Comparative morphology of Van der Vecht's organ in Polistes social parasites: host ecology and adaptation of the parasite

Camouflage strategies are common in insect social parasites. Being accepted into an alien colony as a dominant nestmate favours behavioural and morphological adaptations to mimic a specific odour. In Polistes social parasites, abdominal tegumental glands are involved in this camouflage strategy. These glands secreting cuticular hydrocarbons are connected with a modified cuticular area of the last gastral sternite of female wasps, named Van der Vecht's organ, whose secretion is involved in rank and dominance recognition. The size of this exocrine area has been demonstrated to be under selective pressure in Polistes, as a response to an efficient dominance recognition. Because chemical and behavioural integration differs between parasitic species, we carried out a comparison of Van der Vecht's organ size between the three Polistes social parasites and their respective hosts. The parasites Polistes sulcifer and Polistes semenowi, capable of a rapid chemical mimicry and specialized to exploit a lowland host, also show an enlarged Van der Vecht's organ. Conversely, the parasite Polistes atrimandibularis, specialized on a mountain species and showing a slow chemical integration, has a smaller organ. The time available for the parasite to tune up its chemical mimicry, before the emergence of workers to be accepted as a dominant nestmate, appears to be the most important selective pressure acting on the size of this abdominal organ. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013 , 109 , 313–319.     

Searching factors causing implausible non-monophyly: ssu rDNA phylogeny of Isopoda Asellota (Crustacea: Peracarida) and faster evolution in marine than in freshwater habitats

This contribution addresses two questions: which alignment patterns are causing non-monophyly of the Asellota and what is the phylogenetic history of this group? The Asellota are small benthic crustaceans occurring in most aquatic habitats. In view of the complex morphological apomorphies known for this group, monophyly of the Asellota has never been questioned. Using ssu rDNA sequences of outgroups and of 16 asellote species from fresh water, littoral marine habitats and from deep-sea localities, the early divergence between the lineages in fresh water and in the ocean, and the monophyly of the deep-sea taxon Munnopsidae are confirmed. Relative substitution rates of freshwater species are much lower than in other isopod species, rates being highest in some littoral marine genera (Carpias and Jaera). Furthermore, more sequence sites are variable in marine than in freshwater species, the latter conserve outgroup character states. Monophyly is recovered with parsimony methods, but not with distance and maximum likelihood analyses, which tear apart the marine from the freshwater species. The information content of alignments was studied with spectra of supporting positions. The scarcity of signal (=apomorphic nucleotides) supporting monophyly of the Asellota is attributed to a short stem-line of this group or to erosion of signal in fast evolving marine species. Parametric boostrapping in combination with spectra indicates that a tree model cannot explain the data and that monophyly of the Asellota should not be rejected even though many topologies do not recover this taxon.     

Developmental Changes in the Sterol Composition and the Glycerol Content of Cuticular and Internal Lipids of Three Species of Flies

The glycerol concentration and the composition of cuticular and internal sterols in three medically and forensically important fly species, viz., Musca domestica, Sarcophaga carnaria, and Calliphora vicina, were analyzed. The cuticular and internal lipid extracts were separated by HPLC‐LLSD, after which the sterol fraction was characterized by GC/MS in total ion current (TIC) mode. The cuticular lipids of M. domestica larvae contained seven sterols, while in pupae and females, six sterols were identified. Five sterols were found in the cuticular lipids of M. domestica males. The internal lipids of M. domestica larvae and pupae contained six and seven sterols, respectively, while those of male and female flies contained only five sterols. Sitosterol, cholesterol, and campesterol were the dominant sterols in M. domestica, while campestanol, stigmasterol, sitostanol, and fucosterol were identified in low concentrations or in traces. In contrast, cuticular and internal lipids of S. carnaria and C. vicina contained only cholesterol. Glycerol was identified in all stages of M. domestica, S. carnaria, and C. vicina. For all the three examined fly species, the present study clearly showed species‐specific developmental changes in the composition of cuticular and internal sterols as well as in the glycerol concentration.     

Chemical and behavioral analysis of the cuticular hydrocarbons from Asian citrus psyllid,Diaphorina citri

Huanglongbing (HLB) is the most destructive disease of citrus worldwide. The Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Psyllidae), is the vector of the phloem‐inhabiting bacterium, Candidatus Liberibacter asiaticus, which is presumed to cause HLB in Florida citrus. Laboratory and field studies were conducted to examine the behavioral responses of male and female D. citri to their cuticular extracts. In olfactometer assays, more male D. citri were attracted to one, five, or 10 female cuticular extract equivalent units than blank controls. The results were confirmed in field studies in which clear or yellow traps baited with 10 female cuticular extract equivalent units attracted proportionately more males than clear traps baited with male cuticular extract or unbaited traps. Analyses of cuticular constituents of male and female D. citri revealed differences between the sexes in chemical composition of their cuticular extracts. Laboratory bioassays with synthetic chemicals identified from cuticular extracts indicated that dodecanoic acid attracted more males than clean air. Traps baited with dodecanoic acid did not increase total catch of D. citri as compared with blank traps at the dosages tested; however, the sex ratio of psyllid catch was male biased on traps baited with the highest lure loading dosage tested (10.0 mg).     

An ultrastructural approach to feeding behaviour in Philodina roseola and Brachionus calyciflorus (rotifers) II. The oesophagus

The cuticular oesophagus is a simple expansion of the dorsal pharyngeal wall of the mastax. The ciliary oesophagus is the cellular anterior wall of the stomach lumen, but seems to have the same embryological origin as the pharynx.In Brachionus calyciflorus, its cilia are surrounded by cuticular velums which have the same myelin-like structure and the same function as the buccal velum of Philodina roseola. In all cases, the oesophagus prevents the return of food particles from the stomach to the mastax lumen.     

Application of Two‐Way Hierarchical Cluster Analysis for the Identification of Similarities between the Individual Lipid Fractions of Lucilia sericata

The composition of the cuticular and internal lipids of larvae and pupae of Lucilia sericata was studied using chromatographic techniques. The lipids from both stages of L. sericata had similar free fatty acid (FFA) profiles and also contained alcohols and cholesterol. The range of the number of C‐atoms detected for these classes of compounds was to some extent similar in larvae and pupae, but the relative amounts of each class differed between stages. Saturated as well as unsaturated FFAs with even and odd numbered C‐atom chains were present in both cuticular and internal lipids. The alcohol fractions of L. sericata were represented by free, straight‐chain primary alcohols containing an even number of C‐atoms. The lipid composition of male and female L. sericata adults and the hydrocarbon composition of all stages of L. sericata had previously been analyzed. To have a full overview of the lipid composition and to identify similarities or dissimilarities between the individual lipid fractions in this insect species, two‐way hierarchical cluster analysis (HCA) was performed using also the data from these previous publications. The content of FFA 18 : 1 (n‐9) was noticed to be very high in the cuticular fractions of larvae and pupae as well as in all internal fractions (male, female, larvae, and pupae) and low in the cuticular fractions of male and female imago. The contents of FFAs 16 : 0 and 16 : 1 (n‐9), cholesterol, and the n‐alkanes n‐C₃₁, n‐C₂₉, n‐C₂₇, n‐C₂₅, and n‐C₂₃ varied between particular fractions, whereas the amounts of other compounds were similar in all fractions.     

New findings on the origin of Cavia intermedia,one of the world's rarest mammals

Cavia intermedia, a rare species of cavy found exclusively on Moleques do Sul Island, Brazil, differs from other cavies in genetic and morphological characteristics and has been regarded as phylogenetically closest to Cavia magna. Here, I present morphological evidence for the likely origin of Cavia intermedia from an ancestor shared with Cavia magna: an external female genital organ similar in size and appearance to the male's penis. Masculinization of the female genitalia has been observed in Cavia magna but does not occur in Cavia aperea, which suggests a sister relationship between Cavia magna and Cavia intermedia.     

Early reproduction and increased reproductive allocation in metal-adapted populations of the terrestrial isopod Porcellio scaber

Organisms inhabiting metal-contaminated areas can be stressed by metal exposure and are possibly subject to selection, resulting in increased metal tolerance and changes in growth and/or reproduction characteristics. In a previous study it was found that in the terrestrial isopod Porcellio scaber, sampled from the vicinity of a zine smelter, the body size was small and the brood size was large compared to isopods from a reference area. To assess whether these differences were due to genetic differentiation between strains, isopods collected from a reference wood, a zinc smelter area and a lead mine were cultured on non-polluted food, while growth, reproduction and metal concentrations were studied in first and second laboratory generations. The isopods from the three populations differed in age and weight at first reproduction, although there were hardly any differences in growth. The females of the mine and the smelter population started to reproduce earlier, at a lower weight, which resulted in fewer young per female. However, reproductive allocation (=wight of young relative to the weight of the mother) was higher in mine and smelter isopods. We conclude that the isopods at the metal-contaminated sites have been selected for early reproduction and increased reproductive allocation. The results indicate that populations inhabiting metal-polluted sites have probably undergone evolutionary changes. This study showed that growth and reproduction characteristics of different populations under laboratory conditions may provide information on selection processes in the field.     

Evolution of insect pheromones and their role in reproductive isolation and speciation

The importance for reproductive isolation of a change in the pheromone biosynthetic pathway, resulting in a different pheromone blend, is discussed in Lepidoptera and Diptera. The different sites of pheromone production and the biosynthetic enzymes are briefly reviewed. Two examples of a modification in the pheromone blend leading to reproductive isolation in Lepidoptera are taken as examples: the first, in Ostrinia nubilalis, is involved in the formation of two different populations showing reproductive isolation; the second, in the genus Ostrinia, might be at the origin of the formation of two different species. In both examples, a modification in the function of a desaturase involved in pheromone biosynthesis brings about change in the pheromone blend. In the fruitfly, Drosophila melanogaster, a mutation at a desaturase locus leads to the formation of two populations which differ in their pheromone mixtures and have developed premating isolation. The closely related species, D. melanogaster and D. simulans, differ in their female pheromonal cuticular hydrocarbons. This pheromonal difference is due to two species- and female-specific genes, desatF and eloF. The activity of desatF could account for an effective barrier between these species. All these examples show that a birth and death process of desaturases is at the origin of major shifts in the pheromone blend leading to sexual isolation and speciation.     

Diversity of isopods (Crustacea): new data from the Arctic and Atlantic Oceans

New data on the diversity pattern of isopods (Crustacea) from the northern most part of the North Atlantic and the Arctic Oceans is presented. The pattern of diversity with depth is similar at depths <1000m, but differs considerably below about 1000m. In the Arctic the diversity of isopods (expressed both as numbers of species per sled and expected number of species) increased with increased depth to a maximum at depths of about 320 to 1100m, but then declined towards deeper waters. There was a significant increase in numbers per sled and in the expected number of species with increased depth in the northernmost part of the North Atlantic Ocean. Additionally, changes occurred in the relative composition of the shallow and deep water fauna, with asellote isopods being relatively larger part of the isopod fauna in the Arctic than in the northern most part of the North Atlantic. This indicates major faunistic changes occurring at the Greenland-Iceland-Faeroe Ridge, possibly caused by rapid changes in the temperature. Furthermore, that the low diversity of the Arctic deep-sea is a regional phenomenon, and not a part of a large scale latitudinal pattern in the North Atlantic.     

Pathogenic effect of entomopathogenic nematode-bacterium complexes on terrestrial isopods

In this study, we evaluated the effect of entomopathogenic nematodes (EPNs) Steinernema carpocapsae, Steinernema feltiae and Heterorhabditis bacteriophora, symbiotically associated with bacteria of the genera Xenorhabdus or Photorhabdus, on the survival of eight terrestrial isopod species. The EPN species S. carpocapsae and H. bacteriophora reduced the survival of six isopod species while S. feltiae reduced survival for two species. Two terrestrial isopod species tested (Armadillidium vulgare and Armadillo officinalis) were found not to be affected by treatment with EPNs while the six other isopod species showed survival reduction with at least one EPN species. By using aposymbiotic S. carpocapsae (i.e. without Xenorhabdus symbionts), we showed that nematodes can be isopod pathogens on their own. Nevertheless, symbiotic nematodes were more pathogenic for isopods than aposymbiotic ones showing that bacteria acted synergistically with their nematodes to kill isopods. By direct injection of entomopathogenic bacteria into isopod hemolymph, we showed that bacteria had a pathogenic effect on terrestrial isopods even if they appeared unable to multiply within isopod hemolymphs. A developmental study of EPNs in isopods showed that two of them (S. carpocapsae and H. bacteriophora) were able to develop while S. feltiae could not. No EPN species were able to produce offspring emerging from isopods. We conclude that EPN and their bacteria can be pathogens for terrestrial isopods but that such hosts represent a reproductive dead-end for them. Thus, terrestrial isopods appear not to be alternative hosts for EPN populations maintained in the absence of insects.     

Female genitalia,reproductive cycle and sperm storage in Armadillidium vulgare (Crustacea,Isopoda, Oniscidea)

Light and electron microscopic studies that we have published in the past have reported many aspects of the reproductive process in Armadillidium vulgare with particular emphasis on the reconstruction of female genitalia. Together this body of work provides an almost complete, albeit fragmented picture of these processes and include many data on sperm storage and sperm translocation. Females of A. vulgare have a pair of cuticular genitalia in the lumen of the oviduct. For insemination, these genitalia can receive the copulatory organs of males formed by the elongated tips of the first two pleon endopods. During transitions between reproductive (parturial) and non-reproductive (normal, non-parturial) moult cycles the genitalia undergo intriguing structural changes resulting in two types of genitalia. Throughout their reproductive lifetime, either type is reconstructed after each moult depending on the reproductive phase of the female. In this review, we integrate the events that occur during a reproductive cycle with particular emphasis on the genitalia reconstruction and sperm storage, and discuss functional aspects of the genitalia. Thereby, we provide a case model that can be useful for further studies on genitalia diversity and female reproductive strategies in terrestrial isopods.     

Morphology of the prothoracic discs and associated sensilla of Acanthocnemus nigricans (Coleoptera, Acanthocnemidae)

The Australian ‘little ash beetle’ Acanthocnemus nigricans (Coleoptera, Cleroidea, Acanthocnemidae) is attracted by forest fires. A. nigricans has one pair of unique prothoracic sensory organs and it has been speculated that these organs may play a role in fire detection. Each organ consists of a cuticular disc, which is fixed over an air-filled cavity. On the outer surface of the disc, about 90 tiny cuticular sensilla are situated. The poreless outer peg of a sensillum is 3–5 μm long and is surrounded by a cuticular wall. One ciliary sensory cell innervates the peg. As a special feature, the outer dendritic segment is very short already terminating below the cuticle. A massive electron-dense cylindrical rod, which most probably represents the hypertrophied dendritic sheath, extends through the cuticular canal connecting the tip of the outer dendritic segment to the peg. The dendritic inner segment and the soma are fused indistinguishably. Thin, leaflike extensions of glial cells deeply extend into that conjoint and considerably enlarged compartment which also contains large numbers of mitochondria. In summary, the sensilla of the sensory disc of A. nigricans represent a new type of insect sensillum of hitherto unknown function. The possible role of the prothoracic sensory organ in fire detection is discussed.     

Global diversity of Isopod crustaceans (Crustacea; Isopoda) in freshwater

The isopod crustaceans are diverse both morphologically and in described species numbers. Nearly 950 described species (∼9% of all isopods) live in continental waters, and possibly 1,400 species remain undescribed. The high frequency of cryptic species suggests that these figures are underestimates. Several major freshwater taxa have ancient biogeographic patterns dating from the division of the continents into Laurasia (Asellidae, Stenasellidae) and Gondwana (Phreatoicidea, Protojaniridae and Heterias). The suborder Asellota has the most described freshwater species, mostly in the families Asellidae and Stenasellidae. The suborder Phreatoicidea has the largest number of endemic genera. Other primary freshwater taxa have small numbers of described species, although more species are being discovered, especially in the southern hemisphere. The Oniscidea, although primarily terrestrial, has a small number of freshwater species. A diverse group of more derived isopods, the ‘Flabellifera’ sensu lato has regionally important species richness, such as in the Amazon River. These taxa are transitional between marine and freshwater realms and represent multiple colonisations of continental habitats. Most species of freshwater isopods species and many genera are narrow range endemics. This endemism ensures that human demand for fresh water will place these isopods at an increasing risk of extinction, as has already happened in a few documented cases. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Exoskeletal cuticle of cavernicolous and epigean terrestrial isopods: A review and perspectives

Comparative ultrastructural studies of the integument in terrestrial isopod crustaceans show that specific environmental adaptations of different eco-morphotypes are reflected in cuticle structure. The biphasic molting in isopods is a valuable experimental model for studies of cuticular matrix secretion and degradation in the same animal. The aim of this review is to show structural and functional adaptations of the tergal cuticle in terrestrial isopods inhabiting cave habitats. Exoskeletal cuticle thickness, the number of cuticular layers, epicuticle structure, mineralization, pigmentation and complexity of sensory structures are compared, with greater focus on the well-studied cave trichoniscid Titanethes albus. A large number of thinner cuticular layers in cave isopods compared to fewer thicker cuticular layers in related epigean species of similar body-sizes is explained as a specific adaptation to the cavernicolous life style. The epicuticle structure and composition are compared in relation to their potential waterproofing capacity in different environments. Cuticle mineralization is described from the functional point of view as well as from the aspect of different calcium storage sites and calcium dynamics during the molt cycle. We also discuss the nature and reduction of pigmentation in the cave environment and outline perspectives for future research.     

Mate guarding behaviour in the supralittoral isopod,Ligia dentipes (Oniscidea) from the Andaman and Nicobar Islands

Precopulatory mate guarding is a characteristic feature in the mating behaviour of many Malacostraca, and a necessary prerequisite for those species in which female receptivity for males is restricted to a short period of time after the pubertal/reproductive moult. This study deals with the pre-mate guarding behaviour of the semi-terrestrial isopod Ligia dentipes living in the crevices of coral boulders and rocks in the supralittoral region of the Andaman Islands. As in other isopods, moulting in L. dentipes is biphasic, in which the posterior body part invariably moults first. The guarding male aids the female partner in the removal of the moulted exoskeleton. Mating occurs immediately after the posterior body exuviates. The male leaves the female after copulation and goes in search of another receptive female, demonstrating a polygamous mating system in these isopods. The mated females also re-mate with several other males without mate guarding. Females that had mated several times produced more young, compared to females mated only once in the laboratory. Female receptivity ceases following moulting of the anterior half. Intrasexual encounters among males lead to the large males acquiring receptive females. This study reveals interesting deviations from the general pattern of mate guarding already reported in other isopods and decapods. The evolutionary and ecological significances of mate guarding, intrasexual and intersexual conflicts, found in these semi-terrestrial isopods, are discussed.     

Evolution of sexually dimorphic pheromone profiles coincides with increased number of male‐specific chemosensory organs in Drosophila prolongata

Binary communication systems that involve sex‐specific signaling and sex‐specific signal perception play a key role in sexual selection and in the evolution of sexually dimorphic traits. The driving forces and genetic changes underlying such traits can be investigated in systems where sex‐specific signaling and perception have emerged recently and show evidence of potential coevolution. A promising model is found in Drosophila prolongata, which exhibits a species‐specific increase in the number of male chemosensory bristles. We show that this transition coincides with recent evolutionary changes in cuticular hydrocarbon (CHC) profiles. Long‐chain CHCs that are sexually monomorphic in the closest relatives of D. prolongata (D. rhopaloa, D. carrolli, D. kurseongensis, and D. fuyamai) are strongly male‐biased in this species. We also identify an intraspecific female‐limited polymorphism, where some females have male‐like CHC profiles. Both the origin of sexually dimorphic CHC profiles and the female‐limited polymorphism in D. prolongata involve changes in the relative amounts of three mono‐alkene homologs, 9‐tricosene, 9‐pentacosene, and 9‐heptacosene, all of which share a common biosynthetic origin and point to a potentially simple genetic change underlying these traits. Our results suggest that pheromone synthesis may have coevolved with chemosensory perception and open the way for reconstructing the origin of sexual dimorphism in this communication system.     

Epicuticular compounds and sexual recognition in the ball-roller scarab, Canthon cyanellus cyanellus

We analyzed the role of cuticular compounds in sexual recognition of the carrion ball‐roller scarab, Canthon cyanellus cyanellus (LeConte) (Coleoptera: Scarabaeidae). Males rolled food balls with freeze‐killed females on them as though the females were alive. Acceptance of female cadavers was low when they were washed with solvents and increased significantly when the solvent extract was reapplied to the cadavers. Acceptance of female cadavers by males was significantly higher than that of male cadavers by females. Male cadavers washed with solvent and impregnated with female cuticular compounds were accepted by males as though they were frozen or living females. Similarly, extracted female cadavers impregnated with male cuticular compounds were accepted by females. These results suggest that C. c. cyanellus males recognize females by their cuticular compounds, whereas females use a combination of cuticular compounds and other unknown cues to recognize males.     

Faunistic data and biogeography of terrestrial isopods from Tunisian wetlands

In Tunisia, while wetlands are considered as remarkable habitats for their faunal and floral diversity, few studies on the biogeography and the diversity of terrestrial isopods were performed. To fill this gap, we carried out a field study in the supralittoral zone of 146 Tunisian wetlands belonging to eight types (lagoon, river, lake, sebkha, dam, hill reservoir, beach and chott) and to five bioclimatic zones (humid, subhumid, semi‐arid, arid and Saharan). Field work was carried out in spring 2010. Terrestrial isopods were collected in the morning by hand search each time with the same sampling effort. During the study, 22 species of terrestrial isopods belonging to nine families were collected. Porcellio variabilis, Porcellio laevis, Chaetophiloscia elongata and Armadillidium pelagicum are the most common species identified. From the northern to the southern regions, a gradual decrease in species richness has been shown. The identified species belonged to different biogeographical categories: Mediterranean, Mediterranean‐Atlantic, North Africa with circum‐Sicilian islands, semi‐arid and Saharan species. Some of these species are endemic to Tunisia (Porcellio dominici) or to North Africa (Armadillidium sulcatum, Armadillidium tunisiense, Porcellio marginenotatus and P. variabilis). Terrestrial isopod species differ according to wetland type and bioclimatic zone.