The Sticta Filix Morphodeme (Ascomycota: Lobariaceae) in New
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The Lichenologist 50(2): 185–210 (2018) © British Lichen Society, 2018 doi:10.1017/S0024282917000706 The Sticta filix morphodeme (Ascomycota: Lobariaceae)in New Zealand with the newly recognized species S. dendroides and S. menziesii: indicators of forest health in a threatened island biota? Hannah RANFT, Bibiana MONCADA, Peter J. DE LANGE, H. Thorsten LUMBSCH and Robert LÜCKING Abstract: We present a phylogenetic revision of the Sticta filix morphodeme in New Zealand. This non- monophyletic group of early diverging clades in the genus Sticta is characterized by a stalked thallus with a green primary photobiont and the frequent formation of a dendriscocauloid cyanomorph. Traditionally, three species have been distinguished in New Zealand: S. filix (Sw.) Nyl., S. lacera (Hook. f. & Taylor) Müll. Arg. and S. latifrons A. Rich., with two cyanomorphs separated under the names Dendriscocaulon dendriothamnodes Dughi ex D. J. Galloway (traditionally associated with S. latifrons)andD. dendroides (Nyl.) R. Sant. ex H. Magn. (traditionally associated with S. filix). Sticta lacera was not included in the present study due to the lack of authentic material (all specimens originally identified under that name and sequenced clustered with S. filix); S. filix was confirmed as a distinct species whereas S. latifrons s. lat. was shown to represent two unrelated species, S. latifrons s. str. and the reinstated S. menziesii Hook. f. & Taylor. The cyanomorphs of S. filix and S. latifrons are not conspecific with the types of the names D. dendriothamnodes and D. dendroides, respectively; the D. dendriothamnodes cyanomorph belongs to the Australian taxon Sticta stipitata C. Knight ex F. Wilson, which is not present in New Zealand, whereas the D. dendroides cyanomorph corresponds to a previously unrecognized species with unknown chloromorph, recombined here as Sticta dendroides (Nyl.) Moncada, Lücking & de Lange. Thus, instead of three species (S. filix, S. lacera, S. latifrons)withtheir corresponding cyanomorphs, five species are now distinguished in this guild in New Zealand: S. dendroides (cyanomorph only), S. filix (chloro- and cyanomorph), S. lacera (chloromorph only), S. latifrons (chloro- and cyanomorph) and S. menziesii (chloro- and cyanomorph). A key is presented for identification of the chloromorphs and the dendriscocauloid cyanomorphs of all species. Semi-quantitative analysis suggests that species in this guild are good indicators of intact forest ecosystems in New Zealand and that the two newly recognized species, S. dendroides and S. menziesii, appear to perform particularly well in this respect. The use of lichens as bioindicators of environmental health is not yet established in New Zealand and so, based on our results, we make the case to develop this approach more thoroughly. Key words: Australia, ITS barcoding locus, lichens, photosymbiodemes, species recognition Accepted for publication 19 July 2017 Introduction 2007; Brodo et al. 2001; Wirth et al. 2013; Stenroos et al. 2016). They are usually found Lobariaceae species are among the most in wet, temperate to tropical ecosystems. conspicuous macrolichens (Galloway 1985, Currently, 466 species are accepted in 12 H. Ranft: Riverside Brookfield High School, 160 H. T. Lumbsch: Integrative Research Center, Science Ridgewood Rd, Riverside, IL 60546, USA; & Education, The Field Museum, 1400 South Lake present address: Johns Hopkins University, Baltimore, Shore, Chicago, IL 60605, USA. MD 21218, USA. R. Lücking (corresponding author): Botanic Garden B. Moncada: Licenciatura en Biología, Universidad and Botanical Museum, Freie Universität Berlin, Distrital Francisco José de Caldas, Cra. 4 No. 26D-54, Königin-Luise-Straße 6–8, 14195 Berlin, Germany; Torre de Laboratorios, Herbario, Bogotá D.C., Research Associate, Integrative Research Center, Colombia; Research Associate, Science & Education, Science & Education, The Field Museum, 1400 The Field Museum, 1400 South Lake Shore, Chicago, South Lake Shore, Chicago, IL 60605, USA. Email: IL 60605, USA. [email protected] P. J. de Lange: Environmental and Animal Sciences, Unitec Institute of Technology, Mt Albert, Auckland, New Zealand. Downloaded from https://www.cambridge.org/core. IP address: 52.0.234.91, on 10 Feb 2020 at 14:25:43, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0024282917000706 186 THE LICHENOLOGIST Vol. 50 genera (Galloway & Elix 2013; Moncada (Galloway 1985, 1988, 2007; de Lange et al. et al. 2013a; Galloway 2015; Jaklitsch et al. 2012; de Lange & Galloway 2015), New 2016; Lücking et al. 2017). Except for Zealand lags behind in using lichens tropical species of Sticta (Schreb.) Ach., the (and especially Lobariaceae) for conservation family is richest in the Southern Hemisphere monitoring (Galloway 2008; de Lange et al. (Galloway et al. 1995; Galloway 2001) and 2012). This is all the more remarkable con- has been particularly well studied in New sidering that New Zealand’s native ecosystems, Zealand with 66 species currently accepted like many other island biotas, have been and (Galloway 2007). These correspond to ten continue to be severely affected by humans and genera under the current genus concept, two invasive plants and animals (Atkinson 1989; of them (Ricasolia De Not. and Yoshimuriella Holdaway 1989; King 1990; Anderson 2002; Moncada & Lücking) requiring confirmation Worthy & Holdaway 2002; Tennyson & (see below) and two, namely Dendriscosticta Martinson 2006; Prebble & Wilmshurst 2009; Moncada & Lücking and Lobariella Yoshim., de Lange et al. 2010; Brown et al. 2015). being absent from the country (Moncada Hutcheson et al. (1999) provided a sum- et al. 2016). Additional species have been mary of the use of bioindicators in New reported from oceanic possessions of New Zealand as a means of determining ecosystem Zealand (de Lange & Galloway 2015). health, pointing out the general lack of studies Lichens have long been used as bioindica- and near absence of work with lichens; tors and species in Lobariaceae are excellent observations confirmed by Galloway (2008). indicators of forest health (Rose 1974, 1976, De Lange et al. (2012) published the first ever 1992; Selva 1994, 1996; Zedda 2002). A prime threat assessment for the New Zealand lichen example is Lobaria pulmonaria (L.) Hoffm., biota, including their lichenicolous fungi, which has been extensively studied in this noting that for lichens assessed as ‘threatened’ respect in Europe and North America or ‘at risk’, ecosystem degradation and habitat (Søchting & Christensen 1989; Gauslaa 1994; loss were significant factors in causing their McCune 2000; Campbell & Fredeen 2004; decline. The majority of New Zealand’s Kalwij et al. 2005; Jüriado & Liira 2009; lichen biota listed by de Lange et al. (2012) Scheidegger 2009; Nascimbene et al. 2010; were assessed as ‘data deficient’, one of the Gustafsson et al. 2013; Cansaran-Duman et al. issues being the correct application of names, 2015; Giorgio et al. 2015). Other Lobariaceae especially of presumably cosmopolitan or considered rare and threatened by land use predominantly Northern Hemisphere lichens change in the Northern Hemisphere, including thought to occur in New Zealand, including Sticta fuliginosa (Dicks.) Ach., S. limbata those in Lobariaceae. (Sm.) Ach. and S. sylvatica (Huds.) Ach. Only a small number of studies have so far (Hodkinson et al. 2014; Magain & Sérusiaux addressed the problem of species recognition 2015), offer similar potential for bioindication of in lichen fungi in New Zealand with mole- ecological continuity. Recent phylogenetic stu- cular techniques (Summerfield et al. 2002; dies have shown that species concepts in these Thomas et al. 2002; Parnmen et al. 2012, taxa require critical revision (Moncada et al. 2013; Buckley et al. 2014; Hayward et al. 2014a; Magain & Sérusiaux 2015), which will 2014; Myles 2014; Pino-Bodas et al. 2015). have an influence on their use as bioindicators. Multi-locus approaches or studies using the In the tropics and the Southern Hemisphere, ITS barcoding locus (Schoch et al. 2012) Lobariaceae have been used or mentioned have shown that species delimitation in as potential bioindicators of forest health Lobariaceae is complex and that names based on studies in Cuba (Rosabal et al. 2010), globally applied to certain morphotypes, Colombia (Simijaca-Salcedo et al. 2014; such as Pseudocyphellaria crocata (L.) Vain., Ramírez-Morán et al. 2016), French Guiana Sticta dichotoma Bory and S. fuliginosa, (Normann et al. 2010), Chile (Galloway 1992) represent numerous lineages (Magain et al. and Thailand (Wolseley et al. 1994). Despite a 2012; Moncada et al. 2014a, b; Magain & wealth of literature dealing with Lobariaceae Sérusiaux 2015). As Lobariaceae are Downloaded from https://www.cambridge.org/core. IP address: 52.0.234.91, on 10 Feb 2020 at 14:25:43, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0024282917000706 2018 Sticta filix morphodeme in New Zealand—Ranft et al. 187 conspicuous lichens within the majority of two species have been reported from (south-) New Zealand’s ecosystems and have eastern Australia (Galloway 2001). To tremendous potential for use as bioindica- provide a solid base for the use of this guild tors, testing current species delimitations and its photomorphs as bioindicators, we (Galloway 1985, 1988, 1997, 2007) using assessed its phylogenetic relationships and molecular data is critically needed, enabling species delimitation