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22Новости систематики высших растений Novitates Systematicae Plantarum Vascutarium
2020
51:26-38
ISSN 0568-5443 (print) ISSN 2687-1564 (onLine)
New delimitation of two series within Alchemilla subsection Calycanthum and new subdivision within Alchemilla series Calycinae (Rosaceae)
Новое разграничение рядов в подсекции Calycanthum и новое разделение ряда Calycinae рода Alchemilla (Rosaceae)
A. V. Chkalov
Lobachevsky State University of Nizhny Novgorod Gagarina Ave., 23, Nizhny Novgorod, 603950, Russia [email protected]
https://doi.org/10.31111/novitates/2020.51.26
А. В. Чкалов
Национальный исследовательский Нижегородский государственный университет им. Н. И. Лобачевского пр. Гагарина, 23, Нижний Новгород, 603950, Россия [email protected]
Abstract. The distinctions between Alchemilla ser. Calycinae and A. ser. Elatae are emended, treating the latter group as relict, connected to the Tertiary broadleaved forests of the eastern Mediterranean Basin. Alchemilla ser. Calycinae (in renewed circumscription) is subdivided into seven provisional aggregates according to the specially developed coordinate system (with two axes — a ratio of central zone width to leaf length vs. number of leaf teeth in total). The original key included all the species of the series occurring in the Caucasus and Asia Minor was compiled.
Keywords: aggregates, apomicts, Asia Minor, Caucasus, coordinate system.
Аннотация. Предложено иное разграничение между двумя рядами (Calycinae и Elatae) рода Alchemilla, рассматривающее вторую группу как реликтовую, приуроченную к третичным широколиственным лесам восточной части Средиземноморского бассейна. Ряд Calycinae (в новых границах) подразделен на семь предварительно выделенных агрегатов, руководствуясь оригинальной системой координат (с двумя осями — отношение ширины центральной зоны к длине листа и суммарное количество зубцов листа). Составлен ключ, охватывающий все виды ряда из Кавказа и Малой Азии.
Ключевые слова: агрегаты, апомикты, Кавказ, Малая Азия, система координат.
After morphological bordering of a Calycinae group (Buser, 1893, 1896) of the genus Alchemilla L., many species were described and different conceptions were presented to organise this diversity into a kind of a system.
Alchemilla subsect. Calycanthum (Rothm.) Rothm. ser. Calycinae (Buser) Rothm. is a peculiar group within the genus having very distinct geographical, but not morphological, borders. There are different ways to divide the Calycanthum group. The first, proposed by Rothmaler (1933, 1938): to A. ser. Calycinae, to which only small plants with coriaceous leaves, appressed pubescence on petioles and stems, mostly glabrous leaves and ever glabrous flowers would be placed. All other species with another kind of appearance, with petioles and stems pubescent in different types, glabrous flowers or, alternatively, densely hairy ones, were supposed to be arranged in the group A. ser. Elatae (Rothm.) Rothm. But even with such a narrow concept of A. ser. Calycinae, some misconceptions appeared (Rothmaler,
Поступила в редакцию | Submitted: 08.07.2020
1938): A. venosa Juz. was placed here, despite the fact that it had hairy hypanthia and leaves beneath; similarly, so was A. wischniewskii Rothm., which had densely pubescent leaves beneath and hairy stems throughout, with multilobed leaves (the author noted as the closest to this species A. subsplendens Buser, attributed by himself to A. ser. Elatae).
The second concept was proposed by Juzepczuk (1941). He divided the Calycanthum group (using the name "Calycinae") according to an analogy with subdivision of A. subsect. Alchemilla (also known as "Vulgares"): 1) a group with patent indumentum and differently hairy flowers ("Vulgares: Hirsutae and Imberbes"); 2) a group with dense appressed or sub-appressed indumentum of leaves, petioles, and stems, densely hairy hypanthia ("Pubescentes"); 3) a group with appressed sparse indumentum of stems and petioles, mostly glabrous leaves, mostly glabrous flowers ("Subglabrae Obtusae"); i. e. it meant that A. subsect. Calycanthum should be divided into "cycles"
Принята к публикации | Accepted: 18.11.2020
Oxysepalae, Subsplendentes, and Durae, respectively (all the names invalidly published). The latter group was close in concept to A. ser. Calycinae sensu Rothm., therefore the other two were together approximately equal to A. ser. Elatae. However, during practical usage there were many exceptions to such strict rules. For example, at the border between "Oxysepalae" and "Durae" there are A. tredecimloba Buser and A. undecimloba Juz. (subglabrous, appressed pubescent as "Durae", but large and robust, with multitoothed leaf blade and variably hairy hypanthia as "Oxysepalae"). A. procerrima S. E. Frohner, A. ellenbergiana Rothm., and A. wischniewskii Rothm. have quite dense pubescence on the stem internodes and on the leaves like "Subsplendentes", but glabrous flowers like "Durae". There are many other similar examples.
Very different, non-hierarchical approach was found in the viewpoints of Frohner (1986, 1990). Here, the most important points about Calycinae are as follows:
1. A non-hierarchical system of sections, which reflected a reticulate hybridogenous formation of the genus. It was described as follows: four parental sections (A. sect. Alpinae = A, A. sect. Pentaphylleae = P, A. sect. Ultravulgares = U, A. sect. Erectae = E) stated as basal, the other were formed due to hybridization between these four and their progenies, e. g. PA-, UP-, EU-, EUP-hybrids, and so on. Every of 14 "parental" and "hybrid" groups were endowed with a separate position and equal taxonomic rank of a section.
2. Based on morphology, the author narrowed the circumscriptions of two groups, A. sect. Erectae (A. ser. Elatae Rothm., p. min. p.) and A. sect. Calycinae (A. ser. Calycinae sensu Rothm., p. p.).
3. A. sect. Calycinae was considered EP-hybrid, i. e. a relative of Frohner's A. sect. Coriacea = EUP (A. ser. Subglabrae Pawl., p. p.). Moreover, some species of A. ser. Calycinae sensu Rothm. (A. incisa Buser, A. oth-marii Buser, A. firma Buser) were shifted by the author to A. sect. Coriacea.
To discuss this system substantially, I need to elucidate some important traits of Calycinae (sensu Rothmaler):
1. Typical morphology is: (a) plants of a small size with coriaceous leaves (up to 10-30 cm); (b) leaf blades mostly glabrous (sometimes sparsely hairy beneath); (c) leaf blade (of medium leaves) dissected to 2/5-y2 (central zone 50-60%), with 60-110 teeth in total, 5-6(7) at each side of leaf lobe; (d) indumentum of petioles and stems is poor containing appressed subtle hairs, stems pubescent only on the lower inter-nodes, hairs gradually becoming rare and disappearing in the lower half of stems; (e) flowers with short obconical hypanthia, with almost uniform, approxi-
mately equal (lanceolate, ovate or elongate triangular) sepals and epicalyx segments, longer than hypanthia; glabrous.
2. Ecologically, it is attached to evolutionarily pioneer habitats with low competition: stony substrates, stony slopes usually near a snowline, secondary synan-thropic habitats like roadsides; from dry to over-moist, such as those found along springs or riversides, in open or shadowed habitats, and on sour and alkaline soils (Frohner, 1986, 1990).
3. Biogeographically, it is one of the most clearly-bordered Alchemilla groups. In comparison to A. ser. Alchemilla or A. ser. Pubescentes, or A. ser. Subglabrae, which are widely distributed in temperate Eurasia, it is connected distinctly with the mountain systems of the Mediterranean basin. The highest species richness (in the circumscription sensu Rothmaler) is observed in the Caucasus (15), while only seven, three, and five species occur in the Alps (Atlas..., 2009), Pyrenees (Frohner, 1998), and Carpathians (Pawlowski, 1955; Sytschak, 2011) respectively. Thus, the formation center of Ca-lycinae group is undoubtedly the Caucasus, as indicated by its high morphological diversity and irradiation there (see also below).
Discussing Frohner's system as a whole, I tested Frohner's hypothesis that the Calycinae-group was an EP-hybrid, i. e. it appeared after hybridization between members of A. sect. Erectae (E) and A. pentaphyllea L. — the only member of A. sect. Pentaphylleae (P).
1. A. pentaphyllea has extremely short epicalyx segments (55% of sepal length) and elongate hypanthium. The mean value of those features for A. sect. Erectae and A. sect. Pentaphylleae would be much higher than those of A. ser. Calycinae.
2. None of the parent strains have an appressed indumentum with subtle hairs of petioles and stems (A. pentaphyllea is subappressed hairy with bristly hairs), but it is the key peculiarity of Calycinae group.
3. "Parental" A. pentaphyllea was distributed only in the Alps and there was no obvious reason for its extinction anywhere else. But the diversity centers of Calyci-nae group are located far to the east, in the Caucasus, and, in the sense narrowed by Frohner, it counts in Europe even less species than mentioned above.
4. Ecologically, A. pentaphyllea is a highland species and grows on peaty or sandy overmoist soils often beside a snowline (Frohner, 1990).
Taking geography into consideration, I conclude that it is a neoendemic species attached to evo-lutionarily young widespread habitats that did not have enough time to distribute more widely (for more discussion on this topic see Chkalov, Vorotnikov, 2009). Thus, despite its primitive morphological traits, I can
hardly regard A. pentaphyllea as an ancient parental line and ancestral group for the Calycinae.
The list of weak items of this system may be continued:
1. There are similar circumstances (to above-mentioned Pentaphylleae) with Alpinae, i. e. at least two of "parental lines".
2. If they really were ancient and parental to all Eurasian Alchemilla, as it was supposed by the author, they or their relatives would be widespread to the Central Asia to establish Alchemilla there (to mention, there were no ecological barriers against it). Since they had restricted distribution, Central-Asian Alchemilla would be formed independently of those European "parents".
3. There are two groups of A. ser. Alchemilla, A. aggr. retropilosa Juz. and A. aggr. semilunaris Alechin, which are very numerous in Central Asia and Eastern Europe. In Central Europe, however, the former is presented by only a few species located at the western limits of their distribution areas, and the latter is not yet found. Moreover, a morphometric study showed (Sepp, Paal, 2001) that A. semilunaris Alechin fell out of the pool of other European species, which were supposed close relatives to each other, as well as A. subglobosa C. G. Westerlund (A. aggr. retropilosa).
4. In this isolated system of sections, a place of African Alchemilla is not transparent, neither their relations with closely related genera (Aphanes, Lachemilla). The high rank of the infrageneric taxa (i. e. section) chosen by the author does not correspond to the grade of morphological and biogeographical differences between groups.
Thus, it is highly likely that this system reflects thoroughly circumstances of Alchemilla irradiation in the Alps, maybe in the Western Europe concerning the formation of the endemic groups, their hybridization, etc. In such cases, those groups may be considered as parental, and the principles of Frohner's system work in the destined way. When the most dominated taxonomic groups of Quaternary endemics are different, as in the Caucasus or Central Asia, one must analyse Alchemilla diversity independently in each case.
The aim of this study was to discuss existing systems of the Calycinae group to estimate their attribution to Caucasian Alchemilla and to modify them, if necessary.
Materials and methods
The material of Caucasian Alchemilla in LE and MW was investigated by the author. Many samples were obtained through on-line platforms: B (Herbarium B, 2020); C, M (JSTOR, 2020); E (Herbarium E, 2020); FI (Herbarium FI, 2020); G (Herbarium G, 2020); H
(FinBIF, 2020); K (Herbarium K, 2020); MW (Seregin, 2020); P (Herbarium P, 2020); S (Herbarium S, 2020); W, WU, JE (JACQ, 2020). Calculation of values for two features (i. e. grade of leaf dissection as a ratio of central zone value to leaf length; number of radical leaf teeth in total) of either medium or upper radical leaves, or both (for those that present usually) for species of A. ser. Calycinae was fulfilled. The terminology used in the keys and for the calculation was previously described (Chkalov, 2011, 2015). Zoning of the Caucasus and relevant abbreviations were presented in "Caucasian Flora Conspectus" (Caucasian..., 2009).
Results and discussion
The discussion is partly speculative, but still we need to organize this material to a system; thus, even a preliminary and hypothetic trial is welcome. Primary prepositions for the above-mentioned points are such: for the first item, "no morphology without geography" — a geographical morphological approach according to A. De Candolle, R. Wettstein, V. L. Komarov (Kamelin, 1973); therefore, hierarchical division has a sense, when a morphological group is outlined geographically. Following this preposition, I accept that by now it is hardly possible to subdivide those series hierarchically, although they could be bordered more accurately by this approach.
Also, I use some basic prepositions to arrange the group: (1) this group formed in a hybridogenous way and thus represents reticulate system; (2) I expect that hybridogenous progenies must possess some intermediate traits relatively to parental strains (medium for continual features, or those discrete from the only one parent); (3) the same applies to ecological traits; (4) the features to arrange this group must satisfy some requirements, such as: (a) they should have an evolutionary prospect (i. e. they should have clearly changed in comparison with ancient and more recent groups); (b) the features, bordered geographically or ecologically, or both, are favorable; and (c) they must be easy to obtain and study (e. g. checked and used in casual practice, with in sicco specimens).
How was it formed?
From the information on A. ser. Calycinae outlined above, I conclude that there is no need to look for an ancient parental group, because the progeny is relatively young. It is easy to identify one parent that possesses trait-complex consisting of coriaceous glabrous leaves — poor appressed indumentum — similar long sepals and epicalyx segments: A. ser. Subglabrae. The remarkably similar flower structure with A. ser. Calycinae suggests the above-mentioned
shifts of the species from one group to another. Frohner (1986) also mentioned such an affinity between the two groups (see above).
The second parent is harder to identify, but the key point may be the geography described above. While the Subglabrae group is widely distributed (temperate areas of Eurasia almost throughout), the limited distribution of A. ser. Calycinae may be explained due to restricted distribution of its second parent, possibly A. subsect. Chirophyllum Rothm. (specifically, two groups within it — A. ser. Saxatiles Buser and A. ser. Sericeae Buser) or A. subsect. Alchemilla ser. Pubescentes Buser (partially — affinity of A. sericata Rchb.). A. subsect. Chirophyllum could be the second parent because of deeply dissected leaves and appressed indumentum of some species, flowers with quite short hypanthia. However, their epicalyx segments are too short, teeth on the leaves of Caucasian species are too straight, and apical teeth are about the length of neighboring ones; so that hypothesis cannot be adopted. A. aggr. sericata Rchb. species are the most suitable for the role of parent for A. ser. Calycinae for the following reasons:
1. Morphology: (a) plants mostly of small size; (b) some species have glabrous leaf surfaces (A. acropsila Rothm.), which demonstrates an evolutionary opportunity, despite the fact that they are mostly densely pubescent; (c) leaf blade dissected to 2/5-1/2 (central zone 50-60%), with 50-90 teeth in total, 5-7 at each side of leaf lobe; leaf structure of some species (e. g. A. bombycina Rothm.) is similar to that of A. retinervis Buser; (d) indumentum of stems and petioles is appressed; (e) epicalyx segments are often as long as sepals or nearly so. With an intermixture of Subglabrae traits, the formation of A. ser. Calycinae is quite believable.
2. Ecologically, it is similar to the Calycinae group, growing in habitats with low competition: on stony substrates, slopes, usually dry, and in secondary synanthropic habitats like roadsides. With the addition of eco-traits of Subglabrae group, which is the most vigorous and abundant at high latitudes and in high mountain areas, an ecological specificity of A. ser. Calycinae becomes quite reasonable.
3. Geography. The most suitable species of the A. sericata aggregate are endemic to the Caucasus; thus, it fits into the geography of the Calycinae group mentioned above. Local endemics of this group outside the Caucasus should be discussed thoroughly; it may be the case that they might be a consequence of polytopous speciation (i. e. repeated hybridogenous formation of the similar Calycinae species) or might be a subsequent progeny of some widely distributed Calycinae species.
How to arrange this group?
I have accepted quite limited applying of the hierarchical system within A. subsect. Calycanthum, and still believe that it is possible to arrange these species into a better system.
Some ideas on opportunity of taxonomic groups arrangement through a construction of periodical (coordinate) system were developed by Lyubischev (1982). The most promising way to achieve this is to use a coordinate system applying the features fit to the above-mentioned requirements. The first feature is grade of leaf dissection as a ratio of central zone value to leaf length because it has clear evolutionary prospects, from wholly dissected (compound) leaves (e. g. Potentilla palustris (L.) Scop., some species of the genus Lachemilla (Focke) Rydb. and Alchemilla subsect. Chirophyllum, A. pentaphyllea), to the almost lobeless leaves of A. ser. Elatae s. str. and A. ser. Alchemilla. The second feature is total number of leaf teeth, as its evolutionary prospect is similarly wide: fewer than 10 teeth in Aphanes L. and some Lachemilla, the 20-30 teeth of some A. subsect. Chirophyllum and A. pentaphyllea to the 220 teeth of some A. ser. Elatae s. str. and A. ser. Alchemilla.
I conclude that this approach permits the combination of species into groups with similar appearance and, thus, provide a more holistic analysis of the species similarity. I have achieved such a coordinate system (Fig.), by which several more or less clearly outlined groups have been qualified as preliminary aggregates.
Alchemilla aggr. retinervis Buser (Re).
A. adelodictya Juz., A. ancerensis Kalheber, A. dey-lii Plocek ex Sojak, A. fissa Günther et Schummel, A. hayirliogluorum Kalheber, A. microdictya Juz., A. retinervis Buser, A. retinerviformis Juz., A. rivularis Ponert, A. tiryalensis Pawl.
Geography: Asia Minor and Caucasus (WC: Bel.-Lab., Urup-Teb.; CC: Malk., U. Ter.; EC: Ass.-Arg., U. Sulak., Man.-Samur., Kubin.; WTC: Tuap.-Adl., Abkh., Ing.-Rion., Adzh.; CTC: Kart.-S. Oss.; ETC: Alaz.-Agrich., Murg.-Murovd.; SWTC: Meskh., Dzhav.-U. Akh., Arag.; STC: Sevan, Nakh., Zang., S. Karab.) (8), Balkans (1), Carpathians (2), Alps (1), Pyrenees (1).
Here is the most typical group, with the morphological features described above. However, there is an exception: four species (A. ancerensis, A. hayiriogluo-rum, A. retinerviformis, A. tiryalensis) are strongly pubescent (in most parts, including stems and flowers, but not on the upper surface of leaves), while still having the typical flower structure and the appearance. Logically, it is likely that the latter are also a
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consequence of hybridization between Subglabrae and Seri-catae groups, but have predominantly Sericatae traits. There are many mesoxerophi-lic and xerophilic Alchemilla species in the Caucasus that can compete with this Re-species; thus, such progenies cannot be especially numerous and abundant. Once, other Re-species possessed mostly Sublabrae-trait-complex, and were more successful due to their ecological peculiarity and as a consequence of low competition in the absence of ecologically similar species. That is why I did not separate the four species mentioned above, considering all the species of A. aggr. retinervis are a consequence of the same process. I conclude with confidence that Asia Minor and the Caucasus were a formation center for this most typical Calycinae-group. Only one of widespread European species (A. fissa) belongs here. Likely, it migrated from the primary speciation center and became an ancestor of the European Calycinae-representatives.
Moreover, the inconsistency of the indumentum character of flowers and pedicels is typical in A. subsect. Ca-lycanthum and all its groups (similar variation has also been identified for A. ser. Ela-tae s. str.), and is observed in most aggregates listed below (see Table).
All following groups differ from Re by more numerous teeth of leaf blades. Their hairiness can vary widely, as it does so in the Re-group, but special attention should be paid to an inconsistency between glabrous flowers and densely hairy
v. 1 g
leaves and stems, or vice versa
Alchemilla aggr. betuletorum Rothm. (Be).
A. barbatiflora Juz., A. betuletorum Rothm., A. beyazoglui Kalheber, A. eugenii Pawl., A. procerrima S. E. Fröhner, A. sevangensis Juz. (A. ikizdereensis Kalheber).
Geography: Asia Minor and Caucasus (WC: Bel.-Lab., Urup-Teb.; CC: U. Kum., Malk., U. Ter.; EC: U. Sulak.; WTC: Tuap.-Adl., Abkh., Ing.-Rion., Rion.-Kvir., Adzh.; CTC: Kart.-S. Oss.; ETC: Alaz.-Agrich., Murg.-Murovd.; SWTC: Meskh., Arag.; STC: Sevan.) (4), Iran (1), Carpathians (1).
Here, I focus on the traits most divergent from typical Re-group. These include wide central zone, short lobes (mostly arcuate), stems more densely and higher pubescent, and leaves ± pubescent beneath, hypanthia sometimes hairy (A. barbatiflora, A. sevangensis). As a second parental group, based on these features, I assume members of A. ser. Subglabrae subser. Appressipilae Juz. ex V. N. Tikhom., the affinity of A. obtegens Juz. in particular.
Alchemilla aggr. ellenbergiana Rothm. (El).
A. akdoganica Kalheber, A. bezengiensis Czkalov ined., A. debilis Juz., A. ellenbergiana Rothm., A. oritur-cica Pawl., A. stricta Rothm.
Geography: Asia Minor and Caucasus (WC: Bel.-Lab., Urup-Teb.; CC: Malk., U. Ter.; EC: U. Sulak., Man.-Samur., Kubin.; WTC: Tuap.-Adl., Abkh.; CTC: Kart.-S. Oss., Trial.-L. Kart.; ETC: Alaz.-Agrich., Murg.-Murovd.; SWTC: Meskh., Dzhav.-U. Akh.; STC: Erev., Sevan, Nakh., S. Karab.) (6).
The traits most divergent from those of Re-group are short wide teeth, 6-8 at each side of leaf lobe, lobes rounded at the top, usually semicircular or nearly so, stems more densely and upper pubescent, and leaves ± pubescent beneath (except for A. debilis), hypanthia variously pubescent. Such features — the grade of leaf dissection, larger apical teeth, and the appearance of leaves — point to A. ser. Alchemilla, A. aggr. retropilosa Juz. (for example, A. compactilis Juz. and A. dzhavakhe-tica Juz.) as a probable second parental group.
Alchemilla aggr. dura Buser (Du).
A. dura Buser, A. fallax Buser, A. firma Buser, A. kazbekensis Czkalov ined., A. sericoneura Buser, A. venosula Buser.
Geography: Caucasus (WC: Bel.-Lab., Urup-Teb.; CC: Malk., U. Ter.; EC: U. Sulak., Man.-Samur., Kubin.; WTC: Tuap.-Adl., Abkh., Ing.-Rion., Rion.-Kvir., Adzh.; CTC: Kart.-S. Oss.; ETC: Alaz.-Agrich., Murg.-Murovd.; SWTC: Dzhav.-U. Akh., Arag.; STC: Sevan, Nakh., Megr.-Zan., S. Karab.) (2), Balkans (2), Alps (4), Pyrenees (1).
This is a very interesting group, and mostly European. Likely, there was a parental species of A. ser.
Table. Traits of the Alchemilla series Calycinae aggregates
Traits Aggregates
Re Be El Du Tr Su Ab
Total teeth number in radical leaves 60-82 50-80(90)* [25%] (80)90-130 [27%] (70)80-110 [30%] (80)90-125 [33%] (80)100-130 [30%] (95)105-135 [19%] (100)125-210 [33%]
Central zone to leaf length ratio, % 50-75(79) 52-74* [19%] (72)77-86 [10%] (67)70-78 [10%] 59-75 [18%] 56-73 [20%] (60)64-73 [15%] (61)64-88 [25%]
Angle of leaf sector, grades 42-51(60) 52-74* 33-46 32-47(55) 35-47(55) 40-54(60) 36-48(60) 20-39(43)
Leaf lobe length to leaf length ratio, % 19-34 16-34(42)* 11-25(27) 17-25(30) (10)16-34 22-39 20-36 12-31
Incision depth to standard tooth length ratio 1-3(4) 1-3* 0.5-3 0.5-1.5 1.5-2(3) 0.5-1.5 1.5-3 0.5-4
Standard tooth length to leaf length ratio, % (6)9-16 9-20* 4.5-9 3-10 3-9(13) 5.513 3-10.5 1.5-5(7.5)
Apical tooth length to neighboring teeth length ratio (0.2)0.4-1.1 0.2-0.7* 0.2-0.7(0.9) 0.5-0.8 (0.2)0.4-0.8(1.2) 0.2-0.5(0.7) 0.5-1 0.3-1
Indumentum density Leaves beneath G Ge Ge* S* S D Ge(r) D Ge Ge D(r) Ge D(r) D Ge(r) Ge D S
Hypanthia G D* GS* G GS D GS G G G GS D(r) D GS D GS G
Pedicels G G* GS* G D(r) G G G D(r) D GS G(r) G GS(r)
Stem G S D* D S D S(r) G(r) S D D S D S S D(r) G(r)
Stem indumentum height 0-1/3-1/2 У2*-2/3*-1* 1/-2/-3/-1 '2 '3 '4 У-1/-2/- '3 2 '3 3-4-1 У-1/-2/-3/-! 3234 0(r) y y 1 У-1 У-У-У-2/- ' 4 '3 '2 '3 34-1
Note. Bold borders — the most differentiative traits; * — trait values for the pubescent species of the Re-aggregate; in square brackets — the largest of (max-min)x100/max value of a trait. Indumentum density: G — glabrous; Ge — glabrous leaves with hairy main veins and basal lobes beneath; S — sparsely hairy; D — densely hairy; GS — glabrous and sparsely hairy hypanthia/pedicels in the same plant. Stem indumentum height: 0 — stem glabrous; V ... — stem hairy to the V ... 3/4 of its length; 1 — stem hairy throughout. (r) — rarely detected value. See the text for the aggregate abbreviations.
Calycinae (e. g. A. fissa), and (an)other parent(s) from a group that should be identified. Here, there are orbicular, more rarely orbicular reniform, leaves (angle of sector mostly 40-55°) with a central zone of medium width (near the mean value of Re-group), lobes with large acute teeth. Most of these species look like a representative of the Re-group that acquired more teeth. Moreover, they are similar in appearance to some representatives of A. ser. Subglabrae subser. Denudatae V. N. Tikhom. (e. g., Caucasian A. pseudocartalinica Juz., A. pilicincta Buser (= A. cartalinica Juz.), A. erectilis Juz.; European A. reniformis Buser, A. effusa Buser, A. impexa Buser etc.). I consider this to be a second parental group.
Alchemilla aggr. transcaucasica Rothm. (Tr).
A. grandidens Juz., A. indurata Juz., A. muldaschevii Czkalov ined., A. pseudotranscaucasica Czkalov ined., A. ptyschensis Czkalov ined., A. transcaucasica Rothm.
Geography: Asia Minor and Caucasus (WC: Bel.-Lab., Urup-Teb.; CC: U. Kum., Malk., U. Ter.; EC: Ass.-Arg., U. Sulak., Man.-Samur.; WTC: Tuap.-Adl., Abkh., Ing.-Rion., Rion.-Kvir., Adzh.; CTC: Kart.-S. Oss., Trial.-L. Kart.; ETC: Alaz.-Agrich.; SWTC: Meskh., Dzhav.-U. Akh.; STC: Sevan, Nakh., S. Karab.) (6).
These are very similar in appearance to the Regroup, but differ in having longer lobes with short incisions between, and more numerous, but similarly large curved wide teeth. It likely formed through admixture of some traits of A. ser. Alchemilla subser. Pastorales V. N. Tikhom. or A. ser. Alchemilla subser. Alche-milla (when the species is poorly pubescent in some parts). The more pubescent species (e. g. A. grandidens, A. pseudotranscaucasica) may be progenies of Pastorales group, the others would be closer to A. subser. Alchemilla, the affinity of A. subcrenatiformis Juz. (because of the teeth and leaf form). These are second parental groups.
Alchemilla aggr. subsplendens Buser (Su).
A. cartilaginea Rothm., A. divaricans Buser, A. speciosa Buser, A. subsplendens Buser, A. venosa Juz.
Geography: Asia Minor and Caucasus (WC: Bel.-Lab.; CC: U. Kum., Malk., U. Ter.; EC: Ass.-Arg.; WTC: Tuap.-Adl., Abkh., Ing.-Rion., Rion.-Kvir.; CTC: Kart.-S. Oss.; ETC: Alaz.-Agrich., Murg.-Murovd.; STC: Sevan, Nakh., S. Karab.) (5).
Very pubescent with appressed silky hairs in most parts including pedicels and hypanthia plants, with deep leaf dissection. I suspected its parent was the Seri-catae group, but the many teeth of the leaf blade forced me to look it in the opposite direction. The high teeth number and abundant indumentum are only found from among A. ser. Elatae (a second parental group), such as
A. hirtipedicellata Juz., A. orthotricha Rothm., A. holotri-cha Juz., A. ziganadagensis Pawl. etc. This group is very close to Juzepczuk's "cycle" Subsplendentes, and might be a separate subseries.
Alchemilla aggr. abchasica Buser (Ab).
A. abchasica Buser (A. ayderensis Kalheber), A. buseriana Rothm., A. camptopoda Juz., A. ciminensis Pawl., A. gorcensis Pawl., A. longipedicellata Czkalov ined., A. sciadiophylla Rothm., A. sericoneuroides Pawl., A. tredecimloba Buser, A. undecimloba Juz.
Geography: Asia Minor and Caucasus (WC: Bel.-Lab., Urup-Teb.; CC: U. Kum., Malk., U. Ter.; WTC: Tuap.-Adl., Abkh., Ing.-Rion., Adzh.; CTC: Kart.-S. Oss.; ETC: Alaz.-Agrich., Murg.-Murovd.; SWTC: Arag.; STC: Sevan, S. Karab.) (8), Crimea (1), Carpathians (2), Balkans (1).
As the previous one, the latter group is a progeny of the Elatae group, but the teeth number is dramatically large. That is obviously heterogeneous, because (a) A. ser. Elatae s. str. is variable itself. Here, among the progenies, there are mostly glabrous plants (A. buseri-ana, A. gorcensis, A. tredecimloba, A. undecimloba), some with ± hairy hypanthia; other species (A. longipedicella-ta, A. sciadiophylla, A. ciminensis) have hairy leaves beneath, pubescent stems and hypanthia; some are strongly pubescent, sometimes with almost erectopatent hairs (A. abchasica, A. camptopoda); (b) there is a geographical heterogeneity as consequence of secondary irradiation in other speciation centers (some local Calycinae endemics might have formed by a widespread ancestor of Caucasian origin or by their aboriginal relatives), or this indicates polytopous speciation (i. e. analogous hybridization between some species of Sericatae and Sub-glabrae groups might have taken place). Thus, revision of the latter group as a whole is still needed. As well, if further geographical and morphological specificity is revealed, further hierarchical subdivision will be possible.
The last two aggregates are at the border between series, because of a natural hybridization process, and produce most difficulties in their delimitation. Their geographical and ecological features forced me to shift representatives of these aggregates to A. ser. Calycinae.
Also, there is A. incisa, with a position isolated from all other species. Treating this species as a part of the Calycinae group, I believe its position points to a separate paraphyletic formation, which includes the rest of the species of the probable corresponding aggregate.
Therefore, the system of A. ser. Calycinae, including all of its European representatives, is necessary, but careful attention must be paid to all its representatives. Geography of the aggregates listed above was outlined rather superficially in the case of the European represen-
tatives (e. g. there are about 20-30 species (Pawlowski, 1954, 1957; Plocek, 1983, 1990) of this subsection from Southern and Eastern Europe, which are hard to arrange with any aggregate, as I cannot assess their features) and must be further explored. From my primary observation in the relatives of Calycinae group, a key for the aggregates described above is proposed.
Identification key for aggregates of Alchemilla ser. Calycinae
1. Teeth of leaf blades of medium and upper leaves in total 50-80 (maximum 90).................................A. aggr. retinervis.
+ Teeth of leaf blades of medium and upper leaves in total
80-120 (maximum 135)......................................................... 2.
++ Teeth of leaf blades of medium and upper leaves in total
125-210 ....................................................... A. aggr. abchasica.
2. Central zone of the most medium and upper leaves more
than 76% of leaf length (total range 72-86%).......................
......................................................................A. aggr. betuletorum.
+ Central zone of the most medium and upper leaves less than 74% of leaf length (total range 52-78%), 6-8 teeth
at each side of the leaf lobes................................................... 3.
3. Incisions between lobes short, less than 1.5 of the standard
tooth length ............................................................................... 4.
+ Incisions between lobes profound, more than 1.5 of the standard tooth length .............................................................. 5.
4. Leaf lobes less than 25% of leaf length, apical tooth more
than 0.5 of the standard tooth length .......................................
...................................................................A. aggr. ellenbergiana.
+ Leaf lobes more than 25% of leaf length, apical tooth less
than 0.5 of the standard tooth length .......................................
................................................................ A. aggr. transcaucasica.
5. Hypanthia and pedicels ± hairy, often quite densely, stems pubescent up to the top ......................A. aggr. subsplendens.
+ Hypanthia and pedicels glabrous, stems hairy only in the lower half...............................................................A. aggr. dura.
Applying this scheme (Fig.) and reasoning presented above, the change of the borders between two series of A. subsect. Calycanthum is presented as follows:
Alchemilla ser. Elatae (Rothm.) Rothm., emended here. = A. subsect. Elatae Rothm. 1933, Feddes Repert. 33: 854, p. p. = A. ser. Elatae (Rothm.) Rothm. 1938, Feddes Repert. Beih. 100: 59, p. p.; Plocek, 1982, Preslia, 53: 50, p. max. p.
= A. sect. Calycinae Buser, 1893, in Magnier, Scr. Fl. Sel. 12: 278, p. p., excl. typo; id. 1896, Bull. Herb. Boiss. 4: 758, p. p., excl. typo.
= A. sect. Erectae S. E. Frohner, 1986, Gleditschia, 14: 30, p. max. p.
= A. sect. Alchemilla: S. E. Frohner, 1986, Gleditschia, 14: 34, p. p., non auct. mult.
Found in mesophilic conditions, large plants (up to 70 cm), with a patent, long, sometimes rather bristly indumentum of petioles and stems, radical leaves
with 120-250 teeth in total (from 90 teeth of small specimens from dry habitats, but with non-coriaceous leaves), spreading pubescent beneath (sometimes woolly); with (30)60-140 teeth of cauline leaves, differently pubescent on leaves (above), stems, flowers, and pedicels. Mesophilic plants tied to margins of relict Tertiary broadleaved forests, or derivative meadows. Distributed in the Caucasus, Asia Minor, Balkans, Carpathians (1 species), and Iran (6).
Alchemilla ser. Calycinae (Buser) Rothm., emended here. = A. ser. Calycinae (Buser) Rothm. 1938, Feddes Repert. Beih. 100: 59; Plocek, 1982, Preslia, 53: 50. = A. sect. Calycinae Buser, 1893, in Magnier, Scr. Fl. Sel. 12: 278, p. p., incl. typo; id. 1896, Bull. Herb. Boiss. 4: 758, p. p., incl. typo; S. E. Frohner, 1986, Gleditschia, 14: 35.
= A. subsect. Elatae Rothm. 1933, Feddes Repert. 33: 854, p. p., excl. typo. = A. ser. Elatae (Rothm.) Rothm. 1938, Feddes Repert. Beih. 100: 59, p. p., excl. typo.
= A. ser. Retinerves Plocek, 1982, Preslia, 53: 50.
= A. ser. Venosae Plocek, 1982, Preslia, 53: 50.
= A. sect. Coriacea S. E. Frohner, 1986, Gleditschia, 14: 41, p. min. p.
= A. sect. Erectae S. E. Frohner, 1986, Gleditschia, 14: 30, p. min. p.
= A. sect. Alchemilla: S. E. Frohner, 1986, Gleditschia, 14: 34, p. min. p., non auct. mult.
Usually small plants, or medium (up to 40 cm), found in mesophilic conditions, generally glabrous or with appressed slender indumentum of petioles and stems; if subappressed, then with appressed hairs on the leaves beneath; radical leaves with 50-120 teeth in total (to 220 teeth for A. aggr. abchasica, but still mostly glabrous or with appressed hairs on the leaves beneath), with ever glabrous leaves from above; with 15-40 (to 70-100 for A. aggr. abchasica) teeth of cauline leaves; usually with a coriaceous appearance. Plants of rocky slopes, dry or moist, often near the snowline, banks of watercourses, or of secondary disturbed habitats such as roadsides. Distributed in the Caucasus, Asia Minor, Iran (2), as well as the Balkans, Carpathians, Alps, Pyrenees, and ?Urals.
In the light of suggestions outlined above, several additions to the Rothmaler's concept made by Pawlowski (1954) and Plocek (1982, 1990) need to be discussed.
The Calycinae as proposed above is approximately equal as a whole to the three series presented by Plo-cek (1982): A. ser. Calycinae, A. ser. Retinerves Plocek, A. ser. Venosae Plocek. As it came from Plocek's comparison of A. ser. Retinerves with A. ser. Calycinae, the
latter was supposed to comprise European representatives. Therefore, it should include most of my A. dura aggregate. In that concept, the Retinerves, being supposed as a Caucasian group, united arbitrarily A. dura and A. retinervis, which were in fact rather remote from each other. At the same time, the types of both groups (A. retinervis Buser and A. fissa Günther et Schummel (A. glabra Poiret, non Neygenf.), respectively) belong, in my opinion, to the same aggregate. Thus, I shall consider A. ser. Retinerves synonymic to A. ser. Calycinae. The last A. ser. Venosae Plocek (1982) being barely described, probably, was supposed to embody the rest of our aggregates. However, the only I can say with confidence is that the valid name A. subser. Venosae Plocek (from A. ser. Venosae), which was typified by A. venosa and noted with a reference to Juzepczuk's "cikl" (circle) Subsplendentes, may be applied to my A. aggr. subsplen-dens, but in restricted sense. Shifted by Plocek (1982) A. subser. Semielatae Pawl. (with the type A. gorcensis) from A. ser. Elatae to A. ser. Venosae may be relevant for that part of A. aggr. abchasica, which includes species with no or poor hairiness. Later, endemic of the Carpathians A. ser. Calycinae subser. Serratae Plocek (1990) was also described, which I can hardly discuss having almost no material available — only illustration of A. hyperptycha Plocek (1990: Fig. 16). It might be quite close to A. subser. Semielatae mentioned above and a part of A. aggr. abchasica as well as another group, A. ser. Venosae subser. Pterophyllum Plocek. I can conclude that Re-, Su-, and Ab-aggregates were covered by previous studies with some taxonomic outcomes, which can be employed in the future, especially, for the latter group. The presence of Be-, El-, Du-, and Tr-aggregates have been previously overlooked, and their specifity has not been outlined.
Key for the Alchemilla ser. Calycinae species of the Caucasus and Asia Minor
1. Central zone of most of the leaves 50-70% of leaf length, and stems glabrous at least in the upper 1-2 internodes to glabrous throughout................................................................ 2.
+ Central zone of most of the leaves more than 70% of leaf
length, or, when less than 70%, stems hairy throughout .....
...................................................................................................... 11.
2. Teeth 9-12 at each side of the leaf lobes, lobes triangular at the top, stems hairy almost up to the top with appressed
or patent hairs ................................................. A. tredecimloba.
+ Teeth fewer than 9 at each side of the leaf lobes, lobes different....................................................................................... 3.
3. Stems hairy only in the 1-2 lower internodes (in two, when the first one shortened), or totally glabrous........... 4.
+ Stems hairy above the first lower elongate internode..... 6.
4. Leaf lobes more than Vi of leaf length, teeth to 8 at each side of the leaf lobes, stems hairy at the lowermost internode or glabrous throughout ........................................ 5.
+ Leaf lobes less than V4 of leaf length, teeth to 5 at each side
of the leaf lobes, stems glabrous throughout..........................
.................................................................................A. adelodictya.
5. Stems hairy at the lowermost internode, incisions between lobes less than standard tooth length, teeth to 8 at each side of the leaf lobes..........................................A. ptyschensis.
+ Stems glabrous throughout, incisions between lobes 1-2 of standard tooth length, teeth to 6 at each side of the leaf lobes.....................................................................A. microdictya.
6. Teeth 4-6 at each side of the leaf lobes............................... 7.
+ Teeth 6 and more at each side of the leaf lobes ................. 9.
7. Hypanthia glabrous, stems hairy very sparsely in the lower half only........................................................A. retinervis.
+ Hypanthia ± hairy, stems densely hairy ............................. 8.
8. Indumentum patent, stems hairy only in the lower half .....
.........................................................................A. hayirliogluorum.
+ Indumentum appressed or subappressed, stems hairy also in the upper half...................................................A. ancerensis.
9. Angle of leaf sector 45-60° ...................... A. transcaucasica.
+ Angle of leaf sector 25-40° .................................................. 10.
10. Indumentum erectopatent, leaves densely hairy beneath, leaf lobes rounded at the top .........................A. kazbekensis.
+ Indumentum appressed, leaves glabrous on the surfaces beneath, leaf lobes obtuse at the top ................ A. buseriana.
11. Teeth 9 or more at each side of the leaf lobes .................. 12.
+ Teeth usually fewer than 9 at each side of the leaf lobes .....
...................................................................................................... 16.
12. Angle of leaf sector 25-35° .................................................. 13.
+ Angle of leaf sector 35-45° .................................................. 15.
13. Incisions between lobes 1-3 of standard tooth length........
...................................................................................................... 14.
+ Incisions between lobes less than standard tooth length, leaf lobes obtuse or triangular at the top, leaves mostly plain or nearly so ............................................ A. undecimloba.
14. Leaf lobes obtuse or triangular at the top, leaves mostly
plain, indumentum of stems and petioles very sparse ..........
.................................................................................... A. buseriana.
+ Leaf lobes mostly rounded at the top, leaves undulate, indumentum of stems and petioles dense .. A. camptopoda.
15. Leaf lobes semiovate, parabolic or elongate triangular, leaves mostly reniform ........................................ A. abchasica.
+ Leaf lobes arcuate or shortly trapezoid to widely
parabolic, leaves mostly orbicular or orbicular reniform ....
................................................................................... A. ciminensis.
16. Stems hairy only in the lower internode or totally glabrous..................................................................................... 17.
+ Stems hairy above the lower internode ............................21.
17. Angle of leaf sector 30-45° .................................................. 18.
+ Angle of leaf sector 45-60° .................................................. 19.
18. Some hypanthia sparsely hairy at the base, pedicels to 2(3) of the hypanthia length, upper leaves glabrous beneath (except for basal lobes and main veins) .......................
................................................................................ A. cartilaginea.
+ All hypanthia glabrous, pedicels to 6(8) of the hypanthia
length, upper leaves evenly hairy beneath ..............................
......................................................................... A. longipedicellata.
19. All the petioles hairy (with sub-appressed and erectopatent hairs), leaf lobes shorter than Vi of leaf length ..........................................................................................A. debilis.
+ Most of petioles glabrous (if hairy, with appressed hairs), leaf lobes longer than У4 of leaf length...............................20.
20. Stems glabrous throughout, leaf teeth of ordinary width (length to width ratio 1-2) .........................A. muldaschevii.
+ Stems hairy at the lowest internode, leaf teeth wide (length to width ratio less than 1)..................... A. rivularis.
21. Stems hairy throughout........................................................22.
+ Stems glabrous at least in the upper 1-2 internodes to
glabrous throughout..............................................................33.
22. Teeth 3-5(6) at each side of the lobes ..............................23.
+ Teeth 6-8 at each side of the lobes ....................................27.
23. Lobes longer than Vi of leaf length.....................................24.
+ Lobes shorter than Vi of leaf length ...................................25.
24. Apical teeth less than 1/2 of standard ones, pedicels hairy ... ................................................................................. A. grandidens.
+ Apical teeth more than V2 of standard ones, pedicels glabrous..................................................................A. tiryalensis.
25. Incisions between lobes short, around 1 standard tooth length, indumentum appressed...........................................26.
+ Incisions between lobes long, more than 2 standard tooth lengths, indumentum patent ......................... A. akdoganica.
26. Leaves reniform, evenly hairy beneath......A. bezengiensis.
+ Leaves orbicular or orbicular reniform with glabrous
surfaces beneath...........................................A. retinerviformis.
27. Lobes longer than Vi of leaf length.....................................28.
+ Lobes shorter than Vi of leaf length ...................................30.
28. Hypanthia and pedicels glabrous or nearly so .......................
..............................................................A. pseudotranscaucasica.
+ Hypanthia and pedicels hairy .............................................29.
29. Incisions short (about 1 standard tooth length), teeth nearly obtuse, and almost equally-sized (standard tooth 1-2 times longer than the lowermost tooth) ... A. subsplendens.
+ Incisions long (more than 2 standard tooth length), teeth acute, strongly unequally-sized (standard tooth to 4-5 times longer than the lowermost tooth)........... A. speciosa.
30. Angle of leaf sector 40-50° ..................................................31.
+ Angle of leaf sector 35-45°, apical teeth more than 2/3 of
standard ones, hypanthia and pedicels hairy ..........................
................................................................................ A. sevangensis.
31. Petioles and stems patent hairy, hypanthia glabrous...........
..................................................................................A. beyazoglui.
+ Petioles and stems pubescent with appressed hairs.......32.
32. Apical teeth less than У2 of standard ones, pedicels hairy, hypanthia hairy, incisions long (more than 1.5 standard tooth length) ........................................................ A. divaricans.
+ Apical teeth more than 1/2 of standard ones, pedicels almost glabrous, hypanthia sparsely pubescent or glabrous,
incisions short (less than 1 standard tooth length) .............
......................................................................................... A. venosa
33. Lobes longer than у of leaf length.....................................34
+ Lobes shorter than У4 of leaf length ...................................37
34. Hypanthia ± hairy................................................A. oriturcica
+ Hypanthia glabrous ...............................................................35
35. Angle of leaf sector 45° and more ....................... A. indurata
+ Angle of leaf sector 40° and less .......................................... 36.
36. Indumentum erectopatent, lobes to semicircular ............
................................................................................A. kazbekensis
+ Indumentum appressed, lobes to parabolic .......................
.............................................................................A. wischniewskii
37. Hypanthia ± hairy..................................................................38.
+ Hypanthia glabrous ...............................................................40.
38. Angle of leaf sector 35-45(50)°, leaves dissected to y and more, 7-9-lobed......................................................................39.
+ Angle of leaf sector 25-30°, leaves dissected to y and less, 11-13-lobed.................................................... A. sciadiophylla.
39. Lobes semicircular, rounded at the top, medium and upper leaves glabrous on both surfaces.............................A. stricta.
+ Lobes ± triangular at the top, medium and upper leaves evenly sparsely hairy on both surfaces........A. barbatiflora.
40. Leaf lobes plane, shortly trapezoid to arcuate.................41.
+ Leaf lobes semicircular to semiovate.................................42.
41. Stems to 2/3 hairy, leaves sometimes sparsely hairy beneath, with hairy main veins throughout ...............A. betuletorum.
+ Stems to y-y sparsely hairy, leaves glabrous on the lower surface, with glabrous main veins at the base ........A. dura.
42. Teeth 6-8 at each side of the lobes, leaves glabrous or
sometimes sparsely hairy on the lower surface ......................
..................................................................................A. procerrima.
+ Teeth 4-6 at each side of the lobes, leaves densely hairy on the lower surface............................................A. ellenbergiana.
Conclusion
I have emended the borders between A. ser. Calycinae and A. ser. Elatae, treating the latter group as relict, attached to the Tertiary broadleaved forests of the Eastern Mediterranean basin. In these renewed circumscription, A. ser. Calycinae is subdivided into seven provisional aggregates according to the specially developed coordinate system (with two axes — a ratio of central zone value to leaf length vs. number of leaf teeth in total). The subdivision of the series in a hierarchical way is expected after a proper revision of European representatives and revealing of these aggregates geography in full.
Acknowledgements
I am grateful for the invaluable help with the search and digitization of authentic materials to many colleagues: Dr. J. Müller (Herbarium Haussknecht, Friedrich Schiller University Jena, Germany), Dr. A. Fleischmann, Dr. H.-J. Esser (Botanische Staatssammlung München, Germany), Dr. R. Vogt (Botanischer Garten und Botanisches Museum Berlin-Dahlem, Germany), Dr. M. A. Piirainen (Finnish Museum of Natural History, University of Helsinki, Finland), H. Smith (Royal Botanic Gardens Kew, London, UK), Dr. M. Granisze-wska (University of Warsaw, Poland), Dr. W. Till, M. Hofbauer, D. Reich (University of Vienna, Austria), Dr. L. Cecchi, Dr. Ch. Nepi (Museo di Storia Naturale, Universita degli Studi di Firenze, Italy), Dr. C. Sarthou (Muséum national d'Histoire Naturelle, Paris, France), Dr. L. Gautier, L. Loze (Conservatoire et jardin botaniques (CJB), Genève, Switzerland), Dr. M. Mosulish-vili (Georgian National Museum, Tbilisi, Georgia),
Dr. Ü. Reier, K. Orav (University of Tartu, Estonia). I thank D. G. Melnikov, I. V. Tatanov, A. V. Leostrin, and V. V. Shvanova (V. L. Komarov Botanical Institute of RAS, St. Petersburg) for their help and assistance during this study. I thank the anonymous reviewers whose remarks helped to improve the manuscript. The work is supported by grant RFBR 20-04-00183.
References
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