Range expansion and the possibility of an emerging contact zone between two subspecies of Chiffchaff Phylloscopus collybita ssp.

The Chiffchaff Phylloscopus collybita is represented in Sweden by two different subspecies; the northern well-established abietinus and the southern recently established collybita which has expanded its range northward during the past two decades. At present, an area approximately 500 km wide separates the two subspecies. In order to document differences between the northern and southern populations we compared morphology, vocalisation, habitat choice, and neutral genetic variation in mitochondrial (mt) DNA and at four microsatellite loci of 30 male Chiffchaffs from each subspecies. Our results show significant differences in several morphological traits and in song. Playback experiments revealed a significant difference in aggressive behaviour depending on which population-specific song that was played to the birds. Mitochondrial DNA was geographically structured with ∼90% of the birds carrying a mt haplotype matching their sample population. No allelic differences at the microsatellite loci were found between populations. Our data demonstrate a substantial differentiation between the northern and southern populations despite gene flow, clearly separating them into the subspecies abietinus and collybita .     

Amplified fragment length polymorphism analysis identifies hybrids between two subspecies of warblers

In the western Pyrenees (Southwest France and Northwest Spain), a narrow hybrid zone exists between the common chiffchaff Phylloscopus (collybita) collybita and the Iberian chiffchaff Phylloscopus (c.) brehmii. In this zone, which is approximately 20 km wide, mixed matings and individuals singing the songs of both taxa occur at substantial frequencies (24 and 8.6%, respectively), suggesting frequent hybridization. Previous studies have shown very weak mitochondrial gene flow (Nm = 0.065), whereas four microsatellites suggested much higher nuclear gene flow (Nm = 4.9). In this study we used the amplified fragment length polymorphism (AFLP) method in order to identify hybrids and early backcrosses. We typed 91 birds from both allopatric and sympatric areas for 12 informative AFLP markers (of > 141 polymorphic fragments), obtained by screening 13 AFLP primer combinations. These individuals were previously typed for song (brehmii, collybita or mixed singers), mitochondrial DNA (mtDNA) haplotype and allelic genotypes at four microsatellite loci. Assignment tests demonstrated that in the zone of sympatry, a substantial number of intermediate genotypes existed among the birds previously believed to be pure collybita and brehmii, based on song and mtDNA haplotype. The majority of the mixed singers had intermediate genotypes. Our data suggest that the fraction of the adult population having a hybrid origin (hybrids or backcrosses) is in the order of 10%. With such a frequency of genetic hybrids, there would have been much more mtDNA introgression than observed, had female hybrids been perfectly fertile/viable. This result is consistent with male-biased gene flow and Haldane's rule.     

Wing shape variation of Finnish and Central European Willow Warblers Phylloscopus trochilus and Chiffchaffs P. collybita

Wing shape variation of European Willow Warblers Phylloscopus trochilus and Chiffchaffs P. collybita was studied using indices calculated from wing formulae. Our data were from free-living local populations of P. t. acredula and P. c. abietinus from southern Finland, and P. t. trochilus and P. c. collybita from southwestern Germany.
There were no significant shape differences between the subspecies of the Willow Warbler in which sexual dimorphism was pronounced. The subspecies of the Chiffchaff were significantly different while the sexes were not. There were also significant differences between adult and immature individuals in the autumn. The intrapopulation variation must be considered in attempts to recognize different subspecies or populations in data on birds caught during migration.
In the Willow Warbler at least, the difference between age-groups was larger in males than in females. If the change in wing formula due to the complete pre-nuptial moult is similar in both sexes, juvenile males intermediate between adult females and adult males suffer disproportionately high mortality. It was earlier suggested that sexual selection increases body size in Willow Warbler males, but the present results imply additional selection pressures for increasing sexual dimorphism.     

Speciation of the Paradeontacylix spp. (Sanguinicolidae) of Seriola dumerili. Two new species of the genus Paradeontacylix from the Mediterranean

Two new species of teleost blood fluke belonging to the sanguinicolid genus Paradeontacylix are described from the greater amberjack, Seriola dumerili, i.e. Paradeontacylix ibericus n. sp. from the Iberian Peninsula and Paradeontacylix balearicus n. sp. from the Balearic Islands. P. ibericus n. sp. and P. balearicus n. sp. show morphological similarities with Paradeontacylix kampachi and Paradeontacylix grandispinus respectively, which occur in mixed infection in S. dumerili from Japan. Multivariate analysis of morphometrical data provided statistical evidence for the separation of four species. However, component by component analysis did not show statistically significant differences between P. balearicus and P. grandispinus. Molecular data based on rITS2 and mCO1 gene sequences also supported the separation into four species. Morphological and molecular data were used to examine phylogenetic relationships between Paradeontacylix species from S. dumerili and other species in the genus. The results coincided in revealing two main branches with P. kampachi+P. ibericus and (((P. grandispinus+P. balearicus) Paradeontacylix sanguinicoloides) Paradeontacylix godfreyi). Paradeontacylix odhneri, for which little data are available, was located basal in a separate branch. This is the only species of Paradeontacylix which parasitizes a non-carangid host which might probably explain the separation from the other species. Paired similarities between the Japanese and the Mediterranean species, despite the large geographic distance, could be explained by the speciation of parasite geminate lines before host separation by tectonic events. Consequently, geographic and historical isolation support the morphological and genetic differences leading to the evolution of the new species described here.     

SERPUKHOVIAN CONODONTS FROM NORTHERN SPAIN AND THEIR BIOSTRATIGRAPHIC APPLICATION

Abstract:  Conodont collections from late Viséan–early Bashkirian deposits include numerous elements related to the cosmopolitan species Gnathodus bilineatus (Roundy) in the Cantabrian Mountains. Arnsbergian beds yield the taxa Gnathodus bilineatus leonicus subsp. nov., G. millarensis sp. nov., G. truyolsi sp. nov. and Gnathodus sp. nov. A. Almost all are also found in limestones of similar age in the Pyrenees, and the G. truyolsi conodont Zone is defined for the upper Serpukhovian of the northern Iberian Peninsula. In addition, numerous references to the proposed taxa come from Austria, Eastern Europe, Northern Africa and Asian localities. The increase in gnathodid diversity and abundance in northern Spain is related to a local Arnsbergian drowning episode of the carbonate platform. G. truyolsi is possibly derived from G. b. bilineatus , and there is morphological convergence with Gnathodus kiensis Pazukhin.     

Taxonomic revision of the genus Succisella (Dipsacaceae) in the Iberian Peninsula

A revision of the three endemic Iberian species of Succisella G. Beck ( S. carvalhoana , S. microcephala and S. andreae-molinae ), based on herbarium studies, SEM photographs and field observations, is presented utilizing morphological, palynological, karyological, biogeographical and ecological characters. The distribution of the species in the Iberian Peninsula is shown in a grid map. Full synonymy is given for all taxa.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 351–364.     

Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review

In the fossil deposits of the Late Pleistocene ( c . 115 000–11 500 years ago), five felid species are recorded in Europe: the wildcat Felis silvestris , Eurasian lynx Lynx lynx , Iberian lynx Lynx pardinus , leopard Panthera pardus and cave lion Panthera leo spelaea . In the Holocene, Europe was colonized by F. silvestris , L. lynx and L. pardinus as well as the lion Panthera leo . The status of P. pardus in post-glacial Europe is unknown. So far, only sparse records indicate that P. pardus survived into the early Holocene. During the Late Glacial, both L. lynx and L. pardinus occurred on the Iberian Peninsula. However, from the Holocene, only the Iberian lynx is recorded in this region. There are subfossil records that indicate that L. pardinus also occurred in central and western France until c . 3000 years ago. Surprisingly, with reservations on the determination of the bones (by J. Altuna), both lions and cave lions seem to be recorded in the Iberian Peninsula in the Late Glacial. There are published records of the lion P. leo in the northern Iberian Peninsula from the early Holocene. However, its presence in Europe on the basis of subfossil records was proven initially from the Atlantic period. In Ponto-Mediterranean regions of Europe, the lion is recorded from the Atlantic to the younger sub-Atlantic.     

Male‐biased gene flow across an avian hybrid zone: evidence from mitochondrial and microsatellite DNA

Mating pattern and gene flow were studied in the contact zone between two morphologically very similar Chiffchaff taxa (Phylloscopus collybita, P. brehmii) in SW France and northern Spain. Mating was assortative in brehmii, but not in collybita. Mixed matings were strongly asymmetric (excess of collybita male × brehmii female pairs), but did produce viable offspring in some cases. Sequence divergence of the mitochondrial cytochrome b gene was 4.6%. Haplotypes segregated significantly with phenotype (only five ‘mismatches’ among 94 individuals), demonstrating that mitochondrial gene flow was very restricted. The estimated proportion of F₁ hybrids in the reproductive population was significantly lower than expected under a closed population model, indicating strong selection against hybrids. Genetic typing of 101 individuals at four microsatellite loci also showed significant population differentiation, but nuclear gene flow was estimated to be 75 times higher than mitochondrial gene flow. This strong discrepancy is probably due to unisexual hybrid sterility (Haldane’s rule). Thus, there is a strong, but incomplete, reproductive barrier between these taxa.     

Phylogenetic relationships of the European lacertid genera Archaeolacerta and Iberolacerta and their relationships to some other 'Archaeolacertae' (sensu lato) from Near East, derived from mitochondrial DNA sequences

Parts of the mitochondrial genes coding for 12SrRNA and 16SrRNA (together about 960 bp) were sequenced for all Mediterranean species of 'Mountain lizards' of the genera Archaeolacerta ( sensu lato ) and Iberolacerta . All subspecies of the Iberian species Iberolacerta cyreni and I. monticola were included in this study. In addition, samples of Apathya cappadocica and Darevskia rudis were analysed to elucidate the relationships of the European 'Mountain lizards' to their possible relatives in the Near East. Maximum parsimony and neighbour joining analyses lead to the following major conclusions: (i) the monophyly of the genus Iberolacerta is very well supported; (ii) Archaeolacerta bedriagae (the type species of the genus) is most basal with respect to the ingroup taxa. If we accept Iberolacerta as a genus, Archaeolacerta becames paraphyletic. Therefore, we propose to restrict Archaeolacerta to the type species and to treat A. mosorensis and A. oxycephala provisionally as members of the collective genus Lacerta ; (iii) within the genus Iberolacerta three groups were found: a Pyrenean group, an Iberian group and I. horvathi . The relationships among these groups remain unresolved; and (iv) the Peña de Francia lizards, described originally as a subspecies of I. cyreni , are in fact more closely related to I. monticola .     

A morphometric study of three closely related taxa in the European Isoetes velata complex

Isoetes velata A. Braun is an amphibious plant with Mediterranean and Ibero-Atlantic distribution, showing marked morphological variability. Within the same geographical area, closely related restricted-range taxa are also present, including I. velata ssp. asturicense (restricted to mountainous areas of the north-west Iberian Peninsula) and I. boryana (restricted to the southern French Atlantic coast). These related taxa have unclear taxonomy and are both threatened. We performed a morphometric study of specimens of the three taxa. Our results suggest that I. velata ssp. asturicense is better considered as a distinct species, I. asturicense Laínz (Laínz), distinguishable from the other two taxa by mean megaspore length ( N  ≥ 30) < 360 µm. In contrast, our results do not support consideration of I. boryana as a separate species, and we here propose that this taxon be considered as a variety, I. velata A. Braun var. boryana M.I. Romero & C. Real stat. nov.   © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 459–464.     

Karyological and taxonomic notes on Ophrys (Orchidoideae, Orchidaceae) from the Iberian Peninsula

Karyological information on Iberian Ophrys species is very limited. This paper provides the haploid and diploid chromosome numbers of 11 taxa of sect. Pseudophrys and sect. Ophrys , both of which are well represented in the Iberian Peninsula, and two taxa from Tunisia. The first data on chromosome numbers for O. vasconica (2 n  = 72, 74), O. ficalhoana (2 n  = 36), O. picta (2 n  = 36), O. sphegifera ( n  = 18, 2 n  = 36, 38) and O. passionis (2 n  = 36) are also presented, confirming the stability of the chromosome number in Ophrys . In addition, populations of the group O. omegaifera ( O. dyris and O. vasconica ), together with tetraploidy, pentaploidy and the existence of aneuploid phenomena, are reported for the first time in Iberia. The basic diploid number is always 2 n  = 36. The karyotypes of several species were analysed. Evolutionary trends in Ophrys chromosomes are discussed. Taxonomic and phytogeographical details are provided on several species or groups of species from Iberia.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003 , 142 , 395−406.     

Glacial history and colonization of Europe by the blue tit Parus caeruleus

Mitochondrial control region sequences from European populations of the blue tit Parus caeruleus were used to reveal the Pleistocene history and the post-glacial recolonization of Europe by the species. The southern subspecies, P. c. ogliastrae was found to represent a stable population with isolation-by-distance structure harboring a lot of genetic variation, and the northern subspecies P. c. caeruleus a recently bottlenecked and expanded population. We suggest that after the last Ice Ages, the subspecies have colonized Europe from two different southern refuges following previously proposed general recolonization routes from the Balkans to northern and Central Europe, and from the Iberian Peninsula north- and eastwards. The two subspecies form a wide secondary contact zone extending from southern Spain to southern France.     

A NEW EARLY DEVONIAN THELODONT FROM CELTIBERIA (SPAIN), WITH A REVISION OF SPANISH THELODONTS

Abstract:  A revision of thelodont scales from the Lower Devonian of the Iberian Chains enables their grouping into two taxa: Turinia pagei and T. nachoi sp. nov. These taxa are clearly distinguishable by morphological and histological features; they also have a different stratigraphic range ( T. pagei is restricted to Lochkovian strata, whereas T. nachoi sp. nov. occurs within lower–middle Pragian rocks). The new species is represented by head, transitional (cephalopectoral) and trunk scales.     

Genetic diversity and phylogeography of the European roe deer: the refuge area theory revisited

The extant taxa of central and northern Europe are commonly believed to derive from Pleistocene ancestors, who moved to the north from three separate glacial refugia: the Iberian and Italian peninsulae, as well as the southern Balkans. The issue of postglacial dispersal patterns was addressed through the investigation of population structure and phylogeography of the European roe deer, Capreolus capreolus . The genetic diversity in 376 individuals representing 14 allegedly native populations across their European range was assessed, using ten autosomal microsatellite loci and restriction fragment length polymorphisms of the mitochondrial D-loop and NADH dehydrogenase 1 gene segments. Our results suggest the existence of three major genetic lineages of roe deer in Europe. One comprises populations in the south-western limit of the species' distribution (i.e. Iberia), where an internal substructure splits a northern from a southern sublineage. A second lineage includes populations of southern and eastern Europe, as well as a separate sublineage sampled in central-southern Italy, where the existence of the subspecies Capreolus c. italicus was supported. In central-northern Europe, a third lineage is present, which appeared genetically rather homogeneous, although admixed, and equally divergent from both the eastern and western lineages. Current patterns of intraspecific genetic variation suggest that postglacial recolonization routes of this cervid to northern Europe could be due to range expansion from one or more refugia in central-eastern Europe, rather than proceeding from the Mediterranean areas.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 85–100.     

Monogenea versus Monogenoidea: the case for stability in nomenclature

Monogenea van Beneden, 1858 is the most appropriate name to apply to the class of Platyhelminthes comprising the monogenean flatworms. Monogenoidea sensu Bychowsky (1937) is considered an emendation of Monogenea van Beneden, 1858. Arguments that support the use of Monogenoidea as the valid name of the class are based on erroneous assumptions of authorship and of “priority” and “rank”, as defined in the International Code of Zoological Nomenclature, a document that does not apply to taxa above the family level. In the absence of defined rules, names for higher taxa should be based on a combination of priority, stability and consensus among specialists, with stability and consensus the most important considerations. The priority of Monogenoidea versus Monogenea is questionable, and both nomenclatural stability and consensus among specialists favour the use of Monogenea rather than Monogenoidea.     

Patterns of genetic diversity in related taxa of Antirrhinum L. assessed using allozymes

Allozyme diversity was studied within and among populations of five related taxa of Antirrhinum L. endemic to the Iberian Peninsula ( A. graniticum Rothm. ssp. graniticum , ssp. brachycalyx Sutton and ssp. ambiguum (Lange) Mateu & Segarra, A. boissieri Rothm. and A. onubensis (Fdez. Casas) Fdez. Casas). All of the studied taxa are obligate outcrossing endemic perennial herbs which form isolated populations. However, the taxa vary in range and population sizes, and are found on different soil types. The level and distribution of allozyme diversity differed widely between taxa: A. graniticum ssp. brachycalyx had the lowest level of allozyme diversity (H_T = 0.09), whilst the highest level was detected in A. boissieri (H_T = 0.25). Total variation was partitioned into within- and among-population variation. The proportion attributable to variation within populations varied from about 67% up to 84.3% and 89.5% in A. graniticum ssp. brachycalyx and A. graniticum ssp. ambiguum , respectively. Both these subspecies also showed little population divergence (G_ST = 0.10 and 0.09, respectively) and had high levels of estimated gene flow (Nm = 2.18 and 2.62, respectively). These results are discussed in relation to geographical proximity of populations and habitat continuity. Isolation by distance was not detected in any of the studied taxa. This result suggests that divergence among populations is due to random genetic drift.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 79 , 299–307.     

Speciation mirrors geomorphology and palaeoclimatic history in African laminate-toothed rats (Muridae: Otomyini) of the Otomys denti and Otomys lacustris species-complexes in the 'Montane Circle' of East Africa

We adopted an integrated systematic approach to delimit evolutionary species and describe phylogeographic, morphometric and ecological relationships in Otomys denti (from the Albertine Rift, Southern Rift in Malawi and the northern Eastern Arc Mountains) and Otomys lacustris (from the Southern Rift in Tanzania and Zambia, and the southern Eastern Arc Mountains). Molecular [cytochrome (cyt) b sequences, 1143 bp, N  = 18], craniometric (classical, N  = 100 and geometric, N  = 60) and ecological (Partial Least Squares regression of shape and ecogeographic variables) approaches show a profound, parallel disjunction between two groups: (1) Eastern Arc and Southern Rift (including the Malawi Rift) ( O. lacustris and Otomys denti sungae ) and (2) Albertine Rift ( Otomys denti denti and Otomys denti kempi ) taxa. Within both groups, cyt b sequences or craniometric analysis provided evidence for the differentiation of both southern and northern Eastern Arc from Southern Rift lineages (across the so-called Makambako Gap). Within the Albertine Rift ( denti – kempi ) lineage, populations from individual mountain ranges differed significantly in skull shape (but not size), but were similar genetically. Over-reliance in the past on very few morphological characters (e.g. number of molar laminae) and a polytypic species concept has obscured phylogenetic relationships and species discrimination in this group. We recognize at least three species in this group, and distinct lineages within two of these species. Each species or lineage was endemic to one of three regions: the Albertine Rift, the Malawi Rift or the Eastern Arc. Our result echo conclusions of recent studies of other mammalian and bird taxa and reflect the geomorphology and palaeoclimatic history of the region.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 913–941.     

Morphological divergence among Spanish Common Crossbill populations and adaptations to different pine species

Crossbills (Loxia spp.) provide a classical avian model of ecological specialization on food resources. Previous studies have suggested that morphometric, genetic and vocal diversification among Common Crossbill Loxia curvirostra populations is better explained by ecological distance (use of different conifers) than by geographical distance, indicating that populations have diverged adaptatively. We tested for adaptive divergence in Iberian crossbills using bill and body size measurements of 6082 crossbills from 27 sites, each consisting of a dominant or single pine (Pinus) of four possible species. Crossbills using different pines differed significantly in body size and bill size and shape. There was no correlation between geographical and morphological distance among sampling sites, consistent with the hypothesis that the morphological divergence of Iberian crossbills is shaped by their ecological differences (foraging on alternative conifers) rather than geographical distance. However, for unknown reasons, Common Crossbills foraging on Pinus sylvestris in Iberia have on average much smaller bills than Parrot Crossbills Loxia pytyopsittacus feeding on the same pine species in northern Europe. The extent to which crossbills specialize on Iberian P. sylvestris remains to be established. Specialization on conifers with overlapping geographical distributions may be facilitated by matching habitat choice of crossbills as a function of their local intake rates.     

Chromosome counts in the genus Festuca section Eskia (Poaceae) in the Iberian Peninsula

Chromosome numbers of taxa of Festuca L. section Eskia Willk. in the Iberian Peninsula are given. The levels of ploidy for five taxa are confirmed. Idiograms and karyotypic formulae of the five taxa are presented for the first time. Two levels of ploidy occur in this section: diploid and tetraploid. One taxon, Festuca elegans ssp. merinoi is tetraploid and two other taxa have diploid and tetraploid populations. The remaining two taxa are solely diploid.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 331–337.