Phylogenetic Analysis of Araucariaceae: Integrating Molecules, Morphology, and Fossils

Abstract

Premise of research. Phylogenetic relationships of Araucariaceae (Coniferophyta, Araucariales) are revised on the basis of the first combined data matrix for the family.Methodology. Taxon sampling includes 39 ingroup species (31 extant, 8 fossils) and outgroup species of all the remaining conifer families. Five fossil Araucaria species, one species of the genus Araucarites, and two species of the extinct genera Wairarapaia and Emwadea were included in the analyses. Character sampling includes 23 genomic regions (19 plastid, 2 nuclear, and 2 mitochondrial) and 62 morphological characters (52 discrete and 10 continuous). The phylogenetic analyses were conducted with equally weighted parsimony. Additionally, several analyses under different taxon-and gene-sampling regimes were analyzed for identifying the causes of the long-lasting controversies in the interrelationships of the three extant genera of Araucariaceeae.Pivotal results. Monophyletic Araucariaceae is the sister group of Podocarpaceae, forming the order Araucariales. Monophyly of Araucaria and Agathis is also strongly supported by the data. The results of both molecular and combined analyses indicate that Wollemia and Agathis form a clade (pagathioid clade) sister to Araucaria. Within Araucaria, the analyses support the monophyly of the four currently recognized sections: Araucaria, Bunya, Intermedia, and Eutacta. Results support the monophyly of living and fossil Araucaria (including Araucarites), whereas the remaining extinct genera are placed as the stem of the agathioid clade. In terms of the sensitivity analyses performed, results suggest that inconsistencies among previous results would be related to ingroup sampling.Conclusions. By means of a combined phylogenetic analysis, we have been able to obtain a strongly supported and well-resolved phylogeny of Araucariaceae that includes both living species and fossil species for the group. This study shows the feasibility and usefulness of phylogenetic analyses that incorporate multiple sources of evidence (molecules/morphology, living/fossil species, discrete/continuous characters).

First South AmericanAgathis(Araucariaceae), Eocene of Patagonia

“…Numerous recent papers have produced variable results for relationships within Agathis (e.g., Gilmore and Hill, 1997 ;Setoguchi et al, 1998 ;Stefanović et al, 1998 ;Conran et al, 2000 ;Quinn et al, 2002 ;Stöckler et al, 2002 ;Rai et al, 2008 ;Liu et al, 2009 ;Biffi n et al, 2010 ;Leslie et al, 2012 ;Escapa and Catalano, 2013 ). Critically, the living species that we found to have greatest morphological similarity to the fossils ( A. lenticula ; see Discussion) has never been analyzed in a phylogenetic context with either morphological or molecular data.…”

Fluorescent Banding Pattern of Chromosomes in <i>Araucaria araucana</i>, Araucariaceae

“…angustifolia, with restricted distribution in areas of South America. The South American species were put into the same clade by molecular phylogenetic analysis indicating very closely related species (Setoguchi et al 1998, Escapa andCatalano 2013) and had a different genome size from species distributed in other areas (Zonneveld 2012). These species have very similar karyotypes with a long secondary constriction or large CMA-band.…”

Araucarian leaves and cone scales from the Loreto Formation of Río de Las Minas, Magellan Region, Chile

“…Combined cladistics analysis of Araucariaceae by Escapa and Catalano (2013) clarified monophyly of Eutacta. Although only fossil permineralized cones were used for their analysis, they revealed 5 of 62 scored characters as unambiguous synapomorphies of the section.…”

“…Section Araucaria is represented by A. cartellei Duarte from the Lower Cretaceous of Brazil (Duarte 1993), Araucaria grandifolia Del Fueyo and Archangelsky from the Lower Cretaceous of Argentina (Del Fueyo and Archangelsky 2002), A. pararaucana Panti et al (2007) from the Paleogene of Argentina, and A. nathorstii Dusén (1899) from the Paleogene-Noegene of Argentina and Chile (Berry 1928;Doktor et al 1996;Fiori 1939;Hünicken 1995;Menéndez and Caccavari 1966). Section Bunya is represented by the seed cones Araucaria mirabilis (Spegazzini) Windhausen (1931) from the Jurassic of Argentina (Stockey 1978), although its relationship to section Bunya was not supported by recent phylogenetic analyses based on molecular and morphological data (Escapa and Catalano 2013). Section Eutacta is represented by cone-scales described as Araucarites species; A. minimus Archangelsky (1966), A. baqueroensis Archangelsky (1966), and A. chilensis Baldoni (1979) from the Lower Cretaceous of Argentina and Chile, and Araucaria pichileufulensis Berry (1938) from the Eocene of Argentina.…”