Annonaceae are a pantropically distributed family found predominantly in rainforests, so they are megathermal taxa, whereas Rhamnaceae are a cosmopolitan family that tend to be found in xeric regions and may be classified as mesothermal. Phylogenetic analyses of these families are presented based on rbcL and trnL-F plastid DNA sequences. Likelihood ratio tests revealed rate heterogeneity in both phylogenetic trees and they were therefore made ultrametric using non-parametric rate smoothing and penalized likelihood. Divergence times were then estimated using fossil calibration points. The historical biogeography of these families that are species rich in different biomes is discussed and compared with other published reconstructions. Rhamnaceae and most lineages within Annonaceae are too young to have had their distribution patterns influenced by break-up of previously connected Gondwanan landmasses. Contrasts in the degree of geographical structure between these two families may be explained by differences in age and dispersal capability. In both groups, long-distance dispersal appears to have played a more significant role in establishing modern patterns than had previously been assumed. Both families also contain examples of recent diversification of species-rich lineages. An understanding of the processes responsible for shaping the distribution patterns of these families has contributed to our understanding of the historical assembly of the biomes that they occupy.
The pantropical flowering plant family Annonaceae is the most species‐rich family of Magnoliales. Despite long‐standing interest in the systematics of Annonaceae, no authoritative classification has yet been published in the light of recent molecular phylogenetic analyses. Here, using the largest, most representative, molecular dataset compiled on Annonaceae to date, we present, for the first time, a robust family‐wide phylogenetic tree and subsequent classification. We used a supermatrix of up to eight plastid markers sequenced from 193 ingroup and seven outgroup species. Some of the relationships at lower taxonomic levels are poorly resolved, but deeper nodes generally receive high support. Annonaceae comprises four major clades, which are here given the taxonomic rank of subfamily. The description of Annonoideae is amended, and three new subfamilies are described: Anaxagoreoideae, Ambavioideae and Malmeoideae. In Annonoideae, seven tribes are recognized, one of which, Duguetieae, is described as new. In Malmeoideae, seven tribes are recognized, six of which are newly described: Dendrokingstonieae, Fenerivieae, Maasieae, Malmeeae, Monocarpieae and Piptostigmateae. This new subfamilial and tribal classification is discussed against the background of previous classifications and characters to recognize subfamilies are reviewed. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169, 5–40.
Aim Rain forest-restricted plant families show disjunct distributions between the three major tropical regions: South America, Africa and Asia. Explaining these disjunctions has become an important challenge in biogeography. The pantropical plant family Annonaceae is used to test hypotheses that might explain diversification and distribution patterns in tropical biota: the museum hypothesis (low extinction leading to steady accumulation of species); and dispersal between Africa and Asia via Indian rafting versus boreotropical geodispersal.Location Tropics and boreotropics.Methods Molecular age estimates were calculated using a Bayesian approach based on 83% generic sampling representing all major lineages within the family, seven chloroplast markers and two fossil calibrations. An analysis of diversification was carried out, which included lineage-through-time (LTT) plots and the calculation of diversification rates for genera and major clades. Ancestral areas were reconstructed using a maximum likelihood approach that implements the dispersal-extinction-cladogenesis model. ResultsThe LTT plots indicated a constant overall rate of diversification with low extinction rates for the family during the first 80 Ma of its existence. The highest diversification rates were inferred for several young genera such as Desmopsis, Uvariopsis and Unonopsis. A boreotropical migration route was supported over Indian rafting as the best fitting hypothesis to explain present-day distribution patterns within the family.Main conclusions Early diversification within Annonaceae fits the hypothesis of a museum model of tropical diversification, with an overall steady increase in lineages possibly due to low extinction rates. The present-day distribution of species within the two largest clades of Annonaceae is the result of two contrasting biogeographic histories. The 'long-branch clade' has been diversifying since the beginning of the Cenozoic and underwent numerous geodispersals via the boreotropics and several more recent long-distance dispersal events. In contrast, the 'short-branch clade' dispersed once into Asia via the boreotropics during the Early Miocene and further dispersal was limited.
Aim We test biogeographical hypotheses regarding the origin of Andean-centred plant groups by reconstructing phylogeny in the short-branch clade (SBC) of Annonaceae, and estimating the timing of diversifications in four apparently Andean-centred genera: Cremastosperma R.E.Fr., Klarobelia Chatrou, Malmea R.E.Fr. and Mosannona Chatrou. The SBC includes species distributed in both the Old and New World tropics. A number of the Neotropical genera display 'Andean-centred' distribution patterns, with high species richness on both sides of the Andes mountain range. In particular, we test whether these groups could have originated on the South American continent during the time frame of the Andean orogeny [from c. 23 Ma (Miocene) to the present].Methods Chloroplast DNA sequences were used to reconstruct phylogeny in related Annonaceae taxa plus outgroups, under maximum parsimony and Bayesian inference. The markers rbcL, trnL-trnF and psbA-trnH were sampled for 96 accessions to test the monophyly of each of the genera, and thus whether they might be para-or polyphyletic with respect to related groups distributed across Amazonia. To determine the sister groups of the four genera, the additional markers matK, ndhF, trnT-trnL, trnS-trnG and atpB-rbcL were sampled for 23 of the 96 accessions. Molecular dating techniques (nonparametric rate-smoothing; penalized likelihood; Bayesian inference) were then applied to estimate the age of the crown group of each genus and the age of their sister groups.
C 4 photosynthesis is a fascinating example of parallel evolution of a complex trait involving multiple genetic, biochemical and anatomical changes. It is seen as an adaptation to deleteriously high levels of photorespiration. The current scenario for C 4 evolution inferred from grasses is that it originated subsequent to the Oligocene decline in CO 2 levels, is promoted in open habitats, acts as a pre-adaptation to drought resistance, and, once gained, is not subsequently lost. We test the generality of these hypotheses using a dated phylogeny of Amaranthaceae s.l. (including Chenopodiaceae), which includes the largest number of C 4 lineages in eudicots. The oldest chenopod C 4 lineage dates back to the Eocene/ Oligocene boundary, representing one of the first origins of C 4 in plants, but still corresponding with the Oligocene decline of atmospheric CO 2 . In contrast to grasses, the rate of transitions from C 3 to C 4 is highest in ancestrally drought resistant (salt-tolerant and succulent) lineages, implying that adaptation to dry or saline habitats promoted the evolution of C 4 ; and possible reversions from C 4 to C 3 are apparent. We conclude that the paradigm established in grasses must be regarded as just one aspect of a more complex system of C 4 evolution in plants in general.
The tribe Miliuseae (Annonaceae) comprises six genera distributed in Asia: Alphonsea, Mezzettia, Miliusa, Orophea, Platymitra, and Phoenicanthus. A phylogenetic study to investigate the putative monophyly of the tribe and the intergeneric relationships is presented here. Nucleotide sequences of the plastid gene rbcL, trnL intron, and trnL-trnF intergenic spacer were analyzed from 114 Annonaceae taxa, including 24 Miliuseae species and two outgroups using maximum parsimony and Bayesian inference. The two data sets (rbcL and the trnL-trnF regions) were analyzed separately and in combination. Miliuseae were found to be polyphyletic due to the position of Mezzettia and are part of a large, predominantly Asian and Central-American clade (miliusoid clade). Although intergeneric relationships were poorly resolved, all genera, except Polyalthia, were monophyletic, supporting previous generic delimitation based on morphology. A group of three Polyalthia species seems the most likely sister group of Miliusa. Several infrageneric groups of Miliusa, Orophea, and Polyalthia are supported by both molecular and morphological data. No morphological synapomorphies have yet been found for the miliusoid clade. Molecular clades within the miliusoid clade, however, can be characterized by size and the shape of the outer petals, number of ovules per carpel, and the size of the fruits.
BackgroundThe disproportionate species richness of the world’s biodiversity hotspots could be explained by low extinction (the evolutionary “museum”) and/or high speciation (the “hot-bed”) models. We test these models using the largest of the species rich plant groups that characterise the botanically diverse Cape Floristic Region (CFR): the genus Erica L. We generate a novel phylogenetic hypothesis informed by nuclear and plastid DNA sequences of c. 60 % of the c. 800 Erica species (of which 690 are endemic to the CFR), and use this to estimate clade ages (using RELTIME; BEAST), net diversification rates (GEIGER), and shifts in rates of diversification in different areas (BAMM; MuSSE).ResultsThe diversity of Erica species in the CFR is the result of a single radiation within the last c. 15 million years. Compared to ancestral lineages in the Palearctic, the rate of speciation accelerated across Africa and Madagascar, with a further burst of speciation within the CFR that also exceeds the net diversification rates of other Cape clades.ConclusionsErica exemplifies the “hotbed” model of assemblage through recent speciation, implying that with the advent of the modern Cape a multitude of new niches opened and were successively occupied through local species diversification.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0764-3) contains supplementary material, which is available to authorized users.
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