Despite extensive study of poisonous and venomous organisms and the toxins they produce, a review of the literature reveals inconsistency and ambiguity in the definitions of 'poison' and 'venom'. These two terms are frequently conflated with one another, and with the more general term, 'toxin.' We therefore clarify distinctions among three major classes of toxins (biological, environmental, and anthropogenic or man-made), evaluate prior definitions of venom which differentiate it from poison, and propose more rigorous definitions for poison and venom based on differences in mechanism of delivery. We also introduce a new term, 'toxungen', thereby partitioning toxic biological secretions into three categories: poisons lacking a delivery mechanism, i.e. ingested, inhaled, or absorbed across the body surface; toxungens delivered to the body surface without an accompanying wound; and venoms, delivered to internal tissues via creation of a wound. We further propose a system to classify toxic organisms with respect to delivery mechanism (absent versus present), source (autogenous versus heterogenous), and storage of toxins (aglandular versus glandular). As examples, a frog that acquires toxins from its diet, stores the secretion within cutaneous glands, and transfers the secretion upon contact or ingestion would be heteroglandular-poisonous; an ant that produces its own toxins, stores the secretion in a gland, and sprays it for defence would be autoglandular-toxungenous; and an anemone that produces its own toxins within specialized cells that deliver the secretion via a penetrating wound would be autoaglandular-venomous. Adoption of our scheme should benefit our understanding of both proximate and ultimate causes in the evolution of these toxins.
Sexual differences in morphology, ranging from subtle to extravagant, occur commonly in many animal species. These differences can encompass overall body size (sexual size dimorphism, SSD) or the size and/or shape of specific body parts (sexual body component dimorphism, SBCD). Interacting forces of natural and sexual selection shape much of the expression of dimorphism we see, though non-adaptive processes may be involved. Differential scaling of individual features can result when selection favors either exaggerated (positive allometry) or reduced (negative allometry) size during growth. Studies of sexual dimorphism and character scaling rely on multivariate models that ideally use an unbiased reference character as an overall measure of body size. We explored several candidate reference characters in a cryptically dimorphic taxon, Hadrurus arizonensis. In this scorpion, essentially every body component among the 16 we examined could be interpreted as dimorphic, but identification of SSD and SBCD depended on which character was used as the reference (prosoma length, prosoma area, total length, principal component 1, or metasoma segment 1 width). Of these characters, discriminant function analysis suggested that metasoma segment 1 width was the most appropriate. The pattern of dimorphism in H. arizonensis mirrored that seen in other more obviously dimorphic scorpions, with static allometry trending towards isometry in most characters. Our findings are consistent with the conclusions of others that fecundity selection likely favors a larger prosoma in female scorpions, whereas sexual selection may favor other body parts being larger in males, especially the metasoma, pectines, and possibly the chela. For this scorpion and probably most other organisms, the choice of reference character profoundly affects interpretations of SSD, SBCD, and allometry. Thus, researchers need to broaden their consideration of an appropriate reference and exercise caution in interpreting findings. We highly recommend use of discriminant function analysis to identify the least-biased reference character.
Understanding the behaviors by which animals deploy their venoms has been largely neglected compared to other aspects of the evolution and biology of venomous organisms and their venoms. Yet, behavior has long been recognized as a pacemaker for the evolution of morphological, ecological, life history, and other traits, in large part because behavioral responses can expose organisms to or protect them from novel selection pressures. The importance of behavior is especially evident in that venom most often functions through a behavioral act that generates a wound in a target animal through which the toxic secretion must be introduced. As a limited and costly commodity, venom should be deployed strategically and judiciously by those animals that possess it. The chapter summarizes the major aspects of adaptive venom use in animals, and highlights the best documented examples of strategic venom deployment among spiders. These animals, like other venomous taxa, exhibit four major behavioral strategies. First, they are often highly selective when using their venom, discharging it only under certain conditions. Second, they can modulate the quantity of venom they expend in both predatory and defensive contexts, delivering multiple bites or variable quantities within individual doses. Third, at least one study suggests that spiders possess venom gland heterogeneity and therefore deliver varying venom composition with successive venom expulsions. Finally, some evidence suggests that spiders can strategically target the delivery of their weapon at a particularly vulnerable region of their target. Collectively, the evidence suggests a common theme among spiders and other venomous animals for economization and optimization of venom deployment.
This study presents evidence of pre-mortem traumatic injury and its sequalae on multiple Edmontosaurus annectens skeletal elements recovered from a largely monodominant Cretaceous (Maastrichtian) bonebed. The sample consists of 3013 specimens excavated and prepared from two quarries, of which 96 elements manifest one or more macroscopic bone abnormalities and 55 specimens display pathology attributable to physical trauma. Evidence of traumatic pathology is strongly associated (P < .05) with body region, occurring disproportionately in the caudal vertebrae. Pre-mortem fractures with subsequent bone remodeling and hypertrophic ossification of caudal neural spines are present principally in the middle and mid-distal regions of the tail, while fractures of the vertebral centra are present primarily in the distal tail region. Other skeletal regions, such as chevrons, phalanges of the manus and ribs display unambiguous evidence of healed trauma, but with less frequency than the tail. These findings, in combination with current understanding of hadrosaurian tail biomechanics, indicate that intervertebral flexibility within the middle and mid-distal region of the tail likely rendered these caudal vertebrae more susceptible to the deleterious effects of repeated mechanical stress and subsequent trauma, potentially accompanying running locomotion and other high-impact herd interactions. Healed fractures within the region are also suggestive of accumulated injuries due to a combination of tail usage in defense and possibly accidental bumping/trampling associated with gregarious behavior.
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