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Phylogram derived from Bayesian analysis, based on the chloroplast trnK-matK-psbA region, including coded length mutations. Posterior probabilities (PP) are given above and bootstrap values (BS) obtained from Maximum Parsimony analysis are shown below the branches. Probability values less than 0.5 and 50 for Bayesian and Maximum Parsimony, respectively, are not assigned. Branches interrupted by '//' represent only 50 percent of the actual substitution rate observed. The subgeneric classification within Thottea s.l. (Hou 1981) represents the generic concepts of Duchartre (1864) and Huber (1985). The recently described species from India were assigned to these three groups based on their morphology.  

Phylogram derived from Bayesian analysis, based on the chloroplast trnK-matK-psbA region, including coded length mutations. Posterior probabilities (PP) are given above and bootstrap values (BS) obtained from Maximum Parsimony analysis are shown below the branches. Probability values less than 0.5 and 50 for Bayesian and Maximum Parsimony, respectively, are not assigned. Branches interrupted by '//' represent only 50 percent of the actual substitution rate observed. The subgeneric classification within Thottea s.l. (Hou 1981) represents the generic concepts of Duchartre (1864) and Huber (1985). The recently described species from India were assigned to these three groups based on their morphology.  

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The genus Thottea comprises about 35 species distributed from India throughout Southeast Asia. However, most of the species have a narrow distribution. A first molecular phylogeny based on the chloroplast trnK intron, matK gene and trnK-psbA spacer is presented and confirms the monophyly of the genus according to Hou in 1981. Earlier subdivisions i...

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... Piperales represent the largest angiosperm order outside the eudicots and monocots. This magnoliid clade includes 16 genera with approximately 4200 species that are distributed nearly worldwide (Meng et al., 2002;Quijano-Abril et al., 2006;Wanke et al., 2006;Oelschlägel et al., 2011;Wagner et al., 2012;Frenzke et al., 2015;Sinn et al., 2018;Funez et al., 2019;Hatt et al., 2022). Members are highly diverse in growth and life forms, including parasitism (Wanke et al., 2007;Isnard et al., 2012), making it the most morphologically diverse magnoliid lineage . ...
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Background and AimsThe order Piperales has the highest diversity of growth forms among the earliest angiosperm lineages, including trees, shrubs, climbers and herbs. However, within the perianth-bearing Piperales (Asarum, Saruma, Lactoris, Hydnora, Prosopanche, Thottea and Aristolochia), climbing species only occur in the most species-rich genus Aristolochia. This study traces anatomical and morphological traits among these lineages, to detect trends in growth form evolution and developmental processes.Methods Transverse stem sections of different developmental stages of representatives of Asarum, Saruma, Lactoris, Hydnora, Thottea and Aristolochia were compared and anatomical traits were linked to growth form evolution. Biomechanical properties of representative climbers were determined in three-point bending tests and are discussed based on the anatomical observations. Growth form evolution of the perianth-bearing Piperales was reconstructed by ancestral character state reconstruction using Mesquite.Key ResultsWhile species of Asarum and Saruma are exclusively herbaceous, species of the remaining genera show a higher diversity of growth habit and anatomy. This growth form diversity is accompanied by a more complex stem anatomy and appropriate biomechanical properties. The ancestral growth form of the perianth-bearing Piperales is reconstructed with either a shrub-like or herbaceous character state, while the following three backbone nodes in the reconstruction show a shrub-like character state. Accordingly, the climbing habit most probably evolved in the ancestor of Aristolochia.Conclusions Since the ancestor of the perianth-bearing Piperales has been reconstructed with a herb- or shrub-like habit, it is proposed that the climbing habit is a derived growth form, which evolved with the diversification of Aristolochia, and might have been a key feature for its diversification. Observed anatomical synapomorphies, such as the perivascular fibres in Lactoris, Thottea and Aristolochia, support the phylogenetic relationship of several lineages within the perianth-bearing Piperales. In addition, the hypothesis that the vegetative organs of the holoparasitic Hydnoraceae are most probably rhizomes is confirmed.
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... 85 species; Saruma, monotypic) or holoparasites (Prosopanche and Hydnora), the latter with a highly modified sub-terranean bauplan (e.g. González and Stevenson, 2000;Neinhuis et al., 2005;Ohi-Toma et al., 2006;Wanke et al., 2006Wanke et al., , 2007Tennakoon et al., 2007;Oelschlägel et al., 2011;Naumann et al., 2013;Yao, 2013). ...
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Conference Paper
The predominantly tropical and subtropical genus Aristolochia is known for its large number of climbing species. In addition, comparatively few herbaceous and shrub-like species are known. While climbers and herbaceous species occur in all three clades (subgenera) named Aristolochia, Pararistolochia and Isotrema, shrub-like species have only been reported in Isotrema. Our molecular phylogenies based on chloroplast and nuclear markers indicate that these shrubs are restricted to the monophyletic Central American lineage of the subgenus Isotrema, while its Asian/North American lineage contains exclusively climbing species. Recent field work shows that the rare shrub-like species are neither strictly climbing nor fully self-supporting; they develop a more-or-less upright growth with their branches leaning on the surrounding vegetation. After comparing the anatomy and biomechanics of these species to related climbers within Isotrema, Aristolochia and Pararistolochia, we investigate whether clades that become specialized as climbers can diversify further into self-supporting growth forms. Biomechanical and anatomical investigations reveal some differences between climbing and shrub-like species, but both growth forms show several stem characters typically found in climbers. These include large cross-sectional areas of ray parenchyma and a thick periderm, which are known to promote flexibility. Our study suggests that evolution of growth forms within the genus Aristolochia might be constrained by organizations favoring the climbing habit where shrub-like species do not develop the height, size and stability generally typical of self-supporting shrubs and trees. The insights into growth form evolution within the genus Aristolochia and Isotrema in particular, have encouraged us to enlarge the study on closely related clades. These include Thottea, sister genus to Aristolochia, which contains exclusively shrub-like species, as well as the monotypic genus Lactoris including the small shrub Lactoris fernandeziana, and the holoparasitic genus Hydnora. The latter clades have both been recently placed in close relationship to the genera Aristolochia and Thottea. By studying the anatomy and biomechanics of these lineages in a comparative approach, we intend to enlarge the knowledge about growth form evolution in Piperales.