A New Species of Lindsaea (Lindsaeaceae, Polypodiopsida) from Mt

A new species of Lindsaea (Lindsaeaceae, Polypodiopsida) from Mt. Hamiguitan, Mindanao, Philippines

DIRK NIKOLAUS KARGER1*, SAMULI LEHTONEN2, VICTOR B. AMOROSO3 & MICHAEL KESSLER1

1 Institute of Systematic Botany, University of Zurich, Zollikerstrasse 107, CH-8008 Zurich, Switzerland 2 Department of Biology, University of Turku, FI-20014 Turku, Finland 3Department of Biology, Central Mindanao University, Musuan, Bukidnon, Philippines *Corresponding author: [email protected]

Abstract

A new species of Lindsaea from the Philippines, Lindsaea hamiguitanensis is described. The phylogenetic relationships of L. hamiguitanensis were analysed by sequencing trnL-trnF and trnH-psbA intergenic spacer regions and performing a cladistic analysis of the Lindsaea section Schizoloma. Lindsaea hamiguitanensis groups together with L. bouillodii and L. cf. cambodgensis, from which it differs morphologically by the length of its petioles and the entire pinnules, as well as genetically by three apomorphic substitutions in the trnL-trnF intergenic spacer, and by two apomorphic substitutions in trnH-psbA spacer region.

Introduction

The pantropical fern genus Lindsaea Dryander (1797: 39) comprises ca. 200 species, with about two thirds of the species occurring in the Old World tropics. A few species occur in subtropical Japan and Tasmania. The genus usually occurs in moist forests below 3000 m (Kramer 1971). Most species are terrestrial with some species at higher elevations also growing as epiphytes (Kramer 1971). Some lowland species can also grow on decaying wood (Tuomisto 1998). Mount Hamiguitan Range Wildlife Sanctuary in Davao Oriental, Mindanao, Philippines is a protected area covering 6 834 ha located between 6°40'N to 6°47'N and 126°09'E to 126°13'E. The mountain range is known for its unique characteristics and the largest pygmy ‘bonsai forest’ in the Philippines. Ultramafic rocks form the predominant substrate and weather into soil with an unusually high concentration of iron and magnesium (Amoroso et al. 2009). An inventory of pteridophytes revealed 141 species of ferns and 14 lycophytes on the mountain, with 9 being endemic to the range (Amoroso et al. 2011). During botanical fieldwork in 2009, we collected a distinctive species of Lindsaea which we could not identify using standard references (Kramer 1971). Molecular analyses of this species revealed it as distinct, and therefore it is here described as a new species.

Methods

We investigated the phylogenetic relationships of Lindsaea hamiguitanensis by sequencing trnL-trnF and trnH-psbA intergenic spacer regions and performing a cladistic analysis of Lindsaea sect. Schizoloma, with L. linearis Swartz (1801: 78) as outgroup. Laboratory protocols for DNA extraction, amplification, and sequencing followed Lehtonen et al. (2010). Data for other taxa included here were taken from Lehtonen et al.

Accepted by Marcus Lehnert: 29 May 2012; published online in PDF: 30 May 2012 15 (2010). Sequences were aligned using default options in Muscle (Edgar 2004). Short inverted sequence fragments are present in the studied spacer regions of some Lindsaea species, and they were replaced with their complementary sequences before the analysis, and a separate binary character was coded to recognise the original orientation (Lehtonen et al. 2010). Sequences were concatenated and analysed with unweighted parsimony analysis using TNT (Goloboff et al. 2008). The search consisted of 50 ratchet replicates holding five trees per replicate. Jackknife support indices (Farris et al. 1996) were calculated by repeating the search 1000 times with a removal probability of 37%. GenBank accession numbers for the studied specimens are given in Table 1. Morphological examinations have been conducted on two collections of L. hamiguitanensis, Karger 444 (Z) and Karger 426 (Z).

TABLE 1. DNA vouchers and GenBank accession numbers analysed here. New sequences produced in this study are marked in bold.

Taxon Voucher trnL-trnF trnH-psbA

Lindsaea agatii (Brack.) Lehtonen & Tuomisto Braithwaite 4519 (U) GU478854 GU478523 Lindsaea annamensis K.U.Kramer Averyanov VH2540 (AAU) GU478804 GU478501 Lindsaea austrosinica Ching Averyanov VH4816 (AAU) GU478801 GU478486 Lindsaea blotiana K.U.Kramer 1 Rakotondrainibe 6350 (P) GU478813 GU4785 Lindsaea blotiana K.U.Kramer 2 Rakotondrainibe 4327 (MO) GU478814 GU478511 Lindsaea bouillodii Christ Nielsen 592 (AAU) FJ360993 FJ360903 Lindsaea cf. cambodgensis Christ Iwatsuki 14505 (AAU) GU478840 GU478502 Lindsaea chienii Ching Kramer 7554 (U) FJ360995 FJ360905 Lindsaea dissectiformis Ching Averyanov VH2557 (AAU) FJ360997 FJ360907 Lindsaea ensifolia Sw. 1 Larsen 42363 (AAU) GU478853 GU478522 Lindsaea ensifolia Sw. 2 Kessler 13597 (GOET) n.a. GU478521 Lindsaea fraseri Hook. Streimann 8951 (L) FJ360998 FJ360908 Lindsaea gomphophylla Baker Hotta 12856 (U) GU478820 GU478497 Lindsaea goudotiana (Kunze) Mett. ex Kuhn Deroin 112 (P) GU478818 n.a. Lindsaea grandiareolata (Bonap.) K.U.Kramer Rakotondrainibe 4090 (MO) GU478810 GU478500 Lindsaea hamiguitanensis D.N.Karger & V.B.Amoroso Karger 444 (Z) JQ736811 JQ736810 Lindsaea heterophylla Dryand. Charoenphol 5081 (AAU) GU478855 n.a. Lindsaea javanensis Blume 1 Hennipman 3937 (U) FJ361002 FJ360911 Lindsaea javanensis Blume 2 Poulsen 1714 (TUR) FJ361003 FJ360912 Lindsaea kawabatae Kurata Saiki 1093 (Z) GU478803 GU478499 Lindsaea linearis Sw. Braggins 589 (AAU) FJ361008 FJ360917 Lindsaea madagascariensis Baker Rakotondrainibe 6349 (P) GU478815 GU478512 Lindsaea media R.Br. 1 Gittinss.n. (AAU) GU478842 GU478515 Lindsaea media R.Br. 2 van der Werff 11655 (MO) GU478843 GU478516 Lindsaea millefolia K.U.Kramer 1 Lan en Meeuse 5065 (L) GU478812 GU478513 Lindsaea millefolia K.U.Kramer 2 Guillaumet 4206 (P) GU478811 n.a. Lindsaea orbiculata (Lam.) Mett. ex Kuhn Averyanov VH4814 (AAU) FJ361013 FJ360922 Lindsaea oxyphylla Baker Gautier 2662 (P) GU478816 GU478508 Lindsaea schizophylla (Baker) Christ Sledge 613 (Z) GU478802 GU478498 Lindsaea sp. Laegaard 13835 (AAU) FJ361025 FJ360934 Lindsaea subtilis K.U.Kramer Werff 12855 (MO) GU478817 GU478514 Lindsaea walkerae Hook. Bostock 638 (Z) GU478841 GU478517 Lindsaea vieillardii Mett. McKee 5304 (U) GU478844 GU478496

Our cladistic analysis resulted in three most parsimonious trees of 183 steps, with a CI of 0.87 and RI of 0.93 (Fig. 1). Lindsaea hamiguitanensis grouped together with L. bouillodii Christ (1909: 59) and L. cf. cambodgensis Christ (1909: 58), although this clade did not receive >50% jackknife support. Lindsaea hamiguitanensis differed from these two species by three apomorphic substitutions in the trnL-trnF intergenic spacer, and by two apomorphic substitutions in trnH-psbA spacer region. The studied L. bouillodii and L. cf. cambodgensis specimens had identical sequences. Lindsaea hamiguitanensis shows various morphological characteristics that separate it from L. bouillodii. Rhizomes of L. hamiguitanensis are short-creeping and thinner than in L. bouillodii. The pinnae are entire in L. hamiguitanensis while they are shallowly incised in L. bouillodii. The petioles of L. hamiguitanensis are about twice as long as in L. bouillodii. DNA sequence data of non-coding chloroplast markers revealed that Lindsaea hamiguitanensis differs from its closest relatives. Although the differences are not great, they are still larger than the variation observed between different species in the L. annamensis Kramer (1972: 18) group, for example. The differences in morphological characters in comparison with the closely related species underline the molecular results.

FIGURE 1. Strict consensus tree of three most parsimonious trees found in a combined analysis of trnL-trnF and trnH-psbA intergenic spacer regions. Jackknife support (>50%) is indicated below the branches.

Lindsaea hamiguitanensis D.N.Karger & V.B.Amoroso, sp. nov. (Fig. 2)

Type:—PHILIPPINES. Mindanao: Davao Oriental, Mount Hamiguitan, 6°44'4.16" N, 126° 9'59.88" E, 1100 m, 1 November 2009, Karger 444 (holotype PNH!, isotypes UC!, Z!).

Lindsaea rhizomate breve repente, pinnulis alternis, remotis, utrinque circa 2–5, 1.0–1.8 cm longis, 0.8–1.0 cm latis, non incisis, textura herbacea, colore laete virente, petiolibusque rigidis, 10–30 cm longis, quadrangularibus.

FIGURE 2. A–C. Lindsaea hamiguitanensis D.N.Karger & V.B.Amoroso. A. Habit. B. Rhizome scale. C. Pinnule, abaxial, showing indusium. All from Karger 444 (Z).

18 • Phytotaxa 56 © 2012 Magnolia Press KARGER ET AL. Plants terrestrial. Rhizomes short-creeping, 1.00–1.75 mm diameter, scales ferruginous, very narrowly triangular, to 1.5 mm long. Petioles 10–30 cm long, to 4 times longer than the laminae, sharply quadrangular, dark reddish brown (proximally) to pale, with more or less well defined paler margins. Laminae triangular, 6– 13 cm long, bipinnate to (basally) tripinnate-pinnatifid, rachises reddish brown, with pale margins, sharply quadrangular. Major primary pinnae 4–6 to a side, spreading to moderately ascending, triangular, the largest (at the bases) to 6 cm long, to 4 cm wide, the proximal ones up to their width apart, the distal ones closer to each other. Distal pinnae gradually reduced, with a conform terminal pinna, costae dark green, slightly winged, adaxially sulcate. Proximal primary pinnae basally with some pinnules 1–3 cm long. Pinnules 2–5 on each side, herbaceous, light green when dry, slightly ascending, not contiguous, dimidiate-ovate to subtrapeziform (basally), the larger ones 1–1.8 cm long, 0.8–1 cm wide, 1.5–2 times longer than wide, margins continuous, rarely incised. Pinnules at lower primary pinnae shallowly incised. Distal pinnules gradually reduced, distal ones confluent with the acuminate terminal segment. Veins free, ca. 1.5 mm apart, 1- or 2-forked. Sori continuous, rarely incised on the basiscopic pinnules of the proximal primary pinnae, convex, extending 1/2 to 2/3 of the length of the pinnules along the sides of the pinnules, to 15 mm long, multinerval. Indusia coloured as the pinnules, indusial margins entire, rarely slightly erose, 0.5–0.7 mm wide, not reaching the margin by 1–1.5 times of their width. Spores trilete. Distribution and habitat:—Known only from Mount Hamiguitan, Mindanao, Philippines, where it grows in lower montane rainforest at 1100–1200 m. Etymology:—The species is named for the type locality. Additional specimen examined (paratype):—PHILIPPINES. Mindanao: Davao Oriental, Mount Hamiguitan, 6°44'4.16" N, 126°9'59.88" E, 1100 m, 1 November 2009, Karger 426 (Central Mindanao University Herbarium!, UC!, Z!).

Acknowledgements

We thank the officials of the Protected Areas and Wildlife Bureau (PAWB)-Department of Environment and Natural Resources (DENR), DENR Region XI, Protected Areas Management Board (PAMB) of Mount Hamiguitan Wildlife Sanctuary for the gratuitous permit, Alan R. Smith for initiating and revising this manuscript, and Fulgent Coritico and Elisea ‘Bebet’ Gozun for logistic support. This study was funded by the Swiss National Science Foundation (SNF) and the Claratz Schenkung.

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