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Research Article
Pteris latipinna sp. nov. (Pteridaceae), a new species segregated from Pteris fauriei
expand article infoYi-Shan Chao, Atsushi Ebihara§, Wen-Liang Chiou|, Yao Moan Huang
‡ Kaohsiung Medical University, Kaohsiung, Taiwan
§ National Museum of Nature and Science, Tsukuba-shi, Japan
| Dr. Cecilia Koo Botanic Conservation Center, Pingtung, Taiwan
¶ Taiwan Forestry Research Institute, Taipei, Taiwan
Open Access

Abstract

Pteris fauriei is widely distributed in Eastern Asia and has high morphological variation. Some morphologically similar plants related to this species are difficult to distinguish. We showed that the new Pteris species from Taiwan, previously identified as P. fauriei, can be morphologically distinguished by its wide pinnae, larger terminal pinnae than the lateral pinnae in sterile fronds, and triangular basal segments of the lateral pinnae. It was confirmed that this species is phylogenetically separated from the other East Asian Pteris species, except for a morphologically distinct species P. arisanensis, by means of chloroplast genes, rbcL and matK. The new species is named as Pteris latipinna sp. nov., referring to its wide pinnae. Here, we provide a key to facilitate the identification of the morphologically similar Pteris species in Asia. The morphological descriptions, images, ecology, and distribution are also presented.

Keywords

Pteris, Pteris fauriei, Pteris latipinna, Taiwan, taxonomy

Introduction

Pteris fauriei Hieron. is widely distributed in Eastern Asia. Two varieties of P. fauriei have been confirmed, and both varieties have different cryptic characteristics and prefer different niches. Pteris fauriei Hieron. var. fauriei, apomictic and triploid (2n = 87), usually has herbaceous laminae and prefers cooler sites; P. fauriei var. minor Hieron., sexual and diploid (2n = 58), usually has coriaceous laminae and is found in warmer sites (Huang et al. 2006; Huang et al. 2007). In Taiwan, some undescribed Pteris plants (Fig. 1), usually regarded as P. fauriei var. fauriei, with herbaceous laminae were found in understory of evergreen forests. However, those plants have wider laminae and pinnae than other bipinnatifid Pteris species recorded in Taiwan. Outside Taiwan, P. natiensis Tagawa, a Japanese endemic fern (Iwatsuki 1995), apomictic and diploid (Nakato and Ebihara 2016), is the most morphologically similar species in East Asia.

Figure 1. 

Photographs of Pteris latipinna Y.S.Chao & W.L.Chiou, sp. nov. in Hsinchu, Taiwan. A Habitat. Terminal pinna of sterile frond is larger than the lateral pinna. St, sterile fronds; Fe, fertile fronds B A frond C Concolorous scales a stipe.

Pteris fauriei and morphologically similar Pteris species are phylogenetically close. Chao et al. (2014) revealed that the clade (A1, including those species with bipinnatified laminae mostly) arose more recently than most of other clades in Pteris. Because of similar morphology, the characteristics to delimitate species need to be examined and compared in detail, such as venation, scale color, shapes of pinnae and segments, and pinnae stalked or sessile (Chao et al. 2013).

In this study, we clarified the morphological and phylogenetic characteristics of the undescribed Pteris plants, in comparison with P. fauriei, P. natiensis, and related bipinnatifid Pteris species from East Asia, including P. wulaiensis C.M. Kuo endemic to Taiwan; P. arisanensis Tagawa, P. biaurita L., P. kawabatae Sa. Kurata, P. kiuschiuensis Hieron., and P. oshimensis Hieron. distributed in China and Japan; and P. boninensis H. Ohba, P. laurisilvicola Sa. Kurata, P. satsumana Sa. Kurata, and P. yakuinsularis Sa. Kurata endemic to Japan (Iwatsuki 1995; Liao et al. 2013). On the basis of morphological and molecular data, the taxonomic treatments were applied.

Materials and methods

Morphology

We examined type materials of morphologically similar taxa, including P. fauriei var. fauriei (in herbaria B, BM, KYO, MO, P), P. fauriei var. minor (in herbaria B, BM, KYO, P), and P. natiensis (in herbaria KYO, P). Several morphologically similar species in neighboring areas were also compared, including P. arisanensis, P. biaurita, P. boninensis, P. kawabatae, P. kiuschiuensis, P. laurisilvicola, P. oshimensis, P. satsumana, P. wulaiensis, and P. yakuinsularis.

Phylogenetic analyses

To clarify the phylogenetic relationships of the undescribed plants, 34 other Pteris taxa with bipinnatifid laminae were sampled. Three Pteris species, P. grevilleana, P. longipinna, and P. venusta, were used as outgroups. These bipinnatifid and outgroup species belong to clades A1 and A2, respectively, according to the phylogenetic tree of Pteris (Chao et al. 2014). Vouchers and GenBank accession numbers are listed in Appendix 1. Total genomic DNA was extracted from young fronds, following a modified cetyltrimethylammonium bromide (CTAB) method (Doyle and Doyle 1990). Two chloroplast genes, rbcL and matK, were amplified using the PCR primers for rbcL and matK as per Chao et al. (2014). Alignment was performed with ClustalW (Thompson et al. 1994) and manually edited using BioEdit 7.1.3 (Hall 1999). Gaps were treated as missing data.

Maximum likelihood (ML) analyses were performed using GARLI v.2.0.1019 (Zwickl 2006). Ten independent runs were conducted using automatic termination following 10,000 generations without a significant (lnL increase of 0.01) change in topology. To calculate ML bootstrap support for each node, 1,000 bootstrap replicates were performed with automatic termination at 10,000 generations, under one run.

Results

Morphology

The distinct morphologies that distinguished the undescribed species from other bipinnatifid Pteris species are its wide pinnae, up to 7 cm wide, and fewer pairs of lateral pinnae, only 2–5 pairs (Fig. 1). Furthermore, its terminal pinnae of sterile fronds are larger than the lateral pinnae (Table 1). In Taiwan, these characteristics can separate the undescribed species from P. fauriei var. fauriei and P. fauriei var. minor (these two taxa were illustrated by one of their type materials, Figs S1 and S2, respectively).

An endemic species in Japan, Pteris natiensis (illustrated by holotype, KYO, Fig. S3), also has sterile fronds with slightly larger terminal pinnae than the lateral pinnae. Its pinnae are slightly narrower than those of the undescribed species (3–5 cm vs. 3–7 cm), and the basal pinna-segments are adnate to the rachis whereas they are not adnate to the rachis in the undescribed species (Table 1). Another specific trait of the undescribed species is the triangular (vs. falcate) basal segments of the lateral pinnae, which could be used to identify the new species from other similar species, including P. fauriei and P. natiensis (Table 1). The triangular and falcate basal segments are resulted by the longer costa adnate with the segments of the undescribed species and shorter costa adnate with the segments of the other species, respectively.

Morphological comparisons among Pteris latipinna Y.S.Chao & W.L.Chiou, sp. nov., P. fauriei var. fauriei, P. fauriei var. minor, and P. natiensis.

Species/Characteristics P. latipinna P. fauriei var. fauriei P. fauriei var. minor P. natiensis
Lamina size 15–45 cm long, 15–40 cm wide; length/width ratio about 1 15–40 cm long, 10–35 cm wide; length/width ratio 1.2–1.5 10–30 cm long, 10–25 cm wide; length/width ratio about 1 15–40 cm long, 10–35 cm wide; length/width ratio about 1.1–1.2
Number of lateral pinnae of sterile fronds 2–3(4) pairs 2–7 pairs 2–5 pairs 2–5 pairs
Lateral pinnae of sterile fronds Slightly incurved Straight Straight Incurved
Petiolule Sessile or short-petiolate. Most basal pinna-segments free to the rachis, sometimes adnate Sessile or short-petiolate. Basal pinna-segments free to the rachis Sessile or short-petiolate. Basal pinna-segments free to the rachis Sessile. Basal pinna-segments adnate to the rachis; except basal pinnae
Basal segment of lateral pinnae Triangular Falcate Falcate Falcate
Terminal pinna size of sterile fronds Distinctly wider than lateral pinnae except basal ones Smaller than lateral pinnae Smaller than lateral pinnae Almost the same size as lateral pinnae
Pinna shape Ovate-lanceolate, distinctly narrowed at base Lanceolate, not narrowed at base Lanceolate, not narrowed at base Ovate to lanceolate, more and less narrowed at base
Width of lateral pinna 3–7 cm 2–3.5 cm 1–3 cm 3–5 cm

Phylogeny and chloroplast DNA differences

Genetic data and the accession numbers of the sequences are listed in Appendix 1. The chloroplast DNA (cpDNA) alignment matrix of rbcL (1,278 bp) and matK (900 bp) contained a total of 2,178 characters with 121 parsimony-informative sites. The log-likelihood score for the most likely ML tree was -5304.42470.

The phylogenetic tree (Fig. 2) infers that the Pteris species with bipinnatifid laminae formed one monophyletic group (the clade of ingroup taxa), as revealed in the previous Pteris phylogeny (Chao et al. 2014). The undescribed taxon and P. fauriei were divided into two different clades, Clade I and II. In Clade I, the undescribed taxon shared identical cpDNA sequences with P. arisanensis, although they can be separated by their morphologies, such as venation and lamina shape (Fig. S4). The undescribed taxon cpDNA differed from P. natiensis, P. wulaiensis (Fig. S5), and P. yakuinsularis cpDNA by one nucleotide substitution, and from P. laurisilvicola cpDNA by two nucleotide substitutions. In Clade II, P. fauriei var. fauriei, P. fauriei var. minor, and P. oshimensis shared identical cpDNA sequences.

Figure 2. 

Chloroplast DNA phylogeny of the Pteris latipinna Y.S.Chao & W.L.Chiou, sp. nov. and related taxa. ML bootstrap support values are indicated on each branch.

Both morphological and DNA characteristics support that this taxon is a new species, rather than a variety of P. fauriei. Here, we describe the new species and delimitate P. fauriei var. fauriei and P. fauriei var. minor. The morphology of the new species is presented in Fig. 3 and described below.

Figure 3. 

Illustration of Pteris latipinna Y.S.Chao & W.L.Chiou, sp. nov., based on holotype. A A fertile frond B Venation C Linear, concolorous scale. Basal segments of lateral pinnae are triangular (indicated by arrows).

Taxonomic treatment

Pteris latipinna Y.S.Chao & W.L.Chiou, sp. nov.

Figs 1, 3

Type

TAIWAN. Hsinchu County: Zhudong Town, Wuchihshan, 3 March 2013, Y.-S. Chao 2092 (holotype TAIF!, isotype TAIF!, TNS!).

Description

Rhizomes short, ascending, apex scaly; scales linear lanceolate, 1–4 mm long, 0.2–0.5 mm wide, concolorous, dark brown, entire, apex long-acuminate. Fronds clustered, 30–100 cm long, nearly monomorphic. Sterile fronds 30–70 cm long; stipes green, 2–4 mm thick, 10–30 cm long, base with persistent and scattered scales; grooved on the adaxial side; laminae widely ovate, 15–45cm long, 15–40 cm wide, bipinnatifid; 2–3(4) pairs lateral pinnae, pinna angle against rachis 60–70°, straight, basal pinnae with one pair of exaggerated basiscopic pinnules, terminal pinnae distinctly longer and wider than the lateral except basal ones; pinnae ovate-lanceolate, distinctly narrowed at base, pectinate, 8–21 cm long, 3–7 cm wide, sessile or short-petiolate, apex caudate, 1–4 cm long. Basal segments of the lateral pinnae triangular, the other segments of pinnae falcate, 4–9 mm wide, apex obtuse, margins entire; veins forked, free. Fertile fronds 50–105 cm long; stipes 25–55 cm long; laminae ovate to widely ovate, 20–50 cm long, 20–35cm wide, bipinnatifid; 3–5 pairs lateral pinnae, slightly incurved or straight; terminal pinna usually wider than the lateral; pinnae 8–20 cm long, 2–6 cm wide, 1–4 cm long; segments of pinnae 4–6 mm wide, apex acute or obtuse. Sori along pinna margins, protected by pseudoindusia; spore number 32; spores tetrahedral, tan.

Other specimens examined

TAIWAN. Hsinchu County: Guanxi, Chike Mt., P.-F. Lu 24585, 24586 (TAIF); Jianshi, P.-F. Lu 25108 (TAIF); Pawushan, P.-F. Lu 26666, 26673 (TAIF); Shuitien Logging Trail, L.-Y. Kuo 01 (TAIF). Miaoli County: Sintikusyu, komokwan, Yaiti Simada 5175A (HAST).

Distribution

Taiwan (Fig. 4).

Ecology

In shaded places, understory of evergreen broad leaf forests, below 1,000 m in elevation.

Etymology

The specific epithet ‘latipinna’ refers to its wide pinnae.

Preliminary conservation assessment

We investigated the distribution of P. latipinna Y.S.Chao & W.L.Chiou, sp. nov. in Taiwan. To date, only a few small populations are recorded. However, the available information is inadequate to support the assessment of its extinction risk. According to the IUCN (2012) criteria, the category of Data Deficient (DD) is appropriate.

Figure 4. 

Distribution of Pteris latipinna Y.S.Chao & W.L.Chiou, sp. nov. (black circles) in Taiwan.

Discussion

A new species, P. latipinna Y.S.Chao & W.L.Chiou, sp. nov., growing understory of forests in Taiwan was found and identified in this study. Pteris latipinna is the largest species among the bipinnatifid Pteris species with single-axis in Taiwan. There were 29 Pteris species recorded in the Flora of Taiwan (Shieh 1994), and several new species and new records have been recently found (Chao et al. 2013; Chao et al. 2015; Ebihara et al. 2014; Knapp 2011; Knapp and Hsu 2017). In this study, we describe one more new species, and thus in total, 36 Pteris species, including infraspecies, have been documented in Taiwan.

Although the ploidy of P. latipinna is not known, with the similar morphology and apomitic reproductive mode, it is inferred that those species possibly evolved through a complex reticulate hybridization-polyploidization speciation. Those apomicitic Pteris species have also been suggested with possible hybrid origins (Chao et al. 2012a; Chao et al. 2012b; Walker 1979). Pteris latipinna has 32 spores per sporangium, which is thought as apomictic (Chao et al. 2010; Huang et al., 2006; Nakato 1975; Walker 1979). For those species in the same clade (Clade I) of P. latipinna, it is reported that P. laurisilvicola is diploid and triploid and apomictic (Nakato 1996; Nakato and Ebihara 2016); P. natiensis and P. wulaiensis are diploid (Huang et al. 2011; Kurita 1962; Nakato and Ebihara 2016); P. yakuinsularis are triploid (Nakato and Ebihara 2016); P. arisanensis is tetrapolyploid (Tsai and Shieh 1984). Remarkably, P. latipinna and P. arisanensis have the same cpDNA characteristics although their morphologies are clearly different. They have different lamina shapes (wide ovate for P. latipinna vs. ovate for P. arisanensis) and venation (free veins in P. latipinna vs. costal areolae in P. arisanensis) (Fig. S4). Similarly, in Clade II, P. oshimensis does not morphologically resemble P. fauriei but share identical cpDNA sequences. More cpDNA and nuclear DNA markers are needed to clarify the relationships among these species in P. fauriei complex.

In this study, taxa in Clade I and Clade II compose Pteris fauriei complex because they are morphologically similar and phylogenetically close with Pteris fauriei. All of them are distributed in Asia, mostly in Japan and Taiwan. Interestingly, distributions of most of those species are limited: Pteris latipinna and P. wulaiensis are endemic in Taiwan; P. boninensis, P. natiensis, and P. yakuinsularis are endemic in Japan (Iwatsuki 1995; Shieh 1994). This pattern of distribution implies those species arose in a small area within a short time recently (Chao et al. 2014).

The traits useful for separating P. latipinna from the similar species are used in a key for identification of this species as shown below.

1 Stipes <2 mm thick 2
2 Pairs of lateral pinnae 4–6; basal pinnae shorter or equal to the second basal ones; pinnae narrowest at base P. wulaiensis
2' Pairs of lateral pinnae 6–11; basal pinnae longer than the second basal ones; pinnae widest at base P. oshimensis
1' Stipes 2.5–4 mm thick. 3
3 Laminae widely lanceolate; ratio of length to width approximately 3:2 4
4 Laminae bipinnatifid; the segments extending to 2/3–4/5 of the way toward the costae; venation free or with costal areolae 5
5 Costal areolae arched, few triangular, connective veins with free veinlets. P. biaurita
5' Costal areolae triangular or absent; if present, connected by a pair of furcated veinlets P. arisanensis
4' Laminae bipinnatisect; the segments extending almost to the costae; venation completely free, no costal areolae 6
6 Pinnae caudate with long tail 2–4 cm. P. boninensis
6' Pinnae acute or caudate with short tail 0.5–2 cm 7
7 Scales at stipe base caducous; pinnae sessile P. laurisilvicola
7' Scales at stipe base persistant; pinnae often stalked P. yakuinsularis
3' Laminae widely ovate, ratio of length to width approximately 5:4 8
8 Pinnae sessile except basal ones, with basal pinna-segments adnate to the rachis, pinna angle against rachis nearly 90°, incurved 9
9 Pinnae sometimes suddenly wider at base; segments oblong with rounded apex P. kawabatae
9' Pinnae not wider at base; segments falcate with obtuse apex 10
10 Pinnae nearly oblong, equally wide, 2–3 cm wide P. kiuschiuensis
10' Pinnae ovate-lanceolate to lanceolate, widest at middle, 3–6 cm wide. 11
11 Lateral pinnae 5–6 pairs, pinnae 3–4 cm wide, terminal pinna-segments long, >1 cm P. satsumana
11' Lateral pinnae 2–5 pairs, pinnae 3–6 cm wide, terminal pinna-segments short, <0.5 cm P. natiensis
8' Pinnae stalked to sessile, without basal pinna-segments adnate to the rachis, pinna angle against rachis 60–70°, straight 12
12 Basal segments of lateral pinnae triangular P. latipinna
12' Basal segments of lateral pinnae falcate (P. fauriei) 13
13 64 spores per sporangium; laminae coriaceous P. fauriei var. minor
13' 32 spores per sporangium; laminae herbaceous P. fauriei var. fauriei

Acknowledgments

The authors thank Pi-Fong Lu, for her assistance in collecting plant samples, and Dr. Chuan-Kuei Liao for illustration drawings. The curators of herbaria B, BM, HAST, KYO, MO, P, PE, TAI, TAIF, and TNS are gratefully acknowledged for loan of specimens. This research is supported by Ministry of Science and Technology of Taiwan (MOST103-2621-B-037-001-MY2, MOST106-2311-B-037-005-MY3).

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Appendix 1

Specimen information and GenBank accession numbers.

Taxon Specimen collection number Collection locality GenBank accession numbers for rbcL matK Herbarium for voucher specimen
P. setulosocostulata Y.-S. Chao 1146 Taiwan KF289634 KF289501 TAIF
P. keysseri Y.-S. Chao 1403 Philippines KF289640 KF289510 TAIF
P. mucronulata Y.-S. Chao 1410 Philippines KF289641 KF289511 TAIF
P. pacifica P.I. Forster 27643 Australia KF289647 KF289517 MEL
P. kawabatae Y.-S. Chao 1637 Vietnam KF289655 KF289525 TAIF
P. giasii C. R. Fraser-Jenkins 30176 Bangladesh KF289660 KF289530 TAIF
P. kathmanduensis C. R. Fraser-Jenkins FN35 Nepal KF289663 KF289533 TAIF
P. otaria C. R. Fraser-Jenkins FN26 India KF289666 KF289536 TAIF
P. roseililacina C. R. Fraser-Jenkins FN31911 Nepal KF289669 KF289539 TAIF
P. biaurita P.-F. Lu 17285 Taiwan KF289676 KF289546 TAIF
P. argyraea C. R. Fraser-Jenkins FN145 India KF289684 KF289554 TAIF
P. aspericaulis C. R. Fraser-Jenkins FN36 India KF289685 KF289555 TAIF
P. assamica C. R. Fraser-Jenkins FN5 Nepal KF289686 KF289556 TAIF
P. khasiana C. R. Fraser-Jenkins FN129 India KF289688 KF289558 TAIF
P. praetermissa C. R. Fraser-Jenkins FN64 India KF289692 KF289562 TAIF
P. subindivisa C. R. Fraser-Jenkins FN266 Bhutan KF289700 KF289570 TAIF
P. asperula Y.-C. Liu 9870 Philippines KF289702 KF289572 TAIF
P. dataensis Y.-C. Liu 9973 Philippines KF289703 KF289573 TAIF
P. catoptera G. Rouhan 301 Madagascar KF289714 KF289584 P
P. humbertii F. Rakotondrainibe 5965 Madagascar KF289718 KF289588 P
P. confusa Y.-M. Huang 20061128-A India KF289726 KF289596 TAIF
P. flava M. Kurutok 23 Sabah KF289731 KF289601 KEP
P. perrottei C. R. Fraser-Jenkins FN215 Nepal KF289736 KF289606 TAIF
P. grevilleana Y.-S. Chao 770 (diploid) Taiwan HM582644 KF289484 TAIF
P. venusta Y.-S. Chao 873 Taiwan HM582650 KF289486 TAIF
P. longipinna P.-F. Lu 11383 Taiwan HM582603 KF289495 TAIF
P. laurisilvicola Y.-S. Chao 1848 Japan KF289738 KF289608 TAIF
P. kiuschiuensis Y.-S. Chao 1852 Japan KF289739 KF289609 TAIF
P. satsumana Y.-S. Chao 1853 Japan KF289740 KF289610 TAIF
P. oshimensis Y.-S. Chao 1881 Japan KF289741 KF289611 TAIF
P. yakuinsularis Y.-S. Chao 1906 Japan KF289742 KF289612 TAIF
P. boninensis Y.-S. Chao 1941 Japan KF289743 KF289613 TAIF
P. natiensis Y.-S. Chao 1835 Japan KF289744 KF289614 TAIF
P. arisanensis Y.-S. Chao 1621 Vietnam KF289677 KF289547 TAIF
P. latipinna P.-F. Lu 24585 Taiwan MF416317 MF416323 TAIF
P. latipinna P.-F. Lu 25108A Taiwan MF416318 MF416324 TAIF
P. latipinna P.-F. Lu 25108B Taiwan MF416319 MF416325 TAIF
P. wulaiensis P.-F. Lu 26667-1 Taiwan MF537503 MF537504 TAIF
P. fauriei var. minor Y.-S. Chao 2078 Taiwan MF416320 MF416327 TAIF
P. fauriei var. fauriei Y.-S. Chao 2083 Taiwan MF416321 MF416328 TAIF
P. latipinna Y.-S. Chao 2092 Taiwan MF416322 MF416326 TAIF