The evolution of prey‐wrapping behaviour in spiders

We traced the evolution of silk use by spiders in attacks on prey by combining previous publications with new observations of 31 species in 16 families. Two new prey‐wrapping techniques are described. One, in which the spider holds a tense line (often covered with viscid silk) with both legs IV and applies it to the prey with a simultaneous movement of both legs, may be a synapomorphy linking Theridiidae, Nesticidae, and Synotaxidae. The other, in which the spider stands over the prey and turns in place, is apparently very ancient; it occurs in Theraphosidae, Tengellidae, and Agelenidae. The use of legs IV to wrap prey is described for the first time in Filistatidae and Scytodidae. Using a recent phylogeny of spiders, we propose that prey wrapping with legs IV has evolved convergently at least four times. We propose that prey wrapping originally evolved from egg‐sac construction behaviour.     

Littorina neglecta Bean: ecotype or species?

The taxonomic status of the littoral prosobranch Littorina neglecta has been investigated using animals from the northeast coast of England. Rough periwinkles were collected from barnacle-covered rock platforms near the northern (Ness Point) and southern (Old Peak) ends of Robin Hood's Bay, and from small and large boulders at the southern end. Shape was analysed in shells from all sites, and the activity levels and heat stability of aspartate aminotransferase (Aat) and alanine aminotransferase (Alat) were determined in samples from Peak Steel and a nearby boulder habitat. Of the shells from Ness Point, 98% could be separated visually, using colour, sculpturing and banding, into three categories (the oviparous L. arcana, and the ovoviviparous L. saxatilis and L. neglecta). The categories were confirmed using discriminant analysis on shell measurements and cross-validation indicates at least 70% accuracy. On Peak Steel L. saxatilis breeds at a small size and its shells are visually quite distinct from those of Peak Steel L. neglecta. In discriminant analysis 84% of the latter classify with the Ness Point L. neglecta. The enzyme studies clearly indicate that the activity level of Aat after 1 min exposure to 56°C is significantly different between Peak Steel L. saxatilis and L. neglecta. In general, it appears that there are different physiological responses in the different groups of animals depending on the zone, microhabitat and taxonomic category. The relationship between the 2 barnacle-dwelling, brooding periwinkles is discussed. It is considered that the existence of two distinct, sympatric ecotypes is unlikely, and that there is a true barnacle-dwelling ecotype of L. saxatilis that is morphologically and biochemically distinct from L. neglecta. The possibility of gene flow between these forms is currently being investigated.     

Subspeciation or none? The hardun in the Aegean (Reptilia: Sauria: Agamidae: Laudakia stellio)

A study aimed at testing the contested validity of the subspecies Laudakia stellio daani yielded novel insights into the essence of subspecies. We examined morphologically museum specimens from Greece, Aegean islands, and Anatolia (n?=?118; not all could be used in all analyses). Beyond the conventional mensural, meristic and qualitative characters we quantified 14 coloration characters, thus totalling 34 characters (including sex). Biometry was statistically analysed within and between the two geographically defined presumed subspecies, L. s. daani and L. s. stellio. Excluding or including broken‐tailed specimens changed the outcome of tests. Significant minor directional asymmetry occurred in one of four character‐taxon combinations. Phenetic cluster analysis poorly separated the two presumed subspecies when all characters, including those with discordant variation, were included; after selection of characters, the separation improved. Some biometric characters distinguish the two presumed subspecies, confirming their validity. The associations of significant inter‐character correlations differed between the two subspecies. Additionally, the two differed in parameters reflecting selection pressures and social structure: L. s. stellio is more colourful than L. s. daani; its sexual dimorphism is mainly chromatic, versus mensural in L. s. daani; and its population seems to include many males with underdeveloped callous scales, presumably socially subordinate, versus very few in L. s. daani.     

The first catenicellid (Bryozoa,Ascophora) from Mediterranean shallow waters: a hidden resident or a new immigrant?

Catenicella paradoxa sp. nov. is described from Mediterranean shallow bottoms. Colonies are small, rooted, erect and articulated, each segment either sterile unizooidal or bizooidal, or fertile bizooidal. Basal parts consist of an “articulated basal complex”, sometimes related to thick encrusting tubes, invariably followed by morphologically ordinary zooids belonging to each type of segment. The hypotheses of small separate colonies or subcolonies from a network of encrusting tubes are discussed, but relationships with the substratum remain unclear. The species represents the first known Mediterranean catenicellid, a family taxon with a warm, mostly Australasian present-day distribution. Nevertheless, catenicellids and Catenicella species have a long history in the European area from the Middle Eocene to the Early Pleistocene, with at least four different species. In this context, the meaning of C. paradoxa sp. nov. is discussed, bearing in mind problems of species spreading in relation to global warming and human activities.     

A putative hybrid swarm within O?nopsis foliosa (Asteraceae: Astereae)

O?nopsis foliosa var. foliosa and var. monocephala are endemic to short-grass steppe of southeastern Colorado and until recently were considered geographically disjunct. The only known qualitative feature separating these 2 varieties is floral head type; var. foliosa has radiate heads, whereas var. monocephala heads are discoid. Sympatry between these varieties is restricted to a small area in which a range of parental types and intermediate head morphologies is observed. We used distribution mapping, morphometric analyses, chromosome cytology, and pollen stainability to characterize the sympatric zone. Morphometrics confirms that the only discrete difference between var. foliosa and var. monocephala is radiate versus discoid heads, respectively. The outer florets of putative hybrid individuals ranged from conspicuously elongated yet radially symmetric disc-floret corollas, to elongated radially asymmetric bilabiate- or deeply cleft corollas, to stunted ray florets with appendages remnant of corolla lobes. Chromosome cytology of pollen mother cells from both putative parental varieties and a series of intermediate morphological types collected at the sympatric zone reveal evidence of translocation heterozygosity. Pollen stainability shows no significant differences in viability between the parental varieties and putative hybrids. The restricted distribution of putative hybrids to a narrow zone of sympatry between the parental types and the presence of meiotic chromosome-pairing anomalies in these intermediate plants are consistent with a hybrid origin. The high stainability of putative-hybrid pollen adds to a growing body of evidence that hybrids are not universally unfit.     

An unusual song-pattern in the Chorthippus mollis group (Orthoptera: Acrididae): local variant or hybrid population?

Summary

Published accounts of geographical variation in the songs of Orthoptera are briefly reviewed. An unusual song-pattern produced by an isolated alpine population of a grasshopper belonging to the Chorthippus mollis group is described and illustrated with oscillograms. Both the song and morphological characters of this population are compared with those of typical C. mollis and C. biguttulus. The status of the population is discussed and it is concluded that the balance of evidence favours the hypothesis that it is a hybrid population resulting from a cross between these two species.     

The Eucerotinae: a Gondwanan origin for a cosmopolitan group of Ichneumonidae?

A new genus and species of the formerly monobasic and predominantly amphitropical, ichneumonid subfamily, Eucerotinae, is described from Chile. A phylogenetic analysis of the redefined Eucerotinae is undertaken. Barronia araucaria gen. et sp. n. has a sister-group relationship with Euceros. The southern hemisphere species of the latter genus were found to comprise a monophyletic clade, the sister group to all other Euceros species. Similarly all north temperate Euceros species form a monophyletic clade, the sister group of which occurs in Madagascar. Within the former clade, the North American species form a monophyletic group, the sister species of which is Palaearctic. Several other groups of Palaearctic taxa have a sister-group relationship to this lineage. These results suggest the Eucerotinae had an ancient origin on Gondwanaland in the lower Cretaceous, prior to the separation of the Indo-Madagascar terrane from the Australian-Antarctic-South American tectonic plate. We hypothesize the Palaearctic fauna colonized Eurasia following the impaction of India on to its southern margin, and that the North American fauna is derived from a single circum-polar dispersal from Eurasia. Our results do not support an earlier hypothesis that the Eucerotinae arose as hyperparasitoids of Symphyta. Evidence suggests the group arose as parasitoids of ophioniform ichneumonids attacking lepidopterous larvae feeding on gymnosperms or anemophilous angiosperm trees.     

Fly parasitism in Papuan frogs,with a discussion of ecological factors influencing evolution of life‐history differences

Dipteran parasitism of frogs in the Papuan region is described and aspects of the ecological interaction are detailed. The fly involved, Batrachomyia krausi, is the largest known member of an otherwise Australian genus. Larvae of this species develop in the lateral lymph sacs of frogs of the genus Rana, feed on host blood tissues, occur with high incidence and moderate intensity, yet exhibit minimal virulence. Minimal virulence and high infestation rate seen in this host–parasite system contrast with those reported for Australian congeners. These differences are hypothesized to derive from interaction of the large size of the Papuan parasite with its phylogenetically constrained life‐history. Differences in virulence exhibited between parasitic chloropids versus calliphorids and sarcophagids are discussed as consistent with ecological predictions based on parasite transmission dynamics.     

First record of the genus Leptodactylus (Anura: Leptodactylidae) in Cuba: Leptodactylus fragilis,a biological invasion?

The Neotropical genus Leptodactylus is currently represented by three species in the West Indies (Leptodactylus albilabris, Leptodactylus fallax and Leptodactylus validus). Based on morphological, acoustic and molecular evidence, we document the presence of a fourth species in the Caribbean region, Leptodactylus fragilis (Brocchi, 1877). The species was found at two localities in western Cuba, and molecular data suggest a northern South American origin, possibly Venezuela, for these populations. We discuss the potential invasive status of L. fragilis, based on its known distribution, relative abundance, behaviour and possible impacts on native species of Cuban amphibians.     

Does nutritional status constrain adoption of more costly and less risky foraging behaviour in an Amazonian shelter-building spider?

Individual nutritional status may increase marginal foraging costs and potentially drive animal foraging strategies. Here, we investigated how it might affect foraging strategies of the shelter-building spider Hingstepeira folisecens (Hingston 1932) (Araneidae). This spider can catch prey using two strategies with different relative costs: ‘pulling’ (higher energy expenditure) and ‘pursuing’ (lower energy expenditure). We conducted experiments by offering prey at a fixed distance from the shelter entrance to investigate if spiders with lower nutritional status would exhibit riskier but less costly foraging behaviours more frequently than spiders with better status. The nutritional status did not affect the foraging behaviours adopted by H. folisecens. Additionally, the frequency of both foraging behaviours was similar. Therefore, our results did not provide any evidence that individual nutritional status is a factor driving the selection of foraging behaviours in H. folisecens. We suggest that the malnourished spiders should adopt additional strategies to compensate for the costs of pulling behaviour, especially when prey is far from the shelter. Furthermore, it is possible that other factors, like relative prey size in relation to web size, can help us to understand the threshold that determines when such individuals must adopt one of two possible behaviours.     

Foregut anatomy and predation by Charonia lampas (Gastropoda: Prosobranchia: Neotaenioglossa) attacking Ophidiaster ophidianus (Asteroidea: Ophidiasteridae) in the Açores,with a review of triton feeding behaviour

This paper is the first to describe, in detail, the process of feeding by a species of Charonia. As Charonia lampas commenced attacking its sea star prey, Ophidiaster ophidianus, the latter autotomized its trailing arm and escaped. The pursuing C. lampas ate this arm. The results of this study are not in accord with the established view of feeding by species of Charonia. Firstly, C. lampas did not inject either a venom into its prey to paralyse it nor an acid to access it and, second, nor did it pierce the skeleton of its prey and suck out the tissues, all as reported in the literature. Rather, in the case herein described, C. lampas held and manipulated its prey with the foot and used its taenioglossan radula located at the end of the extendible pleurembolic proboscis to scrape and ingest the soft tissues of the sea star and, subsequently, consume the remaining skeletal components again using the proboscis and contained radula. These two phases of consumption were followed by two matching phases of faeces production. Also described is the foregut anatomy of C. lampas, including, for the first time, the minute jaws and the histology of the salivary glands, and for which comparison is made with those of the known acid-producing structures of representatives of the Cassidae and other Ranellidae. As finally described, predator and prey clearly recognised each other signifying an identifiably intimate relationship. Charonia lampas consumed only the autotomised arm and it is argued that in such a case, as an anti-predation device, autotomy is of survival benefit to both predator and prey.     

Nest-site microhabitat association of red-billed leiothrix in subtropical fragmented forest in central China: evidence for a reverse edge effect on nest predation risk?

Previous studies of nest-site selection on a fine scale may reveal limiting resources within habitat types. The red-billed leiothrix (Leiothrix lutea Scopoli, 1786) is a common bird species that lives in the subtropical forests of Asia. Despite many reports of this species from introduced populations, little information has been obtained from its native range. From 2011 to 2013, we studied nest-site selection of red-billed leiothrix at micro-scales in Daweishan Nature Reserve, Hunan Province, central China. A total of 363 nests were found in five vegetation types. We measured the habitat variables and constructed nest-site selection models for nests found in the forest and scrub-grassland. Among the 18 variables measured in the forest, six variables were selected to construct the nest-site selection model: distance to forest edge (DTE), distance to water (DTW), vegetation comprehensive coverage, tree coverage, bamboo coverage and shrub height. According to Akaike’s information criterion, the best model consisted of five of these variables (excluding vegetation comprehensive coverage), and distance to forest edge, distance to water, tree coverage and bamboo coverage had negative effects on nest-site selection. In scrub-grassland, the DTE, DTW, and bush coverage (BUC) were selected from the 13 variables measured, and, accordingly, the best model consisted of DTE and BUC. Model averaging suggested that BUC had a positive effect on nest-site selection. In contrast, DTE has a reverse effect. In addition, DTE differed significantly between successful and failed nests in forest and scrub-grassland. More successful nests were found near the forest edge. Taken together, these findings emphasise the power of fine-scale habitat selection models in identifying relevant habitat variables with a significant effect on preferred habitat and eventually, breeding success.     

Phoretic behaviour of Attacobius attarum (Roewer, 1935) (Araneae: Corinnidae: Corinninae) dispersion not associated with predation?

Attacobius attarum spiders exhibit a phoretic behaviour on the winged sexual of Atta leaf-cutting ants during their mating flight. However, it is unclear if this behaviour is for dispersion or to facilitate the predation of ants in the new colonies. A nest of Atta sexdens was monitored on the day of the mating flight, and the winged ants, as well as the spiders, were collected. The results obtained corroborate the hypothesis that phoretic behaviour is commonly used for dispersion of the spider A. attarum, predominantly females. Of these spiders, 64 individuals of A. attarium were collected, of which 62 were females (96.9%) and two were males (3.1%). Regarding the winged leaf-cutting ants sampled, 378 females and 361 males were collected, totaling 739 individuals. Of these, 64 individuals (8.7%) had a spider attached to its back for phoretic dispersal and none was observed on the queens after the nuptial flight. In our study, A. attarum females perform phoretic dispersal into the environment on winged leaf-cutting ants but do not settle in the new nests.