Alchemilla L. – Lady's mantle
In Alchemilla, apomixis was documented rather early (Murbeck 1897, 1901, Strasburger 1905), which is of autonomous aposporia type: the embryo sac develops from a cell of the nucellus, but not from the spore-forming tissue, the archespore. No fertilization is necessary for the development of the endosperm.
Only apomictic specimens are known from Germany, sexual species have not yet been documented in Europe. Since no functional pollen is formed, crossings are impossible, and spontaneous hybrids, which make determination difficult in other apomictic genera, such as Rubus or Pilosella, are missing in Germany. Each plant can therefore be assigned to a apomictic micro-species.
So far, there is no reasonable explanation how the large number of groups originated in the Eurasian mountains.
The taxonomy of some species complexes has not yet been fully elucidated. In Germany this concerns: A. fallax, A. obscura, A. othmarii, A. sericoneura and A. versipila.
About 150 species are known from Central Europe. The description of new species is to be expected. Fröhner (1990) assigned the species occurring in Germany to three “basic” sections and nine "secondary" sections, the latter distinguished by a combination of characters of the basic sections. Basic sections are: (1) Alchemilla sect. Erectae S.E. Fröhner (tall, herbaceous perennials, stipules adnate to petiole, calyx and epicalyx long compared to hypanthium, only two naturalized species); (2) Alchemilla sect. Ultravulgaris S.E. Fröhner (medium-sized perennials, stipules free, calyx shorter than hypanthium and epicalyx even shorter than calyx, many species); (3) Alchemilla sect. Alpinae Camus (small, dwarf shrubs with silky pubescence, stipules adnate to petiole, epicalyx short compared to hypanthium, few species). A representative of Alchemilla sect. Pentaphylleae Camus (semi-shrubs or herbaceous perennials, stipules hardly adnate to petiole, leave blades very coarse-toothed, epicalyx small to absent) is presumably parent of hybridogenic species found in Germany (Fröhner 1990). Older classification approaches, according to which only three sections are distinguished in Germany, are presented by Sebald (1992).
The majority of Alchemilla species colonize open, nutrient-rich habitats. Only the species of section Alpinae and Glaciales colonize rocks in alpine habitats. Here, alpine pastures are the most important habitats above the tree line. Otherwise, lady's mantle grows mainly in grassland that is well-supplied with nutrients, but not in heavily fertilized multi-crop grassland. Other important habitats are roadsides. Only a few species, such as A. glaucescens, can be found on nutrient-poor grasslands.
The spread of lady's mantle has been strongly facilitated by grassland farming and the expansion of traffic routes. Many Alchemilla species are typical apophytes (native species that also colonize man-made sites). Lady's mantle is often displaced by running waters, by this many montane species can establish occurrences in lowlands. In addition, Alchemilla species are spread easily through cattle feed or soil material, but also through garden wastes. The ornamental plant A. mollis which is native to southeastern Europe and the Near East, has already acquired a large synanthropic area. The naturalization of further species is to be expected. Hügin (2006: 51) reports a still undescribed species, probably native to the Caucasus, in the northern Black Forest.
Members of the genus are deciduous dwarf shrubs or perennials with above-ground, horizontal shoots. Branching is sympodial. Plants are rhizomatous, often forming woody stocks. Shoots contain pith. Basal leaves are arranged in a rosette, petioled, blades are lobed to palmately compound with dentate margins. In the bud, each lobe of the blade is individually folded. The stipules are membranous, relatively large and function as bud scales. They are adnate in at least halfway along their length to the petiole. The connation of both stipules opposite the petiole is incomplete. This incision is called “ochrea incision" and constitutes an important taxonomic character, which is, however, difficult to recognize in herbarium material. In addition, both stipule auricles may be connate above the petiole, this trait is also important for species identification but difficult to detect in herbarium material. The leaves bear hydathodes at the margin of lobes, from which water is exuded at night (guttation). Stem leaves resemble basal leaves, but petioles are shorter and blades are smaller, with their size decreasing upwards. Inflorescences develop from the leaf axils, usually from leaves of previous years, as closed thyrsus (composed of monopodial basal structure consisting of 3-10 units and the actual sympodial inflorescences consisting of 2-7 cymes). Single inflorescences are usually composed of a few, verticillate monochasia. The flowers are small, with 4 sepals of green or yellowish color arising from a bell- or jug-shaped hypanthium (terminal flower also pentamerous). From the adaxial tips of sepals arises a woolly tuft of hair closing the bud. An epicalyx is almost always present. Petals are missing. The flowers have a distinct discus which secrets nectar. The stamens represent transformed petals, inserted outside the discus alternating with the sepals. The pollen grains are predominantly deformed, only a few high-altitude alpine species missing in Germany have fertile pollen. The flower has one, rarely two carpels with an upright style and a lenticular to capitate, rarely hooked stigma. Nutlets are completely or partially enclosed by hypanthium and are dispersed together with it.
The determination of Alchemilla plants is (very) difficult. On the one hand, the species are similar and show considerable intra-specific variability, so that determination on the basis of certain characters such as number of teeth or indumentum is only possible in exceptional cases. On the other hand, atypical, mostly indeterminable specimens form after browsing or mowing, as well as in autumn. Well-developed plants at the beginning of seed ripening are suitable for identification. When collecting, care must be taken to ensure that coherent plants are herbarized, otherwise there is a high risk of creating mixed specimens. Tools for digging are required for collecting instructive specimens.
Since lady's mantle can often only be determined by a combination of different characters, an extensive familiarization is necessary in order to be able to evaluate the variability of characters accordingly. In addition, characters that are hardly suited for identification keys are of great importance for the determination, such as the green shade of the leaves and the leaf folding. These characters allow determination in the field, but are hardly useful for herbarium specimens. Hügin (2006) published exquisite color illustrations of photos by K. Rasbach, which give a realistic idea of the habit of many species.
Leaf characters usually refer to basal leaves, exceptions are indicated. Characteristics of the stipule auricles are not used, as they are difficult to recognize from herbarium specimens. In several alpine species, the indumentum changes during the vegetation period. It should always be checked whether the hairiness is uniform across season or whether spring leaves have significantly less hairs or are even glabrous.
Unfortunately, hairiness of calyxes and pedicels within an inflorescence, which is important for the determination, is often not uniform. The first flower of an inflorescence that stands between two partial inflorescences is the largest and usually also the one with the densest indumentum of an inflorescence.
There are several modern identification keys available: by S. Fröhner in "Hegi" (Fröhner 1990) and in "Rothmaler" (Fröhner 2011), by F. Grims in the Austrian Excursion Flora (Grims 2005) as well as by W. Lippert in the "Oberdorfer" (Lippert 2001). Determination is nevertheless difficult because characters are often difficult to describe and do often overlap. In many cases, it is not possible to determine herbarium specimens with certainty. Anyone wishing to follow up with the genus cannot avoid the guidance of reference cultivations, whereby the cultivation of alpine species requires considerable horticultural skills. It should also be mentioned that the morphology of many species can change untypically, especially under unsuitable cultivation conditions.
Typification and thus the use of the name Alchemilla vulgaris L. is associated with difficulties. Fröhner (1986) chose a specimen corresponding to Alchemilla acutiloba Opiz as lectotype, which was not the basis for the description of Linné (Jarvis 2007). Original material of Linné (specimen 166.1 in the London Linné Herbarium), which was then selected as type by Purohit & Panigrahi (1991), represents a mixture of probably Alchemilla monticola Opiz and another species, possibly A. glaucescens (Mitt. H. Kalheber). Therefore, the name A. vulgaris is considered by many authors as nomen ambiguum, and its use should be avoided.
Species numbers are strongly correlated with elevation. Thus, in the North German lowlands where only a few species occur, identification is comparably easy. However, these Alchemilla species are rare in this area. But already in the colline and montane regions the number of species considerably increases, and species diversity of the German low mountain ranges is probably not yet completely known. Hügin (2006) was able to significantly extend the knowledge about the distribution of lady's mantle in the Black Forest and the Swabian Alb. Kalheber (1982) clarified the occurrence of Alchemilla propinqua, A. connivens and A. glomerulans. It is still unclear whether plants from the Thuringian Forest that were recognized by Fröhner (2011) as Alchemilla impexa belong to this taxon.
The situation in the Alps, where Alchemilla reaches the highest number of species, is unclear. Optimal habitats are moist, moderately nitrogen-rich pastures in cold-air zones; several species grow in snowbeds. Uncertainties still exist, although Lippert & Merxmüller (1974-1982) have intensively investigated the occurrence of lady's mantle in the German Alpine region. Some of the determinations of this work proved to be erroneous. Within the framework of the Alpine biotope mapping (Urban & Meyer 1992, 2006, 2008), the distribution area could be significantly extended for some species, and Alchemilla fallax was newly recorded for Germany.
The genus name goes back to the word alchemy. The water produced by guttation was used for magic experiments.
Our thanks go to Sigurd E. Fröhner (Dresden) and Heinz Kalheber (Runkel) for the provision and verification of specimens. Botanical State Collection of Munich (M) provided a substantial selection of suitable specimens.
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