Neither Phoebe Nor Cinnamomum - the Tetrasporangiate Species of Aiouea (Lauraceae)

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Neither Phoebe Nor Cinnamomum - the Tetrasporangiate Species of Aiouea (Lauraceae) See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/320588407 Neither Phoebe nor Cinnamomum - the tetrasporangiate species of Aiouea (Lauraceae) Article in Taxon · October 2017 DOI: 10.12705/665.6 CITATION READS 1 167 7 authors, including: Randi Rohde Barbara Rudolph University of Hamburg University of Hamburg 1 PUBLICATION 1 CITATION 35 PUBLICATIONS 274 CITATIONS SEE PROFILE SEE PROFILE Pedro Moraes Jie Li São Paulo State University Xishuangbanna Tropical Botanical Garden 88 PUBLICATIONS 403 CITATIONS 1,162 PUBLICATIONS 9,645 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Studies on Pteridium from South America View project REDS-II View project All content following this page was uploaded by Randi Rohde on 28 October 2017. The user has requested enhancement of the downloaded file. TAXON 66 (5) • October 2017: 1085–1111 Rohde & al. • Transfer of Neotropical “Cinnamomum” to Aiouea (Lauraceae) Neither Phoebe nor Cinnamomum – the tetrasporangiate species of Aiouea (Lauraceae) Randi Rohde,1 Barbara Rudolph,1 Kristina Ruthe,1 Francisco G. Lorea-Hernández,2 Pedro Luis Rodrigues de Moraes,3 Jie Li4 & Jens G. Rohwer1 1 Universität Hamburg, Biozentrum Klein Flottbek, Ohnhorststr. 18, 22609 Hamburg, Germany 2 Instituto de Ecología, Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico 3 Universidade Estadual Paulista “Júlio de Mesquita Filho”, Departamento de Botânica, Av. 24 A 1515, Bela Vista, Rio Claro, São Paulo, CEP 13506-900, Caixa Postal 199, Brazil 4 Plant Phylogenetics and Conservation Group, Centre for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, 88 Xuefu Rd., Kunming, Yunnan 650223, China Author for correspondence: Jens G. Rohwer, [email protected] ORCID JGR, http://orcid.org/0000-0001-8560-9015 DOI https://doi.org/10.12705/665.6 Abstract Cinnamomum is among the largest genera of the Lauraceae, including species from tropical to temperate Asia and from tropical to subtropical America. However, previous studies indicated that Cinnamomum might not be monophyletic in its current circumscription. We therefore re-investigated the genus and possible relatives with an increased taxon sample, particularly of the Neotropical species. Our results, based on sequences of the nuclear ITS region and the chloroplast spacers psbA-trnH and trnG-trnS, show that the Neotropical Cinnamomum species indeed do not form a monophyletic group with the Paleotropical species. Instead, the American species form a clade in which most species of the Neotropical genus Aiouea are nested within several subclades. According to the ITS data the Aiouea/Cinnamomum clade is sister to the likewise predominantly Neotropical Ocotea complex, rather than the two Paleotropical Cinnamomum clades. Therefore, Cinnamomum cannot be upheld in its cur- rent circumscription. As the type C. verum, is from Asia, the Neotropical species need to be placed in a different genus. We decided to transfer them to Aiouea here, because this is the oldest generic name in the Aiouea/Cinnamomum clade, and the only character separating Aiouea from Neotropical Cinnamomum is the number of pollen sacs per anther (di- vs. tetrasporangiate), a feature that is known to be variable within several other genera as well. This transfer requires 42 new combinations and two nomina nova. In addition, six species described in the unpublished Ph.D. thesis of one of the authors (FLH) are validated in Aiouea here, and the only species of the genus Mocinnodaphne is transferred to Aiouea as well. Keywords Aiouea; Cinnamomum; cpDNA; ITS; Lauraceae; Mocinnodaphne; Neotropics; phylogeny; psbA-trnH; trnG-trnS Supplementary Material Electronic Supplement (results of maximum parsimony, Bayesian inference, and maximum likeli- hood analyses of the ITS, cpDNA and combined datasets, and the full descriptions, paratype data and discussions of the newly described species) and DNA sequence alignments, are available in the Supplementary Data section of the online ver- sion of this article at http://ingentaconnect.com/content/iapt/tax INTRODUCTION (Rohwer, 2000; Chanderbali & al., 2001; Chanderbali, 2004; Rohwer & al., 2009; Li & al., 2011). According to the results of The Lauraceae are the largest woody family of the Chanderbali & al. (2001), Groth (2003) and Huang & al. (2016), Magnoliidae and are commonly found in moist tropical and Cinnamomum Schaeff. appears to be one of those doubtfully subtropical forests of Central and South America, Africa, delimited genera. As currently circumscribed (Kostermans, Madagascar, the Canary Islands, Asia, and Australia (Rohwer, 1961; Rohwer, 1993a), it includes an Old World group, predomi- 1993a). The family consists of about 50 genera and approxi- nantly distributed in subtropical to tropical Asia, plus a few spe- mately 2500 to 3500 species as estimated by Rohwer (1993a). cies in Australia and the western Pacific area, as well as a New In the past, morphology-based delimitations of genera appeared World group in Central and South America. Rohwer (1993a) complicated and resulted in divergent classifications based on estimated the total number of species to about 350, including the importance of specific characters emphasized by differ- about 60 American species. Lorea-Hernández (1996), however, ent authors (Rohwer, 1993a; Van der Werff & Richter, 1996). recognized only 47 Neotropical species, and many names for Molecular studies, however, turned out to be largely congruent, Asian species have since been reduced to synonymy, especially and revealed that several traditional genera were polyphyletic by Li & al. (2008). The American species currently placed in Received: 2 Dec 2016 | returned for (first) revision: 20 Jan 2017 | (last) revision received: 22 May 2017 | accepted: 22 May 2017 || publication date(s): online fast track, n/a; in print and online issues, 24 Oct 2017 || © International Association for Plant Taxonomy (IAPT) 2017 Version of Record 1085 Rohde & al. • Transfer of Neotropical “Cinnamomum” to Aiouea (Lauraceae) TAXON 66 (5) • October 2017: 1085–1111 Cinnamomum had been included in Phoebe Nees by Nees (1836 Cinnamomum, but at the same time raised doubts about the – except Cinnamomum montanum (Sw.) Bercht. & J.Presl), current circumscription of Cinnamomum. The Paleotropical Meissner (1864) and Mez (1889). Kostermans (1961) transferred and Neotropical species did not form a monophyletic group, but them to Cinnamomum based on characters of the fruit. Already were rather placed in different clades consisting exclusively of Nees (1836) had noted that there were two distinct groups in either Paleotropical or Neotropical species. A few Neotropical Phoebe (in his circumscription): in the Asian species (now Cinnamomum species appeared nested among Ocotea Aubl., Phoebe s.str.) the fruit does not have a cupule, and its base whereas the majority formed a clade with species of Aiouea is clasped by enlarged, indurate tepals (Fig. 1A), whereas in Aubl. and Mocinnodaphne Lorea-Hern. (Chanderbali & al., the American species there is a cupule of variable size with 2001; Groth, 2003), but the nodes separating these clades were or without remnants of the tepals on its margin, or at least a only weakly supported. thickened and more or less fleshy pedicel (Fig. 1B). The latter Previous molecular studies including Cinnamomum were characters are also found in the Asian species of Cinnamomum conducted using the internal transcribed spacer region (ITS) (Fig. 1C, D). In addition, the American and Asian species cur- of nuclear ribosomal DNA (Chanderbali & al., 2001; Groth, rently placed in Cinnamomum share trimerous flowers with 2003; Rohwer & al., 2009; Lee & al., 2010 [ITS2]) and chloro- six equal tepals, nine fertile stamens, each with (mostly) four plast markers such as matK, trnL-trnF, rpoB, rbcL and rpl16 pollen sacks placed in two pairs above each other, and three (Rohwer, 2000; Chanderbali & al., 2001; Kuo & al., 2010; internal staminodes with sagittate glandular heads. These Sudmoon & al., 2014; Swetha & al., 2014). These markers, floral characters, however, are likely to be plesiomorphic, as however, did not yield substantial support values for the nodes they are also found in the Persea group, the sister group of the separating the main clades of Cinnamomum. Better support core Lauraceae (Cinnamomeae, Laureae). Molecular studies for some of the main clades was found in a recent study by (Rohwer, 2000; Chanderbali & al., 2001; Groth, 2003; Rohwer Huang & al. (2016) using ITS as well as two low-copy nuclear & al., 2009; Huang & al., 2016) confirmed that the American gene regions, the second intron of the LEAFY gene and a part species do not belong to the genus Phoebe and are closer to of the rpb2 gene. This study, however, was focused on the Fig. 1. Fruits of Phoebe and Cinnamomum species. A, Phoebe sp., almost mature (Indonesia, Bogor Botanical Garden); B, Aiouea hirsuta (“Cinnamomum hirsutum”), immature (Brazil, São Paulo, Serra da Cantareira); C, Cinnamomum wilsonii, immature (sect. Cinnamomum; China, Kunming Botanical Garden); D, Cinnamomum glanduliferum, mature (sect. Camphora; China, yard of Kunming Institute of Botany). — Scales: A, Black and white squares 1 cm wide; B–D, scale bar = 1 cm. All photos by Jens G. Rohwer. 1086 Version of Record TAXON 66 (5) • October 2017: 1085–1111 Rohde & al. • Transfer of Neotropical “Cinnamomum” to Aiouea (Lauraceae) Asian species and did not include Neotropical
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