rRNA及MaSp1基因证据不支持圆网蛛类(蜘蛛目:妖面蛛总科+园蛛总科)的单系起源

圆网蛛类(妖面蛛总科 园蛛总科)是否为单系,圆网究竟经历一次进化还是多次进化,这是多年来有争论的、悬而未决的蛛形学难题之一。本文测定了包括妖面蛛总科、园蛛总科和非圆网蛛类等类群在内的9科10种蜘蛛线粒体12S rDNA、16S rDNA及核18S rDNA、28S rDNA等4个基因片段序列,并基于4个基因序列的整合数据,分别通过邻接(NJ)法、最大简约(MP)法、最大似然(ML)法和贝叶斯法(Bayesian)分析,对园蛛总科和妖面蛛总科蜘蛛之间的分子系统关系进行了探讨。系统发生结果表明:1)园蛛总科和妖面蛛总科蜘蛛不是姊妹群,从而支持这两个类群的圆网是平行演化而非同源演化的观点;2)筛器类蜘蛛并非单系发生而为多系发生。另外,依据编码大壶状腺丝蛋白-1(MaSp1)C末端非重复氨基酸序列区段的核酸序列重建的系统发生树也证实圆网蛛类并非单系发生。     

圆网蛛类系统发生与网型进化

本文从妖面蛛总科和园蛛总科系统发生关系、园蛛总科网型不同的蜘蛛间的系统发生关系及蛛网构建行为等几个方面着重介绍了圆网蛛类系统发生及网型进化的研究进展.圆网蛛类系统发生与其网型进化有效地结合、进行综合研究将有助于圆网蛛类的起源及网型多样性的研究.     

Adhesive efficiency of spider prey capture threads

Cribellar capture threads are comprised of thousands of fine silk fibrils that are produced by the spigots of a spider's cribellum spinning plate and are supported by larger interior axial fibers. This study examined factors that constrain the stickiness of cribellar threads spun by members of the orb-weaving family Uloboridae in the Deinopoidea clade and compared the material efficiency of these threads with that of viscous capture threads produced by members of their sister clade, the Araneoidea. An independent contrast analysis confirmed the direct relationship between cribellar spigot number and cribellar thread stickiness. A model based on this relationship showed that cribellar thread stickiness is achieved at a rapidly decreasing material efficiency, as measured in terms of stickiness per spigot. Another limitation of cribellar thread was documented when the threads of two uloborid species were measured with contact plates of four widths. Unlike that of viscous threads, the stickiness of cribellar threads did not increase as plate width increased, indicating that only narrow bands along the edges of thread contact contributed to their stickiness. As thread volume increased, the gross material efficiency of cribellar threads decreased much more rapidly than that of viscous threads. However, cribellar threads achieved their stickiness at a much greater gross material efficiency than did viscous threads, making it more challenging to explain the transition from deinopoid to araneoid orb-webs.     

Capture thread extensibility of orb-weaving spiders: testing punctuated and associative explanations of character evolution

Spider orb-webs contain sticky prey capture threads and non-sticky support threads. Primitive orb-weavers of the Deinopoidea produce dry cribellar threads made of thousands of silk fibrils that surround supporting axial fibres, whereas the viscous threads of modern Araneoidea orb-weavers produce adhesive threads with an aqueous solution that coalesces as droplets around the axial fibres. We have previously shown that the greater diversity of the Araneoidea is phylogenetically significant and attributed this disparity to a number of advantages, considered key innovations, that adhesive thread has over cribellar thread. An important putative advantage of adhesive thread demonstrated by Kohler and Vollrath in their 1995 study is its greater extensibility, a feature that better adapts it to absorb the kinetic energy of a prey strike. However, this conclusion is based on a two-species comparison that does not take advantage of the modern comparative method that requires hypotheses to be tested in a phylogenetic context. Using a transformational analysis to examine threads produced by nine species, our study finds no support for the punctuated explanation that adhesive thread has a greater extensibility than cribellar thread. Instead, it strongly supports the associative null hypothesis that capture thread extensibility is tuned to spider mass and to architectural features of the web, including its capture area, capture spiral spacing, and capture area per radius.     

Adhesive compatibility of cribellar and viscous prey capture threads and its implication for the evolution of orb-weaving spiders

Evolution of orb-weaving spiders that comprise the Orbiculariae clade involved a transition in the composition of prey capture thread that has been challenging to explain. The primitive cribellar threads spun by members of the Deinopoidea subclade resemble the capture threads of their non-orb-web-weaving ancestors and are formed of thousands of fine, dry, protein cribellar fibrils. In contrast, the derived viscous capture threads spun by members of the Araneoidea subclade have regularly spaced, aqueous adhesive droplets. When second instar deinopoid spiderlings emerge from egg sacs they are unable to spin cribellar threads, and, therefore, do not construct orb-webs; whereas second instar araneoids spin capture threads and construct orb-webs. If, as we hypothesize, viscous material evolved to enable second instar spiderlings to construct orb-webs, early araneoids may have spun composite cribellar-viscous capture threads. To examine the functional feasibility of such intermediate capture threads, we compared the adhesion of cribellar threads, viscous threads, and combined cribellar-viscous threads. The stickiness of these combined threads was greater than that of native cribellar or viscous threads alone. The viscous material of Araneus marmoreus threads exhibited a substantial increase in stickiness when combined with cribellar fibrils and that of Argiope aurantia threads a small increase in stickiness when combined with cribellar fibrils. Thus, if early araneoids retained their ability to spin cribellar threads after having evolved glands that produced viscous material, their composite threads could have formed a functional adhesive system that achieved its stickiness at no loss of material economy.     

EVOLUTIONARY SHIFTS IN THE SPECTRAL PROPERTIES OF SPIDER SILKS

We measured the reflectance properties of unpigmented silks spun by a systematic array of primitive (Deinopoidea) and derived (Araneoidea) aerial, web-spinning spiders, as well as silks spun by Araneomorphae and Mygalomorphae spiders that do not spin aerial webs. Our data show that all of the primitive aerial web spinners produce catching silks with a spectral peak in the ultraviolet (UV), and cladistic analysis suggests that high UV reflection is the primitive character state for silk spectral properties. In contrast, all of the derived aerial web spinners produce silks that are spectrally flat or characterized by reduced reflectance in the UV. Correlated with the evolution of these catching silks is a 37-fold increase in species number and apparent habitat expansion. This suggests that the unique silk proteins spun by the araneoids have been important to their ecological and evolutionary diversity.     

Changes in spinning anatomy and thread stickiness associated with the origin of orb-weaving spiders

The cribellum is an oval spinning field whose spigots produce silk fibrils that form the outer surfaces of the primitive prey capture threads found in aerial spider webs. A comparison of the cribella and cribellar capture threads of 13 species of spiders representing seven families (Amaurobiidae, Desidae, Dictynidae, Filistatidae, Neolanidae, Oecobiidae, and Uloboridae) confirms that the stickness of a cribellar thread is directly related to the number of spigots on a spider's cribellum. This comparison also demonstrates that the origin of orb-weaving spiders from ancestors that constructed less highly organized webs was associated with increases in both the weight-specific number of cribellum spigots and the weight-specific stickiness of cribellar prey capture threads. In contrast to other cribellate spiders, the number of cribellum spigots of orb-weaving species of the family Uloboridae scales to spider mass. Thus, the origin of orb-weaving spiders involved not only behavioural changes that stylized and restricted the placement of cribellar threads, but also included morphological changes that increased the stickiness of these capture threads by endowing them with more cribellar fibrils.     

从12S rRNA基因第三结构域序列分析探讨蜘蛛若干重要类群的亲缘关系

本文将12S rRNA基因序列分析应用于研究若干重要蜘蛛类群的系统关系,以对传统的分类研究结论进行验证和补充,并且探讨12S rRNA基因序列分析在蜘蛛系统发生研究中的适用性。根据12S rRNA基因第3结构域构建的分子系统树得出结论:1.圆网类(即妖面蛛总科与园蛛总科)并非单系;2.隙蛛与暗蛛较漏斗蛛具有更近的亲缘关系;3.壁钱和拟壁钱并不近缘;4.有筛器类蜘蛛为复系类群;5.12S rRNA基因第3结构域片段对推断近缘科属间的系统发生关系是有效的遗传标记。     

Orb-weaving spider diversity in the Iberá Marshlands, Argentina

The Iberá Marshlands RAMSAR reserve, in the northeast of Argentina, is one of the largest and most important wetlands of America. In this study we assess orb-weaving spider (Araneae: Orbiculariae) diversity in this reserve, analyzing different facets of local diversity (species richness, diversity, evenness and taxonomic distinctness), and the contribution of species differentiation (beta diversity) among localities and habitat types to the composition of regional diversity. We found 1657 individuals of 59 orb-weaving spider species/morphospecies. Local diversity differs among the three sampled localities. At the habitat level, the different facets of biodiversity followed a clear pattern, where woodlands have higher species richness, diversity, evenness and taxonomic distinctness than savannas. Savanna sites shared a common spider species composition, while woodland communities have high values of complementarity. Thus, beta diversity has a very high contribution to the regional diversity of the orb-weaving spiders in the Iberá Marshlands. We suggest that conservation management in the reserve should be directed towards promoting natural spatial heterogeneity, giving special protection to habitat mosaics in different localities.     

Economics of spider orb-webs: the benefits of producing adhesive capture thread and of recycling silk

1. The replacement of dry, fuzzy cribellar prey capture thread by viscous, adhesive capture thread was a major event in the evolution of orb-weaving spiders. Over 95% of all orb-weaving species now produce adhesive threads.
2. Adhesive thread achieves its stickiness with a much greater material economy than does cribellar thread.
3. Transformational analyses show that, relative to spider mass, adhesive orb-weavers invest less material per mm of capture thread and produce stickier capture threads than do cribellate orb-weavers.
4. The total cost of producing an orb-web that contains cribellar thread is reduced by 32% when a spider recycles its silk and another 34% when these capture threads are replaced by adhesive threads of equal stickiness.
5. The increased economy with which adhesive capture thread achieves its stickiness may have been an important factor that favoured the origin and success of modern orb-weaving spiders that produce adhesive capture threads.     

Foraging strategies of Eriophora transmarina and Nephila plumipes (Araneae: Araneoidea): Nocturnal and diurnal orb-weaving spiders

Abstract The foraging behaviour, web characteristics and prey availability of two sympatric orb-weaving spiders, Nephila plumipes and Eriophora transmarina (Araneae: Araneoidea), are compared. The spiders are similarly sized but have different temporal foraging patterns. Nephila plumipes spins a relatively permanent web and captures most of its prey during the day. Eriophora transmarina only forages at night, spinning a new web every night and usually dismantling it at dawn. These different foraging activities are most likely to be responsible for the observed differences in the types and rates of prey capture: E. transmarina captured mostly Lepidoptera that were more abundant at night than during the day, while N. plumipes captured mostly Hymenoptera that were more abundant during the day than at night. While nocturnal E. transmarina have less time available for foraging than the diurnal N. plumipes, the former has a substantially higher nocturnal prey capture rate. We argue that the difference between the species in their prey capture rates are likely to be due to differences in the architecture of their webs.     

Evidence for diet effects on the composition of silk proteins produced by spiders

Silks are highly expressed, secreted proteins that represent a substantial metabolic cost to the insects and spiders that produce them. Female spiders in the superfamily Araneoidea (the orb-spinning spiders and their close relatives) spin six different kinds of silk (three fibroins and three fibrous protein glues) that differ in amino acid content and protein structure. In addition to this diversity in silks produced by different glands, we found that individual spiders of the same species can spin dragline silks (drawn from the spider's ampullate gland) that vary in content as well. Freely foraging ARGIOPE: argentata (Araneae: Araneoidea), collected from 13 Caribbean islands, produced dragline silk that showed an inverse relationship between the amount of serine and glycine they contained. X-ray microdiffraction of the silks localized these differences to the amorphous regions of the protein that are thought to lend silks their elasticity. The crystalline regions of the proteins, which lend silks their strength, were unaffected. Laboratory experiments with ARGIOPE: keyserlingi suggested that variation in silk composition reflects the type of prey the spiders were fed but not the total amount of prey they received. Hence, it may be that the amino acid content (and perhaps the mechanical properties) of dragline silk spun by ARGIOPE: directly reflect the spiders' diet. The ability to vary silk composition and, possibly, function is particularly important for organisms that disperse broadly, such as Argiope, and that occupy diverse habitats with diverse populations of prey.     

Foraging advantages of mixed-species association between solitary and colonial orb-weaving spiders

This study investigated association between solitary orb-weaving spiders and a colonial orb-weaving spider, Metepeira incrassata (Araneae: Araneidae). Spiders were sampled along transects and an index of species association showed that two of the species were associated more frequently than expected based on a null hipothesis of random co-occurrence. The potential advantages of mixed-species association were investigated by comparing prey-capture success of one of these associates, Nephila clavipes (Araneae: Tetragnathidae), when it occurs alone, in single-species groups, and when associated with M. incrassata colonies. Field observations of prey-capture success by all three of these categories of N. clavipes revealed that individuals in M. incrassata colonies captured significantly more prey than solitaries or individuals in single-species groups. The increase in prey capture by N. clavipes in M. incrassata colonies may result from utilization of a foraging niche which intercepts a diffirent spectrum of prey than that available to single-species groups or solitaries. Related to this enhanced prey consumption is greater fecundity of spiders in association with M. incrassata as compared to solitaries or individuals in single-species groups.     

The material cost and stickiness of capture threads and the evolution of orb-weaving spiders

Prey capture threads are essential to the operation of spider orb-webs because they prevent insects that have been intercepted from escaping before a spider can subdue them. The volume of material invested in a web's capture threads is related to spider weight and is the same for primitive orb-weavers that produce cribellar capture thread and modern orb-weavers that produce adhesive capture thread. However, as adhesive capture thread achieves greater stickiness relative to its volume, adhesive orb-webs have a greater total stickiness and, consequently, a greater prey capture potential than cribellate orb-webs. These differences appear to have favoured the transition from cribellate to adhesive capture threads and the success of adhesive orb-weavers, which include 95% of all orb-weaving species. Differences in the thread economy and the total stickiness of webs constructed by spiders of different weights also suggest that adhesive orb-weavers should grow more rapidly and be capable of attaining a larger size than cribellate orb-weavers.     

Patterns of prey utilization in a web of orb-weaving spiderAraneus pinguis (Karsch)

The pattern of prey utilization of the orb-weaving spider Araneus pinguis was studies by comparing between arthropods restrained in the empty webs (spiders were removed) and those unattacked in the intact webs (spiders were not removed). The number of arthropods was larger in the empty webs than in the intact webs. In the empty webs, web area, mesh width, number of radii, and signal thread length were presumed to affect the number of arthropods left in a single web. As for the intact webs, web area, mesh width, and web-exposure time were important factors. In the empty webs, the density of arthropods decreased away from the hub. On the other hand, the density of unattacked arthropods in the intact webs was the same throughout the web. Arthropods in the empty webs were larger than those in the intact webs. The upper limit in size of unattacked arthropods increased along with the distance from the hub in the intact webs, but not in the empty webs. These results indicate increase in the minimum size of eaten arthropods increased in the former. This positive sizedistance relation may have resulted from the adaptive switching of spiders’ alternative foraging methods (i.e., the rapid attack at encounter and the later eating during web deconstruction) on the basis of the prey profitability.     

A novel property of spider silk: chemical defence against ants

Spider webs are made of silk, the properties of which ensure remarkable efficiency at capturing prey. However, remaining on, or near, the web exposes the resident spiders to many potential predators, such as ants. Surprisingly, ants are rarely reported foraging on the webs of orb-weaving spiders, despite the formidable capacity of ants to subdue prey and repel enemies, the diversity and abundance of orb-web spiders, and the nutritional value of the web and resident spider. We explain this paradox by reporting a novel property of the silk produced by the orb-web spider Nephila antipodiana (Walckenaer). These spiders deposit on the silk a pyrrolidine alkaloid (2-pyrrolidinone) that provides protection from ant invasion. Furthermore, the ontogenetic change in the production of 2-pyrrolidinone suggests that this compound represents an adaptive response to the threat of natural enemies, rather than a simple by-product of silk synthesis: while 2-pyrrolidinone occurs on the silk threads produced by adult and large juvenile spiders, it is absent on threads produced by small juvenile spiders, whose threads are sufficiently thin to be inaccessible to ants.     

Factors governing the stickiness of cribellar prey capture threads in the spider family Uloboridae

The surface of a cribellar prey capture thread is formed of thousands of fine, looped fibrils, each issuing from one of the spigots on an oval spinning plate termed the cribellum. This plesiomorphic capture thread is retained by members of the family Uloboridae, in which its stickiness differs among genera. An examination of five cribellar thread features in nine uloborid species shows that only the number of fibrils that form a thread explains these differences in thread stickiness. Neither the physical features of these fibrils, nor the manner in which they are combined to form threads differs among species. Threads produced by orb-weaving species contain fewer fibrils than those produced by species that build reduced webs. Relative to spider weight, the number of fibrils that form a cribellar thread is greatest in simple-web species of the genus Miagrammopes, less in triangle-web species of the genus Hyptiotes, and least in orb-weaving species representing five genera. A transformational analysis shows that change in the number of cribellum spigots is directly related to change in the stickiness of cribellar thread. This direct relationship between the material invested in a cribellar thread and its stickiness may have been a limiting factor that favored the switch from the dry cribellar threads of uloborids to the adhesive capture threads produced by other orb-weaving families. © 1994 Wiley-Liss, Inc.     

Differences in venom composition between orb-weaving and wandering Hawaiian Tetragnatha (Araneae)

Spider venoms are complex mixtures of toxins that are used primarily for immobilizing prey. There is evidence of chemical variation in spider venoms among close relatives, yet few studies have analysed their evolution within an ecological and phylogenetic framework. On the Hawaiian archipelago, Tetragnatha, a cosmopolitan orb-weaving genus, has undergone a radiation in which a monophyletic lineage has abandoned web-building and become obligately wandering foragers. This study compares venom composition and details of feeding behaviour between orb-weaving and wandering Hawaiian Tetragnatha. Protein gel electrophoresis patterns indicated that relative to orb-weavers, wandering species had a reduced concentration of low molecular weight (<14kDa) components. Both orb-weaving and wandering Tetragnatha captured flying prey (adult lepidopterans, dipterans), but wandering spiders also captured non-flying prey (insect larvae, spiders). There were no distinct differences between orb-weavers and wanderers in prey capture and immobilization sequences, or in the paralytic effects of bites on prey. However, prey bitten by wanderers took longer to be permanently immobilized than prey bitten by orb-weavers. Contrary to predictions, there was no indication that web-loss in this group was associated with an increase in venom potency.     

Impact of predation by spiders on a territorial damselfly (Odonata: Calopterygidae)

Summary Predation by orb-weaving spiders and crab spiders on the damselfly Calopteryx haemorrhoidalis was studied at a small stream in Southern France. One species of orb-weaving spider, Larinioides folium, caught 76% of the damselflies which fell prey to spiders. Displacement experiments on spiders on sections of bank and the positioning of webs in male territories show that the density and distribution of damselflies is not influenced by orbweb density or by the position of webs. Predation rates corresponded to orb-web density, but neither for sex nor for stage was there a relationship with damselfly density. Mean daily predation rates ranged between 0.9% for females and 4.1% for adult males. Predation risk to adult damselflies by orb-weaving spiders was male biased, whereas among tenerals there was no bias. Males were captured more frequently at territories near the water. Captures show a maximum at noon when territorial disputes of adult males were most frequent. After orbwebs were placed within territories predation rate of males was strongly increased. Predation risk to adult females in the direct vicinity of the stream was less than in the bank vegetation where they perch close to orbwebs. The risk of predation by crab spiders, which catch damselflies at their perching sites, was not sex-biased.     

Resting metabolic rates of two orbweb spiders: a first approach to evolutionary success of ecribellate spiders

Spiders are considered conservative with regard to their resting metabolic rate, presenting the same allometric relation with body mass as the majority of land-arthropods. Nevertheless, web-building is thought to have a great impact on the energetic metabolism, and any modification that affects this complex behavior is expected to have an impact over the daily energetic budget. We analyzed the possibility of the presence of the cribellum having an effect on the allometric relation between resting metabolic rate and body mass for an ecribellate species (Zosis geniculata) and a cribellate one (Metazygia rogenhoferi), and employed a model selection approach to test if these species had the same allometric relationship as other land-arthropods. Our results show that M. rogenhoferi has a higher resting metabolic rate, while Z. geniculata fitted the allometric prediction for land arthropods. This indicates that the absence of the cribellum is associated with a higher resting metabolic rate, thus explaining the higher promptness to activity found for the ecribellate species. If our result proves to be a general rule among spiders, the radiation of Araneoidea could be connected to a more energy-consuming life style. Thus, we briefly outline an alternative model of diversification of Araneoidea that accounts for this possibility.