Systematics and phylogeny of the tribe Paragini (Diptera: Syrphidae) based on molecular and morphological characters

We studied the phylogeny and systematics of the tribe Paragini (Diptera: Syrphidae) using morphological and molecular data. The paper presents separate parsimony analyses of both adult morphological characters and partial DNA sequence data from mitochondrial cytochrome c oxidase I and nuclear ribosomal 28S rRNA gene, as well as a combined analysis of all the data. The data set of morphological characters included some features of the male terminalia (i.e. shape of the ejaculatory apodeme; relative position of elements of the aedeagal complex; shape of surstylar apodeme; shape of the aedeagal apodeme) not previously used in the systematics of the Paragini. The trees obtained from separate parsimony analyses of molecular and morphological data produced almost identical topologies. Four lineages are supported by the combined data set, and we establish two new subgenera, i.e. Serratoparagus Vujić et Radenković subgen. nov., and Afroparagus Vujić et Radenković subgen. nov., and redefine Pandasyopthalmus Stuckenberg, 1954 stat. rev. and Paragus Latreille, 1804, stat. rev. The monophyly of the Pandasyopthalmus clade, including the species fitting neither of the current species groups ( jozanus -group) of Paragini, is established. Diagnoses of all known species groups are presented, including a new arrangement of almost all valid species of Paragini.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 507–536.     

Phylogenetic analysis of Stylosanthes (Fabaceae) based on the internal transcribed spacer region (ITS) of nuclear ribosomal DNA

 Phylogenetic relationships in Stylosanthes are inferred by DNA sequence analysis of the ITS region (ITS1–5.8S–ITS2) of the nuclear ribosomal DNA in 119 specimens, representing 36 species of Stylosanthes and 7 species of the outgroup genera Arachis and Chapmannia. In all examined specimens of any particular diploid and (allo)polyploid species, only a single ITS sequence type was observed. This allowed us to identify a parental genome donor for some of the polyploids. In several diploid and polyploid species, different specimens contained a different ITS sequence. Some of these sequence types were present in more than one species. Parsimony analysis yielded several well-supported clades that agree largely with analyses of the chloroplast trnL intron and partially with the current sectional classification. Discordances between the nuclear and cpDNA analyses are explained by a process of allopolyploidization with inheritance of the cpDNA of one parent and fixation of the ITS sequences of the other. S. viscosa has been an important genome donor in this process of speciation by allopolyploidy. Received August 14, 2001; accepted March 4, 2002 Published online: November 14, 2002 Addresses of the authors: Jacqueline Vander Stappen, Steven Van Campenhout and Guido Volckaert (E-mail: guido.volckaert@agr.kuleuven.ac.be), Katholieke Universiteit Leuven, Laboratory of Gene Technology, Kasteelpark Arenberg 21, B-3001 Leuven, Belgium. Jan De Laet, American Museum of Natural History, Division of Invertebrate Zoology, Central Park West at 79th Street, New York 10024–5192, USA. Susana Gama-López, Universidad Nacional Autónoma de México, Unidad de Biología, Tecnología y Protipos (UBIPRO), FES-Iztacala, Laboratorio de Recursos Naturales, Av. de Los Barrios S/N, Colonia Los Reyes Iztacala, Municipio Tlalnepantla, Estado de México, C.P. 54090, México. Present address: Apartado Postal 154, Cto. Parque No. 3, C.P. 53102, México.     

Phylogenetic analysis of the genus Hexachlamys (Myrtaceae) based on plastid and nuclear DNA sequences and their taxonomic implications

Myrtaceae are one of the most species‐rich families of flowering plants in the Neotropics. They include several complex genera and species; Hexachlamys is one of the complex genera. It has not been recognized as a distinct genus and has been included in Eugenia, based on morphological grounds. Therefore, molecular systematic studies may be useful to understand and to help to solve these relationships. Here, we performed a molecular phylogenetic analysis using plastid and nuclear data in order to check the inclusion of Hexachlamys in Eugenia. Plastid (accD, rpoB, rpoC1, trnH‐psbA) and nuclear (ITS2) sequence data were analysed using Bayesian and maximum parsimony methods. The trees constructed using ITS2 and trnH‐psbA were the best able to resolve the relationships between species and genera, revealing the non‐monophyly of Hexachlamys. The molecular phylogenetic analyses were in agreement with previous morphological revisions that have included Hexachlamys in Eugenia. These results reinforce the importance of uniting knowledge and strategies to understand better issues of delimitation of genera and species in groups of plants with taxonomic problems. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 532–543.     

Phylogenetic analysis of Silphium and subtribe Engelmanniinae (Asteraceae: Heliantheae) based on ITS and ETS sequence data

The phylogenetic relationships of Silphium and subtribe Engelmanniinae were examined using DNA sequence data. The internal transcribed spacer (ITS) region and the external transcribed spacer (ETS) region were sequenced for 39 specimens representing the six genera of subtribe Engelmanniinae (Berlandiera, Chrysogonum, Dugesia, Engelmannia, Lindheimera, and Silphium), plus five additional genera identified as closely related to the Engelmanniinae by chloroplast DNA restriction site analysis, and three outgroups. Phylogenetic analysis supported the monophyly of Silphium with Lindheimera as sister. Silphium can be divided into two sections based upon two well-supported clades that correspond to root type and growth form. These results also supported the expansion of subtribe Engelmanniinae to include Balsamorhiza, Borrichia, Rojasianthe, Vigethia, and Wyethia. We hypothesize that subtribe Engelmanniinae originated in Mesoamerica and later radiated to the United States. We suggest that the cypsela complex, which is present in Berlandiera, Chrysogonum, Engelmannia, and Lindheimera, arose only once and was subsequently lost in Silphium.     

Wood anatomy of the tribe Podalyrieae (Fabaceae,Papilionoideae): Diversity and evolutionary trends

Detailed wood anatomical data for 32 species from all nine genera of the tribe Podalyrieae are presented, together with numerical analyses and the mapping of character states onto the latest available molecular phylogeny. It was found that trees (Cadia, Calpurnia and Virgilia) have vessels in small isolated groups, whilst fynbos shrubs (the remaining genera: Amphithalea, Cyclopia, Liparia, Podalyria, Stirtonanthus and Xiphotheca) commonly show highly grouped narrow vessels (frequently in a dendritic pattern), and helical thickening on the vessel walls. Comparisons of the main character state changes with the molecular phylogeny of the tribe show that the wood structure of trees probably represents the basic condition in the tribe; character states present in shrubs appear to have arisen a few times and very likely represent adaptations to seasonal water stress. In general, the wood anatomy is congruent with current subtribal and generic delimitations. Fire-survival strategy is reflected in the rays, with seeders having mostly procumbent cells whilst sprouters have square and upright cells. The close similarity in wood anatomy between Cadia and Calpurnia is in agreement with the transfer of Cadia to the Podalyrieae. A remarkable diversity of crystals was found, including prismatic, acicular and navicular crystals, the last two of which may occur singly or in sheaf-like aggregates.     

Phylogenetic relationships in the tribe Oxyptilini (Lepidoptera,Pterophoridae, Pterophorinae) based on morphological data of adults

The monophyly of the tribe Oxyptilini and phylogenetic relationships of the genera embraced in this tribe were examined using 171 (75 binary and 96 multistate) characters of adult morphology. The study material included 98 species of 30 genera, representing all previously recognized genera of Oxyptilini, together with the genera Sphenarches, Antarches, Diacrotricha, and Cosmoclostis, four species of Oidaematophorini, three species of Platyptiliini, as well as three and two other species belonging to Pterophorini and Exelastini respectively. Two Agdistis species were used as outgroups. The cladistic analysis resulted in six equally parsimonious trees. A majority of the recovered synapomorphic characters have previously been used in the taxonomy of the subfamily. However, 25 novel characters were found. The monophyly of Oxyptilini was supported, although only with homoplastic characters and low amounts of tree confidence; the genera Capperia, Procapperia, Paracapperia, Oxyptilus, Megalorhipida, and Trichoptilus were found to be nonmonophyletic; Sphenarches and Antarches were recovered as members of Oxyptilini; the two genera Cosmoclostis and Diacrotricha were placed out of Oxyptilini, inside the tribe Pterophorini; and close affinity of the genus Dejongia to Stangeia, Stenodacma, Megalorhipida, Trichoptilus, and Buckleria species was revealed. Four new combinations, Cosmoclostis lanceata (Arenberger) comb. nov. , Nippoptilia regulus (Meyrick) comb. nov. , Capperia tadzhica (Zagulajev) comb. nov. , and Buckleria negotiosus (Meyrick) comb. nov. are proposed; Capperia insomnis Townsend was considered as a senior synonym of Procapperia hackeri Arenberger syn. nov. , Buckleria negotiosus (Meyrick) as a senior synonym of Buckleria vanderwolfi Gielis syn. nov. , and Oxyptilus variegatus Meyrick syn. nov. as a junior synonym of Oxyptilus secutor Meyrick. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 484–547.     

Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological andndhF sequence data

Phylogenetic relationships within the tribe Antirrhineae (Scrophulariaceae) are analysed and discussed on the basis of parsimony analyses of morphological andndhF gene sequence data. The results indicate that the tribe Antirrhineae consists of four major groups of genera, theAnarrhinum clade, theGambelia clade, theMaurandya clade, and theAntirrhinum clade. TheAnarrhinum clade, consisting of the Old World bee-pollinated generaAnarrhinum andKickxia, is sister to the rest of the tribe. TheGambelia clade consists of the New World generaGambelia andGalvezia, which are very closely related and pollinated by hummingbirds. TheMaurandya clade consists of one subclade includingMaurandya and a number of related bee- or hummingbird-pollinated New World genera and another subclade with the Old World bee-pollinated generaAsarina andCymbalaria. TheAntirrhinum clade consists mainly of bee-pollinated Old World genera, such asAntirrhinum, Linaria, Chaenorhinum, and their segregates, but also includes the New World generaMohavea andHowelliella, of which the latter is known to be partly pollinated by hummingbirds. It is concluded that hummingbirdpollination has evolved independently within Antirrhineae at least three times from bee-pollinated ancestors.     

Molecular data reveal convergence in fruit characters used in the classification of Thlaspi s. I. (Brassicaceae)

Phylogenetic relationships of 18 Thlaspi s.l. species were inferred from nuclear ribosomal internal transcribed spacer (ITS) sequence data. These species represent all sections of the basic classification system of Schulz primarily based on fruit characters. The molecular phylogeny supported six clades that are largely congruent with species groups recognized by Meyer on the basis of differences in seed coat anatomy, i.e. Thlaspi s. s., Thlaspkeras, Moccaea {Raparia included), Microthhspi, Vania and Neurotropy. Some of these lineages include species which are morphologically diverse in fruit shape (e.g. Thlaspi s. s.: T. arvense - fruits broadly winged, T. ceratocarpum - fruits with prominent horns at apex, T. alliaceum - fruits very narrowly winged). Furthermore, the same fruit shape type is distributed among different clades. For instance, fruits with prominent horns at apex are found in Thlaspi s. s. ( T. ceratocarpum) and Thlaspiceras (T oxyceras). These results clearly indicate convergence in fruit characters previously used for sectional classification in Thlaspi s. l.     

Taxonomic revision of Dictyochales (Dictyochophyceae) based on morphological,ultrastructural, biochemical and molecular data

The taxonomy of the siliceous members of Dictyochales has been based exclusively on the structure of their silica skeletons. In this study, other morphological, ultrastructural, pigment and molecular characters, in addition to the silica skeletons, were used in the systematics of the siliceous members of this group. As very little is known about the ultrastructure of Dictyocha octonaria, cells of both the skeleton‐bearing and naked forms were also studied. A cladistic analysis based on morphological data and a molecular phylogeny based on nuclear coded small subunit ribosomal DNA retrieved a well‐supported monophyletic Dictyochales. D. octonaria and D. speculum were resolved together with strong support. There was no support for a clade for the three species currently placed in Dictyocha; D. fibula is clearly distinct from the other two Dictyocha taxa, D. speculum and D. octonaria. It is highly likely there are two or three undescribed species within D. octonaria/D. speculum clade and two undescribed genera within the Dictyochales, based on the positions of sequences from uncultured eukaryotes present in GenBank. These findings necessitate a taxonomic revision of the three siliceous, skeleton‐bearing species. Because D. fibula is the type species of Dictyocha, we propose that D. octonaria and D. speculum be reassigned respectively to the genus Octactis as O. octonaria Hovasse and O. speculum (Ehrenberg) F. H. Chang, J. M. Grieve & J. E. Sutherland, comb. nov.     

Systematics of African lowland rainforest Praomys (Rodentia, Muridae) based on molecular and craniometrical data

Representatives of the genus Praomys occur throughout the African intertropical zone. It is unclear how many species this genus contains, nor do we know the exact distribution ranges and phylogenetic relationships of these taxa. Using molecular (16S rRNA gene sequencing) and morphological (multivariate craniometry) analyses we clarify the taxonomy and phylogenetic relationships among the Praomys occurring in Africa's lowland tropical rainforests. We studied most species known from this area, based on specimens collected in seven countries (Guinea, Ivory Coast, Cameroon, Central African Republic, Gabon, Congo Republic and Democratic Republic of Congo). In our study, Praomys appears to be monophyletic. Our results identify two species complexes: the jacksoni complex includes at least two species ( P. jacksoni and P. mutoni ) and the tullbergi complex contains at least four species ( P. tullbergi , P. rostratus , P. misonnei , P. petteri ). Although the 16S rRNA gene appears insufficient to resolve the phylogenetic relationships among all the members of the tullbergi species complex, it is suitable for the identification of most of the studied species, and its use has allowed us to redefine the geographical limits of several species.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 539–553.     

Systematics of Amaryllidaceae based on cladistic analysis of plastid sequence data

Cladistic analyses of plastid DNA sequences rbcL and trnL-F are presented separately and combined for 48 genera of Amaryllidaceae and 29 genera of related asparagalean families. The combined analysis is the most highly resolved of the three and provides good support for the monophyly of Amaryllidaceae and indicates Agapanthaceae as its sister family. Alliaceae are in turn sister to the Amaryllidaceae/Agapanthaceae clade. The origins of the family appear to be western Gondwanaland (Africa), and infrafamilial relationships are resolved along biogeographic lines. Tribe Amaryllideae, primarily South African, is sister to the rest of Amaryllidaceae; this tribe is supported by numerous morphological synapomorphies as well. The remaining two African tribes of the family, Haemantheae and Cyrtantheae, are well supported, but their position relative to the Australasian Calostemmateae and a large clade comprising the Eurasian and American genera, is not yet clear. The Eurasian and American elements of the family are each monophyletic sister clades. Internal resolution of the Eurasian clade only partially supports currently accepted tribal concepts, and few conclusions can be drawn on the relationships of the genera based on these data. A monophyletic Lycorideae (Central and East Asian) is weakly supported. Galanthus and Leucojum (Galantheae pro parte) are supported as sister genera by the bootstrap. The American clade shows a higher degree of internal resolution. Hippeastreae (minus Griffinia and Worsleya) are well supported, and Zephyranthinae are resolved as a distinct subtribe. An Andean clade marked by a chromosome number of 2n = 46 (and derivatives thereof) is resolved with weak support. The plastid DNA phylogenies are discussed in the context of biogeography and character evolution in the family.     

The phylogenetic relationships of Morgan's Sphinx, Xanthopan morganii (Walker), the tribe Acherontiini, and allied long-tongued hawkmoths (Lepidoptera: Sphingidae, Sphinginae)

A cladistic analysis is presented of the hawkmoths of the tribe Acherontiini, Morgan's Sphinx (Xanthopan morganii (Walker)), and related genera. The study aims to test the monophyly of tribe Acherontiini; the hypothesis that all taxa with extremely long probosces (some Acherontiini, Meganoton rubescens, Neococytius, Xanthopan) form a monophyletic group, or at least fall within a single reasonably compact clade; and, within this group, to determine whether Xanthopan is more closely related to Acherontiini or to Cocytius and Neococytius. The data set comprises 109 characters derived from adult and immature stage morphology, biology and behaviour. These data were analysed using equal weighting, successive approximations character weighting (SACW) and implied weighting. All weighting schemes agreed on the monophyly of Acherontiini and of a group of genera comprising Amphimoea, Cocytius and Neococytius (the Cocytius group). Several other generic and suprageneric clades were also consistently recovered. However, those hawkmoths with extremely long probosces were never recovered as a monophyletic group. The relationships of Xanthopan were also ambiguous. Equal weighting and SACW placed Xanthopan + Meganoton rubescens (Butler) as sister to the Cocytius group, while implied weighting placed Xanthopan as sister to Acherontiini. This latter relationship is based primarily on shared possession of a pilifer/palp hearing organ. Further analyses suggested the two components of this organ were not biologically independent. Downweighting this feature accordingly resulted in all weighting schemes converging on the topology found by equal weighting. Exclusion of the incomplete subset of immature stage data had no effect under implied weighting but equal weighting and SACW now recovered a Neotropical clade comprising Manduca and the Cocytius group, while Xanthopan was placed with M. rubescens and Panogena. Downweighting the pilifer/palp hearing organ under implied weighting again caused convergence with the equal weighting/SACW results. Thus, the relationships of Xanthopan remain equivocal and further data, particularly from the immature stages, will be required to elucidate its phylogenetic position further. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society, 2002, 135 , 471–527.     

Phylogeny of Ptychostomum （Bryaceae,Musci） inferred from sequences of nuclear ribosomal DNA internal transcribed spacer （ITS） and chloroplast rps4

The phylogeny of Ptychostomum was first spacer （ITS） region of the nuclear ribosomal （nr） DNA DNA rps4 sequences. Maximum parsimony, maximum undertaken based on analysis of the internal transcribed and by combining data from nrDNA ITS and chloroplast likelihood, and Bayesian analyses all support the conclusion that the reinstated genus Ptychostomum is not monophyletic. Ptychostomum funkii （Schwagr.） J. R. Spence （≡ Bryum funkii Schwaigr.） is placed within a clade containing the type species of Bryum, B. argenteum Hedw. The remaining members of Ptychostomum investigated in the present study constitute another well-supported clade. The results are congruent with previous molecular analyses. On the basis of phylogenetic evidence, we agree with transferring B. amblyodon Mull. Hal. （≡ B. inclinatum （Brid.） Turton≡ Bryum archangelicum Bruch ＆ Schimp.）, Bryum lonchocaulon Mull. Hal., Bryum pallescens Schleich. ex Schwaigr., and Bryum pallens Sw. to Ptychostomum.     

Cladistic analysis of the Stephaniellaceae (Marchantiophyta) based on morphological data

This work evaluates the phylogenetic relationships between Stephaniellaceae, Arnelliaceae, Southbyaceae, and Gymnomitriaceae, and between genera and species of the Stephaniellaceae. We analysed morphological data obtained from herbarium specimens and from the literature under the parsimony criterion using equal weightings and implied weightings. The inclusion of Stephaniella and Stephaniellidium within Arnelliaceae, Southbyaceae or Gymnomitriaceae is not supported in this analysis. The clade Stephaniellaceae had a moderately high Jackknife support and the family was characterized by the following synapomorphies: presence of paraphyllia and presence of longitudinal folds in female bracts. Within the Stephaniellaceae, Stephaniella is corroborated as monophyletic and Stephaniellidium is resolved as its sister group. The relationship between Stephaniella uncifolia Winkler and Stephaniella hamata Stephani was strongly supported in both analyses while the relationship between Stephaniella paraphyllina Jack and Stephaniella boliviensis Stephani was weakly supported. With these results, S. boliviensis is proposed as a synonym of S. paraphyllina. We propose changes in the taxonomic position of Stephaniella and Stephaniellidium and in their classification in consideration of the complex morphology of the species and the lack of sexual structures and collections.     

Molecular phylogeny of Phalaenopsis Blume (Orchidaceae) based on the internal transcribed spacer of the nuclear ribosomal DNA

The internal transcribed spacer (ITS1, 5.8S rDNA, and ITS2) region of nuclear ribosomal DNA (nrDNA) was sequenced from 53 species, which represent most of the living species diversity in the genus Phalaenopsis (Orchidaceae). A phylogeny was developed for the genus based on the neighbor-joining and maximum parsimony analyses of molecular data. Results of these analyses provided support for the monophyly of the genus Phalaenopsis and concurred in that the genera Doritis and Kingidium should be treated as being parts of the genus Phalaenopsis as suggested by Christenson (2001). Within the genus Phalaenopsis, neither subgenera Aphyllae nor Parishianae were monophyletic, and they were highly clustered with subgenus Proboscidioides plus sections Esmeralda and Deliciosae of subgenus Phalaenopsis based on ITS data. Those species also have the same characters of morphology of four pollinia and similar biogeographies. Furthermore, neither subgenus Phalaenopsis nor Polychilos was monophyletic. Within the subgenus Phalaenopsis, only section Phalaenopsis was highly supported as being monophyletic. As for the subgenus Polychilos, only section Polychilos was moderately supported as being monophyletic. In conclusion, the present molecular data obtained from the ITS sequence of nrDNA of the genus Phalaenopsis provide valuable information for elucidating the phylogeny of this genus.     

Chemotaxonomic value of anthocyanins in Podalyria and Virgilia (tribe Podalyrieae: Fabaceae)

Anthocyanin pigments responsible for purple and pink flower colours in the tribe Podalyrieae have been identified. The considerable variation in flower colour is not reflected in the chemical variation and the flower pigments are surprisingly conservative. Virgilia flowers always have the acetic acid esters of cyanidin-3-glucoside and peonidin-3-glucoside as major compounds, together with trace amounts of the coumaroyl ester of cyanidin-3-glucoside. Podalyria flowers invariably have the rather more stable coumaroyl ester of cyanidin. The data support a close affinity between the two genera but also show that flower colour is only partially homologous.     

Phylogenetic analysis of Chloraeinae (Orchidaceae) based on plastid and nuclear DNA sequences

The phylogenetic relationships of subtribe Chloraeinae, a group of terrestrial orchids endemic to southern South America, have not been satisfactorily investigated. A previous molecular phylogenetic analysis based on plastid DNA supported the monophyly of Chloraeinae and Gavilea, but showed that Chloraea is non‐monophyletic and that the sole species of Bipinnula analysed is sister to Geoblasta. However, that analysis included only 18 of the 73 species belonging to this subtribe. Here, the phylogenetic relationships of Chloraeinae were assessed by analysing aproximately 7500 bp of nucleotide sequences from nuclear ribosomal internal transcribed spacer (ITS) and plastid DNA (rbcL, matK, trnL‐trnF, rpoB‐trnC) for 42 species representing all four currently accepted genera of Chloraeinae and appropriate outgroups. Nuclear and plastid data were analysed separately and in combination using two different methods, namely parsimony and Bayesian inference. Our analyses support the monophyly of Chloraeinae and their inclusion in an expanded concept of Cranichideae, but none of the genera of Chloraeinae that includes more than one species is monophyletic. Gavilea and Bipinnula are paraphyletic, with Chloraea chica nested in Gavilea and Geoblasta penicillata in Bipinnula. As currently delimited, Chloraea is polyphyletic. The taxonomic changes proposed recently are for the most part not justifiable on phylogenetic grounds, except for recognition of the monotypic genus Correorchis. The lack of resolution for the relationships among species of ‘core’Chloraea suggests a relatively recent diversification of this group. The current generic classification is in need or revision, but additional study is advisable before carrying out further taxonomic changes. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 258–277.     

Phylogeny of Rubus (rosaceae) based on nuclear ribosomal DNA internal transcribed spacer region sequences

We used nuclear ribosomal DNA internal transcribed spacer region (ITS 1 - 5.8S - ITS 2; ITS) sequences to generate the first phylogeny of Rubus based on a large, molecular data set. We sampled 57 taxa including 20 species of subgenus Rubus (blackberries), one to seven species from each of the remaining 11 subgenera, and the monotypic and closely related Dalibarda. In Rubus, ITS sequences are most informative among subgenera, and variability is low between closely related species. Parsimony analysis indicates that Rubus plus Dalibarda form a strongly supported clade, and D. repens may nest within Rubus. Of the subgenera with more than one species sampled, only subgenus Orobatus appears monophyletic. Three large clades are strongly supported: one contains all sampled species of nine of the 12 subgenera; another includes extreme Southern Hemisphere species of subgenera Comaropsis, Dalibarda, and Lampobatus; and a third clade consists of subgenus Rubus plus R. alpinus of subgenus Lampobatus. Rubus ursinus appears to be a hybrid between a close relative of R. macraei (subgenus Idaeobatus, raspberries) and an unidentified subgenus Rubus species. ITS sequences are generally consistent with biogeography and ploidy, but traditionally important morphological characters, such as stem armature and leaf type, appear to have limited phylogenetic value in Rubus.