Diversity and evolutionary patterns of bacterial gut associates of corbiculate bees

The animal gut is a habitat for diverse communities of microorganisms (microbiota). Honeybees and bumblebees have recently been shown to harbour a distinct and species poor microbiota, which may confer protection against parasites. Here, we investigate diversity, host specificity and transmission mode of two of the most common, yet poorly known, gut bacteria of honeybees and bumblebees: Snodgrassella alvi (Betaproteobacteria) and Gilliamella apicola (Gammaproteobacteria). We analysed 16S rRNA gene sequences of these bacteria from diverse bee host species across most of the honeybee and bumblebee phylogenetic diversity from North America, Europe and Asia. These focal bacteria were present in 92% of bumblebee species and all honeybee species but were found to be absent in the two related corbiculate bee tribes, the stingless bees (Meliponini) and orchid bees (Euglossini). Both Snodgrassella alvi and Gilliamella apicola phylogenies show significant topological congruence with the phylogeny of their bee hosts, albeit with a considerable degree of putative host switches. Furthermore, we found that phylogenetic distances between Gilliamella apicola samples correlated with the geographical distance between sampling locations. This tentatively suggests that the environmental transmission rate, as set by geographical distance, affects the distribution of G. apicola infections. We show experimentally that both bacterial taxa can be vertically transmitted from the mother colony to daughter queens, and social contact with nest mates after emergence from the pupa greatly facilitates this transmission. Therefore, sociality may play an important role in vertical transmission and opens up the potential for co‐evolution or at least a close association of gut bacteria with their hosts.     

Patterns of host-plant choice in bees of the genus Chelostoma: the constraint hypothesis of host-range evolution in bees

To trace the evolution of host-plant choice in bees of the genus Chelostoma (Megachilidae), we assessed the host plants of 35 Palearctic, North American and Indomalayan species by microscopically analyzing the pollen loads of 634 females and reconstructed their phylogenetic history based on four genes and a morphological dataset, applying both parsimony and Bayesian methods. All species except two were found to be strict pollen specialists at the level of plant family or genus. These oligolectic species together exploit the flowers of eight different plant orders that are distributed among all major angiosperm lineages. Based on ancestral state reconstruction, we found that oligolecty is the ancestral state in Chelostoma and that the two pollen generalists evolved from oligolectic ancestors. The distinct pattern of host broadening in these two polylectic species, the highly conserved floral specializations within the different clades, the exploitation of unrelated hosts with a striking floral similarity as well as a recent report on larval performance on nonhost pollen in two Chelostoma species clearly suggest that floral host choice is physiologically or neurologically constrained in bees of the genus Chelostoma. Based on this finding, we propose a new hypothesis on the evolution of host range in bees.     

Phylogenetic relationships within Diadasia, a group of specialist bees

We estimated phylogenetic relationships among species of the bee genus Diadasia, a group of new world, specialist bees. We sequenced approximately 2 kb of the mitochondrial genes cytochrome oxidase subunit I and II and tRNA leucine and approximately 1 kb of the nuclear gene elongation factor 1-alpha for 24 North American Diadasia species, 4 South American species, and five outgroup genera. Parsimony analyses of the two data sets were highly congruent. A combined analysis produced a well-resolved phylogenetic hypothesis that supported the monophyly of Diadasia, but not that of traditional subgenera: Diadasia s. str. was paraphyletic in all analyses. With one exception, the North and South American species formed separate clades, supporting previous hypotheses that two lineages of Diadasia have dispersed from South to North America: a more recent dispersal of D. ochracea and an older dispersal of the ancestor to all other North American species. Different species of Diadasia specialize on pollen from at least five different plant families; the phylogeny presented here, along with known host affinities, indicates that host-switching has been rare.     

Genomic changes underlying host specialization in the bee gut symbiont Lactobacillus Firm5

Bacteria that engage in long‐standing associations with particular hosts are expected to evolve host‐specific adaptations that limit their capacity to thrive in other environments. Consistent with this, many gut symbionts seem to have a limited host range, based on community profiling and phylogenomics. However, few studies have experimentally investigated host specialization of gut symbionts and the underlying mechanisms have largely remained elusive. Here, we studied host specialization of a dominant gut symbiont of social bees, Lactobacillus Firm5. We show that Firm5 strains isolated from honey bees and bumble bees separate into deep‐branching host‐specific phylogenetic lineages. Despite their divergent evolution, colonization experiments show that bumble bee strains are capable of colonizing the honey bee gut. However, they were less successful than honey bee strains, and competition with honey bee strains completely abolished their colonization. In contrast, honey bee strains of divergent phylogenetic lineages were able to coexist within individual bees. This suggests that both host selection and interbacterial competition play important roles in host specialization. Using comparative genomics of 27 Firm5 isolates, we found that the genomes of honey bee strains harbour more carbohydrate‐related functions than bumble bee strains, possibly providing a competitive advantage in the honey bee gut. Remarkably, most of the genes encoding carbohydrate‐related functions were not conserved among the honey bee strains, which suggests that honey bees can support a metabolically more diverse community of Firm5 strains than bumble bees. These findings advance our understanding of the genomic changes underlying host specialization.     

FLORAL NECTAR SUGAR COMPOSITION AND POLLINATOR TYPE AMONG NEW WORLD GENERA IN TRIBE ANTIRRHINEAE (SCROPHULARIACEAE)

The floral nectar sugar compositions of 20 New World species from 10 genera and of five interspecific hybrids in tribe Antirrhineae have been analyzed using high-performance liquid chromatography. Species are pollinated by short-tongued bees, long-tongued bees, and hummingbirds. Ornithophily represents the derived condition in the tribe and has arisen independently in subtribes Maurandyinae and Gambeliinae. All nectars analyzed are sucrose-dominant or -rich, except for the hexose-rich nectar of Mohavea breviflora. Despite the predominance of sucrose, floral nectar sugars from species pollinated by different pollen vectors have characteristic constituents. Nectar sugars from flowers visited by hummingbirds average 76.2% sucrose and have compositions remarkably similar to hummingbird nectars analyzed in previous studies of unrelated species. Long-tongued bee nectars average 87% sucrose and differ from shorttongued bee nectars which have the lowest mean sucrose percentage (40.2%). The association of sugar constituent types and principal pollinators is concordant with previous data and supports hypotheses concerning pollinator preferences and the adaptive significance of certain nectar sugar compositions. Within this adaptive framework, phylogenetic constraint is also operative and may explain the predominance of sucrose in nectar sugars, similarities in sugar composition among hummingbird nectars in subtribes Maurandyinae and Gambeliinae, and the similarity of nectar from Galvezia leucantha (long-tongued bee-pollinated) to hummingbird-pollinated species also in subtribe Gambeliinae.     

Host plant preference and performance of the vine weevil Otiorhynchus sulcatus

Abstract 1 The relationship between reproductive performance and preference for potential host plants of the vine weevil is investigated, as shown in tests on contact (or feeding) preference, presented herein, and tests on olfactory preference, published elsewhere. 2 Assessment of reproductive performance shows that the host‐plant range of the adult vine weevil Otiorhynchus sulcatus in Europe is limited to one gymnosperm genus (Taxus sp.) and a broad range of angiosperm plants in two subclasses of the Dicotyledonae, namely Dilleniidae and Rosidae. The successful reproduction on very distantly related plant taxa suggests that the original weevil‐ and plant‐habitat has mediated the current host‐plant range of the vine weevil. 3 Contact‐preference tests with equally suitable hosts, such as Aronia, Fragaria, Euonymus and Taxus, and one less suitable host, Humulus, indicate a mismatch between contact preference and performance and, as far as olfactory preferences are known, these match neither the contact preferences nor the performance. This mismatch may arise because (i) host plant species offered do not occur in weevil habitat in Europe (e.g. Aronia and the cultivated Fragaria come from North America) and (ii) predation (or disease) risks differ among host plants, thereby altering effective reproductive performance. 4 With respect to performance on novel hosts (Thuja, Prunus) and bad hosts (Rhododendron), some between‐individual variation is found within a single population, suggesting that local populations harbour (possibly genetic) variation for adaptation to new hosts. How this variation is maintained in the face of strong selection pressures on local populations of flightless and thelytokous weevils, is an important question for understanding the broad host plant range in the vine weevil.     

Biogeographical origins and diversification of the exoneurine allodapine bees of Australia (Hymenoptera,Apidae)

Aim Early diversification of allodapine bees occurred in Africa c. 50 Ma. They are most abundant in sub‐Saharan Africa and Australia, and one of the oldest phylogenetic divergences in the tribe involves a split between an African + Malagasy clade and an Australian clade. The historical biogeographical scenario for this has been highly problematic, entailing an Eocene dispersal from Africa to Australia, followed by an unresolved, and apparently rapid, set of bifurcations leading to the Australian ‘exoneurine’ genera. Here we use an expanded taxon set of Australian species to explore the timing and historical biogeography of the exoneurine radiation. Location Australia, Africa, Madagascar. Methods One nuclear gene (F2 copy of elongation factor 1α) and two mitochondrial genes (cytochrome c oxidase subunit I and cytochrome b) were sequenced for 33 Australian exoneurine species from all five genera found on the continent, as well as for an additional 37 species from all non‐parasitic genera in the remainder of the tribe. We used Bayesian inference analyses to study phylogenetic topology and penalized likelihood analyses to infer key dates of divergence within the tribe. We also used lineage‐through‐time (LTT) analyses and Bayesian analyses to explore the tempo of radiations and biogeographical history of the exoneurines. Results Results from the phylogenetic analyses were congruent with previous studies, indicating a single colonization event c. 34 Ma, too late for Gondwanan vicariance models, and too early for a Laurasian dispersal route. In contrast to earlier studies, we show that this colonization event did not result in an ancient rapid radiation. However, LTT patterns indicated a rapid radiation of the temperate‐adapted genera Exoneura and Brevineura, but not of the xeric‐adapted genus Exoneurella, from 10 to 6 Ma. Main conclusions Our results indicate a trans‐oceanic dispersal event from Africa to Australia, most likely via Antarctica, with an accelerated diversification of temperate‐adapted lineages during the major Late Miocene event referred to as the ‘Hill Gap’. This is the first study to link radiations in Australian bee faunal elements to changing climate, and differs from many other plant and insect phylogenetic studies by showing increased radiation of temperate clades, rather than xeric clades, with increasing aridification of Australia.     

Resource assurance predicts specialist and generalist bee activity in drought

Many short-lived desert organisms remain in diapause during drought. Theoretically, the cues desert species use to continue diapause through drought should differ depending on the availability of critical resources, but the unpredictability and infrequent occurrence of climate extremes and reduced insect activity during such events make empirical tests of this prediction difficult. An intensive study of a diverse bee–plant community through a drought event found that bee specialists of a drought-sensitive host plant were absent in the drought year in contrast to generalist bees and to specialist bees of a drought-insensitive host plant. Different responses of bee species to drought indicate that the diapause cues used by bee species allow them to reliably predict host availability. Species composition of the bee community in drought shifted towards mostly generalist species. However, we predict that more frequent and extended drought, predicted by climate change models for southwest North America, will result in bee communities that are species-poor and dominated by specialist species, as found today in the most arid desert region of North America.     

Worldwide long‐distance dispersal favored by epizoochorous traits in the biogeographic history of Omphalodeae (Boraginaceae)

Biogeographic dispersal is supported by numerous phylogenetic results. In particular, transoceanic dispersal, rather than vicariance, is suggested for some plant lineages despite current long distances between America and Europe. However, few studies on the biogeographic history of plants have also studied the role of diaspore syndromes in long‐distance dispersal (LDD). Species of the tribe Omphalodeae (Boraginaceae) offer a suitable study system because the species have a wide variety of diaspore traits related to LDD and different lineages conform to patched worldwide distributions on three distant continents (Europe, America and New Zealand). Our aim is to reconstruct the biogeographical history of the Omphalodeae and to investigate the role of diaspore traits favoring LDD and current geographic distributions. To this end, a time‐calibrated phylogeny with 29 of 32 species described for Omphalodeae was reconstructed using biogeographical analyses (BioGeoBEARS, Lagrange) and models (DEC and DIVA) under different scenarios of land connectivity. Character‐state reconstruction (SIMMAP) and diversification rate estimations of the main lineages were also performed. The main result is that epizoochorous traits have been the ancestral state of LDD syndromes in most clades. An early diversification age of the tribe is inferred in the Western Mediterranean during late Oligocene. Colonization of the New World by Omphalodeae, followed by fast lineage differentiation, took place sometime in the Oligocene‐Miocene boundary, as already inferred for other angiosperm genera. In contrast, colonization of remote islands (New Zealand, Juan Fernández) occurred considerably later in the Miocene‐Pliocene boundary.     

Factors affecting parasite prevalence among wild bumblebees

1. Bumblebees are important pollinators in North America and are attacked by a range of parasites that impact their fitness; however, few studies have investigated the extent or causes of parasitism in North America. 2. This study used a 2‐year multi‐site survey of bumblebee parasitism to ask: (i) how common are parasitoid conopid flies and the internal parasites Crithidia bombi and Nosema bombi in Massachusetts; and (ii) what factors are correlated with parasitism? 3. Infection rates by all three parasites were higher in this study than previously documented in North America. Overall, conopids infected 0–73% of bees in each sample, C. bombi infected 0–82% of bees, and N. bombi infected 0–32%. 4. Conopid flies infected female bees more than males and intermediate‐sized bees more than large or small bees. Crithidia bombi infection rates were higher in certain bee species and sites, and exhibited a unimodal pattern of prevalence over time. Nosema bombi parasitism was higher in male than female bees. 5. Infection by N. bombi in two rare bumblebee species was higher than expected based on parasitism rates of common bee species but C. bombi infection was lower. If high prevalence of N. bombi in these bumblebee species is common, parasitism may be a potential cause of their decline. 6. Given the documented effects of these parasites, the high levels of infection may affect bee populations in Massachusetts and threaten the stability of their valuable ecosystem services.     

Honey bees and wild pollinators differ in their preference for and use of introduced floral resources

Introduced plants may be important foraging resources for honey bees and wild pollinators, but how often and why pollinators visit introduced plants across an entire plant community is not well understood. Understanding the importance of introduced plants for pollinators could help guide management of these plants and conservation of pollinator habitat. We assessed how floral abundance and pollinator preference influence pollinator visitation rate and diversity on 30 introduced versus 24 native plants in central New York. Honey bees visited introduced and native plants at similar rates regardless of floral abundance. In contrast, as floral abundance increased, wild pollinator visitation rate decreased more strongly for introduced plants than native plants. Introduced plants as a group and native plants as a group did not differ in bee diversity or preference, but honey bees and wild pollinators preferred different plant species. As a case study, we then focused on knapweed (Centaurea spp.), an introduced plant that was the most preferred plant by honey bees, and that beekeepers value as a late‐summer foraging resource. We compared the extent to which honey bees versus wild pollinators visited knapweed relative to coflowering plants, and we quantified knapweed pollen and nectar collection by honey bees across 22 New York apiaries. Honey bees visited knapweed more frequently than coflowering plants and at a similar rate as all wild pollinators combined. All apiaries contained knapweed pollen in nectar, 86% of apiaries contained knapweed pollen in bee bread, and knapweed was sometimes a main pollen or nectar source for honey bees in late summer. Our results suggest that because of diverging responses to floral abundance and preferences for different plants, honey bees and wild pollinators differ in their use of introduced plants. Depending on the plant and its abundance, removing an introduced plant may impact honey bees more than wild pollinators.     

From Africa via Europe to South America: migrational route of a species‐rich genus of Neotropical lowland rain forest trees (Guatteria,Annonaceae)

Aim Several recent studies have suggested that a substantial portion of today’s plant diversity in the Neotropics has resulted from the dispersal of taxa into that region rather than by vicariance. In general, three routes have been documented for the dispersal of taxa onto the South American continent: (1) via the North Atlantic Land Bridge, (2) via the Bering Land Bridge, or (3) from Africa directly onto the continent. Here a species‐rich genus of Neotropical lowland rain forest trees (Guatteria, Annonaceae) is used as a model to investigate these three hypotheses. Location The Neotropics. Methods The phylogenetic relationships within the long‐branch clade of Annonaceae were reconstructed (using maximum parsimony, maximum likelihood and Bayesian inference) in order to gain insight in the phylogenetic position of Guatteria. Furthermore, Bayesian molecular dating and Bayesian dispersal–vicariance (Bayes‐DIVA) analyses were undertaken. Results Most of the relationships within the long‐branch clade of Annonaceae were reconstructed and had high support. However, the relationship between the Duguetia clade, the Xylopia–Artabotrys clade and Guatteria remained unclear. The stem node age estimate of Guatteria ranged between 49.2 and 51.3 Ma, whereas the crown node age estimate ranged between 11.4 and 17.8 Ma. For the ancestral area of Guatteria and its sister group, the area North America–Africa was reconstructed in 99% of 10,000 DIVA analyses, while South America–North America was found just 1% of the time. Main conclusions The estimated stem to crown node ages of Guatteria in combination with the Bayes‐DIVA analyses imply a scenario congruent with an African origin followed by dispersal across the North Atlantic Land Bridge in the early to middle Eocene and further dispersal into North and Central America (and ultimately South America) in the Miocene. The phylogenetically and morphologically isolated position of the genus is probably due to extinction of the North American and European stem lineages in the Tertiary.     

PHYLOGENY AND THE EVOLUTION OF HOST PLANT ASSOCIATIONS IN THE LEAF BEETLE GENUS OPHRAELLA (COLEOPTERA,CHRYSOMELIDAE)

Species of Ophraella, a North American genus of leaf beetles (Chrysomelidae), feed variously on eight genera in four tribes of Asteraceae. A phylogenetic analysis, based on morphological features and allozymes, was undertaken to deduce the history of host affiliation within the genus. The two data sets are combined to arrive at a provisional phylogeny of the species, onto which host associations are parsimoniously mapped. Among and within the 12 species studied, at least two shifts are postulated to have occurred among congeneric plant species, five between genera in the same tribe, and four between different tribes of Asteraceae. The phylogeny of Ophraella appears not to be congruent with that of its hosts. This and other evidence indicates that many host shifts in Ophraella postdate the divergence of the host plants, a conclusion that may apply commonly to phytophagous insects. A phenetic analysis of the plants' secondary compounds provides modest support for the hypothesis that host shifts are facilitated by commonalities in plant chemistry. A possible trend in host shifts is evident, from chemically simpler to chemically more forbidding plants. The chemical barriers to host shifts in Ophraella appear to require adaptation in both behavior and in physiological attributes. There is no evidence that the host associations of these insects or the divergence in secondary chemistry of their hosts can be attributed to coevolution.     

Oviposition preference and larval performance of North American monarch butterflies on four Asclepias species

Monarch butterflies, Danaus plexippus L. (Lepidoptera: Nymphalidae), occur world‐wide and are specialist herbivores of plants in the milkweed family (Asclepiadaceae). In North America, two monarch populations breed east and west of the continental divide in areas populated by different host plant species. To examine the population variation in monarch responses to different Asclepias species, we measured oviposition preference and larval performance among captive progeny reared from adult butterflies collected in eastern and western North America. Host plant use was evaluated using two milkweed species widely distributed in eastern North America (A. incarnata and A. syriaca), and two species common to western North America (A. fascicularis and A. speciosa). We predicted that exposure to different host plant species in their respective breeding ranges could select for divergent host use traits, so that monarchs should preferentially lay more eggs on, and larvae should perform better on, milkweed species common to their native habitats. Results showed that across all adult female butterflies, oviposition preferences were highest for A. incarnata and lowest for A. fascicularis, but mean preferences did not differ significantly between eastern and western monarch populations. Larvae from both populations experienced the highest survival and growth rates on A. incarnata and A. fascicularis, and we again found no significant interactions between monarch source population and milkweed species. Moreover, the average rank order of larval performance did not correspond directly to mean female oviposition preferences, suggesting that additional factors beyond larval performance influence monarch oviposition behavior. Finally, significant family level variation was observed for both preference and performance responses within populations, suggesting an underlying genetic variation or maternal effects governing these traits.     

The biogeography of Hoffmannseggia (Leguminosae, Caesalpinioideae, Caesalpinieae): a tale of many travels

Aim The flowering plant genus Hoffmannseggia consists of 21 species distributed amphitropically between the arid regions of the south‐western United States and adjacent Mexico, and west‐central South America. This pattern of geographical disjunction is shared by numerous other angiosperm genera and has been the subject of discussions for more than a century with various authors advocating a northern origin for particular taxa and others advocating a southern origin. This study uses a well‐supported phylogeny of a genus with numerous species in each area to address the issues of a northern or southern origin and the facility with which organisms move between the two continents. Location South‐western United States and northern Mexico, northern Chile and Argentina, southern Bolivia, and western Peru. Methods Using DNA sequence data from the nuclear and chloroplast genomes, we generated a phylogenetic hypothesis for all species of Hoffmannseggia rooted with Zuccagnia and Balsamocarpon. Geographical data were optimized on the resultant tree to assess the probable continent of origin for the genus, the pattern of disjunctions between North and South America, and species radiations within the genus. Main conclusions Hoffmannseggia arose in South America and initially split into a suffrutescent (somewhat woody) and an herbaceous clade. Within each of these major clades, there have been at least two exchanges between North and South America. There are no data to support an ancestral pan‐American range for Hoffmannseggia and we therefore ascribe the amphitropical disjunctions to long‐distance dispersal. The phylogeny clearly shows that all dispersals were from South to North America and they occurred at different times and thus the pattern is not the result of a single simultaneous set of dispersals.     

Pollinator size and its consequences: Robust estimates of body size in pollinating insects

Body size is an integral functional trait that underlies pollination‐related ecological processes, yet it is often impractical to measure directly. Allometric scaling laws have been used to overcome this problem. However, most existing models rely upon small sample sizes, geographically restricted sampling and have limited applicability for non‐bee taxa. Allometric models that consider biogeography, phylogenetic relatedness, and intraspecific variation are urgently required to ensure greater accuracy. We measured body size as dry weight and intertegular distance (ITD) of 391 bee species (4,035 specimens) and 103 hoverfly species (399 specimens) across four biogeographic regions: Australia, Europe, North America, and South America. We updated existing models within a Bayesian mixed‐model framework to test the power of ITD to predict interspecific variation in pollinator dry weight in interaction with different co‐variates: phylogeny or taxonomy, sexual dimorphism, and biogeographic region. In addition, we used ordinary least squares regression to assess intraspecific dry weight ~ ITD relationships for ten bees and five hoverfly species. Including co‐variates led to more robust interspecific body size predictions for both bees and hoverflies relative to models with the ITD alone. In contrast, at the intraspecific level, our results demonstrate that the ITD is an inconsistent predictor of body size for bees and hoverflies. The use of allometric scaling laws to estimate body size is more suitable for interspecific comparative analyses than assessing intraspecific variation. Collectively, these models form the basis of the dynamic R package, “pollimetry,” which provides a comprehensive resource for allometric pollination research worldwide.     

Breeding system and pollination by mimicry of the orchid Tolumnia guibertiana in Western Cuba

The mimicry of malpighiaceous oil‐flowers appears to be a recurrent pollination strategy among many orchids of the subtribe Oncidiinae. These two plant groups are mainly pollinated by oil‐gathering bees, which also specialize in pollen collection by buzzing. In the present study, the floral ecology of the rewardless orchid Tolumnia guibertiana (Oncidiinae) was studied for the first time. The orchid was self‐incompatible and completely dependent on oil‐gathering female bees (Centris poecila) for fruit production. This bee species was also the pollinator of two other yellow‐flowered plants in the area: the pollen and oil producing Stigmaphyllon diversifolium (Malpighiaceae) and the polliniferous and buzzing‐pollinated Ouratea agrophylla (Ochnaceae). To evaluate whether this system is a case of mimetism, we observed pollinator visits to flowers of the three plant species and compared the floral morphometrics of these flowers. The behavior, preferences and movement patterns of Centris bees among these plants, as well as the morphological data, suggest that, as previously thought, flowers of T. guibertiana mimic the Malpighiaceae S. diversifolium. However, orchid pollination in one of the studied populations appears to depend also on the presence of O. agrophylla. Moreover, at the two studied populations, male and female pollination successes of T. guibertiana were not affected by its own floral display, and did not differ between populations. The results are discussed in relation to the behavior and preferences of Centris bees, as well as the differential presence and influence of each of the two floral models.     

Intercontinental disjunctions in Cryptotaenia (Apiaceae, Oenantheae): an appraisal using molecular data

Aim The angiosperm genus Cryptotaenia (family Apiaceae, tribe Oenantheae) exhibits an anomalous distribution pattern, with five of its eight species being narrow endemics geographically isolated from their presumed relatives. We examined the monophyly of the genus and ascertained the phylogenetic placements of its constituent members in order to explain their distribution patterns. Location Eastern North America, eastern Asia, the Caucasus, southern Italy, Macaronesia and Africa. Methods In total, 173 accessions were examined for nuclear rDNA ITS sequence variation, representing nearly all major lineages of Apiaceae subfamily Apioideae and seven species of Cryptotaenia. Sampling of tribes Oenantheae, Scandiceae and Pimpinelleae was comprehensive. Phylogenetic analyses included Bayesian, maximum parsimony and neighbour‐joining methods; biogeographical scenarios were inferred using dispersal–vicariance analysis (diva ). Results Cryptotaenia is polyphyletic and includes three distant lineages. (1) Cryptotaenia sensu stricto (C. canadensis, C. japonica, C. flahaultii and C. thomasii) is maintained within tribe Oenantheae; C. canadensis and C. japonica, representing an eastern North American–eastern Asian disjunction pattern, are confirmed to be sister species. (2) Cryptotaenia elegans, endemic to the Canary Islands, is placed within Scandiceae subtribe Daucinae along with two woody endemics of Madeira, Monizia edulis and Melanoselinum decipiens. The phylogeny of these Canarian and Madeiran endemics is unresolved. Either they constitute a monophyletic sister group to a clade comprising some Mediterranean and African species of Daucus and their relatives, or they are paraphyletic to this clade. The herbaceous/woody genus Tornabenea from Cape Verde, once included in Melanoselinum, is not closely related to the other Macaronesian endemics but to Daucus carota. (3) The African members of Cryptotaenia (C. africana, C. calycina and possibly C. polygama) comprise a clade with some African and Madagascan umbellifers; this entire clade is sister group to Eurasian Pimpinella. Main conclusions Elucidating the phylogeny of the biogeographically anomalous Cryptotaenia sensu lato enabled hypotheses on the biogeography of its constituent lineages. Cryptotaenia sensu stricto exhibits a holarctic distribution pattern, with its members occurring in regions that were important glacial refugia. The genus probably originated in eastern Asia and from there dispersed to Europe and North America. For the Macaronesian endemic species –C. elegans, M. edulis and M. decipiens–diva reconstructs either a single dispersal event to Macaronesia from the Mediterranean/African region, or a single dispersal followed by a back‐dispersal to the mainland. The radiation of Tornabenea from Cape Verde followed a second dispersal of Daucinae to Macaronesia. Woodiness in Melanoselinum/Monizia and Tornabenea, therefore, is a derived and independently acquired trait. The African members of Cryptotaenia are derived from an ancestor arriving from the Middle East.     

The origin and evolution of Indomalesian,Australasian and Pacific island biotas: insights from Aglaieae (Meliaceae,Sapindales)

Aim The role of long‐distance dispersal in the Indomalesian, Australasian and Pacific flora is currently hotly debated. The lack of well‐resolved phylogenetic trees for Pacific plants has been a major limitation for biogeographical analysis. Here, we present a well‐resolved phylogenetic tree for the tribe Aglaieae in the mahogany family, Meliaceae, and use it to investigate the origin, evolution and dispersal history of biotas in this area. The subfamily Melioideae, including the tribe Aglaieae (Meliaceae, Sapindales), is a plant group with good representation in the region in terms of biomass and species numbers, wide ecological attributes and known animal vectors. The family has a good fossil record (especially from North America and Europe). Genera and species in the tribe Aglaieae therefore provide an excellent model group for addressing this debate. Location Indomalesia, Australasia, Pacific islands. Methods Results from nuclear internal transcribed spacer ribosomal DNA analyses of 82 taxa, based on sequence alignment guided by secondary structure models, were combined with evidence from fossils and distribution data. We used strict and relaxed molecular clock approaches to estimate divergence times within Aglaieae. Putative ancestral areas were investigated through area‐based and event‐based biogeographical approaches. Information on dispersal routes and their direction was inferred from the investigation of dispersal asymmetries between areas. Results Our study indicates that the crown group of Aglaieae dates back at least to the Late Eocene, with major divergence events occurring during the Oligocene and Miocene. It also suggests that dispersal routes existed during Miocene–Pliocene times from the area including Peninsular Malaysia, Sumatra and Borneo to Wallacea, India and Indochina, and from the area including New Guinea, New Ireland and New Britain further east to the Pacific islands at the peripheries of the distribution range. The origin of the Fijian species dates back to the Pliocene. Main conclusions Dispersal over oceanic water barriers has occurred during geological time and seems to have been a major driving force for divergence events in Aglaieae, with some old Gondwanan land masses (e.g. Australia) colonized only during recent times. Movement from the ancestral area was predominantly towards the east. Extant Fijian species of Aglaia are monophyletic and share morphological features rarely found in species of other areas, suggesting speciation within an endemic clade. Divergence of living taxa from their closest living relatives took place during both the Miocene and the Pliocene, and peaked in the Pliocene. The present‐day distribution of many species in the tribe must therefore have arisen as a result of dispersal rather than vicariance events. Furthermore, colonization from Indomalesia to Australasia and the Pacific has frequently been followed by speciation.     

Life cycle and life strategy features of Puccinia glechomatis (Uredinales) favorable for extending the natural range of distribution

The objective of this study was to find features in microcyclic rust fungi (Uredinales) on wild host plants favorable for extension of the natural range of distribution. Puccinia glechomatis, a leptosporic rust fungus and its herbal host Glechoma hederacea (Lamiaceae), both natives to Eurasia and introduced in North America, were used for this study. Although the host has been known from North America since the beginning of the nineteenth century, the rust fungus was first observed there only in recent years. Favorable features were identified by studying the life cycle of the rust, including nuclear conditions and seasonal characteristics as well as its spread in North America. The life cycle was studied macroscopically by inoculation experiments, by various light microscope techniques, and by scanning electron microscopy. The spread of the pathogen and its host were reconstructed by evaluating host plant herbarium specimens and databases, literature, and field study data. The studies on P. glechomatis show that, generally for microcyclic rust fungi, establishment and potential for spread are based on several favorable features of both the host (e.g., synanthropic occurence and dispersal, genetic stability, regeneration of vegetative plant parts) and the rust fungus (asexual reproduction/genetic stability, homothallism, propagation with host plant, formation of both leptospores and thick-walled teliospores).