Mechanics of trichobothria in orb-weaving spiders (Agelenidae,Araneae)

Summary When a fly is humming at a distance of about one centimetre from an orb-weaving spider (Agelenidae) the trichobothria on the spider's extremities are deflected by air streams and air vibrations. Frequency analysis of the hum of the two prey animals,Drosophila andMusca, shows that the effective sound velocities of the harmonics with frequencies inferior to some five hundred Hz exceeds that of higher frequencies by a factor of at least 5. Biologically relevant resonances would, therefore, have to be looked for in the range of a few hundred Hz. Frequency response diagrams show that single hairs have no resonance between a few Hz and approximately 2 kHz. The maximal relative amplitude of hairs of different lengths shifts from the longer to the shorter hairs with increasing frequency. As this is only a minor effect, however, it appears that there is no frequency discrimination by the mechanical apparatus. Constant air streams with a velocity of 40 mm/s cause hair deflection of about 10 degrees (the hair's bend is neglibible). Similarly, near-field particle velocity of sound fields up to a few hundred Hz is well transmitted. The mechanical directional sensitivity does not depend on the azimuthal angle of deflection. Thus, information about direction and velocity of stationary and near-field air movements is transmitted without deformation by the mechanical apparatus. This is well matched with the fact that the hair is multiply innervated.Supported by grants of the Deutsche Forschungsmeinschaft in the field of the Schwerpunktprogramm Rezeptorphysiologie     

The ontogeny of personality traits in the desert funnel‐web spider,Agelenopsis lisa (Araneae: Agelenidae)

Trait consistency over time is one of the cornerstones of animal personality. Behavioral syndromes are the result of correlations between behaviors. While repeatability in behavior is not a requirement for behavioral syndromes, the two concepts studied together provide a more comprehensive understanding of how behavior can change over ontogeny. The roles of ontogenetic processes in the emergence of personality and behavioral syndromes have received much individual attention. However, the characterization of both individual trait consistency and behavioral syndromes across both sexes, as in our study, has been relatively rare. Ontogeny refers to changes that occur from conception to maturation, and juveniles might be expected to undergo different selection pressures than sexually mature individuals and also will experience profound changes in hormones, morphology, and environment during this period. In this study, we test for behavioral trait consistency and behavioral syndromes across six time points during ontogenetic development in the desert funnel‐web spider (Agelenopsis lisa). Our results indicate behavioral traits generally lack consistency (repeatability) within life stages and across ontogeny. However, penultimate males and mature females do exhibit noticeable mean‐level changes, with greater aggressive responses toward prey, shorter latencies to explore their environment and in the exhibition of risk‐averse responses to predatory cues. These traits also show high repeatability. Some trait correlations do exist as well. In particular, a strong correlation between aggressiveness toward prey and exploration factors is observed in mature males. However, because correlations among these factors are unstable across ontogeny and vary in strength over time, we conclude that behavioral syndromes do not exist in this species. Nevertheless, our results indicate that increased consistency, increasing average trait values, and varying correlations between traits may coincide with developmentally important changes associated with sexual maturation, albeit at different time points in males and females. This period of the life cycle merits systematic examination across taxa.     

Electrophysiology of trichobothria in orb-weaving spiders (Agelenidae,Araneae)

Summary Trichobothria of the house spidersTegenaria atrica (C.L. Koch) andTegenaria derhami (Scopoli) were stimulated by linear deflection of the hair from its resting position to one side. The pulse response of the receptor cell was analyzed. At angular deflection velocities of 10^–21 deg/ms the receptor begins to discharge at an angle of 3 degrees. While the mean pulse rate remains constant during deflections of 10^–210^–1deg/ms the pulse train may be interrupted by repeated breaks. Discharge continues when the hair is bent proximally beyond the bothrium edge. When the hair is bent distally and touches the bothrium edge, however, discharge ceases. Responsible for this phenomenon seems to be an asymmetry of hair suspension. — Repeated deflection leads to logarithmically ascending latency curves and logarithmically falling curves of the pulse numbers. The function coefficients depend on velocities and repetition rates of the deflections. The adaptational effect is heightened by preceding stationary deflection in the direction of the dynamical stimulus. The mean pulse rate as a function of hair deflection velocity increases logarithmically. The mean pulse rate as a function of hair movement direction obeys a cosine law, provided that a given velocity and a definite deflection angle are used.Supported by grants of the Deutsche Forschungsgemeinschaft given to Prof. Dr. P. Görner in the field of the Schwerpunktprogramm RezeptorphysiologieThe present paper is part of a doctoral thesis. My thanks are due to Prof. Dr. P. G6rner for the theme, many valuable discussions and his constant readiness to help.     

Contact sex signals on web and cuticle of Tegenaria atrica (Araneae,Agelenidae)

Qualitative changes in lipid content and composition were examined on the web and cuticle of Tegenaria atrica females in relation to sexual receptivity. In this spider species, 78 different compounds were detected by gas chromatography in the cuticle extract and 50 in the web; 28 identical compounds were present both on the spider silk and the cuticle of the female. The components were long‐chain aliphatic hydrocarbons, fatty acids, and esters. On the web, sexual receptivity was correlated with changes in eleven polar compounds. On the cuticle, sexual receptivity was correlated with changes in eight polar compounds and 26 hydrocarbons. Bioassays demonstrated that the methanol eluate of webs and females were involved in stimulating the sexual behavior of males. Qualitative and/or quantitative changes in hexadecanoic acid, octadecadienoic acid, octadecenoic acid, methyl palmitate, methyl octadecanoate, and n‐tricosane could play a role in the contact sex signals from web and cuticle of T. atrica. Arch. Insect Biochem. Physiol. 40:194–202, 1999. © 1999 Wiley‐Liss, Inc.     

Omega-agatoxins: novel calcium channel antagonists of two subtypes from funnel web spider (Agelenopsis aperta) venom

A new series of polypeptide presynaptic antagonists ("omega-agatoxins") was purified from venom of the funnel web spider Agelenopsis aperta. Physiological data indicate that all of these peptides are antagonists of voltage-sensitive calcium channels. Although all three omega-agatoxins (Aga) described here (omega-Aga-IA, omega-Aga-IB, and omega-Aga-IIA) block insect neuromuscular transmission presynaptically, biochemical data permit their subclassification as Type I and Type II toxins. Type I toxins (omega-Aga-IA and -IB) are 7.5 kDa, have closely related amino acid sequences, and exhibit characteristic tryptophan-like UV absorbance spectra. Complete Edman sequencing of omega-Aga-IA reveals it to be a 66-amino acid polypeptide containing 9 cysteines and 5 tryptophan residues. omega-Aga-IIA, a Type II toxin, is 11 kDa, shows limited amino acid sequence similarity to the Type I toxins, and exhibits mixed tryptophan- and tyrosine-like absorbance. Nanomolar concentrations of omega-Aga-IIA inhibit the specific binding of 125I-labeled omega-conotoxin GVIA to chick synaptosomal membranes while omega-Aga-IA and -IB have no effect under identical conditions. The omega-agatoxins thus are defined as two subtypes of neuronal calcium channel toxins with different structural characteristics and calcium channel binding specificities.     

Use of prey-specific predatory behaviour by North American jumping spiders (Araneae, Salticidae) of the genus Phidippus

Prey-capture behaviour of seven species of Phidippus is studied using two types of prey, house flies (adult Musca domestica ) and caterpillars (cabbage loopers, larval Trichoplusia ni ). Each species is shown to be a versatile predator which uses a different prey-specific prey-capture behaviour depending on whether prey is a fly or a caterpillar: spiders approached the two types of prey differently and leapt on them from different distances. Interspecific differences among Phidippus species are not pronounced, cxcept for variation in the distance from which spiders jumped and the spider's success at capturing a fly on the first try.     

The lace web spiders (Araneae,Phyxelididae) of Madagascar: phylogeny,biogeography and taxonomy

We briefly review the potential history of Madagascar as either a Darwinian or a Wallacean island, summarize the phylogenetic evidence regarding the biogeography of Madagascar spiders, examine the dispersal history of the Madagascar Phyxelididae, and monograph the family in Madagascar. Molecular phylogenetic analyses for 32 Malagasy phyxelidid exemplars, nine confamilial outgroup taxa, and seven other more distant outgroups are performed for three nuclear markers and one mitochondrial genetic marker (28S, 18S, H3 and COI) utilizing Bayesian, maximum‐likelihood and parsimony methods. These analyses suggest that there are 14 species of Phyxelididae that may be recognized from Madagascar, that these may be divided into three genera, and that the Malagasy phyxelidids form a monophyletic group, probably resulting from a single invasion of the island by an ancestor from Africa. Two new genera, ten new species, and two new combinations are proposed: Manampoka atsimo gen. nov., sp. nov. ; Rahavavy gen. nov. , including R. ida sp. nov. and R. fanivelona (Griswold, 1990) comb. nov. and R. malagasyana (Griswold, 1990) comb. nov. ; and Ambohima andrefana sp. nov. , A. antsinanana sp. nov. , A. avaratra sp. nov. , A. maizina sp. nov. , A. ranohira sp. nov. , A. vato sp. nov. , A. zandry sp. nov. and A. zoky sp. nov. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 728–810.     

omega-Aga-I: a presynaptic calcium channel antagonist from venom of the funnel web spider, Agelenopsis aperta

Spider venoms are proving to be important sources of specific ion channel toxins. Venom of Agelenopsis aperta, a funnel web spider, contains a class of polypeptide toxins which blocks neuromuscular synapses at nanomolar concentrations. Detailed physiological analyses of block caused by one of these toxins, omega-Aga-I, show that it suppresses transmitter release at insect and frog neuromuscular junctions and blocks calcium spikes in insect neuronal cell bodies. omega-Aga-I may define a binding site on neuronal calcium channels which is common to both vertebrates and invertebrates.     

Acutipetala, gen. nov., a new genus of funnel-web spiders from northern Thailand (Araneae, Agelenidae)

A new funnel-web spider genus, Acutipetala, gen. nov., is erected to accommodate two new agelenid species known to occur in evergreen forests of northern Thailand: Acutipetala octoginta, sp. nov. (type species,male symbolfemale symbol) and A. donglini, sp. nov. (male symbol). The genus is established on the basis of the distinctive appearance of the genital structures, in which the median apophysis of the male palp is petal-shaped, sharply pointed, and strongly sclerotized, and the truncate embolus is short, originates subapically, and is provided with a hook-shaped apical portion.     

The phylogenetic basis of sexual size dimorphism in orb-weaving spiders (Araneae,Orbiculariae)

Extreme sexual body size dimorphism (SSD), in which males are only a small fraction of the size of the females, occurs only in a few, mostly marine, taxonomic groups. Spiders are the only terrestrial group in which small males are relatively common, particularly among orb-weavers (especially in the families Tetragnathidae and Araneidae) and crab spiders (Thomisidae). We used a taxonomic sample of 80 genera to study the phylogenetic patterns (origins and reversals) of SSD in orb-weaving spiders (Orbiculariae). We collected and compiled male and female size data (adult body length) for 536 species. Size data were treated as a continuous character, and ancestral sizes, for males and females separately, were reconstructed by using Wagner parsimony on a cladogram for the 80 genera used in this study. Of these 80 genera, 24 were female-biased dimorphic (twice or more the body length of the male); the remaining 56 genera were monomorphic. Under parsimony only four independent origins of dimorphism are required: in the theridiid genus Tidarren, in the distal nephilines, in the "argiopoid clade," and in the araneid genus Kaira. Dimorphism has reversed to monomorphism at least seven times, all of them within the large "argiopoid clade." The four independent origins of dimorphism represent two separate instances of an increase in female size coupled with a decrease of male size (involving only two genera), and two separate instances of an increase in female size with male size either remaining the same or increasing, but not as much as females (involving 30 genera). In orb-weaving spiders, far more taxa are sexually dimorphic as a result of female size increase (22 genera) than as a result of male size decrease (two genera). SSD in orb-weaving spiders encompasses several independent evolutionary histories that together suggest a variety of evolutionary pathways. This multiplicity strongly refutes all efforts thus far to find a general explanation for either the origin or maintenance (or both) of SSD, because the different pathways very likely will require distinctly different, possibly unique, explanations. Each pattern must be understood historically before its origin and maintenance can be explained in ecological and evolutionary terms. The most frequently cited example of male dwarfism in spiders, the golden orb-weaving spider genus Nephila (Tetragnathidae), is in fact a case of female giantism, not male dwarfism.     

Purification and characterization of two classes of neurotoxins from the funnel web spider, Agelenopsis aperta

Two classes of paralytic toxins were isolated from the venom of Agelenopsis aperta and their chemical and larvicidal properties characterized. Five acylpolyamine toxins (alpha-agatoxins) of molecular masses 452, 488, 489, 504, and 505 Da produce immediate but reversible paralysis in Manduca sexta following injection. Six insecticidal peptides (mu-agatoxins) produce a gradual but irreversible paralysis. The complete amino acid sequences (36-38 residues) of the mu-agatoxins are presented. These peptides contain eight half-cystines and are quite similar in sequence. At least four of these toxins are amidated at the carboxyl terminus. The secondary structure of one of these toxins (mu-Aga V) was investigated.     

Phylogeny and taxonomy of European funnel‐web spiders of the Tegenaria−Malthonica complex (Araneae: Agelenidae) based upon morphological and molecular data

The taxonomy and systematics of European house spiders, currently constituting the ill‐defined Tegenaria?Malthonica complex (including Aterigena) in the family Agelenidae, are revised. In Europe four monophyletic genera and 81 species are defined. One genus, Eratigena gen. nov. , and seven species are described as new; at species level 17 new synonyms and 20 new combinations are proposed, and the original combination of 14 species is reinstated. Five species could not be placed (incertae sedis) because of insufficient material and one taxon is regarded as ‘nomen dubium’. On the basis of a detailed morphological assessment, 88 characters were chosen for a cladistic analysis. Phylogenetically informative characters include mostly spination patterns as well as spinneret and genital structures. In addition to morphology, three gene sections [cytochrome c oxidase subunit 1 (CO1), nicotinamide adenine dinucleotide dehydrogenase subunit 1 (NADH1) 28S] were analysed. Morphological and molecular analyses were performed individually and in combination applying maximum parsimony and Bayesian tree search methods. In all resulting trees Malthonica and Tegenaria in their present composition are either polyphyletic or paraphyletic. Consequently, we redefined the two genera and erected a new genus, Eratigena gen. nov. Identification keys are provided for the European agelenid genera as well as for the European species of Tegenaria and Eratigena gen. nov. The genera and most of the constituent species are described and illustrated. The new classification has also been applied to some extra European members of the Tegenaria‐Malthonica complex resulting in additional three new synonyms, seven reversals to the original combination, and four new combinations. © 2013 The Linnean Society of London     

Ultrastructural observations of spermatozoa of several tetragnathid spiders with phylogenetic implications (Araneae, Tetragnathidae)

The present study reports on the ultrastructure of spermatozoa and spermatogenesis of 12 tetragnathid spiders (10 Tetragnatha species [T. boydi, T. dearmata, T. extensa, T. montana, T. nigrita, T. obtusa, T. pinicola, T. reimoseri, T. shoshone, T. striata]; Pachygnatha listeri and Metellina segmentata). All species develop typical cleistospermia with a coiled nucleus in the center and a coiled axoneme in the periphery of the cell. Remarkable differences in the sperm ultrastructure of the investigated species comprise the shape of the main sperm cell components (nucleus, acrosomal complex, implantation fossa, and centriolar complex). Within the observed Tetragnatha species, three types of sperms were characterized: T. montana-type, T. boydi-type, and T. striata-type. The highly derivative T. montana-type is characterized by the following remarkable features: an extremely elongated nucleus, shaped like a corkscrew and twisted around the axoneme (before coiling); a deep implantation fossa; a corkscrew-shaped acrosomal vacuole; after the coiling process, the nucleus is coiled five to six times in the center of the spermatozoon and the axoneme is coiled five to six times peripheral to the nucleus. The T. boydi-type hardly differs from the T. montana-type, but is remarkable due to the triangular-shaped nucleus (in cross section). The T. striata-type differs especially by a peculiar acrosomal vacuole with a long, slightly curved process and a short appendix, as well as a nucleus that describes only three loose coils around the axoneme (before coiling). The spermatozoa of Pachygnatha listeri and especially Metellina segmentata differ strikingly from the described Tetragnatha-types and are similar to more primitive araneomorph spermatozoa, such as Hypochilus pococki. The described Tetragnatha-types completely correspond with Okuma's (1988a,b, J Fac Agr Kyushu U 32:165-181, 32:183-213) classification of Tetragnatha species. Furthermore, our results suggest an early derivative systematic position of Pachygnatha within Tetragnathinae and the position of Metellina within the Tetragnathidae.     

Speciation on a conveyor belt: sequential colonization of the hawaiian islands by Orsonwelles spiders (Araneae,Linyphiidae)

Spiders of the recently described linyphiid genus Orsonwelles (Araneae, Linyphiidae) are one of the most conspicuous groups of terrestrial arthropods of Hawaiian native forests. There are 13 known Orsonwelles species, and all are single- island endemics. This radiation provides an excellent example of insular gigantism. We reconstructed the cladistic relationships of Orsonwelles species using a combination of morphological and molecular characters (both mitochondrial and nuclear sequences) within a parsimony framework. We explored and quantified the contribution of different character partitions and their sensitivity to changes in the traditional parameters (gap, transition, and transversion costs). The character data show a strong phylogenetic signal, robust to parameter changes. The monophyly of the genus Orsonwelles is strongly supported. The parsimony analysis of all character evidence combined recovered a clade with of all the non-Kauai Orsonwelles species; the species from Kauai form a paraphyletic assemblage with respect to the latter former clade. The biogeographic pattern of the Hawaiian Orsonwelles species is consistent with colonization by island progression, but alternative explanations for our data exist. Although the geographic origin of the radiation remains unknown, it appears that the ancestral colonizing species arrived first on Kauai (or an older island). The ambiguity in the area cladogram (i.e., post-Oahu colonization) is not derived from conflicting or unresolved phylogenetic signal among Orsonwelles species but rather from the number of taxa on the youngest islands. Speciation in Orsonwelles occurred more often within islands (8 of the 12 cladogenic events) than between islands. A molecular clock was rejected for the sequence data. Divergence times were estimated by using the nonparametric rate smoothing method of Sanderson (1997, Mol. Biol. Evol. 14:1218-1231) and the available geological data for calibration. The results suggest that the oldest divergences of Orsonwelles spiders (on Kauai) go back about 4 million years.     

Studies of morphological and molecular phylogenetic divergence in spiders (Araneae: Homalonychus) from the American southwest, including divergence along the Baja California Peninsula

Comparative phylogenetic and phylogeographic analyses have revealed a pervasive midpeninsular divergence in the mitochondrial genealogies of numerous vertebrate taxa distributed on the Baja California Peninsula. In this study, we extend the investigation of regional vicariance in Baja California to an arthropod taxon by examining patterns of phylogenetic and morphological divergence in the spider genus Homalonychus (Araneae, Homalonychidae). We analyzed data from two mtDNA genes (16S rRNA and NADH dehydrogenase subunit (1) and a nuclear gene (28S rRNA) using maximum parsimony and Bayesian phylogenetic analyses, and also conducted geometric morphometric analyses employing landmark data on male and female genitalia. Genes and morphology both reveal a deep split across the Colorado River and Gulf of California, separating Homalonychus selenopoides on the east side of river from its congener Homalonychus theologus on the west side of the river, including the Baja California Peninsula. Along the north-south axis of the Baja Peninsula, an apparently more recent midpeninsular phylogenetic break is evident within H. theologus in the mitochondrial genome and in female genitalia. However, there is no measurable divergence between northern and southern populations in either nuclear DNA or male genitalia. We suggest that this discordance between datasets reflects either a difference in rates of evolution between male versus female systems, or that male-based nuclear gene flow is obscuring a phylogenetic split that is fixed in the female-based systems. Our findings provide additional support for a midpeninsular Baja divergence event, although the timing and geological evidence for such an event remain elusive.     

A total evidence assessment of the phylogeny of North American euctenizine trapdoor spiders (Araneae, Mygalomorphae, Cyrtaucheniidae) using Bayesian inference

North American trapdoor spiders of the subfamily Euctenizinae (Cyrtaucheniidae) are among the most diverse mygalomorph spiders (trapdoor spiders, tarantulas, and their relatives) on the continent in terms of species numbers and ecological habits. We present a generic level phylogenetic study of the subfamily based on a total evidence approach. Our dataset comprises approximately 3.7 kb of molecular characters (18S and 28S rRNA gene sequences) and 71 morphological characters scored for 32 taxa. When analyzed independently, these data sets, particularly the morphology, depict very different views of mygalomorph and euctenizine relationships, albeit with weak support. However, when these data are combined we recover a tree topology that is supported by high posterior probability for most nodes. The combined data recover a phylogenetic pattern for euctenizines different than previously published and indicate the presence of a narrowly endemic new genus from central California. While euctenizine monophyly is unequivocal, the monophyly of a number of other mygalomorph groups is questionable (e.g., Cyrtaucheniidae, Mecicobothriodina, Rastelloidina). This non-monophyly is noteworthy, as our analysis represents the first employing a total evidence approach for mygalomorphs, a group known to be morphologically conservative.     

Molecular phylogenetics at the population/species interface in cave spiders of the southern Appalachians (Araneae:Nesticidae:Nesticus)

This paper focuses on the relationship between population genetic structure and speciation mechanisms in a monophyletic species group of Appalachian cave spiders (Nesticus). Using mtDNA sequence data gathered from 256 individuals, I analyzed patterns of genetic variation within and between populations for three pairs of closely related sister species. Each sister-pair comparison involves taxa with differing distributional and ecological attributes; if these ecological attributes are reflected in basic demographic differences, then speciation might proceed differently across these sister taxa comparisons. Both frequency-based and gene tree analyses reveal that the genetic structure of the Nesticus species studied is characterized by similar and essentially complete population subdivision, regardless of differences in general ecology. These findings contrast with results of prior genetic studies of cave-dwelling arthropods that have typically revealed variation in population structure corresponding to differences in general ecology. Species fragmentation through both extrinsic and intrinsic evolutionary forces has resulted in discrete, perhaps independent, populations within morphologically defined species. Large sequence divergence values observed between populations suggest that this independence may extend well into the past. These patterns of mtDNA genealogical structure and divergence imply that species as morphological lineages are currently more inclusive than basal evolutionary or phylogenetic units, a suggestion that has important implications for the study of speciation mechanisms.