Phylogeny and the evolution of acoustic communication in extant Ensifera (Insecta, Orthoptera)

Ensifera present an appropriate and interesting model for the study of acoustic communication, because of their diverse signal and communication modalities, and due to their accessibility for field and laboratory studies. Several hypotheses have been proposed to explain the acoustic evolution of Ensifera, but they were elaborated without any reference to a falsifiable phylogeny, and were consequently highly speculative. Similarly, phylogenetic relationships between ensiferan clades have not hitherto been studied using modern standard methodology, and the sole cladistic analysis by Gwynne in 1995 was methodologically flawed. No sound hypothesis therefore currently exists for ensiferan phylogeny, which precludes historical analysis of their communication modalities. In the present paper, the phylogeny is established on the basis of morpho‐anatomical characters and used to analyse the evolution of acoustic communication in this clade by mapping the characters related to auditory and stridulatory structures onto the resultant trees. Cladistic analyses resulted in two equi‐parsimonious cladograms (length 154, C 64, CI 58, RI 61) with the following topologies: (1) [(Grylloidea–Gryllotalpidae) (Rhaphidophoridae (Schizodactylidae (Gryllacrididae ((Stenopelmatidae–Cooloola) (Anostostomatidae (Prophalangopsis (Cyphoderris (Tettigoniidae–Lezina))))))))] (2) [(Grylloidea–Gryllotalpidae)(Rhaphidophoridae (Schizodactylidae (Gryllacrididae–Cooloola–(Stenopelmatidae (Anostostomatidae (Prophalangopsis (Cyphoderris (Tettigoniidae–Lezina))))))))]. According to these topologies, Ensifera were ancestrally devoid of acoustic and hearing systems. An acoustic (tegminal or femoro‐abdominal) apparatus appeared a number of times independently with convergent structures. Similarly, tibial tympana developed several times independently. Moreover, four hypotheses (each according to a definite pattern of character transformation) can be proposed to explain the evolution of acoustic communication in the different ensiferan clades and relate it to a definite communicatory context. These hypotheses do not apply equally to ensiferan subclades. Grylloidea and Gryllotalpoidea could have experienced convergently a direct development of an intraspecific acoustic communication. Acoustic communication in Tettigoniidea has evolved more ambiguously, and may either have resulted from a direct evolution analogous to that having occurred in Gryllidea, or have developed in a completely different behavioural context. Future studies of acoustic communication in the different ensiferan clades will have to take into account the fact that the involved structures most often are not homologous and that their evolution may not have taken place in similar conditions. Different hypotheses of acoustic communication evolution may apply to different clades, and there may be no single explanation for acoustic communication in Ensifera.     

Phylogeny of Ensifera (Hexapoda: Orthoptera) using three ribosomal loci, with implications for the evolution of acoustic communication

Representatives of the Orthopteran suborder Ensifera (crickets, katydids, and related insects) are well known for acoustic signals produced in the contexts of courtship and mate recognition. We present a phylogenetic estimate of Ensifera for a sample of 51 taxonomically diverse exemplars, using sequences from 18S, 28S, and 16S rRNA. The results support a monophyletic Ensifera, monophyly of most ensiferan families, and the superfamily Gryllacridoidea which would include Stenopelmatidae, Anostostomatidae, Gryllacrididae, and Lezina. Schizodactylidae was recovered as the sister lineage to Grylloidea, and both Rhaphidophoridae and Tettigoniidae were found to be more closely related to Grylloidea than has been suggested by prior studies. The ambidextrously stridulating haglid Cyphoderris was found to be basal (or sister) to a clade that contains both Grylloidea and Tettigoniidae. Tree comparison tests with the concatenated molecular data found our phylogeny to be significantly better at explaining our data than three recent phylogenetic hypotheses based on morphological characters. A high degree of conflict exists between the molecular and morphological data, possibly indicating that much homoplasy is present in Ensifera, particularly in acoustic structures. In contrast to prior evolutionary hypotheses based on most parsimonious ancestral state reconstructions, we propose that tegminal stridulation and tibial tympana are ancestral to Ensifera and were lost multiple times, especially within the Gryllidae.     

Male and female genitalia in some Libyan species of Acrididae (Orthoptera: Acridoidea)

A comparative study of male and female genitalia was carried out in thirty‐seven Libyan species representing twenty genera of the family Acrididae. An attempt has been made to describe and illustrate the different structures, namely, epiphallus, aedeagus, subgenital plate, supra‐anal plate and cerci of the male, and spermatheca, ovipositor, subgenital plate, supra‐anal plate and cerci of the female, in Acridids, with an aim to discover their significance in order to make the identification of genera and species, together with other generic characters, more perfect and convenient. Distinct family characters are shield or bridge‐shaped condition of epiphallus; presence or absence of dorso‐lateral appendices, oval sclerites and lophi on epiphallus; divided, undivided or flexured condition of aedeagus; presence or absence of gonopore process on aedeagus; condition of apical and pre‐apical diverticula of spermatheca; presence or absence of glandular pouches of Cornstock and Kellog on female subgenital plate; and rudimentary or well developed condition of egg‐guide. Stable characters for separating the subfamilies are taken to be presence or absence of ancorae on epiphallus, long or short condition of aedeagal sclerites; elongate, slender or short and broad condition of ovipositor valves: presence or absence of Jannone's organs and setae on posterior margin of female subgenital plate; and shape of diverticula of spermatheca. Useful generic characters are shape of male subgenital plate, supra‐anal plate and cerci, broad or narrow condition of bridge, presence or absence of branch of bridge connecting lophi with bridge of epiphallus; mono‐, bi‐ or tri‐lobate condition of lophi of epiphallus, length and upcurved or downcurved condition of apical valve of aedeagus, shape of posterior margin of female subgenital plate, presence of setae on the whole posterior margin or confined to lateral margins only; and toothed, tuberculate or smooth condition of ovipositor valves, length of the lateral apodeme in relation to the dorsal valves. Specific characters are shape of egg‐guide of female subgenital plate, shape of ovipositor valves and apical tips, shape of male supra‐anal plate and cerci, size of anterior and posterior lobes of lophi of epiphallus, size and shape of ancorae, shape of apical valves of aedeagus; and size of apical and pre‐apical diverticula and presence of protuberance on pre‐apical diverticulum.     

Coercive Copulation in the Alpine Bushcricket Anonconotus alpinus Yersin (Tettigoniidae: Tettigoniinae: Platycleidini)

Sexual coercion in the form of forced copulation is widespread in the animal kingdom and has been documented in several insect taxa. In crickets and bushcrickets (sub‐order Ensifera), however, mating typically involves luring acts as opposed to forcing acts. The mating behaviour of the tettigoniid Anonconotus alpinus Yersin, which is described in this paper, appears to be unique amongst the Tettigoniidae (and possibly amongst the Ensifera) in that it is coercive, involving forced matings. Males did not stridulate immediately prior to mating but instead leapt on passing females, using their anal cerci as pincers to maintain a hold on the female's abdomen (which was sometimes damaged in the process). Males appeared to lack a sexual refractory period and attempted to copulate again as little as 18 s following the previous mating.     

A topographical and typological comparison of the rodlike setae of ambulatory dactylopodites in decapod crustaceans

The arrangement and external morphology of the rodlike setae and associated structures located on the dactylopodites of the walking legs of six species of decapod crustaceans are compared. The dactyls of littoral species, represented by the rock crab, Cancer antennarius, and the spiny lobster, Panulirus interruptus, have dense tufts and bands of rodlike setae, as is typical of many decapods, and additionally only a few small plumed setae. The arrangement of setae on the dactyls of the recently discovered Galapagos vent crab, Bythograea thermydron, closely resembles that of C. antennarius. Rodlike and long plumed setae occur in about equal numbers on the dactyls of the pelagic anomuran, Pleuroncodes planipes. The dactyls having the fewest rodlike setae are those of the terrestrial hermit crab, Coenobita perlatus, and those of the kelp crab, Pugettia producta, where flat setae typical of Majidae have replaced most rodlike setae. The presence and structures of the terminal pores in rodlike setae vary intra- and interspecifically, possibly as a function of molt stage. Variations in some features of rodlike setae, such as tip acuity and presence of microsetae and surface sculpting, appear to be related to development. Serrated setae occur on the dactyls of megalopal P. producta but not in later stages. The topography and typology of setae located on the ambulatory dactyls of decapod crustaceans are considered in light of recent interest in using setal characteristics to determine the sensory functions of sensilla and to clarify the phylogeny of arthropod groups.     

The scolopidial accessory organ in the Jerusalem cricket (Orthoptera: Stenopelmatidae)

Multiple mechanosensory organs form the subgenual organ complex in orthopteroid insects, located in the proximal tibia. In several Ensifera (Orthoptera), a small chordotonal organ, the so-called accessory organ, is the most posterior part of this sensory complex. In order to document the presence of this accessory organ among the Ensifera, the chordotonal sensilla and their innervation in the posterior tibia of two species of Jerusalem crickets (Stenopelmatidae: Stenopelmatus) is described. The sensory structures were stained by axonal tracing. Scolopidial sensilla occur in the posterior subgenual organ and the accessory organ in all leg pairs. The accessory organ contains 10–17 scolopidial sensilla. Both groups of sensilla are commonly spatially separated. However, in few cases neuronal fibres occurred between both organs. The two sensillum groups are considered as separate organs by the general spatial separation and innervation by different nerve branches. A functional role for mechanoreception is considered: since the accessory organ is located closely under the cuticle, sensilla may be suited to detect vibrations transferred over the leg's surface. This study extends the known taxa with an accessory organ, which occurs in several taxa of Ensifera. Comparative neuroanatomy thus suggests that the accessory organ may be conserved at least in Tettigoniidea.     

Evolution of antennules of cytheroidean ostracods (Crustacea)

Phylogenetic analysis and fossil records indicate that antennules with whip-like setae are the most plesiomorphic state in cytheroidean ostracods and that antennulae with claw-like setae are derived from antennulae with whip-like setae. Character distributions on the 18S rDNA molecular phylogenetic tree suggest that two phenotypic features of cytheroidean antennules (W/L ratio and claw-like/whip-like setae) have morphological plasticity. These features have evolved as an ethological adaptation rather than due to phylogenetic constraints such as the evolution of shell outline in cytheroideans. However, the species of the Leptocytheridae–Trachyleberididae clade generally have stout, robust antennules, indicating a phylogenetic constraint. The character state of setae (claw/whip) in cytheroidean ostracods is reflected more through their ethology than the W/L ratio of the antennules. On the basis of the present analysis and the fossil record, diversification in the morphology of the antennules seems to have occurred during the early Mesozoic.     

Tarsal attachment devices of the southern green stink bug Nezara viridula (Heteroptera: Pentatomidae)

Based on analyses with cryo‐scanning and transmission electron microscopy, the present study reports on the morphology and ultrastructure of the attachment structures of the green stinkbug Nezara viridula L. (Heteroptera: Pentatomidae), a cosmopolitan pest of different crops in most areas of the world. In addition, the presence and distribution of large proportions of the elastic protein resilin in these structures was revealed by confocal laser scanning microscopy. The attachment structures of each leg comprise two sclerotised claws, a pair of smooth flexible pulvilli and a hairy adhesive pad located at the ventral side of the basitarsus. No sexual dimorphism is evident. Contact areas of resting individuals on a smooth surface show that N. viridula creates contact to the substrate with the ventral surface of (a) the distal portions of the pulvilli, (b) the setae of the hairy adhesive pad, (c) the two paraempodia representing mechanosensory setae, and (d) the tips of the claws. Each pulvillus is a sac‐like structure formed by complex cuticular layers that vary in their structure and resilin content. The dorsal side consists of sclerotised chitinous material, while the ventral cuticle consists mainly of resilin and shows a very thin epicuticle and a thick exocuticle. The setae of the hairy adhesive pad are pointed and socketed. They exhibit a pronounced longitudinal gradient in the material composition, with large proportions of resilin being present in the setal tips. In most of these setae, especially in those of the distal‐most part of the pad, also a transverse gradient in the material composition is visible.     

Copulation behaviour of Glossina pallidipes (Diptera: Muscidae) outside and inside the female, with a discussion of genitalic evolution

If species-specific male genitalia are courtship devices under sexual selection by cryptic female choice, then species-specific aspects of the morphology and behaviour of male genitalia should often function to stimulate the female during copulation. The morphology and behaviour of the complex, species-specific male genitalia of the tsetse fly, Glossina pallidipes Austen, were determined from both direct observations and dissections of flash-frozen copulating pairs; we found that some male genitalic traits probably function to stimulate the female, while others function to restrain her. The male clamps the ventral surface of the female's abdomen tightly with his powerful cerci. Clamping does not always result in intromission. Clamping bends the female's body wall and her internal reproductive tract sharply, posteriorly and dorsally, and pinches them tightly. The male performed sustained, complex, stereotyped, rhythmic squeezing movements with his cerci that were not necessary to mechanically restrain the female and appeared instead to have a stimulatory function. Six different groups of modified setae on and near the male's genitalia rub directly against particular sites on the female during squeezing. The designs of these setae correlate with the force with which they press on the female and the probable sensitivity of the female surfaces that they contact. As expected under the hypothesis that these structures are under sexual selection by female choice, several traits suspected to have stimulatory functions have diverged in G. pallidipes and its close relative, G. longipalpis. Additional male non-genitalic behaviour during copulation, redescribed more precisely than in previous publications, is also likely to have a courtship function. The elaborate copulatory courtship behaviour and male genitalia may provide the stimuli that previous studies showed to induce female ovulation and resistance to remating.     

The evolution of pedipalps and glandular hairs as predatory devices in harvestmen (Arachnida,Opiliones)

Pedipalps are the most versatile appendages of arachnids. They can be equipped with spines (Amblypygi), chelae (Scorpiones), or adhesive pads (Solifugae), all of which are modifications to grasp and handle fast‐moving prey. Harvestmen (Opiliones) show a high diversity of pedipalpal morphologies. Some are obviously related to prey capture, like the enlargement and heavy spination of Laniatores pedipalps. Many Dyspnoi, by contrast, exhibit thin, thread‐like pedipalps that are covered with complex glandular setae (clavate setae). These extrude viscoelastic glue that is used to immobilize prey items. Comparable setae (plumose setae) have previously been found in representatives of both Eupnoi and Dyspnoi, yet comprehensive data on their distribution are lacking. This study examined the distribution and ultrastructure of glandular setae in harvestmen and related them to pedipalpal morphology. Pedipalpal and setal characters were analysed in a phylogenetic framework. We found that glandular setae are synapomorphic for and widespread in the Palpatores clade (Eupnoi plus Dyspnoi). Their occurrence correlates with pedipalp morphology and feeding habit. Remnants of arthropod cuticular structures or secretions, frequently found attached to glandular setae, and behavioural observations, underlined the importance of the setae for capturing and securing prey. We hypothesize that glandular setae evolved as an adaptation to capture small and agile prey, which are hard to catch with a capture basket. Details of ultrastructure indicate that the setae are derived sensilla chaetica, with both a secretory and sensory function. Derived ultrastructural characters of the glandular setae, such as slit‐like channel openings and a globular arrangement of the microtrichia, may increase their effectiveness. The functional role of further pedipalpal modifications, such as apophyses, stalked and hyperbendable joints, and curved segments, as well as sexual dimorphism and ontogenetic polymorphism, are discussed. Some implications of the results obtained for the taxonomic treatment of Phalangiidae are also discussed. These results shed new light on the biology and evolutionary history of this fascinating group of arthropods.     

Tinaminyssus juxtamelloi sp. n. (Acari: Dermanyssidae; Rhinonyssinae) from the nasal passages of the band-tailed pigeon, Columba fasciata, in New Mexico.

Tinaminyssus juxtamelloi sp. n. is described from the nasal passages of Columba fasciata in New Mexico. The new species is most similar to Tinaminyssus melloi (Castro) 1948 and T. turturi (Fain) 1962, but differs in (1) possessing only 5 pairs of ventral opisthosomal setae, (2) presence of 3 pairs of enlarged setae on the dorsal opisthosoma at the posterolateral margin of the podosomal plate (1 pair) and at the lateral margins of the opisthosomal plate (2 pairs), (3) elongate shape and larger size of the poststigmatic plates, and (4) chaetotaxy and solenidiotaxy of the legs, especially tarsus I with a cluster of 4 solenida and 1 club-shaped solenidion on the apex of the dorsum. The relationships of this with allied species of the genus Tinaminyssus from columbiform birds are briefly discussed.     

Position and definition of the genus Paegniodes Eaton, 1881 based on redescription on the type species Paegniodes cupulatus (Eaton, 1871) (Ephemeroptera: Heptageniidae)

The genus Paegniodes Eaton, 1881 and its type species P. cupulatus (Eaton, 1871) have never been described in detail completely. In this article, all stages are described and photographed. Compared to other heptageniid genera, the adults of P. cupulatus have a more colourful body, smaller hindwings but larger titillators. The nymphs of this species have no setae or tubercles on the body and no emarginations on the head capsule. They have smaller lamellae of the first gills, a wider labrum with a median notch, mandibles without setae on outer margins, maxillae with a row of setae and scattered setae on the ventral surface, cerci with mesal setae, and a median filament with setae on both sides. Both imaginal and nymphal stages of the genus have unique characters, so the taxon Paegniodes is recognized at the rank of genus here. It appears closely related to Rhithrogena Eaton, 1881.     

蜣螂与壁虎刚毛的比较及改形对其功能的影响

脱附与粘附是工程作业的两个相互矛盾的难题,土壤和物料的粘附严重降低了机械的作业效率和质量,没有粘附又使某些机械无法工作或运行。目前世界流行的脱附技术,包括向界面注入空气的充气法、向界面注入溶液的充液法、利用机械或超声波的振动法、施加电场的电渗法、施加磁场的磁     

Experimental evidence for friction-enhancing integumentary modifications of chameleons and associated functional and evolutionary implications

The striking morphological convergence of hair-like integumentary derivatives of lizards and arthropods (spiders and insects) demonstrates the importance of such features for enhancing purchase on the locomotor substrate. These pilose structures are responsible for the unique tractive abilities of these groups of animals, enabling them to move with seeming ease on overhanging and inverted surfaces, and to traverse inclined smooth substrates. Three groups of lizards are well known for bearing adhesion-promoting setae on their digits: geckos, anoles and skinks. Similar features are also found on the ventral subdigital and distal caudal skin of chameleons. These have only recently been described in any detail, and structurally and functionally are much less well understood than are the setae of geckos and anoles. The seta-like structures of chameleons are not branched (a characteristic of many geckos), nor do they terminate in spatulate tips (which is characteristic of geckos, anoles and skinks). They are densely packed and have attenuated blunt, globose tips or broad, blade-like shafts that are flattened for much of their length. Using a force transducer, we tested the hypothesis that these structures enhance friction and demonstrate that the pilose skin has a greater frictional coefficient than does the smooth skin of these animals. Our results are consistent with friction being generated as a result of side contact of the integumentary filaments. We discuss the evolutionary and functional implications of these seta-like structures in comparison with those typical of other lizard groups and with the properties of seta-mimicking synthetic structures.     

Epibiotic bacteria associated with the recently discovered Yeti crab, Kiwa hirsuta

The Yeti crab, Kiwa hirsuta Macpherson et al., is the single known species in a recently discovered crab family Kiwaidae (Decapoda: Galatheoidea) from deep-sea hydrothermal vents. Its chelipeds, walking legs and the ventral surface of its cephalothorax are covered with dense setae that, in turn, are covered with clusters of filamentous bacteria, making the crab appear extraordinarily 'hairy'. Electron microscopy revealed dense bacterial clusters attached to the chitinous outer layer of the setae. Molecular phylogenetic analyses revealed the setae-associated bacteria to be dominated by epsilon-Proteobacteria ( approximately 56% of the recovered ribotypes), gamma-Proteobacteria ( approximately 25%) and Bacteroidetes ( approximately 10%). Fluorescence in situ microscopy confirmed the attachment of filamentous epsilon-Proteobacteria on setae, but no specialized morphological structures appeared to exist for bacterial attachment. Key enzymes involved in the reductive tricarboxylic acid cycle (ATP-dependent citrate lyase) and sulfite oxidation or dissimilatory sulfate reduction (bidirectional APS reductase) were detected. Consequently, the potential for carbon fixation and cycling of reduced and oxidized sulfur appear to exist in the dense microflora that grows on the crab's setae.     

Neue Nerillidae (Archiannelida) aus dem Sublitoral der IMordsee und des Mittelatlantik (Nordwest-Afrika)

Bathynerilta marina gen. et sp.n., Bathychaetus heptapous gen. et sp.n. (both off the east coast of Scotland), and Afronerilla hartwigi gen. et sp.n. (off the coast of Mauretania) are described; all from the sublittoral. The main features of Bathynerilla marina are: 8 segments with capillary setae; parapodial cirri on each segment; without anal cirri. The features of Afronerilla hartwigi are: 8 segments with capillary setae; without palps, tentacles, and anal cirri; the parapodial cirri on segment 1 and 8 are absent. The main features of Bathychaetus heptapous are: a pharyngeal bulb with a pair of cuticulary jaw-like structures; 7 segments with capillary setae; parapodial cirri on each segment present; without tentacles.     

Directional sensitivity of wind-sensitive giant interneurons in the cave cricket Troglophilus neglectus.

Unlike the situation in most cockroach and cricket species studied so far, the wind-sensitive cerci of the cave cricket Troglophilus neglectus Krauss (Rhaphidophoridae, Orthoptera) are not oriented parallel to the body axis but perpendicular to it. The effects of this difference on the morphology, and directional sensitivity of cercal giant interneurons (GIs), were investigated. In order to test the hypothesis that the 90 degrees change in cercal orientation causes a corresponding shift in directional sensitivity of GIs, their responses in both the horizontal and vertical planes were tested. One ventral and four dorsal GIs (corresponding to GIs 9-1a and 9-2a, 9-3a, 10-2a, 10-3a of gryllid crickets) were identified. The ventral GI 9-1a of Troglophilus differed somewhat from its cricket homologue in its dendritic arborisation and its directional sensitivity in the horizontal plane. The morphology and horizontal directionality of the dorsal GIs closely resembled that of their counterparts in gryllids. In the vertical plane, the directionality of all GIs tested was similar. They were all excited mainly by wind puffs from the axon-ipsilateral quadrant. The results suggest that directional sensitivity to air currents in the horizontal plane is maintained despite the altered orientation of the cerci. This is presumably due to compensatory modifications in the directional pReferences of the filiform hairs.     

One hundred years of instability in ensiferan relationships

Although Ensifera is a major insect model group, its phylogenetic relationships have been understudied so far. Few phylogenetic hypotheses have been proposed, either with morphological or molecular data. The largest dataset ever used for phylogeny reconstruction on this group is molecular (16S rRNA, 18S rRNA and 28S rRNA sequences for 51 ensiferan species), which has been used twice with different resultant topologies. However, only one of these hypotheses has been adopted commonly as a reference classification. Here we re‐analyse this molecular dataset with different methods and parameters to test the robustness and the stability of the adopted phylogeny. Our study reveals the instability of phylogenetic relationships derived from this dataset, especially for the deepest nodes of the group, and suggests some guidelines for future studies. The comparison between the different classifications proposed in the past 70 years for Ensifera and our results allows the identification of potential monophyletic clades (katydids, mole crickets, scaly crickets + Malgasia, true crickets, leaf roller crickets, cave crickets) and the remaining unresolved clades (wetas, Jerusalem crickets and most of the highest rank clades) in Ensifera phylogeny.     

Surface structures of the antennular flagella of the hermit crab Pagurus alaskensis (Benedict): A light and scanning electron microscopy study

The surface structures of the antennular flagella of Pagurus alaskensis are described in detail. Attention is directed towards the surface morphology of two types of possible sensilla: (1) exoskeletal pores (1.0–3.0 μm in diameter); (2) setae of various kinds. In addition, small (0.1–0.2 μm) pits occur in the exoskeleton which are not considered to be sensory in function. The exoskeletal pores are found at fairly specific locations on both the inner and outer flagella, particularly on the short segments of the outer flagella. Neither the inner nor the outer flagella are bilaterally symmetrical with respect to their setal armature. On the outer flagellum six groups of setae may be distinguished: lateralmesial; dorsal; ventral; accessory; aesthetasc; setae of the distal segment. On the inner flagellum setae of the mesial and lateral rows form distinctive groups. The morphology, orientation and locations of all the flagellar setae are defined and where possible the numbers of the various morphological types within the specific setal groups are given. It is noteworthy that many setal types have obvious apical pores and yet no pores could be found in the chemoreceptive aesthetasc setae. The functions of the various setae are discussed in relation to their topographical position and to existing electrophysiological and behavioral data. Some suggestions are made about future experiments to demonstrate the central connections of specific sensilla or groups of sensilla and to show their significance in the whole animal.