Molecular phylogeny of Cotesia Cameron, 1891 (Insecta: Hymenoptera: Braconidae: Microgastrinae) parasitoids associated with Melitaeini butterflies (Insecta: Lepidoptera: Nymphalidae: Melitaeini)

Phylogenetic relationships among Cotesia Cameron (Braconidae) species parasitising Melitaeini butterflies were examined using DNA sequence data (mitochondrial cytochrome oxidase subunit I and NADH1 dehydrogenase genes, nuclear ribosomal DNA internal transcribed spacer region) as well as 12 microsatellite loci. Molecular data were available from ostensibly six species of Cotesia from 16 host butterfly species in Europe, Asia, and North America. Analysis of the combined sequence data using both maximum parsimony and maximum likelihood revealed two distinct Cotesia clades. In one clade (C. acuminata (Reinhard); C. bignellii (Marshall)) host ranges are apparently narrow and, although Euphydryas (s. lato) is well-utilised, permeation of Melitaea (s. lato) has been slight. In the other clade (C. melitaearum (Wilkinson); C. lycophron (Nixon); C. cynthiae (Nixon)) host utilization across the Melitaeini as a whole is more extensive and the data are consistent with more recent, or active, speciation processes. Neighbour-joining trees calculated separately for the two main clades based on chord distance (DCE) of microsatellite allele frequencies were consistent with phylogenetic trees obtained from the sequence data. Our analysis strongly suggests the presence of several additional, previously unrecognised, Cotesia species parasitising this group of butterflies.     

Does plasticity drive speciation? Host‐plant shifts and diversification in nymphaline butterflies (Lepidoptera: Nymphalidae) during the tertiary

How and why the great diversity of phytophagous insects has evolved is not clear but, if the explanation is the diversity of plants as a resource, colonizations of novel plant taxa can be expected to be associated with higher net speciation rates. In the present study, we make use of recent advances in plant and butterfly systematics to trace the evolution of host-plant utilization in the butterfly subfamily Nymphalinae (tribes Nymphalini, Melitaeini, and the probably paraphyletic 'Kallimini'). A clear historical pattern emerges, with an ancestral host-plant theme of 'urticalean rosids' and two major colonizations of novel distantly-related plant clades. The asterid order Lamiales was colonized by an ancestor of 'Kallimini' + Melitaeini and the family Asteraceae in Asterales was later colonized by Melitaeini butterflies. These colonization events appear to have been followed by increases in the rate of net butterfly diversification. Two not mutually exclusive scenarios to explain such patterns have been suggested: (1) adaptive radiation due to release from competition following host-plant shifts or (2) higher rates of net speciation during a relatively long-lasting potentially polyphagous (plastic) state. In support of the 'plasticity scenario', phylogenetic traces of a long-lasting stage with some potential to feed on more than one host-plant clade can still be seen, despite the ancient age of the colonizations. When angiosperm communities changed after the K/T boundary due to extinctions and subsequent diversification, herbivore taxa that could occupy several alternative niches may have had the greatest opportunity to diversify in turn.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 115–130.     

Relationships among non-native plants, diversity of plants and butterflies, and adequacy of spatial sampling

Non-native invasive species are altering ecosystems in undesirable ways, often leading to biotic homogenization and rapid reduction of evolutionary potential. However, lack of money and time hampers attempts to monitor the outcome of restoration efforts. Hence, it is useful to determine whether relatively limited sampling can provide valid inferences about biological responses to pattern-based and process-based variables that are affected by restoration actions. In the Mojave Desert, invasion of salt-cedar ( Tamarix ramosissima ) has altered vegetational communities and some measures of faunal diversity. We tested whether six vegetation-based predictor variables affected species richness of butterflies in the Muddy River drainage (Nevada, USA). We also explored whether similar conclusions about relationships between vegetation and butterflies could have been obtained by using data from a subset of the 85 locations included in the study. We found that the effect of non-native plants on species richness of butterflies was negligible. Availability of nectar had the greatest independent explanatory power on species richness of butterflies, followed by species richness of plants. In comparison with the full data set, subsamples including 10, 25 and 50% of sites yielded similar conclusions. Our results suggest that relatively limited data sets may allow us to draw reliable inferences for adaptive management in the context of ecological restoration and rehabilitation.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 157–166.     

Trophic differentiation in the phylogenetically young Cyprinodon species flock (Cyprinodontidae, Teleostei) from Laguna Chichancanab (Mexico)

Analysis of the gut contents of six syntopic Cyprinodon species from Laguna Chichancanab, Mexico, shows that the supposed basal form, C. beltrani , feeds on detritus and has the longest gut. All other species have significantly shorter guts. Whereas C. simus exclusively ingests detritus despite short gut length, all other species additionally rely on different benthic invertebrates. Each species has developed specific preferences: C. labiosus (amphipods), C. verecundus (bivalves), C. maya (ostracods and gastropods) and C. esconditus (ostracods and chironomid larvae). However, all of them ingest a minimum of c. 40% detritus. According to Schoener's index, pairwise niche overlap varies from c. 40% to over 95% in the species flock. Contrary to this, there is almost no overlap between the members of the flock and the only other native fish species, Gambusia sexradiata , which feeds nearly exclusively on terrestrial arthropods dropping on the water surface. Recently, the cichlid genus Oreochromis and the characid Astyanax fasciatus gained access to the lake. The cichlids show moderate to high niche overlap, whereas in A. fasciatus it is low. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 125–134.     

Food intake of fruit-feeding butterflies: evidence for adaptive variation in proboscis morphology

Adult butterflies feed from a variety of substrates and have appropriate adaptations. We examined proboscis morphology in a community of fruit-feeding butterflies (Nymphalidae) in a tropical forest in Uganda. These data were supplemented with behavioural observations and measurements of intake rate on natural and artificial substrates. We found no sexual dimorphism in proboscis morphology even though puddling behaviour is usually performed by males. Two main feeding techniques could be distinguished on the basis of behaviour and morphology: the piercing technique, typically found in Charaxinae, and the sweeping technique employed by both Nymphalinae and Satyrinae. These techniques, distinguished in previous studies, are described in more detail and their relative efficiencies are discussed in the context of sexual dimorphism, food-choice and life history evolution.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 333–343.     

A phenetic analysis and revision of Guineo-Congolean rain forest taxa of Chlorophytum (Anthericaceae)

The taxonomy of rain forest species of the genus Chlorophytum in central and west Africa has to date been difficult to resolve. Plants collected in the Guineo-Congolean plant geographical region deposited in several herbaria formed the basis for a morphometric analysis using Principal Component Analyses and including 16 characters. In total, 166 individuals were included from 461 herbarium collections, from which 57 seeds were scanned to investigate the surface structures. Eight species are recognized, including one with three subspecies and another with three varieties. Four new combinations are proposed ( Chlorophytum comosum (Thunb.) Jacques var. bipindense (Engl. & K. Krause) A.D. Poulsen & Nordal, C. comosum (Thunb.) Jacques var. sparsiflorum (Baker) A.D. Poulsen & Nordal, C. lancifolium Baker ssp. cordatum (Engl.) A.D. Poulsen & Nordal, and C. lancifolium Baker ssp. togoense (Engl.) A.D. Poulsen & Nordal), ten new synonyms are introduced and three lectotypes are designated.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 1–20.     

On the confusion surrounding Pareledone charcoti (Joubin, 1905) (Cephalopoda: Octopodidae): endemic radiation in the Southern Ocean

Until recently, all papillated specimens of Pareledone were ascribed to the species Pareledone charcoti (Joubin, 1905), of which P. aurorae (Berry, 1917) was considered a junior synonym. Re-examination of the papillated type material of Pareledone, coupled with extensive fishing over several years off the Antarctic Peninsula, has led to a revision of this position. Seven new species of papillated Pareledone are identified from the Antarctic Peninsula region. They are identified by subtle taxonomic characters, such as the morphology and placement of their papillae, although traditional indices often fail to separate the species. Whilst apparently sympatric, there is some evidence of niche separation of these species with respect to depth. A key is provided for their identification.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 75−108.     

Pinus mugo (Pinaceae) in the Abruzzi Mountains: high morphological variation in isolated populations

Thirty-three individuals of Pinus mugo from the Abruzzian Apennines (Italy), the most isolated population of the species, were examined biometrically to verify the range of trait variation and compared with six other populations sampled in the north-eastern part of the species range. Ten needles and ten cones from each individual were measured and quantified on the basis of 31 traits. The interpopulation comparison revealed a close affinity of the Abruzzian population of P. mugo to three of the six other populations. The other three north-eastern populations, one from the Tatra Mountains and two from the Chornokhora Mountains, appeared to be different. This suggests that these populations originated from other Pleistocene centres of occurrence. The intrapopulation variation of the population from the Abruzzian Apennines is comparable with those of all six populations of P. mugo from the Sudety and Carpathian Mountains. These findings support the hypothesis that isolated but locally abundant plant populations do not necessarily show a reduction in variability resulting from isolation.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 309–316.     

Cuticular lipid diversification in Lasiommata megera and Lasiommata paramegaera: the influence of species, sex, and population (Lepidoptera: Nymphalidae)

Numerous studies have investigated the presence and the effectiveness of volatile pheromones in Lepidoptera. Conversely, very few studies have focused on the composition and the perception of the relatively low volatile components of cuticular mixtures. Yet, cuticular lipids are implied in the recognition processes of several solitary and social insects. In the present study, the cuticular signatures of the satyrid butterflies Lasiommata megera and Lasiommata paramegaera were examined by gas chromatography. General linear model and discriminant analyses on chemical data clearly revealed large differences between sexes, which showed the same diversification pattern in both species. Moreover, a strong diversification between the two species was found, as were differences among populations. These results represent a first step in demonstrating the communicative function of cuticular compounds in the L. megera / paramegaera complex. Moreover, the discrimination among different species and populations on the basis of cuticular mixtures could represent a platform for studying chemotaxonomy and chemical biogeography in butterflies, as already found in several other insect groups.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 703–710.     

Phylogenetic relationships and historical biogeography of tribes and genera in the subfamily Nymphalinae (Lepidoptera: Nymphalidae)

We infer for the first time the phylogenetic relationships of genera and tribes in the ecologically and evolutionarily well‐studied subfamily Nymphalinae using DNA sequence data from three genes: 1450 bp of cytochrome oxidase subunit I (COI) (in the mitochondrial genome), 1077 bp of elongation factor 1‐alpha (EF1‐α) and 400–403 bp of wingless (both in the nuclear genome). We explore the influence of each gene region on the support given to each node of the most parsimonious tree derived from a combined analysis of all three genes using Partitioned Bremer Support. We also explore the influence of assuming equal weights for all characters in the combined analysis by investigating the stability of clades to different transition/transversion weighting schemes. We find many strongly supported and stable clades in the Nymphalinae. We are also able to identify ‘rogue’ taxa whose positions are weakly supported (the different gene regions are in conflict with each other) and unstable. Our main conclusions are: (1) the tribe Coeini as currently constituted is untenable, and Smyrna, Colobura and Tigridia are part of Nymphalini; (2) ‘Kallimini’ is paraphyletic with regard to Melitaeini and should be split into three tribes: Kallimini s.s., Junoniini and Victorinini; (3) Junoniini, Victorinini, Melitaeini and the newly circumscribed Nymphalini are strongly supported monophyletic groups, and (4) Precis and Junonia are not synonymous or even sister groups. The species Junonia coenia, a model system in developmental biology, clearly belongs in the genus Junonia. A dispersal‐vicariance analysis suggests that dispersal has had a major effect on the distributions of extant species, and three biotic regions are identified as being centres of diversification of three major clades: the Palaearctic for the Nymphalis‐group, the Afrotropics for Junoniini and the Nearctic for Melitaeini. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 227–251.     

The Rubiaceae of São Tomé e Príncipe (Gulf of Guinea): taxonomy and conservation

A synoptic treatment of the Rubiaceae of São Tomé e Príncipe (Gulf of Guinea) is presented based on herbarium collections kept at BM, COI, K and LISC. There are 36 genera and 64 species of Rubiaceae represented in the flora of São Tomé e Príncipe. Identification keys are provided for genera, species and infraspecific taxa. A new combination is made in Chassalia . Twenty-nine taxa are endemic to these islands, two of which are also endemic to Annobon. The conservation status of the endemic taxa is assessed and IUCN categories of threat are proposed for ten taxa. Nineteen taxa of Rubiaceae are considered under threat in São Tomé e Príncipe and ten are considered nearly threatened.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 85–114.     

Astragalus sect. Astragalus (Fabaceae) in Iran

Astragalus rahiminejadii , a new species endemic to Iran, is described and illustrated. This species, which belongs to section Astragalus , is confined to the western part of Iran (Prov. Kermanshah) and is known only from a single population. Seed testa morphology as viewed under the scanning electron microscope is discussed and photomicrographs are provided. In addition, the geographical distribution and ecology of the species belonging to this section are discussed.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 363–368.     

Age-related changes in thoracic mass: possible reallocation of resources to reproduction in butterflies

In nectar-feeding butterflies, reproductive potential is usually thought to depend on the size of the reproductive reserves in the abdomen, the adult food quality and, for females, the amount of resources received in the spermatophores at mating. Recent findings show that thorax mass and nitrogen content decrease with age in some butterfly species, and that thorax resources may be used for reproduction in the butterfly Pieris napi , just as in some other insects. In order to determine whether this is a general pattern and ascertain how it relates to the investment of resources in reproduction we studied the dynamics of thorax and abdomen mass changes in 11 Swedish butterfly species. By regressing thorax and abdomen mass on age of field-collected specimens, we show that loss of mass from both the thorax and the abdomen is a common phenomenon among nectar-feeding temperate zone butterflies under natural conditions. We argue that our results indicate that resources from flight muscles can be reallocated to reproduction by these butterflies, thus increasing their reproductive potential. Within species, females use proportionately more resources from the thorax than do males, as expected from the difference in investment of resources in reproduction. Among males we expect species with a higher reproductive investment to have a larger decrease in thorax and abdomen mass, and our data indicate that this is the case. Looking at the change in relative thorax mass, our results suggest that the use of resources from the thorax does not affect flight performance negatively, something that could constrain the use of muscle resources.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 363–380.     

A new species of Achetaria (Plantaginaceae) from south-eastern Brazil

A new species, Achetaria latifolia V.C.Souza, is described and illustrated. It is characterized by its broadly ovate leaves, indument of stems and leaves of noncapitate hairs, and pubescent capsule. Achetaria latifolia is known only in the coastal dune-area of Cabo Frio region, Rio de Janeiro State, Brazil.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 73–75.     

A new species of Ornithogalum (Liliaceae) from South Anatolia, Turkey

Ornithogalum chetikianum Uysal, Ertugrul & Dural sp. nov. (Liliaceae) is described and illustrated from Anatolia, Turkey. The species grows on alpine steppe and stony places in Bozkır (C4 Konya province) in South Anatolia. It is closely related to O. nutans L. and O. demirizianum Malyer & M.Koyuncu, an endemic confined to South Anatolia. Diagnostic morphological characters are discussed. Notes are also presented on the ecology, biogeography and conservation status of the new species.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 501–504.     

Comparative morphology of the leaf epidermis in Schisandra (Schisandraceae)

The leaf epidermis of 127 samples representing ten species within Schisandra Michaux. and one species of the related genus, Kadsura Kaempf. ex Juss., was investigated using light and scanning electron microscopy. Many characters of the leaf epidermis in Schisandra , such as pattern of epidermal cells, type of stomata, shape of guard cell pairs and cuticular ornamentation, are usually constant within species and thus make good characters for studying the relationship between and within genera. A new character, rim number, of the outer stomatal rim in the genus is introduced. It is shown that double outer stomatal rims occur only in evergreen Schisandra species, whereas a single rim occurs in deciduous species. This character supports the classification of Schisandra into two subgenera based on habit and androecial organs. This classification is also supported by additional morphological and molecular taxonomic characters. Kadsura coccinea (Lem.) Smith A. C. is the most primitive taxon in the related genus Kadsura . The outer stomatal rim of this species also has double rims. Combined with morphological and molecular evidence, this suggests that Schisandra and Kadsura are closely related and may share a recent common ancestor.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 39–56.     

A new species of Centaurea L. (Cardueae: Asteraceae) from Turkey

A new species of Centaurea L., C. kizildaghensis (Cardueae: Asteraceae), is described and illustrated from south Anatolia. It is restricted to the serpentine rocky slopes of Kızıl Dağ (C3 Konya–Derebucak, Çamlık), and is closely related to C. pseudokotschyi Wagenitz and C. drabifolioides Hub.-Mor., from which it differs in stem, leaf, and phyllary appendage characters. The diagnostic morphological characters are discussed. A new key is produced for Group G species for the flora of Turkey.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 61–66.