Dispersal history of a spider (Stegodyphus lineatus) across contiguous deserts: vicariance and range expansion

Israel marks a crossroads between three continents encompassing several phytogeographical and zoogeographical zones. In this complex area, the flow of species from different biogeographical regions creates opportunities to study how geographical division and colonization routes affect current distribution and structure of resident populations of organisms associated with desert and arid environments, habitats that may have persisted throughout Pleistocene glacial periods. The present paper analyses the population history of the spider Stegodyphus lineatus in the contiguous Negev and Judean deserts in Israel using allozyme and mtDNA variation. The distinct patterns of variation indicate that Judean and Negev populations are vicariant lineages. The residence time was longer in Judea, where populations were more polymorphic for mtDNA, showed isolation by distance and were less structured than in the Negev. The Negev population, possibly linked to other Mediterranean populations of S. lineatus , consisted of two subdivisions derived from a recent eastward expansion across the central Negev watershed. Despite differences in age and level of structure, all lineages show similar dispersal processes dominated by restricted gene flow. The distribution patterns of allozyme and mtDNA markers are unrelated to geographical patterns of precipitation and vegetation. Rather, they follow large-scale topographic features, namely the water divide between Mediterranean and Afro-Syrian rift drainages and between eastern and western Negev drainages.   © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 739–754.     

Phylogeographical history of the sponge Crambe crambe (Porifera, Poecilosclerida): range expansion and recent invasion of the Macaronesian islands from the Mediterranean Sea

We studied sequence variation in the nuclear ribosomal internal transcribed spacers (ITS-1 and ITS-2) in 111 individuals from 11 populations/localities of the sponge Crambe crambe across the core species range in the western Mediterranean Sea and Atlantic Ocean. We report the first confirmed instance of intragenomic variation in sponges. Phylogeographical, nested clade and population genetic analyses were used to elucidate the species' evolutionary history. The study revealed highly structured populations affected by restricted gene flow and isolation-by-distance. A contiguous range expansion in the whole distribution area of the sponge was inferred. Phylogenetic analyses indicate a recent origin of most sequence types that could be explained by a recent origin of the species or a by recent bottleneck event in the studied area. A recent expansion of the distribution range to the Macaronesian region from the Mediterranean Sea was also detected, suggesting that C. crambe was recently introduced from the Mediterranean Sea to the Atlantic Ocean via human-mediated transport, and that the pattern observed is not the result of a natural biogeographical relationship between these zones.     

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.     

Fistularia commersonii in the Mediterranean Sea: invasion history and distribution modeling based on presence-only records

The bluespotted cornetfish (Fistularia commersonii) (Osteichtyes, Fistulariidae) is considered to be one of the most invasive species of the Mediterranean Sea and Europe but only scattered information exists on its distribution and abundance. Here we collated the available species records, following its first detection in the Mediterranean Sea, in January 2000, until October 2011. A total of 191 observations were used to reconstruct the invasion sequence, to provide estimates of the rate of spread and to construct an environmental suitability model based on six biophysical variables and the maximum entropy approach. The results showed that colonization of the Mediterranean Sea proceeded in parallel along the southern and northern rim of the Basin at speeds that reached 1,000–1,500 km year^?1 with a clear decrease in the rate of spread at the Sicily Strait. The most important explanatory variables for describing the distribution of F. commersonii were mean depth (explaining 32.4 % of the data variance), chlorophyll-a (29.3 %), and salinity (18.4 %). Coastal areas with relatively low chlorophyll-a concentrations and high salinity were the preferred habitat of the bluespotted cornetfish in its invaded range. Conversely, extreme productivity (highly eutrophic or highly oligotrophic), low salinity and cold temperatures provided abiotic resistance to this invasion. Areas of high environmental suitability were identified along the northern coasts of the Levantine Sea, Dodecanese, Sicily Strait and Tyrrhenian Sea. In contrast, the north Aegean Sea, the Adriatic and the Alboran Sea, the Nile Delta, the western coasts of Egypt and Cyrenaica were unfavourable for the invasion. Despite some limits due to the model’s resolution scale, these general predictions provide new insights into the F. commersonii invasion, indicating abiotic factors of primary importance in shaping the distribution of this species in its invaded range.     

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.     

Test of colonisation scenarios reveals complex invasion history of the red tomato spider mite Tetranychus evansi

The spider mite Tetranychus evansi is an emerging pest of solanaceous crops worldwide. Like many other emerging pests, its small size, confusing taxonomy, complex history of associations with humans, and propensity to start new populations from small inocula, make the study of its invasion biology difficult. Here, we use recent developments in Approximate Bayesian Computation (ABC) and variation in multi-locus genetic markers to reconstruct the complex historical demography of this cryptic invasive pest. By distinguishing among multiple pathways and timing of introductions, we find evidence for the "bridgehead effect", in which one invasion serves as source for subsequent invasions. Tetranychus evansi populations in Europe and Africa resulted from at least three independent introductions from South America and involved mites from two distinct sources in Brazil, corresponding to highly divergent mitochondrial DNA lineages. Mites from southwest Brazil (BR-SW) colonized the African continent, and from there Europe through two pathways in a "bridgehead" type pattern. One pathway resulted in a widespread invasion, not only to Europe, but also to other regions in Africa, southern Europe and eastern Asia. The second pathway involved the mixture with a second introduction from BR-SW leading to an admixed population in southern Spain. Admixture was also detected between invasive populations in Portugal. A third introduction from the Brazilian Atlantic region resulted in only a limited invasion in Europe. This study illustrates that ABC methods can provide insights into, and distinguish among, complex invasion scenarios. These processes are critical not only in understanding the biology of invasions, but also in refining management strategies for invasive species. For example, while reported observations of the mite and outbreaks in the invaded areas were largely consistent with estimates of geographical expansion from the ABC approach, historical observations failed to recognize the complex pathways involved and the corresponding effects on genetic diversity.     

The black widow spider genus Latrodectus (Araneae: Theridiidae): phylogeny, biogeography, and invasion history

The spider genus Latrodectus includes the widely known black widows, notorious because of the extreme potency of their neurotoxic venom. The genus has a worldwide distribution and comprises 30 currently recognized species, the phylogenetic relationships of which were previously unknown. Several members of the genus are synanthropic, and are increasingly being detected in new localities, an occurrence attributed to human mediated movement. In particular, the nearly cosmopolitan range of the brown widow, Latrodectus geometricus, is a suspected consequence of human transport. Although the taxonomy of the genus has been examined repeatedly, the recognition of taxa within Latrodectus has long been considered problematic due to the difficulty associated with identifying morphological features exhibiting discrete geographic boundaries. This paper presents, to our knowledge, the first phylogenetic hypothesis for the Latrodectus genus and is generated from DNA sequences of the mitochondrial gene cytochrome c oxidase subunit I. We recover two well-supported reciprocally monophyletic clades within the genus: (1) the geometricus clade, consisting of Latrodectus rhodesiensis from Africa, and its is sister species, the cosmopolitan L. geometricus, and (2) the mactans clade containing all other Latrodectus species sampled, including taxa occurring in Africa, the Middle East, Iberian Peninsula, Australia, New Zealand, and North and South America. Recovery of the geometricus and mactans clades is consistent with previous designations of species groups within the genus based on female genitalic morphology. All L. geometricus sampled, consisting of specimens from Africa, Argentina, North America, and Hawaii, were recovered as a strongly supported monophyletic group with minimal amounts of genetic divergence, corroborating the hypothesis that human transport has recently expanded the range of this species.     

Niche modelling of the Chilean recluse spider Loxosceles laeta and araneophagic spitting spider Scytodes globula and risk for loxoscelism in Chile

In Chile, all necrotic arachnidism is attributed to the Chilean recluse spider Loxosceles laeta (Nicolet) (Araneae: Sicariidae). It is predated by the spitting spider Scytodes globula (Nicolet) (Araneae: Scytodidae). The biology of each of these species is not well known and it is important to clarify their distributions. The aims of this study are to elucidate the variables involved in the niches of both species based on environmental and human footprint variables, and to construct geographic maps that will be useful in estimating potential distributions and in defining a map of estimated risk for loxoscelism in Chile. Loxosceles laeta was found to be associated with high temperatures and low rates of precipitation, whereas although S. globula was also associated with high temperatures, its distribution was associated with a higher level of precipitation. The main variable associated with the distribution of L. laeta was the human footprint (48.6%), which suggests that this is a highly invasive species. Similarly to other species, the distribution of L. laeta reaches its southern limit at the Los Lagos region in Chile, which coincides with high levels of precipitation and low temperatures. The potential distribution of L. laeta in Chile corresponds to the distribution of cases of loxoscelism.     

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.     

The effects of landscape history and time-lags on plant invasion in Mediterranean coastal habitats

Human-driven landscape changes may promote plant invasions by increasing propagule pressure and providing favourable conditions for non-native species to establish and spread. The increase in invasion levels might not be immediate but rather exhibit a time-lag (i.e. invasion debt). Moreover, the relationship between invasion and landscape history (i.e. changes in landscape composition) might be extremely complex as it might also include extinction debts of invasive species currently in regression. In order to understand the effect of landscape history on plant invasion, we studied the invasion level in coastal plant communities affected by intense human-driven landscape alteration in the last 50 years. We identified all non-native plant species within 809 grid cells (250 × 250 m) along ~500 km of the Spanish SW coast. We tested the association of (1) non-native richness conditional on the overall presence of non-natives (at least one species), and (2) the occurrence of the most frequent non-native species, with the percentage of human land-cover categories in 1956, 1991 and 2007 using a multimodel inference approach. We used these models to project future invasion patterns in the region. We found non-native richness to be more associated with land-cover variables in 1956 than in 2007, suggesting the existence of an invasion debt, but not for the overall presence of non-natives. For most frequent species, the effects of past landscape alteration depended on the introduction pathway and the region of origin. Species used in agriculture were more related to past cropland area, while most of the species used in gardening or forestry showed higher affinity for recently altered areas. These results highlight the relevance of the species’ introduction history and landscape history in assessing future long-term invasion trends.     

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.     

Trophic structure of the spider community of a Mediterranean citrus grove: A stable isotope analysis

Spiders are dominant terrestrial predators that consume a large variety of prey and engage in intraguild predation. Although the feeding habits of certain species are well known, the trophic structure of spider assemblages still needs to be investigated. Stable isotope analysis enables characterisation of trophic relationships between organisms because it tracks the energy flow in food webs and indicates the average number of trophic transfers between a given species and the base of the web, thus being a useful tool to estimate the magnitude of intraguild predation in food webs. Using this technique, we studied the trophic groups of spiders and their links within the arthropod food web of a Mediterranean organic citrus grove. We assessed the trophic positions of the 25 most common spider species relative to other arthropod predators and potential prey in the four seasons of the year, both in the canopy and on the ground. The analyses showed great seasonal variation in the isotopic signatures of some arthropod species, as well as the existence of various trophic groups and a wide range of trophic levels among spiders, even in species belonging to the same family. Differences in δ¹⁵N between spiders and the most abundant prey in the grove usually spanned two trophic levels or more. Our findings provide field evidence of widespread intraguild predation in the food web and caution against using spider families or guilds instead of individual species when studying spider trophic interactions.     

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.