Florian M. Steiner

Integrative Taxonomy: A Multisource Approach to Exploring Biodiversity

Good alpha taxonomy is central to biology. On the basis of a survey of arthropod studies that used multiple disciplines for species delimitation, we evaluated the performance of single disciplines. All included disciplines had a considerable failure rate. Rigor in species delimitation can thus be increased when several disciplines chosen for complementarity are used. We present a flexible procedure and stopping rule for integrative taxonomy that uses the information from different disciplines separately. Disagreement among disciplines over the number and demarcation of species is resolved by elucidating and invoking evolutionary explanations for disagreement. With the identification of further promising study organisms and of new questions for in-depth analysis, evolutionary biology should profit from integrative taxonomy. An important rationale is clarity in researcher bias in the decision-making process. The success of integrative taxonomy will further increase through methodological progress, taxonomic training of evolutionary biologists, and balanced resource allocation.

The Evolution of Invasiveness in Garden Ants

It is unclear why some species become successful invaders whilst others fail, and whether invasive success depends on pre-adaptations already present in the native range or on characters evolving de-novo after introduction. Ants are among the worst invasive pests, with Lasius neglectus and its rapid spread through Europe and Asia as the most recent example of a pest ant that may become a global problem. Here, we present the first integrated study on behavior, morphology, population genetics, chemical recognition and parasite load of L. neglectus and its non-invasive sister species L. turcicus. We find that L. neglectus expresses the same supercolonial syndrome as other invasive ants, a social system that is characterized by mating without dispersal and large networks of cooperating nests rather than smaller mutually hostile colonies. We conclude that the invasive success of L. neglectus relies on a combination of parasite-release following introduction and pre-adaptations in mating system, body-size, queen number and recognition efficiency that evolved long before introduction. Our results challenge the notion that supercolonial organization is an inevitable consequence of low genetic variation for chemical recognition cues in small invasive founder populations. We infer that low variation and limited volatility in cuticular hydrocarbon profiles already existed in the native range in combination with low dispersal and a highly viscous population structure. Human transport to relatively disturbed urban areas thus became the decisive factor to induce parasite release, a well established general promoter of invasiveness in non-social animals and plants, but understudied in invasive social insects.

Permanent Genetic Resources added to Molecular Ecology Resources Database 1 April 2010-31 May 2010

This article documents the addition of 396 microsatellite marker loci to the Molecular Ecology Resources Database. Loci were developed for the following species: Anthocidaris crassispina, Aphis glycines, Argyrosomus regius, Astrocaryum sciophilum, Dasypus novemcinctus, Delomys sublineatus, Dermatemys mawii, Fundulus heteroclitus, Homalaspis plana, Jumellea rossii, Khaya senegalensis, Mugil cephalus, Neoceratitis cyanescens, Phalacrocorax aristotelis, Phytophthora infestans, Piper cordulatum, Pterocarpus indicus, Rana dalmatina, Rosa pulverulenta, Saxifraga oppositifolia, Scomber colias, Semecarpus kathalekanensis, Stichopus monotuberculatus, Striga hermonthica, Tarentola boettgeri and Thermophis baileyi. These loci were cross‐tested on the following species: Aphis gossypii, Sooretamys angouya, Euryoryzomys russatus, Fundulus notatus, Fundulus olivaceus, Fundulus catenatus, Fundulus majalis, Jumellea fragrans, Jumellea triquetra Jumellea recta, Jumellea stenophylla, Liza richardsonii, Piper marginatum, Piper aequale, Piper darienensis, Piper dilatatum, Rana temporaria, Rana iberica, Rana pyrenaica, Semecarpus anacardium, Semecarpus auriculata, Semecarpus travancorica, Spondias acuminata, Holigarna grahamii, Holigarna beddomii, Mangifera indica, Anacardium occidentale, Tarentola delalandii, Tarentola caboverdianus and Thermophis zhaoermii.

No sympatric speciation here: multiple data sources show that the ant Myrmica microrubra is not a separate species but an alternate reproductive morph of Myrmica rubra

No aspect of speciation is as controversial as the view that new species can evolve sympatrically, among populations in close physical contact. Social parasitism has been suggested to yield necessary disruptive selection for sympatric speciation. Recently, mitochondrial DNA phylogeography has shown that the ant Myrmica microrubra is closely related to its host, Myrmica rubra, leading to the suggestion that sympatric speciation has occurred. We investigated the relationships between the two ant forms using mitochondrial and nuclear DNA sequences, microsatellite genotyping and morphometrics. Molecular phylogenetic and population structure analyses showed that M. microrubra does not evolve separately to its host but rather shares a gene pool with it. Probability analysis showed that mitochondrial DNA data previously adduced in favour of sympatric speciation do not in fact do so. Morphometrically, M. microrubra is most readily interpreted as a miniature queen form of M. rubra, not a separate species. Myrmica microrubra is not an example of speciation. The large (typical M. rubra) and small (M. microrubra) queen forms are alternative reproductive strategies of the same species. Myrmica microrubra Seifert 1993 is consequently synonymized here with M. rubra Linnaeus, 1758.

Host specificity revisited: New data on Myrmica host ants of the lycaenid butterfly Maculinea rebeli

Larvae of Maculinea rebeli, one of the most endangered European butterflies, are obligatory social parasites of Myrmica ants. At present, this relationship is thought to be highly specific, with Myrmica schencki being regarded as the primary host. Here we present data on six populations from Poland and Austria, including the first record of Myrmica specioides as a host, together with published data from other central European countries, which severely questions the inference that M. schencki is the exclusive host of M. rebeli. Our results indicate that Myrmica sabuleti is the most frequently used host ant in central Europe, whereas M. scabrinodis, M. sulcinodis, M. specioides and M. schencki are used as secondary hosts. Possible explanations for this highly variable host use include (1) regional differences in semiochemicals, behaviour or social structure of the potential Myrmica host species and (2) the existence of different ecological subspecies or cryptic species of M. rebeli. Finally, we emphasize the importance of identifying local host ant species prior to further conservation strategies in order to avoid failure of management programs or even damage to populations on the edge of extinction.

Stable N-isotope signatures of central European ants – assessing positions in a trophic gradient

Abstract.Studies employing stable isotope technology have greatly contributed to understanding trophic relationships of tropical ants, but temperate-zone ants remain under-explored. We studied δ15N values of 43 ant species from three subfamilies sampled across central Europe. After statistically accounting for the effects of elevation and geographical location of habitats, which alter the isotopic composition of nitrogen in ecosystems, significant patterns in N-isotope signatures were detected. These signatures hint at differences across ants in the contribution of plant-derived nitrogen obtained via trophobiosis and nectarivory relative to nitrogen obtained via predation and scavenging. In general, Myrmicinae had higher δ15N values than Formicinae, in line with a greater relative importance of trophobiosis in the latter. The genus Myrmica scored especially high, indicating predominantly predacious nitrogen sources. Remarkably, also the granivore Messor cf. structor had high δ15N values. This suggests that, despite the major portion of food uptake being made up by plant seeds, this ant could derive substantial fractions of its nitrogen budget from feeding on arthropod corpses or vertebrate faeces. Moreover, this highlights that deductions from observed quantities of ingested food on its relative contribution to ants’ matter balance should be accompanied by isotope analyses. At the other end of the spectrum, Camponotus and Plagiolepis had low δ15N values. In line with multiple field observations, this suggests a contribution of trophobiosis not only to their energy, but also to their nitrogen budget. Formica and Lasius had intermediate 15N values, which is in agreement with the current view that these ants have mixed diets with a balance between trophobiosis and predation. A possible influence of endosymbiotic bacteria on the isotope signatures of several genera is discussed. This study provides a first application of stable isotope technology to estimate the role of plant-derived nutrients to the nitrogen budget of a larger range of central European ants. Furthermore, it shows that N-isotope analysis is applicable across extended ecological and geographical gradients. Future studies along this line are promising to complement our current understanding of the nutritional ecology of temperate-zone ants.

Predator complex of the horse chestnut leafminer Cameraria ohridella: identification and impact assessment

Abstract:  The control of Cameraria ohridella Deschka and Dimic (Lepidoptera, Gracillariidae) by natural enemies in Europe is poor. Thus, in the past 15 years epidemic population densities of the moth regularly caused a continuous pre‐mature defoliation of horse chestnut trees, Aesculus hippocastanum L.. Whereas several studies investigated the parasitoid complex of the leafminer and revealed its inefficiency, only little is known about the predators of C. ohridella. The aim of this study was to identify the predator complex in European horse chestnut stands by visual observations and exposure experiments and to assess the impact of different predator guilds on C. ohridella populations by an exclusion experiment. Of all potential predators observed, only blue tits, Parus caeruleus L., great tits, Parus major L., marsh tits, Parus palustris L. and the southern oak bushcricket Meconema meridionale (Costa) were found to prey on the pre‐imaginal leafminer stages. Bushcrickets have to our knowledge never before been recorded preying on leafminers; moreover M. meridionale showed a measurable negative impact on the C. ohridella population. However, birds seem to be of greater importance as predators of C. ohridella than arthropods. Predation rates of birds ranged from 2 to 4% of leafminer populations, which is of the same order of magnitude as published parasitism rates. We conclude that the augmentation of tit densities, for example via nesting boxes in chestnut trees, should be considered as a component of an integrated control strategy against the horse chestnut leafminer.

Assessing ant assemblages: pitfall trapping versus nest counting (Hymenoptera, Formicidae)

Abstract.Pitfall trapping and nest counting are the most common census methods for ant assemblages. We examined the concordance between pitfall catches and nest counts on dry grassland. Spearman rank correlations and non-metric multidimensional scaling of the Bray Curtis similarity index revealed moderate concordance between the data collated by the two methods, but overall method-related differences were considerable. The dissimilarity was influenced by the type of land management, but not by trapping period or plot shape. Trapping success depended on nest density, ground vegetation cover and species-specific traits (inhabited stratum, colony size, foraging distance). Even when these factors were taken into account, the convertibility of pitfall trap and nest density values was unsatisfactory: the census method proved to be crucial in designing ant-ecological studies and interpreting literature data.

Unraveling the chromophoric disorder of poly(3-hexylthiophene)

Significance Ideal photovoltaic cells would be black, absorbing all of the Sun’s radiation, whereas Nature’s machinery for solar energy harvesting—photosynthesis—looks green. Organic semiconductor devices, based on molecular building blocks, lie conceptionally between the extremes of inorganic and photosynthetic light harvesting. How can organic solar cells appear almost black if they are based on molecular units? Using single-molecule spectroscopy, we identify the fundamental electronic building blocks of organic solar cells and reveal that discrete molecule-like transitions scatter over the entire visible spectrum. The fundamental molecular unit is narrowband, but disorder induces a continuum reminiscent of that characterizing highly ordered inorganic crystals. The spectral breadth of conjugated polymers gives these materials a clear advantage over other molecular compounds for organic photovoltaic applications and is a key factor in recent efficiencies topping 10%. However, why do excitonic transitions, which are inherently narrow, lead to absorption over such a broad range of wavelengths in the first place? Using single-molecule spectroscopy, we address this fundamental question in a model material, poly(3-hexylthiophene). Narrow zero-phonon lines from single chromophores are found to scatter over 200 nm, an unprecedented inhomogeneous broadening that maps the ensemble. The giant red shift between solution and bulk films arises from energy transfer to the lowest-energy chromophores in collapsed polymer chains that adopt a highly ordered morphology. We propose that the extreme energetic disorder of chromophores is structural in origin. This structural disorder on the single-chromophore level may actually enable the high degree of polymer chain ordering found in bulk films: both structural order and disorder are crucial to materials physics in devices.

Singlet-triplet annihilation limits exciton yield in poly(3-hexylthiophene).

Control of chain length and morphology in combination with single-molecule spectroscopy techniques provides a comprehensive photophysical picture of excited-state losses in the prototypical conjugated polymer poly(3-hexylthiophene) (P3HT). Our examination reveals a universal self-quenching mechanism, based on singlet-triplet exciton annihilation, which accounts for the dramatic loss in fluorescence quantum yield of a single P3HT chain between its solution (unfolded) and bulklike (folded) state. Triplet excitons fundamentally limit the fluorescence of organic photovoltaic materials, which impacts the conversion of singlet excitons to separated charge carriers, decreasing the efficiency of energy harvested at high excitation densities. Interexcitonic interactions are so effective that a single P3HT chain of order 100  kDa weight behaves like a 2-level system, exhibiting perfect photon antibunching.