Published phylogeny reconstructions of the palm family (Arecaceae) are based on plastid DNA sequences or restriction fragment length polymorphisms (RFLPs), nuclear DNA sequences, morphological characters or a combination thereof, and include between 33 and 90 palm species. The present study represents all previously recognized subfamilies, tribes and subtribes of palms and 161 of the 189 genera. The plastid DNA region mat K was sequenced for 178 palm species and ten commelinid monocot outgroup species, and was combined with new and previously published plastid DNA sequences of trn L-trn F, rps 16 intron and rbcL . The addition of mat K sequences and more taxa resulted in a highly resolved and largely well-supported phylogeny. Most importantly, critical basal nodes are now fully resolved and, in most cases, strongly supported. On the basis of this phylogeny, we have established a new subfamilial classification of the palms, in which five subfamilies are recognized, rather than the six that were included in the previous classification. The circumscriptions of the subfamilies Calamoideae and Nypoideae were corroborated. The phylogeny supported a new circumscription for the subfamily Coryphoideae, including all taxa previously recognized in Coryphoideae with the addition of the tribe Caryoteae, formerly of the subfamily Arecoideae. The phylogenetic analysis also supported a new delimitation for the subfamily Ceroxyloideae that contains the tribes Cyclospatheae and Ceroxyleae, and all genera formerly included in the subfamily Phytelephantoideae, but excludes the tribe Hyophorbeae. Finally, the subfamily Arecoideae was modified to exclude the tribe Caryoteae and to include the tribe Hyophorbeae.
Phylogenetic relationships were determined in the Araucariaceae, which are now found mainly in the Southern Hemisphere. This conifer family was well diversified and widely distributed in both hemispheres during the Mesozoic era. The sequence of 1322 bases of the rbcL gene of cpDNA was determined from 29 species of Araucariaceae, representing almost all the species of the family. Phylogenetic trees determined by the parsimony method indicate that Araucariaceae are well defined by rbcL sequences and also that the monophyly of Agathis or Araucaria is well supported by high bootstrap values. The topology of these trees revealed that Wollemia had derived prior to Agathis and Araucaria. The rbcL phylogeny agrees well with the present recognition of four sections within Araucaria: Araucaria, Bunya, Eutacta, and Intermedia. Morphological characteristics of the number of cotyledons, position of male cone, and cuticular micromorphologies were evaluated as being phylogenetically informative. Section Bunya was found to be derived rather than to be the oldest taxon. Infrageneric relationships of Agathis could not be well elucidated because there are few informative site changes in the rbcL gene, suggesting the more recent differentiation of the species as their fossil records indicate. The New Caledonian Araucaria and Agathis species each formed a monophyletic group with very low differentiation in rbcL sequences among them, indicating rapid adaptive radiation to new edaphic conditions, i.e., ultramafic soils, in the post-Eocene era.
Despite the recovery of paralogues and pseudogenes in a small number of taxa, PRK and RPB2 were both highly informative, producing well-resolved phylogenetic trees with many nodes well supported by bootstrap analyses. Simultaneous analyses of the combined data sets provided additional resolution and support. Two areas of incongruence between PRK and RPB2 were strongly supported by the bootstrap relating to the placement of tribes Chamaedoreeae, Iriarteeae and Reinhardtieae; the causes of this incongruence remain uncertain. The current classification within Arecoideae was strongly supported by the present data. Of the 14 tribes and 14 sub-tribes in the classification, only five sub-tribes from tribe Areceae (Basseliniinae, Linospadicinae, Oncospermatinae, Rhopalostylidinae and Verschaffeltiinae) failed to receive support. Three major higher level clades were strongly supported: (1) the RRC clade (Roystoneeae, Reinhardtieae and Cocoseae), (2) the POS clade (Podococceae, Oranieae and Sclerospermeae) and (3) the core arecoid clade (Areceae, Euterpeae, Geonomateae, Leopoldinieae, Manicarieae and Pelagodoxeae). However, new data sources are required to elucidate ambiguities that remain in phylogenetic relationships among and within the major groups of Arecoideae, as well as within the Areceae, the largest tribe in the palm family.
SummaryWhether sex chromosomes are differentiated is an important aspect of our knowledge of dioecious plants, such as date palm (Phoenix dactylifera). In this crop plant, the female individuals produce dates, and are thus the more valuable sex. However, there is no way to identify the sex of date palm plants before reproductive age, and the sex-determining mechanism is still unclear.To identify sex-linked microsatellite markers, we surveyed a set of 52 male and 55 female genotypes representing the geographical diversity of the species.We found three genetically linked loci that are heterozygous only in males. Male-specific alleles allowed us to identify the gender in 100% of individuals. These results confirm the existence of an XY chromosomal system with a nonrecombining XY-like region in the date palm genome. The distribution of Y haplotypes in western and eastern haplogroups allowed us to trace two male ancestral paternal lineages that account for all known Y diversity in date palm.The very low diversity associated with Y haplotypes is consistent with clonal paternal transmission of a nonrecombining male-determining region. Our results establish the date palm as a biological model with one of the most ancient sex chromosomes in flowering plants.
Chloroplast DNA sequences are of great interest for population genetics and phylogenetic studies. However, only a small set of markers are commonly used. Most of them have been designed for amplification in a large range of Angiosperms and are located in the Large Single Copy (LSC). Here we developed a new set of 100 primer pairs optimized for amplification in Monocotyledons. Primer pairs amplify coding (exon) and non-coding regions (intron and intergenic spacer). They span the different chloroplast regions: 72 are located in the LSC, 13 in the Small Single Copy (SSC) and 15 in the Inverted Repeat region (IR). Amplification and sequencing were tested in 13 species of Monocotyledons: Dioscorea abyssinica, D. praehensilis, D. rotundata, D. dumetorum, D. bulbifera, Trichopus sempervirens (Dioscoreaceae), Phoenix canariensis, P. dactylifera, Astrocaryum scopatum, A. murumuru, Ceroxylon echinulatum (Arecaceae), Digitaria excilis and Pennisetum glaucum (Poaceae). The diversity found in Dioscorea, Digitaria and Pennisetum mainly corresponded to Single Nucleotide Polymorphism (SNP) while the diversity found in Arecaceae also comprises Variable Number Tandem Repeat (VNTR). We observed that the most variable loci (rps15-ycf1, rpl32-ccsA, ndhF-rpl32, ndhG-ndhI and ccsA) are located in the SSC. Through the analysis of the genetic structure of a wild-cultivated species complex in Dioscorea, we demonstrated that this new set of primers is of great interest for population genetics and we anticipate that it will also be useful for phylogeny and bar-coding studies.
A (GA)n microsatellite‐enriched library was constructed and 16 nuclear simple sequence repeat (SSR) loci were characterized in Phoenix dactylifera. Across‐taxa amplification and genotyping tests showed the utility of most SSR markers in 11 other Phoenix species and the transferability of some of them in Elaeis guineensis, 11 species of Pritchardia, Pritchardiopsis jeanneneyi and six species of Astrocaryum. The first to be published for P. dactylifera, these new SSR resources are available for cultivar identification, pedigree analysis, germplasm diversity as well as genetic mapping studies.
Bactridinae include about 150 species of spiny Neotropical palms in five genera that are ecologically important in several vegetation types such as open woodland (Acrocomia), lowland rainforest (Astrocaryum, Bactris), and montane forest (Aiphaness). The subtribe also includes the only exclusively lianescent palm genus in the Neotropics (Desmoncus). We present a fully resolved molecular phylogeny of 41 species of Bactridinae, representing all genera as well as most of the currently ac ‐ cepted infrageneric taxa (subgenera, sections etc.) and recently proposed informal groups. Analyses are based on five plastid DNA regions (matK, trnQ‐rps16, rps16 intron, trnD‐trnT, trnL‐trnF) and three nuclear markers (PRK, RPB2, ITS). A combined dataset was analysed with likelihood and parsimony methods. The results show that all accepted taxa at and above the generic level are monophyletic with high support. Astrocaryum alatum and A. mexicanum, recently segregated into a genus of their own (Hexopetion), form a strongly supported monophyletic group sister to the remaining Astrocaryum species. Desmoncus and Acrocomia are resolved as sister genera, and together they are sister to the remaining Bactridinae. This finding contrasts with that of two previous studies reporting Acrocomia to be sister to the rest of the subtribe. Aiphanes is resolved as sister to Bactris and Astrocaryum. Species‐level relationships recovered within Astrocaryum and Bactris disagree to a large extent with previous morphology‐based infrageneric classifications, suggesting that those characters are homoplasious, particularly within Bactris. A Bayesian dating analysis using the relaxed‐clock model indicates that most genera of Bactridinae diverged during a relatively short period around the Eocene–Oligocene boundary, which might explain the difficulties in resolving the phylogenetic backbone of the group. The mostly Andean genus Aiphanes shows an initial radiation of early lineages in the Oligocene (around 25 Ma ago) corresponding to an early uplift phase of the cordillera. These taxa are nowadays restricted to the mountain forests of Colombia and Ecuador. The main diversification of Andean Aiphanes began in the Miocene (around 11 Ma ago). This study provides the first substantial insight into Bactridinae phylogeny and sets the stage for more comprehensively sampled species‐level studies analysing drivers of diversity of Neotropical palms, speciation patterns, character evolution, or biogeography.
For decades, palynologists working in tropical South America are using the genus Podocarpus as a climate indicator although without referring to any modern data concerning its distribution and limiting factors. With the aim to characterize the modern and past distribution of the southern conifer Podocarpus in Brazil and to obtain new information on the distribution of the Atlantic rainforest during the Quaternary, we examined herbarium data to locate the populations of three Brazilian endemic Podocarpus species: P. sellowii, P. lambertii, and P. brasiliensis, and extracted DNA from fresh leaves from 26 populations. Our conclusions are drawn in the light of the combination of these three disciplines: botany, palynology, and genetics. We find that the modern distribution of endemic Podocarpus populations shows that they are widely dispersed in eastern Brazil, from north to south and reveals that the expansion of Podocarpus recorded in single Amazonian pollen records may have come from either western or eastern populations. Genetic analysis enabled us to delimit regional expansion: between 5° and 15° S grouping northern and central populations of P. sellowii expanded c. 16,000 years ago; between 15° and 23° S populations of either P. lambertii or sellowii expanded at different times since at least the last glaciation; and between 23° and 30° S, P. lambertii appeared during the recent expansion of the Araucaria forest. The combination of botany, pollen, and molecular analysis proved to be a rapid tool for inferring distribution borders for sparse populations and their regional evolution within tropical ecosystems. Today the refugia of rainforest communities we identified are crucial hotspots to allow the Atlantic forest to survive under unfavourable climatic conditions and, as such, offer the only possible opportunity for this type of forest to expand in the event of a future climate change.
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