Evolution,Extinction,Homology and Homoplasy of the Larger Benthic Foraminifera from the Carboniferous to the Present Day,as Exemplified by Planispiral-Fusiform and Discoidal Forms

Examples of evolution,extinction and homoplasy of the larger benthic foraminifera(LBF)occur throughout their history.Since the Carboniferous,LBF have thrived in carbonate-rich tropical and subtropical shallow-marine shelf environments.Their high abundance and diversity are due primarily to their extraordinary ability to inhabit a range of ecological niches and by hosting a variety of symbionts.Attaining relatively large,centimetre-scale sizes,made some forms very specialized and vulnerable to rapid ecological changes.For this reason,some LBF have shown a tendency to suffer periodically during major extinctions,especially when environmental conditions have changed rapidly and/or substantially.This,however,makes them valuable biostratigraphic microfossils and,in addition,gives invaluable insight into the spatial and temporal process of biological evolution,such as convergent/homoplasy and homology/iterative evolution.Here the evolutionary behavior of two important morphological types that occurred throughout the history of the LBF are discussed,namely the planispiral-fusiform test as typified by the fusulinids in the Late Paleozoic and the alveolinids in the Mid-Cretaceous and Neogene,and the three-layered discoid lenticular test as characterized by the orbitoids in the Mid-to Late Cretaceous,the orthophragminids in the Paleogene,and lepidocyclinids in the Oligocene to Quaternary.Understanding the propensity of these forms to convergent and iterative evolution,with the repeated re-occurrence of certain morphological features,is essential in understanding and constructing their phylogenetic relationships more generally within the main groups of the LBF.The insights gained from the history of these LBF have wider implications,and provide a more general understanding of the impacts of climate and ecological changes as driving forces for biological evolution.

Phylogeny,character evolution,and classification of Selaginellaceae(lycophytes)

Selaginella is the largest and most taxonomically complex genus in lycophytes.The fact that over 750 species are currently treated in a single genus makes Selaginellales/Selaginellaceae unique in pteridophytes.Here we assembled a dataset of six existing and newly sampled plastid and nuclear loci with a total of 684 accessions(74%increase of the earlier largest sampling)representing ca.300 species to infer a new phylogeny.The evolution of 10 morphological characters is studied in the new phylogenetic context.Our major results include:(1)the nuclear and plastid phylogenies are congruent with each other and combined analysis well resolved and strongly supported the relationships of all but two major clades;(2)the Sinensis group is resolved as sister to S.subg.Pulviniella with strong support in two of the three analyses;(3)most morphological characters are highly homoplasious but some characters alone or combinations of characters well define the major clades in the family;and(4)an infrafamilial classification of Selaginellaceae is proposed and the currently defined Selaginella s.l.is split into seven subfamilies(corresponding to the current six subgenera t the Sinensis group)and 19 genera(the major diagnosable clades)with nine new species-poor genera.We support the conservation of Selaginella with a new type,S.flabellata,to minimize nomenclatural instability.We provide a key to subfamilies and genera,images illustrating their morphology,their morphological and geographical synopses,a list of constituent species,and necessary new combinations.This new classification will hopefully facilitate communication,promote further studies,and help conservation.