A molecular phylogeny and a revised classification of tribe Lepisoreae (Polypodiaceae) based on an analysis of four plastid DNA regions

Phylogenetic relationships within the palaeotropical tribe Lepisoroideae (Polypodiaceae) were investigated by studying sequence variation of four plastid DNA regions: rbcL, rps4 plus rps4‐trnS IGS, trnL intron plus trnL‐F IGS, rbcL‐atpB IGS plus part of atpB. In total, over 4000 nucleotides were sequenced for 39 species. Seven well‐supported clades were found in the analyses of the combined data set. We provide a new classification of Lepisoroideae by integrating phylogenetic results and known variation of morphological characters. The two small genera Neocheiropteris and Tricholepidium are supported as monophyletic, the genus Paragramma is resurrected and the genera Lepisorus, Neolepisorus, Lemmaphyllum and Lepidomicrosorium are re‐circumscribed. We proposed 14 new combinations, among which Caobangia is treated as a synonym of Lemmaphyllum. A key for identifying the recognized genera is presented. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 28–38.     

Cranial osteology and phylogenetic position of the theropod dinosaur Proceratosaurus bradleyi (Woodward, 1910) from the Middle Jurassic of England

The cranial osteology of the small theropod dinosaur Proceratosaurus from the Bathonian of Minchinhampton, England, is described in detail, based on new preparation and computed tomography (CT) scan images of the type, and only known, specimen. Proceratosaurus is an unusual theropod with markedly enlarged external nares and a cranial crest starting at the premaxillary–nasal junction. The skull is highly pneumatic, with pneumatized nasals, jugals, and maxillae, as well as a highly pneumatic braincase, featuring basisphenoid, anterior tympanic, basipterygoid, and carotid recesses. The dentition is unusual, with small premaxillary teeth and much larger lateral teeth, with a pronounced size difference of the serrations between the mesial and distal carina. The first dentary tooth is somewhat procumbent and flexed anteriorly. Phylogenetic analysis places Proceratosaurus in the Tyrannosauroidea, in a monophyletic clade Proceratosauridae, together with the Oxfordian Chinese taxon Guanlong. The Bathonian age of Proceratosaurus extends the origin of all clades of basal coelurosaurs back into the Middle Jurassic, and provides evidence for an early, Laurasia‐wide, dispersal of the Tyrannosauroidea during the late Middle to Late Jurassic. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009.     

The potential role of chromosome telomere resetting consequent upon sex in the population dynamics of aphids: an hypothesis

Models of population structure have emphasized the importance of sex in maintaining lineages. This is because, despite the well known ‘two‐fold cost of sex’ compared with asex, it is considered that recombination rids the genome of accumulated mutations and increases its potential for adaptive variation. However, asexual lineages of eukaryotic organisms can also rapidly gain genetic variance directly by various mutational processes, thereby proving that so‐called ‘clones’ do not have strict genetic fidelity ( Lushai & Loxdale, 2002 ; Loxdale & Lushai, 2003a ), whereas the variation so produced may well have adaptive advantage during the evolutionary process. This being so, obligated asexuals or cyclical parthenogens that occasionally indulge in sexual recombination (‘rare sex’) cannot be deemed as ‘evolutionary dead‐ends’( Lushai, Loxdale & Allen, 2003a ). In addition, the persistence of asexual lineages (i.e. lineage longevity) may also involve the integrity of the telomere region, the physical end of the chromosomes ( Loxdale & Lushai, 2003b ). In this earlier study on this topic, we argued that the persistence and ultimate senescence of eukaryotic cell lineages (based upon the frequency of ‘capped’ and ‘uncapped’ chromosomes related to telomere functionality; Blackburn, 2000 ) may directly relate to the survival and persistence of lineages of whole asexual organisms. Aphids are a good model system to test this hypothesis because they show a variety of sexual/asexual reproductive strategies, whereas their mode of asexual reproduction is of the mitotic (= apomictic) type. We also suggested that many aphid lineages require occasional or even rare sexual recombination to re‐set telomere length to allow lineages to persist. Ample empirical evidence from diverse taxa, lineages, and different developmental stages now reveals that the telomere states are indeed re‐set by recombination (homologous or meiotic), thereby rejuvenating the lineage in question. The generational clock element of telomeric functionality has also been successfully described in artificially‐induced mammalian clonal systems. It thus appears that telomere function is a central molecular mechanism instigating and promoting lineage continuity per se. By contrast, we hypothesized that other long‐lived asexuals, or the rare category of ancient asexuals such as bdelloid rotifers, have compensatory mechanisms for maintaining chromosome functional integrity, which are somewhat different from conventional telomeric repeats. In the present study, we carry the analogy between eukaryotic cell functionality and aphid lineages a stage further. Here, we hypothesize that the changing frequency of capped and uncapped telomeres, progressing to senescence in a stochastic manner, may be an underlying factor that significantly contributes to population dynamics in asexual lineage evolution. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 719–728.     

On the synonymy of Delamarella Chappuis and Latiremus Božić (Copepoda, Harpacticoida, Latiremidae), including the description of D. obscura sp. nov. from the Black Sea

A new species of Delamarella Chappuis, 1954 (Copepoda, Harpacticoida, Latiremidae), D. obscura sp. nov. , is described from sandy beaches along the western Black Sea coast of Turkey, using both light and scanning electron microscopy. Additional morphological observations of the closely related D. galateae Cottarelli, 1971 are presented, based on topotype material from Sardinia and newly collected specimens from mainland Italy. Previous records of D. karamani Petkovski, 1957 from the Bulgarian Black Sea coast are based on misidentifications and should be attributed to D. obscura sp. nov. All three species are morphologically similar, and criteria based on meristic and ornamentation characters are provided to distinguish them. Examination of copepodid V ♂ intermoult stages provided new insights into the morphology of the highly transformed P4 exopod of the adult male and its homology with reference to the female. A reassessment of the generic distinctiveness of Delamarella and the monotypic genus Latiremus Božić, 1969 revealed that morphological criteria, traditionally employed to separate both genera, are all essentially unsound and based on erroneous reports in the literature. Consequently, Latiremus is relegated to a junior synonym of the type genus, leaving Delamarella and Arbutifera Huys & Kunz, 1988 as the only remaining valid genera in the family.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 263–281.     

Logistic regression in comparative wood anatomy: tracheid types, wood anatomical terminology, and new inferences from the Carlquist and Hoekman southern Californian data set

Despite collecting copious amounts of data, wood anatomists rarely perform appropriate statistical analyses, especially in the case of categorical variables. Nevertheless, anatomists have succeeded in identifying strong ecological trends. We show that, with only a slightly more sophisticated analysis, the strength and significance of 'well-known' associations can be quantified, and new associations pinpointed. Using logistic regression to reanalyse the classic Carlquist and Hoekman data set for the southern Californian flora, we show strong support for the notion that true tracheid presence lowers vessel grouping; in contrast, vasicentric tracheids are associated with a diversity of vessel grouping strategies. We show that statistical models can refine anatomical interpretations by identifying unusual species. For example, Fremontodendron californicum and Baccharis salicifolia (=  B. glutinosa ) were identified as unusual in lacking vasicentric tracheids; a consultation of preparations revealed that they are indeed present. For purposes of ecological wood anatomy, anatomical terminology should reflect cell function; we suggest that terminological systems that yield better predictive power in statistical models such as ours are preferable. Finally, we make recommendations ranging from the statistical, e.g. the need to check assumptions and the need for the inclusion of phylogeny, to the biological, e.g. gathering data expressly designed to test functional hypotheses rather than all of the information in standardized lists.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 331–351.     

A taxonomic revision of the genus Flagenium Baill. (Rubiaceae–Octotropideae)

A taxonomic revision of the genus Flagenium Baill. is presented. Flagenium is endemic to Madagascar and comprises six species; three new species are described here ( F. farafanganensis , F. petrikensis , and F. pedunculatum ) and one species ( F. arboreum ) is transferred into synonymy. Each species is fully described, and summaries of distribution, habitat and ecology, and phenology are given; conservation assessments are provided for each species. Flagenium is characterized by having ovaries with at least two erect and two pendulous ovules per locule, which, depending on the species, can display an additional two to four horizontal ovules per locule arranged between the uppermost and lowermost ovules. In addition, Flagenium has a longitudinally ten-ribbed fruit. These features are potentially unique within the Octotropideae.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 557–570.     

Phylogeny and biogeography of African Biomphalaria (Gastropoda: Planorbidae), with emphasis on endemic species of the great East African lakes

The pulmonate snails of the genus Biomphalaria are widely distributed in the tropics, and they are intermediate hosts of the digenean trematode Schistosoma mansoni that causes intestinal schistosomiasis in humans. Recent molecular evidence suggests that Biomphalaria originated in South America, and following a recent transatlantic migration colonized Africa, where it radiated into the currently recognized 12 species. In the present study we further investigate the internal phylogenetic relationships of African Biomphalaria with emphasis on the dispersal and speciation on the continent, especially in the Great Lakes in East Africa. Our results, based on 16S ribosomal DNA, cytochrome oxidase subunit I (COI), and ribosomal internal transcribed spacer I (ITS1), support the monophyly of an African clade with two separate lineages ( Biomphalaria pfeifferi / Biomphalaria camerunensis and the Nilotic species complex/ Biomphalaria angulosa ). Following the initial colonization of Africa, Biomphalaria spread towards the east where a later radiation occurred in the Lake Victoria basin and the Albertine Rift Valley Lakes. With further dispersal along the River Nile, additional speciation took place giving origin to the North-east African species Biomphalaria alexandrina . Our results present almost no support of the species groups of Mandahl-Barth (except for the pfeifferi group), which is in accordance with other molecular appraisals. Our results suggest that Biomphalaria stanleyi , which is endemic to Lake Albert, is not an ecophenotype of the continental B. pfeifferi as previously suggested by other molecular studies. B. angulosa is sequenced for the first time and it is inferred to have an important phylogenetic position as sister group to the Albertine Rift/Lake Victoria basin radiation.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 337–349.     

Floral anatomy in Dypsis (Arecaceae–Areceae): a case of complex synorganization and stamen reduction

Male (staminate) flowers of Dypsis possess either six or three stamens and/or staminodes, in contrast to most other palms, in which the basic stamen number is six (although polyandry is also common in palms). Significant variation among the tristaminate forms, both in stamen morphology and stamen position with respect to the perianth parts, indicates that stamen reduction from six to three has occurred more than once, and possibly several times within Dypsis. A few species include teratological forms with zygomorphic flowers; for example, Dypsis lantzeana normally possesses three antesepalous stamens, but in some specimens only the median outer (abaxial) stamen is expressed, perhaps indicating incipient zygomorphy correlated with complex synorganization. Inclusion of earlier historic genera such as Neophloga and Chrysalidocarpus within a broadly circumscribed Dypsis appears to be justified, although the informal taxonomic groupings within Dypsis require review, in particular the taxonomic significance of the different types of anther morphology. The discovery here that adnation of staminodes to the pistillode (complex synorganization) occurs in species other than D. mirabilis opens further questions about the taxonomic utility of this character in Dypsis, in which stamen/staminode development and adnation are apparently unusually labile. Such androecial–gynoecial adnation is otherwise rare in palms, as also in other monocots, in which probably the best‐known example occurs in orchids. Septal nectaries are present in some, but not all, staminate flowers in species of Dypsis. Dypsis bejofo is exceptional in that in staminate flowers the pistillode is distally bulbous and bears three prominent modified supralocular septal nectaries. Female (pistillate) flowers in Dypsis are syncarpous, normally pseudomonomerous (as in many other Arecoideae), and possess septal nectaries that effectively delimit the carpel margins and indicate insect pollination. There is a central solid transmitting tissue that extends from the placenta to three stylar canals. The stylar canals of the two sterile carpels are apparently functional, in addition to that of the fertile carpel. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 115?133.     

Pollen morphology of Dioscorea (Dioscoreaceae) and its relation to systematics

This paper adds new data on the pollen and orbicule morphology of 61 Dioscorea L. (Dioscoreaceae) species to the survey of Schols et al . (2001). The results indicate that pollen characters may be significant in infrageneric systematics in Dioscorea . Pollen and orbicule characters are described based on observations with light microscopy, and scanning and transmission electron microscopy, and are critically evaluated and discussed in the context of existing hypotheses of systematic relationships within the genus. Pollen is mostly disulculate (sometimes monosulcate) with a perforate, microreticulate, striate, gemmate, rugulate, or cerebroid perforate sexine. The basal section Stenophora is one of the few sections with monosulcate pollen. Brachyandra , Cardiocapsa , and Seriflorae , three Malagasy sections, are characterized by striate pollen. Pollen morphology strongly supports section Enantiophyllum as a monophyletic group. The correlation between pollen size and tuber type, as suggested previously by P. Su (1987), is confirmed by our data. As found in our earlier survey, orbicules in Dioscorea are mostly spherical with a smooth or spinulose surface.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 375–390.