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石竹及康乃馨
石竹
• 石竹科(Caryophyllaceae)
• 石竹屬(Dianthus)
• 英名:sweet william, pink
• 中名:五彩石竹、美國石竹、
三寸石竹、五寸石竹、美女撫子、
深山櫻
• 和名:なでしこ(nadeshiko)
石竹屬~概況與分布~
• 本屬估計約有200~300個物種皆分布於舊世界(歐亞大陸)
• 以南歐-西亞為種原中心,尤地中海沿岸
(Valente et al., 2010)
花壇石竹的歷史
• 最早在地中海希臘羅馬文化用於祭祀、香氣等
– 希臘哲學家泰奧弗拉斯托斯(Theophrastus)命名
Dianthus,由dios (主神)及anthos (花)組成
– 所描述之物種可能為D. plumarius或D. caryophyllus
– 種名caryophyllus為古希臘文意指丁香樹(Caryophyllus
aromaticus)之乾燥花苞,源其花朵特有之似丁香味
• 當時並非以矮性花壇為主,亦有切花用途
• 石竹在早期中歐地區並非重要花卉(非聖經花卉)
• 直到中世紀晚期(1450左右),才逐漸帶有神聖、
愛情等意義,漸為人所重視
The Unicorn in Captivity
(from the Unicorn Tapestries)
Date: 1495–1505
Culture: South Netherlandish
The Hunters Enter the Woods
(from the Unicorn Tapestries)
Date: 1495–1505
Culture: South Netherlandish
花壇石竹的歷史
• 近代使用D. barbatus為主要育種材料,主要考量
– 枝條頂端聚生多花序且單一花序花朵數多
– 適應性強
• ‘幸好成功了’ ─ 導入D. chinensis血統的故事
– 1960年代後半
– 美國GoldSmith公司育種家Charles James
– 種間雜交種(‘Queen of Hearts’)的優勢
• 不易結種子(花期延長)
• 大花性
• 耐寒
• 生長勢
古文中的石竹
• 司空曙 <雲陽寺石竹花>
一自幽山別,相逢此寺中。
高低俱出葉,深淺不分叢。
野蝶難爭白,庭榴暗讓紅。
誰憐芳最久,春露到秋風。
• 白居易 <牡丹芳 美天子憂農也>
…石竹金錢何細碎,芙蓉芍藥苦尋常。…
瞿麥
[釋名]蘧麥、大菊(出自爾雅)、巨句麥(出自神農本草經)、大蘭(出自名醫
別錄)、石竹(出自日華子本草)、南天竺草。
[敘述]{宏景曰}子頗似麥,故名瞿麥。{時珍曰}按陸佃解韓詩外傳云,生
于兩旁謂之瞿,此麥之穗旁生故也。爾雅作蘧,有渠衢二音。日華本草
云,一名燕麥,一名杜姥草者,誤矣;燕麥即雀麥,雀瞿二字相近,傳
寫之訛爾。
[集解]{別錄曰}瞿麥生太山山谷,立秋採,陰乾。{宏景曰}今出近道,一
莖生細葉,花紅紫赤色可愛,合子葉刈取之。子頗似麥子。有兩種,一
種為大,花邊有叉椏,未知何者是也。今市人皆用小者,復一種葉莖相
似,而有毛,花晚而甚赤。按經云採實,其中子細,燥熱便脫盡矣。{頌
曰}今處處有之,苗高一尺以來,葉尖小、青色,根紫黑色,形如細蔓,
花紅紫赤色,亦似映山紅。二月至五月開,七月結實作穗。子頗似麥。
河陽河中府出者,苗可用;淮甸出者,根細,村民取作刷帚。爾雅謂之
大菊,廣雅謂之茈萎,是也。{時珍曰}石竹葉似地膚葉而尖小,又似出
生小竹葉而細窄。其莖纖細有節,高尺餘,稍閒開花。田野生者花大如
錢,紅紫色,人家栽者花稍小而嫵媚,有細白粉紅紫赤斑斕數色。俗乎
為洛陽花。結石如燕麥,內有小黑子。其嫩苗煠熟水滔過可食。
本草綱目中的訊息
• {宏景曰}子頗似麥,故名瞿麥。{時珍曰}按陸佃解韓詩外
傳云,生于兩旁謂之瞿,此麥之穗旁生故也。
– 因外型似麥又穗生兩旁,故名瞿麥。
• {別錄曰}瞿麥生太山山谷
– 太山可能指大山,也可能指洛陽嵩山一帶。可能喜冷
涼且需水氣足。
• {宏景曰}有兩種,一種為大,花邊有叉椏,未知何者是也。
今市人皆用小者,復一種葉莖相似,而有毛,花晚而甚赤。
• {頌曰}二月至五月開,七月結實作穗。…河陽河中府出者,
苗可用;淮甸出者,根細,村民取作刷帚。
• {時珍曰}田野生者花大如錢,紅紫色,人家栽者花稍小而
嫵媚,有細白粉紅紫赤斑斕數色。
– 南朝(456-536)時已觀察到種類差異。
– 大花者可能為中國石竹,花小者可能為瞿麥。
中國石竹
瞿麥
長萼瞿麥
(Flora of China, 2001)
三種常見石竹類植物之分布 西洋石竹
(亞洲種)
花壇石竹品種
D. barbatus ‘Barbarini’
(GoldSmith by Syngenta, 2016)
Dianthus barbatus ‘Dash’
Dianthus barbatus ‘Sweet’
(PanAmerican Seed, 2016)
Dianthus barbatus ‘プラネット(Branet)’或‘Heritage’
標榜低春化需求
Dianthus barbatus F1 ‘新緋車’
F1 ‘Carousel’
(Takii Seed, 2005)
切花石竹品種
(HilverdaKooij B.V., 2016)
西洋石竹品種比一比
品種
株高
(cm)
幅寬
(cm)
花期
培育期
(週)
GoldSmith©
Barbarini 20-25 25-30 春-夏初 15-18
PanAmerican©
Dash 38-51 30-36 夏季除外 13-17
Sweet 46-91 25-30 春、秋 15-20
Takii©
Heritage 40-50 秋-冬 16-18
F1 Carousel 50-60 冬-春 20-22
適切花
D. chinensis ‘Super Parfait’
Red Peppermint
Raspberry
Strawberry
Red
D. chinensis ‘Venti Parfait’
(GoldSmith by Syngenta, 2016)
D. chinensis F1 ‘Diana’
(Hem Genetics, 2016)
D. chinensis F1 ‘Grace’
(半重瓣品種)
中國石竹品種比一比
品種
株高
(cm)
幅寬
(cm)
瓣性 花期
培育期
(週)
GoldSmith©
Super Parfait 15-20 20-25 單瓣 春 15-18
Venti Parfait 15-20 20-25 單瓣 春 15-18
Hem Genetics©
Diana 15-20 單瓣 春 14-15
Grace 15-20 半重瓣 春 14-15
Dianthus ×barbatus
‘Sprinter’或‘Polar’
Dianthus ×barbatus
F1 ‘Telstar’
(Takii Seed, 2007)
(Takii Seed, 2007)
Dianthus ×barbatus ‘Diamond’
(Sakata Ornamentals, 2016)
D. hybrid
‘Diabunda’
(GoldSmith by Syngenta, 2016)
Dianthus chinensis × barbatus ‘Floral Lace’
D. chinensis × barbatus
‘Ideal Select’
(PanAmerican Seed, 2016)
Dianthus ×barbatus ‘Amazon’ Dianthus ×barbatus ‘Bouquet’
Dianthus ×barbatus ‘Dynasty’
(PanAmerican Seed, 2016)
雜交石竹比一比
株高
(cm)
幅寬
(cm)
花期 培育期
(週)
Takii©
Telstar 25-35 秋至春 11-14
Sakata©
Diamond 18 24 秋至春 10-12
GoldSmith©
Diabunda 20-25 20-25 秋至春 10-12
PanAmerican©
Ideal Select 20-25 20-25 秋至春 9-10
Dianthus hybrid ‘Jolt’ (標榜耐熱)
(PanAmerican Seed, 2016)
(Takii Seed, 2005)
Dianthus superbus Dianthus japonicus
日本地區的石竹(撫子)
• 日本地區產有4種石竹屬物種
– 瞿麥/河原撫子
• 長萼瞿麥/河原撫子(D. superbus var. longicalycinus)
• 瞿麥/伊勢河原撫子(D. superbus var. superbus)
• 高原河原撫子(D. superbus var. speciosus)
• 雲井撫子(D. superbus var. amoena)
– 姬濱撫子(D. kiusianus)
– 濱撫子/日本石竹(D. japonicus)
– 信濃撫子(D. shinanensis)
橘保國(1715-1792) 繪. 畫本野山草.
飯室庄左衛門(1789~1858?). 草花図譜.
收錄於伊藤圭介(1803~1901)編纂. 植物図説雑纂.
岩崎常正. 1921.
本草圖譜
長谷酔華. 1863. 撫子培養手引草
<瞿麥品類圖>
喜齋立祥. 1866. 東都谷中撫子
歌川國貞(1786~1865). 國芳國貞錦繪.
野村正治郎 編. 1931-1932.
御所とき江戸とき. 第4輯
撫子紋家徽
小倉淳 画、広川松五郎 編. 1920.
命ある野乃草 : 画集
日本地區特有之雜交石竹
• 伊勢撫子(D. isensis)
–中國石竹長萼瞿麥
• 鴉撫子(D. nigritus) *可能稀見於京都藥草園
–中國石竹日本石竹
(加茂莊, 2016)
進貢給皇室賞玩的石竹
香石竹的歷史
• 與石竹擁有共同的起點
• 中世紀前以香味為主要目的(以前沒有專門花壇石竹)
• 後漸以重瓣花形、花徑為主要選拔目標
• 中世紀後期(約1450年後)逐漸成為重要花卉
• 早期香石竹的特徵
– 植株柔軟易倒伏
– 一年僅開花一次,不耐寒,需耕起保護、重植
Hans Memling
(Netherlandish, active by 1465–died 1494 Bruges)
Date: 1465–75
香石竹的歷史
1840
1936-1937
before 1750
1846
1852-1856
花期 株型/莖 人物
春季一次 較軟
延長至秋季
稍挺 M. Dalmais
品種多樣化
硬挺 A. Alagatiére
C. Marc
Dailledouze & Zeller
William Sim
‘de Mahon’ (Remontant)  ‘Biohon’
‘Atim’  Flemish carnation
Perpetual carnation


康乃馨品種
(HilverdaKooij B.V., 2016)
康乃馨品種
(HilverdaKooij B.V., 2016)
康乃馨品種
(HilverdaKooij B.V., 2016)
Dianthus – A possible lineage
D. plumarius  D. caryophyllus 
D. barbatus (multiflora)
× D. chinensis (sterile, grandiflora)
D. knappii
D. carthusianorum (multiple head)
Old-fashioned pinks (calyx split)
(select based on scent, flower form, colour)
Carnations (cut flower)
(select based on forcing, flower form, color)
Sweet william, modern pinks (mat-forming)
(select based on fast-growth, low-vernalization, flower color)
D. chinensis
×
D. hybrid
1850
1960
300 BC
1450
學名、中名、英名對照表
學名 中名 英名
D. superbus 瞿麥 Garden pinks, wild pinks
D. barbatus 西洋石竹、美女撫子 Sweet Williams
D. (×)plumarius -- Feathered pinks, heirloom pinks,
grass pinks
D. gratianopolitanus -- Cheddar pinks
D. deltoides -- Maiden pinks
D. monspessulanus -- Fringed pinks
D. chinensis 中國石竹、美國石竹 China pinks
D. ×barbatus 五彩石竹 Sweet Williams, modern pinks
D. ×allwoodii 雜交石竹 Allwood pinks
D. arenarius -- Sand pink
英名來源
• Pinks
– 一說源自德語pfingsten或pinksten
– 一說為鱷齒剪刀(pink-sheared)與花瓣邊緣的聯想
• Gilliflowers
– 源自Gillofloures,意指七月開花(July-flower/jolie-fleur)
– 源自義文garofolo或法語giroflée,意指丁香
– Clove gilliflower西班牙人用於酒類之康乃馨“Sops-in-Wine”
• Carnation
– 一說源自拉丁carnis意為肉,源自其原種花朵顏色
– 一說源自十六世紀用於加冕典禮(coronation)
石竹屬相關資源及育種
年輕的物種
• 石竹科在7000萬
年前由莧科演化
分歧 (箭頭)
• 石竹屬約於距今
120-700萬年前演
化出(下方粉色區塊)
(Valente et al., 2010)
Fig. Log-lineage-through-time plot for genus Dianthus (including Velezia), based on 1000 resampled
trees with missing taxa added using the method of Day et al. (2008). The line in bold corresponds to
the maximum credibility tree from the Bayesian dating analysis (conservative calibration). The
shaded panel highlights the time window in which a dramatic increase in diversification rates took
place, coinciding with a period of profound climatic transformations in the Pleistocene (更新世).
Sect.Armerium(E)
Sect.Verruculosi(E)
Africanclade(A)
Eurasian
radiation(E)
Time(millionyearsbeforepresent)
Velezia(E)
(Valente et al., 2010)








(Valente et al., 2010)
Petrorhagia, Dianthus, Velezia
Allochrusa, Acanthophyllum
Vaccaria, Gypsophila, Saponaria
Agrostemma, Petrocoptis, Lychnis, Silene
Moehringia, Arenaria, Holosteum, Stellaria, Cerastium
Drypis, Honckenya, Sagina, Buffonia
Minuartia, Spergula, Spergularia
Loeflingia, Ortegia, Polycarpon
Paronychia, Herniaria
Telephium, Corrigiola
Mollugo
(Valente et al., 2010)
已知可種間雜交組合
• D. arenarius × D. chinensis
• D. caryophyllus × D.
gallicus
• D. caesius × D. plumarius
• D. chinensis × D. knappii
• D. sylvestris × carnation
• D. chinensis × carnation
• D. caryophyllus × carnation
• D. gallicus × arenarius
• D. knappii × D. sylvestris
• D. knappii × D. arenarius
• D. knappii × carnation
• D. barbatus × D. deltoides
• D. barbatus × D. alpinus
• D. barbatus × D. glacialis
• D. areanarius × carnation
(Kottö and Gairdner, 1931)
其他已知不容易成功組合
不易得種子
• D. ×plumarius × carnation
• D. (×)plumarius × D. knappii
• D. (×)plumarius × D. barbatus
後代雄不稔
• D. ×plumarius × D. knappii
• D. carthusianorum × D. gallicus
• D. barbatus × carnation
• D. arenarius × carnation
• D. ×plumarius × carnation
• D. ×plumarius × D. barbatus
• D. microlepis × D. barbatus
• D. microlepis × D. ×plumarius
(Kottö and Gairdner, 1931)
(Buell, 1953)
(Buell, 1953)
(Nakano and Mii, 1993)
Fig. Greenhouse-grown plants of
parental species and somatic hybrid.
(Nakano and Mii, 1993)
(Nakano et al., 1996)
Fig. Intergeneric somatic hybrid plantlets between Dianthus
barbatus and Gypsophila paniculata (SH-1). Bar = 2 cm.
Fig. Flowering plants and flowers of the parental
species and the hybrid, D. caryophyllus (left), the
hybrid (centre), and D. japonicus (right). Flower
variates in the interspecific hybrids.
(Nimura et al., 2003)
(Nimura et al., 2006a)
Fig. Flowering
plants and
flowers of the
parental species
and the hybrid,
D. ×isensis
(left), a hybrid
(centre), and D.
japonicus
(right). Flower
variates in the
interspecific
hybrids.
Fig. Flowers and pollen from the original hybrid and
natural chromosome-doubling amphidiploid.
(Nimura et al., 2006b)
(Nimura et al., 2006c)
Fig. Flowering plants, flowers, and flower
variations from a cross between carnation and D.
×isensis.
其他石竹屬種間雜交案例
• 康乃馨(黃) × D. plumarius (白)粉紅
• D. plumarius (白) × D. knappii 黃
• D. giganteus (白) × D. carthusianorum (粉)淺粉
(Gatt et al., 1998: Lee et al., 2005)
Fig. Biosynthetic
pathways of anthocyanins
and related flavonoids
(Sasaki et al., 2013)
Fig. Representative flower colors of
carnations that accumulate different
flavonoids/anthocyanin molecules.
(Sasaki et al., 2013)
紫色康乃馨 by Suntory©
田中良和 博士Moon Series
(Fukui et al., 2003;Suntory©, 2016)
Hardwork
…
1992
基因選殖
1996
成品
1997
販售
紫色康乃馨What’s the trick?
• 轉殖品種挑選
– 以‘Cerise West Pearl’ (粉紅色鑲白邊品種)
• 基因選殖
– F3’5’H  DFR  cyt P450
• 一定要Delphinidin嗎?
–pel 3,5-diglucoside
• 其他要素
–花瓣細胞形狀
康乃馨
‘Nazareno’
(Sasaki et al., 2013)
臺灣的石竹
• 臺灣產有4種石竹
–玉山石竹D. pygmaeus
–清水山石竹D. seisuimontanus
–巴陵石竹D. palinensis
–瞿麥
• 瞿麥D. superbus var. superbus
• 長萼瞿麥D. superbus var. longicalycinus
玉山石竹
(李, 2004)
(Flora of Taiwan)
清水山石竹
(李, 2004)
巴陵石竹
(李, 2004)
(Flora of Taiwan)
瞿麥
(李, 2004)
長萼瞿麥
(李, 2004)
(Flora of Taiwan)
臺灣石竹屬物種檢索表
1. 葉線形………….玉山石竹
1. 葉線-狹長形
2. 花瓣裂<1/4;萼先端銳尖
………………...巴陵石竹
2. 花瓣裂≥1/4;萼先端圓滑
3. 瓣稜基部具腺毛
……………..... 長萼瞿麥
3.瓣稜基部不具腺毛
……………………..瞿麥
1. 株高≤15 cm;葉線形
…………………..玉山石竹
1. 株高≥25 cm;葉線-狹長形
2. 花瓣裂<1/4
………………...巴陵石竹
2. 花瓣裂>1/4
3. 莖及葉灰白..清水山石竹
3. 莖及葉綠色
4. 萼紅紫色…………瞿麥
4. 萼綠色(略長).長萼瞿麥
by Flora of Taiwan by 李(2004)
臺灣原生石竹屬物種比較
原生
海拔(m)
花期
株高
(cm)
花色
玉山石竹 >3000 夏 6-15 粉紅、粉紫、白
巴陵石竹 600-1800 春至秋 25-45 粉
清水山石竹 2000-2400 夏至秋 30-50 粉
瞿麥 600-3100 全年 30-60 粉、白
長萼瞿麥 1500-2500 全年 40-70 粉、白
西洋石竹 -- -- 25-30 --
中國石竹 -- -- 15-20 --
雜交石竹 -- -- 25-35 --
臺灣研發-石竹‘巴陵紫雲’
• 中興大學
• 日本石竹 × 巴陵石竹
• 單朵花壽命可達13天
• 切花壽命可達1個月
• 具香味
臺灣研發-石竹‘臺東1號-香粉雲’
• 中興大學移轉
• 中國石竹 × 長萼瞿麥
臺灣研發-石竹‘臺東2號-香怡夏’
• (可能為)中國石竹 × 清水山石竹
石竹類栽培習性
生產週數
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
18-2113-15 15-21/11-12
22-2317/19 17-18 / 19-20 (5-9 a.m.有5-10負DIF)
22-2317/19 17-18 / 19-20 (5-9 a.m.有5-10負DIF)
21-2418-20 15
21-2417-18 10-16 / 4-10
22-2317/19 17-18 / 19-20 (5-9 a.m.有5-10負DIF)
18-2115/13 15-24/11-15
西洋石竹
Dash
Barbarini
中國石竹
Super Parfait
Diana
雜交石竹
Telstar
Diabunda
Ideal Select
石竹喜好涼溫,高溫導致徒長
可利用負DIF調節株高
(GoldSmith, 2016;Hem Genetics, 2016;PanAmerican, 2016;Takii, 2016)
移植後置於23oC較14oC者提前30天開花,但花較少
14.4 17.2 20.0 22.8
(Erwin, 2005)
石竹為日中性或非絕對長日植物
(Meng and Runkle, 2014)
Species Temp (oC) Light intensity (fc)
Dianthus barbatus 20 650
Lactuca sativum 25 900-1350
Pelargonium sp. 20 350-550
Pisum sativum 14 400
Salvia sp. 20 300
Silene latifolia 25 800
Stellaria media 25 920
Tropaeolum minus 20 250-450
Viola tricolor 25 1000
Table. Light Intensities at which plants bloomed and produced seed.
(Hendricks and Harvey, 1924)
石竹喜好較強光度
石竹為耐旱、不耐濕之植物
(洪, 2014)
石竹的育種目標
• 耐熱(延長利用期間) *尚未有品種比較報告
• 早花(減少涼溫需求) *尚未有定量及比較報告
• 耐逆境、抗病(多濕/病害)
• 株高(依利用目的而變,兩個極端)
• 同一系列內齊全的市場喜好花色
• 花徑
• 新性狀(蔓性、花瓣質感*、花序形態)
近年重要種原
• D. japonicus (日本石竹)
–被認為具有耐熱性及耐海洋氣候特性
–具一定抗病力
• D. sylvestris group (林石竹)
–原生在南歐地區且多為乾旱地帶
未來研究方向
• 溫度對石竹屬物種/品種生育之影響
• 比較石竹屬物種/品種間對涼溫之需求
• 介質水分對石竹屬物種/品種生育之影響
• 石竹屬耐濕熱性狀(若存在)之導入
• 評估雜交石竹後代之耐熱性
• 評估雜交石竹後代之耐濕性
儘管尚未有科研報告..但已有耐熱品種
*但可能沒有大量商業化Trailing

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Pinks and carnations

Hinweis der Redaktion

  1. 石竹屬植物一般估計有200至300個原生種,分布於舊世界的歐亞大陸,即圖中標示橘黃色塊部分。大致上以南歐-中亞為種原中心,其中又以地中海沿岸為重要之原生地,向東最遠至日本亦有分布。事實上,學者指出石竹屬染色體數在物種間具保守性,皆為2n = (2x =) 30,種原間雜交障礙不強,甚至可在種原重疊分布處觀察到自然雜交種之存在,是故咸認對石竹屬以植物標本及外型進行物種鑑定並不具良好意義,也因此石竹屬植物在分子生物學資料庫建立完善前不易做出種原鑑定。 圖中粉紅色圈起範圍為石竹屬的歐亞分布區域,並且少數可能經由東非向南分布,最遠達南非地區。但據估計,有60%石竹屬物種集中分布在地中海沿岸(綠色)及伊朗-小亞細亞區(藍色),而這僅占了不到分布總區域的10%。
  2. 在這幅15世紀末-16世紀初的掛毯畫中,背景圖案隱藏著石竹。
  3. 西洋石竹D. barbatus,其種名意指綴飾有鬚毛,一個可能是指本屬物種花冠面近花冠管部分會有突出的短毛,也有可能是形容其花瓣末端叉裂。 在1960年代後期,當時GoldSmith公司的老闆Glenn Goldsmith警告育種家詹姆士先生,這個種間雜交組合很有可能會失敗得不到種子。但幸好GoldSmith老闆錯了,而順利得到的種間雜交後代就被命名為‘Queen of Hearts’,並且成功贏得1971年的全美園藝選(All-America Selection Award)花卉組獎。儘管這個西洋石竹與中國石竹的雜交組合得到了具良好開花表現的後代,但其不稔無法結種子。總結此一雜交組合後代之特性,包括來自中國石竹親本的大花性、西洋石竹親本的耐寒性與生長勢,且因不易結種子,相較於以往的花壇石竹品種容易因為大量結籽而在移植後一個半月至兩個月就失去觀賞價值,‘Queen of Hearts’能夠在整個花季期間開花不斷,打破當時一般人對石竹美麗但短暫的印象。 2004 Year of the Dianthus
  4. Jan 30, 1971 The Home Garden - Two Zinnias Head List of All-America Flowers BY ART KOZELKA / Garden Editor Two showy zinnias that feature large, double blooms on dwarf-size plants head the list of six new flowers which have been named as All-America Selections for 1971. A new hybrid sweet corn was the only vegetable winner for 1971. The zinnias are Peter Pan Plum, which is a fine, lavender-rose or plum color, and Peter Pan Pink, a coral pink beauty with a salmon-rose center. The two award-winning flowers have the added distinction of scoring more points in the All-America trials than any other flowers in the almost 40-year history of All-America Selections. Perhaps as newsworthy as the announcement of these zinnia award winners is the fact that they were created by Charles L. Weddle of Palisades, Colo., who is still affectionately known as "Mr. Petunia" for his origination of dozens of outstanding petunias, including Comanche, Prima Donna, and many others that have won All-America awards. It was only four years ago that Weddle switched his hybridizing talents from petunias toward the improvement of zinnias. Just a year later, he produced the popular Wild Cherry Zinnia which won All-America honors. * * * Weddle's new award-winning Peter Pan varieties are certain to be a boon to home gardeners who like to mass zinnias in beds and borders. The plants remain compact, maturing at 12 to 14 inches high, and flaunt their colorful, 3-to-4-inch blooms in profusion throughout the summer. As with all zinnias, the Peter Pans grow easily from seed, which may be started in peat pots indoors and transplanted to the garden. Quite often these potted plants will bear flowers when only 6 inches tall, which means there can be color in the garden right after they have been transplanted. Seeds, of course, also can be sown directly outdoors after soil is warmed if there is no hurry for blooms. * * * The other 1971 flower award winners are Queen of Hearts, a brilliant, scarlet dianthus; Little Darling, a sturdy new, low-growing snapdragon; Silver Puffs, the first dwarf hollyhock; and Southern Belle, a huge-flowering perennial hibiscus. Queen of Hearts, which is the first F1 [first generation] hybrid dianthus, grows 12 to 15 inches tall and about as wide and yields masses of single-flowered blooms 1 to 1 1/2 inches across. Seed can be sown directly in the garden borders. The plants also are excellent in pots for patio display. * * * Little Darling should bring snaps into even greater favor with gardeners for landscape border plantings. Because they grow only 12 inches tall and are compact in habit, they withstand windy locations better than the taller growing varieties. They come in a mixture of snap colors with the new open-face flowers popularized by Bright Butterflies, a previous, taller growing All-America winner. Both Queen of Hearts dianthus and Little Darling snaps were developed by Goldsmith Seeds, Inc. Fluffy, double, silvery-pink flowers are only one of the virtues of Silver Puffs, the award winning hollyhock. Growing to a height of only 2 feet, it is tailor made for home gardens. It never needs staking, and it branches well, with blooms forming on the side branches as well as main branches. It was originated by Sluis en Groot, a Dutch seed firm. Not too many gardens boast hibiscus plantings, but Southern Belle should change this. Plants grow 5 to 6 feet tall and produce fantastically large, single flowers of crimson, red, rose, pink, white, and white with a red eye. It is not uncommon for the blooms to measure 10 inches across. The plants die back each winter, but come back again the following season. For first year blooms on this unusual perennial, seed should be started early indoors. The new hibiscus was developed by Sakata & Co., a Japanese seed firm. The 1970 award-winning vegetable is Early Xtra Sweet, an F1 hybrid sweet corn. It won the praise of All-America judges for its extra sugar content. It is much like its predecessor, Illini Xtra Sweet, but matures two weeks earlier and is more uniform. Ears are 7 to 9 inches long and 1 1/2 to 2 inches in diameter, with 12 to 16 rows of kernels on each cob. You can enjoy this corn over a long season by making several plantings a few weeks apart. Plantings should be made as far away as possible from other corn varieties; otherwise pollen from the other varieties will keep it from developing its extra sugar content. The All-America Selections, which are sponsored by the garden seed industry of the United States and Canada, evaluate the merits of new flower and vegetable varieties for two years in trial gardens throughout North America.
  5. 根據Flora of China及Flora of Taiwan之記述,臺灣及中國大陸地區共產17種石竹,其中包括中國石竹(D. chinensis)、瞿麥(D. superbus)、長萼瞿麥(D. longicalyx)三種分布最為廣泛,又中國石竹與瞿麥皆有納為中藥材使用之記錄,其餘石竹屬物種有3種特別出自臺灣,也就是巴陵石竹(D. palinensis)、玉山石竹(D. pygmaeus)及清水山石竹(D. seisuimontanus),西洋石竹(D. barbatus var. asiaticus)則僅產於吉林省與北韓及俄羅斯交界處,且因與歐洲產的西洋石竹有些許不同(歐洲產者較肥壯、葉片較寬且厚、植株較大、花較多),故被認定屬於亞洲變種。剩下的10種石竹屬物種則僅分布在新疆(尤其北部)及西藏等與中亞交界地帶。 我們可以根據這張將Flora of China中,依記述物種分布省分所繪製之分布圖觀察到,中國石竹與瞿麥的分布地重疊,因此不可見粉紅色區域,中國石竹大約僅分布在黃河以北地區,而瞿麥則仍持續向華中華南地區分布,但大約在福建、廣東一帶即不再有分布之記錄,相較之下,長萼瞿麥則更南、向海南及臺灣地區分布。其中,以山東、河南、陝西、寧夏、甘肅、山西、河北、吉林、內蒙一區,為三個物種的共同重疊區域。而位在河南省的古都洛陽一直是中國文化的中心,可以推測古人在這裡或鄰近地區可以同時觀察到這三種最為常見的石竹種類,因此在本草綱目中會記述有花大、花小之分。 有趣的是,在Flora of Taiwan的記述中,將瞿麥及長萼瞿麥視為同種,長萼瞿麥為變種(variety)而非種(species),並且認定臺灣產的瞿麥亦為變種應稱為臺灣瞿麥(var. taiwanensis)。但二者又以長萼瞿麥分布之海拔較高(1000-2100 m),臺灣瞿麥分布海拔較低且少見,採集紀錄多半在東半部山區草地中(800-1800 m)。不論如何,Flora of Taiwan中所有的石竹物種多半分布在山區,僅巴陵石竹生長在海拔600 m的臺灣北部(桃園巴陵),又即使是Flora of China中分布最南的長萼瞿麥,最低也分布至海拔800 m地帶,顯示本屬植物多半仍喜冷涼氣候。
  6. 這類花簇成球的石竹,雖然以正式學名應該稱為西洋石竹(D. barbatus),但國內在販售本類商品時,多半稱為美女撫子,這是沿用日本種苗商目錄上的漢字美女なでしこ而來。麻煩的是,D. deltoides有時也以美女撫子這個名字販售。西洋石竹在花壇石竹當中多半花朵較小,但單一花序有非常多的花朵,且幾乎是同時開放,因此會形成花球的樣子。但對臺灣花壇布置來說,這樣一球一球的特性並不符合我們的喜好。
  7. 西洋石竹品種中也有可做為花壇背景使用,或兼可剪取切花的品種,但在臺灣幾不可見。
  8. 這是荷蘭公司HilverdaKooij開發的切花石竹品種(其實本來是兩家公司,但後來合併成一家)。
  9. 在這邊就將前面幾個介紹的西洋石竹品種特性整理成表以比較,可以看到若以臺灣花壇布置喜歡較低矮的品種而言,只有GoldSmith公司的Barbarini可以符合要求,但誠如前面提到的,臺灣的花壇布置很少選用花球型的西洋石竹品種,因此僅供參考。
  10. 有別於西洋石竹品種,中國石竹品種多半低矮,反而少見株高較高者,可能是因為其單一花序花朵數較西洋石竹少,因此花朵排放較接近平面狀,適合花毯型景觀設計,是故沒有育種公司會反向操作選拔較高、缺乏觀賞價值的後代。GoldSmith公司的Super Parfait一直以來都是臺灣地區有穩定少量生產的石竹品種,一方面是因為植株低矮、強壯,一方面是該公司此品系主打具有眼睛(eye)斑紋的雙色花色(只有Red是幾乎單色),在眾多石竹產品排開時,大花及強對比色很容易吸引消費者的目光。而另一個較新推出的Venti Parfait則是強調可以高密度生產,而品質不致因此下滑。 就像西洋石竹被稱為美女撫子一樣,中國石竹因為主要流通品種Super Parfait來自美國,因此被農民稱為美國石竹,而穗耕公司沿用日本種苗目錄的用字,則有三寸石竹、五寸石竹等名稱。 *老品種為‘Parfait’,近年則是‘Super Parfait’,可能是改良花色、生長習性及穩定度。
  11. 另外,由荷蘭Hem Genetics選育的Diana系列也是一個曾經推行過的老品種,近年則比較少見。此外,該公司還有一個Grace中國石竹品種,則是以半重瓣為賣點,但也較少出現在市場上。
  12. 就這張比較表而言,其實兩家公司的產品差別不大,在臺灣地區以Super Parfait較常見的原因,可能是荷蘭公司的種子較貴、不易湊單或對方不願意出貨到台灣等因素。
  13. 雖然剛剛看到了不少石竹品種,但其實這邊開始才是所謂花壇石竹的主戰場,也就是以西洋石竹和中國石竹等材料進行種間雜交所得到的雜交石竹品種。首先看到的是臺灣地區最重要的品種,由Takii公司所推出的Telstar,一般稱為天王星,雖然花色至少有10種,但臺灣地區主要只栽培雙色和一部分較淺的單色品種,非常少見白色,主要是因為民眾容易將白色和喪事聯想在一起。在天王星之後,Takii公司也推出Polar,在臺灣曾以北極星之名販售,但農民並未有良好反應,是故天王星一直都是臺灣地區重要的石竹品種。
  14. 毯系(矮牽牛中也有Carpet這個品系名)是Takii公司早年推出的品種,現在已被淘汰,可能是因為低溫需求較天王星更多的緣故。
  15. Sakata公司也算是日本花卉育種公司的一大龍頭,但是說到石竹品種選拔方面,Sakata公司的品種相對而言市佔率似乎不高(而且只有一個品系),僅在康乃馨部分有種子品系,是供應盆花康乃馨種子的公司之一。
  16. *老品種為‘Ideal’,新品種為‘Ideal Select’
  17. PanAmerican公司推出的‘Amazon’及‘Bouquet’都是屬於外型比較類似西洋石竹的雜交石竹品種,株高也偏高,甚至Bouquet品系的商品介紹上指出可以剪下做為切花利用。而 ‘Dynasty’則是重瓣品種。
  18. PanAmerican公司近年著眼全球氣候變遷,不斷推出標榜耐熱的新品種,就連石竹也不例外。但這種外型的石竹品種在臺灣能有多少市場還有待觀望。
  19. 除了前面提到的西洋石竹、中國石竹和雜交石竹,其實在東亞地區還有一些小具市場的石竹類,但多半是以切花利用為主。包括河原撫子(也就是瞿麥)及紅撫子或稱深山櫻(也就是日本石竹)。不過穗耕種苗公司也將部分日本來的中國石竹品種稱為日本石竹,應有所區分。
  20. *加茂莊. 2016. 加茂花菖蒲園の伊勢撫子. <<http://www.kamoltd.co.jp/kakegawa/nadeshi.html>> 伊勢撫子は、江戸時代に三重県松阪地方で改良が進んだナデシコの一種で古典園芸植物と言えます。別名、大阪撫子、御所撫子と言い、三重県の天然記念物に指定されています。 セキチクの改良種、あるいはセキチクとカワラナデシコの交雑種より改良されたとも、中国から渡来したと思われるカラナデシコが、薩摩、大阪と伝わるうち に変異を起こし、伊勢(松阪)を経て江戸に伝わったものとも考えられています。江戸時代に光格天皇はことのほかこの撫子をお気に召し、京都市内の宝鏡寺に 下げ渡されたものが、こんにちでも保存栽培伝されています。
  21. 在1750年左右前,法國地區的香石竹仍然只有在春天會開花,不耐溽暑及寒冬,冬天必須挖起植株保護。在1750年後,正確的人及時間已不可考,則出現一種稱為Remontant的香石竹,已經具有花期較長的特性,其中的‘de Mahon’或‘Mahon’或名‘St. Martin’的品種,則是意指可以持續開花到11月聖馬汀節時的品種。法國里昂(Lyon)的園丁M. Dalmais先生以剛剛所提到的 ‘de Mahon’和另一個香石竹‘Biohon’或‘Bichon’(比熊犬)雜交,獲得‘Atim’,並再與Flemish carnation雜交,獲得的後代持續進行雜交與回交,最後得到一系列花期較長且株型較挺直的香石竹品種(1840-1846推出),使香石竹的花期延長,更能為人利用。 在Dalmais先生之後,則由M. Schmidt承繼這批植物材料,繼續雜交選育出不少優良後代,但在1850左右被迫放棄這項工作,而由A. Alagatiére承接。Alagatiére先生是現代香石竹另一位重要的推手,他從這批材料中選拔出莖梗非常硬、直的後代,在後續的幾年中被稱為tree carnation,也就是有別於以往莖柔軟、無法撐托大而重瓣花朵的香石竹。在1852到1856年間,位在美國紐約州的Charles Marc先生和Dailledouze&Zeller公司,競爭選拔自法國可能是Alagatiére先生育成的種子材料,推出許多新的香石竹品種。 在1936或1937年美國緬因州的William Sim先生推出‘William Sim’香石竹,這是近代許多香石竹品種的來源親本,其容易芽變特性,使香石竹的新品種如同雨後春筍一般在世界各地被選拔而出。‘William Sim’原本是一個深紅色的重瓣品種,但在幾年間,陸續選出粉紅色、白色、淺黃色鑲邊等花色的變異株。
  22. 單瓣型在臺灣販售時常稱為雜交石竹,認為是取康乃馨與其他原生種雜交所得,但有可能僅僅是選拔過程中產生的特殊型。
  23. *Midwest Gardening <<http://www.midwestgardentips.com/dianthus_allwood_hybrids.html>>
  24. Monticello <<https://www.monticello.org/site/house-and-gardens/pinks-gilliflowers-carnations-exalted-flowers>> Ward, C.W. 1903. The American Carnation – How to grow it. Applewood Books, Bedford, MA, USA. p.17-18.
  25. 分子生物學利用細胞核染色體上轉錄出核醣體RNA (ribosomal RNA, rRNA),其中小次級單元序列和大次級單元序列間之轉錄區間,以及葉綠體或質體中負責轉錄轉運RNA (transfer RNA, tRNA)及其他RNA之相關基因序列,此類基因具有保守性,因此其序列變異程度可以做為推測演化時間之依據。 Valente等人(2010)分析104個石竹屬物種及近緣屬(Velezia、Petrohagia及Saponaria)之ITS1-5.8S-ITS2、trnK-matK、psbA-trnK及trnH-psbA序列。並且利用已報導的植物演化分歧時間及化石等進行推估。結果顯示盡管莧科與石竹科在七千萬年前已演化分歧,但石竹科的石竹屬卻約在距今120-700萬年才演化形成。
  26. 而根據石竹屬內物種序列之變異,並多次建立演化樹,則顯示大約在距今150~200萬年前開始多樣化速率(diversification rate)明顯提升,而此時恰伴隨更新世時期的乾燥氣候,顯示這樣的氣候環境可能有助於石竹屬物種的多樣化。
  27. 根據這張演化分枝樹圖可以觀察到:在Valente等人(2010)參試的104個石竹屬及1個Velezia物種,共可分為五個演化分支,然而不論是哪一個演化分支群,多樣化都是發生在約150~200萬年前。當中又以歐亞擴散(Eurasian radiation)分支群的多樣化最為豐富,現代花壇石竹與康乃馨的重要親本如D. plumarius、D. barbatus、D. chinensis、D. superbus、D. knappii等都囊括在內。 Sect. Armerium: D. corymbosus, D. deltoides, D. tymphresteus, D. viscidius, D. myrtinervius, D. tenuiflorus, D. armeria, D. diffusus Sect. Verruculosi: D. strictus, D. cyri, D. tripunctatus Velezia: Velezia rigida African clade: D. leptoloma, D. longiglumis, D. crinitus, D. serrulatus, D. laingsburgesis, D. excelsus, D. albens, D. zeyheri, D. thunbergii, D. bolusii, D. caespitosus pectinatus, D. caespitosus caespitosus, D. mooiensis, D. basuticus, D. micropetalus, D. namaensis Eurasian radiation: D. barbatus, D. callizonus, D. collinus, D. giganteiformis, D. glacialis, D. henteri, D. integer, D. leucopheus, D. libanotis, D. petraeus, D. pinifolius, D. serotinus, D. strymonis, D. subacaulis, D. trifasciculatus, D. turkestanicus, D. eretmopetalus, D. serratifolius, D. plumarius, D. superbus (Russia), D. anatolicus, D. calocephalus, D. micranthus, D. erinaceus, D. giganteus, D. crinitus, D. orientalis, D. cretaceus, D. zederbaueri, D. gallicus, D. hyssopifolius, D. furcatus, D. pavonius, D. pallens, D. zonatus, D. pygamaeus, D. superbus (Japan), D. stenopetalus, D. rupicola, D. sylvestris, D. arrosti, D. ciliatus, D. haematocalyx, D. carmelitarum, D. biflorus, D. capitatus, D. cruentus, D. gracilis, D. knappii, D. fischeri, D. chinensis, D. subulosus, D. versicolor, D. microlepis, D. gredensis, D. serrulatus, D. lusitanus, D. multiaffinis, D. pyrenaicus, D. langeanus, D. laricifolius, D. charidemi, D. legionensis, D. toletanus, D. seguieri, D. pungens, D. multiceps, D. lusitanus, D. crassipes, D. costae, D. cintranus, D. broteri, D. benarnensis, D. anticarius, D. algetanus
  28. 這是前一頁演化分枝樹中歐亞擴散部分的其中一種可能的親緣關係圖。假設以地中海沿岸為種原中心向外演化,並且將分布最為廣泛常見的D. barbatus為基礎分枝點,則依親緣關係由近到遠,分別為D. plumarius (花壇及切花石竹的共同親本)、瞿麥(俄羅斯種)、玉山石竹及瞿麥(日本種)、D. knappii、D. carthusianorum,最後為中國石竹。令人驚訝的是玉山石竹和日本種瞿麥雖然在以往的親緣分類上是不同的種,但在演化時間及親緣關係上卻顯示是相當親密的。毫無意外的西洋石竹和中國石竹的親緣關係較遠,也因此當年GoldSmith公司預測這兩個物種的種間雜交可能不容易成功,這對當時植物分子親緣鑑定還不普遍而言,是相當準確的推測。
  29. 根據上一頁石竹科演化分歧時間,我們也可以進一步推斷出石竹屬與其他石竹科植物之可能親緣關係。例如Dianthus與Petrorhagia及Velezia是在同一個演化分支當中,Petrorhagia (膜萼花屬)可能較早演化出,而Velezia則是較Dianthus晚點演化出之物種。再向前有園藝栽培的物種則是王不留行(Vaccaria)、滿天星(或稱霞草Gypsophila)及皂質草(Saponaria)。在1996年有一篇文章發表,利用電擊使去細胞壁的石竹及滿天星細胞融合,雜交體細胞可以分裂、生成枝條,但全數非常低矮(dwarf)且早熟開花,即使出瓶也無法正常生長,因此並未能獲得實際有利用價值的雜交後代。 而更早演化出的則包括麥仙翁(梅花石竹, Agrostemma)、Petrocoptis、剪秋羅(Lychnis)、雪輪(或稱蠅子草, Silene)等。更早之前演化出的物種多半是小型草本植物,甚至是一些現在全球常見的雜草。
  30. 以PEG融合西洋石竹與中國石竹,瓶內苗不斷開花且極端簇生。後有出瓶持續觀察。 *千葉大學
  31. 以PEG法融合康乃馨及中國石竹體細胞,深粉中國石竹與紅康乃馨體細胞雜交後代花色為紅。
  32. 日本新潟科技大學和千葉大學的中野等人將滿天星‘Bristol Fairy’和西洋石竹原生質體脫出,共同培養於KM8p培養基中,然後交替以50V 2MHz交流電(20 s)和1kV直流電(30 ms)電擊兩回。結果僅滿天星和雜交體細胞有發生細胞分裂及再生。在培養2個月後才能觀察到可見的細胞叢,繼代300個癒傷組織團中,僅8個再生綠色芽原體,其中又僅2個生成枝條,再次繼代後凡是置於無生長調節劑1/2 MS培養基者,皆生長停滯、死亡,添加GA3可使枝條抽長。體細胞雜交後代的花朵形似石竹,但即使持續繼代2年,也未獲得可正常生長的枝條。
  33. 只有以香石竹為親本與日本石竹雜交可取得後代,且不需倚賴胚拯救 種間雜交後代外型介於兩親本之間,但花色齊一,為洋紅色(reddish purple)。
  34. 以多倍體化處理康乃馨與日本石竹種間雜交後代(不稔),可恢復稔性,且植株較為粗壯。
  35. 伊勢石竹與日本石竹雜交 後代自然多倍體化,產生具稔性花粉
  36. 伊勢石竹與康乃馨雜交,後代有重瓣、單瓣、似伊勢石竹(三倍體)等變化 伊勢石竹具早花特性,且可導入雜交後代中 兩向雜交有特定康乃馨品種為親本才能取得種子
  37. 李祖文. 2004. 台灣產石竹科石竹亞科植物之分類研究. 國立師範大學生命科學系碩士論文.
  38. 在Flora of Taiwan中並未將清水山石竹列入。 兩位作者都將葉片線形做為將玉山石竹與其他原生種石竹區分之性狀,而李(2004)則強調株高可以做為重要的指標,因為玉山石竹即使在人工栽培環境下生長,株高也低矮。另外,萼筒長度在17-20 mm,較其他石竹屬物種在25 mm以上,為另一可利用的物種區別性狀。兩位作者都認同在剩下的3~4個原生石竹物種中,花瓣頂緣淺裂,不超過花瓣長度1/4為一可信的鑑別物種依據,可將巴陵石竹自其他植株較高的原生種分出。而其餘2~3個物種中,皆具有明顯膨大的節,但又以清水山石竹最為明顯,且清水山石竹葉片表面綠色具白粉,基生葉緊密排列(簇生特性)。而瞿麥及長萼瞿麥在Flora of Taiwan中主張長萼瞿麥為花瓣基部具有腺毛,瞿麥則否,但李(2004)觀察野外實際植株狀態則指出腺毛可能不是理想的鑑別依據,而採取與Flora of China相同的鑑別依據,即長萼瞿麥的萼為綠色,瞿麥則具有紅斑或為紫紅色。
  39. Erwin, J. 2005. Factors affecting flowering in ornamental plants. In: M.B. McDonald and F.Y. Kwong (eds.). Flower seeds: Biology and technology. CABI Publishing, Cambridge, MA, USA.
  40. 雖然這裡明確寫出必須是在30oC一個月間都開花,但實際是1999年在伊利諾州(West Chicago)田間(氣溫記錄為日/夜溫30/17)進行耐熱比較試驗,種植於5加侖盆,直接日曬,新品種可以生長良好、開花,而Telstar則33%死亡率,且有葉燒發生。
  41. 夏季平均溫
  42. 冬季平均溫,耐寒帶(hardiness zone)圖