陳新艷, 馬良, 周承源, 陳彬, 陳世品*
(1. 三明市園林中心,福建 三明 365000;2. 福建衛(wèi)生職業(yè)技術(shù)學院,福州 350101;3. 福建農(nóng)林大學林學院,福州 350002;4. 華東野生瀕危資源植物保育中心,上海辰山植物園,上海 201602)
全世界漆樹科(Anacardiaceae)植物超過60 屬,600 多種[1]。漆樹屬[Toxicodendron(Tourn.) Mill.]是比較小的屬,約30 種,分布于亞洲東部和北美至中美,我國17 種,主要分布于長江以南各省區(qū)[2–4]。漆樹屬植物分泌的乳液含漆酚,易引起皮膚過敏。部分植物具有重要的經(jīng)濟價值,例如漆樹割取的天然生漆,在藥用、化工、工業(yè)生產(chǎn)以及工藝制品等方面都具有重要的作用[5–6]。
文獻記載福建省產(chǎn)漆樹屬植物共6 種:少葉漆(T.oligophyllumS.L. Tang, Liang Ma & S.P.Chen)[3]、刺果毒漆藤[T.radicanssubsp.hispidum(Engl.) Gillis]、毛漆樹[T.trichocarpum(Miq.) O.Kuntze]、木蠟樹[T.sylvestre(Sieb. et Zucc.) O.Kuntze]、漆[T.vernicifluum(Stokes) F. A. Barkl.]和野漆[T.succeda-neum(L.) O. Kuntze][7]。2019 年以來,筆者在全省進行植物資源調(diào)查期間,在福建省泰寧縣和邵武縣丹霞山發(fā)現(xiàn)1 種漆樹屬植物,在海拔300~500 m 的林下或灌木叢中。該種均為灌木狀, 樹形上與小漆樹相近,不同之處在于葉更長,被微毛,葉背紫紅色;花序長不超過葉長1/3;花瓣不具暗褐色脈紋。該種與木蠟樹、少葉漆相近,不同之處在于樹高不超過1.8 m,為落葉灌木;小葉常具粗鋸齒;聚傘總狀花序,長度不超過葉長1/3;花瓣不具暗褐色脈紋。經(jīng)查閱相關(guān)文獻和標本[3,6],與相關(guān)種進行了形態(tài)特征比較(表1)。結(jié)果表明,該種與漆屬其他種存在可以明顯區(qū)分的特征,可以作為獨立的種。同時,又開展了分子系統(tǒng)學研究。根據(jù)前人的分子系統(tǒng)學研究結(jié)果[1,3],本研究共選取29 種物種,其中外類群9 種;使用的分子標記包括核基因片段ITS 和2 個質(zhì)體基因片段(trnL-F 和ndhF), 共獲得26 條ITS、29 條trnL-F 和28 條ndhF;相關(guān)樣品的憑證標本和序列數(shù)據(jù)GenBank 登錄號見表2?;诤嘶騃TS 矩陣、質(zhì)體基因片段聯(lián)合矩陣和核基因質(zhì)體聯(lián)合矩陣,分別構(gòu)建最大似然法(maximum likelihood, ML)、最大簡約法(maximum parsimony,MP)和貝葉斯推斷法(bayesian inference, BI)系統(tǒng)發(fā)育樹。結(jié)果表明,除基于質(zhì)體基因片段聯(lián)合矩陣構(gòu)建的系統(tǒng)發(fā)育樹外,其余所有系統(tǒng)發(fā)育樹均支持該新種與木蠟樹以高支持率構(gòu)成姐妹關(guān)系(BSML≥89,BSMP≥85, PP≥0.99)。綜上,該種為漆樹屬一新種,現(xiàn)予以報道。
表1 紅背漆、木蠟樹、少葉漆和小漆樹的形態(tài)特征對比Table 1 Morphological comparison of Toxicodendron purpureum, T. sylvestre, T. oligophyllum and T. delavayi
表2 本研究所用物種的標本信息及GenBank 登錄號Table 2 Voucher information and GenBank accession No. for taxa used in this study
紅背漆新種 圖2, 3
圖1 基于核基因質(zhì)體聯(lián)合矩陣(ITS+trnL-F+ndhF)構(gòu)建的最大似然法(ML)、最大簡約法(MP)和貝葉斯推斷法(BI)系統(tǒng)發(fā)育樹。A: 核基因ITS 序列(局部);B: 質(zhì)體片段聯(lián)合矩陣(trnL-F+ndhF)(局部); 節(jié)點旁數(shù)字為自展值和貝葉斯后驗概率(BPML, BPMP, PP); –: MP/ML 和貝葉斯樹間的拓撲結(jié)構(gòu)存在不一致。Fig. 1 Phylogenetic tree obtained by ML, MP and BI analysis of nuclear and plastid combined matrix (ITS+trnL-F+ndhF). A: Separate combined nrITS (part of tree); B: Separate combined plastid (trnL-F+ndhF) (part of tree); the data near the nodes are bootstrap percentages and Bayesian posterior probabilities (BPML,BPMP, PP); –: Node is inconsistent between the topology of the MP/ML trees and the Bayesian tree.
圖2 紅背漆。A: 植株; B: 莖; C: 葉背; D: 葉; E: 葉柄基部; F: 雄花; G: 雌花; H: 果序。Fig. 2 Toxicodendron purpureum S. P. Chen, Xin Y. Chen & L. Ma. A: Plant; B: Stem; C: Leaf abaxially; D: Leaf; E: Petiole base; F: Male flower; G: Female flower: H: Infructescence.
圖3 紅背漆。A: 植株; B~C: 葉; D: 果序; E: 果實; F: 雄花; G: 雌花。(蘭雪靜繪圖)Fig. 3 Toxicodendron purpureum S. P. Chen, Xin Y. Chen & L. Ma. A: Plant; B-C: Leaf; D: Infructescence; E: Fruit; F: Male flower; G: Female flower. (Drawn by Lan Xuejing)
ToxicodendronpurpureumS. P. Chen, Xin Y.Chen & Liang Ma,sp.nov.
The new species is similar toT.delavayi, but is distinguished fromT.delavayiby leaves 25 cm (vs to 13 cm); leaflets papery, abaxially purplish red, rare green (vs abaxially glaucous, leaflet petiolule 1–2 mm);inflorescence as long as 1/3 pinnately compound leaf(vs as long as pinnately compound leaf); petals without brown featherlike venation pattern (vs petals with brown featherlike venation pattern).
China.Fujian Province (福建省), Shaowu County(邵武縣), Jiangshi Provincial Nature Reserve (邵武將石省級自然保護區(qū)), under the forest, elevation 327 m,2 May 2021,XinY.Chen&L.MaFAFU20210502(holotype: FJFC); China. Fujian Province (福建省),Taining County (泰寧縣), Zhukou Town (朱口鎮(zhèn)),Xufang Village (許坊村), elevation 340 m, 9 May 2023,B.Chen&Z.W.ZhuCB04558 (CSH).
落葉灌木,高0.4~1.8 m;具白色乳汁,在空氣中暴露會變黑;莖粗1~1.6 cm,具橢圓形突起小皮孔,小枝紫色或綠白色,常被白粉,被微柔毛。奇數(shù)羽狀復(fù)葉互生,長達25 cm;葉柄長4~8 cm,基部明顯枕狀;小葉3~5 對,對生,紙質(zhì),卵狀披針形或披針形,長3~6 cm,寬2~3 cm,先端尾尖或漸尖,基部偏斜,楔形,下延,全緣或1/3 處以上具粗鋸齒,兩面微被毛,上面綠色,成長葉背面紫紅色,稀綠色;側(cè)脈10~17 對,兩面突起;小葉柄長2~5 mm。雌雄異株,花序聚傘總狀,長5~8 cm,約為葉長的1/3,無毛,總梗纖細,長4~5 cm,苞片披針形,長約1 mm;花小,黃綠色,徑約2 mm,花梗無毛,長約2.5 mm;花萼無毛,裂片5,卵形,先端鈍,長約0.6 mm;花瓣長圓形,先端鈍,長約2 mm,開花時外卷;雄花的雄蕊長2 mm,花絲線形,伸出外卷的花冠外,花藥長圓形,長約1 mm,雌蕊退化僅余扁平痕跡;雌花的花藥狹卵形,花絲藏于花冠內(nèi),雌蕊比花瓣短,長約1.7 mm;花盤圓形,無毛,子房圓形,徑約1 mm。核果偏斜,壓扁,先端偏于一側(cè),長約8 mm,寬6~7 mm,外果皮薄,具光澤,無毛,中果皮蠟質(zhì),果核堅硬?;ㄆ? 月—5 月,果期6 月—8 月。
Deciduous shrub, 0.4–1.8 m tall; white latex,turning black upon air exposure; stem 1–1.6 cm in diam, with small oval lenticels, branchlets purple or green white, often whiting, puberulous. Leaves blade imparipinnately compound, to 25 cm; petiole 4–8 cm,base obvious occipital shape; leaflets 3–5, opposite,papery, leaflet blade ovate–lanceolate to lanceolate,3–6 cm×2–3 cm, apex acute or acuminate, base oblique, cuneate, decurrence, margin entire or 1/3 with coarse serrations on top, sparsely pubescent on both surfaces, adaxiall green, abaxially purplish red, rare green, lateral veins 10–17 pairs, prominent on both surfaces; leaflet petiolule 2–5 mm. Inflorescence cymose racemose, 5–8 cm, as long as 1/3 pinnately compound leaf, glabrous, peduncle slender, 4–5 cm;dioecious, bracts lanceolate, ca. 1 mm; flowers small,yellowish green, 2 mm in diam; pedicel glabrous, ca.2.5 mm; calyx glabrous, lobes ovate, obtuse apically,ca. 0.6 mm; petals oblong, ca. 2 mm, lobes ovate,obtuse apically, ca. 2 mm, roll out during flowering;stamens of male flowers ca. 2 mm; filaments linear,above corolla, anthers oblong, ca. 1 mm, pistil degeneration with only flat marks; anthers of female flowers narrowly ovate, filaments hidden within corolla, pistil shorter than petals, ca. 1.7 mm; disk glabrous, ovary globose, ca. 1mm in diam. Drupe oblique, compressed, apex eccentric, ca. 8 mm×6–7 mm,glabrous and lucid; epicarp thin, separating; mesocarp thick, waxy. Flowering in April to May and fruiting in June to August.
因該新種小葉葉背常為紫紅色,稀綠色,故命名為紅背漆(Toxicodendronpurpureum)。紅背漆目前只發(fā)現(xiàn)于福建省邵武縣將石省級自然保護區(qū)和泰寧縣丹霞山地區(qū),生長于丹霞山闊葉林下或山頂灌木叢中,調(diào)查區(qū)域種群數(shù)量大,居群性狀穩(wěn)定。該種均為灌木,調(diào)查的十幾個居群,尚未發(fā)現(xiàn)高度超過1.8 m,小葉常有粗鋸齒,花序不超過葉長的1/3,花瓣不具暗褐色脈紋,通過這些特征易與福建省廣布常見的木蠟樹區(qū)分開。而近年在福建省發(fā)現(xiàn)的少葉漆為喬木狀,葉背有白粉,花序與葉等長等特征與該新種相區(qū)分。與同為灌木狀的小漆樹不同之處在于葉長達25 cm,被微毛,葉背紫紅色,稀綠色;花序長不超過葉長1/3;花瓣不具暗褐色脈紋。更多照片保存于“自然標本館”(https://www.cfh.ac.cn/album/ShowSpAlbum.aspx?spid=96041),可供進一步對照比較。