Journal of Ecology ( IF 5.3 ) Pub Date : 2024-05-28 , DOI: 10.1111/1365-2745.14341 Anthony J. Davy 1 , Alan J. M. Baker 2 , Arthur J. Willis 3
Sea Campion. Caryophyllaceae, subfamily Caryophylloideae. Silene uniflora Roth is a long-lived, mat-like perennial herb, with leafy shoots remaining green at the base and overwintering just above ground level.
Main tap-root producing secondary roots that vary from descending nearly vertically to a shallower spreading system. Stems procumbent to ascending, (5)10–30(60) cm, with short internodes, lower parts giving rise to new shoots to form loose or compact tufts; prostrate stems sometimes rooting adventitiously at nodes. Leaves entire, non-petiolate, opposite, decussate, glaucous and usually glabrous; laminae variable in shape and size, from narrowly oblanceolate, elliptic to linear, (10)15–30(50) × (1)3–6(10) mm. In coastal populations, leaf margin often ciliolate. Flowering stems mostly with three flowers, but varying from a single terminal one rarely to seven, in a little-branched dichasial cyme (unequal development of the pair of lateral branches not uncommon). Bracts herbaceous, sometimes with a scarious margin, narrowly lanceolate, channelled. Pedicels often elongated. Flowers 20–25 mm diameter, erect, perfectly actinomorphic. Sepals 5, joined for most of their length; calyx campanulate to urceolate or subglobose, inflated with anastomosing purplish veins, 14–21 mm long, varying in diameter from inflated (13 mm) to cylindric (8 mm); teeth c. 3 mm. Petals white or slightly purplish, 5, free, with erect narrow claw mostly 12–16 mm long; laminae of petals normally 10–13 mm long, often overlapping and fairly deeply bilobed, hiding the calyx mouth. Corona (a crown-like ring of appendages between the petals and stamens), at the throat of the corolla, represented by two small but distinct scales or flaps per petal. Stamens normally 10, but some may be represented by vestiges only. The usually white filaments arise from the carpophore (the internode between the calyx and corolla, 3.0–4.2 mm). Anthers 1.5–2.5 mm, often purple (with anthocyanin). Styles 3(−5), free to the base. Ovary 1-celled, but septate at base. Capsules 5–10 × 4–8 mm, subglobose and papery, enclosed in the calyx, with a wide mouth and six strongly recurved teeth. Seeds blackish, rounded-reniform, 1.2–2.0 × 1.0–1.3 mm, tubercled or ‘armadillo’ (with flat plates). Air-dry mass of seed c. 1.4 mg. This information is largely derived from Marsden-Jones and Turrill (1957), who provide extensive details of variation in flower and seed structure in many populations (both British and from elsewhere), Clapham et al. (1987) and Sell and Murrell (2018).
The close relationship between Silene vulgaris (Moench) Garcke and S. uniflora has long been recognized, but the latter was afforded specific rank by Roth (1794). Following Withering (1796), British floras referred it to S. maritima for nearly 200 years (Babington, 1881; Clapham et al., 1962; Hooker, 1884). However, Clapham et al. (1987) and the first edition of Flora Europaea (Chater & Walters, 1964) demoted it again to one of eight subspecies of S. vulgaris. Subsequently, in the second edition of Flora Europaea (Chater et al., 1993), S. uniflora was restored to specific rank, with four subspecies: subsp. uniflora, subsp. thorei (Dufour) Jalas, subsp. prostrata (Gaudin) Chater & Walters and subsp. glareosa (Jordan) Chater & Walters; of these, only subsp. uniflora occurs in Britain. The basis for this treatment is described by Chater and Walters (1990), who refer to five of 14 characters given by Marsden-Jones and Turrill (1957) of value ‘for diagnostic purposes’. These features are winter habit, inflorescence, flower shape, corona and teeth of the capsule. Although, unlike subsp. uniflora, subspp. prostrata and glareosa are hemicryptophytes, they have the other four diagnostic characters of S. uniflora. Subspecies prostrata and subsp. glareosa have inclined, slightly zygomorphic flowers and distinctive leaf shape, subsp. prostrata occurring on mountain rocks in S. Europe and subsp. glareosa usually on calcareous screes from the Pyrenees to the E. Carpathians. Subsp. thorei, with erect, actinomorphic flowers, is wholly maritime, on coastal sands of W. France (and now extinct in N. Spain). Currently, only three subspecies are recognized by Plants of the World Online (2024): subsp. uniflora, subsp. thorei (Dufour) Jalas and subsp. petraea (C. Hartm.) Jonsell & H.C. Prent. The last of these, featuring small-flowered, procumbent plants with few-flowered inflorescences, is endemic to the islands of Öland and Gotland in the Baltic.
Silene uniflora and S. vulgaris were the subject of a remarkable, seminal study in the development of ‘experimental taxonomy’, elaborated in Marsden-Jones and Turrill (1957). The considerable similarity between the two species has led to some confusion, especially in the older literature. Silene uniflora has quite often been referred to as S. vulgaris under the earlier names of the latter of S. cucubalus Wibel and S. inflata Sm., especially in plants growing on metalliferous soils. Distinguishing features of S. uniflora and S. vulgaris include, in the former, distinct coronal scales, herbaceous bracts and recurved teeth on the capsule, contrasted with small coronal scales, scarious bracts and erect teeth on the capsule in S. vulgaris. Many distinctive characters are tabulated by Baker and Dalby (1980). Both seed morphology (testa ornamentation and seed shape) and allozymes show a separation between Silene vulgaris and S. uniflora, and support the taxonomic treatment of ‘petraea’ as a subspecies of Silene uniflora (Runyeon & Prentice, 1997b).
Silene as currently circumscribed is a diverse genus of more than 700 species with an intricate taxonomic history. A recent large-scale analysis of its molecular phylogeny (Jafari et al., 2020) places S. vulgaris (and therefore presumably S. uniflora) in subgenus Behenantha, section Behenantha.
Studies of genetic variation in S. uniflora were pioneered by the early allozyme study of Baker et al. (1975), who failed to find any association between habitat and gene frequency. Subsequent work has focused on its genetic differentiation from S. vulgaris (see Sections 8.2 and 10), using morphometrics (Runyeon-Lager & Prentice, 2000), allozyme variation (Prentice & Giles, 1993; Runyeon & Prentice, 1996, 1997a, 1997b) and plastid DNA markers (Prentice et al., 2011).
Silene uniflora is a showy, native, perennial herb, widespread on coastal shingle, sands and cliffs around Britain and Ireland but is also very localized inland on mountains and waste from metal mining.
中文翻译:
英国和爱尔兰的生物区系:Silene uniflora‡
海坎皮恩。石竹科,石竹亚科。 Silene uniflora Roth 是一种长寿、席状的多年生草本植物,其叶芽在基部保持绿色,并在地面上方越冬。
主要的主根产生次生根,其变化范围从几乎垂直下降到较浅的蔓延系统。茎平卧至上升,(5)10-30(60)厘米,节间短,下部生出新枝,形成松散或紧凑的丛生;匍匐茎有时在节处不定生根。叶全缘,无叶柄,对生,交叉,有白霜并且通常无毛;叶片的形状和大小各不相同,从狭倒披针形、椭圆形到线形,(10)15–30(50)×(1)3–6(10)毫米。在沿海种群中,叶缘通常有纤毛。花茎大多有三朵花,但从很少的单一顶生花到七朵花不等,形成小分枝的二歧聚伞花序(一对侧枝不等发育并不罕见)。草质的苞片,有时具粗糙的边缘,狭披针形,沟状。花梗通常拉长。花直径20-25毫米,直立,完全辐射对称。萼片5,大部分长度相连;花萼钟状到瓶形或近球形,膨胀有网状紫色脉,长14-21毫米,直径从膨胀的(13毫米)到圆柱形(8毫米)不等;牙齿c. 3毫米。花瓣白色或略带紫色,5枚,离生,具直立狭瓣,大多长12-16毫米;花瓣的薄片通常长 10-13 毫米,经常重叠且相当深的双裂,隐藏萼口。花冠(花瓣和雄蕊之间的冠状附属物环),位于花冠的喉部,由每个花瓣两个小但不同的鳞片或瓣片代表。雄蕊通常为10,但有些可能仅由残迹代表。通常白色的花丝源自果柄(花萼和花冠之间的节间,3.0-4.2 毫米)。花药1.5-2.5毫米,通常紫色(含花青素)。花柱 3(−5),离基部。 子房1室,但基部有隔膜。蒴果5-10×4-8毫米,近球形和纸质,封闭在花萼内,有宽口和六个强烈下弯的齿。种子淡黑色,圆形肾形,1.2-2.0 × 1.0-1.3 毫米,有结节或“犰狳”(有平板)。种子的风干质量c . 1.4 毫克。这些信息主要来自 Marsden-Jones 和 Turrill ( 1957 ),他们提供了许多种群(英国和其他地方)花和种子结构变异的广泛细节,Clapham 等人。 ( 1987 ) 以及 Sell 和 Murrell ( 2018 )。
Silene vulgaris (Moench) Garcke 和S. uniflora之间的密切关系早已得到认可,但后者由 Roth ( 1794 ) 给予特定的排名。在 Withering( 1796 年)之后,英国植物区系将其称为S. maritima近 200 年(Babington, 1881 ;Clapham 等人, 1962 ;Hooker, 1884 )。然而,克拉彭等人。 ( 1987 ) 和第一版《欧洲植物志》(Chater & Walters, 1964 ) 再次将其降级为S. vulgaris的八个亚种之一。随后,在《欧洲植物志》第二版(Chater et al., 1993 )中, S. uniflora被恢复到特定等级,有四个亚种: subsp. subsp. 。单花亚种thorei (Dufour) Jalas,亚种。 prostrata (Gaudin) Chater & Walters 及其亚种。 glareosa (约旦)Chater & Walters;其中,只有亚种。 uniflora出现在英国。 Chater 和 Walters ( 1990 ) 描述了这种治疗的基础,他们提到了 Marsden-Jones 和 Turrill ( 1957 ) 给出的具有“诊断目的”价值的 14 个字符中的 5 个。这些特征是冬季习性、花序、花形、花冠和蒴果的齿。虽然,与亚种不同。单花亚种prostrata和glareosa是半隐植物,它们具有S. uniflora的其他四个诊断特征。前列腺亚种和亚种。 glareosa有倾斜的、稍微左右对称的花朵和独特的叶子形状,亚种。 产于南欧山岩上的prostrata及其亚种。 Glareosa通常生长在从比利牛斯山脉到东喀尔巴阡山脉的钙质碎石上。亚种。雉鱼具有直立的辐射对称花,完全是海洋性的,生长在法国西部的沿海沙滩上(现已在西班牙北部灭绝)。目前,《世界植物在线》( 2024 )仅认可三个亚种:亚种。单花亚种thorei (Dufour) Jalas 及其亚种。 petraea (C. Hartm.) Jonsell & HC Prent。最后一种以小花、平卧植物和很少花的花序为特色,是波罗的海厄兰岛和哥特兰岛的特有种。
Silene uniflora和S. vulgaris是“实验分类学”发展中一项引人注目的开创性研究的主题,Marsden-Jones 和 Turrill ( 1957 ) 对此进行了详细阐述。这两个物种之间相当大的相似性导致了一些混乱,特别是在较古老的文献中。 Silene uniflora通常被称为S. vulgaris,后者的早期名称为S. cucubalus Wibel 和S. infata Sm.,特别是在含金属土壤上生长的植物中。 S. uniflora和S. vulgaris的显着特征包括,前者有明显的冠状鳞片、草质苞片和蒴果上的下弯齿,而S. vulgaris则有小的冠状鳞片、可怕的苞片和直立的蒴果齿。 Baker 和 Dalby( 1980 )列出了许多独特的特征。种子形态(种皮装饰和种子形状)和等位酶均显示Silene vulgaris和S. uniflora之间的分离,并支持将“ petraea ”作为Silene uniflora亚种的分类处理(Runyeon & Prentice, 1997b )。
目前所界定的硅烯是一个拥有 700 多个物种的多样化属,具有复杂的分类历史。最近对其分子系统发育的大规模分析(Jafari 等人, 2020 )将S. vulgaris (因此可能是S. uniflora )置于Behenantha亚属Behenantha组。
Baker 等人的早期等位酶研究开创了单花S. uniflora遗传变异的研究。 ( 1975 ),他未能发现栖息地和基因频率之间的任何关联。随后的工作集中于其与普通链球菌的遗传分化(参见第 8.2 和 10 节),使用形态计量学(Runyeon-Lager & Prentice, 2000 )、等位酶变异(Prentice & Giles, 1993 ;Runyeon & Prentice, 1996、1997a 、 1997b ) )和质体 DNA 标记(Prentice 等, 2011 )。
Silene uniflora是一种艳丽的本土多年生草本植物,广泛分布于英国和爱尔兰周围的沿海卵石、沙滩和悬崖上,但也非常集中于内陆山区和金属采矿废料中。