Conservation of Flower-arthropod Associations in Remnant African Grassland Corridors in an Afforested Pine Mosaic

While there has been much debate on the role of landscape corridors or linkages for animal movement and residency, there is little information on their role in maintaining ecological associations. We assess here the arthropod assemblages on flowers, seed heads and leaves of indigenous plants in a network of remnant grassland corridors across an alien pine-planted landscape in southern Africa. The results showed that the arthropod-plant (flowers, seed heads and leaves) associations remained, and were mostly quantitatively similar throughout the network of corridors. Narrow corridors (20–100 m wide) deep within the plantation-grassland mosaic still had high biodiversity value. The most threatening factor to flower-arthropod associations was disturbance, particularly cattle grazing. That the host plants were actually present was, in itself, an indicator that disturbance nevertheless was low. The associations are only lost when the plants are lost, and not when the plants are present but under some disturbance pressure. Although these corridors, which amount to about a third of the total area, mitigate the effects of pine afforestation, they can also be conduits for heavy human impact. This means that these corridors, even narrow and deep ones, are only of maximum value when there is minimal anthropogenic impact.     

Out-of-Africa again: A phylogenetic hypothesis of the genus Charaxes (Lepidoptera: Nymphalidae) based on five gene regions

Despite the long popularity of Charaxes among collectors and researchers, their evolutionary history is largely unknown. The current and accepted species groupings and relationships within the genus are based exclusively on adult morphology and life histories. Here, we examine the monophyly and evolutionary affinities of the species-groups within the genus Charaxes and explore how they relate to members of their closest genera (Euxanthe, Polyura and Palla) using 4167 bp of sequence data from five (1 mitochondrial and 4 nuclear) gene regions. Within the proposed phylogenetic framework, we estimate ages of divergence within the genus and also reconstruct their historical biogeography. We included representatives of all known species-groups in Africa and Asia, all known species of Euxanthe and Palla and two exemplar species of Polyura. We found the genus Charaxes to be a paraphyletic group with regard to the genera Polyura and Euxanthe, contrary to the earlier assumption of monophyly. We found that 13 out of 16 morphologically defined species-groups with more than one species were strongly supported monophyletic clades. Charaxes nichetes is the sister group to all the other Charaxes. Polyura grouped with the Zoolina and Pleione species-groups as a well-supported clade, and Euxanthe grouped with the Lycurgus species-group. Our results indicated that the common ancestor of Charaxes diverged from the common ancestor of Palla in the mid Eocene (45 million years ago) in (Central) Africa and began diversifying to its extant members 15 million years later. Most of the major diversifications within the genus occurred between the late Oligocene and Miocene when the global climates were putatively undergoing drastic fluctuations. A considerable number of extant species diverged from sister species during the Pliocene. A dispersal–vicariance analysis suggests that many dispersal rather than vicariance events resulted in the distribution of the extant species. The genus Polyura and the Indo-Australian Charaxes are most likely the results of three independent colonizations of Asia by African Charaxes in the Miocene. We synonymize the genera Polyura (syn. nov.) and Euxanthe (syn. nov.) with Charaxes, with the currently circumscribed Charaxes subdivided into five subgenera to reflect its phylogeny.     

Captive rearing of Puget blue butterflies (Icaricia icarioides blackmorei) and implications for conservation

The use of captive rearing to promote recovery of endangered butterflies has substantially increased over the last decade. These programs have the potential to play a significant role in butterfly population recovery, but the effects of captive conditions are poorly understood and rarely are traits of captive individuals assessed relative to traits in their founding populations. To develop rearing protocols and investigate possible effects of captive conditions, we reared Puget blue butterflies (Icaricia icarioides blackmorei), a subspecies closely related to the endangered Fender’s blue (I. i. fenderi) which is limited to Oregon, USA. We reared individuals from two wild populations in Washington, USA to investigate two approaches for egg collection (collect eggs in the wild vs. collect eggs from adult females which were brought to a greenhouse for oviposition) and three diapause environments (in indoor facilities at two independent locations vs. outdoors in enclosures). Survival from egg to adult was similar across all captive groups which survived past diapause and was less than 10%. Captive reared individuals were lighter and had smaller wings and shorter body lengths than their founding populations for both sites. Based on our findings, we recommend that rearing programs compare characteristics of reared individuals to individuals from the founding population to quantify possible effects of captive conditions, diapause individuals in natural environments, and for programs with survival rates similar to rates in the wild, consider alternatives to augment declining populations and reintroduce historic ones.     

Movement of two grassland butterflies in the same habitat network: the role of adult resources and size of the study area

Abstract. 1.  Movement patterns of two butterfly species (meadow brown Maniola jurtina L. and scarce copper Lycaenae virgaureae L.) were studied in a 172 ha area within a landscape with a high percentage of suitable habitats for mark–release–recapture experiments.
2.  Adult resource density, but not patch size or larval food plant abundance, influenced the numbers and the fractions of residents, emigrants, and immigrants.
3.  Differences between species were observed in movement frequency and maximum distances moved but not in mean distances moved.
4.  The scarce copper showed much greater movement ability than expected from the results of published studies. This is believed to be a result of the comparatively large size of the study area and the high cover of suitable habitat (>50%).
5.  The mean and maximum distances travelled by butterflies reflected differences in the size of the study area.     

Assignment and evaluation of ground beetle (Coleoptera: Carabidae) assemblages to sites on different scales in a grassland landscape

Carabid beetles and environmental parameters were investigated in 52 grassland sites with three replicate pitfall traps in each site and in the valley of the River Eider in Schleswig–Holstein (northern Germany) with 61 pitfall traps. Environmental parameters included water content of soil, sand content, organic matter content and pH. Ground beetle assemblages were derived by detrended correspondence analyses (DCA) and characterised by the specific environmental conditions as means for each assemblage. On the regional scale including all investigated sites of Schleswig–Holstein, five assemblages were differentiated. On the local scale including the investigated sites in the valley of the River Eider, three assemblages were found corresponding well with those found on the regional scale. Environmental conditions at the sites of the five assemblages were correlated with land use data, soil types, and water level stages provided by three maps of a geographic information system (GIS). The GIS maps were combined to develop smaller areas with land use, soil type and water level stage information. The characteristic environmental conditions were assigned to each area to derive the spatial distribution of the five ground beetle assemblages. Spatial prediction was correct for 65% of investigated sites. The potential area of each assemblage was estimated for the valley. The different grassland areas were evaluated as potential habitats for ground beetle species comparing total species richness with the regional species richness of each assemblage. The comparison shows that species richness in the evaluated assemblages is relatively low compared to the regional potential.     

An analysis of the phenotypic effects of certain colour pattern genes in Heliconius (Lepidoptera: Nymphalidae)

The colour patterns of Heliconius butterflies are built up from an array of serially homologous pattern elements known as the nymphalid groundplan. An analysis of the phenotypic effects of ten genetic loci from H. melpomene and H. cydno reveals that each alters the expression either of a single element of the groundplan or of an entire row of serially homologous elements. Five of the ten loci affect the size (or presence/absence) of specific pattern elements, two affect the colour in which a pattern element is expressed, two affect pattern-inducing activity of the wing veins, and one appears to affect an overall threshold for pattern determination. Three of the ten loci have identical effects on homologues of the fore- and hindwing. We show that most of the apparently large and qualitative phenotypic effects of these genes can be readily explained by relatively small and quantitative changes in the dimensions or positions of specific pattern elements.     

Survival of refrigerated Tatochila butterflies (Lepidoptera: Pieridae) as an indicator of male nutrient investment in reproduction

Summary Tatochila butterflies show prolonged survival in cold (2° C) storage. Mating significantly reduces survival in males, suggesting that nutrients may be directed into spermatophores or maintenance under storage conditions; this provides a potential index of male nutrient investment in reproduction.     

Persistence of species in a fragmented urban landscape: the importance of dispersal ability and habitat availability for grassland butterflies

Some species cope with, and survive in, urban areas better than others.From a conservation viewpoint it is important to understand why some species arerare or are excluded in the urban landscape, in order that we might take actionto conserve and restore species. Two ecological factors that might explain thedistribution and abundance of butterfly species in the urban landscape aredispersal ability and the availability of suitable habitat. The influence ofthese factors was assessed by examining the distribution and genetic structureof four grassland butterfly species in the West Midlands conurbation, UK. Thefour species differ in their distribution and abundance, mobility and habitatspecificity. No significant fit to the isolation-by-distance model was found forany of the study species at this spatial scale. MeanF _ST values revealed a non-significant level ofpopulation structuring for two species, Pieris napi (L.)and Maniola jurtina (L.), but moderate and significantpopulation differentiation for Pyronia tithonus (L.) andCoenonympha pamphilus (L.). Results suggest that thesespecies are limited more by the availability of suitable habitat than by theirability to move among habitat patches. Conservation strategies for thesegrassland species should initially focus on the creation and appropriatemanagement of suitable habitat. More sedentary species that have already beenexcluded from the conurbation may require a more complex strategy for theirsuccessful restoration.     

A mitochondrial-DNA-based phylogeny for some evolutionary-genetic model species of Colias butterflies (Lepidoptera, Pieridae)

We study the phylogenetic relationships among some North American Colias ("sulfur") butterflies, using mitochondrial gene sequences (ribosomal RNA, cytochrome oxidase I+II) totaling about 20% of the mitochondrial genome. We find that (1) the lowland species complex shows a branching order different from earlier views; (2) several montane and northern taxa may be more distinct than in earlier views; (3) one morphologically conservative Holarctic assemblage, C. hecla, is differentiated at the molecular-genetic level into at least three taxa which occupy distinct positions in the phylogeny and are sisters to diverse other taxa. These conclusions, constituting phylogenetic hypotheses, are supported by parsimony, maximum-likelihood, and Bayesian reconstruction algorithms. They are tested formally, by interior branch tests and paired-site tests, against alternative hypotheses derived from conventional species and subspecies naming combinations. In all cases our hypotheses are supported by these tests and the conventional alternatives are rejected. The "barcoding" subset of cytochrome oxidase I sequence identifies only some of the taxa supported by our full data set. Comparison of genetic divergence values among Colias taxa with those among related Pierid butterflies suggests that species radiations within Colias are comparatively younger. This emerging Colias phylogeny facilitates comparisons of genetic polymorphism and other adaptive mechanisms among taxa, thereby connecting micro- and macro-evolutionary processes.     

Importance of historical factors on species richness and composition of butterfly assemblages (Lepidoptera: Rhopalocera) in a northern Iberian mountain range

ABSTRACT. The butterfly assemblages of three vegetation types (pasture, heathland and rocky outcrop with Genista spp.) and two geographical orientations (a northern slope and a sheltered gorge) were studied along an altitudinal gradient in the Picos de Europa in northern Spain. In order to study the effects or historical climatic changes on current butterfly assemblages, it was examined if changes in richness and faunal composition of assemblages were joined with changes in their biogeographical composition. The three vegetation types showed characteristic butterfly assemblages, but no difference in species richness and biogeographical composition was found. Species richness in the sheltered gorge was higher than in the northern slope. Faunal composition also varied between orientations and there were more widespread and Mediterranean species in the sheltered gorge than in the northern slope. Species richness declined with increasing altitude, though a midaltitudinal peak was observed. There was a faunal segregation between high and low localities. Species richness of widespread and Mediterranean butterflies decreased as altitude increased, whereas the reverse was true for montane species. Moreover, biogeographical elements differed in their climatic tolerances measured as altitudinal ranges. Therefore, changes in richness and composition of butterfly assemblages between both slopes and along the altitudinal gradient were joined in part with variation in their biogeographical composition. These results suggest that current species richness and composition of butterfly assemblages in the Picos de Europa might be the consequence of differential colonization of refuges during the past climatic changes.     

The proximate control of pupal color in swallowtail butterflies: implications for the evolution of environmentally cued pupal color in butterflies (Lepidoptera: Papilionidae)

The environmentally cued production of cryptic green/yellow or brown/melanized pupae is widespread in butterflies, occurring in the Nymphalidae, Pieridae, and the Papilionidae subfamily Papilioninae. The dimorphism is controlled by the hormone pupal melanization reducing factor (PMRF). In the nymphalid Inachis io dibutryl cAMP mimics PMRF, and inhibits pupal melanization. However, in the papilionid Papilio polyxenes PMRF stimulates browning, suggesting that the control of pupal color by PMRF has evolved independently in the swallowtail and nymphalid-pierid lineages. We examined this hypothesis by using ligatures to prevent hormone release in five species representing three Papilioninae tribes. One species, Papilio glaucus, produces only brown pupae. Ligatures resulted in green cuticle posterior to the ligature in all five swallowtail species, including P. glaucus, suggesting that the mode of action of PMRF is the same in the three tribes. We also found that in P. polyxenes injections of dibutryl cAMP into prepupal larvae mimic the effect of PMRF, by causing dose-dependent pupal browning. Our results support the hypothesis that the control of pupal color by PMRF has evolved independently in the two lineages. The observation that green pupal color can be induced in P. glaucus by ligature indicates that environmentally cued pupal color could evolve by facultative inhibition of PMRF release.     

Correspondence analysis of shell morphology in the African freshwater snail Biomphalaria pfeifferi (Kraus 1848) (Pulmonata: Gastropoda)

A correspondence analysis of shell measurements taken from 521 widely dispersed specimens of the African aquatic pulmonate snail Biomphalaria pfeifferi suggests the existence of eight morphological groups. These groups appear to relate to either ecophysiological factors or to factors associated with the stability of the freshwater system rather than to aspects of geographic distribution and genetic isolation.     

Parasitism of orangestriped oakworm (Lepidoptera: Saturniidae) eggs in the urban landscape

Orangestriped oakworm, Anisota senatoria (J. E. Smith), has caused widespread defoliation of oak trees in the urban landscape of southeastern Virginia since 1985. Egg masses were collected from 1988 to 1990 to determine the impact of native egg parasites on A. senatoria populations. The most abundant egg parasite was Aprostocetus new sp. and mean egg mass parasitism was 24.6%. The eupelmid Anastatus hirtus (Ashmead), a new host record, parasitized a mean of 11.7% of A. senatoria egg masses. The encyrtid Ooencytrus sp., a new host record, had a mean egg mass parasitism of 0.09%. Inundative releases of Trichogramma minutum (Riley) in 1989 and 1990 did not increase parasitism rates and mean egg mass parasitism was 2.3%. Parasitism of first generation A. senatoria egg masses was higher compared with second generation. The four egg parasites collected in this study parasitized 30% of A. senatoria egg masses and within egg mass parasitism was 7.9%. These relatively low parasitism rates may partially explain the presence of consistently high A. senatoria populations in southeastern Virginia.     

Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera: Nymphalidae)

The historical biogeography of insects in South America is largely unknown, as dated phylogenies have not been available for most groups. We have studied the phylogenetic relationships and historical biogeography of a subtribe of butterflies, Phyciodina in the family Nymphalidae, based on one mitochondrial gene (COI) and two nuclear gene regions (EF-1alpha and wingless). The subtribe comprises 89 species mainly found in tropical South America, with a few species in North America and the Greater Antilles. We find that the enigmatic genus Antillea is sister to the rest of Phyciodina, and suggest that it should be included in the subtribe. Several genera are found to be polyphyletic or nested within another genus, and are proposed to be synonymised. These are Dagon, Castilia, Telenassa and Janatella, which we propose should be synonymised with Eresia. Brazilian "Ortilia" form an independent lineage and require a new genus name. The diversification of Phyciodina has probably taken place over the past about 34 MYA. The ancestral phyciodine colonised South America from North America through a possible landspan that connected the Greater Antilles to South America about 34MYA. A vicariance event left the ancestral Antillea on the Greater Antilles, while the ancestral 0e on South America colonised the Guyanan Shield and soon after the Brazilian Shield. We hypothesise that the Brazilian Shield was an important area for the diversification of Phyciodina. From there, the ancestor of Anthanassa, Eresia and Tegosa colonised NW South America, where especially Eresia diversified in concert with the rising of the Andes beginning about 20 MYA. Central America was colonised from NW South America about 15 MYA by the ancestors of Anthanassa and Phyciodes. Our study is the first to use a dated phylogeny to study the historical biogeography of a group of South American species of butterflies.     

A new methodology for compiling national Red Lists applied to butterflies (Lepidoptera, Rhopalocera) in Flanders (N-Belgium) and the Netherlands

The compilation of the Red Lists of butterflies in Flanders and the Netherlands was based on two criteria: a trend criterion (degree of decline) and a rarity criterion (actual distribution area). However, due to the large difference in mapping intensity in the two compared periods, a straightforward comparison of the number of grid cells in which each species was recorded, appeared inappropriate. To correct for mapping intensity we used reference species that are homogeneously distributed over the country, that have always been fairly common and that did not fluctuate in abundance too much during this century. For all resident species a relative presence in two compared periods was calculated, using the average number of grid cells in which these reference species were recorded as a correction factor. The use of a standardized method and well-defined quantitative criteria makes national Red Lists more objective and easier to re-evaluate in the future and facilitates the comparison of Red Lists among countries and among different organisms. The technique applied to correct for mapping intensity could be useful to other organisms when there is a large difference in mapping intensity between two periods.     

Evaluation of community indices in seasonal assemblages of butterflies (Lepidoptera) at different frequency of transect count

Community indices were evaluated at different frequency of transect counts of butterflies by randomization. Randomized samples were obtained by re-arranging 1, 3, 6, 12 and 18 counts from 18 counts at a study site (Hiraoka), and 1, 2, 3, 5 and 15 from 15 at another site (Narukawa), in 1994. Morisita's new estimate of species richness, QM gave a good estimate when the number of counts was five or more. QM estimated from the sum of both assemblages (58 species) was 70.7, while 73 species were recorded at least once from 1990 to 1994. The Octave method always resulted in an underestimation. Diversity indices, Shannon's H, Simpson's D and , Hurlbert's E(Sn), Itow's b, and Morisita's H gave good estimates when the number of counts was five or more. Temporariness of univoltine species in flight period made a bias in the estimates. We concluded that the count is needed at least six times (once a month) and the estimates were improved by separating the multivoltine species from the univoltine species. Intensive counting during the flight period of the univoltine species was recommended.     

The evolution of ecologically important characters in Heliconius butterflies (Lepidoptera: Nymphalidae): a cladistic review

The biology of Heliconius butterflies has provided a rich source of data to test theories of ecological genetics, coevolution and community ecology. Many putatively adaptive characters have been discussed with reference to a phylogenetic hypothesis based on a variety of morphological and life-history traits interpreted from an evolutionary taxonomic perspective. Here, alternate interpretations of characters on the traditional tree and a more recent mitochondrial DNA cladogram with a substantially different topology are compared and contrasted. It is shown that many characters ostensibly providing support for the traditional phylogenetic hypothesis are almost equally parsimoniously distributed and in some cases more parsimoniously distributed on the mtDNA tree than on the tree inferred from those characters. Discussion of alternate evolutionary scenarios based on the mDNA-based topology is presented for pupal mating, pollen feeding, foodplant coevolution, and other ecologically significant features.     

Molecular phylogeny of African hinge‐back tortoises (Kinixys): implications for phylogeography and taxonomy (Testudines: Testudinidae)

We examine the phylogeography, phylogeny and taxonomy of hinge‐back tortoises using a comprehensive sampling of all currently recognized Kinixys species and subspecies and sequence data of three mitochondrial DNA fragments (2273 bp: 12S rRNA, ND4 + adjacent DNA coding for tRNAs, cytb) and three nuclear loci (2569 bp: C‐mos, ODC, R35). Combined and individual analyses of the two data sets using Bayesian and Maximum Likelihood methods suggest that the savannah species of Kinixys are paraphyletic with respect to the rainforest species K. homeana and K. erosa, and that the rainforest species may be derived from a savannah‐living ancestor. The previously recognized savannah species K. belliana was a conglomerate of three deeply divergent clades that we treat here as distinct species. We restrict the name K. belliana (Gray, 1830) to hinge‐back tortoises ranging from Angola to Burundi, while five‐clawed hinge‐back tortoises from the northernmost part of the formerly recognized range of K. belliana, together with four‐clawed tortoises from West Africa, are assigned to the species K. nogueyi (Lataste, 1886). These two species are allied to K. spekii, whereas Southeast African and Malagasy hinge‐back tortoises formerly lumped together with K. belliana represent the distinct species K. zombensis Hewitt, 1931, which is sister to K. lobatsiana. The latter two species together constitute the sister group of the rainforest species K. homeana and K. erosa. Mitochondrial data suggest that K. natalensis has a basal phylogenetic position in a clade embracing K. belliana sensu stricto, K. nogueyi and K. spekii, while nuclear data and the two data sets combined favour a sister group relationship of K. natalensis to all other hinge‐back tortoises. Phylogeographic structure is present in all wide‐ranging species and correlates in K. homeana and K. erosa with the Dahomey Gap and former rainforest refugia. The Malagasy population of K. zombensis is weakly differentiated from its South African conspecifics and further sampling is needed to determine whether there is support for the subspecific distinctness of Malagasy tortoises.     

Phylogeographic pattern and regional evolutionary history of the maize stalk borer Busseola fusca (Fuller) (Lepidoptera: Noctuidae) in sub-Saharan Africa

Busseola fusca (Fuller) (Lepidoptera: Noctuidae) is one of the major cereal pests in sub-Saharan Africa. Previous phylogeographic investigations on samples collected in Kenya, Cameroon and West-Africa showed the presence of three main clades (W, KI, KII) originated from populations isolated in West and East Africa around one million years ago. Demographic and phylogenetic analyses suggested that this event was followed by local demographic expansion and isolation by distance. These hypotheses were tested by a more comprehensive sampling across B. fusca’s geographic range in Africa. Comparisons of sequences of partial mitochondrial DNA gene (cytochrome b) from 489 individuals of 98 localities in southern, central, eastern and western African countries confirmed the presence of the three main clades. Phylogenetic, F-statistics, demographic parameters and nested clade phylogeographic analyses confirmed that the clades experienced geographic and demographic expansion with isolation by distance after their isolation in three refuge areas. The geographic range of clade KII, already known from East to Central sub-Saharan Africa was extended to Southern Africa. Mismatch distribution analysis and the negative values of Tajima’s D index are consistent with a demographic expansion hypothesis for these three clades. Significant genetic differentiations were revealed at various hierarchical levels by analysis of molecular variance (AMOVA). Hypotheses about the geographic origin of the three main clades are detailed.     

Impact of an exotic parasitoid on Plutella xylostella (Lepidoptera: Plutellidae) population dynamics, damage and indigenous natural enemies in Kenya

Diadegma semiclausum (Hellén) (Hymenoptera: lchneumonidae), an exotic diamondback moth parasitoid, was released in two pilot areas (Werugha in Coast Region and Tharuni in Central Province) in Kenya. Fifteen month before release, observations on the diamondback moth, Plutella xylostella (Linnaeus), and local natural enemy population dynamics and pest damage were initiated in both areas and continued for three years after release. The P. xylostella population was bimodal with higher records during dry seasons. At Werugha, the peak population of P. xylostella was 16.8 per plant (October 2001); at Tharuni it was 12.8 (February 2002). Populations at Werugha declined from three months after release and decreased from 5.4 per plant (before release) to 0.8 (year 3 after release). Concurrently, average damage (1.9 to 1.5) (on a 0-5 scale), proportion of attacked plants (72 to 31%) and proportion of plants in damage group >2 (plants with head damage) decreased (21.4 to 5.3%), while total parasitism increased from 14.4 (before) to 52.5% (year 3 after release, 90% due to D. semiclausum). At Tharuni, D. semiclausum was only recovered 3 months after release. Average populations of P. xylostella declined from 5.9 per plant (before release) to 2.4 (year 3 after release) and damage scores from 2.3 to 1.7. The proportion of plants in damage group >2 declined from 39.7 to 4.5% while overall parasitism increased from 4.2 to 40.6% (98.3% by D. semiclausum). Four species of indigenous parasitoids (Diadegma mollipla (Holmgren), Oomyzus sokolowskii (Kurdjumov), Apanteles sp. and Itoplectis sp., all primary parasitoids) were almost completely displaced by D. semiclausum. Possible reasons for the different parasitoid development between the two release areas and the displacement of the indigenous species are discussed.