Abstract
The genus Gomphrena comprises about 120 species in the Americas and 35 in Australia. Previous research revealed that Gossypianthus, Lithophila and Philoxerus are closely related but the monophyly of Gomphrena remained unresolved. Our aim was to clarify phylogenetic relationships in Gomphrena and allies based on a thorough sampling of species and to reconstruct the evolution of morphological characters including C4 photosynthesis, and to explore the disjunction of the Australian taxa. We generated datasets of plastid (matK-trnK, trnL-F, rpl16) and nrITS representing 45 taxa of Gomphrena plus relatives and analysed them with parsimony, likelihood and Bayesian methods. Ancestral states of phenotypic characters were reconstructed with BayesTraits. BEAST was employed for divergence time estimates using an extended Amaranthaceae–Chenopodiaceae dataset to place fossil calibration points. Gossypianthus is closely related to a Gomphrena radiata–G. umbellata–G. tomentosa clade and G. meyeniana, whereas Lithophila and Philoxerus appear as successive sisters of the Australian species of Gomphrena. The majority of Andean species appears in a large clade including annual and perennial species. The Cerrado species Gomphrena mollis and G. rupestris, which are C3, constitute an early-branching lineage, whereas the core Gomphrena clade is C4 and has the inner two sepals strongly compressed as synapomorphy. A major subclade evolved inflorescences with subglobose paracladia in a whorl, supported by pseudanthial leaves. Whereas the core Gomphrena clade started to diversify around 11.4 Ma (8.45–14.5 95% highest posterior density [HPD]) the Australian lineage split at only 4.8 Ma (2.61–7.18 HPD). Our detailed phylogenetic analysis of Gomphrena depicts 10 major lineages including segregate genera. We hypothesize that an adaptation to costal habitats was followed by long-distance dispersal to Australia. We also propose a revised genus concept of Gomphrena including Gossypianthus, Lithophila and Philoxerus, considering that these small segregate genera were based on states of vegetative characters exhibiting adaptations to specific habitats rather than phylogeny and overall morphology.Citation: Ortuno Limarino T. & Borsch Th. 2020: Gomphrena (Amaranthaceae, Gomphrenoideae) diversified as a C4 lineage in the New World tropics with specializations in floral and inflorescence morphology, and an escape to Australia. – Willdenowia 50: 345–381. doi: https://doi.org/10.3372/wi.50.50301Version of record first published online on 31 August 2020 ahead of inclusion in December 2020 issue.
Highlights
Gomphrena L. is one of the largest genera of the Ama ranthaceae with c. 120 species (Townsend 1993; Müller & Borsch 2005) and includes subshrubs, perennial and annual herbs, which are native in the New World tropics and subtropics (Holzhammer 1955, 1956; Eliasson 1988) and Australia (Palmer 1998)
The situation is different in Philoxerus, which cannot be separated from Gomphrena as it just differs by the general lack of stamen tube appendages or filament appendages, an androecium morphology that occurs in some Australian species of Gomphrena
All plastid trees converge on a deep split into a lineage comprising Hebanthe, Pfaffia and allies and a lineage including Gomphrena and allies, with Xerosiphon and Froelichia branching as successive sisters to the remaining taxa (Fig. 1, 2)
Summary
Gomphrena L. is one of the largest genera of the Ama ranthaceae with c. 120 species (Townsend 1993; Müller & Borsch 2005) and includes subshrubs, perennial and annual herbs, which are native in the New World tropics and subtropics (Holzhammer 1955, 1956; Eliasson 1988) and Australia (Palmer 1998). 120 species (Townsend 1993; Müller & Borsch 2005) and includes subshrubs, perennial and annual herbs, which are native in the New World tropics and subtropics (Holzhammer 1955, 1956; Eliasson 1988) and Australia (Palmer 1998). 33 species predominantly occur in the arid and semi-arid regions of western, central and northeastern Australia (Palmer 1998). The morphology of the inner two sepals can differ and has been regarded as one of the diagnostic characters of Lithophila and Philoxerus (Townsend 1993). In these studies the term “tepals” was used for the perianth organs. More recent work converges on the opinion that these organs are sepals (e.g. Ronse de Craene & Brockington 2013)
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