Scavenging by spiders (Araneae) and its relationship to pest management of the brown recluse spider

Experiments reported in Sandidge (2003; Nature 426: 30) indicated that the brown recluse spider, Loxosceles reclusa Gertsch & Mulaik, preferred to scavenge dead prey over live prey and that the spiders were not detrimentally affected when fed insecticide-killed crickets. Extrapolations made in subsequent media coverage disseminating the results of this research made counter-intuitive statements that pesticide treatment in houses would increase brown recluse populations in homes. This information was presented as if the scavenging behavior was specialized in the brown recluse; however, it was more likely that this behavior has not been well studied in other species. To provide a comparison, the current laboratory study examined the likelihood of non-Loxosceles spiders to scavenge dead prey. Of 100 non-Loxosceles spiders that were tested (from 11 families, 24 genera, and at least 29 species from a variety of spider hunting guilds), 99 scavenged dead crickets when offered in petri dishes. Some of the spiders were webspinners in which real-world scavenging of dead prey is virtually impossible, yet they scavenge when given the opportunity. Therefore, scavenging is a flexible opportunistic predatory behavior that is spread across a variety of taxa and is not a unique behavior in brown recluses. These findings are discussed in relation to pest management practices.     

Tracking a medically important spider: climate change, ecological niche modeling, and the brown recluse (Loxosceles reclusa)

Most spiders use venom to paralyze their prey and are commonly feared for their potential to cause injury to humans. In North America, one species in particular, Loxosceles reclusa (brown recluse spider, Sicariidae), causes the majority of necrotic wounds induced by the Araneae. However, its distributional limitations are poorly understood and, as a result, medical professionals routinely misdiagnose brown recluse bites outside endemic areas, confusing putative spider bites for other serious conditions. To address the issue of brown recluse distribution, we employ ecological niche modeling to investigate the present and future distributional potential of this species. We delineate range boundaries and demonstrate that under future climate change scenarios, the spider's distribution may expand northward, invading previously unaffected regions of the USA. At present, the spider's range is centered in the USA, from Kansas east to Kentucky and from southern Iowa south to Louisiana. Newly influenced areas may include parts of Nebraska, Minnesota, Wisconsin, Michigan, South Dakota, Ohio, and Pennsylvania. These results illustrate a potential negative consequence of climate change on humans and will aid medical professionals in proper bite identification/treatment, potentially reducing bite misdiagnoses.     

Hyaluronidase and esterase activities of the venom of the poisonous brown recluse spider

Venom of Loxosceles reclusa free from impurities was expressed from venom glands collected by microdissection. Polyacrylamide gel electrophoresis of the venom at pH 8.3 demonstrated 7 or 8 major plus 3 or 4 minor components. Upon electrophoresis at pH 4.9 two major components plus 3 or 4 minor components were noted. Monophoretic hyaluronidase prepared by Sephadex gel filtration and electrophoresis at pH 8.3 exhibited optimum activity from pH 5.0 to 6.6. Sodium dodecyl sulfate gel electrophoresis of purified hyaluronidase revealed two components with estimated molecular weights of 33,000 and 63,000. The purified hyaluronidase exhibited activity against chondroitin sulfate, types A, B, and C at approximately 20–30% of that upon hyaluronic acid. The enzyme was inhibited 10–20% by the heavy metal ions, Fe⁺³ and Cu⁺². Rabbit antivenom inhibited the spreading effect of whole venom in vivo and completely inhibited hyaluronidase in vitro.Incorporation of [¹⁴C]leucine into the spider venom led to the separation of hyaluronidase from the dermonecrotic activity of the venom.The venom demonstrated activity against carbobenzoxy-l-tyrosine-p-nitrophenyl ester and β-naphthylacetate which was inhibited approximately 65% by 2.5 × 10^?3m levels of EDTA and EGTA but not by 2.5 × 10^?4mo-phenanthroline. The esterase activity resisted concentrations of p-chloromercuribenzoate which totally inactivated papain. The venom appeared devoid of collagenase, dipeptidase, acetylcholinesterase, phosphodiesterase, ribonuclease A, and deoxyribonuclease.     

Daily thermal preference variation of the sand recluse spider Sicarius thomisoides (Araneae: Sicariidae)

Preferential temperature as a physiological feature is crucial for spiders, since it determines the selection of key habitats for their survival and reproduction. In this work, we study the daily and geographical variation of the preferential temperature of the spider Sicarius thomisoides subjected to different degrees of daily thermal oscillation in their habitats. Preferred temperatures differ between coastal and inland populations, but in both cases, there is a marked bimodality in the daily pattern of temperature preference, with two peaks per day that would be given by the changes in the hours of activity. These nocturnal spiders select higher temperatures in the evening (active period) and select lower temperatures during late morning (resting period). In laboratory, spiders have preferred temperatures that differ from those found in their habitats, so they must tolerate or compensate non-preferred temperatures by active thermoregulation in natural conditions.     

Secondary retinae of a primitive jumping spider,Yaginumanis (Arachnida,Araneida, Salticidae)

Summary Retinae of the secondary eyes of a primitive salticid spider, Yaginumanis sexdentatus (Yaginuma 1967) are described at the ultrastructural level. The structures of the anterior lateral, posterior lateral and posterior median eyes are identical. Receptor somata lie in the retinal cups. Each receptor bearing twin rhabdomeres is ensheated by (i) much-divided processes of non-pigmented glial cells whose somata lie distally in the retinal cups; and (ii) four processes of pigmented glial cells whose somata lie basally, below the receptive segments. Pigment granules in the latter are concentrated in the basal retina, and are not present at the level of the rhabdoms. The present findings support the placement of Yaginumanis in a newly erected Subfamily Spartaeinae by Wanless (1984), because of the likelihood of homology in the fine structural organisation of the secondary retinae of this genus and of the genus Portia.     

Proprioceptors and sensory nerves in the legs of a spider,Cupiennius salei (Arachnida,Araneida)

Summary Retrograde CoS-impregnation was used to trace and map the course of sensory nerves and the distribution and innervation of the various proprioceptor types in all leg segments of Cupiennius salei, a Ctenid spider.1. Sensory nerve branches. In both the tibia and femur, axons of all proprioceptor types ascend in just two lateral nerves which do not merge with the main leg nerve until they reach the next proximal joint region. In the short segments — coxa, trochanter, patella, and tarsus — axons of the internal joint receptors often run separately from those of the other sensilla. Axons of the large lyriform slit sense organ at the dorsal metatarsus and of the trichobothria join with only a few hair axons and form their own nerve branches (Figs. 1, 2, 3).2. Proprioceptors. Each of the seven leg joints is supplied with at least one set of the well-known internal joint receptors, slit sensilla (single slits and lyriform organs), and long cuticular hairs. In addition, we found previously unnoticed hair plates on both sides of the coxa, near the prosoma/coxa joint; they are deflected by the articular membrane during joint movements (Fig. 4).3. Sensory cells and innervation. CoS-impregnation shows that each slit of the slit sense organs — be it a single slit or several slits in a lyriform organ — is innervated by two bipolar sensory cells (Fig. 6). We also confirm previous reports of multiple innervation in the internal joint receptors and in the long joint hairs and cuticular spines.Most of the ascending nerve branches run just beneath the cuticle for at least a short distance (Fig. 5); hence they are convenient sites for electrophysiological recordings of sensory activity even in freely walking spiders.     

Red blood cell lysis induced by the venom of the brown recluse spider: the role of sphingomyelinase D.

Red blood cell lysis induced by the venom of Loxosceles reclusa, the brown recluse spider, may be related to the hemolytic anemia observed in several cases of spider envenomation. These investigations demonstrate that the venom of the brown recluse spider contains a calcium-dependent, heat-labile hemolysin of molecular weight approximately 19,000. The pH optimum for the hemolytic reaction was 7.1, and the optimum calcium concentration for venom-induced lysis was observed within the range of 6 to 10 mm. Sheep red blood cells were more susceptible to the spider hemolysin than human red blood cells, although both types exhibited appreciable lysis. Digestion of sheep red blood cell membranes with partially purified venom lysin resulted in degradation of the sphingomyelin component. However, reaction of the membranes with the venom lysin produced no release of water-soluble phosphate, and no free fatty acids were generated. These results indicate that the sphingomyelin-degrading activity of the venom is not a phospholipase C- or a phospholipase A₂-type activity. Sphingomyelin was employed as substrate for the venom hemolysin, and the organic and aqueous fractions of the reaction mixtures were analyzed by thin-layer chromatography. Analysis of the organic fraction revealed a phosphate-containing product with the solubility and chromatographic characteristics of N-acylsphingosine phosphate (ceramide phosphate), and analysis of the aqueous fraction demonstrated the presence of choline. The isolation and identification of these products indicate that the sphingomyelin of the red cell membrane is hydrolyzed by a sphingomyelinase D-type activity expressed by the partially purified venom hemolysin. A close correspondence between the hemolytic and sphingomyelinase D activities was observed when the partially purified hemolysin was further characterized in polyacrylamide gel electrophoresis at pH 8.3 and pH 4.9. The hemolytic and sphingomyelinase activities were coincident within the electrophoretic pattern at both pHs. The results presented demonstrate conclusively a direct lytic action of brown recluse venom upon red blood cells and report for the first time the presence of sphingomyelinase D in spider venom.     

Model studies on the mechanical significance of grouping in compound spider slit sensilla (Chelicerata,Araneida)

Summary The mechanical implications of various types of slit arrangements found among the strain-sensitive slit sensilla in the arachnid exoskeleton (Fig. 3) were studied by measuring the deformation of model slits, cut into plastic discs, under static load applied in the plane of the disc and from varying directions (Figs. 1, 2).1. Close parallel, lyriform arrangements. Compression of slits (adequate stimulus) reaches much higher values than dilatation. It is highest with the load direction at right angle to the slit axes. Also, in the majority of slits the range of load angles resulting in compression is considerably larger than that leading to dilatation. Length distribution and lateral shift of slits in the models have a pronounced effect on slit deformability (Figs. 4-5): (a) In the oblique bar arrangement with slits of equal length and regular lateral shift (Fig. 4A) deformation of all slits is very similar at all load directions. In all slits compression results from a range of load angles larger than 120°. (b) In arrangements with a regular increase in slit length and a triangular outline shape deformability differs greatly among the slits at all load angles (Fig. 4B). (c) The slit configuration with a heartshaped outline (Fig. 4C) is peculiar for the large spread of load angles at which the compression of the different slits is highest. — These properties recommend different arrangements for the solution of different strain measuring problems, with for instance, the particular need of a wide angular working range (arrangement a), of a large spectrum of absolute sensitivities (b), or of the analysis of load direction (c).2. Angle and distance between slits. Due to the mechanical directionality inherent in an elongated slit the divergence of slit axes within a group of slits is likely to indicate the importance of the analysis of strain direction (Fig. 6). The mechanical interaction between closely neighbouring slits decreases with their distance from each other. In a parallel arrangement of equally long slits it disappears if the distance is about 1.5 times the slit length (Fig. 7).3. Aiming towards a mechanical model which would explain the complex deformation found in a lyriform organ, we consider the outline of the organ as a hole traversed by beams of material. Slit deformation can be calculated from the elastic lines of the beams which separate the slits and information drawn from photoelastic experiments (Figs. 8-11).     

Biotechnological applications of brown spider (Loxosceles genus) venom toxins

Loxoscelism (the term used to define accidents by the bite of brown spiders) has been reported worldwide. Clinical manifestations following brown spider bites are frequently associated with skin degeneration, a massive inflammatory response at the injured region, intravascular hemolysis, platelet aggregation causing thrombocytopenia and renal disturbances. The mechanisms by which the venom exerts its noxious effects are currently under investigation. The whole venom is a complex mixture of toxins enriched with low molecular mass proteins in the range of 5–40 kDa. Toxins including alkaline phosphatase, hyaluronidase, metalloproteases (astacin-like proteases), low molecular mass (5.6–7.9 kDa) insecticidal peptides and phospholipases-D (dermonecrotic toxins) have been identified in the venom. The purpose of the present review is to describe biotechnological applications of whole venom or some toxins, with especial emphasis upon molecular biology findings obtained in the last years.     

Neuroanatomy of the central nervous system of the wandering spider,Cupiennius salei (Arachnida,Araneida)

Summary In Cupiennius salei (Ctenidae), as in other spiders, the central nervous system is divided into the supraoesophageal ganglion or brain and the suboesophageal ganglia (Fig. 1). The two masses are interconnected by oesophageal connectives. The brain gives off four pairs of optic and one pair of cheliceral nerves. From the suboesophageal ganglia arise a pair of pedipalpal, four pairs of leg, and several pairs of opisthosomal nerves (Fig. 2). 1. Cell types. In the brain a total of 50900 cells were counted, in the suboesophageal ganglia 49000. They are all monopolar cells, found in the ganglion periphery and may be classified into four types: (a) Small globuli cells (nuclear diameter 6–7 m) forming a pair of compact masses in the protocerebrum (Fig. 10b); (b) Small and numerous cells (cell diameter 12–20 m) with processes forming the bulk of the neuropil in the brain and suboesophageal ganglia; (c) Neurosecretory cells (cell diameter ca. 45 m) in the brain and suboesophageal ganglia; (d) Large motor and interneurons (cell daimeter 40–112 m), mostly in the suboesophageal ganglia (Figs. 10a and c). 2. Suboesophageal mass. The cell bodies form a sheet of one to several cell layers on the ventral side of each ganglion and are arranged in groups. Three such groups were identified as motor neurons, four as interneurons. At the dorsal, dorso-lateral, and mid-central parts of the ganglion there are no cell somata. The fibre bundles arising from them form identifiable transverse commissural pathways (Fig. 9b). They form the fibrous mass in the central part of the suboesophageal mass.Neuropil is well-formed in association with the sensory terminations of all major nerves (Fig. 9a). As these proceed centrally they break up into five major sensory tracts forming five layers one above the other. There are six pairs of additional major longitudinal tracts arranged at different levels dorsoventrally (Fig. 8). They ascend into the brain through the oesophageal connectives and terminate mostly in the mushroom bodies and partly in the central body. 3. Protocerebrum. Fine processes of the globuli cells form the most important neuropil mass in the fibrous core, called the mushroom bodies. These consist of well developed glomeruli, hafts, and bridge which are interconnected with the optic masses of the lateral eyes and most fibre tracts from the brain and suboesophageal mass (Fig. 7). The median eye nerves form a small optic lamella and optic ganglia, connected to the central body through an optic tract. Each posterior median and posterior lateral eye nerve ends in large optic lamellae (Fig. 13a). These are connected through chiasmata to a large optic mass where fibres from globuli cells form conspicuous glomeruli. There are 10–12 large fibres (diameter 9 m) of unknown origin on each side, terminating in the optic lambella of the posterior lateral eye.The central body, another neuropil mass (Fig. 13b) in the protocerebrum, is well developed in Cupiennius and located transversely in its postero-dorsal region (Fig. 10d). It consists of two layers and is interconnected with optic masses of the median and lateral eyes through optic tracts. Fibre tracts from the brain and suboesophageal mass join the central body.     

Observations on the leg receptors ofCiniflo (Araneida: Dictynidae)

Summary The curved, blunt-tipped hairs on the legs ofCiniflo have a structure characteristic of contact chemoreceptors. Using a hair tip recording technique, it has been possible to confirm that these sensilla do respond to contact stimulation by certain chemical substances (Figs. 1 and 3). A few experiments were also performed onTegenaria (Fig. 2). So far, positive responses to some monavalent salts (Figs. 1 and 2) and hydrochloric acid (Fig. 3) have been established, involving perhaps 5 to 6 chemoreceptor units in all. However, each sensillum is known to have 19 chemoreceptor cells and thus most of the reaction spectrum of the sensillum remains unknown. The suggestion that, in contrast to insect contact chemoreceptors (which usually have only 4–7 sensory units), some of the dendrites may be very specific receptor units and are perhaps involved in the detection of contact pheromones or other equally specific substances, is discussed.One of the authors (DJH) would like to thank the Science Research Council for a research studentship, during which this work was carried out. Thanks are also due to Mr. J. Scott, Mr. C. Gilbert and Mr. R. Stevenson for their excellent technical help.     

The immigration of spiders (Araneida) into a new polder

Abstract. (1) The immigration of spiders into the Lauwerszeepolder (constructed in 1969) was studied during four years in four areas. The spider fauna was sampled weekly and 65 000 specimens of eighty-four species were caught using strip traps, window traps, simple pitfall traps and fences.
(2) About sixty species from all kinds of habitats were caught rarely; nineteen species were caught in numbers that suggest that they had established populations in the study areas.
(3) The four successful pioneer species, plus two marsh-dwelling species, were most abundant during 1969 and 1970. The 1971 catch was dominated by species from saline habitats; they became less abundant in 1972 when species from non-saline habitats increased. Only halotolerant species established populations.
(4) No evidence was found that dispersing spiders select certain areas. The differences between the spider fauna of the differennt study areas arise from degrees of success in colonization by immigrants. Pioneer species are least influenced by the abiotic environment.
(5) A greater aeronautic dispersal power is found in species inhabiting unstable habitats than in species from stable habitats.
(6) Aeronautic activity in adult linyphiids (s.l.) is not restricted to a special season, but is related to definite phases of the phenology.
(7) Male and female erigonids are equally active in aeronautic behaviour. In erigonids and linyphiids intense aeronautic activity coincides with great ground activity; on the ground males are more active than females.