Botanical Studies (2011) 52: 211-218.
SYSTEMATICS
Cotoneaster rosiflorus (Rosaceae), a new species from Taiwan
Kun-Cheng CHANG1, Chih-Chiang WANG2, Shu-Lin DENG3,4, Yoshiko KONO5, Fu-Yuan LU3, and
Ching-I PENG5*
lDepartment of Forestry, National Chung Hsing University, Taichung 402, Taiwan
2Department of Tourism and Leisure Management, National Penghu University, Penghu 880, Taiwan
3Graduate Institute of Forestry and Natural Resources, National Chiayi University, Chiayi 600, Taiwan
4Chungpu Research Center, Taiwan Forestry Research Institute, Chiayi 600, Taiwan
5Herbarium (HAST), Biodiversity Research Center, Academia Sinica, Nangang, Taipei 115, Taiwan
(Received December 8, 2009; Accepted November 8, 2010)
ABSTRACT. Cotoneaster rosiflorus K. C. Chang & F. Y Lu, a new species of Rosaceae subfam. Spirae-oideae, tribe Pyreae (formerly subfamily Maloideae) from Taiwan is described. A taxonomic treatment, line drawings, color photographs and pollen SEM micrographs are provided to illustrate the species. A somatic chromosome number of 2n = 68 is reported for the first time for C. rosiflorus. Cotoneaster rosiflorus occurs mainly at 2,500-3,500 m altitude in central Taiwan. It resembles C. morrisonensis vegetatively, but is distinct by the erect to slightly spreading pink petals, pink or reddish filaments, 3-5 styles, and 3-5 pyrenes.
Keywords: Cotoneaster horizontalis; Cotoneaster morrisonensis; Cotoneaster rosiflorus; Maloideae; New species; Rosaceae; Spiraeoideae; Taiwan.
INTRODUCTION
Cotoneaster (Rosaceae) is a genus of about 90 species widespread in temperate Asia (except Japan), Europe and North Africa (Yu and Lu, 1974; Lu and Brach, 2003), although other authors consider the number of species 260 (Mabberley, 2008) or closer to 400 (Fryer and Hylmo, 2009). In 1911 Hayata described from Taiwan three species of Cotoneaster, namely C. formosana Hayata, C. koidzumii Hayata and C. taitoensis Hayata, all of which were synonymized under Pyracantha koidzumii (Hayata) Rehd (Ohashi, 1993). Subsequently, Hayata, (1913, 1915) published three species, C. konishii Hayata, C. morrisonensis Hayata and C. rokujodaisanensis Hayata, in 'Icones Plantarum Formosanarum.' Masamune (1932) treated C. rokujodaisanensis as a synonym of C. mor-risonensis, which was followed by Kanehira (1936), Li (1963), Liu and Su (1977) and Ohashi (1993). Only two species of Cotoneaster, C. konishii and C. morrisonensis, were recognized in the second edition of Flora of Taiwan (Ohashi, 1993). Hsieh and Huang (1997), in their revision of the genus Cotoneaster in Taiwan, considered C. mor-risonensis and C. rokujodaisanensis distinct, and added a neglected species, C. horizontalis Decne. In 2001 Fryer and Hylmo published a new species, C. hualiensis J. Fryer & B. Hylmo, from this island. Lai and Hsieh (2001, 2003)
added two neglected species, C. subadpressus Yu and C. apiculatus Rehd. & Wils., to the flora of Taiwan. Lu et al. (2005) documented another new distributional record, C. dammeri C. K. Schneid., in Taiwan. More recently, Chang et al. (2009) clarified the taxonomic confusion involving Photinia kudoi Masamune and recognized Cotoneaster bullatus Bois for Taiwan. Fryer and Hylmo (2009) pub­lished a new species, C. nantouensis Fryer & Hylmo, from Taiwan. We checked the protologue and examined a type specimen image of C. nantouensis (E. H. Wilson 10072!, MO) and hereby consider it to be a synonym of C. subad-pressus. In this study we report yet an additional new spe­cies, C. rosiflorus K. C. Chang & F. Y Lu, which occurs at 2,500-3,500 m altitude in Central Mountain Ranges on this island. The number of recognized species of Cotoneaster in Taiwan has increased drastically in the recent decade, which is unusual for a woody genus on this island.
MATERIALS AND METHODS
Cryo scanning electron microscopy
Fresh leaves of Cotoneaster rosiflorus K. C. Chang & F. Y. Lu, C. horizontalis Decne and C. morrisonensis Hayata were dissected and attached to a stub. The samples were frozen with liquid nitrogen slush, then transferred to a sample preparation chamber at -160°C. After 5 min, when the temperature rose to -130°C, the samples were frac­tured. The samples were etched for 10 min at -85°C. After coating at -130°C, the samples were transferred to the
*Corresponding author: E-mail: bopeng@sinica.edu.tw.
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SEM chamber and observed at -160°C with a cryo scan­ning electron microscope (FEI Quanta 200 SEM/Quorum Cryo System PP2000TR FEI). Vouchers are deposited at HAST (Cotoneaster rosiflorus, Peng 21493) and TCF (C. horizontalis: Chang 2486; C. morrisonensis, Chang 2684).
Chromosome cytology
Root tips were pretreated with 2 mM 8-hydroxy-quinoline solution at 15-18°C for 8-9 h and fixed with ethanol-acetic acid (3:1) at about 4°C for over 24 h. Fixed materials were macerated and stained in 10:1 mix­ture of 2% acetic orcein and 1N HCl for chromosomal observation. The plant used in this study (Cotoneaster rosiflorus, Peng 21493, HAST) was collected in the field
and cultivated in the experimental greenhouse of Aca-demia Sinica.
Pollen observation
The anthers of fresh flowers of Cotoneaster rosiflorus (Chang 2487, holotype at HAST) were carefully removed with tweezers, then sequentially dehydrated with alcohol (30%, 5 min; 40%, 5 min; 50%, 5 min; 60%, 5 min; 70%, 5 min; 80%, 5 min; 95%, 5 min). The anthers were then broken with a dissection needle under the stereomicro-scope to scatter the pollen. Pollen were kept in a desiccator with silica gel over night. Finally, the pollen was ob-served under the scanning electron microscope (HITACHI S-3000N) and photographs were taken after gilt.
Figure 1. Cotoneaster rosiflorus K. C. Chang & F. Y. Lu. A, Habit; B, Leaf, adaxial surface; C, Leaf, abaxial surface; D, Flower; E, Dissected flower; F, Petal; G, Stamens; H, Styles; I, Pome; J, Pome, dissected to show pyrenes.
CHANG et al. — Cotoneaster rosiflorus, a new species from Taiwan
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Figure 2. Cotoneaster rosiflorus K. C. Chang & F. Y Lu. A, Habit, flowering; B, Habit, fruiting; C, Flower, viewed from above; D, Flower, longitudinal section; E, Petal; F, Immature pome; G, Immature pome, longitudinal section; H, Immature pome, cross section; I, Ripe pome; J, Pyrenes.
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NEW SPECIES
Cotoneaster rosiflorus K. C. Chang & F. Y Lu, sp. nov.-
TYPE: TAIWAN. Nantou Hsien, Hsinrengang, 2,300 m altitude. 23 June 2004, K. C. Chang 2487 (holotype:
HAST; isotypes: A, MO, TCF).粉紅花舖地蜈蚣
(Figures 1, 2, 3A-D)
Haec species affinis Cotoneastro morrisonensi Hayata, sed planta habitu semi-decidua, foliis chartaceis usque subcoriaceis, petalis erectis, roseis, stylis (3-)5, pyrenis (3-)5 differt.
Semi-evergreen low shrubs less than 50 cm tall, branch-lets grayish brown to grayish, terete, initially yellowish appressed-pilose, glabrescent. Leaves chartaceous, rotund,
Figure 3. SEM-micrographs of leaf blade of Cotoneastesr rosiflorus (A-D), C. horizontalis (E-H) and Cotoneaster morrisonensis (I-L). A, E, I, Adaxial surface; B, F, J, Cross section; C, G, K, Abaxial surface, showing trichomes; D, H, L, Abaxial surface, showing sto-matal apparatus.
CHANG et al. — Cotoneaster rosiflorus, a new species from Taiwan
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or subglobose, 5-8 mm long, 4-6 mm across, initially appressed-pilose, glabrous when old, the ripe pome sum­mit rounded or angular, base rounded; sepals persistent on summit of ripe fruit, greenish-brown to dark red, thick and fleshy, coalescent; pedicels 1.5-4 mm, pilose. Pyrenes 3-5, 4-5 mm long, 2-2.7 mm across, outer surfaces strongly ru­gose, summit smooth, style or style scar remains 3/4 from base. Flowering May to July; fruiting June to December.
Distribution and Habitat. Endemic to Taiwan, oc­curring in relative abundance at 2,500-3,500 m altitude in the Central Mountain Range, usually on semi-shaded arid sites, over conglomerate, in fields or rocky places, at margins of broadleaved or pine forests (Figure 4). Coto-neaster rosiflorus is not in the threatened category. Coto­neaster rosiflorus is a creeping shrub. Its branches always hang down or climb in the crags. It is usually found on semi-open, gentle to steep eastern and southern aspects, often co-occurring with Pinus taiwanensis Hayata, Mis-canthus sinensis Anders., Rubus rolfei Vidal or Cotoneas-ter apiculatus Rehd. & Wils., etc. in the field.
Chromosome cytology. A somatic chromosome number of 2n = 68 was observed for Cotoneaster rosiflorus (Fig­ure 5). The 68 chromosomes at mitotic metaphase varied gradually from 0.4 to 1 fim in length. As positions of the centromeres were not ascertainable, the karyotype analysis was not possible.
The genus Cotoneaster belongs to subfamily Spirae-oideae tribe Pyreae subtribe Pyrinae (formerly subfam­ily Maloideae. see Potter et al., 2007; Campbell et al., 2007), which has a basic chromosome number of x =
17 (Raven, 1975; Campbell et al., 2007). Somatic chro­mosome numbers of 2n = 34, 51, 68, 85 and 102 (e.g.,
Kroon, 1975; Klotz and Krugel, 1983; Krugel, 1992;
Zhou and Wu, 1999; Zhou et al., 2000) were known in plants of the genus, representing diploids, triploids, tet-raploids, pentaploids and hexaploids respectively. Our report of 2n = 68 suggests that C. rosiflorus is a tetra-ploid. This is the second cytological report on Taiwanese
Figure 4. Latitudinal and altitudinal distributions of Cotoneaster rosiflorus in Taiwan.
ovate or elliptic, 0.4-3 cm long, 0.4-1.7 cm wide, apex api-culate, rounded, obtuse or acute, rarely emarginated, base obtuse to broadly cuneate, margins recurved; upper surface initially appressed pilose on veins, glabrous, shiny, with slightly impressed veins, veinlets inconspicuous when old, lower surface persistently densely tomentose; veins 3-5 pairs; petiole 1.6-3 mm long, pilose; stipules mostly cadu­cous, lanceolate, 1.4-2.3 mm long, abaxially puberulent. Inflorescence usually 1-flowered, rarely 2- or 3-flowered, flower 7-9 mm long (including hypanthium), 3-5 mm across, pedicel 1.7-4 mm long, densely tomentose; bracts linear-lanceolate or lanceolate, abaxially puberulent, 2-3 mm long. Hypanthium campanulate, abaxially puberulent; sepals triangular, acute at apex, 1.3-1.5 mm long, 1.2-1.7 mm wide, puberulent abaxially and on margins. Petals 5, erect or slightly spreading, pink, subrotund, 2.5-4 mm long, 1.9-2.8 mm wide, margin erose dentate, slightly clawed at base. Stamens 7-18, ca. 2.4 mm long, filaments pink, 1.5-2.2 mm long, anthers ellipsoid, purple or pur­plish red, ca. 0.5 mm long. Ovary pilose apically; styles (3-)5, free, ca. 2.4 mm long, glabrous. Fruits light yellow­ish orange to red, light red at maturity, obovoid, ellipsoid,
Figure 6. SEM-micrographs of pollen grain of Cotoneaster rosi­florus. A, Equatorial view; B, Polar view; C, Ornamentation on the equatorial view.
Figure 5. Somatic chromosomes at mitotic metaphase of Coto­neaster rosiflorus (2n = 68, from Peng 21493, HAST).
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Cotoneaster. Fryer and Hylmo (2001) indicated triploid chromosome count for C. hualiensis that occurs also in Central Mountain Range.
Additional specimens examined. TAIWAN. HSINCHU HSIEN: Luchangdashan ('Mt. Rokujodaishan'), 10 Nov 1983, Lu 13460 (TAIF), 3 Aug 2003, Chang 1544 (CHIA), 11 Aug 2003, Chang 1611 (CHIA), 13 Jun 2004, Chang, 2465, 2486 (CHIA). TAICHUNG HSIEN: Mt. Shen-machenshan, 25 Jun 1994, Wang 968 (HAST); Mt. Shaun-shan, 13 Jul 1983, Lin &Hsieh s. n. (TAIF). HUALIEN HSIEN: Tayuling, 4 Dec 1997, Liu 775 (TAIF); Piluchi, 17 Jul 1980, Ou et al. 5538 (TCF), 9 Nov 1986, Yang & Yang 3200 (TNM), 5 Jun 1998, Su s. n. (TNM), 4 Sep 1988, Lu 12806 (TAIF); Pilu log road, 4 Dec 1991, Wang s. n. (TNM), 7 May 2000, Chung 2777 (TAIF). NANTOU HSIEN: Yuanfeng, 26 Jun 1979, Tu s. n. (TCF), 17 Oct 1982, Ohashi et al. 12553 (TAI), 17 Jul 1996, Lin 179 (TAIF), 8 Aug 2003, Chang 1594 (CHIA), 26 Oct 2003, Chang 2175 (CHIA); Hsinrengang, 15 June 1993, Chiu 1955 (HAST, TNM), 13 July 1993, Wang w00050 (TNM), 12 Apr 2000, Cheng 3018 (TAIF), 5 Jul 2000, Cheng 3296 (TAIF), 28 Jan 2003, Chang 583 (CHIA), 27 May 2004,
Wang s. n. (TCF), 11 June 2008, Peng 21493 (HAST); Tsuifeng, 20 Jun 1981, Ou et al. s. n. (TNM), 22 Sep 1996, Lin s. n. (TAIF); Yunhai, 13 Oct 1992, Liu 15 (HAST),
27 Nov 1998, Wang et al. 3823 (TNM), 8 May 2004, Tunghai Collecting Team 9305097 (TNM); Tienchih, 25 Aug 1984, Tateishi et al. 18120 (TAI), 2 Jan 2003, Chang
418 (CHIA); Mt. Nengkaoshan, 23 Aug 1929, Sasaki s. n.
(TAIF); Tungpu, 19 May 1969, Hsu 5364 (TAI). CHIAYI
HSIEN: Tatachiaanpu, 11 Nov 1990, Wu et al. s. n. (TNM), 17 May 1991, Lin & Lin s. n. (TNM), 7 Aug 1991, Leu 1179 (HAST); Mt. Lulmshan, 15 Jul 1990, Ou et al. s. n.
(TNM), 9 Nov 1986, Chen et al. 900008 (TNM), 25 Dec
2002, Chang 344 (CHIA), 18 Jun 2004, Wu s. n. (CHIA). KAOHSIUNG HSIEN: Tienchih, 21 May 1992, Wang 1150 (HAST).
Notes. Cotoneaster rosiflorus is similar to C. morri-sonensis and C. horizontalis vegetatively. A comparison of salient morphological features of the three species is shown in Figure 3 and Table 1. Six species of Taiwanese Cotoneaster assume prostrate or decumbent habit. The rest (four species) are erect shrubs or small trees. To aid in identification, we provide a key to the six creeping spe-
Table 1. Comparisons of Cotoneaster rosiflorus, C. horizontalis and C. morrisonensis.
C. rosiflorus
C. horizontalis
C. morrisonensis
Habit
Semi-evergreen
Deciduous
Evergreen
Leaf
Shape
Rotund, ovate or elliptic
Obovate, ovate, broadly elliptic to subrotund
Elliptic to oblong
Size (mm)
4-30 x 4-17
5-14 x 4-9
6-18 x 3-12
Margins
Recurved
Plane
Recurved
Adaxial surface sculpture
(SEM)
Shortly roughly striate (Figure
3A)
Shortly roughly striate (Figure 3E) Finely striate (Figure 3I)
Palisade tissue
1 or 2 layers (Figure 3B)
1 or 2 layers (Figure 3F)
3 layers (Figure 3J)
Abaxial surface sculpture
(SEM)
Forked striate (Figure 3D)
Not striate, slightly rugose (Figure
3H)
Shortly forked striate (Figure
3L)
Abaxial trichome
Sparse (Figure 3C)
Glabrous (Figure 3G)
Relatively dense (Figure 3K)
Stomata
Sparse (Figure 3D)
Sparse (Figure 3H)
Dense (Figure 3L)
Flower
Petals
Erect or slightly spreading
Erect
Spreading
Color
Pink
Red
White
Filaments
Pink or reddish
Red
White
Anthers
Purple or purplish red
White
Purple
Styles
(3-) 5
2 (3)
2 (3)
Fruit
Shape
Obovoid, ellipsoid or subglobose Obovoid to subglobose
Ellipsoid or obovoid
Pyrenes
(3-) 5
2 (3)
2 (3)
Pollen
Ornamentation
Prominently rugose (Figure 6)
Prominently rugose and perforate (Hsieh & Huang, 1997: Figures 3-8)
Striate (Hsieh & Huang, 1997:
Figures 9-14)
CHANG et al. — Cotoneaster rosiflorus, a new species from Taiwan
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cies of Cotoneaster currently known in Taiwan. Fryer and Hylmo (2009) chose a Taiwanese endemic species C. rokujodaisanensis as a type to establish a new section Rokujodaisanense. Unfortunately, their description of C. rokujodaisanensis tallies with C. rosiflorus, the new spe­cies we reported in this article. Upon detailed examination of the type collection (fruiting specimen that is devoid of flowers) of Cotoneaster rokujodaisanensis at TAIF and plants in flower collected from the type locality, we are
convinced that C. rokujodaisanensis has leathered and mostly elliptic leaves and white, spreading petals, which did not conform to the description of Fryer and Hylmo (2009). The color photograph of C. rokujodaisanensis in their book turned out to be bona-fide C. rosiflorus. Co-toneaster rosiflorus is cultivated massively by Hualien District Agricultural Research and Extension Station in Taiwan as a promising ornamental plant.
KEY TO CREEPING TAXA OF COTONEASTER IN TAIWAN
1a. Petals fully spreading, white; filaments white; plants evergreen; veinlets on upper leaf surface reticulate or not.
2a. Leaf veinlets markedly reticulate.
3a. Styles 4 or 5; fruits obovoid to globose; pyrenes 4 or 5 ............................................................................... C. dammeri
3b. Styles 2(3); fruits ellipsoid-ovoid or ellipsoid; pyrenes 2 (3)................................................................ C. morrisonensis
2b. Leaf veinlets not reticulate, only primary and secondary veins visible ................................................. C. rokujodaisanensis
1b. Petals erect or slightly spreading, red or pink; filaments red or pink; plant deciduous or semi-evergreen; veinlets on leaf upper surface not reticulate.
4a. Anthers purplish; filaments pink; styles (3-)5; pyrenes (3-)5 ............................................................................C. rosiflorus
4b. Anthers white; filaments red; styles 2(3); pyrenes 2(3).
5a. Leaves chartaceous to subcoriaceous; stems horizontally spreading, distichously branched .................. C. horizontalis
5b. Leaves thick coriaceous; stems tufted on ground, irregularly branched ................................................ C. subadpressus
Acknowledgments. We thank Qiner Yang (IBSC) for improving the Latin diagnosis; Anthony R. Brach (A/ GH) for providing literature and the image of holotype specimen of Cotoneaster nantouensis; curators of the herbaria cited who facilitated our examination of Cotoneaster specimens; Li-Fen Lii for the handsome line drawings; and Shin-Ming Ku for helpful discussion and technical assistance on the color plate and SEM micrograph plate. This work was supported in part by grants from Academia Sinica, Taiwan to Ching-I Peng.
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台灣產枸子屬(薔薇科)一新種：粉紅花舖地蜈蚣
張坤城1 王志強2 鄧書麟3'4 河野淑子5 呂福原3 彭鏡毅5
1國立中興大學森林學系
2國立澎湖科技大學觀光休閒學系
3國立嘉義大學林業暨自然資源研究所
4林業試驗所中埔研究中心
5中央研究院生物多樣性研究中心植物標本館(HAST)
本文報導台灣產枸子屬(Cotoneaster) 一新種：粉紅花舖地蜈蚣（C. rosiflorus K. C. Chang & F. Y.
Lu)，並提供形態描述、手繪圖、花果照片及葉片與花粉的掃瞄式電子顯微鏡照片以供辨識。本種主要
分布於台灣中部海拔2,500-3,500 m之山區，其形態近似玉山舖地蜈蚣(C. morrisonensis)或平枝枸子(C.
horizontalis)，但本種的花瓣直立或微開展、花瓣與花絲粉紅色、及花柱與小核3-5可與之區分。
關鍵詞：平枝枸子；玉山舖地蜈蚣：粉紅花舖地蜈蚣；蘋果亞科；新種；薔薇科；繡線菊亞科；台灣。