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On Ingeae Systematics of Synandrous Mimosoids Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1969 On Ingeae Systematics of synandrous mimosoids JULIA FERM ACTA UNIVERSITATIS UPSALIENSIS ISSN 1651-6214 ISBN 978-91-513-1013-8 UPPSALA urn:nbn:se:uu:diva-420681 2020 Dissertation presented at Uppsala University to be publicly examined in Friessalen, EBC, Norbyvägen 14, Uppsala, Friday, 20 November 2020 at 10:00 for the degree of Doctor of Philosophy. The examination will be conducted in English. Faculty examiner: Tiina Särkinen (Royal Botanic Garden Edinburgh). Abstract Ferm, J. 2020. On Ingeae Systematics of synandrous mimosoids. Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1969. 50 pp. Uppsala: Acta Universitatis Upsaliensis. ISBN 978-91-513-1013-8. Background: The Ingeae tribe (Caesalpiniodeae, Fabaceae) has a long history of taxonomic complexity with genera being merged and split, and many species with former placements in several different genera. Moreover, phylogenetic studies have shown that the tribe is non- monophyletic with the genus Acacia (of tribe Acacieae) nested within Ingeae. This problem of non-monophyly is also reflected at the generic and specific level of ingoid taxa. Phylogenetic relationships have been difficult to resolve, with unsettled generic delimitations as a result. In this thesis, I investigated the systematics and phylogeny of the Ingeae-Acacia complex, with reflections on taxonomy and biogeography. Methods: Molecular data of plastid (matK, psbA, trnL-trnF, ycf1) and nuclear (ETS, ITS) DNA sequences were analysed using Bayesian inference and Ultrafast Bootstrap in order to investigate phylogenetic relationships of the Ingeae-Acacia complex. Results: In paper I, Marmaroxylon was shown to be included in Zygia and Zygia inundata in Inga. Marmaroxylon magdalenae, M. ocumarense and Zygia sabatieri were not recovered in the Zygia-Marmaroxylon clade and therefore left without a placement in any genus. In paper II, Zapoteca was shown to be monophyletic but the subgenera comprising more than one species, and four species, were non- monophyletic. In paper III, a new subgeneric classification of Zapoteca and an identification key to the subgenera are presented, as well as a phytogeographical review of the genus. Zapoteca formosa subsp. schottii and Z. formosa subsp. gracilis were recognized as distinct species. In paper IV, Afrocalliandra and Calliandra were recovered as the earliest diverging lineage within the Ingeae-Acacia complex, and the other taxa possessing the same pod-type as Calliandra, i.e. Sanjappa, Thailentadopsis, Viguieranthus and Zapoteca, were shown to be more closely related to each other and other ingoid genera than to Calliandra. Discussion and conclusions: Phylogenetic relationships of the Ingeae-Acacia complex are possible to resolve with a broad sampling and a combination of several nuclear and plastid informative DNA sequences. Taxonomic revisions are, however, needed for several ingoid genera, as well as for the entire Ingeae tribe since it currently is non-monophyletic with respect to Acacia. Shared morphological characters are not always indicative of common ancestry and older morphology-based classifications do not always reflect the evolutionary history of the group. One example is the “Calliandra-pod” fruit type. While it has often been argued to indicate close relationships, I show that this seemingly specific type of pod occurs in several unrelated genera. Keywords: Fabaceae, Ingeae, Systematics, Phylogenetics, Taxonomy, Morphology Julia Ferm, Department of Organismal Biology, Systematic Biology, Norbyv. 18 D, Uppsala University, SE-75236 Uppsala, Sweden. © Julia Ferm 2020 ISSN 1651-6214 ISBN 978-91-513-1013-8 urn:nbn:se:uu:diva-420681 (http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-420681) Ruusu on ruusu, Ja oli aina ruusu. Mutta teoria menee nyt Että omena on ruusu, Ja päärynä on, ja niin on Luumu, luulisin. Rakas tietää vain Mikä todistaa seuraavaksi ruusun. Olet tietysti ruusu - Mutta ollaan kaiki aina ruusuja. - Robert Frost List of Papers This thesis is based on the following papers, which are referred to in the text by their Roman numerals. I Ferm J., Korall P., Lewis G. P. & Ståhl B. (2019) Phylogeny of the Neotropical legume genera Zygia and Marmaroxylon and close relatives. Taxon. 68(4): 661-672. II Ferm J. (2019) A preliminary phylogeny of Zapoteca (Faba- ceae: Caesalpinioideae: Mimosoid clade). Plant Systematics and Evolution. 305(5): 341-352. III Ferm J. & Ståhl B. A revised classification of the Neotropical genus Zapoteca (Caesalpinioideae; Fabaceae), with one new sub- genus and two new species combinations. [Manuscript submitted for publication.] IV Ferm J., Ståhl B., Wikstöm N. & Rydin C. Phylogenetic relation- ships of the mimosoid Ingeae–AcaCia complex (Fabaceae), based on plastid and nuclear data. [Manuscript.] Published papers were reproduced with permission from the respective pub- lishers. Contents Introduction ................................................................................................... 11 Systematics ............................................................................................... 11 The Ingeae-AcaCia complex .................................................................... 11 Selected ingoid genera .............................................................................. 15 Fieldwork ...................................................................................................... 19 Objectives ...................................................................................................... 23 Materials & Methods ..................................................................................... 25 Taxon sampling ........................................................................................ 25 Laboratory work ....................................................................................... 26 Phylogenetic analyses ............................................................................... 27 Summary of papers ........................................................................................ 29 Phylogeny of the Neotropical legume genera Zygia and Marmaroxylon and close relatives. (Paper I) ..................................................................... 29 A preliminary phylogeny of Zapoteca (Fabaceae: Caesalpinioideae: Mimosoid clade). (Paper II) ...................................................................... 31 A revised classification of the Neotropical genus Zapoteca (Caesalpinioideae; Fabaceae), with one new subgenus and two new species combinations. (Paper III) ............................................................. 32 Phylogenetic relationships of the mimosoid Ingeae–AcaCia complex (Fabaceae), based on plastid and nuclear data. (Paper IV) ....................... 34 Concluding remarks ...................................................................................... 37 Svensk sammanfattning ................................................................................. 39 Systematik ................................................................................................ 39 Den mimosoida tribusen Ingeae ............................................................... 39 Zygia och Marmaroxylon ......................................................................... 40 Zapoteca och närbesläktade (?) taxa ........................................................ 41 Slutsats ...................................................................................................... 43 Acknowledgements ....................................................................................... 45 References ..................................................................................................... 47 Abbreviations DNA Deoxyribonucleic acid PCR Polymerase Chain Reaction s.l. sensu lato (in the broad sense) s.s. sensu stricto (in the narrow sense) sp. Species (singular) spp. Species (plural) subsp. Subspecies subg. Subgenera ser. Series sect. Section LPWG Legume Phylogeny Working Group POWO Plants of the World Online Introduction Systematics Systematics is the study of evolutionary relationships of taxa at all levels, of groups of organisms and lineages over time. These relationships can be visu- alized through phylogenetic trees, which illustrates common ancestry and di- versification in a straight forward way that is easy to interpret. Systematics includes also taxonomy, i.e. the formal classification and naming of taxa. Nowadays phylogenetic analysis is an important foundation for taxonomic work and evolutionary studies. With new techniques for the production of enormous amounts of DNA data, the possibilities for addressing and answer- ing evolutionary questions has increased well beyond expectation in a short period of time. By using modern phylogenetic analysis as a basis we can con- struct classifications that actually reflect the evolutionary relationships amongst taxa. With a robust phylogeny as a basis, questions on for example diversification dynamics, biogeography and morphological evolution can be addressed. The occurrence of the same morphological characters in different clades can be due to convergent evolution in distantly related taxa, or represent an ancestral state that has been lost in some lineages. Although morphological characters might not always provide an easily understandable indication of kinship it can often still be useful for understanding evolution
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