The spinning apparatus of Uloboridae in relation to the structure and construction of capture threads (Arachnida,Araneida)

Summary Besides the two axial fibers and the mass of cribellum fibrils, a third component is present in the capture threads of uloborids. This is a substructure originating from the paracribellum. It probably helps to fasten the axial fibers in their position. The axial fibers are secreted from the two glandulae pseudoflagelliformes whose spigots are situated on the posterior spinnerets. It is hypothesized that the cribellum fibrils become jammed and thus fixed between the axial fibers by periodical abduction and adduction of these spinnerets.     

Selected aspects of spinning apparatus development in Araneus cavaticus (Araneae,Araneidae)

In the first half of this century, several workers observed small, seemingly glandular structures attached to the ampullate glands of spiders. Hence, they were termed accessory ampullate glands. In juvenile Araneus cavaticus, two pairs of these structures are present (starting at least with third instars), one pair attached to the major ampullate (MaA) glands and the other pair attached to the minor ampullate (MiA) glands. In adults, two pairs of accessory MaA glands and two pairs of accessory MiA glands are present. The two latter-formed pairs of accessory ampullate glands are clearly the remnants of those ampullate glands which atrophy shortly after adulthood is reached. Morphological similarities between these accessory ampullate glands and those present in juveniles provide an indication that the latter also have their origin in functional ampullate glands. A reduction in the number of ampullate glands following the last molt occurs in many spiders. The reason(s) for these reductions is unknown. In penultimate spiders close to ecdysis, we have observed that while the larger pairs of MaA and MiA glands (those that are retained in the adult) are undergoing molt-related changes which apparently render them nonfunctional, their smaller counterparts are seemingly unaffected and functional. This raises the possibility that the principal role of the smaller ampullate glands may be to assume functions during the pre-ecdysial period which are normally in the domain of the larger ampullate glands. If true, then their degeneration after the last molt would make economic sense. The presence of cylindrical spigots in juvenile females starting with fourth instars is documented.     

The spinning apparatus of Polenecia producta (Araneae,Uloboridae): Structure and histochemistry

Summary The spinning apparatus of the uloborid spider Polenecia producta was studied to complete previous studies on the same family of spiders. The structure of spinnerets and spigots, under scanning electron microscopy, and the main anatomical and histochemical characteristics of the spinning glands of adult females and males are described. In addition some observations on the spinning apparatus at three successive stages of development are made. There are nine kinds of silk glands in Polenecia, i.e. one more (aciniform — B glands) than found in other uloborids. The spinning apparatus of Polenecia is, therefore, the most complex so far known. It is also more complex than that presently known of Araneoidea. The characteristics of the spinning glands of Polenecia are compared with those of other uloborids. Present knowledge of the spinning apparatus of uloborids leads to a renewed discussion of the origin of the orb web in this family and in araneids. It is concluded that these two types of orb webs emerged from independent evolutionary processes.     

The ontogeny of the spinning apparatus of Tengella perfuga (Araneae: Zoropsidae)

Silk is the most recognizable trait of spiders, and silk use has changed throughout spider evolutionary history. While morphology of the adult silk spigot has been a useful character for systematics, few studies have examined the ontogeny of the spinning apparatus, and none of these included cribellate spiders. Here, we report the first published full ontogeny of the spinning apparatus of a cribellate spider, Tengella perfuga. We found the presence of expected spigots: major ampullate gland and piriform gland spigots on the anterior lateral spinneret, minor ampullate gland and aciniform gland spigots on the posterior median spinneret, and aciniform gland spigots on the posterior lateral spinneret. Females, but not males, possessed cylindrical gland spigots on both the posterior median and lateral spinnerets. Spiderlings did not possess a functioning cribellum until the third instar. The cribellum grew with increasing numbers of spigots, but functionality was lost in adult males. Most intriguingly, second instars possessed a distinct triad of pre‐spigots on the posterior lateral spinneret. From the third instar onward, these structures formed the modified spigot along with two flanking spigots (in females) or formed nubbins (in males). We suggest that the modified spigot serves as the source of axial lines in the cribellate silk produced in T. perfuga. We also compare spigot ontogeny from previous studies of ecribellate spiders. These comparisons warrant further exploration using the recent spider tree of life in a phylogenetic comparative analysis of spigot ontogeny datasets, which could yield evidence for homologous spigots across the Araneomorphae, notably the Araneoidea and the Retrolateral Tibial Apophysis (RTA) clades.     

Functional associations between the cribellum spinning plate and capture threads of Miagrammopes animotus (Araneida,Uloboridae)

Summary Uloborid cribellar silk consists of torus-shaped puffs. In Miagrammopes animotus the width of these puffs is about 36% that of the cribellum of the spider and shows a 2.3-fold increase in surface area during development. The cribellar spigot number increase 5.7-fold during development, although, relative to spider mass, it decreases by 34%. Cribellum width is the best predictor of both cribellar silk puff width and length and is as good a predictor of puff surface area as is cribellum surface area. Relative to cribellum width, the length of the calamistrum comb responsible for drawing fibrils from the cribellum changes little during development. The attachment points of cribellar silk to a parallel frame thread become more widely spaced during development, although the number of puffs they delimit changes little.     

Morphology and function of the spinning apparatus of the wolf spider Pardosa amentata (Cl.) (Araneae,Lycosidae)

The histology of the four types of silk gland, occurring in the wolf spider Pardosa amentata are given. The changes in epithelium height, lumen breadth and size of the granules were studied in relation to silk production in the sub-adult and adult stage of the life-cycle.     

Developmental changes in the spinning apparatus over the life cycle of wolf spiders (Araneae: Lycosidae)

Spiders are characterized by their spinning activity. Much of the current knowledge of the spinning apparatus comes from studies on orb web spiders and their relatives, whereas wolf spiders have been more or less neglected in this respect. Therefore, we studied developmental changes in the spinning apparatus of four wolf spiders (Tricca lutetiana, Arctosa alpigena lamperti, Pardosa amentata, and Xerolycosa nemoralis) throughout their life cycles. Each of these lycosids has a stenochronous life cycle, but of varied length (from 1 to 3 years) and number of instars (from seven to ten). Use of the spinning apparatus begins in the first instar, after leaving the egg sac. Secondary ampullate, all piriform, and all but four aciniform glands are tartipore‐accommodated. The tartipores, collared openings through which silk gland ducts pass during proecdysis, appear on the spinning field starting with the second instar. Tartipore‐accommodated glands can function during proecdysis and their evolution corresponds with the way spiders secure themselves when molting. We suggest that the function of aciniform silk in juvenile wolf spiders is to serve as an ancillary “scaffold” supporting the spider's body during ecdysis.     

Functional organization of the spinning apparatus of Cyrtophora citricola with regard to the evolution of the web (Araneae,Araneidae)

Summary The spinning apparatus of Cyrtophora citricola closely corresponds to that of orb-weaving Araneidae, two peculiarities excepted. Firstly the spigots of the piriform glands differ extremely in size, the smallest of them being numerous and having a unique location on the anterior spinnerets. Secondly, the triad complex (on the posterior spinnerets) used by other Araneidae for producing gluey capture threads is lacking. Both these characteristics are correlated with the construction of a fine meshed sheet of dry silk by Cyrtophora instead of orbwebs with capture spirals. The sheet can be understood as being a very much enlarged central area of orb-webs. Since vestiges of triads could be found in early developmental stages of C. citricola, the origin of the meshed sheet from orb-webs with gluey capture threads is clearly demonstrated. The paper includes a study on how the spider produces thread attachments by means of the secretions of the piriform glands.     

An organic coating keeps orb‐weaving spiders (Araneae,Araneoidea, Araneidae) from sticking to their own capture threads

More than 95% of orb‐weaving spider species ensure prey capture success by producing viscous threads equipped with gluey droplets. However, this trap may bear serious risks for the web‐inhabiting spider as well. The obvious question, how a spider avoids getting stuck in its own capture spiral, has gained little scientific attention up till now. In 1905, the French naturalist Jean‐Henry Fabre concluded from anecdotal observation that orb‐weaving spiders protect themselves by a fatty surface coating. Here, we test this hypothesis by indirectly measuring the force necessary to detach an autotomized spider’s leg from the capture spiral of its own web (here called ‘index of adhesion’, IOA). Three groups of legs, each of the species Araneus diadematus Clerck, 1757 and Larinioides sclopetarius (Clerck, 1757), were tested. One was left untreated, one was washed with distilled water (H₂O), and one was washed with the organic solvent carbon disulphide (CS₂). In both species, we found a weak IOA between the spider leg and the gluey capture spiral in untreated and water‐washed legs without significant differences between the two. The IOA approximately doubled, when spider legs had been washed with carbon disulphide prior to measurement, that is, the CS₂‐washed legs stuck significantly more strongly than the untreated and water‐washed legs. These results provide indirect evidence for a protective anti‐adhesive organic coating on the spider’s body surface and so support Fabre’s hypothesis.     

Fine structural analysis on triad spinning spigots of an orb‐web spider's capture threads

Capture threads of the golden orb‐web spider Nephila clavata are produced from the silks of a pair of triad spinning units composed of a flagelliform gland (FLG) and two aggregate glands (AGG). In N. clavata, arrangement of the triad spigots is closely related to coating an axial supporting fiber with sticky aqueous droplets on a continuous and consistent basis for capture thread production. The central spigot of FLG and peripherally located AGG spigots are aligned along a single plane, and both have bullet‐type spigots with flexible segments. In particular, the pear‐shaped spigot of the AGG with a wide‐aperture nozzle provides not only sufficient luminal space for controlling transient storage of the aqueous gluey substance but also an effective spatial system that thoroughly coats the axial fibers with a viscous aqueous solution.     

Multiple bradykinin-related peptides from the capture web of the spider Nephila clavipes (Araneae, Tetragnatidae)

Three bradykinin-related peptides (nephilakinins-I to -III) and bradykinin itself were isolated from the aqueous washing extract of the capture web of the spider Nephila clavipes by gel permeation chromatography on a Sephacryl S-100 column, followed by chromatography in a Hi-Trap Sephadex-G25 Superfine column. The novel peptides occurred in low concentrations and were sequenced through ESI-MS/MS analysis: nephilakinin-I (G-P-N-P-G-F-S-P-F-R-NH2), nephilakinin-II (E-A-P-P-G-F-S-P-F-R-NH2) and nephilakinin-III (P-S-P-P-G-F-S-P-F-R-NH2). Synthetic peptides replicated the novel bradykinin-related peptides, which were submitted to biological characterizations. Nephilakinins were shown to cause constriction on isolated rat ileum preparations and relaxation on rat duodenum muscle preparations at amounts higher than bradykinin; apparently these peptides constitute B2-type agonists of ileal and duodenal smooth muscles. All peptides including the bradykinin were moderately lethal to honeybees. These bradykinin peptides may be related to the predation of insects by the webs of N. clavipes.     

Microstructure of the silk apparatus of the comb-footed spider, Achaearanea tepidariorum (Araneae: Theridiidae)

The microstructural organization of the silk‐spinning apparatus of the comb‐footed spider, Achaearanea tepidariorum, was observed by using a field emission scanning electron microscope. The silk glands of the spider were classified into six groups: ampullate, tubuliform, flagelliform, aggregate, aciniform and pyriform glands. Among these, three types of silk glands, the ampullate, pyriform and aciniform glands, occur only in female spiders. One (adult) or two (subadult) pairs of major ampullate glands send secretory ductules to the anterior spinnerets, and another pair of minor ampullate glands supply the median spinnerets. Three pairs of tubuliform glands in female spiders send secretory ductules to the median (one pair) and posterior (two pairs) spinnerets. Furthermore, one pair of flagelliform glands and two pairs of aggregate glands together supply the posterior spinnerets, and form a characteristic spinning structure known as a “triad” spigot. In male spiders, this combined apparatus of the flagelliform and the aggregate spigots for capture thread production is not apparent, instead only a non‐functional remnant of this triad spigot is present. In addition, the aciniform glands send ductules to the median (two pairs) and the posterior spinnerets (12–16 pairs), and the pyriform glands feed silk into the anterior spinnerets (90–100 pairs in females and 45–50 pairs in males).     

Synchronized and rhythmical activity during the prey capture in the social spiderAnelosimus eximius (Araneae,Theridiidae)

Summary Anelosimus eximius is a social spider species of South America. Many individuals share the same web and participate in prey capture, taking some ten seconds to locate the prey in the silky structures. In the laboratory, we analyzed the movements of each spider which took part in the pursuit, and showed that they were both synchronized and rhythmical. Spiders alternate simultaneous periods of immobility (involving 100% of the attacking individuals) and activity (involving at least 70% of the spiders).The results are discussed with reference to the model developed by Goss and Deneubourg (1988) suggesting that autocatalysis may be the motor of certain synchronized and rhythmical activities in social arthropods.     

Prey capture by the crab spider Misumena calycina (Araneae: Thomisidae)

Summary Crab spiders Misumena calycina (L.) in pasture rose Rosa carolina flowers regularly attacked bumble bees, smaller bees, and syrphid flies that visited these flowers. Attacks reached a maximum rate of over 20/h during mid morning, but only 1.6% of the most important prey item, bumble bees, were captured. The next most important food source, the most frequently taken item, syrphid flies Toxomerus marginatus (Say), were captured in 39% of the attempts. Since these flies have a biomass only 1/60th that of bumble bees, they comprised a much less important food source than did bumble bees. Spiders would obtain over 7% more food by specializing on bumble bees than by attacking all insect visitors, and as much as 20% more food at certain times of the day. However, they did not show a tendency to specialize at any time.     

Pollen structure, tetrad cohesion and pollen-connecting threads in Pseuduvaria (Annonaceae)

The structure of the pollen of 42 species of Pseuduvaria (Annonaceae) is described. The pollen is consistently inaperturate, isopolar and radially symmetrical. Four basic patterns of exine sculpturing are identified: rugulate, verrucate, scabrate and psilate. The exine stratification of one representative species, P. macrocarpa , is shown to be entirely ectexinal. The ectexine consists of a discontinuous outer tectal layer, a columellar infratectal layer, and an inner lamellar foliated foot layer; the intine is very thin and fibrillar. The pollen is invariably released as acalymmate tetrads, in which the tectum is absent from the proximal walls. The individual pollen grains within the tetrads are connected by crosswall cohesion, involving both exine and intine; this form of cohesion has not hitherto been reported in the Annonaceae. In addition, pollen grains of neighbouring tetrads are connected in two different ways, viz. short exine connections and non-sporopollenin pollen-connecting threads. Neither of these cohesion mechanisms has previously been reported for the genus. The function of the various forms of cohesion between pollen grains and tetrads in Pseuduvaria is discussed as a mechanism to enhance the efficiency of pollination by enabling the fertilization of multiple ovules following a single pollinator visit.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 69−78.     

On the spider genus Amaurobius (Araneae, Amaurobiidae) in India and Nepal

A new species, Amaurobius koponenisp. n., is described from Himachal Pradesh on the basis of a male specimen. A key to all five genera of Amaurobiidae that occur in Asia is provided. Four species from India and Nepal incorrectly assigned to Amaurobius are transferred to three genera of Titanoecidae: Anuvinda milloti (Hubert, 1973), comb. n., Pandava andhraca (Patel & Reddy, 1990), comb. n., Pandava nathabhaii (Patel & Patel, 1975), comb. n., and Titanoeca sharmai (Bastawade, 2008), comb. n.     

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.     

Ultrastructure and function of the adhesion–capture apparatus of Stenus species (Coleoptera, Staphylinidae)

 The elongated labium of rove beetles of the genus Stenus forms an adhesion–capture apparatus that enables the animal to catch fast fleeing prey, for example, collemboles. Structural details of this labium have been reinvestigated by using transmission electron microscopy, and the functional model of the capture mechanism has been refined. The openings of glandular ductules have been found to be located at the outer margin of the sticky cushions formed by the paraglossae. These cushions can be expanded by hemolymph pressure and are compressed when the tip of the protrusible labium hits the prey. Endocuticular fibers stabilize the cushions internally and determine the shape of the cushions in both the expanded and the compressed state. Within the membranous connecting tube that connects the prementum with the head capsule, the existence of an extra inner membrane has been confirmed. It is formed by a portion of the epidermis that has become detached from the cuticle. The most important part of the functional model of the protrusion of the labium is that the membranous connecting tube turns itself inside out distally, but the extra inner membrane does so proximally. During protrusion of the labium by hemolymph pressure, the prementum is accelerated during the initial phase of the process, and the nerves, muscles, tracheae, and glandular ductules that are attached to it are passively drawn into the labium. The mechanoreceptive function of setae at the distal end of the prementum has been confirmed on the basis of their ultrastructural characteristics. Accepted: 4 September 1998