Diverse filters to sense: great variability of antennal morphology and sensillar equipment in gall-wasps (Hymenoptera: Cynipidae).

Comparative studies on antennal sensillar equipment in insects are largely lacking, despite their potential to provide insights into both ecological and phylogenetic relationships. Here we present the first comparative study on antennal morphology and sensillar equipment in female Cynipoidea (Hymenoptera), a large and diverse group of wasps, with special reference to the so-called gall-wasps (Cynipidae). A SEM analysis was conducted on 51 species from all extant cynipoid families and all cynipid tribes, and spanning all known life-histories in the superfamily (gall-inducers, gall-inquilines, and non-gall associated parasitoids). The generally filiform, rarely clavate, antennal flagellum of Cynipoidea harbours overall 12 types of sensilla: s. placoidea (SP), two types of s. coeloconica (SCo-A, SCo-B), s. campaniformia (SCa), s. basiconica (SB), five types of s. trichoidea (ST-A, B, C, D, E), large disc sensilla (LDS) and large volcano sensilla (LVS). We found a great variability in sensillar equipment both among and within lineages. However, few traits seem to be unique to specific cynipid tribes. Paraulacini are, for example, distinctive in having apical LVS; Pediaspidini are unique in having ≥3 rows of SP, each including 6-8 sensilla per flagellomere, and up to 7 SCo-A in a single flagellomere; Eschatocerini have by far the largest SCo-A. Overall, our data preliminarily suggest a tendency to decreased numbers of SP rows per flagellomere and increased relative size of SCo-A during cynipoid evolution. Furthermore, SCo-A size seems to be higher in species inducing galls in trees than in those inducing galls in herbs. On the other hand, ST seem to be more abundant on the antennae of herb-gallers than wood-gallers. The antennal morphology and sensillar equipment in Cynipoidea are the complex results of different interacting pressures that need further investigations to be clarified.

Breaking up the wall: metal-enrichment in Ovipositors, but not in mandibles, co-varies with substrate hardness in gall-wasps and their associates.

The cuticle of certain insect body parts can be hardened by the addition of metals, and because niche separation may require morphological adaptations, inclusion of such metals may be linked to life history traits. Here, we analysed the distribution and enrichment of metals in the mandibles and ovipositors of a large family of gall-inducing wasps (Cynipidae, or Gall-Wasps) (plus one gall-inducing Chalcidoidea), and their associated wasps (gall-parasitoids and gall-inquilines) (Cynipidae, Chalcidoidea and Ichneumonoidea). Both plant types/organs where galls are induced, as well as galls themselves, vary considerably in hardness, thus making this group of wasps an ideal model to test if substrate hardness can predict metal enrichment. Non-galler, parasitic Cynipoidea attacking unconcealed hosts were used as ecological "outgroup". With varying occurrence and concentration, Zn, Mn and Cu were detected in mandibles and ovipositors of the studied species. Zn tends be exclusively concentrated at the distal parts of the organs, while Mn and Cu showed a linear increase from the proximal to the distal parts of the organs. In general, we found that most of species having metal-enriched ovipositors (independently of metal type and concentration) were gall-invaders. Among gall-inducers, metals in the ovipositors were more likely to be found in species inducing galls in woody plants. Overall, a clear positive effect of substrate hardness on metal concentration was detected for all the three metals. Phylogenetic relationships among species, as suggested by the most recent estimates, seemed to have a weak role in explaining metal variation. On the other hand, no relationships were found between substrate hardness or gall-association type and concentration of metals in mandibles. We suggest that ecological pressures related to oviposition were sufficiently strong to drive changes in ovipositor elemental structure in these gall-associated Hymenoptera.

Antennal sensillar equipment in closely related predatory wasp species (Hymenoptera: Philanthinae) hunting for different prey types.

Despite its potential value in phylogenetic and ecological studies, the morphology of antennal sensilla has rarely been compared quantitatively within the Apoidea. Here, through a scanning electron microscopy analysis, we provide an inventory of different types of antennal sensilla and compare their morphology across 10 species of predatory wasps (Crabronidae: Philanthinae) including species that hunt exclusively either on beetles or on bees to feed their larvae. A sensilla-free area was found on the apical flagellomer of all but two species, and its shape and size appear to be useful for separating Philanthini from Cercerini within the subfamily. A total of eight types of sensilla (sensilla placoidea, sensilla basiconica, two types of pit organs, sensilla coelocapitula and three types of sensilla trichoidea) were found in all species, and an additional rarer type (grooved peg sensilla) was found only in three bee-hunting species and for first time in the genus Cerceris. Certain morphological features confirmed the separation of the apoid wasps from the rest of the Apoidea (i.e., bees). A cluster analysis based on the sizes of the different types of sensilla suggested that, overall, sensilla morphology is not a useful taxonomic tool, and thus, other factors likely determine interspecific variability. One candidate factor is the prey type, given some differences in the presence, density, size and distribution of certain types of olfactory sensilla between beetle-hunters and bee-hunters. This hypothesis needs to be further tested quantitatively using a larger species set, more individuals per species, additional sensilla features, and a correction for phylogeny.

Ossification vesicles with calcium phosphate in the eyes of the insect Copium teucrii (Hemiptera: Tingidae).

Arthropod eyes are built of repeating units named ommatidia. Each single ommatidium unit contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The insect Copium eye ommatidia include additional calcium-phosphate deposits, not described in insects to date, which can be examined today using a combined set of modern microscopy and spectroscopy techniques. Teucrium gnaphalodes L'Her plants, growing in central Spain, develop galls induced by Copium insects. A survey of C. teucrii adult specimens resulted in surprising environmental scanning electron microscopy (ESEM) images, showing that their bright red eyes contain a calcium-phosphate mineralization. A complete survey of Copium eye specimens was performed by ESEM using energy-dispersive spectroscopy, backscattered electron detector and cathodoluminescence (CL) probes, field emission scanning electron microscopy, micro-Raman spectroscopy, and confocal laser scanning microscopy in order to learn ommatidia features, such as chemical composition, molecular structure, cell membrane, and internal ommatidium eye fluids and calcium-phosphate distribution deposits. The CL panchromatic images distinguish between the calcium-phosphate ommatidium and calcium-phosphate setae, which are more apatite rich. They show Raman bands attributable to bone tissue apatite biomaterials, such as bone, collagen, lipids, and blood, i.e., peptides, amide-S, amide-II, amide-III, and cytochrome P-450 scc. The chemical composition of both galls and leaves of T. gnaphalodes was determined by gas chromatography-mass spectrometry (GC-MS) of their extracts. The spectrometric and microscopic images reveal that the calcium-phosphate mineralization is formed and constrained to Copium ommatidia, which are both matrix vesicles generating mixtures of apatite collagen and operational compound eyes of the insect.

The importance of beta diversity in local gall-inducing arthropod distribution.

Many studies over the past twenty years have documented the richness of arthropod galling species around the world, and some have proposed hypotheses to explain local and global patterns of galling species richness. However, few studies have been directed toward understanding how the gall-inducing species are locally distributed. The aim of this study was to determine the distribution of gall-inducing arthropods species at Coiba National Park, a tropical habitat on the Pacific coast of Panama. Our results suggest that more gall-inducing species had an aggregated distribution, and gall-inducing arthropod diversity shows a strong beta diversity component. Geographic distance was not correlated with similarity in gall-inducing species composition between the studied sites. This fact has important implications when trying to estimate gall-inducing arthropod richness and general patterns, and could cause contradictory results for hypotheses that attempt to explain the local and global patterns of galling species richness.

The importance of beta diversity in local gall-inducing arthropod distribution

Many studies over the past twenty years have documented the richness of arthropod galling species around the world, and some have proposed hypotheses to explain local and global patterns of galling species richness. However, few studies have been directed toward understanding how the gall-inducing species are locally distributed. The aim of this study was to determine the distribution of gall-inducing arthropods species at Coiba National Park, a tropical habitat on the Pacific coast of Panama. Our results suggest that more gall-inducing species had an aggregated distribution, and gall-inducing arthropod diversity shows a strong beta diversity component. Geographic distance was not correlated with similarity in gall-inducing species composition between the studied sites. This fact has important implications when trying to estimate gall-inducing arthropod richness and general patterns, and could cause contradictory results for hypotheses that attempt to explain the local and global patterns of galling species richness.