Effects of within‐colony competition on body size asymmetries and reproductive skew in a social spider

Reproductive partitioning is a key component of social organization in groups of cooperative organisms. In colonies of permanently social spiders of the genus Stegodyphus less than half of the females reproduce, while all females, including nonreproducers, perform suicidal allo‐maternal care. Some theoretical models suggest that reproductive skew is a result of contest competition within colonies, leading to size hierarchies where only the largest females become reproducers. We investigated the effect of competition on within‐group body size variation over six months in S. dumicola, by manipulating food level and colony size. We found no evidence that competition leads to increased size asymmetry within colonies, suggesting that contest competition may not be the proximate explanation for reproductive skew. Within‐colony body size variation was high already in the juvenile stage, and did not increase over the course of the experiment, suggesting that body size variation is shaped at an early stage. This might facilitate task specialization within colonies and ensure colony‐level reproductive output by early allocation of reproductive roles. We suggest that reproductive skew in social spiders may be an adaptation to sociality selected through inclusive fitness benefits of allo‐maternal care as well as colony‐level benefits maximizing colony survival and production.     

Argas (Persicargas) persicus (Oken, 1818) (Ixodida: Argasidae) in Sicily with considerations about its Italian and West-Mediterranean distribution

Recently, in the province of Trapani (Western Sicily), some overwintering specimens of the argasid tick Argos (Persicargas) persicus (Oken, 1818) were observed and collected. Morphological and genetic analysis were utilized in order to reach a definitive identification. The species was found in two semi-natural sites where, having been found repeatedly, its presence does not appear accidental. Moreover the characteristics of the Sicilian findings seem to exclude a human-induced spread. This record, the first regarding Sicily and South Italy, is discussed together with the previous doubtful citations for Italy. These findings revalue not only all the old citations for Italy but also the hypothesis that the Mediterranean distribution of this argasid is of a natural origin.     

Reproductive state and care giving in Stegodyphus (Araneae: Eresidae) and the implications for the evolution of sociality

Cooperative brood care is a rare phenomenon in spiders and is restricted to a few social species, including three in the genus Stegodyphus. Brood care in Stegodyphus begins with regurgitation feeding followed by matriphagy: the young consume the body fluids of their mother causing her to die quickly. Whether such an extreme form of maternal care can become a communal task should depend on physiological or historical preconditions. I investigated whether femaleStegodyphus lineatus feed young or allow matriphagy according to their own reproductive state. Broods of young of two age classes (2 or 10 days after hatching) were isolated or fostered out to adult females that were unmated, had eggs or had young. Growth and survival of females and broods were followed over 21 days. The timing of matriphagy depended on the interaction between age of young and state of the foster mother. All broods that were fostered out to females with young grew and survived. Two-day-old young did not survive when isolated or fostered out to unmated females, but some survived and gained weight when placed with foster mothers that cared for egg sacs. Young of 10 days of age grew when fostered out to females with eggs but did not grow or lost weight when isolated or fostered out to unmated females. Survival among 10-day spiderlings was relatively high in all groups but differed significantly between treatments (young isolated or fostered out to unmated females or females with eggs) and control (left with the mother). The results show that these spiders will care for young from other females only when they are in the right developmental state. Such a constraint can have important consequences for the evolution of allomaternal care in social species: unless such a mechanism is overcome, nonreproductives cannot help in brood care. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Intersexual cooperation during male clasping of external female genitalia in the spider Physocyclus dugesi (Araneae, Pholcidae)

Sexual conflict may influence the shape and evolution of body structures that males use to grasp females during mating. Not only sexual coercion but also intersexual cooperation may be involved during clasping behavior. Among pholcid spiders, secondary sexual modifications of the male chelicerae, such as apophyses with spines or tooth-like processes, function to grasp the female by specific parts of her external genitalia such as grooves or apophyses of the epigynum. We analyzed how the female and the male respond when their structures for clasping are experimentally modified in the pholcid Physocyclus dugesi. We used three treatment groups for virgin females that differed in the manipulation of the epigynum apophyses (uncovered, partially covered, and fully covered by a plaster) and two groups of males (uncovered and fully covered cheliceral apophyses). We found that females are mainly cooperative to courting males not only when the female genital apophyses were experimentally covered but also when the male cheliceral apophyses were covered. The current data also indicate behavioral flexibility in males during courtship, especially when they had difficulty in genital intromission. Our experimental results, together with previous observational studies, support a modulated-cooperative scenario between the sexes for cheliceral clasping and genital intromission in pholcid spiders.     

Ticks Feeding on Humans: A Review of Records on Human-Biting Ixodoidea with Special Reference to Pathogen Transmission

In this article, literature records of argasid and ixodid ticks feeding on humans worldwide are provided in view of increased awareness of risks associated with tick bites. Ticks can cause paralyses, toxicoses, allergic reactions and are vectors of a broad range of viral, rickettsial, bacterial and protozoan pathogens. Approximately 12 argasid species (Argas and Ornithodos) are frequently found attached to humans who intrude into tick-infested caves and burrows. Over 20 ixodid tick species are often found on humans exposed to infested vegetation: four of these are Amblyomma species, 7 Dermacentor spp., 3 Haemaphysalis spp., 2 Hyalomma spp. and 6 Ixodes species. Personal protection methods, such as repellents and acaricide-impregnated clothing are advised to minimize contact with infected ticks. Acaricidal control of ixodid ticks is impractical because of their wide distribution in forested areas, but houses infested with soft ticks can be sprayed with acaricidal formulations. Attached ticks should be removed without delay. The best way is to grasp the tick as close to the skin as possible with fine tweezers and pull firmly and steadily without twisting. Finally, despite the fact that most people who are bitten destroy the offending tick in disgust, it is recommended that they preserve specimens in ethanol for taxonomic identification and detection of pathogens by molecular methods.     

Group living and inbreeding depression in a subsocial spider

Social spiders are unusual among social organisms in being highly inbred-males and females mature within their natal nest and mate with each other to produce successive generations. Several lines of evidence suggest that in spiders inbred social species originated from outbred subsocial ancestors, a transition expected to have been hindered by inbreeding depression. As a window into this transition, we examined the fitness consequences of artificially imposed inbreeding in the naturally outbred subsocial spider Anelosimus cf. jucundus. Subsocial spiders alternate periods of solitary and social living and are thought to resemble the ancestral system from which the inbred social species originated. We found that inbreeding depression in this subsocial spider only becomes evident in spiders raised individually following the end of their social phase and that ecological and demographic factors such as eclosion date, number of siblings in the group and mother's persistence are more powerful determinants of fitness during the social phase. A potential explanation for this pattern is that maternal care and group living provide a buffer against inbreeding depression, a possibility that may help explain the repeated origin of inbred social systems in spiders and shed light on the origin of other systems involving regular inbreeding.     

Description of adults and nymph, and redescription of the larva, of Ornithodoros marinkellei (Acari:Argasidae), with data on its phylogenetic position

The argasid tick Ornithodoros marinkellei Kohls, Clifford, and Jones, 1969 was described 4 decades ago based on larval specimens collected from bats (Pteronotus spp.) in Colombia and Panama. Thereafter, larval O. marinkellei parasitizing bats were reported from Venezuela, Guyana, and Brazil. Herein, we describe the adults and nymph, and redescribe the larva of O. marinkellei based on specimens recently collected in the western Brazilian Amazon region. In contrast to all other known adult argasids, the idiosoma of both males and females of O. marinkellei is covered with sclerotized plaques. The idiosoma of the nymph of O. marinkellei is entirely micromamillated, and differs from the adults by the absence of plaques. The larva of O. marinkellei is morphologically similar to the larvae of the 2 other species belonging to the subgenus Subparmatus , i.e., Ornithodoros viguerasi Cooley and Kohls, 1941 and Ornithodoros mormoops Kohls, Clifford, and Jones, 1969 . Because of the long and narrow dorsal plate, the larva of O. marinkellei is readily distinguished from O. viguerasi and O. mormoops. Comparison of our larvae from Brazil with O. marinkellei paratype specimens from Colombia confirmed their taxonomic identification. However, a few morphological differences, particularly in the size of the gnathosoma, were observed. Further studies are necessary to clarify whether O. marinkellei is a complex of different species, or a single species represented by morphologically polymorphic, and geographically distinct populations. Partial mitochondrial 16S rDNA gene sequences were generated for O. marinkellei specimens from Brazil, and compared with available homologous sequences in GenBank. Phylogenetic analyses revealed O. marinkellei to be distinct from the remaining argasid species available in GenBank, including other bat-associated tick species that are found in sympatry with O. marinkellei in the Neotropical region.     

New reports of Antricola guglielmonei and Antricola delacruzi in Brazil, and a description of a new argasid species (Acari)

Adults of 3 tick species (Acari: Argasidae) identified as Antricola guglielmonei, Antricola delacruzi, and Carios rondoniensis n. sp. were collected on bat guano in a cave in the state of Rond?nia, western Amazon, Brazil. Adults of C. rondoniensis possess a unique combination of characters that distinguish them from all described adults in the Argasidae, i.e., a large spiracular plate densely filled with small goblets, a well-developed flap covering the female genital opening, and palpi containing several tufts of long setae on articles 2 and 3. Unlike Ornithodoros or other Carios species, adults of C. rondoniensis have a scooplike hypostome devoid of denticles, as in Antricola spp. Conversely, the presence of a pair of long posthypostomal setae, and a slitlike transverse fissure at the capsule opening of the Haller's organ, are characters of C. rondonensis that are also found in species of Carios and Ornithodoros, but not in Antricola species. Molecular analyses inferred from a portion of the 16S rRNA mitochondrial gene indicate that C. rondoniensis is phylogenetically closest to species of Carios, followed by species of Antricola, and then Ornithodoros. Because the highest bootstrap value linking C. rondoniensis to Carios spp. was 62%, further phylogenetic studies are needed to better evaluate the taxonomic status of the former species.     

Maternal care and subsocial behaviour in spiders

While most spiders are solitary and opportunistically cannibalistic, a variety of social organisations has evolved in a minority of spider species. One form of social organisation is subsociality, in which siblings remain together with their parent for some period of time but disperse prior to independent reproduction. We review the literature on subsocial and maternal behaviour in spiders to highlight areas in which subsocial spiders have informed our understanding of social evolution and to identify promising areas of future research. We show that subsocial behaviour has evolved independently at least 18 times in spiders, across a wide phylogenetic distribution. Subsocial behaviour is diverse in terms of the form of care provided by the mother, the duration of care and sibling association, the degree of interaction and cooperation among siblings, and the use of vibratory and chemical communication. Subsocial spiders are useful model organisms to study various topics in ecology, such as kin recognition and the evolution of cheating and its impact on societies. Further, why social behaviour evolved in some lineages and not others is currently a topic of debate in behavioural ecology, and we argue that spiders offer an opportunity to untangle the ecological causes of parental care, which forms the basis of many other animal societies.     

A reassessment of argasid tick salivary gland ultrastructure from an immuno-cytochemical perspective

Previous morphological and histochemical studies of argasid tick salivary glands indicated that they were less complex than ixodid salivary glands, with only three granular cell types. The present study shows that there exist at least four different granular cell types in the salivary glands of the argasid tick Ornithodoros savignyi, based on immuno-localization of the anti-hemostatic factors, apyrase and savignygrin. Both anti-hemostatic factors were localized to dense core granule type 'a' and to granule type 'b', that shares a similar homogenous morphology with non-labeled granule type 'd'. Furthermore, the major tick salivary gland proteins (TSGPs), previously implicated in granule biogenesis, were localized to all the granular cell types. This indicates that granular cell types with different morphologies can express the same proteins, while cell types that show similar morphologies may not express the same proteins. Argasid tick salivary glands seem to be more complex than previously thought and might not be amenable to morphological classification alone. Alternative classification methodologies that rely on physical expression patterns of the salivary gland proteome might be more reliable as markers for a specific granular cell type.     

Tick species infesting ruminants in Italy: ecological and bio-climatic factors affecting the different regional distribution

The distribution of tick species able to infest ruminants in Italy is briefly reviewed on the basis of published records and new data obtained in several surveys carried out mainly in northern Italy. The most frequent and abundant tick species found on ruminants throughout Italy are Ixodes ricinus and Dermacentor marginatus, the first gradually replacing its association with sheep by the association with wild ruminants. Rhipicephalus bursa was mainly found on cattle while R. turanicus, which is considered one of the most sheep-associated species, was found only on wild boar. Haemaphysalis punctata, H. sulcata and H. parva are the other tick species quite frequently found on small ruminants when they are reared on pastures mainly in central-southern Italy. However, the land exploitation of these last decades has dramatically reduced the diversity of Italian environment and significantly modified the distribution and the abundance of ticks causing a heavy spread of the tick species characterized by a strong adaptability such as I. ricinus and R. sanguineus. If this trend continues, it is possible to hypothesise that few "resistant" tick species will gradually replace most of the others. This hypothesis seems to be confirmed by the high percentage observed for I. ricinus (57.4%) and R. sanguineus (35.73%) out of 8,712 specimens collected from several different hosts and from the environment.     

The biology of Portia fimbriata, a web-building jumping spider (Araneae, Salticidae) from Queensland: intraspecific interactions

Portia fimbriata (Doleschall) is an unusual salticid because it spins webs and uses its own webs and those of other species in predation. However, the courtship and threat displays of this species are more like those of typical, cursorial salticids than like typical web-building spiders. During male-female interactions, males perform leg-waving and leg-shaking displays, with legs I and II extended stiffly forward, while distant from the females. Before mounting they tap the females with their legs; and after mounting they tap, scrape and stroke the females' abdomens. Copulation takes place on or near the female's web or on the webs of other species. Hunched-legs displays, with legs I–III highly flexed and held to the side of the body, occur in male-male interactions. Also, during male-female and female-female interactions, females perform hunched-legs displays, strike, charge, ram, embrace, grapple with and leap at conspecifics. Sometimes they lose legs while grappling. After female-female interactions associated with maternal webs, if the resident decamps, the intruder eats the eggs left behind on a suspended leaf, spins a new egg-case over the destroyed one, and oviposits. Adult and sub-adult males co-habit in webs with sub-adult females for as long as 48 days and mate when the spiders mature.     

Task specialization in two social spiders,Stegodyphus sarasinorum (Eresidae) and Anelosimus eximius (Theridiidae)

Understanding the social organization of group‐living organisms is crucial for the comprehension of the underlying selective mechanisms involved in the evolution of cooperation. Division of labour and caste formation is restricted to eusocial organisms, but behavioural asymmetries and reproductive skew is common in other group‐living animals. Permanently, social spiders form highly related groups with reproductive skew and communal brood care. We investigated task differentiation in nonreproductive tasks in two permanently and independently derived social spider species asking the following questions: Do individual spiders vary consistently in their propensity to engage in prey attack? Are individual spiders' propensities to engage in web maintenance behaviour influenced by their previous engagement in prey attack? Interestingly, we found that both species showed some degree of task specialization, but in distinctly different ways: Stegodyphus sarasinorum showed behavioural asymmetries at the individual level, that is, individual spiders that had attacked prey once were more likely to attack prey again, independent of their body size or hunger level. In contrast, Anelosimus eximius showed no individual specialization, but showed differentiation according to instar, where adult and subadult females were more likely to engage in prey attack than were juveniles. We found no evidence for division of labour between prey attack and web maintenance. Different solutions to achieve task differentiation in prey attack for the two species studied here suggest an adaptive value of task specialization in foraging for social spiders.     

Tick cell lines: tools for tick and tick-borne disease research

Over 40 cell lines are currently available from 13 ixodid and one argasid tick species. The successful isolation and propagation of several economically important tick-borne pathogens in tick cell lines has created a useful model to study interactions between tick cells and these viral and bacterial disease agents. Tick cell lines have already proved to be a useful tool in helping to define the complex nature of the host-vector-pathogen relationship. With the availability of genomics tools, tick cell lines will become increasingly important as a complement to tick and tick-borne disease research in vivo once genetic transformation and gene silencing using RNA interference become routine.     

Species composition, distribution, and ecological preferences of the ticks of grazing sheep in north-central Spain

The distribution and ecological preferences of tick (Acari: Ixodidae) parasites of grazing sheep in the region of Aragón (north-central Spain) were surveyed on flocks four times a year and mapped into a 5 x 5 km grid. Nine tick species were found. These were species of the Rhipicephalus sanguineus group (about 95% of them Rhipicephalus turanicus Pomerantsev, in 91% of cells of the grid), Rhipicephalus bursa Canestrini & Fanzago (79% of cells), Dermacentor marginatus (Sulzer) (58% of cells), Haemaphysalis punctata Canestrini & Fanzago (74% of cells) and Ixodes ricinus (Linnaeus) 14% of cells. Other species weakly represented in the surveys were Dermacentor reticulatus (Fabricius), Haemaphysalis sulcata Canestrini & Fanzago and Hyalomma m. marginatum Koch. Data on temperature, Normalized Difference Vegetation index (NDVI), topography, vegetation categories and plant productivity were used to build models of distribution and abundance of D. marginatus, H. punctata, R. bursa and R. turanicus. The occurrence models largely incorporated climatic variables and had good discrimination ability (P < 0.0001 for every modelled species, correct classification rate or sensitivity within 0.89 and 0.99), whereas the abundance models had a lower explanatory power. These models are relevant in the understanding of the variables composing the main distribution patterns, but they are unable adequately to predict the density. Abundance models produce good predictions in cells with low tick density, whereas poor correlation is observed in sites with high tick abundance. Several causes may be responsible for this low predictive power of the abundance models. Model output might be sensible to host density, to local farming practices, or to the size of the grid used to refer the results of the survey. In the latter case, small patches may support locally important populations of ticks, influencing largely the results of the survey. These patches of particular abiotic conditions, or supporting large host densities, may have been undetected at the resolution of the survey, thus obscuring the impact of the predictive variables.     

Immunocytochemical mapping of FMRFamide-like peptides in the argasid tick Ornithodoros parkeri and the ixodid tick Dermacentor variabilis

FMRFamide-like immunoreactivity was studied in the argasid tick Ornithodoros parkeri and the ixodid tick Dermacentor variabilis using immunocytochemistry based on the peroxidase-antigeroxidase method. FMRFamide-like immunoreactive cells are widely distributed in various regions of the tick synganglion including protocerebral, cheliceral, stomodeal, palpal, pedal I–IV, and opisthosomal regions in both species. However, there is one layer of immunoreactive cells located on the dorsal surface of the postoesophageal part of the synganglion that is found only in D. variabilis. Besides the immunoreactivity within the cell body and its axons, the neuropile and the neural lamella (the extracellular sheath of the synganglion) are rich in immunoreactive materials. Some coxal muscles are innervated by the FMRFamide-like immunoreactive processes of the nerve from the pedal ganglion.     

Colour change ability and its effect on prey capture success in female Misumenoides formosipes crab spiders

1. Changing between white and yellow body colour in certain crab spider species has been interpreted as an adaptation for matching the background colour where they hunt and thereby remaining cryptic to prey and/or their own predators. The potential costs and benefits of colour change in female Misumenoides formosipes Walckenaer were investigated via assessment of prey opportunities and capture success, in conjunction with the tendency for and rate of colour change on different backgrounds. 2. It was tested whether being matched or mismatched to their background affected foraging by moving females between white and yellow inflorescences. Female colour was quantified in digital photos using the Lab colour space component of Adobe photoshop , providing the first empirical assessment of the rate of colour change for a crab spider species. 3. Insect visits (potential prey) on inflorescences with and without spiders and prey capture success with females matched and mismatched to their background were quantified. 4. Yellow females abandoned white inflorescences, whereas white females remained on and underwent colour change on yellow inflorescences. This difference supported the notion that the costs of colour change differ depending on the starting colour. Female departures from white flowers were apparently not due to a lack of insect visitation, as white inflorescences had higher visitation rates than did yellow inflorescences, even in the presence of spiders. 5. An increase in the prey capture success of females who transitioned from white to yellow body colour on a yellow background supported the hypothesis that colour matching functions to deceive prey.     

Offspring recognition by mother crab spiders with extreme maternal care

Maternal care is provided by several spider species, but there are no reports of mother spiders recognizing their young, which suggests that maternal care can be exploited by unrelated individuals. Diaea ergandros, a crab spider with extreme, sacrificial maternal care, does accept unrelated spiderlings (ca. 43.9% of spiderlings) into its nest in areas of high nest density. However, a field and a laboratory experiment with mother spiders and natural and adoptive spiderlings demonstrated that mothers did recognize their own offspring. Recognition was not expressed in survival as adopted (unrelated) spiderlings had similar survival rate to that of natural offspring. Instead it was displayed in growth; mother D. ergandros caught large prey items for their own offspring, but not for adopted spiderlings, and so natural offspring grew more than adopted spiderlings. Also, mothers produced trophic oocytes, which are important for the sacrificial care that influences spiderling survival, only when they lived with their own offspring.     

Variability of mitochondrial cytochrome oxidase first subunit gene (<Emphasis Type="Italic">COI</Emphasis>) fragments in several tick species of the marginatus group (Ixodidae,Amblyomminae, <Emphasis Type="Italic">Dermacentor</Emphasis>)

This paper analyses variability of the nucleotide sequence of the mitochondrial cytochrome oxidase first subunit gene fragments in four tick species of the marginatus group—Dermacentor marginatus, D. silvarum, D. niveus, and D. nuttalli. It is noted that, with the exception of D. niveus, the clustering of mitotypes in the marginatus group does not show a clear correspondence of the species identity of ticks, which had preliminarily been established based on morphological characters.     

Cannibalism and kin recognition in Delena cancerides (Araneae: Sparassidae), a social huntsman spider

Social behaviour in spiders is rare: of the 39 000 species of spiders known, only 23 are considered to be cooperatively social. Delena cancerides is a social species of the huntsman spider that is endemic to Australia. This species is virtually unique among social spiders, having evolved social behaviour in the absence of a snare web. It is thought that this form of social behaviour in D. cancerides has evolved via the sub-social route, that is, the extension of an ancestrally occurring period of maternal care and the delayed dispersal of juveniles. Most social spiders show no aggression towards non-kin conspecifics, prompting suggestions that spiders cannot recognize kin; however, D. cancerides individuals are highly aggressive towards conspecifics introduced from outside their own colony. In order to determine whether selective aggression in D. cancerides has its basis in kin recognition, tolerance behaviour was assessed in the context of kinship and size. We observed that, in general, juveniles preferred to starve than engage in cannibalism of any conspecifics, related or not. However, where cannibalism did occur, non-kin were preferentially eaten, indicating that this species is clearly capable of kin recognition. Size thresholds were also established, below which juveniles are tolerated by adults and above which aggressive interactions leading to death occur. We conclude that kin recognition and juvenile dispersal explain the uncharacteristically high levels of genetic polymorphism in this species.