Phylogenetic positions of seven poorly known species of Ferula (Apiaceae) with remarks on the phylogenetic utility of the plastid trnH-psbA,trnS-trnG,and atpB-rbcL intergenic spacers

Ferula L. is one of the most species-rich and taxonomically difficult genera of Apiaceae. In this study, we obtained nrDNA ITS sequences of seven poorly known species of Ferula (Ferula anatolica, Ferula sp. (tentatively identified as F. candelabrum by collectors), F. drudeana, F. huber-morathii, F. marmarica, F. talassica, and F. tunetana) and explored their phylogenetic positions using 148 ITS sequences of the subtribe Ferulinae from GenBank. Five of these newly sequenced species fall into three groups, corresponding to clades recognized in earlier molecular studies. Ferula sp. are added to clade, which is mostly composed of Central Asian species. This placement showed that identification as F. candelabrum was erroneous. The second clade, which is mostly composed of Mediterranean taxa, includes two species from North Africa: F. marmarica and F. tunetana. Despite the well-supported monophyly of this clade, the relationships inside this group need to be revised, as broadly distributed F. communis is paraphyletic with respect to other species. Ferula drudeana and F. huber-morathii, two narrow endemics from Turkey, are placed in the Central Asian clade. Two species, F. anatolica and F. talassica, do not fall into any of the recognized clades. In addition, we examined the sequence variation of three potentially highly variable pDNA regions, the trnH-psbA, trnS-trnG, and atpB-rbcL intergenic spacers, for a subset of 18 specimens. The resulting pDNA and ITS based phylogenetic trees were incongruent, as supported by significant ILD tests. The cause of this incongruence can be manifold, including hybridization, a lack of a phylogenetic signal, and homoplastic substitutions. Our analyses suggest that only trnS-trnG can be added to the list of pDNA markers used for phylogenetic studies of Ferula, as it has the highest number of parsimony informative characters and is easy to amplify from degraded material.     

REPRODUCTIVE MORPHOLOGY OF QUERCUS (FAGACEAE)

Details of inflorescence, floral, and fruit morphology have been studied in more than 120 species of Asiatic and American Quercus. Of the two subgenera, subgenus Cyclobalanopsis has fewer species but greater diversity of reproductive morphology than subgenus Quercus. Some character states of subgenus Cyclobalanopsis, such as more numerous stamens, male flowers sometimes grouped in dichasia, abortive ovules well developed, prominent intrusive septae in the nut, and lamellate cupules, are shared with Lithocarpus. The diversity of cupular sizes, coverage, and ornamentation raises questions about the adaptive nature, ecological function, and phylogeny of the cupule, which clearly has by now evolved as a structure with its own qualities.     

Phylogenetic relationships within the genus Sargassum (Fucales, Phaeophyceae), inferred from ITS-2 nrDNA, with an emphasis on the taxonomic subdivision of the genus

Sequences from the ribosomal DNA internal transcribed spacer‐2 (ITS‐2) were compared among species of Sargassaceae including the genera Sargassum and Hizikia. Species of different subgenera and sections of Sargassum were used to assess the taxonomic relationships within the genus, especially the subdivisions of the subgenus Bactrophycus. Sequences were aligned in accordance with their common secondary structure. Phylogenetic trees were constructed using neighbor‐joining, maximum likelihood and maximum parsimony methods with three species of Turbinaria as outgroups. The resulting phylogenetic trees showed that the genus Sargassum is divided into three clades corresponding to the subgenera Phyllotrichia, Sargassum and Bactrophycus. This last subgenus is further divided into four distinct groups: a Spongocarpus clade, a Teretia clade, a Hizikia clade, and a Halochloa/ Repentia clade. The position of the section Phyllo‐cystae, excluded from the subgenus Bactrophycus and included within the subgenus Sargassum is once again confirmed by the present study. Current results strongly support the assignation of Hizikia fusiformis to the genus Sargassum. Based on morphological differences and a distinct position in the molecular trees, Hizikia should be recognized as a section in the subgenus Bactrophycus so that Hizikia (Okamura) Yoshida, stat. nov. is proposed. A remarkably low divergence of ITS‐2 sequences was observed for the species in the sections Repentia and Halochloa, suggesting very recent radiation of these species. The subgenus Sargassum is divided into three clades corresponding to the three known sections: Acanthocarpicae, Malacocarpicae and Zygocarpicae, previously recognized by the morphology of receptacles. The position of Sargassum duplicatum, S. carpophyllum, S.yendoi, S. piluliferum and S. patens within the subgenus Sargassum is discussed.     

Phylogenetic analysis of the Brazilian microendemic Paepalanthus subgenus Xeractis (Eriocaulaceae) inferred from morphology

Paepalanthus subgenus Xeractis (Eriocaulaceae) comprises 28 recognized species endemic to the Espinhaço Range, in Minas Gerais state, Brazil. Most species of the subgenus are restricted to small localities and critically endangered, but still in need of systematic study. The monophyly of the subgenus has already been tested, but only with a few species. Our study presents the first phylogenetic hypothesis within the group, based on morphology. A maximum parsimony analysis was conducted on a matrix of 30 characters for 30 terminal taxa, including all species of the subgenus and two outgroups. The biogeographical hypotheses for the subgenus were inferred based on dispersal–vicariance analysis (DIVA). The analysis provided one most‐parsimonious hypothesis that supports most of the latest published subdivisions (sections and series). However, some conflicts remain concerning the position of a few species and the relationships between sections. The distribution and origin(s) of microendemism are also discussed, providing the ground for conservation strategies to be developed in the region. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 137–152.     

Phylogenetic analysis of eastern Asian and eastern North American disjunct Lespedeza (Fabaceae) inferred from nuclear ribosomal ITS and plastid region sequences

Lespedeza (tribe Desmodieae, Fabaceae) follows a disjunct distribution in eastern Asia and eastern North America. Phylogenetic relationships among its species and related taxa were inferred from nuclear ribosomal internal transcribed spacer (ITS) and plastid sequences (trnH‐psbA, psbK‐psbI, trnK‐matK and rpoC1). We examined 35 species of Lespedeza, two of Kummerowia and one of Campylotropis, the sole constituents of the Lespedeza group. An analysis of these data revealed that the genus Campylotropis is sister to the other two genera. However, we were unable to resolve the relationships between Kummerowia and Lespedeza in the strict consensus trees of parsimony analyses based on plastid and combined DNA data. In the genus Lespedeza, the Old World subgenus Macrolespedeza is monophyletic, whereas the transcontinental subgenus Lespedeza is paraphyletic. Monophyly of eastern Asian species and of North American species is strongly supported. Although inconsistent with the traditional classification, this phylogenetic finding is consistent with seedling morphology. Three subgroups recognized in subgenus Macrolespedeza were unresolved in our phylogenetic trees. An incongruence length difference (ILD) test indicated that the two partitions (nuclear ITS and plastid sequences) were significantly incongruent, perhaps because of hybridization between species in Lespedeza. Most of the primary clades of tribe Desmodieae are Asian, implying that the relatively few New World ones, such as those in Lespedeza, are more recently derived from Asia. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 221–235.     

Phylogenetic relationships of Sargassum (Sargassaceae, Phaeophyceae) with reference to a taxonomic revision of the section Phyllocystae based on ITS-2 nrDNA sequences

Sequences of the end of the 5.8S gene and the internal transcribed spacer 2 (ITS‐2) of nuclear ribosomal DNA have been determined for 19 species of the brown algal genus Sargassum (Sargassaceae), representing three subgenera and eight sections (sections are in parentheses): Phyllotrichia, Bactrophycus (Teretia, Spongocarpus, Halochloa and Repentia) and Sargassum (Acanthocarpicae, Malacocarpicae, Zygocarpicae) to assess the taxonomic position of the section Phyllocystae traditionally included within the Bactrophycus. The sequence of Myagropsis myagroides (Mertens ex Turner) Fensholt (Sargassaceae) was used as an outgroup. Sequences of ITS‐2 were analyzed using neighbor‐joining, parsimony and maximum likelihood methods. The results showed the existence of three clades in Sargassum, corresponding to the three subgenera. The subgenus Phyllotrichia is positioned near the outgroup. Two robust clades were obtained, one corresponding to the subgenus Bactrophycus and the other to the subgenus Sargassum. Sargassum mcclurei Setchell and Sargassum quinhonense Nguyen, the two Phyllocystae investigated, are close to species belonging to the section Zygocarpicae in the subgenus Sargassum. A transfer of the section Phyllocystae to the subgenus Sargassum is therefore proposed on the basis of molecular data (ITS‐2) and morphological data (receptacles and basal leaf).     

Angelica oncosepala and Heracleum yunnanense are synonyms and refer to a species of Tetrataenium (Apiaceae)

Heracleum yunnanense Franchet is a poorly known taxon known only from the type collection. Angelica oncosepala Handel‐Mazzetti was originally described as a member of Angelica and later placed in Heracleum as Heracleum oncosepalum (Handel‐Mazzetti) Pimenov & Kljuykov. The generic taxonomy of Apiaceae has been largely based on the morphology and anatomy of the fruit. In this study, various characters of A. oncosepala are found to be identical with H. yunnanense. In addition, we discovered that H. yunnanense and A. oncosepala have well developed narrowly keeled dorsal and intermediate ribs, broadly winged marginal ribs, 1 vittae in dorsal valleculae, 2 in lateral valleculae, 4 on commissure and linear vittae, as do members of Tetrataenium, but not congruent with species of Heracleum. Phylogenetic analyses using concatenated nuclear ITS and ETS sequences also suggested that A. oncosepala and H. yunnanense should be merged and placed in Tetrataenium. The species name H. yunnanense is the oldest available and Tetrataenium yunnanense (Franch.) Manden is only invalidly published, so we propose the new combination Tetrataenium yunnanense (Franch.) Manden. ex Q. Y. Xiao & X. J. He, and treat A. oncosepala, H. oncosepalum and H. yunnanense as its synonyms.     

The subgenus Scaptodrosophila (Diptera: Drosophilidae)

Four major subgenera, Drosophila, Sophophora, Hirtodrosophila and Scaptodrosophila, and several minor subgenera, have been recognized in the genus Drosophila. The hitherto little studied subgenus Scaptodrosophila, possessing a complicated synonymy, contains 144 named species, or about 11% of the total described in the genus; it is likely that many more Scaptodrosophila species await discovery. Scaptodrosophila probably originated in tropical Asia, and the greatest Scaptodrosophila faunas occur in Asia, south-east Asia, New Guinea, Australia and Africa, with very few species in north and south America and Europe. Scaptodrosophila species usually possess a pair of prescutellar bristles, a propleural bristle, and three large subequal sternopleural bristles; other morphological features are more variable. Ecological information concerning the majority of Scaprodrosophila species is scanty or absent, but known feeding or breeding sites include tree sap, fungi, fruit, flowers and 1eaf litter; larvae of at least one species are gall forming in plant tissues. The establishment of species groups is complicated by lack of detailed knowledge of many species, but six groups containing four or more species are recognized.     

Molecular evolution of tephritid fruit flies in the genus Bactrocera based on the cytochrome oxidase I gene

Fruit flies of the genus Bactrocera (Diptera: Tephritidae) are one of the major economically important insects in Asia and Australia. Little attention has been given to analyses of molecular phylogenetic relationships among Bactrocera subgenera. By using mitochondrial cytochrome oxidase I gene (COI) sequences, the phylogenetic relationships among four subgenera, Asiadacus, Bactrocera, Hemigymnodacus, and Zeugodacus, were investigated. Nucleotide diversity within subgenera ranged from 11.7 to 12.4%, and the net divergence among subgenera ranged from 11.2 to 15.7%. Phylogenetic trees calculated from both maximum parsimony and neighbor-joining phylogenetic analysis methods were highly congruent in terms of tree topologies. Phylogenetic analysis of mitochondrial COI sequences suggests that tephritid fruit fly species, which attack cucurbit plants, that is, Asiadacus, Hemigymnodacus and Zeugodacus, were more closely related to each other than to fruit fly species of the subgenus Bactrocera, which attack plants of numerous families. Our data supports previous classification of Bactrocera based on morphological characters. However, the phylogenetic tree showed the polyphyletic of fruit flies in subgenus Zeugodacus. Possible causes of speciation among fruit flies species in this genus were also discussed.     

Chemotaxonomic Studies in Ferula of China on the Basis of Flavonoids

Sixteen species of Ferula L. (Umbelliferae) were examined for their leaf flavonoids, and at least 9 glycosides of quercetin, kaempferol and apigenin are detected in 13 species. While most of the constituents reported are common in the Umbelliferae, the unusual one, apigenin-6, 8-C-diglucoside, which was found in Ostericum Hoffm. in an earlier survey has been detected in F. moschata (F. sumbul) The three species (F. sinkiangensis, F. fukangensis and F. krylovii) from which flavonoids have not been found, are all medicinal plants with strong garliky odor due to a high concentration of essential oils containing disulfides. The subgenus Peucedanoides is found to be rich in quercetin glycosides, and the aglycon pattern seems to have some relationship with habitat conditions in the genus: the species occurring in comparatively humid habitats are often large herbs and often characterized by the presence of quercetin, while those growing in dry habitats contain apigening kaepferol, as well as quercetin. Nevertheless, any conclusion with certainty needs examining more species of the genus. It is interesting to note that F. moschata, from which the glycosylapigenin was found, and F. pseudooreoselinum, which probably has a trace amount of kaempferol glycoside, are peculiar in Ferula in their inflorescence morphology. The phytochemic results also raise a suspicion against their systematic position within the genus. F. bungeana, previously placed in the subgenus Peucedanoides, has relatively great similarities with F. syreitschkowii of the subgenus Narthex in their general morphology and fruit anatomy. In the present survey apigenin-7-glycoside is detected from both, and thus it seems more suitable to transfer the former species into the subgenus Narthex. The same compounds have been found in F. licentiana and F. tunshanica, which also share the characters of morphology and anatomy. The fact does support the view that the lattershould be included in F. licentiana as a variety.     

Molecular phylogeny of the subgenus Holothuria (Selenkothuria) Deichmann, 1958 (Holothuroidea: Aspidochirotida)

The subgenus Selenkothuria comprises 12 species of tropical shallow water sea cucumbers that share morphological features, such as rods in the body wall and tube feet, modified tentacles for suspension feeding, and cryptic colours. The taxonomic status of this taxon has been controversial, but currently it is accepted as a subgenus of the genus Holothuria. Phylogenetic analyses of mitochondrial genes [cytochrome c oxidase subunit 1 (COI), 16S RNA] of ten species of Selenkothuria and related subgenera showed the polyphyly of this subgenus; monophyly was rejected by a likelihood ratio test. A geographical split divides the species of this subgenus into three different groups: one Indo‐West‐Pacific (IWP) group and two American groups. The IWP group is more closely related to Holothuria (Semperothuria) cinerascens and to other subgenera such as Roweothuria, Holothuria, and Vaneyothuria, whereas the two American groups are more closely related to each other and to some species of the subgenus Halodeima. These results suggest multiple parallel originations and diversification of ossicle morphology within the subgenus Selenkothuria. The current scheme of subgenera for the genus Holothuria is not supported, suggesting the need for a new classification. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 109–120.     

Pollen morphology and phylogenetic relationships in neotropical Phyllanthus (Euphorbiaceae)

In order to provide new insights into phylogenetic relationships among the neotropical taxa of Phyllanthus , 28 illustrations are provided of the pollen grains of 22 selected species studied from 11 sections of the subgenera represented in the neotropics. Special attention has been given to subgenus Conami because of its variability in pollen morphology: of eight species illustrated, the apertures are diploporate colpi in three species and pores in five species; exine ornamentation is vermiculate in two species and pilate in the other six species. The six species in the neotropical sections Pityrocladus and Microglochidion (subgenus Emblica ) are characterized by prolate grains with an increased number of colpi (4–8). Of particular interest are species in which the pollen exine is clypeate (with exine shields); clypeate pollen grains are illustrated in two species of subgenus Xylophylla and in one species of section Cyclanthera that has unique exine shields with single central pila. The pollen of the one Brazilian phylloclade-bearing species illustrated (in section Choretropsis ) has 3-colporate grains with reticulate exine, typical for subgenus Phyllanthus , and very different from the clypeate grains of the West Indian phylloclade-bearing species in section Xylophylla . This pollen evidence clearly demonstrates homoplasy in the origin of phylloclades in Phyllanthus . Pollen morphological data suggest that the neotropical taxa of Phyllanthus have arisen following colonization from Africa (subgenus Kirganelia ) and Asia (subgenus Emblica ). © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 325–338.     

One and the same: integrative taxonomic evidence that Bactrocera invadens (Diptera: Tephritidae) is the same species as the Oriental fruit fly Bactrocera dorsalis

The invasive fruit fly Bactrocera invadens Drew, Tsuruta & White, and the Oriental fruit fly Bactrocera dorsalis (Hendel) are highly destructive horticultural pests of global significance. Bactrocera invadens originates from the Indian subcontinent and has recently invaded all of sub‐Saharan Africa, while B. dorsalis principally occurs from the Indian subcontinent towards southern China and South‐east Asia. High morphological and genetic similarity has cast doubt over whether B. invadens is a distinct species from B. dorsalis. Addressing this issue within an integrative taxonomic framework, we sampled from across the geographic distribution of both taxa and: (i) analysed morphological variation, including those characters considered diagnostic (scutum colour, length of aedeagus, width of postsutural lateral vittae, wing size, and wing shape); (ii) sequenced four loci (ITS1, ITS2, cox1 and nad4) for phylogenetic inference; and (iii) generated a cox1 haplotype network to examine population structure. Molecular analyses included the closely related species, Bactrocera kandiensis Drew & Hancock. Scutum colour varies from red‐brown to fully black for individuals from Africa and the Indian subcontinent. All individuals east of the Indian subcontinent are black except for a few red‐brown individuals from China. The postsutural lateral vittae width of B. invadens is narrower than B. dorsalis from eastern Asia, but the variation is clinal, with subcontinent B. dorsalis populations intermediate in size. Aedeagus length, wing shape and wing size cannot discriminate between the two taxa. Phylogenetic analyses failed to resolve B. invadens from B. dorsalis, but did resolve B. kandiensis. Bactrocera dorsalis and B. invadens shared cox1 haplotypes, yet the haplotype network pattern does not reflect current taxonomy or patterns in thoracic colour. Some individuals of B. dorsalis/B. invadens possessed haplotypes more closely related to B. kandiensis than to conspecifics, suggestive of mitochondrial introgression between these species. The combined evidence fails to support the delimitation of B. dorsalis and B. invadens as separate biological species. Consequently, existing biological data for B. dorsalis may be applied to the invasive population in Africa. Our recommendation, in line with other recent publications, is that B. invadens be synonymized with B. dorsalis.     

Seed and capsule morphology of Iranian perennial species of Euphorbia (Euphorbiaceae) and its phylogenetic application

Of the 480 species of Euphorbia subgenus Esula, c. 290 occur in the Mediterranean and Irano‐Turanian regions. Turkey and Iran are the most species‐rich countries in Asia with 83 and 74 species, respectively. Following our previous paper on annual species of Iranian Euphorbia, we studied the quantitative and qualitative macro‐ and micromorphological traits of seeds and capsules of 47 perennial species, including E . ferdowsiana sp. nov. , E . sulphurea sp. nov. and E. glareosa, as a first report from Iran. A key for all Iranian perennial Euphorbia spp. based on seed and capsule morphology is provided. The phylogenetic relationships of Iranian species based on internal transcribed spacer (ITS) nuclear and ndhF plastid regions are updated and used for the characterization of the synapomorphies of each clade. Capsule shape, seed shape, seed surface and shape of the caruncle have been found to be homoplastic, whereas the presence or absence of granulate elements on seed surfaces represents a phylogenetically important trait for section delimitation. The capsule surface is synapomorphic for several sections, including Helioscopia (tuberculate‐verrucose), Myrsiniteae (vesiculate) and Esula (granulate), and seed shape is synapomorphic for sections Helioscopia (ellipsoidal), Myrsiniteae (ovoid‐quadrangular) and Herpetorrhizae (pseudo‐hexahedral). Reversals have also taken place in some features, including capsule surface (E. mazandaranica, E. altissima) and seed shape (E. densa, E. aleppica). It seems that ecarunculate seeds are plesiomorphic in sections Helioscopia (E. eriophora) and Herpetorrhizae (E. consanguinea and E. turczaninowii). © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 00 , 000–000.     

Taste sensitivity and divergence in host plant acceptance between adult females and larvae of Papilio hospiton

On the island of Sardinia the lepidopteran Papilio hospiton uses Ferula communis as exclusive host plant. However, on the small island of Tavolara, adult females lay eggs on Seseli tortuosum, a plant confined to the island. When raised in captivity on Seseli only few larvae grew beyond the first–second instar. Host specificity of lepidopterans is determined by female oviposition preferences, but also by larval food acceptance, and adult and larval taste sensitivity may be related to host selection in both cases. Aim of this work was: (i) to study the taste sensitivity of larvae and ovipositing females to saps of Ferula and Seseli; (ii) to cross‐compare the spike activity of gustatory receptor neurons (GRNs) to both taste stimuli; (iii) to evaluate the discriminating capability between the two saps and determine which neural code/s is/are used. The results show that: (i) the spike responses of the tarsal GRNs of adult females to both plant saps are not different and therefore they cannot discriminate the two plants; (ii) larval L‐lat GRN shows a higher activity in response to Seseli than Ferula, while the opposite occurs for the phagostimulant neurons, and larvae may discriminate between the two saps by means of multiple neural codes; (iii) the number of eggs laid on the two plants is the same, but the larval growth performance is better on Ferula than Seseli. Taste sensitivity differences may explain the absence of a positive relationship between oviposition preferences by adult females and plant acceptance and growth performance by larvae.     

The bumblebees of the subgenus Subterraneobombus: integrating evidence from morphology and DNA barcodes (Hymenoptera,Apidae, Bombus)

Although bumblebees have received a lot of attention, some taxonomic problems have persisted for many years. One particularly obdurate case has been the species of the subgenus Subterraneobombus. We revise the bees of this subgenus by integrating evidence from both morphology and, for a 5% subsample, from DNA (cytochrome c oxidase subunit 1, CO1) barcodes from pinned museum specimens. We apply a reciprocal illumination procedure: (1) taxa recognized previously from morphology are used to stratify samples for DNA subsampling; (2) DNA barcodes from these subsamples are used to recognize groups of phylogenetically related specimens; and (3) for these groups, we re‐examine morphological characters in order to recognize and diagnose species. A total of 3854 specimens from 1535 samples from across the geographic range of the subgenus throughout the Holarctic and northern Oriental regions are identified to 11 species. This includes one species newly recognized from Mongolia, Bombus mongolensis Williams sp. nov. Taxon concepts are modified substantially for four species, seven lectotypes are designated, and four new synonyms are recognized. The prevailing usage of Bombus distinguendus is maintained as valid by designating Bombus elegans as a nomen oblitum and designating B. distinguendus as a nomen protectum. Identification keys and colour‐pattern diagrams are provided, and geographic distributions, elevational ranges, and phenological activity periods are described to characterize the species. An estimate of the biogeographic history is reconstructed with dispersal–vicariance analysis. In this study, DNA barcode data have been a cost‐effective source of additional characters for diagnosing groups of specimens. The barcode data contributed directly to recognizing the one new species, of which females remain difficult to identify from morphology alone. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 813–862.     

Tropical Asian species show that the Old World clade of ‘spiny solanums’ (Solanum subgenus Leptostemonum pro parte: Solanaceae) is not monophyletic

The tropical Asian taxa of the species‐rich genus Solanum (Solanaceae) have been less well studied than their highly diverse New World relatives. Most of these tropical Asian species, including the cultivated brinjal eggplant/aubergine and its wild progenitor, are part of the largest monophyletic Solanum lineage, the ‘spiny solanums’ (subgenus Leptostemonum or the Leptostemonum clade). Here we present the first phylogenetic analysis of spiny solanums that includes broad sampling of the tropical Asian species, with 42 of the 56 currently recognized species represented. Two nuclear and three plastid regions [internal transcribed spacer (ITS), waxy, ndhF‐rpL32, trnS‐trnG and trnT‐trnF] were amplified and used to reconstruct phylogenetic relationships using maximum likelihood and Bayesian methods. Our analyses show that Old World spiny solanums do not resolve in a single clade, but are part of three unrelated lineages, suggesting at least three independent introductions from the New World. We identify and describe several monophyletic groups in Old World solanums that have not been previously recognized. Some of these lineages are coherent in terms of morphology and geography, whereas others show considerable morphological variation and enigmatic distribution patterns. Tropical Asia occupies a key position in the biogeography of Old World spiny solanums, with tropical Asian taxa resolved as the closest relatives of diverse groups of species from Australia and Africa.     

Phylogenetic analysis of Allium subg. Melanocrommyum infers cryptic species and demands a new sectional classification

Allium subgenus Melanocrommyum (Alliaceae) from Eurasia comprises about 150 mostly diploid species adapted to arid conditions. The group is taxonomically complicated with different and contradictory taxonomic treatments, and was thought to include a considerable number of hybrid species, as the taxa show an admixture of assumed morphological key characters. We studied the phylogeny of the subgenus, covering all existing taxonomic groups and their entire geographic distribution. We analyzed sequences of the nuclear rDNA internal transcribed spacer region (ITS) for multiple individuals of more than 100 species. Phylogenetic analyses of cloned and directly sequenced PCR products confirmed the monophyly of the subgenus, while most sections were either para- or polyphyletic. The splits of the large sections are supported by differences in the anatomy of flower nectaries. ITS data (i) demand a new treatment at sectional level, (ii) do not support the hypotheses of frequent gene flow among species, (iii) indicate that multiple rapid radiations occurred within different monophyletic groups of the subgenus, and (iv) detected separately evolving lineages within three morphologically clearly defined species (cryptic species). In two cases these lineages were close relatives, while in Allium darwasicum they fall in quite different clades in the phylogenetic tree. Fingerprint markers show that this result is not due to ongoing introgression of rDNA (ITS capture) but that genome-wide differences between both lineages exist. Thus, we report one of the rare cases in plants where morphologically indistinguishable diploid species occurring in mixed populations are non-sister cryptic species.