Chromosome differentiation and the radiation of the butterfly subgenus Agrodiaetus (<i>Lepidoptera</i>: Lycaenidae: <i>Polyommatus</i>) : a molecular phylogenetic approach

The subgenus Agrodiaetus which is distributed extensively in the Palaearctic region and especially in Southwest Asia is a species-rich group of blues which are often very similar morphologically but show a large variation in their chromosome numbers (n=10-125). The karyology of 64 taxa was successfully studied and the previously unknown chromosome numbers of 17 taxa were revealed. By means of molecular techniques it was possible to clarify the phylogeny of this group for the first time. A. damon, a species which is extremely widespread in the Palearctic region and has a constant chromosome number of n=45, is the sister-species of all other species of Agrodiaetus that have been tested. Presumably the colonization of Europe took place before the main radiations in the Anatolian-Caucasian-Iranian region. The greatest differences in the number of chromosomes were found in allopatrically distributed sister-species. It was surprising that morphologically extremely similar sympatric species with different numbers of chromosomes were not closely related, but often belonged to totally different clades, whereas some closely related sympatric species differed greatly in wing colours. This leads to the conclusion that changes in the number of chromosomes do not lead to sympatric speciation, but instead appear as a by-product of allopatric speciation and that such young species can only occur in sympatry after a sufficient differentiation in their phenotype to exclude erroneous matings. The comparison of mitochondrial with nuclear DNA sequences has also shown that hybridizations in Agrodiaetus are rare events. Supposedly they do not occur more frequently in Agrodiaetus than in other Lycaenidae. Only one of the specimen tested, which already drew our attention by its intermediate wing coloration, after molecular testing proved to be a hybrid of two nearly related but quite differently coloured species.Die in der Paläarktis und besonders in Vorderasien weit verbreitete Untergattung Agrodiaetus ist eine sehr artenreiche Gruppe von Bläulingen, welche sich morphologisch oft sehr ähnlich sehen, sich aber durch eine extrem starke Variation ihrer Chromosomenzahlen (n=10-125) auszeichnen. 64 Taxa konnten karyologisch erfolgreich untersucht und die bisher unbekannte Chromosomenzahl von 17 Taxa aufgedeckt werden. Mit Hilfe molekularer Techniken ist es nun erstmals gelungen, die Phylogenie dieser Gruppe weitgehend aufzuklären. Demnach ist A. damon, eine Art mit extrem weiter Verbreitung in der Paläarktis und konstanter Chromosomenzahl (n=45), die Schwesterart aller übrigen untersuchten Agrodiaetus-Arten. Die Besiedlung Europas erfolgte vermutlich schon vor den Hauptradiationen im anatolischkaukasich-iranischen Raum. Die größten Unterschiede in den Chromosomenzahlen finden sich bei allopatrisch verbreiteten Schwesterarten. Überraschend war, dass morphologisch extrem ähnliche sympatrisch vorkommende Arten mit unterschiedlichen Chromosomenzahlen nicht näher miteinander verwandt sind, sondern oft sogar zu ganz verschiedenen Verwandtschaftsgruppen gehören, wohingegen sich einige sehr nahe verwandte sympatrisch vorkommende Arten stark in der Flügelfärbung unterscheiden. Daraus lässt sich der Schluss ziehen, dass Veränderungen der Chromosomenzahlen nicht zu sympatrischer Artbildung führen, sondern stattdessen ein Nebenprodukt allopatrischer Artbildung darstellen und dass solche jungen Arten erst dann wieder in Sympatrie auftreten können, wenn sie sich auch phänotypisch ausreichend differenziert haben, um Fehlpaarungen zu vermeiden. Der Vergleich von mitochondrialen mit Kern-DNA-Sequenzen hat auch gezeigt, dass Hybridisationen bei Agrodiaetus seltene Ereignisse darstellen. Sie treten vermutlich nicht viel häufiger auf als bei anderen Lepidopteren-Gattungen. Nur einer der untersuchten Falter, der bereits durch eine intermediäre Färbung aufgefallen war, erwies sich nach molekularer Untersuchung als Hybrid zwischen zwei nahe verwandten, aber recht unterschiedlich gefärbten Arten

Does the DNA barcoding gap exist? – a case study in blue butterflies (Lepidoptera: Lycaenidae)

BACKGROUND: DNA barcoding, i.e. the use of a 648 bp section of the mitochondrial gene cytochrome c oxidase I, has recently been promoted as useful for the rapid identification and discovery of species. Its success is dependent either on the strength of the claim that interspecific variation exceeds intraspecific variation by one order of magnitude, thus establishing a "barcoding gap", or on the reciprocal monophyly of species. RESULTS: We present an analysis of intra- and interspecific variation in the butterfly family Lycaenidae which includes a well-sampled clade (genus Agrodiaetus) with a peculiar characteristic: most of its members are karyologically differentiated from each other which facilitates the recognition of species as reproductively isolated units even in allopatric populations. The analysis shows that there is an 18% overlap in the range of intra- and interspecific COI sequence divergence due to low interspecific divergence between many closely related species. In a Neighbour-Joining tree profile approach which does not depend on a barcoding gap, but on comprehensive sampling of taxa and the reciprocal monophyly of species, at least 16% of specimens with conspecific sequences in the profile were misidentified. This is due to paraphyly or polyphyly of conspecific DNA sequences probably caused by incomplete lineage sorting. CONCLUSION: Our results indicate that the "barcoding gap" is an artifact of insufficient sampling across taxa. Although DNA barcodes can help to identify and distinguish species, we advocate using them in combination with other data, since otherwise there would be a high probability that sequences are misidentified. Although high differences in DNA sequences can help to identify cryptic species, a high percentage of well-differentiated species has similar or even identical COI sequences and would be overlooked in an isolated DNA barcoding approach

ITS2 secondary structure improves phylogeny estimation in a radiation of blue butterflies of the subgenus Agrodiaetus (Lepidoptera: Lycaenidae: Polyommatus )

<p>Abstract</p> <p>Background</p> <p>Current molecular phylogenetic studies of Lepidoptera and most other arthropods are predominantly based on mitochondrial genes and a limited number of nuclear genes. The nuclear genes, however, generally do not provide sufficient information for young radiations. <it>ITS2 </it>, which has proven to be an excellent nuclear marker for similarly aged radiations in other organisms like fungi and plants, is only rarely used for phylogeny estimation in arthropods, although universal primers exist. This is partly due to difficulties in the alignment of <it>ITS2 </it>sequences in more distant taxa. The present study uses <it>ITS2 </it>secondary structure information to elucidate the phylogeny of a species-rich young radiation of arthropods, the butterfly subgenus <it>Agrodiaetus</it>. One aim is to evaluate the efficiency of <it>ITS2 </it>to resolve the phylogeny of the subgenus in comparison with <it>COI </it>, the most important mitochondrial marker in arthropods. Furthermore, we assess the use of compensatory base changes in <it>ITS2 </it>for the delimitation of species and discuss the prospects of <it>ITS2 </it>as a nuclear marker for barcoding studies.</p> <p>Results</p> <p>In the butterfly family Lycaenidae, <it>ITS2 </it>secondary structure enabled us to successfully align sequences of different subtribes in Polyommatini and produce a Profile Neighbour Joining tree of this tribe, the resolution of which is comparable to phylogenetic trees obtained with <it>COI+COII </it>. The subgenus <it>Agrodiaetus </it>comprises 6 major clades which are in agreement with <it>COI </it>analyses. A dispersal-vicariance analysis (DIVA) traced the origin of most <it>Agrodiaetus </it>clades to separate biogeographical areas in the region encompassing Eastern Anatolia, Transcaucasia and Iran.</p> <p>Conclusions</p> <p>With the inclusion of secondary structure information, <it>ITS2 </it>appears to be a suitable nuclear marker to infer the phylogeny of young radiations, as well as more distantly related genera within a diverse arthropod family. Its phylogenetic signal is comparable to the mitochondrial marker <it>COI </it>. Compensatory base changes are very rare within Polyommatini and cannot be used for species delimitation. The implementation of secondary structure information into character-based phylogenetic methods is suggested to further improve the versatility of this marker in phylogenetic studies.</p

Monitoring Klimawandel und Biodiversität - Konzeption

Die im Heft 24 der Schriftenreihe präsentierten Grundlagen werden im vorliegenden Heft 25 zu einer Konzeption vervollständigt. Teilbereiche der folgenden bestehenden Monitoringprogramme sind für eine Einbeziehung in das konzipierte Monitoring Klimawandel und Biodiversität besonders geeignet: FFH-, SPA-, Tagfalter-, Brutvogel- und Wasserrahmenrichtlinien-Monitoring sowie Forstliches Umweltmonitoring. Es werden acht Module vorgestellt, welche bestehende Monitoringprogramme für ein umfassendes Klimawandel-Biodiversitätsmonitoring ergänzen bzw. bisher nicht untersuchte Aspekte abdecken können. Für die Auswertung der Daten wurden zwei komplexe Kernindikatoren entwickelt und anhand realer Datensets getestet, der Community Temperature Index (CTI) und der Areal Index (AI). Beide zeigen für die Artengruppen der Tagfalter und Libellen innerhalb Sachsens einen Anstieg, der unterstreicht, dass die Erhöhung der Jahresmitteltemperaturen in den letzten Jahrzehnten bereits zu Veränderungen in diesen Artengemeinschaften geführt hat

Life history and biogeography of the enigmatic mantid Nilomantis floweri (Mantodea, Nanomantidae)

The biology and distribution patterns of the Sahelian mantid species Nilomantis floweri are still insufficiently known. For the first time, records are confirmed of this species from Iran and the distribution map of its native range is updated. Records are compiled from the Sahel zone of North Africa, the Arabian Peninsula, and Iran. Detailed information on its biology, oothecal characteristics, male genitalia variation, and intraspecific molecular diversity in the mitochondrial gene cytochrome c oxidase are provided, and ecological niche modelling was used to gain insight into the overall species distribution and understand its climatic niche limits. Genetic analysis revealed only one haplotype shared between Iran and Oman. The Iranian populations likely represent two distinct clusters, both more related to the diverse Oman haplotypes than to each other. Based on new data, N. floweri appears to be mostly associated with coastal areas in southwestern Asia, with the vast majority of records found along the Red Sea, Persian Gulf, and Oman Gulf coasts. This distribution contrasts markedly with N. floweri records in the Sahel, where most collections have been reported in the transitional zone between the southern Sahara and arid thorn savannah, far off the coast. This study contributes to a comprehensive understanding of this still enigmatic mantid species

A new comprehensive trait database of European and Maghreb butterflies, Papilionoidea

Trait-based analyses explaining the different responses of species and communities to environmental changes are increasing in frequency. European butterflies are an indicator group that responds rapidly to environmental changes with extensive citizen science contributions to documenting changes of abundance and distribution. Species traits have been used to explain long- and short-term responses to climate, land-use and vegetation changes. Studies are often characterised by limited trait sets being used, with risks that the relative roles of different traits are not fully explored. Butterfly trait information is dispersed amongst various sources and descriptions sometimes differ between sources. We have therefore drawn together multiple information sets to provide a comprehensive trait database covering 542 taxa and 25 traits described by 217 variables and sub-states of the butterflies of Europe and Maghreb (northwest Africa) which should serve for improved trait-based ecological, conservation-related, phylogeographic and evolutionary studies of this group of insects. We provide this data in two forms; the basic data and as processed continuous and multinomial data, to enhance its potential usage

Integrating three comprehensive datasets shows that mitochondrial DNA variation is linked to species traits and paleogeographic events in European butterflies

Understanding the dynamics of biodiversity, including the spatial distribution of genetic diversity, is critical for predicting responses to environmental changes, as well as for effective conservation measures. This task requires tracking changes in biodiversity at large spatial scales and correlating with species functional traits. We provide three comprehensive resources to understand the determinants for mitochondrial DNA differentiation represented by i) 15,609 COI sequences and ii) 14 traits belonging to 307 butterfly species occurring in Western‐Central Europe and iii) the first multi‐locus phylogenetic tree of all European butterfly species. By applying phylogenetic regressions we show that mitochondrial DNA spatial differentiation (as measured with Gst, G'st, D and Dst) is negatively correlated with species traits determining dispersal capability and colonization ability. Thanks to the high spatial resolution of the COI data, we also provide the first zoogeographic regionalization maps based on intraspecific genetic variation. The overall pattern obtained by averaging the spatial differentiation of all Western‐Central European butterflies shows that the paradigm of long‐term glacial isolation followed by rapid pulses of post‐glacial expansion has been a pervasive phenomenon in European butterflies. The results and the extensive datasets we provide here constitute the basis for genetically‐informed conservation plans for a charismatic group in a continent where flying insects are under alarming decline

An updated checklist of the European Butterflies (Lepidoptera, Papilionoidea)

This paper presents an updated checklist of the butterflies of Europe, together with their original name combinations, and their occurrence status in each European country. According to this checklist, 496 species of the superfamily Papilionoidea occur in Europe. Changes in comparison with the last version (2.6.2) of Fauna Europaea are discussed. Compared to that version, 16 species are new additions, either due to cryptic species most of which have been discovered by molecular methods (13 cases) or due to discoveries of Asian species on the eastern border of the European territory in the Ural mountains (three cases). On the other hand, nine species had to be removed from the list, because they either do not occur in Europe or lost their species status due to new evidence. In addition, three species names had to be changed and 30 species changed their combination due to new evidence on phylogenetic relationships. Furthermore, minor corrections were applied to some authors¿ names and years of publication. Finally, the name Polyommatus ottomanus Lefèbvre, 1831, which is threatened by its senior synonym Lycaena legeri Freyer, 1830, is declared a nomen protectum, thereby conserving its name in the current combination Lycaena ottomana.VL was supported by grant N 14-14-00541 from the Russian Science Foundation to the Zoological Institute of the Russian Academy of Sciences and ZF by grant 14- 36098G from the Czech Science Foundation

The direct drivers of recent global anthropogenic biodiversity loss

Effective policies to halt biodiversity loss require knowing which anthropogenic drivers are the most important direct causes. Whereas previous knowledge has been limited in scope and rigor, here we statistically synthesize empirical comparisons of recent driver impacts found through a wide-ranging review. We show that land/sea use change has been the dominant direct driver of recent biodiversity loss worldwide. Direct exploitation of natural resources ranks second and pollution third; climate change and invasive alien species have been significantly less important than the top two drivers. The oceans, where direct exploitation and climate change dominate, have a different driver hierarchy from land and fresh water. It also varies among types of biodiversity indicators. For example, climate change is a more important driver of community composition change than of changes in species populations. Stopping global biodiversity loss requires policies and actions to tackle all the major drivers and their interactions, not some of them in isolation.Fil: Jaureguiberry, Pedro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; ArgentinaFil: Titeux, Nicolas. German Centre For Integrative Biodiversity Research (idiv) Halle-Jena-Leipzig; Alemania. Luxembourg Institute Of Science And Technology; Luxemburgo. Helmholtz Zentrum Für Umweltforschung; AlemaniaFil: Wiemers, Martin. Helmholtz Zentrum Für Umweltforschung; Alemania. Senckenberg Gesellschaft Für Naturforschung; AlemaniaFil: Bowler, Diana E.. German Centre For Integrative Biodiversity Research (idiv) Halle-Jena-Leipzig; Alemania. Universitat Jena; Alemania. Helmholtz Zentrum Für Umweltforschung; AlemaniaFil: Coscieme, Luca. Hot Or Cool Institute; AlemaniaFil: Golden, Abigail S.. University of Washington; Estados Unidos. German Centre For Integrative Biodiversity Research (idiv) Halle-Jena-Leipzig; Alemania. Department Of Marine And Coastal Sciences; Estados UnidosFil: Guerra, Carlos A.. German Centre For Integrative Biodiversity Research (idiv) Halle-Jena-Leipzig; Alemania. Martin Luther University Halle Wittenberg; AlemaniaFil: Jacob, Ute. Universität Oldenburg; Alemania. Alfred-Wegener-Institut Helmholtz-Zentrum Für Polar- Und Meeresforschung; AlemaniaFil: Takahashi, Yasuo. Institute For Global Environmental Strategies; JapónFil: Settele, Josef. German Centre For Integrative Biodiversity Research (idiv) Halle-Jena-Leipzig; Alemania. University Of The Philippines, Los Baños; Filipinas. Helmholtz Zentrum Für Umweltforschung; AlemaniaFil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; ArgentinaFil: Molnár, Zsolt. Institute Of Ecology And Botany; HungríaFil: Purvis, Andy. Imperial College London; Reino Unido. Natural History Museum; Reino Unid

Arbeitsmarktwirkungen sind gering: Reformvorschläge zur Grundsicherung für Arbeitsuchende

"Das IAB äußert sich in dieser Stellungnahme zu ausgewählten Aspekten des Gesetzentwurfes der Regierungsfraktionen CDU/CSU und FDP sowie der Anträge der Fraktionen Bündnis 90/Die Grünen, Die Linke und SPD. Sie betreffen die Begründung des Regelbedarfs (insbesondere zur Bestimmung der 'verdeckten Armut'), die neue Hinzuverdienstregelung, die Forderungen nach einem allgemeinen Mindest-lohn, die Förderung der Integration und Teilhabe von Langzeitarbeitslosen am Arbeitsleben, den Vorrang für Ausbildung für Menschen unter 25 Jahren im SGB II und die Einführung eines 'Sozialen Arbeitsmarkts'. Die voraussichtlichen Arbeitsmarktwirkungen, die sich aus der Umsetzung des Entwurfes zu einem Gesetz zur Ermittlung von Regelbedarfen und zur Änderung des Zweiten und Zwölften Buches Sozialgesetzbuch ergeben, sind nach Ansicht des IAB gering. Aus der Regelsatzanpassung resultiert auch keine nennenswerte Verschlechterung der Anreize zur Aufnahme einer Beschäftigung im Niedriglohnbereich. Die Anpassung wird zudem keine größeren fiskalischen Belastungen verursachen. Von der Reform der Freibeträge bei Erwerbstätigkeit im SGB II werden ebenfalls keine signifikanten Arbeitsmarktwirkungen und Zusatzkosten ausgehen." (Autorenreferat)"In this statement, the IAB comments on selected aspects of a draft law submitted by the governing parliamentary parties of the Christian Democrats (CDU/CSU) and Liberals (FDP), and on petitions filed by the Green (Bündnis 90/Die Grünen) and Left (Die Linke) parties and the Social Democrats (SPD). These propositions are concerned with needs for regulation (in particular for the identification of 'concealed' poverty), the new unemployment benefit top-up regulation, claims for a general minimum wage, promotion of the integration and participation of long-term unemployed people in the working life, the priority of education for people under 25 within the scope of the Social Code II, and the introduction of a 'social labour market'. The IAB suggests that foreseeable labour market effects, resulting from the implementation of a law on the identification of benefit recipients’ needs and on changes in the Social Codes II and XII, are rather small. Neither does the adjustment of regu-lation decrease incentives to take up work in the low-wage sector. The adjustment, furthermore, will not create higher fiscal burdens. Neither will the reform of tax exemption (of those topping up Unemployment Benefit II transfers by earnings from minor employment) have significant effects on the labour market or lead to addi-tional costs." (author's abstract