West Palearctic Research Articles

A contribution to the knowledge of the genus Niphargus (Arthropoda: Crustacea: Amphipoda) in northwestern Iran

Niphargus Schiödte, 1849 is the most diverse subterranean amphipod genus in the West Palearctic, distributed between Ireland and Iran. The exploration of Niphargus diversity in the Zagros Mountains in Iran has begun a decade ago, and the number of newly described species is increasing fast. Recently, we collected several specimens of Niphargus from springs located in the northern parts of the Zagros Mountains in Iran. Detailed morphological observations and molecular phylogenetic analyses using 28S rDNA and COI markers revealed that these animals comprise two taxonomic units distinct from all hitherto described species. We hypothesize they should be treated as new species, named as Niphargus qalatianensis sp. nov. and Niphargus arasbaranensis sp. nov. Molecular phylogenetic analyses have shown that N. qalatianensis sp. nov. is nested within a monophyletic Middle Eastern clade known from previous studies. By contrast N. arasbaranensis sp. nov., is with low support associated to European species and suggests that the Iranian karst was not colonized two but three times. To gain a more comprehensive understanding of the intricate biogeographical history of the genus Niphargus in the Middle East, additional sampling in the region is imperative.

A phylogenomic monograph of West-Palearctic Nomada (Hymenoptera: Apidae)

Abstract We reconstruct the phylogeny of the most speciose genus of cuckoo bees, genus Nomada Scopoli, 1770, using 221 species from throughout its distribution, yet with a strong emphasis on the West Palearctic. For phylogenetic reconstruction, we sequenced ultraconserved elements, allowing robust phylogenetic estimates with both concatenation and coalescent-based methods. By integrating extensive information on Nomada host records, we study macroevolutionary patterns of host associations, transitions, and phylogenetic conservatism. Using Bayesian divergence time estimates, we assess the historical biogeography of the genus, focusing on the West Palearctic. Our results show that Nomada likely originated in the Eastern Mediterranean and Near Eastern region, and likely expanded its range to a near-global distribution from there. We recovered long-standing phylogenetic conservatism in the host usage of Nomada and provided strong statistical evidence for an ancestral host association with Andrena and its most recent common ancestor. However, host transitions occurred multiple times independently in the natural history of Nomada, and species of the genus are brood parasites in at least 5 genera and 4 different families of bees in the Old World. At last, we systematically revise the taxonomy of the Old World Nomada by integrating morphological study with our well-supported phylogenetic estimates. We re-establish the genus Acanthonomada Schwarz, 1966, stat. res., as a distinct, second genus in the tribe Nomadini. We recognize 13 subgenera for Nomada, 9 of which are described as new: Afronomada Straka and Bossert, subgen. nov., Colliculla Straka, subgen. nov., Gestamen Straka, subgen. nov., Hungias Straka, subgen. nov., Mininomada Straka, subgen. nov., Nomacolla Straka, subgen. nov., Nomonosa Straka, subgen. nov., Plumada Straka, subgen. nov., and Profuga Straka, subgen. nov. Aside from the subgenus Nomada s.s., we reinstitute 3 previously synonymized subgenera: Heminomada Cockerell, 1902, stat. res., Holonomada Robertson, 1903, stat. res., and Hypochrotaenia Holmberg, 1886 stat. res. A total of 15 subgeneric names are formally synonymized with the newly established subgeneric concepts.

ANALYSIS OF THE AMPHIPOD FAUNA OF CONTINENTAL RESERVOIRS IN THE NORTHWEST OF RUSSIA’S EUROPEAN PART

The faunal composition of Amphipoda crustaceans of mainland water bodies in northwestern Russia is analyzed. To compile a checklist of amphipod species, data from original collections made in 1998–2021, the collections of the Zoological Institute, and literary information are used. To date, 29 species of amphipods have been recorded from the continental waters of the region. An arealogical analysis of the faunal composition reveals six groups or distribution patterns: Holarctic, West Palearctic, and North Atlantic estuaries, and further three groups of emigrants (settlers) from the Baikal-Siberian, Ponto-Caspian, and Arctic regions. The greatest species richness is recorded in estuarine sections of Baltic Sea rivers in the territories of the Leningrad and Kaliningrad regions, as well as fresh water bodies of the Republic of Karelia. In terms of species richness, estuarine species amphi-Atlantic in distribution and representatives of allochthonous faunas (invasive species) from the Ponto-Caspian basin dominate the region. The most common in the region are representatives of Holarctic and Palearctic patterns, as well as species of Arctic and Baikal-Siberian origins. Among them, the following species are most often to be recorded in lakes: Gammarus lacustris, Monoporeia affinis, Palaseopsis quadrispinosa and Gmelinoides fasciatus, vs G. zaddachi, G. oceanicus, G. tigrinus and Pontogammarus robustoides in estuarine brackish waters. Human-mediated introductions, both intentional and accidental, must have facilitated the appearance of invasive species in the fauna of Russia’s northwestern region. In the future, an increase in species richness in the region can be expected to occur in two ways: through the introduction of marine species into fresh waters and due to speciation.

Taxonomic notes and key to the West Palearctic Antocha (Antocha) Osten Sacken, 1860 (Diptera, Limoniidae) with description of a new species from Morocco.

The Mediterranean Region of the West Palearctic is one of the most species-rich biomes in the world, hosting a high level of endemism and relict species with important conservation value. The North Africa Atlas Mountains (spanning Morocco, Algeria and Tunisia) belong to a poorly-investigated region of the Mediterranean area, with overlooked aquatic biodiversity; hence, a number of species still remain to be discovered. The subgenus Antocha (Antocha) Osten Sacken, 1860 is recorded for the first time from Africa, with a description of A. (A.) staryi Keresztes & Mabrouki sp. nov. from hilly regions of the Middle Atlas region, Morocco. The unique design of the male terminalia differentiates well the newly-discovered species from its closely-related and range-restricted A. (A.) phoenicia Thomas and Dia, 1982. This is in contrast with the high intraspecific and geographically poorly-defined variability of the widespread A. (A.) vitripennis (Meigen, 1830), for which morphological variability of male genital structures is discussed.Illustrations of male genital parts, distribution data and key to the species from the West Palearctic area are also provided.

Formation of the Holarctic Fauna: Dated molecular phylogenetic and biogeographic insights from the Quedius-lineage of Ground-Dwelling Rove Beetles (Coleoptera, Staphylinidae)

Although the Holarctic fauna has been explored for centuries, many questions on its formation are still unanswered. For example, i) what was the impact of the uplift of the Himalaya and Tibetan Plateau?, ii) what were the timings and climate of the faunal bridges connecting the Nearctic and Palearctic regions?, and iii) how did insect lineages respond to the late Paleogene global cooling and regional aridification? To answer these, we developed a phylogenetic dataset of 1229 nuclear loci for a total of 222 species of rove beetles (Staphylinidae) with emphasis in the tribe Quediini, especially Quedius-lineage and its subclade Quedius sensu stricto. Using eight fossils for calibrating molecular clock, we estimated divergence times and then analysed in BioGeoBEARS paleodistributions of the most recent common ancestor for each target lineage. For each species we generated climatic envelopes of the temperature and precipitation and mapped them across the phylogeny to explore evolutionary shifts. Our results suggest that the warm and humid Himalaya and Tibetan Plateau acted as an evolutionary cradle for the Quedius-lineage originating during the Oligocene from where, in the Early Miocene, the ancestor of the Quedius s. str. dispersed into the West Palearctic. With the climate cooling from the Mid Miocene onwards, new lineages within Quedius s. str. emerged and gradually expanded distributions across the Palearctic. In Late Miocene, a member of the group dispersed to the Nearctic region via Beringia before the closure of this land bridge 5.3 Ma. Paleogene global cooling and regional aridification largely shaped the current biogeographic pattern for Quedius s. str. species, many of them originating during the Pliocene and shifting or contracting their ranges during Pleistocene.

Two new species of Centroptilum Eaton, 1869 from North Africa (Ephemeroptera, Baetidae).

Based on recently collected larvae from Algeria and Morocco, the species delimitation within the genus Centroptilum Eaton, 1869 in that region is validated. Two new species are described and illustrated, one from north-eastern Algeria, and one from North Morocco, using an integrated approach with morphological and molecular evidence. A table summarising the morphological differences between the new species and Centroptilumluteolum (Müller, 1776) from Central Europe is provided. Further, molecular evidence for additional undescribed species of Centroptilum in other regions of the West Palearctic is provided and discussed.

Tracing the invasion of a leaf-mining moth in the Palearctic through DNA barcoding of historical herbaria

The lime leaf-miner, Phyllonorycter issikii is an invasive micromoth with an unusually higher number of haplotypes in the invaded area (Europe, Western Siberia) compared to its putative native region (East Asia). The origin of the genetic diversity in the neocolonized region remains unclear. We surveyed over 15 thousand herbarium specimens of lime trees (Tilia spp.) collected across the Palearctic over a period of 252 years (1764–2016) looking for preserved larvae within the archival leaf mines. We found 203 herbarium specimens with leaf mines of Ph. issikii collected in East Asia, one of them dating back to 1830, i.e. 133 years before the description of the species. In contrast, only 22 herbarium specimens collected in the West Palearctic in the last three decades (1987–2015) carried leaf mines. DNA barcoding of archival specimens revealed 32 haplotypes out of which 23 were novel (not known from modern populations) and found exclusively in East Asia. Six haplotypes are shared between both native and invaded areas and only two were responsible for the recent invasion of the Western Palearctic. The remarkable number of newly discovered haplotypes in archival populations supports East Asia as the native region and the source area of invasion.

Diversity of Bacterial Biota in Capnodis tenebrionis (Coleoptera: Buprestidae) Larvae.

The bacterial biota in larvae of Capnodis tenebrionis, a serious pest of cultivated stone-fruit trees in the West Palearctic, was revealed for the first time using the MiSeq platform. The core bacterial community remained the same in neonates whether upon hatching or grown on peach plants or an artificial diet, suggesting that C. tenebrionis larvae acquire much of their bacterial biome from the parent adult. Reads affiliated with class levels Gammaproteobacteria and Alphaproteobacteria (phylum Proteobacteria ca. 86%), and Actinobacteria (ca. 14%) were highly abundant. Most diverse reads belong to the families Xanthomonadaceae (50%), Methylobacteriaceae (20%), Hyphomicrobiaceae (9%), Micrococcaceae (7%) and Geodermatophilaceae (4.5%). About two-thirds of the reads are affiliated with the genera Lysobacter, Microvirga, Methylobacterium, and Arthrobacter, which encompass species displaying cellulolytic and lipolytic activities. This study provides a foundation for future studies to elucidate the roles of bacterial biota in C. tenebrionis.

High cryptic diversity of bitterling fish in the southern West Palearctic

South-east Europe, along with the adjacent region of south-west Asia, is an important biodiversity hotspot with high local endemism largely contributed by contemporary continental lineages that retreated to southern refugia during colder Quaternary periods. We investigated the genetic diversity of the European bitterling fish (Rhodeus amarus) species complex (Cyprinidae) across its range in the western Palearctic, but with a particular emphasis in the region of Balkan, Pontic and Caspian refugia. We genotyped 12 polymorphic microsatellite loci and a partial sequence of mitochondrial gene cytochrome b (CYTB) for a set of 1,038 individuals from 60 populations. We used mtDNA sequences to infer phylogenetic relationships and historical demography, and microsatellite markers to describe fine-scale genetic variability and structure. Our mtDNA analysis revealed six well-supported lineages, with limited local co-occurrence. Two lineages are distributed throughout central and western Europe (lineages “A” and “B”), with two zones of secondary contact. Another two lineages were restricted to the Ponto-Aegean region of Greece (lineages “C” and “D”) and the final two lineages were restricted south of the Caucasus mountains (lineage “E” from the Black Sea watershed and lineage “F” from the Caspian watershed). A signal of recent expansion was revealed in the two widespread lineages and the Ponto-Aegean lineage “C”. The geographic distribution of clusters detected by nuclear microsatellites corresponded well with mitochondrial lineages and demonstrated finely sub-structured populations. A profound population structure suggested a significant role of genetic drift in differentiation among lineages. Lineage divergence in the Ponto-Aegean and Caspian regions are substantial, supporting the validity of two described endemic species (Rhodeus meridionalis as lineage “D” and Rhodeus colchicus as lineage “E”) and invite taxonomic evaluation of the other two southern lineages (Thracean “C” and Caspian “F”).

DNA barcode data reveal biogeographic trends in Arctic non-biting midges.

Chironomid flies (non-biting midges) are among the most abundant and diverse animals in Arctic regions, but detailed analyses of species distributions and biogeographical patterns are hampered by challenging taxonomy and reliance on morphology for species-level identification. Here we take advantage of available DNA barcode data of Arctic Chironomidae in BOLD to analyse similarities in species distributions across a northern Nearctic - West Palearctic gradient. Using more than 260 000 barcodes representing 4666 BINs (Barcode Index Numbers) and 826 named species (some with interim names) from a combination of public and novel data, we show that the Greenland chironomid fauna shows affinities to both the Nearctic and the West Palearctic regions. While raw taxon counts indicate a strong Greenland - North American affinity, comparisons using Chao's dissimilarity metric support a slightly higher similarity between Greenland and West Palearctic chironomid communities. Results were relatively consistent across different definitions of species taxonomic units, including morphologically determined species, BINs, and superBINs based on a ∼4.5% threshold. While most taxa found in Greenland are shared with at least one other region, reflecting circum-Arctic dispersal, our results also reveal that Greenland harbours a small endemic biodiversity. Our exploratory study showcases how DNA barcoding efforts using standardized gene regions contribute to an understanding of broad-scale patterns in biogeography by enabling joint analysis of public DNA sequence data derived from diverse prior studies.

Patterns of size variation in bees at a continental scale: does Bergmann's rule apply?

Body size latitudinal clines have been widley explained by the Bergmann's rule in homeothermic vertebrates. However, there is no general consensus in poikilotherms organisms in particular in insects that represent the large majority of wildlife. Among them, bees are a highly diverse pollinators group with high economic and ecological value. Nevertheless, no comprehensive studies of species assemblages at a phylogenetically larger scale have been carried out even if they could identify the traits and the ecological conditions that generate different patterns of latitudinal size variation. We aimed to test Bergmann's rule for wild bees by assessing relationships between body size and latitude at continental and community levels. We tested our hypotheses for bees showing different life history traits (i.e. sociality and nesting behaviour). We used 142 008 distribution records of 615 bee species at 50 × 50 km (CGRS) grids across the West Palearctic. We then applied generalized least squares fitted linear model (GLS) to assess the relationship between latitude and mean body size of bees, taking into account spatial autocorrelation. For all bee species grouped, mean body size increased with higher latitudes, and so followed Bergmann's rule. However, considering bee genera separately, four genera were consistent with Bergmann's rule, while three showed a converse trend, and three showed no significant cline. All life history traits used here (i.e. solitary, social and parasitic behaviour; ground and stem nesting behaviour) displayed a Bergmann's cline. In general there is a main trend for larger bees in colder habitats, which is likely to be related to their thermoregulatory abilities and partial endothermy, even if a ‘season length effect’ (i.e. shorter foraging season) is a potential driver of the converse Bergmann's cline particularly in bumblebees.

ФАУНА РУЧЕЙНИКОВ (TRICHOPTERA, INSECTA) ЦЕНТРАЛЬНОГО И СЕВЕРНОГО КАЗАХСТАНА

Caddis flies of Kazakhstan are studied rather poorly. As for 2018, 151 species of 59 genera of 17 families among 14548 species of World fauna are known in Kazakhstan. The largest numbers of species are found in mountain and submontane regions of Tian Shan and Altay (Southern and Eastern Kazakhstan), caddis flies of Central and Northern Kazakhstan are studied much worse. Nowadays are known 22 caddis flies species from Central Kazakhstan and 13 species from North Kazakhstan. Caddis flies research in 2015-2016 made it possible to expand the list of caddis Central Kazakhstan to 1 species, list caddis North Kazakhstan – to 4 species. Molanna albicans (Zetterstedt, 1840) and Grammotaulius nitidus (Mueller, 1764) are recorded for Kazakhstan at first. Detected species are limnophilous and widely distributed in Holarctic, Transpalearctic and sometimes Transpalearctic-Oriental Regions. Comparative analysis showed, that Central and Northern Kazakhstan are uniform in the biogeographical aspects, which are close to Southern trans-Ural (West Palearctic). The resemblance to South-Western Siberia is much lower. The article is illustrated by original photographs of the male genitalia of these species. Key words: Trichoptera, fauna, male genitals, Central Kazakhstan, North Kazakhstan.

<strong>Keynote: The Trichoptera fauna of Asia</strong>

Many foreign and indigenous scientists have contributed extensively to our knowledge of Asian caddisflies. At the time of the Trichoptera Symposium in 2012, the number (1,090 spp.) and density (43 spp./Gm2) of Trichoptera species in the East Palearctic Region were the least for any of the Earth’s 7 biogeographic regions as defined by the Trichoptera World Checklist; the number (5,313 spp.) and density (405 spp./Gm2) in the Oriental Region were the greatest. Limnephilidae, Rhyacophilidae, and Leptoceridae are the most speciose families in the East Palearctic Region; Leptoceridae, Hydropsychidae, and Hydroptilidae in the Oriental Region. In the East Palearctic Region, 288 fossil species have been reported, most of which are ichnotaxa, sedimentary impressions of larval cases for which the family is unknown; in the Oriental Region, only two fossil species are known, both from Burmese amber. Based on cosmopolitan species, the distinction between the East Palearctic and Oriental Regions is as great or greater than average; the distinction between the East and West Palearctic Regions is less than average but between the East Palearctic and the Nearctic Region is greater; the fauna of the Oriental Region is conspicuously endemic. Future increases in knowledge of the Asia Trichoptera fauna will result from accelerated collecting in poorly explored regions; increasing the rate of identifying and describing new and poorly known species, especially with use of modern molecular, computational, and networking tools and especially focusing on descriptions of larvae in the Oriental Region; mentoring young Asian Trichoptera scientists for excellence in research quality and productivity; and promoting international collaborations for theoretical and applied science in Asia.

Morphology versus DNA barcoding: two sides of the same coin. A case study of Ceutorhynchus erysimi and C. contractus identification.

Genotyping of 2 well-known weevil species from the genus Ceutorhynchus (Coleoptera: Curculionidae) distributed in west Palearctic, C. erysimi and C. contractus, revealed phenotype versus genotype inconsistencies in a set of 56 specimens (25 C. erysimi and 31 C. contractus) collected from 25 locations in Serbia and Montenegro. An analysis of the mitochondrial cytochrome oxidase subunit I gene (COI), widely used as a barcoding region, and a nuclear gene, elongation factor-1α (EF-1α), revealed stable genetic divergence among these species. The average uncorrected pairwise distances for the COI and EF-1α genes were 3.8%, and 1.3%, respectively, indicating 2 genetically well-segregated species. However, the genetic data were not congruent with the phenotypic characteristics of the studied specimens. In the first place, C. erysimi genotypes were attached to specimens with phenotypic characteristics of C. contractus. Species-specific PCR-RFLP assays for the barcoding gene COI were applied for the molecular identification of 101 additional specimens of both morphospecies (33 C. erysimi and 68 C. contractus) and were found to confirm this incongruity. The discrepancy between the genetic and morphological data raises the question of the accuracy of using a barcoding approach, as it may result in misleading conclusions about the taxonomic position of the studied organism. Additionally, the typological species concept shows considerable weakness when genetic data are not supported with phenotypic characteristics as in case of asymmetric introgression, which may cause certain problems, especially in applied studies such as biological control programs in which the biological properties of the studied organisms are the main focus.

Amphibians crossing the Bering Land Bridge: Evidence from holarctic treefrogs (Hyla, Hylidae, Anura)

Based on an updated, time-calibrated phylogeny and applying biogeographical model testing and diversification analysis, we re-examined systematics and biogeography of the Holarctic treefrog genus Hyla with a focus on the East Asian species. We analyzed four mitochondrial genes (12S and 16S rRNA, tRNALeu, ND1) and one nuclear gene (POMC) for 192 samples representing 30 species of Hyla. Based on our results we suggest that H. ussuriensis is a synonym of H. japonica. Specimens from Sakhalin and Kunashir Islands might represent a cryptic species within H. japonica. We confirm earlier hypotheses that the genus Hyla originated during the Eocene to Early Oligocene and that Eurasian species originated from two independent dispersal events from North America via the Bering Land Bridge. Middle Eocene to Oligocene dispersal gave rise to the most recent common ancestor of the West Palearctic H. arborea-group and the East Palearctic, newly defined, H. chinensis-group. The Northeast Asian H. japonica-group resulted from a second wave of colonization from the Nearctic. A trans-Atlantic dispersal route could be excluded. Dispersal of the H. arborea-group to the western Palearctic coincides with the closure of the Turgai Strait at the end of the Oligocene. Diversification of Hyla decreased at the end of the Middle Miocene, possibly coinciding with the end of the Mid Miocene Climatic Optimum and the advent of cooler and drier climates in the Northern Hemisphere.

Pleistocene‐driven diversification in freshwater zooplankton: Genetic patterns of refugial isolation and postglacial recolonization in Leptodora kindtii (Crustacea, Cladocera)

In this study we investigate the extent to which successive range fragmentations, postglacial expansions, and recolonizations influenced intraspecific genetic diversity and patterns of diversification in freshwater zooplankton. Specifically, we explore the global phylogeography of the Holarctic predatory cladoceran Leptodora kindtii Focke, 1844. Phylogenetic analyses based on the mitochondrial cytochrome c oxidase subunit I recover seven deeply divergent lineages (11.8–20.0%) comprising two sister clades within the Nearctic and east Palearctic and three related clades within the west Palearctic. Strong continental endemism was also supported by the nuclear 18S ribosomal gene. Intra‐continental divergence levels (< 6%) suggest that many speciation events in Leptodora were initiated by glacial isolation. Demographic and network analyses indicate that in the Nearctic and west Palearctic, regions heavily affected by the Pleistocene glaciations, Leptodora persisted in both classic (e.g., Cascadia, Mississippi, southern Europe) and cryptic refugia (e.g., northeastern Europe, Carpathian basin). Strong signatures of late Pleistocene range expansions and secondary contact were observed in most clades. Deeper intraspecific phylogeographic structure occurs across much finer geographic scales in unglaciated regions of the east Palearctic than across glaciated Nearctic and west Palearctic regions. Leptodora's ecological and biological characteristics, such as a reliance on temperate, permanent lakes, weak dispersal ability, and specific geological settings likely shaped the complex pattern of diversification in this important planktonic predator. Our data suggest the Pleistocene glaciations were important in the diversification of Holarctic freshwater zooplankton.

Quaternary rodent and insect faunas of the Urals and Western Siberia: Connection between Europe and Asia

Abstract The biogeographic groups of rodent (arvicoline) and beetle species in the modern fauna of the Urals and Western Siberia based on the longitudinal patterns of their present-day distribution are established. Occurrence of species of the Transpalearctic, West Palearctic (European), East-Palearctic, and Central-Palearctic faunal groups is estimated at different stages of the Late Quaternary development of the present-day fauna of the region. The border between the European and East-Palearctic faunal groups represents a vast territory where the ranges of the species included in those groups intersect. The border territory includes the Ural Mountains and the lowlands lying west and east of it (the East-European Plain and the West Siberian Plain). The biomes of those territories represent the North-Eurasian corridors providing migrations of species from the different faunal complexes in the west–east, east–west, and north–south directions. The importance of the study area for faunal correlations throughout the continent is determined by presence of the key arvicoline taxa traditionally used for correlation purposes (taxa Dicrostonyx, Lagurus, Microtus gregalis), and by the gradual change in faunal composition along the latitudinal gradient of environment at any given stage of the Quaternary. Transzonal faunal correlations within the Urals and Western Siberia integrating the evolutionary morphological studies of the key taxa and assessment of the faunal successions may serve as a basis for Trans-Eurasian correlation.

Phylogeny and biogeography of Chinese Heracleum (Apiaceae tribe Tordylieae) with comments on their fruit morphology

The Sino-Himalayan Mountain Region is a major center of diversity of Heracleum (Apiaceae, Tordylieae, Tordyliinae), yet the phylogenetic relationships among its rich diversity of endemic taxa are largely unknown. In this study, we (1) investigate the monophyly of Heracleum and ascertain the phylogenetic placements of those species of Heracleum from China that fall outside of the group; (2) estimate phylogenetic relationships within Heracleumsensu stricto, particularly among those species occurring in the Hengduan Mountain Region of southwestern China; and (3) reconstruct the biogeographic history of Heracleum sensu stricto. To achieve these objectives, Bayesian inference and maximum-parsimony analyses of the nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) region and five noncoding chloroplast DNA (cpDNA) loci (rps16-trnK, rps16 intron, trnQ-rps16, rpl32-trnL, and psbA-trnH), separately and in combination, were carried out. Fruit morphology of the Chinese Heracleum species was examined. Statistical dispersal-vicariance analysis (S-DIVA) was used to reconstruct the biogeographic history. In total, 114 accessions were considered, which included 88 species representing eight genera of subtribe Tordyliinae, 28 of the 29 species of Heracleum occurring in China, and 4 species of Angelica as outgroups. The genus Heracleum is monophyletic upon the exclusion of nine species of Heracleum from China and the inclusion of the Caucasian monotypic genera Mandenovia and Symphyoloma. These nine excluded species show affinities to Tetrataenium, Semenovia, and Angelica. The species of Heracleum endemic to southwestern China form a moderately to strongly supported clade; however, major discordance between the ITS- and cpDNA-derived phylogenies, lack of resolution in the trees, and polyphyly of several species preclude unequivocal hypotheses of phylogenetic relationship. The distinctly clavate vittae, shorter than the length of the mericarp, is a diagnostic feature for Heracleumsensu stricto. Heracleum originated in the West Palearctic, with three possible migration routes leading to its present-day distribution. Major topological conflicts between the ITS and cpDNA trees may be due to interspecific hybridization and/or incomplete lineage sorting. The taxonomic limits of many Chinese Heracleum species remain unclear, possibly because of recent radiation in the Hengduan Mountains.

Eight New Microsatellite Loci of the Western PalearcticHyles euphorbiaeComplex (Lepidoptera, Sphingidae)

We describe eight new micro satellite loci for hawkmoths of the Hyles euphorbiae complex. They are polymorphic (except for one locus in one population) with 2–18 alleles per locus, an expected heterozygosity between 0.14 and 0.94, and an observed heterozygosity between 0.10 and 0.75. As typical for Lepidoptera, the yield of new loci was low due to the presence of microsatellite gene families and variable flanking regions. These microsatellites provide informative results in population studies of the West Palearctic H. euphorbiae complex since cross amplification for H. euphorbiae and H. tithymali was successful for all but one locus.

Correlations of geographic distribution and temperature of embryonic development with the nuclear DNA content in the Salamandridae (Urodela, Amphibia)

We used flow cytometry to measure the nuclear DNA content in erythrocytes of 27 salamandrid species. Across these species, diploid genome size varied more than 2 fold (51.3-104.4 pg). According to genome size and geographic distribution, 3 groups of newt species were recognized: West Palearctics with smaller amounts of nuclear DNA; Nearctic, with intermediate values; and East Asiatic, with higher genome sizes. Viviparous West Palearctic salamanders differed from most of the oviparous West Palearctic newts in possessing larger genome sizes. The nuclear DNA content strongly correlates with species range limits. At the same temperature, embryos of salamandrid species with larger genome sizes have a markedly longer developmental time than those with smaller genomes. We present an analysis of the relationships between the amount of nuclear DNA and water temperature at the breeding sites.