lakes was nearly 20-fold higher compared to adjacent lakes with trout, showing that fish reduce availability of aquatic prey to amphibious and terrestrial consumers. Although trout cannot prey on adult frogs due to gape limitation, foraging post-metamorphic frogs are 10 times more abundant in the absence of trout, suggesting an important role for competition for prey by trout in highly unproductive alpine watersheds. Most Sierran lakes contain fish, and those that do not are usually small isolated ponds; in our study, these two lake types supported the lowest densities of post-metamorphic frogs, and these frogs were less reliant on local, benthic sources of productivity. Since Rana muscosa was formerly the most abundant vertebrate in the Sierra Nevada, the reduction in energy flow from lake benthos to this consumer due to fish introductions may have had negative consequences for its numerous terrestrial predators, many of which have also declined. We suggest that disruptions of trophic connections between aquatic and terrestrial food webs are an important but poorly understood consequence of fish introduction to many thousands of montane lakes and streams worldwide and may contribute to declines of native consumers in riparian habitats.     

Threat-sensitive generalization of predator recognition by larval amphibians

Despite the importance of acquired predator recognition in mediating predator–prey interactions, we know little about the specific characteristics that prey use to distinguish predators from non-predators. Recent experiments with mammals and fish indicate that some prey lacking innate predator recognition have the ability to display anti-predator responses upon their first encounter with those predators if they are similar to predators that the prey has recently learned to recognize. This phenomenon is referred to as generalization of predator recognition. In this experiment, we documented for the first time that larval amphibians (woodfrog, Rana sylvatica) have the ability to generalize the recognition of known predators to closely related novel predators. Moreover, we demonstrated that this ability is dependent on the level of risk associated with the known predator. When red-bellied newt, Cynops pyrrhogaster (known predator), was paired with simulated low risk, tadpoles displayed fright responses to newts and novel tiger salamanders, Ambystoma tigrinum, but not to novel African clawed frogs, Xenopus laevis. However, when the newt was paired with simulated high risk, tadpoles generalized their responses to both tiger salamanders and African clawed frogs. Larval anurans seem to have a wider generalization frame than other animals.     

Prevalence of malformed frogs in Kaoping and Tungkang river basins of southern Taiwan

In this study we found many amphibians with bizarre appearances, known as malformations in Pingtung County southern Taiwan. For this investigation we collected frogs inhabiting the Kaoping and Tungkang river watersheds between February 2006 and June 2007. Among the total number of 10,909 normal frogs (i.e., anurans) collected during the investigation period, the Indian rice frogs (Rana limnocharis) account for the greatest number next is the Chinese bullfrog (Rana rugulosa). Of all the 244 captured malformed frogs, the Indian rice frog account for the greatest proportion. These malformed frogs have their main distribution in upstream areas of these two rivers. Our result indicates that the appearance rate of malformed frogs is 1.8% in the upstream reaches of the Kaoping River and 2.6%, and 0.8%, respectively in the upstream and midstream reaches of the Tungkang river. The most-commonly-found malformation is the lack of palms, followed by the lack of appendages, exostosis, and a malformed appendicular. It is, therefore, reasonable to speculate that the causes for the malformation may be related to the increased organic pollutants and agricultural chemicals used in the upstream reaches of these two rivers.     

Combining optimal defense theory and the evolutionary dilemma model to refine predictions regarding plant invasion

Optimal defense theory posits that plants with limited resources deploy chemical defenses based on the fitness value of different tissues and their probability of attack. However, what constitutes optimal defense depends on the identity of the herbivores involved in the interaction. Generalists, which are not tightly coevolved with their many host plants, are typically deterred by chemical defenses, while coevolved specialists are often attracted to these same chemicals. This imposes an "evolutionary dilemma" in which generalists and specialists exert opposing selection on plant investment in defense, thereby stabilizing defenses at intermediate levels. We used the natural shift in herbivore community composition that typifies many plant invasions to test a novel, combined prediction of optimal defense theory and the evolutionary dilemma model: that the within-plant distribution of defenses reflects both the value of different tissues (i.e., young vs. old leaves) and the relative importance of specialist and generalist herbivores in the community. Using populations of Verbascum thapsus exposed to ambient herbivory in its native range (where specialist and generalist chewing herbivores are prevalent) and its introduced range (where only generalist chewing herbivores are prevalent), we illustrate significant differences in the way iridoid glycosides are distributed among young and old leaves. Importantly, high-quality young leaves are 6.5x more highly defended than old leaves in the introduced range, but only 2x more highly defended in the native range. Additionally, defense levels are tracked by patterns of chewing damage, with damage restricted mostly to low-quality old leaves in the introduced range, but not the native range. Given that whole-plant investment in defense does not differ between ranges, introduced mullein may achieve increased fitness simply by optimizing its within-plant distribution of defense in the absence of certain specialist herbivores.     

Pre-encounter versus post-encounter inducible defenses in predator–prey model systems

It has been reported that, in order to reduce mortality, prey are able to change their phenotype in response to cues released from predators. These short-time responses constitute effective antipredator strategies in variable environments, and involve changes in morphology, behavior, physiology or life-history traits of prey individuals belonging to a wide spectrum of taxa. Defenses can be classified into pre-encounter and post-encounter, depending on the phase of the predation process in which they take place. Also, inducible defenses should be costly.Despite the current knowledge of inducible defenses at the individual level, our understanding of their dynamic consequences at the population and community level is limited. In this work we construct and analyze numerically a predator–prey system, parameterized from published experimental data, in which prey exhibit inducible defenses of the type pre-encounter (affecting attack rate) or post-encounter (affecting handling time) and entailing either metabolic or feeding costs. The above assumptions were analyzed over a gradient of resource availability.Our results indicated that both types of cost have a similar effect on the dynamics of the model system, but we expect that different costs will produce different outcomes in a more complex model community. Conversely, pre-encounter and post-encounter IDs define domains of attraction with different size and shape within the studied sections of the multidimensional parameter space. Roughly speaking, post-encounter IDs determine a more rich dynamics when plausible parameter values are chosen, and the effect of resource density is different if the ID is handling-time based or attack-rate based. In agreement with previous works, our analyses indicate that IDs can damp population oscillations and prevent the paradox of enrichment.     

Exposure to metals (Ca,K, Mn) and road salt (NaCl) differentially affect development and survival in two model amphibians

ABSTRACT

Human activity has led to widespread chemical alteration of natural environments. Aquatic ecosystems are especially susceptible to chemical changes, including those caused by runoff and invasive species. Here, we examined the effects of water chemistry, specifically sodium chloride as well as three metals (Ca, K, and Mn) known to differ between native and invasive wetland plant species’ leaf tissues, on the development of two model amphibians: the native Northern leopard frog, Lithobates pipiens, and the non-native African clawed frog, Xenopus laevis. We exposed frog eggs to metal treatment solutions and measured time to hatching (TTH), and following hatching, we exposed tadpoles to a lethal concentration of sodium chloride and recorded time to death (TTD). We found that increasing metal concentrations generally resulted in acceleration of TTH for Xenopus tadpoles, but had no effect on leopard frogs. However, increasing metal concentrations (Ca, Mn) increased leopard frog tadpole susceptibility to NaCl (decreased TTD), while increasing metal concentrations (Ca, K) generally resulted in decreased Xenopus tadpole susceptibility to NaCl. Overall, our data suggest that invasive amphibians may be more tolerant to chemical changes than native amphibians, including those driven by the introduction of invasive plant species.     

Effects of Introduced Salmonids on a Montane Population of Iberian Frogs

Abstract:  Amphibians are declining worldwide because of multiple factors, including human-mediated introduction of fishes into naturally fishless areas. Although several studies have focused on the effect of exotic fishes on native amphibians breeding in ponds or lakes, little is known about their effects on stream-breeding species. We studied the effects of introductions of native brown trout ( Salmo trutta ) and exotic brook trout ( Salvelinus fontinalis ) on the stream-breeding, endemic Iberian frog (  Rana iberica ) in a protected area in central Spain. We assessed occurrence patterns of tadpoles and salmonids and compared habitat use of the three species. We also determined experimentally whether chemical cues from salmonids elicited antipredator behavior in tadpoles. Finally, we assessed the relative influence of tadpole habitat preferences, differences in salmonid species, and invasion geography on tadpole occurrence. Despite widely overlapping habitat preferences, tadpoles and trout did not coexist, with the former restricted to fishless habitats. Tadpoles detected chemical cues from both trout species and reacted by decreasing their activity, although the response toward the native brown trout was stronger. The residual distribution of Iberian frogs in Peñalara is better explained by the geography of fish invasions than by the fish species involved. Measures such as fish extirpation from certain areas, aimed at recovering lost habitat and improving connectivity among remaining populations of Iberian frogs, seem critical for the species' long-term survival in central Spain.     

Jasmonic acid treatment and mammalian herbivory differentially affect chemical defenses and growth of wild mustard (Brassica kaber)

Summary. Jasmonic acid (JA) is a wound-related hormone found in most plants that, when applied exogenously, can induce increases in levels of chemical defenses in patterns similar to those induced by mechanical damage or insect feeding. Relative to responses to insect and pathogen attack, chemical responses of herbaceous plants to mammalian herbivore attack have been little studied. In a field experiment, we compared the effects of JA treatment and naturally occurring mammalian herbivory on the expression of trypsin inhibitors, glucosinolates, peroxidase activity and growth of wild mustard (Brassica kaber). Exogenous JA significantly increased trypsin inhibitor activity and glucosinolate concentration, and moderately increased peroxidase activity in the eighth true leaves of five-week-old plants, relative to untreated controls. In contrast, levels of these chemical defenses in the eighth true leaves or in regrowth foliage of plants that had ∼80% of their leaf area removed by groundhogs (Marmota monax) did not differ from that in undamaged and untreated controls. Although exogenous JA significantly elevated levels of chemical defenses, it did not affect height of plants through the season and only slightly reduced time to first flower. Groundhog herbivory significantly reduced height and delayed or abolished flowering, but these effects were not substantial unless coupled with apical meristem removal. We hypothesize that the lack of effect of groundhog herbivory on chemical defenses may be due in part to the speed and pattern of leaf area removal by groundhogs, or physiological constraints caused by leaf area loss. Despite having no effect on chemical defense production, leaf area loss by groundhogs was more costly to growth and fitness than the effects of JA application in this study, but only substantially so if coupled with apical meristem removal. We suggest that in general, costs of defense production in plants are likely to be minimal when compared to the risk of losing large amounts of leaf area or primary meristematic tissue. Thus, if they are effective at deterring herbivory, the benefits of inducible defense production likely outweigh the costs in most cases. Received 20 December 2000; accepted 3 May 2001     

Pattern of lionfish envenomations reported to Texas poison control centers, 1998–2006

The purpose of this study was to describe a large number of lionfish envenomations reported to poison control centers. Cases were lionfish envenomations reported to Texas poison control centers during 1998–2006. The distribution of cases was determined for selected parameters. A total of 188 cases were identified. Of the cases with a known location of envenomation, 94% involved the finger or hand. The patient was male in 72% of the cases, and the mean age was 32 years (range 4–65 years). The most frequently reported adverse clinical effects were dermal pain (89%) and a puncture or wound (66%). The most commonly reported treatments were immersion in hot water (84%) and tetanus antitoxin/status (48%). Patients with lionfish envenomations reported to Texas poison control centers were most often adults and males. The reported adverse clinical effects were typical of lionfish envenomations, and the treatments were generally consistent with the recommended guidelines.     

Seaweed sex pheromones and their degradation products frequently suppress amphipod feeding but rarely suppress sea urchin feeding

Summary. A diverse group of brown seaweeds produce bouquets of C₁₁ metabolites, some of which act as pheromones that cue gamete release or attract sperm to eggs following release. We demonstrate that these C₁₁ metabolites and their degradation products also frequently and strongly deter feeding by the herbivorous amphipod Ampithoe longimana, but rarely by the herbivorous sea urchin Arbacia punctulata. Across the range of concentrations tested, seven of twelve C₁₁ metabolites or mixtures that we tested deterred feeding by the amphipod, but only two of eleven deterred the sea urchin. For those compounds where we could rigorously contrast the magnitude of deterrence against the amphipod with the magnitude of deterrence against the urchin, the amphipod was deterred significantly more than the urchin by five of six metabolites. Thus, C₁₁ compounds were more frequently and more strongly deterrent to the amphipod than to the sea urchin. These findings for C₁₁ metabolites conflict with previous investigations, where other classes of seaweed chemical defenses have been shown to deter feeding by large mobile herbivores like urchins and fishes but to be relatively ineffective against mesograzers, especially the species of amphipod that we used here. Our results suggest that C₁₁ metabolites are unusual among the known seaweed chemical defenses in that they are especially effective against mesograzers, which often consume seaweed spores, zygotes, and juveniles. The high concentrations of C₁₁ metabolites in brown algal eggs could allow these defenses to be especially important in defending gametes, zygotes, or young sporelings from herbivorous mesograzers. Received 26 February 1998; accepted 9 April 1998.     

两种光照生境下4种常绿阔叶树的单位叶面积干重、光合能力与化学防御物质含量比较

亚热带常绿阔叶林下分布着可塑性强的耐荫植物,其叶片应对外界干扰的防御对策至少有物理性防御和化学性防御两种,这两种防御代价在一定的昆虫取食压力下在理论上应该存在权衡.为检验这种权衡是否普遍存在,分别测定了重庆缙云山4个常绿物种——光叶山矾(Symplocos lancifolia)、四川山矾(S.setchuensis)、四川毛蕊茶(Camellialawii)和细枝柃(Eurya loquaiana)在林窗和林下两种对照光环境下个体叶片的光饱和下的净光合速率、叶面积干重和部分化学防御物质含量.结果表明,4个物种生活在林窗的个体单位叶面积干重、叶片氮含量和单位面积净光合速率通常高于林下的个体,但是单位质量和单位面积的总酚含量和C/N值却低于林下的个体.蛋白质和非结构性碳水化合物含量在不同生境中没有显著差异.因为不同物种叶片对光照条件的响应不同,不同参数的变化在两种生境间的差异并不完全一致.本研究结果在一定程度上暗示了物种在林窗和林下分别倾向于采取物理和化学性防御对策,并初步证实了两种防御强度之间存在权衡关系.图4参40     

Assessing the antipredatory defensive strategies of Caribbean non-scleractinian zoantharians (Cnidaria): is the sting the only thing?

The relative importance of chemical, nematocyst, and nutritional defenses was examined for 18 species of Caribbean sea anemones (actinarians), zoanthids, and mushroom polyps (corallimorpharians) from the Florida Keys and the Bahamas Islands, 2008–2010. Feeding assays were performed using the fish Thalassoma bifasciatum with artificial foods containing crude organic extracts of cnidarian tissues. A novel behavioral assay using brine shrimp nauplii was used to assess nematocyst defenses. The nutritional quality of cnidarian tissues was examined using bomb calorimetry and soluble protein assays. In general, actinarians invested in nematocyst defenses, zoanthids in either nematocyst or chemical defenses, and corallimorpharians lacked both, except for 1 of 3 species that was chemically defended. Relative to other coral reef invertebrates, cnidarian tissues had similar caloric values but lower soluble protein concentrations. Trade-offs between chemical and nematocyst defenses were observed for 65% of species, while habitat and behavior provided a likely explanation for undefended species.     

Conservation of Deep Pelagic Biodiversity

Abstract:  The deep ocean is home to the largest ecosystems on our planet. This vast realm contains what may be the greatest number of animal species, the greatest biomass, and the greatest number of individual organisms in the living world. Humans have explored the deep ocean for about 150 years, and most of what is known is based on studies of the deep seafloor. In contrast, the water column above the deep seabed comprises more than 90% of the living space, yet less than 1% of this biome has been explored. The deep pelagic biota is the largest and least-known major faunal group on Earth despite its obvious importance at the global scale. Pelagic species represent an incomparable reservoir of biodiversity. Although we have yet to discover and describe the majority of these species, the threats to their continued existence are numerous and growing. Conserving deep pelagic biodiversity is a problem of global proportions that has never been addressed comprehensively. The potential effects of these threats include the extensive restructuring of entire ecosystems, changes in the geographical ranges of many species, large-scale elimination of taxa, and a decline in biodiversity at all scales. This review provides an initial framework of threat assessment for confronting the challenge of conserving deep pelagic biodiversity; and it outlines the need for baseline surveys and protected areas as preliminary policy goals.     

Constitutive and induced defenses to herbivory in above- and belowground plant tissues

A recent surge in attention devoted to the ecology of soil biota has prompted interest in quantifying similarities and differences between interactions occurring in above- and belowground communities. Furthermore, linkages that interconnect the dynamics of these two spatially distinct ecosystems are increasingly documented. We use a similar approach in the context of understanding plant defenses to herbivory, including how they are allocated between leaves and roots (constitutive defenses), and potential cross-system linkages (induced defenses). To explore these issues we utilized three different empirical approaches. First, we manipulated foliar and root herbivory on tobacco (Nicotiana tabacum) and measured changes in the secondary chemistry of above- and belowground tissues. Second, we reviewed published studies that compared levels of secondary chemistry between leaves and roots to determine how plants distribute putative defense chemicals across the above- and belowground systems. Last, we used meta-analysis to quantify the impact of induced responses across plant tissue types. In the tobacco system, leaf-chewing insects strongly induced higher levels of secondary metabolites in leaves but had no impact on root chemistry. Nematode root herbivores, however, elicited changes in both leaves and roots. Virtually all secondary chemicals measured were elevated in nematode-induced galls, whereas the impact of root herbivory on foliar chemistry was highly variable and depended on where chemicals were produced within the plant. Importantly, nematodes interfered with aboveground metabolites that have biosynthetic sites located in roots (e.g., nicotine) but had the opposite effect (i.e., nematodes elevated foliar expression) on chemicals produced in shoots (e.g., phenolics and terpenoids). Results from our literature review suggest that, overall, constitutive defense levels are extremely similar when comparing leaves with roots, although certain chemical classes (e.g., alkaloids, glucosinolates) are differentially allocated between above- and belowground parts. Based on a meta-analysis of induced defense studies we conclude that: (1) foliar induction generates strong responses in leaves, but much weaker responses in roots, and (2) root induction elicits responses of equal magnitude in both leaves and roots. We discuss the importance of this asymmetry and the paradox of cross-system induction in relation to optimal defense theory and interactions between above- and belowground herbivory.     

Complex chemical communication in the crazy ant <Emphasis Type="Italic">Paratrechina longicornis</Emphasis> Latreille (Hymenoptera: Formicidae)

Summary. The formicine ant Paratrechina longicornis is known for its extremely opportunistic foraging behaviour. Only a single trail pheromone source, the rectum, was previously described from this ant. Our detailed examination of this ant’s chemical communication system revealed the presence of at least four sources of pheromones. Rectum, poison sac, and Dufour gland contain orientation components with decreasing effectiveness and persistence (in the sequence mentioned) as well as attractants with increasing effectiveness. Furthermore, the mandibular gland contains repellents, and a releaser of defensive behaviour. This set of various signals of different strength and persistence allows an extraordinary degree of flexibility and efficiency in the collective behaviour of P. longicornis, especially food exploitation, and thus may contribute to this insect’s overall ecological success.