日輪果屬，越南唇形科黃芩亞科一新屬

李波, Thanh Son Hoang, Maxim S. Nuraliev, Vu Anh Tai, Andrey N. Kuznetsov, Svetlana P. Kuznetsova, 向春雷

日輪果屬，越南唇形科黃芩亞科一新屬

李波1, Thanh Son Hoang2, Maxim S. Nuraliev3,4, Vu Anh Tai5, Andrey N. Kuznetsov3, Svetlana P. Kuznetsova3, 向春雷6*

(1. 中國科學(xué)院西雙版納熱帶植物園, 云南 勐臘 666303; 2., Hanoi 10000, Vietnam; 3., Hanoi 122000, Vietnam; 4., Moscow 119234, Russia; 5., Hanoi 10000, Vietnam; 6. 中國科學(xué)院昆明植物研究所, 昆明 650201)

描述并繪制了越南唇形科一新屬及新種：日輪果屬(Bo Li, C.L. Xiang, T.S. Hoang & Nuraliev)和日輪果(Bo Li, C.L. Xiang, T.S. Hoang & Nuraliev)。日輪果屬因具攀援藤本習(xí)性，花大且花冠為純白色，花萼近輻射對(duì)稱、5 深裂、裂片長(zhǎng)且在果期增大并開展，果實(shí)頂端具放射狀的瘤狀突起等特征而明顯區(qū)別于黃芩亞科的其他5屬。目前，該屬僅發(fā)現(xiàn)于越南東南部沿海的平定省、富安省、慶和省和寧順省，常生于海岸邊干旱的低地矮林中。

新分類群；特有屬；冬紅屬；膜萼藤屬；唇形目

自2016年起，筆者開始與越南同行合作開展越南唇形科植物分類學(xué)研究，先后發(fā)現(xiàn)新物種越南豆腐柴(Bo Li[1]、香簡(jiǎn)草(Prain)和保亭花(C.Y. Wu & S. Chow)在越南的分布新資料等[2–3]。在整理越南唇形科分類學(xué)文獻(xiàn)資料時(shí),[4]中所記載的第7393號(hào)物種引起了筆者的強(qiáng)烈關(guān)注。該種被鑒定為膜萼藤屬(Wall. ex Griff.)一未知種。通常而言，膜萼藤屬物種具有頂生或腋生的聚傘圓錐花序、花微小(長(zhǎng)不足5 mm)、花冠漏斗狀、4枚花冠裂片自喉部開展、花萼鐘狀、具4個(gè)小齒或淺裂、在果期增大呈透明的囊狀而完全包裹果實(shí)[5–6]。但從該物種的插圖和描述看，其單花腋生、花大型(長(zhǎng)4～6 cm)、花冠明顯二唇形且具5枚花冠裂片、花萼5深裂、裂片近輻射對(duì)稱且在果期增大并開展等特征與膜萼藤屬的形態(tài)差異巨大。

為進(jìn)一步明確該未知物種的真實(shí)身份，我們廣泛檢索了各類文獻(xiàn)資料和網(wǎng)絡(luò)在線資源，尤其是基于Pham-Hoang[4]所記載的原始分布地點(diǎn)“C?u?á, Nhatrang [筆者注：芽莊市，越南(Khanh Hoa Province)]”而將檢索區(qū)域重點(diǎn)聚焦在越南中南部沿海各省，最終在寧順省(Ninh Thuan Province)主山國家公園(Nui Chua National Park)官網(wǎng)上查閱到了兩張?jiān)撐锓N的野外照片(http://vqgnuichua.vn/ Upload/thuvienanh/DDS_14532_Hymenopyramis_04.JPG; http://vqgnuichua.vn/Upload/thuvienanh/DDS_ 14532_Hymenopyramis_10.JPG)，并在法國自然歷史博物館(Muséum National d’Histoire Naturelle)標(biāo)本館中檢索到2010年采自該國家公園的1份標(biāo)本(http://coldb.mnhn.fr/catalognumber/mnhn/p/p03977758)，二者均被鑒定為“sp.”。上述照片和標(biāo)本的發(fā)現(xiàn)為我們?cè)谝巴獠杉撐锓N提供了重要線索。

2021年，Maxim S. Nuraliev等在開展越南植物區(qū)系與植物多樣性研究時(shí)，在芽莊市珍珠島(Hon Tre Island)發(fā)現(xiàn)了該種的野生居群，并分別采集了標(biāo)本及分子材料。2022年， Thanh Son Hoang等相繼在平定省(Binh Dinh Province)和富安省(Phu Yen Province)海岸邊發(fā)現(xiàn)了該種大量野生居群。2022年7月至9月，當(dāng)?shù)谝蛔髡呃畈ú┦筷懤m(xù)收到采自慶和省和平定省的標(biāo)本實(shí)物時(shí)，確認(rèn)該未知種并非膜萼藤屬成員，并據(jù)其果實(shí)表面有顯著的放射狀瘤狀突起(果實(shí)表面具瘤狀或翅狀突起是黃芩亞科的共衍征之一[7])而推測(cè)該種與黃芩亞科存在明顯聯(lián)系。這一推測(cè)進(jìn)而得到了通訊作者向春雷博士實(shí)驗(yàn)室葉綠體系統(tǒng)發(fā)育基因組學(xué)分析結(jié)果的支持，確定其與冬紅屬(Retz.)構(gòu)成高支持率的姊妹群關(guān)系(未發(fā)表數(shù)據(jù))。綜合考慮形態(tài)學(xué)和分子系統(tǒng)學(xué)研究證據(jù)，確定該物種為唇形科黃芩亞科一未被描述的新屬，即日輪果屬(Bo Li, C.L. Xiang, T.S. Hoang & Nuraliev)，本文特此報(bào)道。

1 材料和方法

形態(tài)描述依據(jù)Maxim S. Nuraliev和Thanh Son Hoang采集的標(biāo)本及拍攝的野外照片，墨線圖的繪制參照號(hào)標(biāo)本，使用直尺和游標(biāo)卡尺進(jìn)行性狀測(cè)量，分布地點(diǎn)基于文獻(xiàn)和標(biāo)本采集信息及野外觀察記錄。憑證標(biāo)本分別存放在中國科學(xué)院華南植物園標(biāo)本館(Herbarium of SouthChina Botanical Garden, Chinese Academy of Sciences,IBSC)、中國科學(xué)院昆明植物研究所標(biāo)本館(Herbarium of Kunming Institute of Botany, Chinese Academy of Sciences, KUN)、中國科學(xué)院西雙版納熱帶植物園標(biāo)本館(Herbarium of Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, HITBC)、上海辰山植物園標(biāo)本館(Herbarium of Shanghai Chenshan Botanical Garden, CSH)、莫斯科國立米哈伊爾·瓦西里耶維奇·羅蒙諾索夫大學(xué)標(biāo)本館(Herbarium of M.V. Lomonosov Moscow State University, MW)和越南林業(yè)科學(xué)院標(biāo)本館(Herbarium of Vietnamese Academy of Forest Sciences, VAFS)。

2 結(jié)果

日輪果屬(黃芩亞科) 新屬

Bo Li, C.L. Xiang, T.S. Hoang & Nuraliev, gen. nov. (subfamily Scutellarioideae)

Type:Bo Li, C.L. Xiang, T.S. Hoang & Nuraliev

Diagnosis:is a distinct genus within the subfamily Scutellarioideae of Lamiaceae, which differs from the five other currently recognized genera of the subfamily by large flowers with pure white corolla, sub-actinomorphic calyx with five long lobes which are accrescent and spreading in fruits, strongly 2-lipped corolla with a relatively short tube, abruptly inflated corolla throat, median lobe of lower corolla lip much larger than its lateral lobes and than lobes of upper lip, obconical fruit splitting into 4 irregularly tetrahedral nutlets, and nutlets radially tuberculate at apex.is most closely related toaccording to the phylogenomic analysis of Lamiaceae (C.L. Xiang, unpublished data) but is readily distin- guished from the latter morphologically as indicated above.

Etymology: The generic nameis derived from the Greek(which means) and(which means), referring to the nutlets that are radially tuberculate at apex.

Note:is a monotypic genus currently known to be endemic to Vietnam.

日輪果屬因具攀援藤本習(xí)性；花冠大、顯著二唇形、冠筒為純白色且冠筒短而彎曲、在喉部急劇膨大成囊狀、花冠下唇中裂片顯著大于其他裂片;花萼近輻射對(duì)稱、5深裂、裂片長(zhǎng)且在果期增大并開展；果實(shí)為倒圓錐狀且完全分裂為4個(gè)小堅(jiān)果, 果實(shí)頂端具放射狀的瘤狀突起而明顯區(qū)別于黃芩亞科其他5屬。葉綠體系統(tǒng)發(fā)育基因組學(xué)分析結(jié)果顯示日輪果屬與冬紅屬為姊妹群(未發(fā)表數(shù)據(jù))，但二者存在上述一系列明顯形態(tài)特征差異而極易區(qū)分。

日輪果屬的名稱意指該屬果實(shí)頂端具放射狀的瘤狀突起，形似行星齒輪系傳動(dòng)裝置中的外齒輪形象，即太陽輪。

該屬僅包含日輪果一種，目前僅知特產(chǎn)于越南。

日輪果 新種圖1～3

Bo Li, C.L. Xiang, T.S. Hoang & Nuraliev, sp. nov. Figs. 1–3.

Type: VIETNAM. Khanh Hoa Province: Nha Trang City, Hon Tre Island, near Dam Bai marine climate station, growing in disturbed low coastal forests, 109°17′11″ E, 12°11′29″ N, elev. ca. 50 m a.s.l., 28 May 2021, Maxim S. Nuraliev,[holotype: IBSC (barcode No. 0896075!), isotypes: KUN: (barcode No. 1580435!), MW:(barcode No. MW0595841!)].

圖2 日輪果花果形態(tài)特征。A: 花, 頂面觀; B: 花, 側(cè)面觀; C: 花, 背面觀; D: 花冠管、雄蕊和花柱; E: 花冠, 側(cè)面觀; F: 花冠, 頂面觀; G: 花萼和果實(shí); H: 花萼, 側(cè)面觀; I: 成熟果實(shí), 頂面觀; J: 幼果, 側(cè)面觀; K: 子房橫切面; L: 小堅(jiān)果及合生面。(照片由Thanh Son Hoang和Maxim S. Nuraliev提供)

Description:Woody climbers or scandent shrubs, evergreen, 0.5–5 m long. Shoot erect or ascending when young but scandent when mature, often drooping gracefully to the ground, branching at nearly right angle. Branches distinctly 4-angled in cross section with obviously thickened to slightly winged ridges, green with brown ridges, glabrescent; branchlets slender, minutely pubescent when young, subglab- rescent. Mature nodes with light brown interpetiolar ridges covered with dense short hairs. Leaves opposite, simple; petiole 0.8–1.7 cm long, pubescent when young, glabrescent; leaf blade papery, entire on mature shoots, mostly shallowly 3-lobed on young shoots, ovate to indistinctly rhombic, 2.3–5.5 cm× 1.2–2.8 cm, hairy when young but completely glabre- scent on both sides, base broadly truncate, apex obtuse and shallowly emarginate (and sometimes with a small apical mucro), lateral vein 3–4 pairs, adaxially flat, abaxially slightly raised. Inflorescence a pendu- lous axillary cyme, usually 1-flowered, rarely 2- or 3- flowered; peduncles up to 0.5 cm long, sparsely pub- escent; pedicels 0.9–1.8 cm long, sparsely pubescent; bracteoles minute, linear, caducous. Calyx light green, campanulate, sub-actinomorphic, 1.3–1.9 cm long, papery, sparsely hairy outside (abaxially), densely minutely hairy inside (adaxially), covered with numerous yellow sessile glands, densely net-veined with 10 primary veins, 5-lobed to almost 1/2 of length; tube funnel-shaped, 4–6 mm long; lobes subequal, train- gular, acute at apex, 0.9–1.3 cm long, 0.6–0.8 cm wide at base; calyx persistent in fruits, 3.8–4.3 cm in diameter apically with lobes becoming spreading and accrescent to 1.9 cm long (fruiting calyx in shape of slightly zygomorphic star). Corolla pure white, zygo- morphic, strongly 2-lipped, tubular-infundibular, 3.3– 4.2 cm long; tube ca. 0.6 cm long, 4 mm wide at base, abruptly dilated to 1.2 cm wide at throat, sparsely minutely glandular pubescent outside; lips glabrous; upper (adaxial) lip 2-lobed, lobes triangular-ovate, equal, 0.5–0.8 cm×0.7–0.9 cm; lower (abaxial) lip 3- lobed, lobes unequal, median lobe straight and erect, subelliptic to oblong-obovate, 1.6–1.9 cm long, 1.6– 1.8 cm wide at the widest place, lateral lobes at right angle to median lobe (vertically oriented), nearly erect to recurved backwards, semicircular, 0.6 cm long and wide. Stamens 4, didynamous, appressed to adaxial side of corolla; filaments attached to corolla throat or slightly below, white, distinctly arcuate towards apex, glandular-pubescent near apex; posterior stamens exerted from the upper corolla lip, 1.7–1.9 cm long; anterior stamens included in corolla throat, ca. 1.2 cm long; anthers dorsifixed, yellow, with prominent con- nective, V-shaped (with divergent thecae), dehiscent longitudinally. Ovary superior, yellow, nearly cylin- drical, about 3 mm long and wide, glabrous, covered with scattered white sessile glands (more densely so towards base), longitudinally 4-lobed, each lobe 1- ovuled. Style subterminal (placed at center of ovary between its lobes), exerted from the upper corolla lip (reaching anthers of posterior stamens), white and apically slightly yellowish, 2.1–2.4 cm long, glabrous, curved downwards at apex; stigma unequally 2-cleft with stigma lobes ca. 2 mm long. Fruit deeply enclosed by calyx tube, green when young but brown when mature, obconical, glabrous, with radially tuberculate flat upper surface, completely splitting into 4 nutlets; nutlets dry, irregularly tetrahedral, ca. 6 mm long, outer side longitudinally ridged; attachment-scar lateral, ca. 2/3 as long as nutlet.

圖3 日輪果墨線圖。A: 花枝; B: 莖; C: 花冠, 側(cè)面觀; D: 花冠展開, 示雄蕊; E: 子房和花柱; F: 花萼; G: 果實(shí)頂面觀; H: 小堅(jiān)果合生面。(劉運(yùn)笑根據(jù)主模式NUR 3358繪制)

Phenology: Flowering and fruiting is recorded in January, May, June, and July. We suppose that the species is flowering and fruiting nearly all year round.

Etymology: The specific epithet “” refers to the ecological preferences of the new species, which is confined to the specific coastal vegetation.

Distribution and habitat: The new species is currently known fromDinh, Phu Yen, Khanh Hoa,and Ninh Thuan provinces of Vietnam (Fig. 4). It is confined to low and dry spiny coastal forests and shrubby vegetation, occurring up to 250 m a.s.l. Based on its occurrences and habitat, we suppose that the species is widely distributed in the coastal areas of south-central Vietnam.

Additional specimens examined (paratypes): VIETNAM. Binh Dinh Province: Phu My District, My Thanh Commune, Vinh Loi Village, semi-deciduous thorny scrubs on coastal sandy hill, 109°12′12′′ E, 14° 07′50′′ N, elev. ca. 10 m a.s.l., 5 July 2022, Vu Anh Tai & Thanh Son Hoang,(VAFS!); Khanh Hoa Province: Nha Trang City, Hon Tre Island, territory of Dam Bai Marine Climate Station, disturbed margin of forest, not far from seashore, 109°17′27′′ E, 12°11′52′′ N, elev. ca. 30 m a.s.l., 22 May 2021, Maxim S. Nuraliev,[CSH: (barcode No. CSH0198329!), MW: (barcode No. MW0595842!, MW0595843!), HITBC: (barcode No. HITBC0094275!)]; Ninh Thuan Province: Ninh Hai District, Nui Chua National Park, Botanical Garden, E slope, at Vung Olim Campsite, dry forests on the eastern flange of the Cotuy Range, 109°11′10′′ E, 11°42′48′′ N, elev. 100 m a.s.l., 8 January 2010, D. Djendoel Soejarto, N.T. Trung, T.N. Ninh & T.T. Trinh,[P: (barcode No. P03977758!)].

Other field records: Phu Yen Province: Tuy Hoa City, 3 June 2023, Thanh Son Hoang,(photo LE: LE01093262!); VIETNAM. Khanh Hoa Province: Nha Trang City, Hon Tre Island, near Dam Bai Marine Climate Station, in dry low forests on slope, not far from seashore, 109°17′20′′ E, 12°11′35′′ N, elev. ca. 40 m a.s.l., 7 January 2021, Maxim S. Nuraliev & Dmitry F. Lyskov,(photos LE: LE01093260!); bushland on plakor near dirt road, 109°16′52′′ E, 12° 11′38′′ N, elev. 220 m a.s.l., 28 May 2021, Maxim S. Nuraliev,(photos LE: LE01093261!).

圖4 日輪果在越南的地理分布(紅色五角星示)

Conservation status: Based on the wide distri- bution ofin coastal forests of south-central Vietnam discussed above and no signi- ficant threats observed for the documented popula- tions, we provisionally assess the conservation status of the species as “Least Concern” (LC) according to the IUCN Red List categories and criteria[8].

常綠攀緣狀灌木，高0.5～5 m。莖幼時(shí)直立或斜升，老時(shí)攀緣，常披散，懸垂地面，明顯四棱形且棱顯著增厚或略呈翅狀。老枝綠色但棱為棕色, 小枝近直角展開，柔毛漸脫落至光滑。小枝細(xì)弱, 被細(xì)柔毛至近無毛。節(jié)密被細(xì)柔毛，具橫向、淺棕色的葉柄縫隙聯(lián)合線。單葉對(duì)生，葉柄長(zhǎng)0.8～1.7 cm, 疏被柔毛。葉片紙質(zhì)，倒卵形至略微菱形，長(zhǎng)2.3～5.5 cm,寬1.2～2.8 cm，幼時(shí)3淺裂或全緣，疏被柔毛，老時(shí)漸脫落至完全無毛，葉基寬楔形，葉尖鈍或偶為漸尖，近全緣或偶微缺刻，側(cè)脈3～4對(duì)，在下面較為明顯。腋生聚傘花序常具單花或偶為3花；花序梗短，長(zhǎng)不過0.5 cm，被疏柔毛；花梗長(zhǎng)0.9～1.8 cm, 被疏柔毛；小苞片早落，微小，線形。花萼鐘狀, 淺綠色，近輻射對(duì)稱，長(zhǎng)1.3～1.9 cm，紙質(zhì)，外面疏被細(xì)柔毛，內(nèi)面更密，具10條主脈，網(wǎng)脈細(xì)密而明顯；萼筒短漏斗狀，長(zhǎng)0.4～0.6 cm；萼齒裂至花萼1/2處，裂片近相等，窄倒卵形至披針形，長(zhǎng)0.9～ 1.3 cm，基部寬0.6～0.8 cm；花萼果期宿存，萼片明顯增大，直徑3.8～4.3 cm，萼片開展，長(zhǎng)可達(dá)1.9 cm。花冠純白色，明顯二唇形，管狀漏斗形，長(zhǎng)3.3～ 4.2 cm；冠筒短而細(xì)，長(zhǎng)約0.6 cm，基部寬約0.4 cm，在喉部急劇擴(kuò)大呈囊狀，寬約1.2 cm，外面疏被細(xì)小腺毛；上唇2 裂，裂片等大，寬卵圓形，長(zhǎng)0.5～ 0.8 cm，寬0.7～0.9 cm；下唇3裂，中裂片最大, 近橢圓形至橢圓狀倒卵形，長(zhǎng)1.6～1.9 cm, 最寬處寬1.6～1.8 cm，側(cè)裂片與中裂片近直角，直立或后彎，半圓形。雄蕊4，二強(qiáng)雄蕊，花絲生于花冠喉部或略低位置，急劇地向上膝狀彎曲，花絲頂部被腺毛; 后對(duì)雄蕊自花冠上唇伸出，長(zhǎng)1.7～1.9 cm;前對(duì)雄蕊內(nèi)藏，長(zhǎng)約1.2 cm；花藥背著，黃色，藥室“V型”叉開，縱裂。子房上位，黃色，圓柱形, 長(zhǎng)約0.3 cm，疏被白色、無柄的盾狀腺毛，4室，每室具1胚珠。花柱近頂生，自花冠上唇伸出，與上對(duì)雄蕊近等長(zhǎng)，長(zhǎng)2.1～2.4 cm，無毛，頂部略帶黃色，向下彎曲; 柱頭為不等2 裂，裂片長(zhǎng)約2 mm。果實(shí)幾乎被萼管所包藏，倒圓錐狀，無毛，頂端截平，輪廓圓形, 具放射狀排列的瘤狀突起，成熟時(shí)完全分裂為4枚小堅(jiān)果；小堅(jiān)果幼時(shí)綠色，成熟時(shí)棕色，干燥，呈不規(guī)則四面體狀，長(zhǎng)約6 mm，外側(cè)面具縱棱脊；合生面約長(zhǎng)達(dá)小堅(jiān)果長(zhǎng)度的三分之二(圖1～3)。

基于標(biāo)本信息和我們的野外觀察數(shù)據(jù)，發(fā)現(xiàn)采自1月、5月、6月和7月的標(biāo)本都是盛花且有果的狀態(tài)，故推測(cè)日輪果有可能是全年開花結(jié)果。

越南：慶和省，芽莊市，珍珠島，海岸邊低地矮林，東經(jīng)109°17′11′′ E，北緯12°11′29′′ N，海拔約50 m，2021年5月28日，Maxim S. Nuraliev,(主模式：中國科學(xué)院華南植物園標(biāo)本館, IBSC, 0896075!；等模式：中國科學(xué)院昆明植物研究所標(biāo)本館，KUN, 1580435!；莫斯科國立米哈伊爾·瓦西里耶維奇·羅蒙諾索夫大學(xué)標(biāo)本館，MW, MW0595841!)。

種加詞“”意指其生長(zhǎng)于海岸邊低地矮林中的特殊生境。據(jù)目前所知，日輪果僅發(fā)現(xiàn)于越南東南沿海的平定省、富安省、慶和省和寧順省。盡管周邊省份類似生境中或許還有其他居群，但仍有待于進(jìn)一步確認(rèn)。就當(dāng)前所掌握的信息而言，日輪果的居群并未受到明顯可見的生存威脅。依據(jù)國際自然聯(lián)盟紅色名錄評(píng)估標(biāo)準(zhǔn)，我們暫時(shí)將其瀕危等級(jí)評(píng)估為“無危”[8]。

3 討論

黃芩亞科是唇形科12亞科之一[9–10]，此前包括5屬，即保亭花屬(C.Y. Wu & S. Chow)、冬紅屬、瘤果梓屬(Vatke)、黃芩屬(L.)和火梓屬(Kotschy ex Hook. f.)，其中前3屬為單型屬，黃芩屬包含約300種，為該亞科最大的屬，而火梓屬包含約20種[11]。日輪果屬的發(fā)現(xiàn)，使得該亞科的屬數(shù)增加至6屬。該亞科在形態(tài)上具有較高的多樣性[12]，但也存在兩個(gè)易于識(shí)別的共衍征，即花萼具發(fā)達(dá)的網(wǎng)脈和小堅(jiān)果具瘤狀或翅狀附屬物[7,10]，日輪果屬的相關(guān)形態(tài)特征與此高度吻合。黃芩亞科的花萼總體上呈現(xiàn)出增大與聯(lián)合的趨勢(shì)，其中保亭花屬和日輪果屬花萼具有5枚裂片，冬紅屬花萼擴(kuò)展為近輻射對(duì)稱的圓盤，僅偶見5枚小齒[6]，而黃芩屬、瘤果梓屬和火梓屬的花萼則上下二唇完全聯(lián)合為全緣[11]。相反，聚傘花序在該亞科有明顯的簡(jiǎn)化趨勢(shì)，即具3花的聚傘花序僅中央花發(fā)育而呈單花腋生狀態(tài)，或進(jìn)而由此構(gòu)成總狀花序或聚傘圓錐花序[7,9]。日輪果屬和黃芩屬及火梓屬部分種類的花序表現(xiàn)為前者，而保亭花屬、瘤果梓屬和冬紅屬則為后者。花冠管形態(tài)在該亞科也存在兩種狀態(tài)，一類是花冠管為管狀漏斗形，基部窄，中部彎曲，頂部擴(kuò)大為喉部，明顯長(zhǎng)于花萼, 保亭花屬、冬紅屬、黃芩屬和瘤果梓屬為此類形態(tài); 另一類是花冠管短而直，喉部劇烈擴(kuò)大，短于花萼, 日輪果屬和火梓屬花冠管則為此類型。地理分布上, 瘤果梓屬和火梓屬特產(chǎn)非洲，保亭花屬、日輪果屬和冬紅屬則特產(chǎn)東南亞熱帶亞熱帶地區(qū)，而黃芩屬近全球分布[10,12]。日輪果屬的發(fā)現(xiàn)為解析黃芩亞科的性狀演化和生物地理格局成因提供了全新的視角(未發(fā)表數(shù)據(jù))。

隨著該新屬的發(fā)表，黃芩亞科已增至6屬，為方便檢索，本文提供該亞科6屬的檢索表。

黃芩亞科各屬檢索表

Key to the genera of the subfamily Scutellarioideae

1. 花萼輻射對(duì)稱，圓盤狀，全緣或具不明顯的小齒，小堅(jiān)果略為肉質(zhì)；

Calyx actinomorphic, patelliform, with entire to shallowly dentate margin; nutlets somewhat fleshy ………冬紅屬

1. 花萼近輻射對(duì)稱至兩側(cè)對(duì)稱，絕非圓盤狀，小堅(jiān)果干燥；

Calyx sub-actinomorphic to zygomorphic, clearly lipped or lobed; nutlets dry …………………………………………………2

2. 花萼明顯5裂，小堅(jiān)果側(cè)面具縱棱；

Calyx clearly 5-lobed; nutlets longitudinally ridged ………………………………………………………………………………3

3. 花萼二唇形，上下唇全緣，小堅(jiān)果無縱棱；

Calyx 2-lipped, lips entire; nutlets not ridged ………………………………………………………………………………………4

4. 葉互生，頂生總狀花序，小堅(jiān)果基部具小柄，頂部具不規(guī)則瘤狀突起；

Leaves alternate; inflorescence a many-flowered terminal raceme; nutlets attached to receptacle by slender stalks, apically irregularly tuberculate ………………………………………………………………………………………保亭花屬

3. 葉對(duì)生，腋生簡(jiǎn)化的聚傘花序，小堅(jiān)果基部無柄，頂部具放射狀瘤狀突起；

Leaves opposite; inflorescence a 1–3-flowered axillary cyme; nutlets not stalked, apically radially tuberculate ……………………………………………………………………………………………………………………………………日輪果屬

4. 花萼上唇常具盾片或囊狀突起；

Posterior lip of calyx usually folded to produce a scutellum or conspicuously saccate…………………………黃芩屬

5. 花萼無盾狀或囊狀突起；

Calyx without scutellum and not saccate …………………………………………………………………………………………5

6. 花萼果期膨大；小堅(jiān)果具翅狀附屬物；

Calyx inflated in fruits; nutlets not tuberculate but usually with a wing of stiff ray-like projections ………………火梓屬

5. 花萼果期不膨大；小堅(jiān)果背側(cè)面具瘤狀突起，無翅；

Calyx not inflated; nutlets not winged, dorsal surface tuberculate ……………………………………………瘤果梓屬

致謝 感謝中國科學(xué)院華南植物園標(biāo)本館劉運(yùn)笑女士繪制墨線圖、上海辰山植物園劉夙博士提供命名法參考意見、中國科學(xué)院昆明植物研究所陳亞萍博士分享系統(tǒng)發(fā)育分析結(jié)果及審稿專家和責(zé)任編輯提供寶貴的修改意見。

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gen. nov. (Scutellarioideae, Lamiaceae) from Coastal Southern Vietnam

LI Bo1, Thanh Son HOANG2, Maxim S. NURALIEV3,4, Vu Anh TAI5, Andrey N. KUZNETSOV3, Svetlana P. KUZNETSOVA3, XIANG Chunlei6*

(1. Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla 666303, Yunnan, China; 2. Silviculture Research Institute, Vietnamese Academy of Forest Sciences,Hanoi 10000, Vietnam; 3. Joint Russian-Vietnamese Tropical Scientific and Technological Center, Hanoi 122000, Vietnam; 4. Department of Higher Plants, Biological Faculty, M.V. Lomonosov Moscow State University,Moscow 119234, Russia; 5. Institute of Geography, Vietnam Academy of Science and Technology,Hanoi 10000, Vietnam; 6. Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences,Kunming 650201, China)

Bo Li, C.L. Xiang, T.S. Hoang & Nuraliev andBo Li, C.L. Xiang, T.S. Hoang & Nuraliev, a new genus and a new species of Scutellarioideae (Lamiaceae) endemic to coastal areas of south-central Vietnam, are described and illustrated.is distinct from the other five genera of Scutellarioideae by having climbing habit, large flowers with pure white corolla, sub-actinomorphic calyx with five long lobes which are accrescent and spreading in fruits, and nutlets radially tuberculate at apex. The genus is currently only known from four provinces of VietnamDinh, Phu Yen, Khanh Hoa,and Ninh Thuan).

New taxa; Endemic genus;;; Lamiales

10.11926/jtsb.4827

2023-02-20

2023-07-27

國家自然科學(xué)基金項(xiàng)目(32160047)；The Scientific Project of the State Order of the Government of Russian Federation to Lomonosov Moscow State University (121032500082-2)；云南省基礎(chǔ)研究計(jì)劃重點(diǎn)項(xiàng)目(202001AS070016)資助

This work was supported by the National Natural Science Foundation of China (Grant No. 32160047), the Scientific Project of the State Order of the Government of Russian Federation to Lomonosov Moscow State University (Grant No. 121032500082-2), and the Project for Key Basic Research Planning in Yunnan (Grant No. 202001AS070016).

李波(1984年生)，男，博士，研究方向?yàn)榇叫慰葡到y(tǒng)發(fā)育與分類學(xué)修訂。E-mail: hanbolijx@163.com

. E-mail: xiangchunlei@mail.kib.ac.cn