鞘蕊属
鞘蕊属(学名:Coleus),或作鞘蕊花属,是一个一年生或多年生的草本或灌木植物的属。部分物种属于多肉植物,长有具肉质或块状的块茎或块根。 分布于旧世界(东半球)的热带与亚热带。 有关鞘蕊属、彩叶草属及马刺花属三者的关系很混乱:前两属的物种曾经被并归马刺花属,但结果使马刺花属在支序亲缘分析之下变成了马刺花亚族(Plectranthinae)的一个多系群。
鞘蕊花属 | |
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左手香(Coleus amboinicus;syn. Plectranthus amboinicus) | |
科学分类 | |
界: | 植物界 Plantae |
演化支: | 维管植物 Tracheophyta |
演化支: | 被子植物 Angiosperms |
演化支: | 真双子叶植物 Eudicots |
演化支: | 菊类植物 Asterids |
目: | 唇形目 Lamiales |
科: | 唇形科 Lamiaceae |
亚科: | 荆芥亚科 Nepetoideae |
族: | 罗勒族 Ocimeae |
属: | 鞘蕊花属 Coleus Lour.[1] |
异名[1] | |
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现时最新的分类学处理重新建立鞘蕊属,212个物种分别来自马刺花属、密穗花属及排香草属;Equilabium在系统发育研究支持下从马刺花属独立出去成为一个属[2]。
鞘蕊属物种常被用作园艺植物而种植,当中尤以彩叶草最为常见。一方面,彩叶草的斑斓叶片使之成为一种受欢迎的园艺植物,而近年兴起的生酮饮食,亦因为彩叶草的根部丰富的营养素而使之大受欢迎。彩叶草喜光,但不适宜太阳直射,而是非直射的光线:因为直射的太阳光会破坏叶片上的色素,使叶片颜色变得单一;所以在户外时,彩叶草应完全遮光,只能照射日出时最柔和的日光[3]。
其他长有块根的鞘蕊属植物有被当作食粮而种植,计有Coleus esculentus、Coleus rotundifolius及Coleus maculosus subsp. edulis。
分类学
鞘蕊属最早由若昂·德理路于1790年描述。以形态学特征为指导基础,将它由马刺花属(首先由夏尔·莱里捷在1788年描述)独立出来;其四条雄蕊融合在一起,而不是从基底起就已分开。1962年,约翰·肯尼斯·莫顿留意到雄蕊融合在一起的物种比过往所认知的其实更常见及广泛分布,所以把鞘蕊属物种合并进马刺花属,而彩叶草属(Solenostemon)及一些其他的属仍维持区隔。[4]大多数的来源都跟随莫顿进行鞘蕊属的合并;有些保持彩叶草属的独立,但其他则将它跟着鞘蕊属一起并入马刺花属。[2]
在2004年针对唇形科荆芥亚科罗勒族(Ocimeae)进行的初步研究表明,马刺花亚族(Plectranthinae)为单系群,有两个主要演化支:一个包含鞘蕊属的模式种,并包含彩叶草属;另一个包含马刺花属的模式种以及其他属,因此广义上马刺花属不是单系群。在2018年进行的更详细研究得出类似的结论,并建议应重新确认鞘蕊属(包含彩叶草属),对马刺花属进行更狭义的定义,并建立一个新的属Equilabium,主要来自热带非洲的前马刺花属物种演化支。2019年提供了实现此方法所需的许多新的双名组合。[2]
支序亲缘
裴顿等人(2019)[2]基于其2018年的研究总结[5],出版了马刺花亚族(Plectranthinae)各属的支序亲缘图。下图中被裴顿等人接纳并构成广义的马刺花属的三个属以黄色底显示:
罗勒族 Ocimeae |
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物种
Paton et al. (2019)列出本属物种有294种,当中大多数均由马刺花属转移过来[2]。截至2020年7月[update],《世界植物在线》列出本属25个物种[1]。部分物详列如下[2]:
- Calchas parvifolius (Benth.) P.V.Heath, 1999
- Calchas platystomoides (Robyns & Lebrun) P.V.Heath, 1999
- Coleus abyssinicus (Fresen.) A.J.Paton
- Coleus acariformis (P.I.Forst.) P.I.Forst.
- Coleus achankoviliensis Smitha & A.J.Paton
- Coleus actites (P.I.Forst.) P.I.Forst.
- Coleus adenanthus (Dalzell & A.Gibson) A.J.Paton
- Coleus adenophorus (Gürke) A.J.Paton
- Coleus aegyptiacus (Forssk.) A.J.Paton
- Coleus affinis (Gürke) A.J.Paton
- Coleus albicalyx (Suddee) Suddee
- Coleus aliciae (Codd) A.J.Paton
- Coleus alloplectus (S.T.Blake) P.I.Forst. & T.C.Wilson
- Coleus alpinus Vatke
- Coleus altanmouiensis (T.C.Wilson, P.I.Forst. & M.A.M.Renner) T.C.Wilson & P.I.Forst.
- 到手香 Coleus amboinicus Lour.,异名:Plectranthus amboinicus)又名:左手香、过手香
- Coleus amicorum (S.T.Blake) P.I.Forst. & T.C.Wilson
- Coleus amiculatus (T.C.Wilson, P.I.Forst. & M.A.M.Renner) T.C.Wilson & P.I.Forst.
- Coleus amoenus (P.I.Forst.) P.I.Forst.
- Coleus anamallayensis Bedd.
- Coleus anamudianus (Smitha & Sunojk.) Smitha
- Coleus angolensis (G.Taylor) A.J.Paton
- Coleus angulatus (Hedge) A.J.Paton & Phillipson
- Coleus apoensis Elmer
- Coleus apreptus (S.T.Blake) P.I.Forst. & T.C.Wilson
- Coleus apricus (P.I.Forst.) P.I.Forst.
- Coleus arabicus Benth.
- Coleus arenicola (P.I.Forst.) P.I.Forst.
- 银叶到手香 Coleus argentatus (S.T.Blake) P.I.Forst. & T.C.Wilson;又名:银叶马刺花、银叶香茶菜,异名:Plectranthus argentatus
- Coleus argenteus (Gamble) A.J.Paton
- Coleus argentifolius (Ryding) A.J.Paton
- Coleus aromatium Benth.
- Coleus articulatus (I.M.Johnst.) A.J.Paton
- Coleus ater A.J.Paton
- Coleus auriglandulosus (A.J.Paton) A.J.Paton
- Coleus australis (R.Br.) A.J.Paton
- Coleus autranii Briq.
- Coleus barbatus (Andrews) Benth. ex G.Don,异名:Plectranthus barbatus
- Coleus bariensis (Ryding) A.J.Paton
- Coleus batesii (Baker) A.J.Paton
- Coleus batianoffii (P.I.Forst.) P.I.Forst.
- Coleus bellus (P.I.Forst.) P.I.Forst.
- Coleus betonicifolius (Baker) A.J.Paton
- Coleus bifidus (A.J.Paton) A.J.Paton
- Coleus bipartitus (P.I.Forst.) P.I.Forst.
- Coleus bishopianus (Gamble) Smitha & A.J.Paton
- Coleus blakei (P.I.Forst.) P.I.Forst.
- Coleus bojeri Benth.
- Coleus bolavenensis Suddee, Tagane & Rueangr.[6]
- Coleus botryosus (A.J.Paton) A.J.Paton
- Coleus bourneae (Gamble) Smitha & A.J.Paton
- 光萼鞘蕊花 Coleus bracteatus Dunn
- Coleus brazzavillensis A.Chev.
- Coleus buchananii (Baker) Brenan
- Coleus burorum Chiov.
- Coleus caillei A.Chev.
- Coleus caillei A.Chev. ex Hutch. & Dalziel
- Coleus calaminthoides (Baker) A.J.Paton
- Coleus calcicola Murata
- Coleus caldericola (P.I.Forst.) P.I.Forst.
- Coleus calycinus (Benth.) A.J.Paton
- Coleus cambodianus (Murata) A.J.Paton
- Coleus caninus (Roth) Vatke,异名:Plectranthus caninus
- Coleus caninus Paton, Mwanyambo, Govaerts, Smitha, Suddee, Phillipson, Wilson, Forster & Culham, 2019
- 肉叶鞘蕊花 Coleus carnosifolius (Hemsl.) Dunn
- Coleus casamancicus A.Chev.
- Coleus cataractarum (B.J.Pollard) A.J.Paton
- Coleus caudatus (S.Moore) E.Downes & I.Darbysh.
- Coleus caudatus S.Moore
- Coleus celsus A.J.Paton
- Coleus centraliafricanus A.J.Paton
- Coleus chevalieri Briq.
- Coleus ciliatus (Bramley) A.J.Paton
- Coleus circinnatus (Hedge) A.J.Paton & Phillipson
- Coleus clandulosus Hook.f.
- Coleus coeruleus Gürke
- Coleus comosus Hochst. ex A.Rich., 1850,异名:Plectranthus ornatus (sold as "Coleus canina" or "scaredy cat plant")
- Coleus comosus Hochst. ex Gürke
- Coleus confertiflorus (A.J.Paton) A.J.Paton
- Coleus congensis (Gürke) A.J.Paton
- Coleus congestus (R.Br.) A.J.Paton
- Coleus conglomeratus (T.C.E.Fr.) Robyns & Lebrun
- Coleus crassus (N.E.Br.) Culham
- Coleus cremnus (B.J.Conn) A.J.Paton
- Coleus cucullatus (A.J.Paton) A.J.Paton
- Coleus cuneatus Baker f.
- Coleus cyanophyllus (P.I.Forst.) P.I.Forst.
- Coleus cylindraceus (Hochst. ex Benth.) A.J.Paton
- Coleus daviesii E.A.Bruce
- Coleus decimus (A.J.Paton) A.J.Paton
- Coleus decurrens Gürke
- Coleus deflexifolius (Baker) A.J.Paton
- Coleus defoliatus (Hochst. ex Benth.) A.J.Paton
- Coleus densus (N.E.Br.) A.J.Paton
- Coleus descampsii (Briq.) A.J.Paton
- Coleus dewildemanianus (Robyns & Lebrun) A.J.Paton
- Coleus dichotomus (A.J.Paton) A.J.Paton
- Coleus dinteri (Briq.) A.J.Paton
- Coleus divaricatus A.J.Paton
- Coleus diversus (S.T.Blake) P.I.Forst. & T.C.Wilson
- Coleus dolichopodus (Briq.) A.J.Paton
- Coleus dumicola (P.I.Forst.) P.I.Forst.
- Coleus dysophylloides (Benth.) A.J.Paton
- Coleus elliotii (S.Moore) A.J.Paton
- Coleus eminii (Gürke) A.J.Paton
- Coleus engleri (Briq.) A.J.Paton
- Coleus erici-rosenii (R.E.Fr.) A.J.Paton
- 食用鞘蕊花 Coleus esculentus ,异名:Plectranthus esculentus)
- Coleus eungellaensis A.J.Paton
- Coleus excelsus (P.I.Forst.) P.I.Forst.
- Coleus exilis (A.J.Paton) A.J.Paton
- Coleus fasciculatus (P.I.Forst.) P.I.Forst.
- Coleus ferricola Phillipson, O.Hooper & A.J.Paton
- Coleus foetidus (Benth.) A.J.Paton
- Coleus foliatus (A.J.Paton) A.J.Paton
- 兰屿小鞘蕊花 Coleus formosanus
- Coleus forsteri (Benth.) A.J.Paton
- Coleus fragrantissimus (P.I.Forst.) P.I.Forst.
- Coleus fredericii ,异名:Plectranthus welwitschii
- Coleus gamblei (Smitha & Sunojk.) Smitha
- Coleus geminatus (P.I.Forst.) P.I.Forst.
- Coleus gibbosus A.J.Paton
- Coleus gigantifolius (Suddee) Suddee
- Coleus gillettii (J.K.Morton) A.J.Paton
- Coleus glabriflorus (P.I.Forst.) P.I.Forst.
- Coleus globosus (Ryding) A.J.Paton
- Coleus goetzenii (Gürke) A.J.Paton
- Coleus gossweileri A.J.Paton
- Coleus gracilipedicellatum (Robyns & Lebrun) A.J.Paton
- Coleus graminifolius (Perkins) A.J.Paton
- Coleus grandidentatus (Gürke) A.J.Paton
- Coleus graniticola (A.Chev.) A.J.Paton
- Coleus gratus (S.T.Blake) P.I.Forst. & T.C.Wilson
- Coleus graveolens (R.Br.) A.J.Paton
- Coleus guerkei (Briq.) A.J.Paton
- Coleus habrophyllus (P.I.Forst.) P.I.Forst.
- Coleus hadiensis (Forssk.) A.J.Paton
- Coleus hairulii Kiew, 2021
- Coleus hallii (J.K.Morton) A.J.Paton
- Coleus harmandii (Doan ex Suddee & A.J.Paton) A.J.Paton
- Coleus helferi (Hook.f.) A.J.Paton
- Coleus hereroensis (Engl.) A.J.Paton
- Coleus hijazensis (Abdel Khalik) A.J.Paton
- Coleus huberi Regel
- Coleus humulopsis (B.J.Pollard) A.J.Paton
- Coleus hymalis (J.R.I.Wood) A.J.Paton
- Coleus idukkianus (J.Mathew, Yohannan & B.J.Conn) Smitha
- Coleus igniarioides (Ryding) A.J.Paton
- Coleus ignotus (A.J.Paton) A.J.Paton
- Coleus inselbergii (B.J.Pollard & A.J.Paton) A.J.Paton
- Coleus insignis (Hook.f.) A.J.Paton
- Coleus insularis (P.I.Forst.) P.I.Forst.
- Coleus intraterraneus (S.T.Blake) P.I.Forst. & T.C.Wilson
- Coleus kanyakumariensis (Shinoj & Sunojk.) Smitha
- Coleus kirkii (Baker) A.J.Paton
- Coleus kisanfuensis DeWild.
- Coleus klossii (S.Moore) P.I.Forst. & T.C.Wilson
- Coleus kottalensis A.Chev.
- Coleus koulikoroensis A.J.Paton
- Coleus kunstleri (Prain) A.J.Paton
- Coleus laetus (P.I.Forst.) P.I.Forst.
- Coleus lageniocalyx Briq.
- Coleus lanceolatus (Bojer ex Benth.) A.J.Paton & Phillipson
- Coleus lancifolius (Bramley) A.J.Paton
- Coleus lanuginosus Hochst. ex Engl., 1892
- Coleus lateriticola (A.Chev.) Phillipson, O.Hooper & A.J.Paton
- Coleus lateriticola A.Chev.
- Coleus latifolius Hochst. ex Engl., 1892
- Coleus laxus (T.C.Wilson & P.I.Forst.) T.C.Wilson & P.I.Forst.
- Coleus leiperi (P.I.Forst.) P.I.Forst.
- Coleus linearifolius (J.K.Morton) A.J.Paton
- Coleus livingstonei A.J.Paton
- Coleus longipetiolatus Gürke
- Coleus longipetiolatus Gürke ex Engl., 1894
- Coleus luengerensis Gürke
- Coleus lyratus (A.Chev.) Roberty
- Coleus macrostachys Benth.
- Coleus maculosus (Lam.) A.J.Paton
- 马达加斯加鞘蕊花 Coleus madagascariensis ,异名:Plectranthus madagascariensis)又名:马达加斯加延命草、马达加斯加香茶菜
- Coleus magnificus P.I.Forst. & A.J.Paton
- Coleus marrubatus (J.K.Morton) A.J.Paton
- Coleus megacalyx (A.J.Paton) A.J.Paton
- Coleus megadontus (P.I.Forst.) P.I.Forst.
- Coleus melleri (Baker) A.J.Paton & Phillipson
- Coleus meyeri (Gürke) A.J.Paton
- Coleus minor (J.K.Morton) A.J.Paton
- Coleus minutiflorus (Ryding) A.J.Paton
- Coleus minutus (P.I.Forst.) P.I.Forst.
- Coleus mirus (S.T.Blake) P.I.Forst. & T.C.Wilson
- Coleus mitis (R.A.Clement) A.J.Paton
- Coleus monostachyus (P.Beauv.) A.J.Paton
- Coleus montanus Hochst. ex Ces.
- Coleus mutabilis (Codd) A.J.Paton
- Coleus myrianthellus Briq.
- Coleus namuliensis E.Downes & I.Darbysh.
- Coleus neochilus (Schltr.) Codd
- Coleus nepetifolius (Baker) A.J.Paton
- Coleus nervosus Briq.
- Coleus niamniamensis (Gürke) A.J.Paton
- Coleus nigericus (Alston) A.J.Paton
- Coleus nitidus (P.I.Forst.) P.I.Forst.
- Coleus niveus (Hiern) A.J.Paton
- Coleus omahekense Dinter
- Coleus omissus (P.I.Forst.) P.I.Forst.
- Coleus orbicularis Baker
- Coleus orthodontus (Gürke) A.J.Paton
- Coleus otostegioides (Schweinf. ex Gürke) A.J.Paton
- Coleus pallidus (Wall. ex Benth.) A.J.Paton
- Coleus parensis Gürke
- Coleus parishii (Hook.f.) A.J.Paton
- Coleus parvicalyx (A.J.Paton) A.J.Paton
- Coleus parvifolius Benth.
- Coleus penicillatus (A.J.Paton) A.J.Paton
- Coleus penzigii Schweinf. ex Penz.
- Coleus perrieri (Hedge) A.J.Paton & Phillipson
- Coleus petiolatissimus Briq.
- Coleus petraeus (Backer ex Adelb.) A.J.Paton
- Coleus petricola (J.Mathew & B.J.Conn) A.J.Paton
- Coleus petrophilus Gürke
- Coleus phulangkaensis (Suddee, Suphuntee & Saengrit) Suddee
- Coleus plantagineus (Hook.f.) A.J.Paton
- Coleus platyphyllus (A.J.Paton) A.J.Paton
- Coleus polystachyus (Benth.) A.J.Paton
- Coleus porcatus (van Jaarsv. & P.J.D.Winter) A.J.Paton
- Coleus porphyranthus (T.J.Edwards & N.R.Crouch) A.J.Paton
- Coleus prittwitzii (Perkins) A.J.Paton
- Coleus prostratus (Gürke) A.J.Paton
- Coleus psammophilus (Codd) A.J.Paton
- Coleus pseudospeciosus (Buscal. & Muschl.) A.J.Paton
- Coleus pulchellus (P.I.Forst.) P.I.Forst.
- Coleus pumilius Blanco, 1837
- Coleus rafidahiae Kiew, 2021
- Coleus recurvata (Ryding) A.J.Paton
- Coleus rhodesianum (N.E.Br.) A.J.Paton
- Coleus robustus (Hook.f.) A.J.Paton
- 圆叶鞘蕊花 Coleus rotundifolius (Poir.) M.R.Almeida, 2003,异名:Plectranthus rotundifolius
- Coleus ruandensis (De Wild.) A.J.Paton
- Coleus rutenbergianus (Vatke) A.J.Paton & Phillipson
- Coleus sahyadricus (Smitha & Sunojk.) Smitha
- Coleus sallyae (A.J.Paton) A.J.Paton
- Coleus sanguineus (Britten) A.J.Paton
- Coleus saxicola Gürke
- Coleus saxorum (J.Mathew, Yohannan & B.J.Conn) Smitha
- Coleus scaber (Benth.) A.J.Paton
- Coleus schizophyllus (Baker) A.J.Paton
- Coleus schliebenii (Mildbr.) A.J.Paton
- Coleus scruposus A.J.Paton
- 彩叶草 Coleus scutellarioides (L.) Benth.,异名Coleus blumei, Plectranthus scutellarioides
- Coleus scutellarioides Elmer
- Coleus sessilifolius (A.J.Paton) A.J.Paton
- Coleus shoolamudianus (Sunil & Naveen Kum.) Smitha & A.J.Paton
- Coleus sigmoideus (A.J.Paton) A.J.Paton
- Coleus socotranus (Radcl.-Sm.) A.J.Paton,异名:Plectranthus socotranus
- Coleus sphaerocephalus (Baker) A.J.Paton
- Coleus splendens (P.I.Forst.) P.I.Forst.
- Coleus splendidus A.Chev.
- Coleus steenisii (H.Keng) A.J.Paton
- Coleus stenostachys (Baker) A.J.Paton & Phillipson
- Coleus strictipes (G.Taylor) A.J.Paton
- Coleus strobilifer (Roxb.) A.J.Paton
- Coleus stuhlmannii (Gürke) A.J.Paton
- Coleus suaveolens (S.T.Blake) P.I.Forst. & T.C.Wilson
- Coleus subscandens Gürke
- Coleus suffruticosus (Wight) A.J.Paton
- Coleus sylvestris (Gürke) A.J.Paton & Phillipson
- Coleus tetradenifolius (A.J.Paton) A.J.Paton
- Coleus thalassoscopicus (P.I.Forst.) P.I.Forst.
- Coleus togoensis (Perkins) A.J.Paton
- Coleus tomentifolius (Suddee) Suddee
- Coleus torrenticola (P.I.Forst.) P.I.Forst.
- Coleus triangularis (A.J.Paton) A.J.Paton
- Coleus trullatus (A.J.Paton) A.J.Paton
- Coleus ulugurensis Gürke
- Coleus unguentarius (Codd) A.J.Paton,异名:Plectranthus unguentarius
- Coleus velutinus (Trimen) A.J.Paton
- Coleus venteri (van Jaarsv. & Hankey) A.J.Paton
- Coleus ventosus (P.I.Forst.) P.I.Forst.
- Coleus venustus (P.I.Forst.) P.I.Forst.
- Coleus verticillatus (Baker) A.J.Paton
- Coleus veyretiae (Guillaumet & A.Cornet) A.J.Paton & Phillipson
- Coleus villosus (Forssk.) A.J.Paton
- Coleus wallamanensis (T.C.Wilson & P.I.Forst.) T.C.Wilson & P.I.Forst.
- Coleus wugensis Gürke
- Coleus xerophilus (Codd) A.J.Paton
- Coleus xylopodus (Lukhoba & A.J.Paton) A.J.Paton
- Coleus yemenensis A.J.Paton
- Coleus zombensis (Baker) Mwany.
- 杂交种:
- 归入本属的物种:
- Rabdosiella leemannii N.Hahn
- Saccostoma urticifolium Wall. ex Voigt
- Solenostemon pubescens A.Chev.
图库
参考文献
- ^ 1.0 1.1 1.2 Coleus Lour.. Plants of the World Online. Royal Botanic Gardens, Kew. [2020-07-28]. (原始内容存档于2020-10-27).
- ^ 2.0 2.1 2.2 2.3 2.4 2.5 Paton, Alan J.; Mwanyambo, Montfort; Govaerts, Rafaël H.A.; Smitha, Kokkaraniyil; Suddee, Somran; Phillipson, Peter B.; Wilson, Trevor C.; Forster, Paul I. & Culham, Alastair. Nomenclatural changes in Coleus and Plectranthus (Lamiaceae): a tale of more than two genera. PhytoKeys. 2019, 129: 1–158. PMC 6717120 . PMID 31523157. doi:10.3897/phytokeys.129.34988.
- ^ Coleus Care. Gardening Know How. 2019-07-15 [2019-07-15]. (原始内容存档于2019-07-15).
- ^ Morton, J.K. Plectranthus. Hedberg, I.; Kelbessa, E.; Edwards, S.; Demissew, S. & Persson, E. (编). Flora of Ethiopia and Eritrea 5. Addis Ababa: The National Herbarium, Addis Ababa University: 586–598. 2006. ISBN 978-91-971285-6-8., cited in Paton et al. 2019
- ^ Paton, A.; Mwanyambo, M. & Culham, A. Phylogenetic study of Plectranthus, Coleus and allies (Lamiaceae): Taxonomy, distribution and medicinal use (PDF). Botanical Journal of the Linnean Society. 2018, 188 (4): 355–376 [2020-12-22]. doi:10.1093/botlinnean/boy064. (原始内容存档 (PDF)于2020-11-30).
- ^ Somran Suddee, Shuichiro Tagane, Phetlasy Souladeth, Deuantar Kongxaisavath, Sukid Rueangruea, Yoshihisa Suyama and Eizi Suzuki. 2020. Coleus bolavenensis (Lamiaceae), A New Species from Laos. Thai Forest Bulletin (Botany). 48(1), 82-85. DOI: 10.20531/tfb.2020.48.1.14
外部链接
- EcoPort on P. rotundifolius (synonym of Coleus rotundifolius)
- PROTAbase on P. rotundifolius (页面存档备份,存于互联网档案馆) (synonym of Coleus rotundifolius)
- EcoPort on P. esculentus (with tuber photo) (synonym of Coleus esculentus)
- Lamiales Newsletter on P. esculentus (页面存档备份,存于互联网档案馆) (synonym of Coleus esculentus)
- Little-known African Tubers (页面存档备份,存于互联网档案馆) (discusses C. rotundifolius and C. esculentus)