Siegfried Kehl

Predation by pioneer water beetles (Coleoptera, Dytiscidae) from sandpit ponds, based on crop-content analysis and laboratory experiments

Crop-content analyses of 898 pioneer beetles, belonging to five dytiscid species of sandpit ponds, (Coleoptera: Dytiscidae) were performed to identify possible nutritionally-based niches and competitive effects. Agabus nebulosus, Nebrioporus canaliculatus, Scarodytes halensis, Hygrotus confluens and Hydroglyphus geminus mainly feed on chironomid larvae and planktonic crustacea, especially Copepoda. However, a large amount of filamentous algae was found in the crops of A. nebulosus and H. confluens fed on Rotatoria, especially Keratella, while the small hydroporine H. geminus fed on the small nauplius larvae of Copepoda. Experiments were used to test the relative feeding preference of S. halensis for Copepoda and Cladocera and for dead or living plankton. Dead plankton was preferred, whereas the size of the plankton was important in the selection of Copepoda or Cladocera. The crop clearance times for different food items were also ascertained: no differences in digestion times of chironomid larvae, Chaoborus larvae and plankton were found.

Diversity of Symbiotic Organs and Bacterial Endosymbionts of Lygaeoid Bugs of the Families Blissidae and Lygaeidae (Hemiptera: Heteroptera: Lygaeoidea)

ABSTRACT Here we present comparative data on the localization and identity of intracellular symbionts among the superfamily Lygaeoidea (Insecta: Hemiptera: Heteroptera: Pentatomomorpha). Five different lygaeoid species from the families Blissidae and Lygaeidae (sensu stricto; including the subfamilies Lygaeinae and Orsillinae) were analyzed. Fluorescence in situ hybridization (FISH) revealed that all the bugs studied possess paired bacteriomes that are differently shaped in the abdomen and harbor specific endosymbionts therein. The endosymbionts were also detected in female gonads and at the anterior poles of developing eggs, indicating vertical transmission of the endosymbionts via ovarial passage, in contrast to the posthatch symbiont transmission commonly found among pentatomoid bugs (Pentatomomorpha: Pentatomoidea). Phylogenetic analysis based on 16S rRNA and groEL genes showed that the endosymbionts of Ischnodemus sabuleti, Arocatus longiceps, Belonochilus numenius, Orsillus depressus, and Ortholomus punctipennis constitute at least four distinct clades in the Gammaproteobacteria. The endosymbiont phylogeny did not agree with the host phylogeny based on the mitochondrial cytochrome oxidase I (COI) gene, but there was a local cospeciating pattern within the subfamily Orsillinae. Meanwhile, the endosymbiont of Belonochilus numenius (Lygaeidae: Orsillinae), although harbored in paired bacteriomes as in other lygaeoid bugs of the related genera Nysius, Ortholomus, and Orsillus, was phylogenetically close to “Candidatus Rohrkolberia cinguli,” the endosymbiont of Chilacis typhae (Lygaeoidea: Artheneidae), suggesting an endosymbiont replacement in this lineage. The diverse endosymbionts and the differently shaped bacteriomes may reflect independent evolutionary origins of the endosymbiotic systems among lygaeoid bugs.

Characterization of an obligate intracellular bacterium in the midgut epithelium of the bulrush bug Chilacis typhae (Heteroptera, Lygaeidae, Artheneinae).

ABSTRACT Many members of the suborder Heteroptera have symbiotic bacteria, which are usually found extracellularly in specific sacs or tubular outgrowths of the midgut or intracellularly in mycetomes. In this study, we describe the second molecular characterization of a symbiotic bacterium in a monophagous, seed-sucking stink bug of the family Lygaeidae (sensu stricto). Chilacis typhae possesses at the end of the first section of the midgut a structure which is composed of circularly arranged, strongly enlarged midgut epithelial cells. It is filled with an intracellular endosymbiont. This “mycetocytic belt” might represent an evolutionarily intermediate stage of the usual symbiotic structures found in stink bugs. Phylogenetic analysis based on the 16S rRNA and the groEL genes showed that the bacterium belongs to the Gammaproteobacteria, and it revealed a phylogenetic relationship with a secondary bacterial endosymbiont of Cimex lectularius and free-living plant pathogens such as Pectobacterium and Dickeya. The distribution and ultrastructure of the rod-shaped Chilacis endosymbiont were studied in adults and nymph stages using fluorescence in situ hybridization (FISH) and electron microscopy. The detection of symbionts at the anterior poles of developing eggs indicates that endosymbionts are transmitted vertically. A new genus and species name, “Candidatus Rohrkolberia cinguli,” is proposed for this newly characterized clade of symbiotic bacteria.

Surviving submerged - Setal tracheal gills for gas exchange in adult rheophilic diving beetles

The gas exchange in adult diving beetles (Coleoptera: Dytiscidae) relies on a subelytral air store, which has to be renewed in regular intervals at the water surface. The dive duration varies from a few minutes to 24 h depending on the species, activity, and temperature. However, some species remain submerged for several weeks. Stygobiont species do not ascend to the surface and gas exchange of these species remains unclear, but it is assumed that they require air filled voids for respiration or they use cutaneous respiration. In this study, we investigate the gas exchange in the running water diving beetle Deronectes aubei, which survive submerged for over 6 weeks. The diffusion distance through the cuticle is too great for cutaneous respiration. Therefore, the dissolved oxygen uptake of submerged beetles was determined and an oxygen uptake via the rich tracheated elytra was observed. Fine structure analyses (SEM and TEM) of the beetles showed tracheated setae mainly on the elytral surface, which acts as tracheal gills. Prevention of the air bubble formation at the tip of the abdomen, which normally act as physical gill in Dytiscidae, resulted in no effect in oxygen uptake in D. aubei, but this was the sole way for a submerged Hydroporus palustris to get oxygen. The setal gas exchange technique explains the restriction of D. aubei to rivers and brooks with high oxygen concentration and it may also be used by subterran living diving beetles, which lack access to atmospheric oxygen. The existence of setal tracheal gills in species in running water which are often found in the hyporheic zone and in stygobiont species supports the known evolution of stygobiont Dytiscidae from species of the hyporheic zone. For species in running water, setal tracheal gills could be seen as an adaptation to avoid drifting downstream by the current. J. Morphol., 2009.

First Sex Pheromone of the Order Strepsiptera: (3R,5R,9R)-3,5,9-Trimethyldodecanal in Stylops melittae Kirby, 1802

The twisted-wing parasites (Strepsiptera) are an unusual and small order of insects with about 600 known species. As obligate endoparasitoids, they develop and spend most of their lives living in other insects. Adults show an extreme sexual dimorphism: The free-living males have large eyes, branched antennae, reduced forewings, and well developed hind wings, while the neotenic females of most species lack all external characters that normally define an insect, remain endoparasitic, and only extrude the cephalothorax from the host. Due to the males’ short life span of only a few hours, there must be an efficient means of mate finding. This is believed to be mediated by chemical cues released by virgin females. Here, we report the first identification and synthesis of a female-produced strepsipteran sex pheromone, (3R,5R,9R)-3,5,9-trimethyldodecanal, from Stylops melittae, a species parasitizing andrenid bees. We found this highly EAD-active compound to be present in cephalothoraxes of and released from unmated females, and synthetic samples proved to be extremely attractive when offered in the field during the swarming period of the males. The structural features of this new natural compound may further support the re-establishment of the Strepsiptera as the closest living relatives of the Coleoptera.

The Phlesirtes complex (Orthoptera, Tettigoniidae, Conocephalinae, Conocephalini) reviewed: integrating morphological, molecular, chromosomal and bioacoustic data.

The tettigoniid genus Phlesirtes Bolivar and its allies are reviewed. Morphological, ecological and molecular data prompt the erection of the new genus Chortoscirtesgen.n. with type species Xiphidion meruense Sjöstedt. The genera Phlesirtes, Chortoscirtes, Karniella and Naskreckiella are characterized by morphological characters supported by molecular, acoustic, ecological and chromosomal data. Four species, Chortoscirtes pseudomeruensissp.n., C. masaicussp.n., C. puguensissp.n. and C. serengetisp.n., are described from localities in northern and coastal Tanzania and one Karniella, K. crassicercasp.n., is described from Uganda. The following comb n. are proposed: Phlesirtes kibonotensis (Sjöstedt) and Phlesirtes kilimandjaricus (Sjöstedt). Subtribal status is proposed for the four investigated African genera. A key to the Chortoscirtes species is provided.

A new genus of African Karniellina (Orthoptera, Tettigoniidae, Conocephalinae, Conocephalini): integrating morphological, molecular and bioacoustical data

Melanoscirtes gen.n. is established within Karniellina. The members of this subtribe are small conocephaline bush crickets, confined to Africa. Melanoscirtes is erected on Phlesirtes kibonotensis, a species restricted to forest clearings and forest edge in the submontane and montane zones of Mt. Kilimanjaro. A subspecies, M. kibonotensis uguenoensis, is described from the North Pare mountains, a mountain range of the Eastern Arc adjacent to Mt. Kilimanjaro. Further species of Melanoscirtes occur on other mountain ranges of the northern branch of the Eastern Arc mountains of northern Tanzania and southern Kenya. The South Pare mountains harbour M. shengenae; the West Usambaras, M. usambarensis, and the Taita Hills, M. taitensis. All species and subspecies of Melanoscirtes exhibit a similar morphology and occupy analogous habitats on the respective mountains. The song patterns for all species found within this genus are very similar, and this, together with evidence from molecular data, suggests that allopatric speciation is the reason for the biogeographic pattern found in this genus. A key for the subspecies and species of Melanoscirtes is provided.

Climatic fluctuations and orogenesis as motors for speciation in East Africa: case study on Parepistaurus Karsch, 1896 (Orthoptera)

Mechanisms of speciation of flightless grasshoppers in mountainous and coastal East Africa are inferred considering (i) phylogenies estimated with a combination of molecular markers (16S rRNA locus, COI and H3), (ii) ecological data and (iii) the geographic distribution of Parepistaurus species. The study suggests that coastal taxa of Parepistaurus belong to ancestral lineages from which evolved the high diversity of species found in the Eastern Arc Mountains of Tanzania and Kenya, which are geologically ancient mountain formations. Network analyses and a molecular clock approach, calibrated with the geological age of the volcanoes, suggested that speciation was boosted by climatic fluctuations affecting large areas of East Africa. With the aridification beginning 2.8 Ma, forest taxa were isolated due to forest fragmentation and populations were separated by extended grasslands, which are avoided by Parepistaurus species. However, a humid period between 2.7 and 2.5 Ma triggered a spread of coastal taxa along the Eastern Arc Mountains. Forests expanded again and riparian vegetation along rivers draining into the Indian Ocean probably served as corridors for the dispersal of coastal taxa to the hinterland. The inland volcanoes such as Mount Kilimanjaro are therefore good time markers because their geological age is known, limiting the available time for speciation processes of mountainous Parepistaurus in the area to a maximum of about 1–2 Ma. A third humid but cold period between 1.1 and 0.9 Ma probably further boosted the spread of several flightless and montane‐adapted Orthoptera taxa.

Characterization and localization of Rickettsia sp. in water beetles of genus Deronectes (Coleoptera: Dytiscidae)

In the present study, Rickettsia sp. was detected in four water beetles of the genus Deronectes (Dytiscidae) for the first time. Rickettsiae were found in 100% of examined specimens of Deronectes platynotus (45/45), 39.4% of Deronectes aubei (28/71), 40% of Deronectes delarouzei (2/5) and 33.3% of Deronectes semirufus (1/3). Analysis of 16S rRNA gene sequences revealed a phylogenetic relationship with rickettsial isolates of Limonia chorea (Diptera), tentatively classified as members of the basal ancestral group. Phylogenetic analysis of the gltA (citrate synthase) gene sequences showed that Deronectes symbionts were closest to bacterial symbionts from spiders. Ultrastructural examinations revealed typical morphological features and intracellular arrangements of rickettsiae. The distribution, transmission and localization of Rickettsia sp. in D. platynotus were studied using a diagnostic PCR assay and FISH. Eggs from infected females of D. platynotus were all Rickettsia-positive, indicative of a vertical transmission.

Characterization and localization of Rickettsia sp. in water beetles of genus Deronectes

In the present study, Rickettsia sp. was detected in four water beetles of the genus Deronectes (Dytiscidae) for the first time. Rickettsiae were found in 100% of examined specimens of Deronectes platynotus (45/45), 39.4% of Deronectes aubei (28/71), 40% of Deronectes delarouzei (2/5) and 33.3% of Deronectes semirufus (1/3). Analysis of 16S rRNA gene sequences revealed a phylogenetic relationship with rickettsial isolates of Limonia chorea (Diptera), tentatively classified as members of the basal ancestral group. Phylogenetic analysis of the gltA (citrate synthase) gene sequences showed that Deronectes symbionts were closest to bacterial symbionts from spiders. Ultrastructural examinations revealed typical morphological features and intracellular arrangements of rickettsiae. The distribution, transmission and localization of Rickettsia sp. in D. platynotus were studied using a diagnostic PCR assay and FISH. Eggs from infected females of D. platynotus were all Rickettsia-positive, indicative of a vertical transmission.