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Abildgaardieae, Cyperaceae) À a New Genus from Sub-Saharan Africa

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Abildgaardieae, Cyperaceae) À a New Genus from Sub-Saharan Africa South African Journal of Botany 128 (2020) 326À332 Contents lists available at ScienceDirect South African Journal of Botany journal homepage: www.elsevier.com/locate/sajb Zulustylis (Abildgaardieae, Cyperaceae) À a new genus from sub-Saharan Africa A.M. Muasyaa,c,*, P. Goetghebeurb, I. Larridonb,c a Bolus Herbarium, Department of Biological Sciences, University of Cape Town, Private Bag X3, Rondebosch 7701, South Africa b Systematic and Evolutionary Botany Lab, Department of Biology, Ghent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium c Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, United Kingdom ARTICLE INFO ABSTRACT Article History: The generic placement of two African species currently included in the genus Fimbristylis, F. hygrophila and F. Received 1 May 2019 variegata, has been contentious. These two species, which have been known for the past half century, are gla- Revised 2 September 2019 brous tufted perennials whose spikelets have semi-distichous to spiral glume arrangement, bisexual flowers, Accepted 22 November 2019 and a pistil with an enlarged and deciduous style base. While the two species fit the gross morphology of Available online 5 December 2019 Fimbristylis, they differ in lacking the characteristic Kranz anatomy. We present phylogenetic inference of the Edited by AR Magee Abildgaardieae based on DNA sequence data and map photosynthetic type. C4 photosynthesis has arisen twice in Abildgaardieae, in the Bulbostylis and the Fimbistylis clades, with a C3 clade (including Arthrostylis) Keywords: sister to rest of tribe and a C3 grade (the two African Fimbristylis species, Trachystylis, Actinoschoenus) sepa- C3 photosynthesis rating the two C4 clades. The new genus Zulustylis is described here, to segregate the two aberrant African Fimbristylis hygrophila Fimbristylis species, and its taxonomy presented. Fimbristylis variegata Kranz anatomy Crown Copyright © 2019 Published by Elsevier B.V. on behalf of SAAB. All rights reserved. New combinations Phylogenetic inference Taxonomy 1. Introduction 1 G, 1H), and can be distinguished from Scirpeae by the unique embryo types (Fimbristylis-, Bulbostylis-, and Abildgaardia-type) and A number of changes in generic circumscription and classification consistent lack of perianth segments. Goetghebeur (1998) classified in the Cyperaceae have been made within the last two decades. The six genera in Abildgaardieae (Abildgaardia Vahl, Bulbostylis, Crosslan- majority of these have been caused by the shifting paradigm to recog- dia W.Fitzg., Fimbristylis, Nelmesia Van der Veken, Nemum Desv.), all nize monophyletic taxa (Humphries and Linder, 2009). During this sharing presence of multiple florets per spikelet and distinctly thick- period, increased use of molecular data in reconstructing phyloge- ened style bases in the majority of the taxa. At the same time, Goet- netic relationships has revealed the need to revise the classification ghebeur (1998) classified four other genera (Actinoschoenus Benth., at several suprageneric ranks, and the family is currently classified Arthrostylis R.Br., Trachystylis S.T.Blake, Trichoschoenus J.Raynal), shar- into two subfamilies (Cyperoideae and Mapanioideae) and into 18 ing a similarly thickened style base, in tribe Schoeneae due to their tribes (Muasya et al., 2009; Semmouri et al., 2019). Nearly every one spikelets having increasingly longer glumes but only the penultimate of the tribes has had changes in classification, with species and even glume (or rarely 2) bearing a bisexual floret (Figs. 1A, 1E, 1F). The lat- genera moved from tribe to tribe, the most drastic being the reclassi- ter four genera have previously been placed in the segregate tribe fication of the monotypic South African genus Hellmuthia Steud. from Arthrostylideae (Goetghebeur, 1986; Bruhl, 1995). subfamily Mapanioideae (Goetghebeur, 1998) into the Cyperoideae Recent phylogenetic studies (e.g. Ghamkhar et al., 2007; Simpson (Muasya et al., 2009). et al., 2007; Muasya et al., 2009; Semmouri et al., 2019; Roalson et al., Tribe Abildgaardieae is predominantly pantropical, comprising c. 2019a) have shown that Arthrostylis and allied genera are part of the 540 species, with highest diversity in Bulbostylis Kunth and Fimbristy- Abildgaardieae clade. Furthermore, Crosslandia has been demon- lis Vahl (WCSP, 2019). Abildgaardieae was circumscribed (Lye, 1973) strated to be a lineage within Fimbristylis (Ghamkhar et al., 2007; to include taxa that have many-flowered bisexual spikelets (Figs. 1B, Roalson et al., 2019b), and that Nemum (and perhaps Nelmesia) are lineages within Bulbostylis (Muasya et al., 2009; Hinchliff and Roal- son, 2013; Spalink et al., 2016; Semmouri et al., 2019; Roalson et al., * Corresponding author at: Bolus Herbarium, Department of Biological Sciences, University of Cape Town, Private Bag X3, Rondebosch 7701, South Africa. 2018, 2019a). The generic placement of two African species (Fimbris- E-mail address: [email protected] (A.M. Muasya). tylis hygrophila Gordon-Gray, F. variegata Gordon-Gray; Figs. 1C, 1D) https://doi.org/10.1016/j.sajb.2019.11.027 0254-6299/Crown Copyright © 2019 Published by Elsevier B.V. on behalf of SAAB. All rights reserved. A.M. Muasya et al. / South African Journal of Botany 128 (2020) 326À332 327 We first inferred the gene trees for each of the four regions sepa- rately to identify potential incongruence. As there was little conflict, the matrices of the four regions were concatenated for the down- stream analyses. PartitionFinder 2.1.1 (Lanfear et al., 2012) was used to determine an appropriate data-partitioning scheme from potential partitions that were defined a priori (in this case, each marker was treated as a separate partition, as was each codon in the two coding markers), as well as the best-fitting model of molecular evolution for each partition, using the Bayesian Information Criterion. The optimal data-partitioning scheme was ITS, ndhF codon 1 + rbcL codon 1, ndhF codon 2, ndhF codon 3, rbcL codon 2, rbcL codon 3, trnL-trnF. Maxi- mum likelihood (ML) analyses of the optimally partitioned data were performed using RAxML 8.2.10 (Stamatakis, 2014) with model set to GTR (gamma) for each partition. The search for an optimal ML tree Fig. 1. Photographic illustration of the inflorescence morphology in the Abildgaar- dieae. Arthrostylis aphylla (A), Bulbostylis atrosanguinea (B), Zulustylis hygrophila (C), was combined with a rapid bootstrap analysis of 1000 replicates. Par- Zulustylis variegata (D), Actinoschoenus repens (E), Trachystylis stradbrokensis (F), Abil- titioned analyses were conducted using Bayesian Inference (BI) in dgaardia ovata (G), Fimbristylis ferruginea (H). Vouchers: A, Brass 18669 (K); B, Luke MrBayes 3.2.6 (Ronquist et al., 2012). Based on the PartitionFinder 4599 (K); C, Venter 2832 (K); D, Huntley 734 (K); E, Robinson 5977 (K); F, Blake 22673 analysis, the GTR (invgamma) model of sequence evolution was set (K); G, Verdcourt 3253 (K); H, Ward 7924 (K). Scale bar = 1 mm. for the ITS partition; the GTR (gamma) model for the ndhF codon 1+rbcL codon 1 partition and the ndhF codon 3 partition; the F81 has remained uncertain. These species were described as part of Fim- model (propinv) for the ndhF codon 2, rbcL codon 2 and rbcL codon 3 bristylis (Gordon-Gray, 1966) and were later transferred into Abil- partitions; and the F81 model (invgamma) for the trnL-trnF partition dgaardia (Lye, 1971). As Abildgaardia has been considered to be a in the concatenated data set. The analysis was allowed to run for synonym of Bulbostylis (e.g. Lye, 1971, 1973; Haines and Lye, 1983), 100 million generations across four independent runs with four these species have by extension also been viewed as belonging to chains each, sampling every 10,000 generations. Convergence, associ- fi Bulbostylis. The uncertainty in af nity is epitomized by the segrega- ated likelihood values, effective sample size (ESS) values and burn-in tion of these two species as the only C3 species of Fimbristylis by values of the different runs were verified with Tracer 1.6 (Rambaut fi Bruhl (1995) and by Gordon-Gray (1995) who classi ed one of the and Drummond, 2014). The first 25% of the trees from all runs were species in Abildgaardia (A. hygrophila (Gordon-Gray) Lye) and the excluded as burn-in before making a majority-rule consensus of the other in Fimbristylis (F. variegata). Molecular phylogenetic studies 30,000 posterior distribution trees using the “sumt” function. All phy- (Besnard et al., 2009; Hinchliff and Roalson, 2013; Bouchenak-Khe- logenetic analyses were run using the CIPRES portal (http://www. ladi et al., 2014; Spalink et al., 2016; Semmouri et al., 2019) have phylo.org/; Miller et al., 2010), and were executed for both full and shown that these sister species are not part of the clade including reduced sampling alignments. Trees were drawn using FigTree 1.4.3 Abildgaardia and Fimbristylis. We investigate the taxonomic place- (http://tree.bio.ed.ac.uk/software/figtree/) and adapted in Adobe ment of these two sub-Saharan African species of Abildgaardieae and Photoshop CS5. revise their taxonomy. 3. Results 2. Materials and methods Since the topologies of the concatenated ML and BI trees were Representative herbarium specimens of tribe Abildgaardieae were entirely congruent, we here provide the BI tree with both the BI pos- studied to identify distinguishing morphological features. In addition, terior probability values (PP) and ML bootstrap values (BS) (Fig. 2). all available specimens of the two target species were studied in Analysis of the individual markers show strong support for the Abil- selected herbaria (BOL, BR, GENT, K, NBG, NU, PRE; codes follow dgaardieae sensu lato. The first diverging lineage has Arthrostylis (sis- Thiers 2019+) as well as consultation of key literature (e.g. Gordon- ter to Fimbristylis composita) as sister to rest of tribe; then strongly Gray, 1995; Goetghebeur, 1998). Gross morphological observations supported lineages observed consistently comprise Bulbostylis were made to enable species descriptions, with the aid of dissecting (including Nemum), a grade with three lineages (Fimbristylis hygro- microscope and measurements made using a ruler or digital calipers. phila sister to F.
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