Article

Mimicry complex in two central European zodariid spiders (Araneae: Zodariidae): How Zodarion deceives ants

April 2002
Biological Journal of the Linnean Society 75(4):517-532

DOI:10.1046/j.1095-8312.2002.00043.x

Authors:

Stano Pekar

Masaryk University

Jiří Král

Charles University in Prague

Ant-eating spiders, Zodarion germanicum and Z. rubidum, were found to resemble ants structurally (size, colour, setosity) and behaviourally (ant-like movement, antennal illusion). Zodarion germanicum mimics large dark ants, such as Formica cinerea, whereas Z. rubidum resembles red ants, e.g. Myrmica sabuleti. Thus, these spiders are generalized Batesian mimics. The two spiders use aggressive mimicry during prey capture. When a spider carries a captured ant it will try to pass by approaching ants using special deceiving behaviour, which is based on imitation of ants’ nestmate recognition. The spider first taps the antennae of the curious ant with its front legs (transmitting a tactile cue), then exposes its prey (the ant corpse) which the ant antennates (thus the corpse transmits an olfactory cue). The distal part of the front legs of Zodarion are almost without macrosetae similar to the antennae of ants. Additionally, all the other legs are covered with flattened incised setae, which imitate the dense setosity of ants’ limbs. These remarkable microstructural imitations are believed to improve imitation of tactile signals by spiders. Moreover, by tapping, zodariids can presumably recognize the approaching intruder and decide whether to undertake the risk of deception or to run away. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 517–532.

The isolated locality of Zodarion rubidum Simon, 1914 (Araneae: Zodariidae) in Poland

Article

Full-text available

Jan 2006

Life-history constraints in inaccurate Batesian myrmecomorphic spiders (Araneae: Corinnidae, Gnaphosidae)

Article

Full-text available

Apr 2011

Accurate Batesian mimicry is known to impose constraints on some traits of the mimic, such as foraging or reproductive behaviour. It is not known whether life-history traits of inaccurate Batesian mimics are constrained as well. We studied selected lifehistory traits of three spider species, Liophrurillus flavitarsis, Phrurolithus festivus (both Corinnidae), and Micaria sociabilis (Gnaphosidae), that are inaccurate mimics of ants. Namely, we were interested in how myrmecomorphy (ant-like resemblance) constrains their circadian activity, trophic niche and reproductive behaviour. The spiders were found to have diurnal activity like their models, whereas their close relatives have nocturnal activity. The three mimics do not catch ants, nor do they use food resources of ants, but catch various tiny invertebrates that occur in the vicinity of their models. Their trophic niche seems to be constrained by occurrence among ants. Absence of courtship and long lasting copulation, in a position that does not provide protective resemblance, do not seem to be constrained by mimicry in the three species. Comparative analysis of fecundity in mimetic and non-mimetic spiders showed that clutch size is also not constrained. Unlike in accurate mimics, life-history traits of inaccurate myrmecomorphs appear not to be constrained.

Natural history and karyotype of some ant-eating zodariid spiders (Araneae, Zodariidae) from Israel

Article

Full-text available

Apr 2005
J ARACHNOL

Natural history, including phenology, circadian activity, mimicry, reproduction, prey specialization and karyotype was studied in the zodariid spiders Trygettus sexoculatus. Zodarion cyrenaicum, Z. lutipes and Z. nitidum (Zodariidae. Zodariinae) found in Israel. The spiders were active throughout the year, with maximum seasonal activity in the summer. Two distinct reproductive periods were found for Z cyrenaicum and Z nitidum, one in May and the other in November Individuals of all species studied were observed hunting only in the morning. Three zodariid species were found to generally mimic ants: Trygettus sexoculatus mimicked tiny yellow-brown ants such as Monomorium niloticum, Z. cyrenuicum mimicked large black ants such as Messor arenarius. and Z. lutipes mimicked large yellow-brown ants such as Camponotus fellah, The zodariids observed were able to subdue Various ant species, from the subfamilies Formicinae, Myrmicinae and Dolichoderinae. Trygettus sexoculatus appeared to specialize on Monomorium sp., Z lutipes on Camponotus sp. and Z cyrenaicum on Messor sp. ants, i.e., the same ant species they imitate. When bitten by zodariids. Formicinae and Dolichoderinae ants were paralyzed much more quickly than Myrmicinae. Female zodariid paralyzed ants faster than juveniles and males. Courtship and mating were observed only in Z lutipes and were found to he similar to other Zodarion species. The mean fecundity for all three Zodarion species ranged from 38-45 eggs per egg sac, thus being higher than reported in central European species. Females of all three species guarded egg sacs inside of their retreats. Karyotypes of studied Zodarion spiders were similar to the karyotypes of other zodariid spiders in terms of the diploid number (26 in Z cyrenaicum and 25 in both Z. lutipes and Z nitidum), sex chromosome systems and morphology of chromosomes. Most of the data indicate that the Zodarion species of this study have a close affinity to a group of Western European Zodarion species.

Specialized exploitation of ants (Hymenoptera: Formicidae) by spiders (Araneae)

Article

Full-text available

Aug 2012

Ants are a dominant resource in the spider's world, and spiders have a variety of ways of exploiting this resource. Two broad domains of exploitation are reviewed, namely specializing on ants for food and specializing on ants for models to mimic. Exploiting of ants as a source of food includes preying on worker ants and also taking food out of the ant's mandibles. Experiments have revealed numerous examples of spiders that specialize on ants by deploying ant-specific prey-capture behaviour. Consistent with other evidence that predatory versatility is widespread among spiders, many of the spiders that specialize at preying on ants sometimes adopt alternative tactics for capturing ants and are also proficient at targeting other prey. The venom, enzymes and sensory systems of spiders can also be specialized for preying on ants. Many spiders adopt Batesian mimicry of ants for protection against predators that readily eat spiders but have an aversion to ants. For these spiders, one of the costs of mimicking ants is attracting the unwanted attentions of spiders that specialize at preying on ants. Sometimes spiders solve this problem by making use of a conditional anti-predator strategy of resembling ants by default but switching to behaviour unlike an ant when ant-eating predators are encountered. Batesian mimicry of ants is sometimes communal (i.e., ant mimics living in groups appear more formidable because of the group's resemblance to a group of ants) and communal Batesian mimicry can then be deployed as a part of an aggressive-mimicry strategy. Ant-averse spiders may abandon their broods when confronted by a swarm of ants and likewise they flee when confronted by a swarm of communal ant-mimics, with the mimics then feeding on the unguarded broods. Other spiders use Wasmannian mimicry based on acquiring the cuticular hydrocarbons of ants as a means of safely mingling with the ants and then robbing the ants of their broods.

Ecological specialization and reproductive isolation among closely related sympatric ant‐eating spiders

Article

Full-text available

Jul 2022
J ANIM ECOL

Biological divergence results from several mechanisms. Defensive mechanisms, such as Batesian mimicry, can cause reproductive isolation via temporal segregation in foraging activity, particularly, in species that closely associate with their model. This seems to be the case of ant‐eating spiders, which can be inaccurate Batesian mimics of their prey. Here, we focused on Zodarion nitidum, which has two forms occurring in sympatry, black and yellow. Given the expected noticeable impact of their colour differences on the spiders' interactions with their potential predators and prey, we investigated whether these morphotypes have diverged in other aspects of their biology. We measured the two morphotypes' phenotypic resemblance to a mimetic model, tested whether they were protected from predators, investigated their circadian activity, surveyed the prey they hunted, modelled their distributions, performed crossing experiments and estimated their degree of genetic differentiation. We found that the black morphotype is ant‐like, resembling Messor ants, and it was not distinguishable from their ant models by four potential predators. In contrast, the yellow morphotype seems to use predator avoidance as a defensive strategy. Additionally, the two morphotypes differ in their circadian activity, the yellow morphotype being nocturnal and the black one being diurnal. The two morphotypes hunt and associate with different ant prey and possess marked differences in venom composition. Finally, crossing trials showed complete pre‐mating isolation between the two morphotypes, but there was no evidence of genetic (mitochondrial data) or environmental niche differentiation. We conclude that the two morphotypes show evidence of a deep differentiation in morphological, behavioural, physiological and ecological traits that evolved together as part of the spider's diverging lifestyles.

No ontogenetic shift in the realised trophic niche but in Batesian mimicry in an ant-eating spider

Article

Full-text available

Jan 2020

In predators an ontogenetic trophic shift includes change from small to large prey of several different taxa. In myrmecophagous predators that are also mimics of ants, the ontogenetic trophic shift should be accompanied by a parallel mimetic change. Our aim was to test whether ant-eating jumping spider, Mexcala elegans, is myrmecomorphic throughout their ontogenetic development, and whether there is an ontogenetic shift in realised trophic niche and their mimetic models. We performed field observations on the association of Mexcala with ant species and investigated the natural prey of the ontogenetic classes by means of molecular methods. Then we measured the mimetic similarity of ontogenetic morphs to putative mimetic models. We found Mexcala is an inaccurate mimic of ants both in the juvenile and adult stages. During ontogenesis it shifts mimetic models. The mimetic similarity was rather superficial, so an average bird predator should distinguish spiders from ants based on colouration. The realised trophic niche was narrow, composed mainly of ants of different species. There was no significant difference in the prey composition between ontogenetic stages. Females were more stenophagous than juveniles. We conclude that Mexcala is an ant-eating specialist that reduces its prey spectrum and shifts ant models during ontogenesis.

A contribution to the Slovenian spider fauna – III

Article

Full-text available

Nov 2016

Abstract. The study reports on first records of two spider species for Slovenian fauna, Zodarion rubidum and Prinerigone vagans from the Bela krajina region in southeastern Slovenia. Regarding their presence in the neighbouring countries and distribution in Europe, both species could be considered as expected. The finding of Z. rubidum during night time in the urban area demonstrates negligence of sampling at unconventional time and in anthropogenic habitats. Finding two new records during short term field survey also indicates undersampling of the spider fauna and supports the need for further faunistic work in the field of arachnology.

Trophic specialisation in a predatory group: The case of prey-specialised spiders (Araneae)

Article

Aug 2014
BIOL REV

Predators appear to be less frequently specialised (i.e. adapted to restricted diet) on their prey than herbivores, parasites or parasitoids. Here, we critically evaluate contemporary evolutionary hypotheses that might be used to explain the evolution of specialised foraging in predators. We propose a unifying concept within which we define four types of trophic categories using ecological (diet breadth) and evolutionary (degree of adaptations) contexts. We use data on spiders (Araneae), the most diversified order of terrestrial predators, to assess applicability of frameworks and evolutionary concepts related to trophic specialisation. The majority of spider species are euryphagous but a few have a restricted prey range, i.e. they are stenophagous. We provide a detailed overview of specialisation on different prey types, namely spiders, crustaceans, moths, dipterans, ants, and termites. We also review the available evidence for trophic adaptations, classified into four categories: behavioural, morphological, venomic and metabolic. Finally, we discuss the ecological and evolutionary implications of trophic specialisation and propose avenues for future research.

Dangerous prey is associated with a type 4 functional response in spiders

Article

Jun 2013
ANIM BEHAV

Prey can defend themselves against predators in many different ways. Social insects, such as ants, possess particularly effective defensive systems. Some predators are better adapted to prey defence than others. We compared the capture and defence efficiency in three spider species that differ in their level of myrmecophagy. We used three ant species differing in body size and aggression in a functional response experimental set-up that measured capture frequency at different prey densities. We found a type 4, dome-shaped functional response, and we propose a new mechanistic model to describe this type. Estimated parameters (searching efficiency, handling time, inhibition by prey) were then compared among spider and ant species to quantify density-dependent defensive effects on the predator's capture efficiency. We also compared survival of spiders during experiments. We found that myrmecophagous Zodarion spiders hunted ants with the highest capture efficiency and had the highest survival, suggesting that these spiders are adapted to living with high densities of ants. Polyphagous Xysticus spiders captured ants with lower efficiency and had the lowest survival, indicating that these spiders are adapted to the capture of solitary ant workers. Polyphagous nonanteating Pardosa spiders did not capture ants but had high survival, and are apparently adapted to living with high densities of ants. The new proposed model of the type 4 functional response can be applied to other predator–prey systems in which the prey is dangerous and a decrease in predator hunting has a similar dependence on prey density.

Spider-Ant Associations: An Updated Review of Myrmecomorphy, Myrmecophily, and Myrmecophagy in Spiders

Article

Full-text available

Apr 2012

Paula E Cushing

This paper provides a summary of the extensive theoretical and empirical work that has been carried out in recent years testing the adaptational significance of various spider-ant associations. Hundreds of species of spiders have evolved close relationships with ants and can be classified as myrmecomorphs, myrmecophiles, or myrmecophages. Myrmecomorphs are Batesian mimics. Their close morphological and behavioral resemblance to ants confers strong survival advantages against visually hunting predators. Some species of spiders have become integrated into the ant society as myrmecophiles or symbionts. These spider myrmecophiles gain protection against their own predators, live in an environment with a stable climate, and are typically surrounded by abundant food resources. The adaptations by which this integration is made possible are poorly known, although it is hypothesized that most spider myrmecophiles are chemical mimics and some are even phoretic on their hosts. The third type of spider-ant association discussed is myrmecophagy—or predatory specialization on ants. A table of known spider myrmecophages is provided as is information on their biology and hunting strategies. Myrmecophagy provides these predators with an essentially unlimited food supply and may even confer other protections to the spiders.

An ant‐mimicking ant on an oceanic archipelago: Camponotus guanchus mimics Crematogaster alluaudi—An analogy with the situation of Camponotus lateralis (Hymenoptera: Formicidae)

Article

Full-text available

Aug 2024

Mimicry, that is, the imitation of any unpalatable or defensive species by another, has been of central interest to evolutionary research since Darwin's lifetime. Two ant species, Camponotus guanchus Santschi, 1908 and Crematogaster alluaudi Emery, 1893, endemic to the Canary Islands, occur in two color‐morphs: While the head of workers is always reddish and the gaster blackish, the mesosoma (inclusive waist) is either fully reddish or fully blackish. In addition to the obvious morphological and coloration similarities, we provide evidence of mimicry: (i) Ca. guanchus was found only within the area of Cr. alluaudi. (ii) Color morphs are geographically non‐randomly distributed: Workers of both species from 16 localities of syntopic occurrences shared in eight cases a blackish and in eight cases a reddish mesosoma. Hence, Ca. guanchus mimics both local color‐morphs of Cr. alluaudi. We consider a fascinating analogy with the Mediterranean mimicry system in Camponotus lateralis (Olivier, 1792) and its model species of the Crematogaster scutellaris (Olivier, 1792) group on an island scale. Additionally, we present two endemic bug species, Perenotus stysi (Ribes et al., 2008) and P. malobae Roca‐Cusachs & Goula, 2016, as mimics of those Cr. alluaudi workers having a reddish mesosoma. Our distribution, coloration, frequency, and behavioral data as well as the analogy with Ca. lateralis and the Cr. scutellaris group suggest a Batesian‐mimicry system in which Ca. guanchus, Perenotus stysi, and P. malobae mimic the unpalatable and aggressive Cr. alluaudi as an antipredator adaptation.

Living with the enemy: behavioral study of Myrmecicultor chihuahuensis Ramírez, Grismado & Ubick (Araneae: Myrmecicultoridae)

Article

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Apr 2024

Spiders of Iran – Systematics, diversity and distribution

Thesis

Full-text available

Apr 2023

Alireza Zamani

Spiders (Araneae) are the largest order of Arachnida and the sixth most speciose order of Animalia, comprising more than 50,000 extant species as well as over 1,400 species known from fossils. Despite this immense diversity that has been estimated to comprise 120,000–200,000 species, our knowledge of their systematics and distribution remains rather incipient. While attempts to evaluate the diversity and distribution patterns of spiders have been made for the Neotropical and a few other faunas, most other regions remain historically neglected. The aim of this dissertation is to explore the systematics, diversity and distribution of spiders of Iran, a highly interesting region from a zoological and biogeographical point of view that unfortunately has been poorly investigated regarding its invertebrate fauna. For this purpose, I examined more than 9,000 specimens that were either collected during expeditions to numerous regions and ecosystems across the country or deposited in several natural history collections from around the world. As a result of these efforts, which were initiated in 2013 and mostly carried out in collaboration with researchers from various countries, a total of 11 genera and 147 species of Iranian spiders were described as new to science, and 419 taxa (i.e., 13 families, 87 genera and 319 species) were recorded in Iran for the first time. The total number of newly described and recorded species (i.e., 466 species) constitutes almost half (i.e., 49.83%) of the currently known Iranian species diversity of this group (i.e., 935 species). Amongst 147 species described, 137 are known only from Iran, representing 63.72% of all species currently considered endemic to this country (i.e., 215 species). These results were published in a total of 87 publications, 38 of which were published after the beginning of my doctoral studies at UTU in 2019. This dissertation, however, is primarily based on only nine taxonomic articles (i.e., I–IX), all published after 2019 and highlighting some of the more important findings. Because of the newly obtained taxonomic information and an extensive database of all published records, it was possible to conduct a comprehensive review of spatial variation in the diversity patterns of spiders in Iran. This was the main objective of publication X, which also included an evaluation of the effect of sampling bias on the current understanding of the distribution of Iranian spiders. The analyses showed that the diversity of Iranian spiders remains inadequately studied and is heavily affected by the Linnean shortfall (i.e., gaps in taxonomic knowledge), despite a remarkable improvement in taxonomic research on this fauna since the beginning of the 21st century. There are 935 spider species in the 1,648,195 km2 of Iran. Comparing the number of species per area of 171 countries and other political regions indicated that Iran was in position 132, whereas many considerably smaller and less ecologically diverse countries were in lower positions. It was also found that this fauna clearly suffers from a severe Wallacean shortfall (i.e., lack of knowledge of species distributions), as approximately 85% of Iran lacks a single record of spiders. There is a highly uneven distribution of records throughout the country and its provinces and ecoregions, with most of the records situated near large cities. A high correlation was found between the number of records of spiders and the number of records of plants and other animals in Iran, indicating that the noted shortfalls are indeed corroborated by other taxa. Finally, it is suggested that to gain a more complete picture of the diversity of Iranian spiders, future collecting efforts should be primarily in the form of extensive systematic surveys instead of opportunistic sampling, and ideally targeting lesser sampled areas and ecoregions. Once a satisfactory amount of information regarding the taxonomy and distribution of species becomes available, it will be possible to properly assess the conservation status and risk factors that affect these species and to identify areas of higher conservation and management priority.

Defences of Arachnids: diversified arsenal used against range of enemies

Article

Jul 2022
ENTOMOL GEN

Arachnida represent a hyperdiverse group of terrestrial arthropods. Although arachnids are mostly predators, they often fall prey to other predators. Here, the diversity of enemies and the defences of arachnids which are used against their predators is reviewed. The main predators of arachnids are other arachnids, followed by insects, mammals and birds. Almost all types of defences has evolved in this class, but countershading, and elusiveness await to be discovered in Arachnida. Overall, empirical evidence is rare but has slowly been accumulating over the last decade. Anachoresis is used most frequently across all orders; only in Acari, Araneae, Opiliones, and Solifugae it is rivalled by background matching, and Batesian mimicry. The number of different types of defences used by an order is positively correlated with the number of predatory groups preying on it. The major gaps are identified and future avenues for investigation – cases which deserve special attention because these may reveal completely new phenomena – are proposed. It is concluded that this group offers a very diverse array of defences which has not been sufficiently studied; thus, this review aims to stimulate further research.

Dangerous visions: a review of visual antipredator strategies in spiders

Article

Full-text available

Jan 2022
EVOL ECOL

Many animals use visual traits as a predator defence. Understanding these visual traits from the perspective of predators is critical in generating new insights about predator–prey interactions. In this paper, we propose a novel framework to support the study of strategies that exploit the visual system of predators. With spiders as our model taxon, we contextualise these strategies using two orthogonal axes. The first axis represents strategies using different degrees of conspicuousness to avoid detection or recognition of the spider and deter predator attacks. The second axis represents the degree of honesty of the visual signal. We explore these issues with reference to the three main vision parameters: spectral sensitivity, visual acuity, and temporal resolution, as well as recent tools to study it, including multispectral digital imaging.

Gut content metabarcoding of three widespread Iberian ant-eating spiders reveals specialisation on the same abundant harvester ants

Article

Dec 2021
ECOL ENTOMOL

Trophic specialists often follow unique evolutionary pathways, show potential applications in biological control, and suffer from increased ecological vulnerability. The myrmecophagous spider genus Zodarion, highly diversified across the Mediterranean, is among the few predatory groups showing strict stenophagy. Most Zodarion species display narrow distributions, but a few of them occur across hundreds to thousands of kilometres, raising questions on the causes of their comparatively higher distributional success. In this study, through extensive geographic sampling and the metabarcoding of gut contents, we explored the diets of the three most widely distributed Iberian species of Zodarion: Zodarion alacre and the two lineages of Zodarion styliferum. Our data support the three species as strictly myrmecophagous, showing diets largely based on the granivorous Messor ants. Auxiliary prey includes other ants of the subfamilies Myrmicinae, Formicinae, and Dolichoderinae. The diet breadth of both Z. styliferum lineages was slightly narrower than that of Z. alacre, coinciding with their higher physiological and behavioural efficiency for predating on Messor. We discard that the relatively large distribution range of our three focal Zodarion species is related to breaking the boundaries of myrmecophagy, and instead argue that such success could be favoured by their specialisation on Messor ants, which are also abundant and widespread across the Iberian Peninsula.

Phylogenomics and loci dropout patterns of deeply diverged Zodarion ant-eating spiders suggest a high potential of RAD-seq for genus-level spider phylogenetics

Article

Oct 2021
CLADISTICS

RAD sequencing yields large amounts of genome-wide data at a relatively low cost and without requiring previous taxon-specific information, making it ideal for evolutionary studies of highly diversified and neglected organisms. However, concerns about information decay with phylogenetic distance have discouraged its use for assessing supraspecific relationships. Here, using Double Digest Restriction Associated DNA (ddRAD) data, we perform the first deep-level approach to the phylogeny of Zodarion, a highly diversified spider genus. We explore the impact of loci and taxon filtering across concatenated and multispecies coalescent reconstruction methods and investigate the patterns of information dropout in reference to both the time of divergence and the mitochondrial divergence between taxa. We found that relaxed loci-filtering and nested taxon-filtering strategies maximized the amount of molecular information and improved phylogenetic inference. As expected, there was a clear pattern of allele dropout towards deeper time and mitochondrial divergences, but the phylogenetic signal remained strong throughout the phylogeny. Therefore, we inferred topologies that were almost fully resolved, highly supported, and noticeably congruent between setups and inference methods, which highlights overall inconsistency in the taxonomy of Zodarion. Because Zodarion appears to be among the oldest and most mitochondrially diversified spider genera, our results suggest that ddRAD data show high potential for inferring intra-generic relationships across spiders and probably also in other taxonomic groups.

Increased speed of movement reduced identification of Batesian ant-mimicking spiders by surrogate predators

Article

Full-text available

Nov 2021
ANIM COGN

Stano Pekar

In Batesian mimicry, the mimic gains protection from predators by imitating a noxious model. Some myrmecomorphic species use ants as models as ants have strong defensive capabilities. Ants are highly mobile models, and besides colour, shape, and size, mimics also imitate their movement. Yet, former studies focused mainly on static traits. Here, I tested the hypothesis that artificially increased speed of movement reduces the probability of the mimic being identified. First, images of 14 myrmecomorphic spider species and their models were used for humans to rank their mimetic accuracy. Humans were used as surrogate predators to obtain scores for each mimetic pair. In the second experiment, the effect of movement playback speed on the probability of identification was investigated, again using humans. Videos of mimics were played at different speeds, and the identification probability was recorded. While ants were correctly identified at any playback speed, the identification of myrmecomorphic spiders declined with increasing playback speed. In other words, the latency to correct identification increased with playback speed. Overall, mimics with higher accuracy scores were more difficult to identify while moving. The natural speed of movement of accurate mimics was similar to that of inaccurate ones. Movement is thus an important trait for myrmecomorphic species.

Poor performance of DNA barcoding and the impact of RAD loci filtering on the species delimitation of an Iberian ant-eating spider

Article

Oct 2020
MOL PHYLOGENET EVOL

Genomic data provide unprecedented power for species delimitation. However, current implementations are still time and resource consuming. In addition, bioinformatic processing is contentious and its impact on downstream analyses is insufficiently understood. Here we employ ddRAD sequencing and a thorough sampling for species delimitation in Zodarion styliferum, a widespread Iberian ant-eating spider. We explore the influence of the loci filtering strategy on the downstream phylogenetic analyses, genomic clustering and coalescent species delimitation. We also assess the accuracy of one mitochondrial (COI) and one nuclear (ITS) barcode for fast and inexpensive species delineation in the group. Our genomic data strongly support two morphologically cryptic but ecologically divergent lineages, mainly restricted to the central-eastern and western parts of the Iberian Peninsula, respectively. Larger matrices with more missing data showed increased genomic diversity, supporting that bioinformatic strategies to maximize matrix completion disproportionately exclude loci with the highest mutation rates. Moderate loci filtering gave the best results across analyses: although larger matrices returned concatenated phylogenies with higher support, middle-sized matrices performed better in genetic structure analyses. COI displayed high diversity and a conspicuous barcode gap, revealing 13 mitochondrial lineages. Mitonuclear discordance is consistent with ancestral isolation in multiple groups, probably in glacial refugia, followed by range expansion and secondary contact that produced genomic homogenization. Several apparently (unidirectionally) introgressed specimens further challenge the accuracy of species identification through mitochondrial barcodes in the group. Conversely, ITS failed to separate both lineages of Z. styliferum. This study shows an extreme case of mitonuclear discordance that highlights the limitations of single molecular barcodes for species delimitation, even in presence of distinct barcode gaps, and brings new light on the effects of parameterization on shallow-divergence studies using RAD data

THE IMPACT OF PREDATION BY THE MYRMECOPHAGOUS SPIDER ZODARION ELEGANS (ARANEAE: ZODARIIDAE) ON THE ACTIVITY PATTERN OF THE MEDITERRANEAN HARVESTER ANT MESSOR WASMANNI (HYMENOPTERA: FORMICIDAE)

Article

Sep 2016

Tanja Traxler

This study was designed to determine the effect of predation by the zodariid, obligate myrmecophagous spider Zodarion elegans (Simon, 1873) on the activity pattern and behavior of the Mediterranean harvester ant Messor wasmanni Krausse, 1910. Previous field studies suggested that the activity patterns of M. wasmanni colonies are modified not only by microclimatic conditions, but also by biological factors such as the presence of predators. The present study clearly demonstrates that M. wasmanni colonies respond in two different ways to spider predation. The ants 1) attack the invader, and 2) temporarily stop all foraging activities and close nest entrances with pebbles and other materials. This causes obligate myrmecophagous spiders such as Z. elegans to leave such inactive colonies and move to active ones nearby. Moreover, the laboratory experiments show that the capture of a single ant is in every case sufficient to trigger cessation of foraging activities and nest closure. After the spider attacks, entrances remained closed for up to six days. Perturbation experiments show that events that directly affect workers engaged in one task outside the nest also alter the activity intensity of task performance inside the nest. All exterior workers remain inside the nest when entrances are closed. Inside the nest, only a small proportion of marked foragers switch to food processing, while the majority was inactive or performed grooming during the inactive phase outside the nest.

A contribution to the Slovenian spider fauna - IV

Article

Full-text available

Jun 2019

Sixteen spider species, new to the Slovenian fauna, have been recorded: Aphantaulax trifasciata, Araneus triguttatus, Arctosa cinerea, Cnephalocotes obscurus, Helophora insignis, Heriaeus hirtus, Nomisia exornata, Nurscia albomaculata, Pardosa agricola, Pardosa prativaga, Pardosa proxima, Philodromus longipalpis, Runcinia grammica, Theridula gonygaster, Xysticus ninnii and Zora armillata. Recent records on presumably extinct endemic species Nesticus idriacus are also provided, as well as suggestion to its transfer to the IUCN category rare (R).

Evolution of ant-eating specialization in the basal lineage of Zodariidae (Araneae): the trophic ecology of South American Leprolochus birabeni Mello-Leitão

Article

Apr 2018

Most spiders are euryphagous predators. However, there are also a few species with a stenophagous diet, foraging on only a single prey taxon. This strategy could have resulted from ecological opportunity or from evolutionary specialization, which is typical for phylogenetically advanced species. In Zodariidae, myrmecophagy is considered a derived trophic strategy. Therefore, the main objective of this study was to investigate the trophic ecology of Leprolochus birabeni (Zodariidae), whose rather basal position in the phylogeny predicts euryphagy. We studied the realized niche (potential and actual prey) and the fundamental niche (capture success), capture efficiency and behavioural adaptations in males and females of this species. Observations in nature revealed a narrow trophic niche composed only of ants. The most frequently captured prey in the field were two species of Acromyrmex ants, and this was also confirmed in the DNA gut content analysis. Acceptance experiments revealed that both females and males refused to catch prey other than ants. We found nocturnal activity in L. birabeni to be correlated significantly with the activity of one of their preferred prey, Acromyrmex lobicornis. In conclusion, our results show that L. birabeni is a stenophagous ant-eating specialist with a high preference for Acromyrmex ants.

A new ant-eating spider genus Suffascar (Araneae:Zodariidae) endemic to Madagascar: A considerable extension of the dual femoral organ clade

Article

Sep 2017

The new zodariid genus Suffascar, gen. nov. is described from Madagascar. It is apparently endemic to the island, common and speciose. Twelve new species were discovered and are here described: Suffascar albolineatus, sp. nov., S. fianara, sp. nov., S. fisheri, sp. nov., S. fitzpatrickae, sp. nov., S. gigas, sp. nov., S. griswoldi, sp. nov., S. macromma, sp. nov., S. micromma, sp. nov., S. nonus, sp. nov., S. scutatus, sp. nov., S. sufficiens, sp. nov. and S. tofti, sp. nov. It is well known that Zodariidae with femoral organs are specialised consumers of ants or termites, although the exact role of these glands still remains obscure. Suffascar is part of the 'dual femoral organ clade' (DF), which also includes Asceua Thorell, 1887, Suffasia Jocqué, 1991, and Suffrica Henrard & Jocqué, 2015. Using a morphology-based phylogenetic analysis, we test whether zodariids sharing the dual femoral organ are part of the same group as the 'single femoral organ clade' (SF) or form a separate monophyletic group. It appears that SF and DF constitute clearly monophyletic entities with a sister-group relation. These groups may be considered as distinct subfamilies and, together, they may turn out to be sister to the Storeninae (which then could be re-established). An analysis that focuses on these questions is underway. http://zoobank.org/urn:lsid:zoobank.org:pub:281E1357-FAEE-4300-B1BB-F0D43E724E91

The amber worlds: paleobiology, paleoenvironments and paleogeography of resin-producing ecosystems.

Article

May 2015

Vincent Perrichot

L'ambre constitue une fenêtre exceptionnelle sur les écosystèmes du passé, et son étude peut intéresser des disciplines aussi variées que la paléobotanique, la paléoentomologie, la paléomicrobiologie, ou la géochimie organique et inorganique. Ce mémoire présente la démarche scientifique et une synthèse des travaux de recherche menés depuis quinze ans sur différents gisements du Crétacé, de l'Eocène, et du Miocène. Trois principaux axes de recherche sont exposés et accompagnés d'une sélection de publications, portant sur des aspects biologiques et géologiques: l'origine et l'histoire évolutive des insectes Hyménoptères ; l'évolution paléoécologique et paléogéographique des écosystèmes résinifères ; et des préservations exceptionnelles trouvées dans l'ambre. Les orientations des futurs travaux de recherche sont brièvement envisagées.

The evolutionary ecology of deception

Article

Full-text available

Jun 2015
BIOL REV

Through dishonest signals or actions, individuals often misinform others to their own benefit. We review recent literature to explore the evolutionary and ecological conditions for deception to be more likely to evolve and be maintained. We identify four conditions: (1) high misinformation potential through perceptual constraints of perceiver; (2) costs and benefits of responding to deception; (3) asymmetric power relationships between individuals and (4) exploitation of common goods. We discuss behavioural and physiological mechanisms that form a deception continuum from secrecy to overt signals. Deceptive tactics usually succeed by being rare and are often evolving under co-evolutionary arms races, sometimes leading to the evolution of polymorphism. The degree of deception can also vary depending on the environmental conditions. Finally, we suggest a conceptual framework for studying deception and highlight important questions for future studies. © 2015 Cambridge Philosophical Society.

A coat of many scents: Cuticular hydrocarbons in multitrophic interactions of fig wasps with ants

Article

Full-text available

Aug 2015
ACTA OECOL

The fig-fig wasp system of Ficus racemosa constitutes an assemblage of galler and parasitoid wasps in which tritrophic interactions occur. Since predatory ants (Oecophylla smaragdina and Technomyrmex albipes) or mostly trophobiont-tending ants (Myrmicaria brunnea) were previously shown to differentially use volatile organic compounds (VOCs) from figs as proximal cues for predation on fig wasps, we examined the response of these ants to the cuticular hydrocarbons (CHCs) of the wasps. CHC signatures of gallers were distinguished from those of parasitoids by the methyl-branched alkanes 5-methylpentacosane and 13-methylnonacosane which characterised trophic group membership. CHC profiles of wasp predator and wasp prey were congruent suggesting that parasitoids acquire CHCs from their prey; the CHC composition of the parasitoid Apocrypta sp 2 clustered with that of its galler host Apocryptophagus fusca, while the CHC profile of the parasitoid Apocryptophagus agraensis clustered with its galler prey, the fig pollinator Ceratosolen fusciceps. In behavioural assays with ants, parasitoid CHC extracts evoked greater response in all ant species compared to galler extracts, suggesting that parasitoid CHC extracts contain more elicitors of ant behaviour than those of plant feeders. CHCs of some wasp species did not elicit significant responses even in predatory ants, suggesting chemical camouflage. Contrary to earlier studies which demonstrated that predatory ants learned to associate wasp prey with specific fig VOCs, prior exposure to fig wasp CHCs did not affect the reaction of any ant species to these CHCs.

Behavioral manipulation outside the world of parasites

Book

Jun 2012

Evolutionary implications of deception in mimicry and masquerade

Article

Full-text available

Feb 2014

Ximena Nelson

Aggressive mimicry occurs when an organism resembles some aspect of another organism (the model) in order to obtain prey through its deceptive resemblance. This may function either through the overt response of the receiver or through the lack of response of the receiver. Reviewing selected examples, I discuss some of the difficulties in ascribing a model for the mimic. I also discuss how a single animal can have multiple ploys in its armoury of deceptive signals, thus belonging within two or more categories of deceptive signalling. In addition to aggressive mimicry, these may include crypsis or camouflage, masquerade (mimicry of inanimate objects), and Batesian or protective mimicry. Each of these examples of deception has multiple evolutionary pathways, and some deceptive signals may be more costly to receivers than others, but no single organism is subject to a single selection pressure, leading to the reality that many evolutionary pathways contribute to the diversity we see around us. New technologies are opening new channels of investigation into deceptive signaling in many different sensory modalities, and this is reflected in the recent increase in studies investigating the structure and function of deceptive signals. In turn, these studies are beginning to expose the fascinating complexity of deceptive signaling systems, allowing us to discover the myriad, non-mutually exclusive, solutions that can be selected for to obtain prey

Predatory behavior of two European ant-eating spiders (Araneae, Zodariidae)

Article

Full-text available

Apr 2004
J ARACHNOL

Stano Pekar

Prey specialization and the predatory behavior of two European ant-eating zodariid spiders, Zodarion germanicum and Zodarion rubidum, were studied in detail. The spiders were offered 12 ant species and seven other insects (termites, beetles, aphids, silverfish, flies, crickets and grasshoppers). Study spiders turned out to be ant specialists as they were able to subdue many ant species but ignored all other insects, except termites, which they attacked but rarely subdued. The best capture success was obtained with medium-sized ants (e.g. Lasius and Formica). The predatory behavior of the zodariid spiders involves an attacking and a handling phase separated by a period of waiting at a safe distance. The attacking phase consisted of a very rapid lunge from the rear, followed by a bite on the most extended ant leg. After an attack, the spider retreated to a safe distance, perhaps an indication that natural selection has favored such caution in the presence of an aggressive prey. The spider waited until the ant ceased moving. Such predatory behavior, which limits contact with the predator and prey, is clearly an effective means of handling a dangerous prey.

How spiders practice aggressive and Batesian mimicry

Article

Full-text available

Aug 2012

To understand communication, the interests of the sender and the receiver/s of signals should be considered separately. When our goal is to understand the adaptive significance of specific responses to specific signals by the receiver, questions about signal information are useful. However, when our goal is to understand the adaptive significance to the sender of generating a signal, it may be better to envisage the receiver's response to signals as part of the sender's extended phenotype. By making signals, a sender interfaces with the receiver's model of the world and indirectly manipulates its behaviour. This is especially clear in cases of mimicry, where animals use deceptive signals that indirectly manipulate the behaviour of receivers. Many animals adopt Batesian mimicry to deceive their predators, or aggressive mimicry to deceive their prey. We review examples from the literature on spiders to illustrate how these phenomena, traditionally thought of as distinct, can become entangled in a web of lies [Current Zoology 58 (4): 620-629, 2012].

Predatory characteristics of ant-eating Zodarion spiders (Araneae: Zodariidae): Potential biological control agents

Article

Aug 2005

Stano Pekar

Some ant species are important nuisance pests. Mechanical and chemical strategies are currently used to control them. Biological control, based on the use of a natural enemy, has not yet been proposed. The aim of this study was to investigate the predatory efficiency of ant-eating Zodarion spiders as possible biological control agents of ants. The investigation included identification of prey specificity, study of the functional response, and the consumption rate. Two species were used, Zodarion rubidum and Zodarion styliferum. Z. rubidum was found to be a Formicinae specialist as it was more efficient in the paralysis of Lasius and Formica ants (both Formicinae) than in the paralysis of Monomorium, Tetramorium, and Messor ants (all Myrmicinae). The functional response of Z. rubidum was Type II in the small arena and Type IV in the large arena. Although spiders killed up to 24 ants during 12h, only a few prey items were actually consumed. The overkilling thus amounted to 80–100% and was due to specialized predatory behavior, in which ants are consumed some time after being immobilized. Spiders usually started to feed only if no living ant was around. Juvenile spiders of Z. rubidum consumed at most two Lasius ants consecutively, the handling time being prolonged with the later prey captured. During 26 days juveniles of Z. styliferum captured 21 ants, but their exploitation of ant biomass decreased with time. It is concluded that Zodarion spiders possess some features typical of biological control agents, however, further experiments are necessary to assess their real potential.

Nowe dane na temat lokalizacji Zodarion rubidum SIMON, 1914 (Araneae, Zodariidae) w Polsce

Article

Full-text available

Jan 2009

The paper presents the information about a new locality of Zodarion rubidum SIMON, in the eastern outskirts of Świnoujście (NW Poland). The discovered station is the first locality in Poland where a larger number of Z. rubidum individuals was collected. This fact shows that this invasive spider species became a stable component of the Polish araneofauna. The station discovered in Świnoujście is its farthest northwards site of occurrence in Europe.

Comparison of natural histories and karyotypes of two closely related ant‐eating spiders, Zodarion hamatum and Z. italicum (Araneae, Zodariidae)

Article

Full-text available

Sep 2005
J NAT HIST

Based on morphological characters, two species of the genus Zodarion, Z. hamatum and Z. italicum, were placed into the italicum group. The aim of this study was to evaluate similarity of these two species in respect of various aspects of their natural history and karyotypes. Both species had similar phenological patterns, being univoltine, but some specimens reached adulthood the same year before winter. Both representatives had nocturnal activity and were able to capture various ant species. Zodarion hamatum seems to be specialized on Myrmicinae ants, and Z. italicum on Formicinae ants. Zodarion hamatum showed more agile predatory behaviour than Z. italicum presumably due to specialization on Myrmicinae ants, which have lower population density. Both species are generalized mimics of bicoloured (orange‐dark brown) ants, such as Lasius emarginatus; they performed similar courtship and mating. The two study species exhibited contrasting reproductive strategies. Zodarion hamatum spiders had higher fecundity than Z. italicum as a result of larger body size but Z. italicum had larger eggs than Z. hamatum. This is presumably an adaptation to different availability of prey ants. Both species had the same diploid chromosome number, sex chromosome system and pattern of X chromosome heteropycnosis in testes. Results support a very close relationship between these two Zodarion species.

[Arachnida] Spinnentiere

Chapter

Full-text available

Jan 2008

Datasets of traits of zodariid spiders (Araneae: Zodariidae)

Article

Full-text available

Aug 2024

Stano Pekar

Species traits are essential for inferences on ecology and the evolution of organisms. Spiders are the most abundant and diversified terrestrial predators, playing an important role in a range of ecosystem services. Here, I present datasetse on all traits of zodariid spiders, which are known to be free-living ground-dwellers occurring on all continents (except Antarctica) with the highest species diversity in Australia. I collated the data from published resources. The datasets includes nearly 100 000 trait records on all (90) genera and almost all species (1249) of the family. The majority of the 88 traits collected are morphometric, followed by those relating to ecology, reproduction, and physiology. Morphometric traits were available for the majority of species. Other trait classes were only available for some species. I provide a standardized classification of selected categorical traits (habitat, microhabitat, retreat type, circadian activity, prey, primary defensive, and predators). This is the first complete database of traits of a whole spider family, which is available through the World Spider Trait database.

A contribution to the Slovenian spider fauna – IV

Article

Full-text available

Jun 2019

We provide a list of 171 spider species from 27 families recently recorded in Slovenia. Among them are first records of Nigma flavescens, Walckenaria alticeps, Pelecopsis parallela, Erigone autumnalis and Micaria subopaca for Slovenia and a second record of Zodarion rubidum, a species just recently added to the Slovenian spider fauna.

Spinnen und andere Nachahmer, Vortäuscher und Räuber

Chapter

Aug 2023

An arachnid’s guide to being an ant: morphological and behavioral mimicry in ant-mimicking spiders

Article

Dec 2022

Batesian mimicry imposes several challenges to mimics and evokes adaptations in multiple sensory modalities. Myrmecomorphy, morphological and behavioral resemblance to ants, is seen in over 2000 arthropod species. Ant-like resemblance is observed in at least 13 spider families despite spiders having a distinct body plan compared to ants. Quantifying the extent to which spiders’ shape, size, and behavior resemble model ants will allow us to comprehend the evolutionary pressures that have facilitated myrmecomorphy. Myrmaplata plataleoides are thought to closely resemble weaver ants, Oecophylla smaragdina. In this study, we quantify the speed of movement of model, mimic, and non-mimetic jumping spiders. We use traditional and geometric morphometrics to quantify traits such as foreleg size and hindleg size, body shape between the model ant, mimic, and non-mimics. Our results suggest that while the mimics closely resemble the model ants in speed of movement, they occupy an intermediate morphological space compared to the model ants and non-mimics. Ant-mimicking spiders are better at mimicking ants’ locomotory movement than morphology and overall body shape. Some traits may compensate others, suggesting differential selection on these mimetic traits. Our study provides a framework to understand the multimodal nature of mimicry and helps discern the relative contributions of such traits that drive mimetic accuracy in ant-mimicking spiders.

Trophic specialization of a newly described spider ant symbiont, Myrmecicultor chihuahuensis (Araneae: Myrmecicultoridae)

Article

Full-text available

Aug 2022

The spider Myrmecicultor chihuahuensis Ramrez, Grismado & Ubick (Myrmecicultoridae) was described in 2019 and hypothesized to be a myrmecophile, living inside the nests of Novomessor (Myrmicinae) and perhaps also Pogonomyrmex (Myrmicinae) ants. To test the hypothesis that M. chihuahuensis are chemical mimics of their host ants, we carried out behavioral bioassays to observe interactions between the spiders and the host ants. We compared the cuticular hydrocarbon (CHC) profiles of the spiders and the ants. We discovered that this new species of spider is a myrmecophage, displaying hunting strategies similar to other myrmecophagous spiders, particularly those in the genus Zodarion Walckenaer, 1826 (Zodariidae). The CHC profile of M. chihuahuensis did not match the profile of the ants with which it associates, Novomessor albisetosus (Mayr), but ants and spider shared several compounds, potentially involved in species recognition.

An arachnid′s guide to be an ant: morphological and behavioural mimicry in ant-mimicking spiders

Preprint

Full-text available

Nov 2021

Batesian mimicry imposes several challenges to mimics and evokes adaptations in multiple sensory modalities. Myrmecomorphy, morphological and behavioral resemblance to ants, is seen in over 2000 arthropod species. Ant-like resemblance is observed in at least 13 spider families despite spiders having a distinct body plan compared to ants. Quantifying the extent to which spider's shape, size, and behavior resemble model ants will allow us to comprehend the evolutionary pressures that have facilitated myrmecomorphy. Myrmaplata plataleoides are 'accurate' mimics of the weaver ants, Oecophylla smaragdina. In this study, we quantify the speed of movement of model, mimic, and non-mimetic jumping spiders. We use traditional and geometric morphometrics to quantify traits such as foreleg and hindleg size, body shape between the model ant, mimic, and non-mimics. Our results suggest that while the mimics closely resemble the model ants in speed of movement, they occupy an intermediate morphological space compared to the model ants and non-mimics. We suggest that ant-mimicking spiders are better at mimicking ant's locomotory movement than morphology and overall body shape. Our study provides a framework to understand the multimodal nature of mimicry and helps discern the relative contributions of such traits that drive mimetic accuracy in ant-mimicking spiders.

Disguises and the Origins of Clothing

Article

Full-text available

Dec 2021
HUM NATURE-INT BIOS

William Buckner

Thermoregulation is often thought to be a key motivating factor behind the origins of clothing. Less attention has been given, however, to the production and use of clothing across traditional societies in contexts outside of thermoregulatory needs. Here I investigate the use of disguises, modesty coverings, and body armor among the 10 hunter-gatherer societies in the Probability Sample Files (PSF) within the Human Relations Area Files (HRAF) World Cultures database, with a particular focus on disguise cases and how they compare with strategies of deception across other taxa. The employment of disguises-defined as altering one's appearance for purposes of deceiving conspecifics or other animals-is noted for eight of the 10 societies, with their use occurring in contexts of hunting, religious or cult practices, and war or interpersonal violence. Most hunter-gatherer disguises demonstrated clear similarities to cases of visual deception found in other species, with the majority of examples fitting categories of animal mimicry, masquerading as plants, disrup-tive coloration (camouflage), or background matching (camouflage), while disguises unique to humans involved the impersonation of culture-specific "spirit-beings." Clothing for modesty purposes (nine societies) and body armor (six societies) are also noted. I propose that strategic initiatives by individuals or groups to disguise or conceal themselves represents one possible initial pathway to the cultural evolution of clothing. There are likely multiple potential (nonexclusive) social and functional pathways to the emergence of clothing outside of thermoregulatory needs.

Predation by avian predators may have initiated the evolution of myrmecomorph spiders

Article

Full-text available

Aug 2021

Myrmecomorphy is a strategy utilized by a variety of species, among which spiders are the most common. It is supposed that myrmecomorphy tends to be selected by predator avoidance of preying on ants rather than by blind ant workers. To date, this hypothesis has been tested mainly on invertebrate predators (mantises and spiders). We are the first to test whether an imperfect myrmecomorph spider (Phrurolithus festivus) gains protection against avian predators (wild adult great tits—Parus major) through its appearance. In a set of preferential trials, we showed that the ant model and the myrmecomorph spider are equally well protected against attack, though the attacked myrmecomorphs are usually eaten. This suggests that the mimicry of the myrmecomorph spiders is effective against avian predators and works in a Batesian manner. In this study, we have provided evidence toward the evolution of myrmecomorphy in response to selective pressure elicited by visually-oriented predators like birds.

Booysen & Haddad, 2021. Revision and molecular phylogeny of the spider genus Micaria Westring, 1851 (Araneae, Gnaphosidae) in the Afrotropical Region

Article

Mar 2021
ZOOTAXA

The genus Micaria Westring, 1851 (Araneae, Gnaphosidae) is a group of small (1.85–5 mm) ant-like spiders that can be distinguished from other gnaphosids by their piriform gland spigots that are similar in size to the major ampullate gland spigots. According to the World Spider Catalog, there are 105 species of Micaria in the world, of which only three species are known from the African part of the Afrotropical Region, namely M. chrysis (Simon, 1910), M. tersissima Simon, 1910 and M. beaufortia (Tucker, 1923). The objectives of this study were to revise Micaria in the Afrotropical Region, providing new and updated records for each of the species, evaluating the relationships between them using COI barcoding data, and providing information on their biology, mimetic relationships and feeding ecology. These objectives were met by collecting fresh material from the KwaZulu-Natal, Western Cape, Northern Cape and Free State provinces in South Africa. Fresh material of M. tersissima and M. chrysis were collected from their type localities, Komaggas and Port Nolloth (Northern Cape Province), respectively, for identification and DNA analyses. COI sequences generated, together with those sourced from Barcode of Life Data Systems (BOLD) and GenBank, were aligned using the CulstalW alignment algorithm in the Mega X software, and molecular phylogenetic analyses were performed using MrBayes for Bayesian Inference (BI) and RaxML for maximum likelihood (ML) analyses. Morphological examination of the collected and voucher material yielded 17 new species for the Afrotropical Region, namely M. basaliducta sp. nov. (♀, ♂, South Africa), M. bimaculata sp. nov. (♀, ♂, Mauritania), M. bispicula sp. nov. (♀, ♂, Namibia, South Africa), M. durbana sp. nov. (♀, ♂, South Africa, Zambia), M. felix sp. nov. (♀, ♂, Cameroon, Ethiopia, Malawi, Mozambique, Namibia, South Africa, Zambia, Zimbabwe), M. gagnoa sp. nov. (♀, ♂, Côte d’Ivoire, Mozambique, Mozambique, Tanzania), M. koingnaas sp. nov. (♂, South Africa), M. lata sp. nov. (♂, Namibia, South Africa), M. laxa sp. nov. (♀, South Africa), M. mediospina sp. nov. (♂, South Africa), M. parvotibialis sp. nov. (♀, ♂, Senegal), M. plana sp. nov. (♀, ♂, Ethiopia), M. quadrata sp. nov. (♀, Ethiopia), M. quinquemaculosa sp. nov. (♀, ♂, Namibia, South Africa), M. rivonosy sp. nov. (♀, ♂, Madagascar), M. sanipass sp. nov. (♂, South Africa) and M. scutellata sp. nov. (♂, South Africa). Furthermore, both sexes of M. beaufortia, as well as the male of M. tersissima, are redescribed. Both sexes of M. chrysis are described for the first time, as this species was only known from a juvenile. Of the previously known species, M. beaufortia (Botswana, Ethiopia, Lesotho, Namibia, South Africa, Zimbabwe) and M. chrysis (Côte d’Ivoire, Ethiopia, Lesotho, Namibia, South Africa, Tanzania) are widespread in the Afroptropics, while M. tersissima is only known from South Africa. Both the Bayesian inference and the maximum likelihood analysess recovered Micaria (sensu lato) as monophyletic with the inclusion of the subopaca group. The pulicaria species group was recovered as polyphyletic in both the BI and ML analyses. Four Afrotropical species, as well as the M. rossica Thorell, 1875/M. foxi Gertsch, 1933 group, formed a clade sister to M. formicaria (Sundevall, 1831). Eight of the Afrotropical species now have COI barcoding data uploaded to BOLD.

The Structure of Mimicry

Chapter

Full-text available

Jan 2017

Timo Maran

A property that makes mimicry both fascinating and difficult to comprehend is its diversity. Mimetic resemblances can occur as colours and forms in the visual medium, as imitations of hissing , buzzing and other sounds, or as similarities of chemical components in pheromones . Mimicry can be based on different ecological relations (predation , parasitism, symbiosis , competition) and the number and composition of involved species can vary to a great extent. Mimicry can take place inside organisms at the cellular level, a phenomenon that is known as molecular mimicry. In some cases, mimicry requires the coordinated behaviour of several individuals to create or enhance the mimetic effect (e.g. myrmecomorph ic jumping spiders aggregating to emphasise their resemblance with ants, Nelson and Jackson 2009). Indeed, there seems to be little in common between the cases of fully behavioural mimicry, such as the mimic octopus Thaumoctopus mimicus , in which the cephalopod uses its tentacles to create sporadic imitations of flatfish, sea snakes , lionfish and other sea creatures, and the stillness of the perfect resemblance between the eggs of the common cuckoo Cuculus canorus and those of reed warblers, pipits, redstarts and other hosts species, whose nests the cuckoo uses to lay its eggs.

Evolutionary implications of deception in mimicry and masquerade

Article

Jan 2014

X.J. Nelszon

Ant mimicry in the spider Myrmecotypus iguazu (Araneae: Corinnidae), with notes about myrmecomorphy in spiders

Article

Full-text available

Nov 2013

We describe the mimetic relationship between the ant-like spider Myrmecotypus iguazu Rubio & Arbino 2009 (Araneae: Corinnidae) and the carpenter ant Camponotus sericeiventris Guérin (Hymenoptera: Formicidae), studied in a subtropical rainforest in Iguazú National Park, Argentina. The morphological adaptations, aspects of coloration, and behavior responsible for the ant-like appearance in M. iguazu (the mimic) provide strong evidence that its model is C. sericeiventris. Both field observations and field and laboratory experiments suggest that this spider is a Batesian mimic.

An Inventory of Alien Species and Their Threat to Biodiversity and Economy in Switzerland

Book

Full-text available

Sep 2005

Cockroach as the Earliest Eusocial Animal

Article

Aug 2010
ACTA GEOL SIN-ENGL

Peter Vršanský

: A completely preserved cockroach Sociala perlucida gen. et sp. nov. (Blattida: Socialidae fam. nov.) is described from the Mesozoic (Albian Early Cretaceous) amber of Archingeay in France. It is categorized within the new family Socialidae, originating from the Mesozoic cockroach family Liberiblattinidae, and representing the sister group to the most basal known eutermite families. Numerous direct and indirect morphological evidence (such as a unique narrow body and pronotum with reduced coloration, legs without carination and with numerous sensillar pitts (forelegs) as adaptations to life in nests analogical to the living termitophillous species; enlarged head, long palps for communication, a general venation modified in a direction towards termites (subcosta and radial vein [R] approximated, R branches approximated, simplified, cross-veins reduced), and possibly also the development of the breaking forewing sutura for detaching wings after marriage flight), suggest its eusocial mode of life. Thus, the first eusociality originated within cockroaches, prior to giving birth to termites; termites and Cryptocercus are not directly related. The group of eusocial cockroaches makes termite morphotaxon polyphyletic, but also in this case, the erection of a new order for mastotermites (including Cratomastotermitidae) and/or the inclusion of eusocial, morphological cockroaches within termites appears counterproductive, thus an example of such a general exception to the taxonomical procedure is provided.

Spiders and allies - Arachnida. In: Invasive alien species in Switzerland

Chapter

Full-text available

Dec 2006

Globalization increases trade, travel and transport and is leading to an unprecedented homogenization of the world’s biota by transport and subsequent establishment of organisms beyond their natural barriers. Some of these alien species become invasive and pose threats to the environment and human economics and health. This report on alien biota in Switzerland lists about 800 established alien species and characterises 107 invasive alien species (IAS) in Fact Sheets: five mammals, four birds, one reptile, three amphibians, seven fish, four molluscs, 16 insects, six crustaceans, three spiders, two «worms», seven fungi, one bacteria, and 48 plants. A general chapter explains some common patterns in pathways, impacts and management, and gives recommendations for the management of alien species. The main body of the report is organised into taxonomic groups, and includes an overview, lists of alien species, Fact Sheets on the invasive species, and an evaluation of the status, impacts, pathways, control options and recommendations. The Fact Sheets summarize information on the invasive species under the headings taxonomy, description, ecology, origin, introduction, distribution, impact, management and references.

Fidelity of vocal mimicry: Identification and accuracy of mimicry of heterospecific alarm calls by the brown thornbill

Article

Mar 2013

Avian vocal mimicry has been studied for decades, but little is known about its function or requirements for accurate imitation. Furthermore, progress is hampered by the difficulty in identifying which vocalizations are indeed mimetic. We tested historical claims of vocal mimicry in the brown thornbill, Acanthiza pusilla, using a combination of human and computer methods to identify mimicry, followed by comparisons of acoustic similarity with model vocalizations. We recorded vocalizations of brown thornbills and sympatric heterospecifics while undisturbed and during mist net capture or the presence of natural or model predators. We then cross-validated human classification of mimicry with computer classification based on spectrographic measurements and spectral cross-correlation. Finally, we quantified the accuracy of the most common imitations. Brown thornbills predominantly imitated alarm calls given by heterospecifics towards aerial predators, which function in these models to provoke immediate flight by receivers. Human and computer-based methods produced consistent results when identifying and classifying mimicry. Mimicked aerial alarms were not perfect imitations of their corresponding model alarms, but did retain specific acoustic properties previously shown to be important for provoking immediate alarm responses. Although less accurate mimicry may reflect physiological constraints, we suggest that mimetic function, perhaps startling predators, only requires mimicry to retain features of model alarms that provoke immediate alarm responses by receivers. Understanding what factors influence the acoustic structure of mimetic vocalizations is essential in understanding the evolution of vocal mimicry, particularly with accumulating evidence that mimetic function does not always require perfect resemblance in other sensory modalities.

Revision of the genus Zodarion Walckenaer, 1833 part II. Western and Central Europe, including Italy (Araneae, Zodariidae)

Article

Full-text available

Jan 1997

Robert Bosmans

Ecology and diet of Psammoduon deserticola (Simon) (Araneae: Zodariidae)

Article

Full-text available

Jan 1999

Quelques aspects de la biologie de Zodarion rubidum Simon, 1918

Article

Full-text available

Jan 1990

Jean-Marc Couvreur

Ant-hunting behaviour in spiders with emphasis on Strophius nigricans (Thomisidae)

Article

Full-text available

Jan 1985

Myrmecomorphy and Myrmecophily in Spiders: A Review

Article

Full-text available

Jun 1997

Paula E Cushing

Myrmecomorphs are arthropods that have evolved a morphological resemblance to ants. Myrmecophiles are arthropods that live in or near ant nests and are considered true symbionts. The literature and natural history information about spider myrmecomorphs and myrmecophiles are reviewed. Myrmecomorphy in spiders is generally considered a type of Batesian mimicry in which spiders are gaining protection from predators through their resemblance to aggressive or unpalatable ants. Selection pressure from spider predators and eggsac parasites may trigger greater integration into ant colonies among myrmecophilic spiders. /// Los mirmecomorfos son artrópodos que han evolucionado desarrollando una semejanza morfológica a las hormigas. Los Myrmecófilos son artrópodos que viven dentro o cerca de nidos de hormigas y se consideran verdaderos simbiontes. Ha sido evaluado la literatura e información de historia natural acerca de las arañas mirmecomorfas y mirmecófilas. El myrmecomorfismo en las arañas es generalmente considerado un tipo de mimetismo Batesiano en el cual las arañas están protegiéndose de sus depredadores a través de su semejanza con hormigas agresivas o no apetecibles. La presión de selección de los depredadores de arañas y de parásitos de su saco ovopositor pueden inducir una mayor integración de las arañas mirmecófílas hacia las colonias de hormigas.

A comparative study of the biology and karyotypes of two Central European zodariid spiders (Araneae, Zodariidae)

Article

Full-text available

Jan 2001

A comparison of the biology and karyotypes of Zodarion germanicum and Zodarion rubidum (Araneae, Zodariidae) which occur in central Europe was carried out. Surprisingly, these species were found to differ in a number of characters such as pattern of activity, reproduction and karyotypes. Zodarion germanicum was observed to be diurnal, whereas Z. rubidum is nocturnal. Courtship and mating were markedly longer and more complex in Z. germanicum than in Z. rubidum. Females of Z. germanicum produced only one or two successive egg sacs including 17 eggs on average which they would guard, while females of Z. rubidum produced up to 5 egg sacs each having 4 eggs that they abandoned. The two species differ from each other also in number of chromosomes and the sex chromosome system. Results suggest these species belong to distant evolutionary lineages within the genus Zodarion.

Inter-specific associations involving spiders: Kleptoparasitism, mimicry and mutualism

Article

Full-text available

Jan 1993

Mark A Elgar

Many spiders have life-styles that involve a relatively close and prolonged association with another species; for example, between a specialist predator and its prey species, or a species may rely on another for either protection from predators or providing a suitable place to live. In asymmetric relationships, where individuals of one species benefit at the expense of the other, each species may act as a selection pressure on the other species. This can result in the evolution of specific adaptations and counter-adaptations that are evident in at least three kinds of inter-specific associations between spiders, kleptoparasitism, mimicry and mutualism are reviewed here. -from Author

Exploitation of an Ant Chemical Alarm Signal by the Zodariid Spider Habronestes bradleyi Walckenaer

Article

Full-text available

Jan 1996

Intraspecific signals are vulnerable to exploitation by predators that are not the targets of the signal. This cost has been documented for several acoustic, visual and chemical signals, but not for chemical alarm pheromones. We reveal a novel form of exploitation of an ant alarm pheromone by the cursorial spider Habronestes bradleyi (Zodariidae), a specialist predator of the highly territorial and aggressive meat ant Iridomyrmex purpureus. We demonstrate experimentally that H. bradleyi locates I. purpureus workers engaged in territorial interactions with conspecifics, by using the alarm pheromone of the ants as a cue. The spiders are attracted to an airborne cue, identified as the alarm pheromone 6-methyl-5-hepten-2-one, which is produced by injured or alarmed I. purpureus workers but not by inactive I. purpureus, or injured workers of other sympatric ant species. These data demonstrate a novel cost of producing alarm pheromones.

Natural history of the spider genus Lutica (Araneae, Zodariidae)

Article

Jan 1995

Martina Giselle Ramirez

A generic revision of the spider family Zodariidae (Araneae)

Article

Jan 1991

Rudy Jocqué

Zodariidae

Article

Jan 1953

H. Wiehle

Further observations on the relation between an ant, Cataglyphis bicolor (F.) (Hymenoptera: Formicidae) and a spider, Zodarion frenatum (Simon) (Araneae: Zodariidae)

Article

Jan 1977

RD Harkness

Tutelina similis (Araneae: Salticidae): An ant mimic that feeds on ants

Article

Jan 1983
J KANSAS ENTOMOL SOC

K. Wing

Histoire naturelle des araignees. Tome 1, Fasicule 3

Book

Jan 1893

E Simon

Histoire Naturelle des Araignées1

Book

Jan 1897

Eugene. Simon

The ants

Book

Jan 1990

Evolutionary origins of displays used in aggressive mimicry by Portia, a web-invading araneophagic jumping spider (Araneae: Salticidae)

Article

Jan 1990

Portia is a genus of jumping spiders which invade alien webs and use aggressive mimicry to prey on the resident spiders. Portia’s aggressive mimicry repertoire includes numerous different vibratory displays. Portia’s aggressive mimicry displays are compared to its grooming and disturbance behaviours, and hypotheses are discussed concerning the evolution of specific aggressive mimicry displays from specific grooming and disturbance behaviours. The contexts in which Portia and a variety of other salticids and spiders from other families groom and perform disturbance behaviours are investigated. Special attention is given to the conflict theory of ritualisation which has been useful in studies of other animals. This theory is not found to be very useful, however, for understanding the evolution of Portia’s aggressive mimicry displays.

Resampling: The New Statistics

Article

Jan 1995

Simon JL

A review of the anti-predator devices of spiders

Article

Jan 1995

John L. Cloudsley-Thompson

A Bolas-Throwing Spider

Article

Jan 1903

Charles E. Hutchinson

Spiders (Araneae) as bird food

Article

Jan 1997

The adaptive bases of ant-mimicry in a Neotropical aphantochilid spider (Araneae: Aphantochilidae)

Article

Jun 1984
BIOL J LINN SOC

The aphantochilid spider Aphanlochilus rogersi accurately mimics black ants of tribe Cephalotini, and is commonly found in the neighbourhood of its models' nests. The mimic seems to be a specialized predator of this type of ant, rejecting any insect offered as prey other than cephalotines. In the field, A. rogersi was observed preying on the model species Zacryptocerus pusillus. In captivity, the spider preyed on the models Z. pusillus and Z. depressus, as well as on the yellow non-model Z. clypeatus. Recognition of correct prey by A. rogersi appears to be based primarily on visual and tactile stimuli. Capturing ant prey from behind was the most common attack tactic observed in A. rogersi, and is probably safer than frontal attacks, as in this case the spider can be bitten on the legs before the ant is immobilized. Aphanlochilus rogersi, when feeding on the hard-bodied ant models, uses the ant corpses as a ‘protective shield’ against patrolling ants of the victim's colony and resembles an ant carrying a dead companion. Certain types of mimetic traits in A. rogersi (close similarity to ant models in integument texture and pilosity of body and legs), together with ‘shielding behaviour’, are thought to function as ant-deceivers, facilitating the obligatory intimate contact the mimic must make with cephalotines in order to capture a prey among other ants. The close similarity in the arrangement of dorsal spines, body shape, integument brightness and locomotion, together with antennal illusion, is regarded as a strategy of A. rogersi for deceiving visually-hunting predators that avoid its sharp spined ant models. It is proposed that ant-mimicry in A. rogersi has both an aggressive and a Batesian adaptive component, and evolved as a result of combined selective pressures exerted both by Cephalotini ant models (through defensive behaviour towards the mimics which attack them) and predators that avoid cephalotines (through predatory behaviour toward imperfect mimics). This suggestion is schematized and discussed in terms of two tripartite mimicry systems.

Colony dispersion and relationships of meat ants Iridomyrmex purpureus and allies in an arid locality in south Australia

Article

Mar 1983

The distributions of three “forms”, actually sibling species, of meat ant were studied near Morgan, South Australia, an area with mean annual rainfall of 230 mm. Colony dispersion is described for a “blue form” and for formpurpureus sens. strict. which occurs in this locality far beyond its normal low rainfall limit. An unusually large colony ofpurpureus had a territory covering 10 ha with more than 85 nests and over 1,500 individual nest entrances. Worker interactions between forms, and within and between colonies within forms are described. The ant fauna of the locality as a whole and the faunas accompanying differentIridomyrmex species were investigated by means of hand collections and pitfall trapping. Diversity (species — richness) and the occurrence of widely adapted ant species are related to vegetation type and associatedIridomymex. Catches of pitfall traps suggested that the numbers of individuals and species of other taxa active on the soil surface, predominantly Collembola, were affected by numbers of ants, particularlyIridomyrmex species and other, widely adapted ants.Iridomyrmex also influence associated fauna at an evolutionary level. The presence of zodariid spiders which are sibling species-specific colour mimics and predators of meat ants is noted. Possible cases of mimicry of meat ants by ants of the genusCamponotus are recorded and discussed.

Adult male bolas spider retain juvenile hunting tactics

Article

Jan 1997

Bolas spiders in the genus Mastophora exhibit extreme sexual size dimorphism. In temperate regions, the diminutive males become adults about 2 months before females mature. Late-instar and adult females attract certain male moths by aggressive chemical mimicry of those moth species' sex pheromones. While hunting, these larger female spiders hang from a horizontal silken line and capture moths by swinging a “bolas” (i.e., a sticky globule suspended on a thread) at the approaching moths. Small, early-instar bolas spiders of both sexes attract moth flies in the genus Psychoda, which they capture without using a bolas or web. Instead, they position themselves along leaf margins and use their front two pairs of legs to grab approaching prey. The predatory habits of adult male bolas spiders have never been reported. Our field experiments demonstrated that adult males of the bolas spider Mastophora phrynosoma attract adult male Psychoda phalaenoides. Each year during our 3-year study, significantly more P. phalaenoides were captured on sticky traps baited with live adult male M. phrynosoma than on unbaited control traps. Thus, the tiny adult male bolas spiders retain the juvenile hunting tactic of attracting psychodid flies, while female bolas spiders switch from hunting psychodid flies as spiderlings to hunting moths when the female spiders become older and larger.

Mimicry, Aposematism and Realted Phenomena in Animals and Plants : Bibliography 1800-1990 /

Article

The prey of the mud-dauber wasp, Sceliphron spirifex (Linnaeus), in Central Africa.

Jan 1988
309

Jocqué

Jocqué R. 1988. The prey of the mud-dauber wasp, Sceliphron spirifex (Linnaeus), in Central Africa. Newsletter of the British Arachnological Society 6/7: 309-312.

Revision of the myrmecophilous tribe Deremini III: the remainder of the genera with notes on behavior, ultrastructure, glands and phylogeny (Coleoptera, Staphylinidae).

Jan 1979
141

Kistner

Kistner DH. 1979. Revision of the myrmecophilous tribe Deremini III: the remainder of the genera with notes on behavior, ultrastructure, glands and phylogeny (Coleoptera, Staphylinidae). Sociobiology 3(3): 141-391.

The ants A bolas throwing spider

Jan 1903
172

B Hölldobler
Wilson
Eo

Hölldobler B, Wilson EO. 1990. The ants. Berlin: Springer Hutchinson CE. 1903. A bolas throwing spider. Scientific American 89/10: 172.

Histoire naturelle des araigneés Resampling: the new statistics Wadsworth: Resampling Stats. Vane-Wright RI. 1976. A unified classification of mimetic resemblances

Jan 1993
25-56

E Simon

Simon E. 1893. Histoire naturelle des araigneés. Tome 1, 2nd edn. Paris. Simon JL. 1993. Resampling: the new statistics, 2nd edn. Wadsworth: Resampling Stats. Vane-Wright RI. 1976. A unified classification of mimetic resemblances. Biological Journal of the Linnean Society 8: 25–56.

The book of spiders and scorpions

R Preston-Mafham

Preston-Mafham R. 1990. The book of spiders and scorpions. Quantum Books.

Morphological adaptation for ant-mimicry in spiders

Jan 1972
221-226

J Reiskind

Reiskind J. 1972. Morphological adaptation for ant-mimicry in spiders. In: Folk C, ed. Proceedings of the Fifth International Congress on Arachnology. Brno: Institute of Vertebrate Zoology, 221-226.

The use of 'shelters' by an ant hunting spider, Zodarium frenatum Simon, as receptacles for eggs. Entomologist's Monthly Magazine

Jan 1995
46

R D Harkness

Harkness RD. 1995. The use of 'shelters' by an ant hunting spider, Zodarium frenatum Simon, as receptacles for eggs. Entomologist's Monthly Magazine 131: 46.

The building and use of ‘shelters’ by a hunting spider (Zodarion frenatum Simon) in Greece.

Jan 1977
161

Harkness

Harkness RD. 1977. The building and use of 'shelters' by a hunting spider (Zodarion frenatum Simon) in Greece. Zoologische Anzeiger 199: 161-163.

Mistress of deception.

Jan 1996
103

Jackson

Jackson RR. 1996. Mistress of deception. National Geographic 190/5: 103-115.

Biológia našich mravčíkov (Araneae: Zodariidae) a pavúkov v sadoch

S Pekár

Pekár S. 2000. Biológia našich mravčíkov (Araneae: Zodariidae) a pavúkov v sadoch [Biology of two zodariid spiders (Araneae: Zodariidae) and of the spiders in orchards]. PhD Thesis, Charles University.

Oxford: Numerical Algorithms Group

Jan 1987

R J Baker

Baker RJ. 1987. GLIM 3.77 Reference Manual, 2nd edn. Oxford: Numerical Algorithms Group.

The comity of spiders II

Jan 1941

W S Bristowe

Bristowe WS. 1941. The comity of spiders II. London: Ray Society.

Le comportement de ‘presentation d’un leurre’ chez Zodarion rubidum (Araneae, Zodariidae)

J M Couvreur

Couvreur JM. 1990a. Le comportement de 'presentation d'un leurre' chez Zodarion rubidum (Araneae, Zodariidae). In: Célérier ML, Heurtault J, Rollard C, eds. Proceedings of the 12th European Colloquium of Arachnology, Paris. Bulletin de la Société européene Arachnologique 1: 75-79.

The guests of British ants, their habits and life histories. London: Routledge and Sons Elgar MA. 1993. Inter-specific associations involving spiders: kleptoparasitism, mimicry and mutualism

Jan 1927
411-430

H Donisthorpe

Donisthorpe H. 1927. The guests of British ants, their habits and life histories. London: Routledge and Sons Elgar MA. 1993. Inter-specific associations involving spiders: kleptoparasitism, mimicry and mutualism. Memoirs of the Queensland Museum 33(2): 411-430.

Gestalt und Lebensweise der 'Ameisenspinne' Synageles venator (Lucas)

Jan 1971
317-334

W Engelhardt

Engelhardt W. 1971. Gestalt und Lebensweise der 'Ameisenspinne' Synageles venator (Lucas). Zugleich ein Beitrag zur Ameisenmimikryforschung. Zoologische Anzeiger 185: 317-334.

Katalóg Pavúkov Slovenska. Catalogue of Slovakian Spiders. Bratislava: Ústav krajinnej ekológie SAV

Jan 1999

P Gajdoš
J Svatoň
K Sloboda

Gajdoš P, Svatoň J, Sloboda K. 1999. Katalóg Pavúkov Slovenska. Catalogue of Slovakian Spiders. Bratislava: Ústav krajinnej ekológie SAV.

UNCERT: Analyses of sequential events by Markov chains

E D Hailman
J P Hailman

Hailman ED, Hailman JP. 1993. UNCERT: Analyses of sequential events by Markov chains, Version alpha. A DOS-compatible software. Available at http://gila.cisab. indiana.edu/pub/.

The ants. Berlin: Springer Hutchinson CE. 1903. A bolas throwing spider

Jan 1990
SCI AMER
172

B Hölldobler
E O Wilson

Hölldobler B, Wilson EO. 1990. The ants. Berlin: Springer Hutchinson CE. 1903. A bolas throwing spider. Scientific American 89/10: 172.

Gestalt und Lebensweise der ‘Ameisenspinne’Synageles venator (Lucas). Zugleich ein Beitrag zur Ameisenmimikryforschung.

Jan 1971
317

Engelhardt

Quelques aspects de la biologie de Zodarion rubidum, Simon, 1914.

Couvreur

The use of ‘shelters’ by an ant hunting spider, Zodarium frenatum Simon, as receptacles for eggs.

Jan 1995
46

Harkness

Myrmecomorphy: morphological and behavioral mimicry of ants.

Jan 1993
351

McIver

Mimicry complex in two central European zodariid spiders (Araneae: Zodariidae): How Zodarion deceives ants

Abstract

No full-text available

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The adaptive bases of ant-mimicry in a Neotropical aphantochilid spider (Araneae: Aphantochilidae)