Nomenclature in the dock. Overclassification leads to instability: a case study in the horticulturally important genus Cyclamen (Myrsinaceae)

Nine different classifications have been produced in the last 70 years for the horticulturally valuable genus Cyclamen , a small genus with fewer than 30 species. These classifications, generated by intuitive methods and cladistic analyses, incorporated a total of four infrageneric ranks above that of species and were based on data from morphology, cytology and DNA sequencing. Our results, based on cladistic analyses of three independent data sources − nrDNA ITS, cpDNA trn L intron and morphological data − reveal good resolution only in nrDNA sequence data. However, when these three data sources are combined they provide stronger resolution and support for three major clades, only one of which, subgenus Psilanthum , has been consistently supported in previous classifications. The differing infrageneric classifications produced in Cyclamen result from varying taxon sampling, differing interpretation of morphological data, changes in the sources and analysis of data, and inconsistent application of names. Extensive subdivision of small genera in the absence of adequate data that could provide evidence for consistent patterns of relationship is premature and leads to a proliferation of names.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 339–349.     

Morphological differentiation among closely related species with disjunct distributions: a case study of Mediterranean Cyclamen L. subgen. Psilanthum Schwarz (Primulaceae)

In Cyclamen subgenus Psilanthum , previous authors have recognized three species ( C. repandum, C. balearicum and C. creticum ), with C. repandum subdivided into three subspecies (ssp. repandum , ssp. peloponnesiacum and ssp. rhodense ), and two varieties (var. vividum and var. peloponnesiacum ) within C. repandum ssp. peloponnesiacum . To examine the validity of this classification and the degree of morphological differentiation among closely related endemic taxa with disjunct distributions, we undertook analyses of three quantitative and eight qualitative morphological traits of flowers and leaves, most of which have previously been used without statistical comparison in the classification of the different taxa. Cyclamen balearicum showed high levels of morphological differentiation from all other taxa in subgenus Psilanthum consistent with its specific status. In contrast, morphological differentiation between C. creticum and subspecies of C. repandum was similar to that among the three subspecies of C. repandum . When compared with a recent molecular phylogeny of this subgenus, our results suggest that C. creticum may best be described as a geographical subspecies of C. repandum . The two varieties of C. repandum ssp. peloponnesiacum showed few significant differences for individual traits and their overall morphology is very similar to each other. The morphology of plants at three sites on Corsica strongly suggest, consistent with molecular data, the occurrence of hybrid populations between C. balearicum (outside of its previously recognized distribution) and local C. repandum .  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 133–144.     

Relationships, classification and evolution of Scleranthus (Caryophyllaceae) as inferred from analysis of morphological characters

Study of floral anatomy, micromorphology, palynology and onotogeny has revealed new characters for phylogenetic analysis in the genus Scleranthus . Cladistic analysis of these characters, along with those previously available, suggests that the genus consists of Eurasian/Mediterranean and Australasian sister clades. Gynoecial morphology and development are closely similar in all species, suggesting the genus is monophyletic despite its disjunct northern and southern hemisphere distribution. Variation in pollen:ovule ratios and their implications for the evolution of Scleranthus species are also discussed and it is concluded that a range of breeding strategies intermediate between autogamy and xenogamy exists in the genus. Single-stamened species of Scleranthus are likely to be obligate autogams, despite their comparatively high pollen:ovule ratios in relation to autogamic species of other genera.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 15–29.     

Hymenaea mexicana sp. nov. (Leguminosae: Caesalpinioideae) from Mexican amber indicates Old World connections

Hymenaea mexicana sp. nov. is described in amber originating from mines of Chiapas, Mexico. The species is characterized by clawed showy petals with cordate bases and glabrous, verrucose ovaries with long hirsute hairs at their bases and margins on one side. This species is closely related to the extinct Hymenaea protera from Dominican amber, and together with it, is placed in the primitive section Trachylobium of the genus Hymenaea in the caesalpinoid legumes. The closest extant species to H. mexicana is H. verrucosa , a relict surviving in East Africa. These results challenge previous views that Mexican amber was produced by the South American species H. courbaril – H. intermedia , which belong to the advanced section of the genus.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 125–132.     

Genets and 'genet-like' taxa (Carnivora, Viverrinae): phylogenetic analysis, systematics and biogeographic implications

The subfamily Viverrinae is a taxon of uncertain systematic status. This study consists of cladistic analyses based on morphological characters of specimens belonging to the genera Genetta , Osbornictis , Poiana and Prionodon . Two levels of analysis are carried out, one concerning generic relationships (intergeneric analysis) and one dealing with the interrelationships of species within the genus Genetta (intrageneric analysis). In the first analysis, different outgroups were used in order to test the ingroup topology.
With regard to the intergeneric analysis, Osbornictis , Poiana and Prionodon , together with Genetta johnstoni , constitute a monophyletic group (including Nandinia ), which is the sister-group of a clade formed by the other species of genets. Thus, the genus Genetta is regarded as paraphyletic. Prionodon appears to be a derived taxon. The Poiana – Prionodon clade is well supported, especially by ultrastructural hair characters. The cladogram topology in the intrageneric analysis indicates an ecological transition from the rain forest genets to the savanna genets. This supports a rain forest origin of the genus Genetta , a conclusion which may be generalized to the entire study group. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 317–334.     

DNA phylogeny supports revised classification of Salmothymus obtusirostris

Salmothymus obtusirostris (soft-muzzled trout) is endemic to the South Adriatic drainage. Owing to its unusual appearance, which resembles both trout and grayling, it has been initially classified as a separate genus. However, this classification is ambiguous and has never been firmly established. We have studied mtDNA (control region and cytochrome b gene) and nuclear DNA (a part of LDH C*1 gene) variation between soft-muzzled trout from the upper part of the River Neretva, Bosnia and Herzegovina, and other salmonid representatives in order to examine how the current classification is congruent with molecular data. On the basis of sequence identity of mtDNA control region among several genera (i.e. Salmo , Oncorhynchus , Salvelinus , Acantholingua , Brachymystax , Thymallus and Coregonus ) a close relationship between Salmothymus , Salmo and Acantholingua was established. Phylogenetic analysis on a combined data set of mitochondrial and nuclear DNA, supported by 100% bootstraping, indicated that S. obtusirostris and A. ohridana are sister taxa which exhibit a closer relationship to S. trutta than to S. salar . This finding refutes the current classification, which recognizes S. obtusirostris as separate genus, and instead suggests its reclassification on the species level as Salmo obtusirostris .  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 399–411.     

Seed micromorphology in the genus Neotinea Rchb.f. (Orchidaceae, Orchidinae)

The ultrastructure of testa seed in the genus Neotinea (Orchidaceae, Orchidinae) was examined for the first time. The morphology of the seed and of the anticlinal and periclinal walls was analysed by scanning electron microscopy. Quantitative data concerning the length and width of the seed and embryo, seed and embryo volume, free air space, and number of cells along the longitudinal axis are presented. In all species, the seeds are fusiform in shape with transverse ridges on the inner periclinal walls. This ornamentation pattern is characteristic for the genus Neotinea . It is a good diagnostic value supporting the monophyly of this genus, which has recently been proposed by several authors.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 133–140.     

Fossil palm flowers in Dominican and Mexican amber

Twelve palm flowers in one piece of Dominican amber and a single flower in a second piece are described as Trithrinax dominicana sp. nov. (Thrinacinae: Coryphoideae), representing the first fossil record of this genus. There are no members of Trithrinax in the Greater Antilles today. Two staminate flowers in Mexican amber are described as Socratea brownii sp. nov. (Iriarteinae: Arecoideae) and represent the first fossils of this genus. A third palm flower in Mexican amber, possibly belonging to the subtribe Euterpeinae, is characterized. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 57–61.     

Checklist of Commelinaceae of Equatorial Guinea (Annobón,Bioko and Río Muni)

A checklist of Commelinaceae of Equatorial Guinea, comprising 46 taxa in 12 genera, is presented. The best represented genus is Palisota, with 11 species. Bibliographical references for Commelinaceae from Equatorial Guinea have been gathered and checked. Eleven species of Commelinaceae are recorded for the first time in the country. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 106–122.     

Phylogenetic relationships, taxonomy, character evolution and biogeography of the lacertid lizards of the genus Takydromus (Reptilia: Squamata): a molecular perspective

Historical relationships were inferred for the oriental lizards of the genus Takydromus Daudin 1802 (Lacertidae) on the basis of DNA sequences. Of the 17 species currently recognized for the genus, 13 species represented by 42 specimens from 29 localities were examined. Maximum-likelihood and maximum-parsimony analyses of data for 829 aligned sites from parts of the mitochondrial 12S and 16S rRNA genes yielded relationships that, while showing no substantial discrepancy with each other, were strikingly different from a currently prevailing phylogenetic hypothesis from a parsimony analysis of morphological characters. Based on the results of these molecular analyses, supplemented by results of the morphological analysis that offered robust evidence for positions of two additional species ( T. khasiensis and T. sylvaticus ), the following interrelationships were hypothesized as the most preferred phylogeny: ( kuehnei ( sexlineatus khasiensis ))( tachydromoides (( smaragdinus ( sauteri ( dorsalis sylvaticus ))) ( amurensis ((( formosanus wolteri ) hsuehshanensis )( toyamai ( septentrionalis stejnegeri )))))). These interrelationships indicate: (1) invalidity of Platyplacopus Boulenger 1917 , which was recently resurrected as a subgenus of Takydromus on the basis of the morphological analysis; (2) homoplasy in states of some morphological characters, such as green dorsal coloration, that were considered as synapomorphs of certain nodes in the morphological analysis; and (3) involvement of the genus in a series of vicariances in both the continental and insular parts of eastern Eurasia. Due to the paucity of available samples, phylogenetic status of T. intermedius and T. haughtonianus remain to be examined in future studies.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 76 , 493–509.     

Cytogenetic studies on Brazilian Xyris species (Xyridaceae)

Cytological features are presented for the first time for 12 species and one natural hybrid from 16 Brazilian populations of the genus Xyris , including members of two sections ( Xyris and Nematopus ). The results are compared with a review of all available chromosome numbers previously reported for the genus. All analysed species show semi-reticulate interphase nuclei and prophase to prometaphase chromosomes with heterogeneous condensing behaviour. Chromosome numbers varied between 2 n = 26 and 2 n ≈ 108. In Brazil especially, the karyological differentiation of Xyris seems to be generally characterized by dysploid and polyploid increase of the chromosome number, possibly after interspecific hybridization. This is the first cytological study including species of the section Nematopus and the first karyological investigation including South American species of the genus Xyris . © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 245–252.     

Karyological studies in Tripleurospermum (Asteraceae, Anthemideae) from north-east Anatolia

Chromosome number and morphology in 14 taxa belonging to 19 populations of Tripleurospermum Sch. Bip. were studied using karyological and numerical taxonomical techniques. Data on chromosome measurements were analysed using cluster analysis. Chromosome numbers of these taxa are 2 n  = 2 x  = 18, 4 x  = 36 and 5 x  = 42–48. Seven records are new, two are not consistent with previous counts, and the remainder confirm the very limited previous data (one to three records). A new ploidy level (pentaploidy) is reported for the first time for the genus. Some correlations between ploidy levels and morphological characters are noted and several systematic and evolutionary aspects of the genus are discussed in the light of karyological data.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 146 , 427–438.     

Multiple glacial refugia and postglacial colonization routes inferred for a woodland geophyte,Cyclamen purpurascens: patterns concordant with the Pleistocene history of broadleaved and coniferous tree species

The phylogeographical history of mid‐altitude woodland herbs that depend on moist and shaded forest habitats is poorly understood. Here, we analysed the genetic structure of Cyclamen purpurascens, a mountainous calcicolous perennial, to test hypotheses regarding its glacial survival in single or multiple refugia and postglacial colonization routes, and to explore how they are congruent with the histories inferred for temperate trees and other mountainous herbs. We gathered AFLP data and chloroplast DNA sequences (trnD‐trnT region) from 68 populations spanning the entire distribution range (the Jura Mountains, Alps, western Carpathians, Dinarides). Both genetic markers revealed two main phylogeographical groups (phylogroups) in C. purpurascens. Additionally, AFLP data detected a more detailed structure of five phylogroups: two widespread, showing east?west geographical separation, and three local ones, restricted to somewhat disjunct, marginal regions of the species range. We suggest that C. purpurascens survived the last glaciation in two main regions, the foothills of the Southern Limestone Alps and the Karst area of the north‐western Dinarides, and possibly also in microrefugia in the Western Carpathians. The glacial persistence and colonization routes of this woodland herb are highly concordant with those inferred for several temperate trees, especially the European beech. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 741–760.     

Mitochondrial genes: signals and noise in the phylogenetic reconstruction of the annual killifish genus Cynolebias (Cyprinodontiformes, Rivulidae)

The genus Cynolebias (Cyprinodontiformes: Rivulidae) is a locally endemic and speciose group of Neotropical fishes with an annual life cycle. Members of the genus vary greatly in morphology and behaviour, and extensive interspecific karyotypic divergence has been documented among species from Uruguay, Argentina and Rio Grande do Sul, Brazil. We present a molecular systematic hypothesis of the relationships between these Cynolebias species based on phylogenetic analysis of a combined dataset containing 1825 base pairs of DNA sequence, representing three mitochondrial genes. The protein-coding cytochrome -b gene, the 12S and 16S rRNA mitochondrial genes, alone and in combination, yield robust support for monophyly within Cynolebias . Furthermore, our analyses identify two major Cynolebias clades, one of which contains at least four monophyletic groups. Corrected mtDNA genetic distances range from 5.2 to 17.5% between Cynolebias species, and application of a molecular clock suggests the occurrence of two pulses of cladogenesis, one in the late Miocene and another in the Pliocene–Pleistocene. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 49–59.     

Biogeography and pattern of gene flow among Barbus species (Teleostei: Cyprinidae) inhabiting the Italian Peninsula and neighbouring Adriatic drainages as revealed by allozyme and mitochondrial sequence data

Genetic relationships among 24 Italian and Adriatic populations of barbs (genus Barbus ) were assessed using electrophoretic analysis of allozymes and mitochondrial DNA sequences of the cytochrome b gene. Results obtained with both markers were concordant, but they were not congruent with the current morphology-based systematics and taxonomy. Populations assigned to the same nominal taxa ( B. caninus , B. petenyi and B. rebeli ), were very divergent in both allozymes and mtDNA, indicating that these populations deserve recognition as different units for conservation and management. On the other hand, the two fluvio-lacustrine taxa considered as distinct species (i.e. B. plebejus and B. tyberinus ) are genetically very close to each other, showing no clear differences at either allozymes or mtDNA. The population of B. caninus from Pellice River carried allozyme alleles and mtDNA specific for B. plebejus , indicating a genetic introgression towards the former species. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 83–99.     

Morphological and pollen differentiation in Solanum cardiophyllum ssp. cardiophyllum and S. cardiophyllum ssp. ehrenbergii

A multivariate analysis to differentiate morphologically the populations of wild potatoes Solanum cardiophyllum ssp. cardiophyllum and S. cardiophyllum ssp. ehrenbergii was carried out. An analysis of the morphology and the viability of pollen of these potato populations was also made. The results of the morphometric analysis indicate that both subspecies are phenetically different. The pollen grain shape and size in ssp. cardiophyllum are different in northern and southern populations. Pollen diameter is significantly different between subspecies. Based on these results we propose that these taxa should be considered as two separate species.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 415–426.