J Haflori Bot. Lab. No. 100: 69- 76 (Aug. 2006)

SHEVOCKIA (), A NEW GENUS WITH TWO SPECIES FROM SOUTHEAST ASIA

1 10HANNES ENROTH AND MENGCHENG 112

ABSTRACT. Shevockia Enroth & M.C. Ji gen. novo (Neckeraceae, ) is described from SE Asia. It has two species, S. inunctocarpa Enroth & M.C. Ji sp. novo (generitype) from Yunnan Province of China, and S. anacamptolepis (Miil!. Ha!.) Enroth & M.C. Ji comb. nov., which has a wider distribution in Asia. Shevockia is morphologically close to , but differs especially in several gametophyte characters: I) distinctly asymmetric leaves, 2) often undulate upper parts of stem leaves (less so in branch leaves), 3) relatively weak and short, sometimes bifurcate costa, and 4) presence of numerous large, leaf-like pseudoparaphyllia around the branch primordia. Shevockia in­ unctocarpa is illustrated by line drawings and a photograph.

Enroth (1994a) recognized 23 genera in the moss family Neckeraceae. That number has changed and is going to change more in the future, as more rigorous phylogenetic studies are carried out. For example, the results of Tsubota et al. (2002) suggest that the genus Homaliadelphus, placed in the Neckeraceae since its inception, belongs elsewhere and is closely related to Bissetia, another genus which has previously been placed in the Neckeraceae (cf. Enroth, 1992). On the other hand, genera such as Taiwanobryum and Echinodium, previously placed in other families, seem to be close to some of the ­ ceous genera (Buck et aI., 2000; Tsubota et aI., 2002). A close relationship between the families Neckeraceae and was shown by Quandt and Huttunen (2004), and the detailed phylogeny and systematics of the genera are under study by those authors in cooperation with the first author of the present paper. Not only have studies on previously known taxa shed new light on the and systematics of the Neckeraceae, but taxa new to science are also being discovered and de­ scribed. The PhD-project of the second author yielded four new Asian species of Neckera, one of which has so far been published (Ji and Enroth, 2005). The novelty reported in this paper resulted from examination of fresh material collected in Yunnan Province in South China. That material also solved a taxonomic problem regarding generic placement of a pe­ culiar SE Asian species, Pinnatella anacamptolepis. The genus Pinnatella was mono­ graphed by Enroth (1994b).

Shevockia Enroth & M.C. Ji , gen. novo Genus haec Pinnatellae proxime, sed praecipue foliis asymmetric is et aliquantum un­ dulatis, costa comparate brevi et debili, pseudoparaphyllis numerosis et longis, et seta laevi differt. Generitype: Shevockia inunctocarpa Enroth & M.C. Ji.

I Department of Biological and Environmental Sciences and the Botanical Museum, P.O. Box 7, F1-000 14 University of Helsinki, Finland. 2 College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, Jiangsu, China. 70 J. Hattori Bot. Lab. No. 100 2 0 0 6

Shevockia inunctocarpa Enroth & M.C. li, sp. novo (Figs. 1-4) Species praesertim Pinnatellae acacamptolepi subsimilis, sed statura robustiore, folia caulorum et ramorum manifeste majora et folia stipitis patentia (non squarrosa). gregarious, slightly glossy, younger parts sordid-green, older parts often brown­ ish, stipitate-frondose. Stolons creeping, bearing small, pale, somewhat spreading leaves and partly covered with a dark rhizoid tomentum, rhizoids smooth, reddish brown, profuse­ ly branched. Stems to ca. 10 cm high, sometimes unbranched, usually irregularly or subpin­ nately branched, lower parts with rhizoid tomentum between stipe leaves; in cross-section elliptic, with 2- 3 layers of small cortical cells with thick, brownish walls, grading to slight­ ly larger and thinner-walled subcortical cells and these to still larger medullary cells with thinner walls; without a central strand. Flagelliform branches sometimes produced, ftexu­ ous, microphyllous and easily detachable. Stipe leaves to ca. 2.0 X 1.2 mm, broadly ovate, the base with broad and long decurrencies, apex acute; margins recurved, entire or slightly denticulate near apex; costa faint, often double; laminal cells elongate, linear-oblong throughout and with relatively thin, porose walls. Stem leaves somewhat complanate and spreading, basal parts plicate, upper parts often undulate, asymmetric, ovate, ca. 2.8(- 3.4) X 1.3(- 1.5) mm, base decurrent, decurrencies to ca. 400.um long, apex acute. Branch leaves similar but relatively narrower, ca. 1.4(- 1.7) XO.5(-O.7) mm, margins often broadly recurved to above midleaf, dentate near apex, entire elsewhere. Costa single but

Fig. I. Shevockia inunctocarpa. a and b. Habit sketches. J. ENROTH & M. JI: Shevockia, a new moss genus from southwest Asia 71

Fig. 2. Shevockia inunctocarpa. a. Portion of shoot woth sporophytes. b. Portion of stipe. c. Mature sporophyte and perichaetium. d. Operculate capsule (damaged). e. Calyptra. 72 1. Hattori Bot. Lab. No. 100 200 6

Fig. 3. Shevockia inunctocarpa. Sporophytes on branch.

quite frequently unevenly furcate, relatively faint, reaching to midleaf, sometimes up to 3/4 leaf length. Leaf cells smooth, relatively thick-walled, walls rather distinctly porose throughout; apical laminal cells mostly (sub )rhomboid to oval or oblong, 15-40 X 7- 12 f.1m, median laminal cells mostly oblong, 40- 70 X 6- 12 f.1m, basal laminal cells oblong, 45- 80 X 9- 12 f.1m, a few rows of cells in leaf margins shorter than the corresponding lami­ nal cells, alar cells fairly indistinct, short-rectangular to irregular. Pseudoparaphyllia nu­ merous, foliose, mostly lanceolate, to ca. 500 J.lm long. Presumably dioicous (perigonia not seen). Post-fertilization inner perichaetial leaves to ca. 2.5 mm long, mostly unicostate, from ovate base abruptly narrowed above midleaf to a lanceolate, spreading acumen with more or less dentate margins. Vaginula bearing nu­ merous archegonia and long, filiform, hyaline paraphyses. Seta ca. 3 mm long, yellow to brownish, smooth. Capsules erect, broadly ovoid, usually strongly lustrous when young, chestnut brown, ca. 1.6 X 1.0 mm; apophysal stomata none; exothecial cells moderately in­ crassate, variable in shape, from nearly isodiametric to oblong or irregularly angular; annu­ lus none. Peristome double, inserted below mouth of capsule; exostome teeth lanceolate, grayish-yellow, ca. 400 f.1m long, with a distinct, slightly zig-zag median line, densely spiculose-papillose throughout; endostome segments nearly as long as exostome teeth, slightly brighter yellow, narrowly lanceolate, with elongate median perforations, more strongly spiculose-papillose than exostome teeth; basal membrane ca. 50- 60 f.1m high, cilia none. Spores 22 f.1m diameter, faintly papillose. Operculum 0.6 mm high, obliquely rostrate from a conic base. Calyptra 1.5 mm, smooth, cucullate. J. ENROTH & M. JI: Shevockia, a new moss genus from southwest Asia 73

60 Jllll I c

e

30 Jll11 I

N 9 h Fig. 4. Shevockia inunctocarpa. a. Stem leaves. b & d. Stipe leaves (b from lower part of stipe, d from upper). c. Branch leaf. e. Pseudoparaphyllia. f. Apex of stem leaf. g. Margin at midleaf. h. Median laminal cells. i. Alar region. j. Post-fertilization perichaetialleaves. k. Peristome. Scale bars: 1 mm for a- d and j, 200 pm for e, 30 pm for f- i, 60 pm for k. 74 1. Hattori Bot. Lab. No. 100 200 6

Type: China: Yunnan Province, Fugong County, Gaoligong Shan Range, southern end of the Hengduan Shan. Nu Jiang (Salween River) watershed. Yaping Pass Road at large waterfall, 4.3 km above Shibali Forestry Field Station, 27°1O'38.4"N, 98°45' 19.9"E, elev. 2700 m. Mixed forest with Aeer, Rhododendron, Tsuga , and Larix on slope from road to river. On hardwood trunk. 3 May 2004, lames R. Shevoek 25325 & Xuezhong Fan , mixed with Neekera pennata (holotype KUN!, isotypes CAS! , E! , H!, msc, KRAM! , MO! , NICH! , NY1 , UC!). Etymology: We are pleased to name the genus after Mr. Jim Shevock, who collected the specimens and sent them to us for study. The epithet is composed of the Latin words in­ unctus = anointed, glossy as if oiled (Steam, 1986), and -carpus = -fruit, -fruited (in Greek comp.), referring to the peculiarly lustrous capsules.

Shevockia anacamptoiepis (Miil\' Ha\.) Enroth & M.C. Ji, comb. novo Neckera anacamptolepis Mull. Hal., Syn. Muse. Frond. 2: 663. 1851 ; Thamnium anacamptolepis (Mull. Hal.) Kindb. , Hedwigia 41 : 251. 1902; Pinnatella anacamptolepis (Mull. Hal.) Broth., Nat. Pflanzenfam. 1(3): 857. 1906; Porotriehum anaeamptolepis Mull. Hal. ex M. Fleisch., Musci Fl. Buitenzorg 3: 913 . 1908, nom. inval. in synon. Lectotype (designated by Enroth 1994b): Indonesia, Java, Blume s. n. (L! "hb. AI. Br."). Illustrations: Noguchi (1989: 723 , fig . 319B), Enroth (I 994b: 47, figs. 3d, 18). For synonymy see Enroth (1994b). Distribution (Enroth 1994b): Sri Lanka, India?, China, Malaysia, Thailand, Vietnam, Indonesia, Philippines, Papua New Guinea, Taiwan, Japan (Kyushu, Shikoku). Shevockia appears to be closely related to Pinnatella, but S. inunctocarpa and S. anacamptolepis share some taxonomically important gametophyte characters not encoun­ tered in Pinnatella. The characters are I) distinctly asymmetric leaves, 2) often undulate upper parts of stem leaves (less so in branch leaves), 3) relatively weak and short, some­ times bifurcate costa, and 4) presence of numerous large, leaf-like pseudoparaphyllia around the branch primordia. These characters therefore rendered S. anacamptolepis quite anomalous in Pinnatella, but at that time (Enroth, 1994b) the first author did not know where else to place it. Discovery of S. inunctocarpa solved the long-lasting problem in an unexpected yet most satisfactory way, even if the sporophytes of S. anacamptolepis remain unknown: these two species belong in a separate genus, for which there was no previously published name available. A very peculiar feature in Shevockia inunctocarpa we have not observed elsewhere in the Neckeraceae (or in any moss in fact) are the often strongly lustrous capsules, appearing almost varnished (cf. Fig. 3). Not all capsules have this feature, however, and it seems that the lustre often, but not always, vanishes when the capsules grow older. Although Shevockia anacamptolepis was nested within Pinnatella in the c1adistic analysis (Enroth, I 994b; Hyvonen and Enroth, 1994), the support for the clades was weak and in the semi strict and strict consensus trees the species was in an unresolved group of taxa. The above-mentioned gametophytic synapomorphies between S. inunctocarpa and S. anacamptolepis would be uninformative (autapomorphies) if the latter species was placed in Pinnatella. The characters 1- 3 listed above, anomalous in Pinnatella, are quite common in the genus Neckera. However, both species of Shevockia have a completely different habit from 1. ENROTH & M. 11: Shevockia, a new moss genus from southwest Asia 75

Neckera, mainly the distinctly stipitate plants, with spreading (s. inunctocarpa) to squar­ rose (s. anacamptolepis) stipe leaves with re curved margins. There is in Asia a group of stipitate, robust Neckera species (cf. Enroth, 1996), but the plants have obtuse or truncate, more strongly and regularly undulate, and almost symmetric leaves, and their stipe leaves are always tightly appressed, have obtuse-apiculate rather than acute apices, and plane mar­ gins. Stipe leaf characters are taxonomically important at the generic and also partly at the specific level in the Neckeraceae, but they have been largely neglected. The resemblance between Shevockia and Asian stipitate species of Neckera is but superficial. Shevockia inunctocarpa shoud also be compared with Noguchiodendron sphaero­ carpum (Nog.) Ninh & Pocs, the single species of its genus, known from the Himalayan re­ gion (Bhutan, Nepal, Sikkim in India) and Thailand (Ninh and Pocs, 1981). It differs from Shevockia in several characters, such as the presence of a central strand in the stem, ap­ pressed stipe leaves with plane margins, incurved upper parts of the fronds, and strongly dentate leaf apices. Porotrichum fruticosum (Mitt.) Jaeg., which has a disjunct distribution in SE Asia (e.g. Gangulee, 1976, as Thamnobryumfruticosum), differs from S. inuncto­ carpa mainly in the ca. 2.3 mm long seta, non-undulate leaves, and much stronger costa. The latter two characters also easily distinguish Pfruticosum from S. anacamptoiepis.

NOTES ON THE HABITATS AND DISTRIBUTION OF SHEVOCKIA Most of the information regarding S. inunctocarpa was kindly provided by Mr. Jim Shevock. The Gaoligongshan Mountain Reserve in Yunnan, in which Shevockia inunctocarpa was found, covers an area of more than 400000 hectares ranging along the China-Myan­ mar border. It is a biodiversity hot spot in Yunnan and was declared part of the UNESCO World Biosphere's Protection Network in 2001. Elevation of the Gaoligongshan ranges from around 600 meters along the lower Nu Jiang (Salween River) to over 5800 meters near the border with Tibet, the mean elevation being ca. 2600 m. The steepness of the area has provided for a high level of intact or minimally disturbed landscapes. In the Reserve there occur 4303 species of seed plants, 699 species of vertebrate ani­ mals and 1690 species of insects. Of these, 61 species and 81 animal species are list­ ed in China's Redbook as rare and endangered. The area has probably served as a Pleistocene refugium, which partly accounts for the high species diversity (sources: http://www.ens-newswire.comlens/may2005/2005-05-04-0 l.asp; http://whc.unesco.org / archive/advisory_body3valuationll083.pdt). The habitat of Shevockia inunctocarpa is a river drainage basin with a dense mixture of hardwoods (especially Acer and Rhododendron spp.) and an overstory of Picea, Larix and scattered Tsuga chinensis. A shrubby bamboo (Fargesia sp.) is fairly common in the understory. cover is abundant both on the ground and as epiphytes. The type lo­ cality is about 4 km above an old logging camp called Shibali, yet the area appears fairly undisturbed. The vegetation at this altitude (2700 m) is essentially temperate, which is in accordance with the fact that S. inunctocarpa was growing mixed with the mainly northern hemisphere temperate species . Shevockia anacamptoiepis is a tropical-subtropical species (Enroth, 1994b). In the 76 J. Hattori Bot. Lab. No. 100 2 0 0 6 tropical regions of SE Asia it thrives approximately between 500 and 2300 meters in mon­ tane rain forests, and reaches the southern parts of Japan, where it has been found below lOO meters. Like S. inunctocarpa, it is an epiphyte, but occasionally grows also on rocks. Sporophytes are unknown for Shevockia anacamptolepis, thus it probably does not re­ produce or disperse by spores, while the specimens of S. inunctocarpa have numerous sporophytes. However, the former has a wide distribution in SE Asia, from Sri Lanka through Indonesia to New Guinea and through the Philippines to southern Japan, and the latter is known only from one locality. Shevockia inunctocarpa may be a climatic relict sur­ viving only in the Gaoligongshan Reserve. Shevockia anacamptolepis is known to repro­ duce only from flagelliform, easily detachable branches (Enroth, 1994b), which can clearly disperse only over short distances, thus its wide distribution in the tropical SE Asia must also be relict.

ACKNOWLEDGEMENT

We thank Or. Halina Bednarek-Ochyra for drawing figures I and 2, and Mr. Markku Lehtonen for taking the photograph of Shevockia inunctocarpa.

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