Western Palaearctic Species Research Articles

The phylogenetic relationships betweenDryocosmus,Chilaspisand allied genera of oak gallwasps (Hymenoptera, Cynipidae: Cynipini)

AbstractOak gallwasps (Hymenoptera; Cynipidae, tribe Cynipini) are cyclically parthenogenetic insects that induce galls on specific plant hosts in the family Fagaceae. Understanding the processes underlying the evolution of specific oak associations requires knowledge of the phylogenetic relationships among oak gallwasp genera. Although three major lineages of oak gallwasps have been identified, the status and relationships of several species‐poor but biologically significant genera remain unresolved. Two such genera areChilaspisandDryocosmus, whose western palaearctic species all gall oaks in the sectionCerris.Dryocosmusis particularly significant biologically because it includes: (a) the only palaearctic gallwasp to gall chestnuts,Castanea, and (b) nearctic species. The oak sectionCerrisis wholly absent from the nearctic, and the relationship between palaearctic and nearcticDryocosmusis significant for patterns of host plant evolution in the tribe as a whole. We examined the relationships betweenChilaspis,Dryocosmusand other oak cynipid genera using cladograms from sequence data for two mitochondrial loci (cytochromecoxidase subunit I and cytochromeb) and two nuclear loci (the 28S ribosomal gene regions D2 and D3–5). Our analyses support the following conclusions: (1) palaearcticChilaspisandDryocosmusspecies form an intermingled monophyletic group. (2) We propose thatChilaspisMayr, 1881 is asyn.n.ofDryocosmusGiraud, 1859 and propose the nameD. mayrias acomb.rev.for the species previously namedC. mayri, andD. nitidusandD. israeliascomb.n.ofC. nitidaandC. israeli, respectively. (3) We reassess the utility of morphological characters previously regarded as diagnostic for these genera. (4) Two species previously known only from a single generation represent two halves of a single species lifecycle.Dryocosmus nervosusis here designated asyn.n.ofD. cerriphilus. (5) The nearctic speciesD. favuslies outside the palaearcticChilaspis/Dryocosmusclade, andDryocosmusas currently recognized is not a monophyletic group. (6)Dryocosmus/Chilaspisis closely related to the other oak gallwasp taxa (Aphelonyx,Plagiotrochus,Pseudoneuroterus,Trichagalma, and someNeuroterusspecies) galling sectionCerrisoaks. This implies an early branching evolution of this oak association within this group, and supports previous work showing the rarity of oak gallwasp host shifts.

A rangewide phylogeography of Hermann's tortoise, Testudo hermanni (Reptilia: Testudines: Testudinidae): implications for taxonomy

Hermann's tortoise (Testudo hermanni), the best‐known western Palaearctic tortoise species, has a rare natural distribution pattern comprising the Mediterranean areas of the Iberian, Apennine, and Balkan Peninsulas, as well as Sicily, Corsica and Sardinia. The western part of this range is traditionally considered habitat for T. h. hermanni, while T. h. boettgeri occurs in the Balkans. Taxonomy of this tortoise has been challenged in recent years, with the two subspecies being considered full species and the central Dalmatian populations of T. h. boettgeri being considered a third species, T. hercegovinensis. Using an mtDNA fragment approximately 1150 bp long (cytochrome b gene and adjacent portion of tRNA‐Thr gene), we investigated mtDNA diversity with regard to contrasting concepts of two subspecies or three species. Seven closely related haplotypes were identified from the western Mediterranean and 15 different, in part much‐differentiated, haplotypes from the Balkans. Western Mediterranean haplotypes differ from Balkan haplotypes in 16–42 mutation steps. One to seven mutation steps occur within western Mediterranean populations. Balkan haplotypes, differing in 1−37 nucleotides, group in parsimony network analysis into three major assemblages that display, in part, a similar degree of differentiation to that of western Mediterranean haplotypes relative to Balkan haplotypes. Rates of sequence evolution are different in both regions, and low divergence, palaeogeography and the fossil record suggest a slower molecular clock in the western Mediterranean. While monophyly in western Mediterranean haplotypes is well‐supported, conflicting evidence is obtained for Balkan haplotypes; maximum parsimony supports monophyly of Balkan haplotypes, but other phylogenetic analyses (Bayesian, ML, ME) indicate Balkan haplotypes could be paraphyletic with respect to the western Mediterranean clade. These results imply a process of differentiation not yet complete despite allopatry in the western Mediterranean and the Balkans, and suggest all populations of T. hermanni are conspecific. In the western Mediterranean no clear geographical pattern in haplotype distribution is found. Distribution of Balkan haplotypes is more structured. One group of similar haplotypes occurs in the eastern Balkans (Bulgaria, Republic of Macedonia, Romania and the Greek regions Evvia, Macedonia, Peloponnese, Thessaly and Thrace). Two distinct haplotypes, differing in eight to nine mutation steps from the most common haplotype of the first group, are confined to the western slope of the Taygetos Mts. in the Peloponnese. Yet another group, connected over between four and 23 mutation steps with haplotypes of the eastern Balkan group, occurs along the western slope of the Dinarid and Pindos Mts. (Istria, Dalmatia, western Greece). Taygetos haplotypes are nested within other haplotypes in all phylogenetic analyses and support for monophyly of the other Balkan groups is at best weak. We conclude that using the traditional two subspecies model should be continued for T. hermanni. Phylogeographies of T. hermanni and Emys orbicularis, another codistributed chelonian, are markedly different, but share a few similarities. Both were forced to retreat to southern refuges during Pleistocene glaciations. With the advent of Holocene warming, E. orbicularis underwent rapid range expansion and temperate regions of Europe and adjacent Asia were recolonized from refuges in the Balkans and the northern Black Sea Region. By contrast, T. hermanni remained more or less confined to refuges and nearby regions, resulting in a much smaller range, and allopatric and parapatric distribution of haplotype groups and clades. MtDNA lineages are more diverse in E. orbicularis than they are in T. hermanni on southern European peninsulas, indicating several distinct glacial refuges in close proximity and extensive intergradation during Holocene range expansion for E. orbicularis. In T. hermanni it is likely that only on the Balkan Peninsula was more than one refuge located, corresponding to the parapatric ranges of haplotype groups currently there. On the old western Mediterranean islands Corsica and Sardinia no differentiated (E. orbicularis) or only weakly differentiated haplotypes (T. hermanni) occur, even though there is evidence for the presence of both species on Corsica since at least the Middle Pleistocene. High mountain chains constitute major barriers separating distinct mtDNA clades or groups in each species.

Two new species of Sciophila Meigen (Diptera: Mycetophilidae) from Turkey, with a key to the Western Palaearctic species of the S. lutea Macquart group

Two new species, Sciophila muglolutea sp. n. and S. turkolutea sp. n., from southeastern Turkey are described and illustrated. The second record of S. delphis Chandler, previously known only from the holotype, is reported. A key to the Western Palaearctic species of the S. lutea Macquart group is provided, and data on the distribution of the Palaearctic species are presented.

Skull shape in thegenus Apodemus: phylogenetic conservatism and/or adaptation to local conditions

We studied morphological variation among western Palaearctic species of woodmice (genusApodemus). Twenty one dental and skull variables were measured and evaluated using multivariate statistical approaches. A total of 501 specimens of the following 9 species of wood mice were examined:A. hermonensis, A. hyrcanicus, A. uralensis (=microps),A. flavicollis, A. sylvaticus, A. epimelas, A. mystacinus, A. peninsulae, A. agrarius. Species occupying large geographic areas were represented by two or three geographically distant populations. The analyses, based both on original and size adjusted data, revealed congruence between morphological evolution and phylogenetic relationships.

Dryocosmus kuriphilus

<i>Dryocosmus kuriphilus</i>

The Western Palaearctic species of Stenomicra Coquillett (Diptera, Periscelididae, Stenomicrinae), with description of a new species of the subgenus Podocera Czerny

Stenomicra (Podocera) soniae sp. n. is described from Central and Southeast Europe, and compared with morphologically similar species. The type material of Stenomicra (Podocera) delicata (Collin, 1944) is revised and the species redescribed. Both European species of Podocera Czerny are illustrated and their biology and distribution reviewed with a number of new records. A key to the four Western Palaearctic species of Stenomicra is presented.

Morphological discrimination of six species of the genusAnuraphis(Hemiptera: Aphididae), including description of a new species

AbstractThe most relevant morphometric characteristics of six species of the genusAnuraphisDel Guercio were compared.Anuraphis shaposhnikovisp. nov.is described and its morphological differences from the closely related speciesAnuraphis subterraneaare presented. The new species was collected in Sicily and in the central area of the Italian peninsula onMagydaris pastinacea(Lam.) Paol. (Apiaceae) andOpopanax chironium(L.) Koch (Apiaceae), which are its secondary host plants. A key to the viviparous morphs (apterae and alatae) of the seven western Palaearctic species living on secondary hosts is provided. Discriminant functions have been derived to separate both apterae and alatae ofA. shaposhnikoviandA. subterranea.

Water mites of the genus Atractides Koch, 1837 (Acari: Parasitengona: Hygrobatidae) in the western Palaearctic region: a revision

The western Palaearctic species of the water mite genus Atractides are revised, based on the available type material, data published in the literature, and additional material from various collections. This paper includes the following: (1) a survey of the classification history of the genus, with description of all important morphological features, including a brief discussion of the apomorphic characters of Atractides and its presumed sister-group relations; (2) documentation of variability and seasonal changes of morphological indices for a selected species, A. nodipalpis; (3) a detailed description and/or discussion of all published taxa, with designation of lectotypes, definition of diagnostic character combinations, and identification of species dubiae; (4) a dichotomic determination table for the accepted species in the study area. The main object of this study is to animate world-wide systematic and ecological work on this important genus, which is currently hampered by taxonomic confusion. Above species level, the following synonyms are confirmed: Tetramegapus, Taeniomegapus, Megapoides = Atractides; Rhynchomegapus = Tympanomegapus. The following new synonyms are proposed: Octomegapus, Iranomegapus = Atractides. Tympanomegapus and Polymegapus are redefined and accepted as distinct subgenera, while Maderomegapus is degraded to subgenus level. © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society, 2003, 138, 141–378.

Una especie nueva de &lt;i&gt;Tillus&lt;/i&gt; Olivier, 1790 (Coleoptera, Cleridae) de la Península Ibérica

A new species of Tillus Olivier, 1790 (Coleoptera: Cleridae) from the Iberian Peninsula Tillus ibericus sp. nov. is described from the Comunidad Autonoma de Madrid (Spain) and it is compared to the western Palaearctic species of the genus Tillus Olivier, 1790. Tillus ibericus is easily distinguished from all other species of the genus by the structure of the elytral sculpture and its characteristic coloration. This new species shares a general habitus and characters from the antennae, and general body stucture with T. flabellicornis Fairmaire, 1866, a species from northern Africa. All preexisting records of T. flabellicornis in the Iberian Peninsula correspond to T. ibericus.

Estimating the number of species not yet described and their characteristics: the case of Western Palaearctic dung beetle species (Coleoptera, Scarabaeoidea)

To determine the degree of completion of Western Palaearctic dung beetlespecies inventory, as well as to ascertain some of the main geographical andmorphological characteristics associated with the probability of new speciesdescription, the available taxonomic and biogeographic information of the threeScarabaeoidea families that are associated with dung was analyzed. An asymptotic fit of the cumulative number of described species is used to estimatethe approximate number of species not yet described. Variation partitioning andhierarchical decomposition techniques were used to explore the influence ofbody size, geographical range size and geographical location variables on theprocess of species description. Results indicate that the inventories of WesternPalaearctic Scarabaeidae and Geotrupidae species are almost complete, but thataround 16% of Aphodiidae species have yet to be described. As the joint effectof geographical distribution and location is the main determinant of the speciesdescription rate in this family, it was predicted that new dung beetle specieswould be found in smaller areas of the southern and eastern Western Palaearcticregion. The species body size accounted for a negligible fraction in thevariation of the year of description. Both the geographical range size and thegeographical location are essential variables negatively correlated with theyear of species description, whose pure and combined effect can account for highpercentages in the year of description variation (from 41 to 56%). The locationof most of the taxonomic workforce in areas where broader-ranged generalist speciesprevail in the assemblages is the probable reason for this pattern.

Genetic variation and evolution in the genus Apodemus (Muridae: Rodentia)

Genetic variation was studied using protein electrophoresis of 28–38 gene loci in 1347 specimens of Apodemus agrar-ius, A. peninsulae, A. flavicollis, A. sylvaticus, A. alpicola, A. uralensis, A. cf. hyrcanicus, A. hermonensis, A. m. mystacinus and A. m. epimelas, representing 121 populations from Europe, the Middle East, and North Africa. Mean values of heterozygosity per locus for each species ranged from 0.02 to 0.04. Mean values of Nel's genetic distance (D) between the taxa ranged from 0.06 (between A. flavicollis and A. alpicola) to 1.34 (between A. uralensis and A. agrarius). The highest values of D were found between A. agrarius and other Apodemus species (0.62-1.34). These values correspond to those generally observed between genera in small mammals. Our data show that A. agrarius and A. peninsulae are sister species, well-differentiated from other taxa. High genetic distance between A. m. mystacinus and A. m. epimelas leads us to consider them distinct species and sister taxa to other Western Palaearctic species of the subgenus Sylvaemus. The data also suggest a recent separation of members of the latter group from a common ancestor, and subsequent rapid radiation, making it difficult to infer phylogenetic relationships. Some taxonomic implications of the results are discussed further.© 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 395–419.

Molecular Systematics and Phylogeny of Old and New World Ratsnakes, Elaphe Auct., and Related Genera (Reptilia, Squamata, Colubridae)

The phylogenetic relationships of the Holarctic ratsnakes (Elaphe auct.) are inferred from portions of two mitochondrial genes, 12S rRNA and COI. Elaphe Fitzinger is made up of ten Palaearctic species. Natrix longissima Laurenti (type species) and four western Palaearctic species (hohenackeri, lineatus, persicus, and situla) are assigned to Zamenis Wagler. Its phylogenetic affinities with closely related genera, Coronella and Oocatochus, remain unclear. The East Asian Coluber porphyraceus Cantor is referred to a new genus. This taxon and the western European Rhinechis scalaris have an isolated position among Old World ratsnakes. Another new genus is described for four Oriental species (cantoris, hodgsonii, moellendorffi, and taeniurus). New World ratsnakes and allied genera are monophyletic. Coluber flavirufus Cope is referred to Pseudelaphe Mertens and Rosenberg. Pantherophis Fitzinger is revalidated for Coluber guttatus L. (type species) and further Nearctic species (bairdi, obsoletus, and vulpinus). Senticolis triaspis is the sister taxon of New World ratsnakes including the genera Arizona, Bogertophis, Lampropeltis, Pituophis, and Rhinocheilus. The East Asian Coluber conspicillatus Boie and Coluber mandarinus Cantor form a monophyletic outgroup with respect to other Holarctic ratsnake genera and are referred to Euprepiophis Fitzinger. Three Old World species, viz. Elaphe (sensu lato) bella, E. (s.l.) frenata, and E. (s.l.) prasina remain unassigned. The various groups of ratsnakes (tribe Lampropeltini) show characteristic hemipenis features.

Two interesting psocids from Corsica (Psocoptera: Caeciliusidae) with a checklist of the Western Palaearctic species of the family

a checklist of the Western Palaearctic species of the family. - Valenzuela labrostylus sp. n. is described and illustrated (female only). The generic placement of this species remains doubtful because of the presence of labral stylets, which are absent in all other known Valenzuela species. The holotype of Caecilius corsicus Kolbe, 1882 is redescribed and the synonymy of this name with rhenanus Tetens, 1891 is proposed: Valenzuela corsicus (Kolbe) becomes the valid name of the most common Mediterranean species of Caeciliusidae. The 15 presently recognized Western Palaearctic species of this family are listed.

Pupae in nomenclature and identification: West Palaearctic Orthocladius s.str. (Diptera: Chironomidae) revised

Abstract. Orthocladius (Orthocladius) is a subgenus of one of the more speciose genera amongst the Holarctic chironomids of the subfamily Orthocladiinae. Although many names have been applied, adult males are difficult to distinguish. However, the immature stages often are known, and pupae discriminate taxa reliably.Based predominantly upon examination of exuviae, Orthocladius atripluma Kieffer and Orthocladius mitisi Goetghebuer are syn.n. of Orthocladius glabripennis (Goetghebuer); Orthocladius smolandicus Brundin a syn.n. of Orthocladius holsatus Goetghebuer; Orthocladius lenzi Kieffer a syn.n. of Orthocladius oblidens (Walker); Orthocladius rhyacobius Kieffer, Orthocladius rhyacophilus Kieffer and Orthocladius excavatus Brundin are syn.n. of Orthocladius obumbratus Johannsen; Orthocladius saxicola Kieffer and Orthocladius curtiseta Sæther are syn.n. of Orthocladius rubicundus (Meigen).Orthocladius (Orthocladius) vaillanti is described as sp.n., based upon distinctive pupal exuviae. Lectotypes are designated for Orthocladius rhyacobius Kieffer, Orthocladius rhyacophilus Kieffer, Orthocladius rivinus Kieffer, Chironomus rubicundus Meigen and Orthocladius saxicola Kieffer.A key to the thirteen western palaearctic species is given.

A revision of the western Palaearctic species of Urophora Robineau-Desvoidy (Diptera: Tephritidae).

A key is provided to twenty‐four western Palaearctic species of Urophora Robineau‐Desvoidy. The hosts of twenty‐three species which attack Asteraceae are listed, including those being used or investigated as possible weed biocontrol agents. The species are divided into four species groups and the differing host relationships and types of galls induced by these groups are discussed. U.lopholomae sp.n. and U.affinis ssp. calcitrapae ssp.n., associated with Centaurea (Lopholoma) spp. and C. (Calcitrapa) spp. respectively, are described. U. algerica (Hering) and U.sjumorum (Rohdendorf) are both treated as subspecies of U. quadrifasciata (Meigen). U.pontica is given full specific status and U.hispanica is removed from synonymy. The following new synonymies are made (junior synonyms in parentheses): U. angustifascia (Hering) (=Euribia phaeocera Hering); U. cardui (Linnaeus) (=U. reaumurii Robineau‐Desvoidy, lectotype designated); U. jaceana (Hering) (=E.conyzae Hering); U.maura (Frauenfeld) (=E. tecta Hering); U. mauritanica Macquart (=U. lejura Rondani, Trypeta macrura Loew); U.solstitialis (Linnaeus) (=E.sonderupi Hering, U. veruata Rondani); U.stylata (Fabricius) (=E.pia Hering, U. vulcaanica Rondani); U. terebrans (Loew) (=E. approximata Hering, T. eriolepidis Loew, E. manni Hendel). The possibility that U. quadrifasciata is a species complex is discussed; it is also suggested that U.affinis and U.jaceana represent the morphological extremes of a complex. The misuse of the name Musca stylata Fabricius in the genus Myopites Blot is noted.

Rheosmittia(Diptera: Chironomidae): a generic validation and revision of the western Palaerctic species

The chironomid (Diptera) genus Rheosmittia gen. nov. is validated with the designation of a type-species by Brundin, in this paper. Generic diagnoses are given for larva, pupa, male and female imagines. The western Palaearctic species of Rheosmittia are revised, with lectotype designations for R. spinicornis Brundin and R. languida Brundin and the description and/or redescription of imagines of both these species and of R. halvorseni sp. nov. The larva and pupa of R. spinicornis are described and the possible confusion between the pupae of R. halvorseni and R. languida, both of which are described, is noted. The systematic placement of Rheosmittia close to Parakiefferiella is confirmed, and a sister-group relationship with Gynnidiocladius and Lopescladius is suggested. The ecology of the psammophilic (sand-dwelling) larvae of Rheosmittia is briefly discussed, and it is observed that the genus is Holarctic in distribution.

A taxonomic revision of the species of Alloclemensia n.gen. (Lepidoptera: Incurvariidae s. str.)

AbstractThe genus Alloclemensia n.gen. (type-species: Alloclemensia maculata n.sp.), with four species in the Palaearctic region and one species in the Nearctic region, is described and its species are revised. The genus comprises five species: mesospilella (Herrich-Schäffer) [= trimaculella (Herrich-Schäffer)], devotella (Rebel) [= muchei (Soffner), n.syn.] and three new species: americana from North America and unifasciata and maculata from Japan. The monophyly of the genus is demonstrated and the primary dichotomy within the genus is considered to lie between the two western Palaearctic species plus the North American species and the two eastern Palaearctic species. Keys to the species are provided and all species and their genitalia are described and illustrated. Two new combinations and one new synonymy are established and two lectotypes are designated.