Webs of theridiid spiders: construction, structure and evolution

Understanding the web construction behaviour of theridiid (comb-footed) spiders is fundamental to formulating specific evolutionary hypotheses and predictions regarding the reduction of orb-webs. We describe for the first time in detail the web construction behaviour of Achaearanea tepidariorum , Latrodectus geometricus , Theridion sisyphium and T. varians as well as webs of a range of other theridiids. In our survey we distinguish four major web types. Among webs with gumfooted lines, we distinguish between webs with a central retreat ( Achaearanea -type) and those with a peripheral retreat ( Latrodectus -type). Among webs without gumfooted lines, we distinguish between those which contain viscid silk ( Theridion -type) and those with a sheet-like structure, which do not ( Coleosoma -type). Theridiid gumfoot-webs consist of frame lines that anchor them to surroundings and support threads which possess viscid silk. Building of gumfooted lines constitutes a unique stereotyped behaviour and is most probably homologous for Nesticidae and Theridiidae. Webs remained in place for extended periods and were expanded and repaired, but no regular pattern of replacement was observed. We suggest that the cost of producing and maintaining viscid silk might have led to web reduction, at least in theridiids.  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 78 , 293−305.     

Social behaviour in a web-building lynx spider, Tapinillus sp. (Araneae: Oxyopidae)

Social behaviour involving cooperative prey capture and communal feeding is reported for the first time in the spider family Oxyopidae (lynx spiders), in a web-building species of the genus Tapinillus. This social spider inhabits communal webs that may contain several dozen individuals, including adults of both sexes and juveniles of different cohorts. Its colonies occur in clusters and appear to be long-lived, much like those of non-territorial permanently social species such as Anelosimus eximius (Theridiidae) or Agelena consociata (Agelenidae). However, unlike colonies of these other cooperative spiders, the colonies of the social Tapinillus do not have highly female-biased sex ratios. The possible explanations for this difference are discussed.     

Effect of abiotic factors on the foraging strategy of the orb‐web spider Argiope keyserlingi (Araneae: Araneidae)

Abstract Environmental conditions such as light level, background contrast and temperature might influence a spider's prey capture success and risk of predation. Thus it may often be advantageous for spiders to adjust web‐building behaviour in response to variation in these environmental conditions. This hypothesis was examined in a study of the construction of webs and web decorations (conspicuous strands of silk at the hub of the web) of the orb‐web spider Argiope keyserlingi. Web decorations are thought to have one or more separate functions. They may attract prey, deter predators or advertise the web to oncoming birds, thus preventing web damage. In this series of experiments, relationships between weather parameters and the construction of webs and web decorations were considered. In complementary laboratory experiments, A. keyserlingi spiders were exposed to two different light levels (700 and 90 lx), background contrasts (black and white) and temperature conditions (20 and 26°C). Of the available weather parameters, only temperature was significantly related to web decorating behaviour but not to web size. In the laboratory, temperature also influenced web‐decorating behaviour, and spiders in dim light (700 lx) constructed larger webs and longer decorations. Background contrast did not significantly alter web size or web decorations. These data suggest that when prey availability is reduced at low temperatures, spiders may use web decorations to attract prey to the web. Similarly, in dim light, spiders may build more and larger decorations to increase the visual signal to approaching prey or to advertise the web to oncoming birds.     

Memory of distances and directions moved as cues during temporary spiral construction in the spiderLeucauge mariana (Araneae: Araneidae)

The spider Leucauge marianaprobably uses the presence of temporary spiral (TSP) lines already in place to determine sites of attachment of currently produced loops of temporary spiral, but less rigidly than has been previously supposed for orb weavers. Memory of distances and directions traveled recently to and from TSP lines is implicated by the fact that adjustments to experimental and natural discontinuities in previous TSP lines occur gradually rather than being abrupt. Distances and directions traveled along both radii and previous TSP lines correlate with relative amounts of adjustment. Body size may also be used as a reference measure, but not in the simple, inflexible way suggested by R. W. G. Hingston (A Naturalist in Himalaya,Small, Maynard, Boston, 1920). Tensions on radii are not used as cues in any simple way, and may not be used at all, since experimental changes in tensions produced effects consistent with resulting changes in thread positions but inconsistent with tension differences.     

Extraordinary web and silk properties of Cyrtarachne (Araneae, Araneidae): a possible link between orb-webs and bolas

Cyrtarachne is an orb-weaving spider of the sub-family Cyrtarachninae (Araneidae), which includes the triangle-web-building Pasilobus and the bolas spiders. We found that web and thread characteristics of Cyrtarachne differed greatly from those of typical orb-webs. Web diameter, sticky spiral spacing, breaking strength and stickiness of thread, thread diameter and droplet diameter were significantly different from those of other members of Araneidae. It is especially worth noting that the diameter was approximately four times, and the breaking strength seven to ten times larger in Cyrtarachne viscid threads than in those of other araneids. Kinetic energy-absorbing ability of Cyrtarachne threads was much greater than in that of other species, and close to the amount of kinetic energy generated by flying moths. Viscid material of threads was peculiar because its adhesiveness decreased to zero in a few hours. Moreover, SEM photos revealed them to be covered with thin scales of material, while threads of other araneids were smooth. These two facts suggest that the viscid material of Cyrtarachne threads may be different from those of other orb-weavers. As web-building, hunting behaviour and prey composition of different species of Cyrtarachninae arc quite similar to each other, we hypothesize that these extraordinary web and thread characteristics of Cyrtarachne are shared by the other members of this sub-family. Because these characteristics differ in many ways from those of typical araneid orb-webs, there appears to have been a great leap in evolution between Cyrtarachne and the other Araneidae.     

Predatory and nesting behaviour of Cocalus gibbosus,a spartaeine jumping spider (Araneae: Salticidae) from Queensland

Abstract

Cocalus gibbosus was studied in the field in Queensland and in the laboratory. This is the first behavioural study of a species from the spartaeine genus Cocalus. C. gibbosus often omitted elements which are usually present in the predatory sequences of typical salticids and tended to lunge at prey from close range rather than leap from afar. Experiments showed that C. gibbosus prefers moths to other prey. In nature, C. gibbosus moulted and oviposited on silk sheets spun against tree trunks, and in the laboratory on sides of cages or blocks of wood, but this species never built an enclosing nest like typical salticids nor a large prey-catching web like some other spartaeines. C. gibbosus stalked across alien webs to catch spiders and insects, but it did not make vibratory signals. It did not stick to cribellate or ecribellate glue on alien webs. The behaviour of C. gibbosus is compared to that of other spartaeine salticids.     

Comparative biology of Portia africana,P. albimana,P. fimbriata,P. labiata,and P. shultzi,araneophagic, web-building jumping spiders (Araneae: Salticidae): Utilisation of webs,predatory versatility,and intraspecific interactions

Abstract

Portia is a behaviourally complex and aberrant salticid genus. The genus is of unusual importance because it is morphologically primitive. Five species were studied in nature (Australia, Kenya, Malaysia, Sri Lanka) and in the laboratory in an effort to clarify the origins of the salticids and of their unique, complex eyes. All the species of Portia studied were both web builders and cursorial. Portia was also an araneophagic web invader, and it was a highly effective predator on diverse types of alien webs. Portia was an aggressive mimic, using a complex repertoire of vibratory behaviour to deceive the host spiders on which it fed. The venom of Portia was unusually potent to other spiders; its easily autotomised legs may have helped Portia escape if attacked by its frequently dangerous prey. Portia was also kleptoparasitic and oophagic when occupying alien webs. P. fimbriata from Queensland, where cursorial salticids were superabundant, used a unique manner of stalking and capturing other salticids. The display repertoires used during intraspecific interactions were complex and varied between species. Both visual (typical of other salticids) and vibratory (typical of other web spiders) displays were used. Portia copulated both on and away from webs and frequently with the female hanging from a dragline. Males cohabited with subadult females on webs, mating after the female matured. Adult and subadult females sometimes used specialised predatory attacks against courting or mating males. Sperm induction in Portia was similar to that in other cursorial spiders. Portia mimicked detritus in shape and colour, and its slow, mechanical locomotion preserved concealment. Portia occasionally used a special defensive behaviour (wild leaping) if disturbed by a potential predator. Two types of webs were spun by all species (Type 1, small resting platforms; Type 2, large prey-capture webs). Two types of egg sacs were made, both of which were highly aberrant for a salticid. Responses of different species and both sexes of Portia were quantitatively compared for different types of prey. Many of the trends in behaviour within the genus, including quantitative differences in predatory behaviour, seemed to be related to differences in the effectiveness of the cryptic morphology of Portia in concealing the spider in its natural habitat (‘effective crypsis’). The results of the study supported, in general, Jackson & Blest’s (1982a) hypothesis of salticid evolution which, in part, proposes that salticid ancestors were web builders with poorly developed vision and that acute vision evolved in conjunction with the ancestral spiders becoming proficient as araneophagic invaders of diverse types of webs.     

Spinning activity of the spider Trogloneta granulum (Araneae, Mysmenidae): web, cocoon, cocoon handling behaviour, draglines and attachment discs

The spider species Trogloneta granulum, which in the wild lives inside scree slopes, builds three-dimensional orb webs. During egg-laying and egg sac building, the females stay with their dorsa down at the central part of the web. In this process, the hub is used as a platform. The threads of the hub are not incorporated into the silk cover of the egg sac. The silk wall of the egg sac is very thin, with all the silken threads constituting it having a uniform ultrastructure. The silk wall of the egg sac and the spinnerets are permanently linked by a dragline. Draglines produced by T. granulum affect the direction of movements of the female carrying its cocoon. Egg sacs are handled using draglines. The low number of piriform glands leads to the formation of very simple attachment discs, which fix the individual threads to the substratum. Thread bundles are attached to the substratum by means of accumulated attachment discs.     

Phylogenetic relationships of antlered flies, Phytalmia Gerstaecker (Diptera: Tephritidae): the evolution of antler shape and mating behaviour

Abstract  The genus Phytalmia (antlered flies, Diptera: Tephritidae) contains remarkable flies with elaborate male head projections known as antlers. The antlers are used in antagonistic intraspecific interactions between males competing to occupy oviposition sites. Phylogenetic relationships between the seven known species of Phytalmia were chosen to be investigated in order to determine the current monophyly of the genus and to assess the evolution of secondary sexual characters and associated behaviours, especially male antlers, fore-femoral, spines and stilting. A phylogenetic analysis of Phytalmia was conducted using two closely related species from the same tribe (Phytalmiini): Sessilina nigrilinea (Walker) and Diplochorda minor Malloch, and one species from a different tribe (Dacini): the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) as outgroups. A total of 1259 base pairs of mitochondrial DNA (COII and 16S) and 35 morphological characters were included in the analysis, using parsimony and maximum likelihood inference methods. A phylogenetic tree generated from combined data was used to infer the evolution of antler shapes and associated behaviours in the genus. The results of this paper support the monophyly of Phytalmia , with P. cervicornis Gerstaecker the sister group of the remaining species. The phylogeny suggests a general decrease in antler complexity over evolutionary time; however, the behavioural sequence of male flies becomes more complex over time, with steps (e.g. stilting) being added to the repertoire during antagonistic encounters. Finally, there are strong correlations between functionally constrained morphological and behavioural characters, therefore allowing the authors to make predications for species whose behaviours are unknown based on morphological attributes and the authors' hypothesised evolution of Phytalmia .     

Organization of the spinnerets and spigots in the orb web spider,Argiope bruennichi (Araneae: Araneidae)

Although the basic taxonomic characteristics usually remain unchanged, some spinning apparatuses undergo consistent adaptive variations. As the presence of additional protuberances known as nubbins and tartipores have caused disagreements regarding some Araneidae spiders, more detailed definitions on the cuticular structures have recently been proposed. Reflecting this definition, microstructural organization of silk spinning apparatuses in the orb web spider Argiope bruennichi were reconsidered using field emission scanning electron microscopy. Among the seven kinds of functional spigots in females, it was revealed that two types (major ampullates and pyrifoms) are located on anterior spinnerets and another five types are distributed on median (minor ampullates, tubuliforms and aciniforms) or posterior (tubuliforms, flagelliforms, aggregates and aciniforms) spinnerets, respectively. In addition to functional spigots, cuticular remnants of the nubbins and the tartipores were found on the spinning fields, but the number of tartipores on each spinneret varied among individuals based on maturity. Nevertheless, three kinds of cuticular protuberances of ampullate silk glands were clearly visible at both the anterior and median spinnerets.     

Normal and atypical nesting behaviour of Copris lunaris (L.): comparison with related species (Coleoptera, Scarabaeidae)

Abstract. 1. The lifecycle, mating and larval behaviour of Clunaris are described. Adults appeared in the autumn and nested in the following spring. The female beetle remained in the nest with the brood and could nest again the following year.
2. Nesting was initiated when virgin females were mated in the spring. Brood balls were formed by techniques sirnilar to those used by Scarabaeini. The female beetle left the nest soon after the first imagos broke out of the brood balls.
3. Nesting behaviour was readily modified by external conditions. Many parts of the sequence could be repeated or omitted. The female beetle left the nest if the brood was removed, but she remained for longer than usual if younger brood was substituted near the end of the normal nesting period.
4. Certain experimental conditions released behaviour patterns typical of other species. These were formation of superficial nests or of two-chambered nests, oviposition before completing the brood ball, and coating of the brood balls with soil (all found in other Coprina), as well as ball rolling and ball burial (found in Scarabaeini). The results are discussed in relation to the evolution of Copris nesting behaviour.     

Biogeography of the grasses (Poaceae): a phylogenetic approach to reveal evolutionary history in geographical space and geological time

The grasses (Poaceae) are the fifth most diverse family of angiosperms, including 800 genera and more than 10 000 species. Few phylogenetic studies have tried to investigate palaeo‐biogeographical and palaeo‐ecological scenarios that may have led to present‐day distribution and diversity of grasses at the family level. We produced a dated phylogenetic tree based on combined plastid DNA sequences and a comprehensive sample of Poaceae. Furthermore, we produced an additional tree using a supermatrix of morphological and molecular data that included all 800 grass genera so that ancestral biogeography and ecological habitats could be inferred. We used a likelihood‐based method, which allows the estimation of ancestral polymorphism in both biogeographical and ecological analyses for large data sets. The origin of Poaceae was retrieved as African and shade adapted. The crown node of the BEP + PACCMAD clade was dated at 57 Mya, in the early Eocene. Grasses dispersed to all continents by approximately 60 million years after their Gondwanan origin in the late Cretaceous. PACCMAD taxa adapted to open habitats as early as the late Eocene, a date consistent with recent phytolith fossil data for North America. C₄ photosynthesis first originated in Africa, at least for Chloridoideae in the Eocene at c. 30 Mya. The BEP clade members adapted to open habitats later than PACCMAD members; this was inferred to occur in Eurasia in the Oligocene. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 543–557.     

Olfactory host‐finding behaviour of Oulema melanopus (Coleoptera: Chrysomelidae) and its parasitoid,Tetrastichus julis (Hymenoptera: Eulophidae)

Behavioural responses to the host‐associated olfactory cues have not been completely understood for the cereal leaf beetle, Oulema melanopus, and its principal parasitoid, Tetrastichus julis. We, therefore, investigated the role of olfactory cues in the host‐finding behaviour of these species using olfactory bioassays. Behavioural responses of O. melanopus to odours emanating from intact host plants (wheat, oat, barley) vs. a clean‐air control were tested using multichoice and two‐choice bioassays. For T. julis, responses of naïve and experienced adult female wasps to odours associated with the faecal coat of O. melanopus larvae were measured under multichoice and two‐choice conditions. Our results indicate that olfactory cues are involved in the host‐finding behaviour of both O. melanopus and T. julis. Olfactory responses of O. melanopus were influenced by the sex of the beetle and the physiological stage of adults (reproductively active vs. in reproductive diapause). Females respond to olfactory cues in greater proportions than males, and reproductively active, overwintered adults show greater responsiveness than teneral adults in reproductive diapause. Behavioural responses to cues emanating from different crop species were different in multichoice bioassays but not in two‐choice bioassays. Further, we report for the first time that the olfactory cues associated with the faecal coat of O. melanopus evoke host‐finding behaviour of its parasitoid, T. julis. Naïve female wasps are more likely to use these cues to locate the potential host than experienced females. The results of this investigation provide insights into host finding by both the species and the nature of behavioural response brought about by olfactory stimuli, and the results can help to design strategies to improve parasitoid activity by enhancing the crop environment to generate cues for host finding and to manage O. melanopus populations.     

The biology of Tauala lepidus,a jumping spider (Araneae: Salticidae) from Queensland: display and predatory behaviour

Abstract

The display and predatory behaviour of Tauala lepidus Wanless, an abundant salticid in north Queensland rainforests, was investigated in the laboratory and, to a lesser extent, in nature. T. lepidus leapt and walked into alien webs to catch spiders and insects, and was captured and fed on by other spiders. Females ate each other’s eggs. During intraspecific interactions, a complex repertoire of displays was used. Courtship versatility occurred, each individual male having a conditional strategy of different behaviours depending on whether the female is at or away from her nest, and whether she is adult or subadult. Yet other combinations of displays occurred during male-male and female-female interactions. Apparently, pheromones on nests and draglines of females released male courtship. Abdomen twitching, a behaviour common to the display repertoires of many salticids, was an especially complex and pervasive behaviour of T. lepidus. T. lepidus also twitched its abdomen when it contacted alien webs and preyed on other species of spiders. The behaviour of T. lepidus is compared to that of Jacksonoides queenslandica Wanless, a species from the same group (Astieae).     

Muscular anatomy of the giant whipscorpion Mastigoproctus giganteus (Lucas) (Arachnida: Uropygi) and its evolutionary significance

Skeletal muscles in the whipscorpion Mastigoproctus giganteus are surveyed and compared with those of several other to clarify the evolutionary morphology and phylogenetic relationships of arachnids. Representatives from 90 muscle groups are described and illustrated, and their possible functions are proposed. Principal results of this analysis include new proposed homologies for the anterior opisthosomal appendages and sclerites in tetrapulmonate arachnids (that is, Trigonotarbida, Araneae, Amblypygi, Uropygi), the discovery that muscular attachments in arthropods can shift from the mesodermal endosternite to the ectodermal exoskeleton, a reconstruction of the evolutionary transformations associated with the apparent uncoupling of pharyngeal and locomotor complexes in the prosoma of Pedipalpi (that is, Amblypygi and Uropygi), and an expanded list of unique synapomorphies supporting the sister-group status of Amblypygi and Uropygi.     

The moss mites of Yes Tor, Dartmoor, Devon (Acari: Oribatida) and their evolutionary significance

A species list of oribatid mites collected from the summit of Yes Tor, Dartmoor, is given, together with notes on their distribution. Yes Tor lies south of latitude 51° in that part of Britain which remained ice free during the Pleistocene. The species found on Yes Tor are compared with the species recorded from the summits of 11 mountains lying north of latitude 53°, which were all ice-covered at the period of maximum glaciation. In the case of Yes Tor and these 11 northern montane sites, three similar faunal elements have been identified arctic-alpine species, European-alpine species and a group of British montane species that are also common in the lowlands. The view is advanced that the similarity of the montane oribatid fauna of Yes Tor with that of the mountains to the north of latitude 53° has resulted from the probable movements of the mites during Quaternary and postglacial times over an area extending from the Arctic Circle to the Mediterranean. It is suggested that these movements, while being largely spontaneous, have also been aided by transportation agents, both physical and biological.     

The biology of Euryattus sp. indet., a web-building jumping spider (Araneae, Salticidae) from Queensland: utilization of silk, predatory behaviour and intraspecific interactions

The biology of an aberrant saltioid spider, Euryattus Thorell sp. indet., is described from observations in a Queensland rain forest and the laboratory. Pronounced morphological and behavioural changes occur during post-embryological development. Juveniles spin webs, but adult females make 'suspension nests' by suspending a curled-up leaf by heavy guylines from a rock ledge or vegetation. Adult males spin neither webs nor suspension nests. Females oviposit inside their suspension nests, but if denied access to leaves for suspension, they spin and oviposit in webs similar to those spun by juveniles. The flat, papery egg sacs of Euryattus are atypical for a salticid, being more like the egg sacs of many of the Gnaphosidae. Intraspecific display behaviour has characteristics in common with typical salticids, but also includes unique features. Male courtship includes vibratory displays performed on the surface of the suspended leaf. Mating occurs inside the curled-up leaf. Males co-habit with sub-adult females in suspension nests. Females take over suspension nests of other females and eat each other's eggs. Normal locomotion, intraspecific interactions, and predatory behaviour are characterized by frequent leaping. Euryattus routinely makes long and accurate leaps on to prey, including flying insects intercepted in mid-air and spiders and insects located in alien webs. The phylogenetic implications of the unusual characteristics of Euryattus are considered.     

The enigmatic Otway odd-clawed spider (Progradungula otwayensis Milledge, 1997, Gradungulidae,Araneae): Natural history,first description of the female and micro-computed tomography of the male palpal organ

The recently described cribellate gradungulid Progradungula otwayensis Milledge, 1997 is endemic to the Great Otway National Park (Victoria, Australia) and known from only one male and a few juvenile specimens. In a recent survey we recorded 47 specimens at several localities across the western part of the Great Otway National park. Our field data suggest that this species is dependant on the microclimate in the hollows of old myrtle beech trees since other hollow trees were very much less inhabited. Furthermore, we describe the female for the first time and study the male palpal organ by using X-ray microtomography. The female genitalia are characterized by eight spermathecae which are grouped in two quartets. The spermophor resembles the general organization of gradungulids, but is similar to Kaiya Gray, 1987 by a convoluted appearance within the embolus. The muscle 30 is located in the cymbium and resembles the organization of other non-entelegyne Araneomorphae.     

Predatory behaviour of the social orb-weaver spider,Geratonephila burmanica n. gen., n. sp. (Araneae: Nephilidae) with its wasp prey,Cascoscelio incassus n. gen., n. sp. (Hymenoptera: Platygastridae) in Early Cretaceous Burmese amber

The present work shows predatory behaviour of the social orb-weaver spider, Geratonephila burmanica n. gen., n. sp. (Araneae: Nephilidae) against a parasitic wasp, Cascoscelio incassus n. gen., n. sp. (Hymenoptera: Platygastridae) in Early Cretaceous Burmese amber. An adult male and juvenile of G. burmanica in the same web provide the first fossil evidence of sociality in spiders. The spider is characterised by a pedipalp with a hemispherical tegulum, a subtegulum curved at 180°and an apical spiralled embolas-conductor bent approximately 45°at midpoint. The male wasp is characterised by an ocellar tubercle, 12-segmented antennae with a feeble five-segmented clava, thick sensilla trichodea curvata with rounded ends on the claval antennomeres, a short uncus, a short post-marginal vein and a nebulose radial sector (Rs) vein extending from the uncus to the costal margin of the forewing. This is the first fossil evidence of spider sociality and a fossil spider attacking prey trapped in its web.     

Skeletomuscular anatomy of the harvestman Leiobunum aldrichi (Weed, 1893) (Arachnida: Opiliones: Palpatores) and its evolutionary significance

Skeletal muscles of the North American harvestman Leiobunum aldrichi are exhaustively surveyed and compared with other chelicerates to clarify the evolutionary morphology and phylogenetic relationships of arachnids. Representatives of 104 muscle groups are described and illustrated, and their possible functions are proposed. Comparisons of the feeding apparatus of L. aldrichi with that of other opilions, especially Sim (Cyphophthalmi) and Acromares (Laniatores), and two scorpion genera ( Centruroides, Pandinus ) indicate that the pharyngeal apparatus in L. aldrichi is derived and that its ability to accommodate large food particles is a secondary rather than primitive condition. Comparisons reveal several possible synapomorphies between Opiliones and Scorpiones suggesting that these orders may be sister groups. Apparently unique synapomorphies include an extrinsic cheliceral muscle that arises from the carapace and inserts on the second cheliceral article (deutomerite); an epistome divided into distal and proximal parts by a transverse sulcus; pharyngeal dilator muscles supported by a peripharyngeal skeleton formed by one dorsomedial and two ventrolateral epistomal processes, the latter also with muscular attachments to the endosternite; a specialized preoral chamber (stomodieca) derived from extensions (coxapophyses) of the coxae of the pedipalp and first two leg pairs; internal processes associated with the coxapophyses that serve, in part, as an attachment for muscles operating the coxa-trochanter joints, and lateral endosternal suspensor muscles that insert on the arthrodial membrane between the leg coxae. These are the first observations providing explicit support for an Opiliones-Scorpiones clade.