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Turczaninowia 19 (4): 114-128 (2016) DOI: 10.14258/turczaninowia.19.4.16 http://turczaninowia.asu.ru
ISSN 1560-7259 (print edition)
TURCZANINOWIA
ISSN 1560-7267 (online edition)
Dedicated to the memory of Rudolf Vladimirovich Kamelin (1938— 2016), Dr. Biol. Sci., Prof., corresponding member of Russian Academy of Sciences, renown Soviet and Russian botanist, expert in Potentilla, Rosaceae, and a founder of Altai botanical school (Barnaul).
Посвящается светлой памяти доктора биологических наук, члена-корреспондента РАН, профессора Рудольфа Владимировича Камелина (1938-2016), выдающегося советского и российского ботаника, знатока рода PotentШa, семейства Rosaceae, и основателя алтайской школы ботаники.
УДК 582.734.4(57.06)
A system of subtribe Potentillinae J. Presl (Rosaceae Juss.)
A. A. Kechaykin, A. I. Shmakov
South-Siberian Botanical Garden, Altai State University, Lenin str. 61, Barnaul, 656049, Russia
E-mail: [email protected]
Key words, new combination, typification, new sections, Argentina Hill, Horkelia Cham. et Schltdl., Potentilla L.
Summary. A new system of subtribe Potentillinae J. Presl down to the rank of sections is presented. The Fragari-astrum and Tormentilla groups of the genus Potentilla L. is classified here as distinct genera, supported by results of molecular studies and morphological differences. Most of species of these groups formerly included in Potentilla are transferred to the separate genera Fragariastrum Heist. ex Fabr. and Tormentilla L. New combinations in Argentina Hill, Horkelia Cham. et Schltdl., and Potentilla are also validated; some sections of these genera are lectotypified. The need of a new typification of Potentilla with subsequent conservation of the name in order to avoid nomenclatural instability is briefly discussed; finally, five new sections within Argentina are described.
Ключевые слова: новые комбинации, типификация, новые секции, Argentina Hill, Horkelia Cham. et Schltdl.,
Potentilla L.
Аннотация. Представлена новая система подтрибы Potentillinae J. Presl в мировом объёме до ранга секций. Группы Fragariastrum и Tormentilla рода Potentilla L. на основании результатов молекулярных исследований и морфологических различий рассматриваются здесь в качестве самостоятельных родов. Большинство видов этих групп, ранее включавшихся в Potentilla, переводятся в отдельные роды Fragariastrum Heist. ех Fabr. и Tormentilla L. В родах Argentina Hill, Horkelia Cham. et Schltdl. и Potentilla приводятся новые комбинации, а также лектотипификация отдельных таксонов. Кратко обсуждается необходимость новой типификации рода Potentilla последующей консервацией этого названия во избежание номенклатурной нестабильности. В роде Argentina описывается пять новых секций.
Поступило в редакцию 12.10.2016 Submitted 12.10.2016
Принято к публикации 20.11.2016 Accepted 20.11.2016
Система подтрибы Potentillinae J. Presl (Rosaceae Juss.)
А. А. Кечайкин, А. И. Шмаков
Южно-Сибирский ботанический сад, Алтайский государственный университет, пр. Ленина, 61, Барнаул, 656049, Россия
Introduction
Development of molecular-genetic methods in the end of XX century prompted phylogenetic research in Rosaceae Juss. and Potentilla s. l., in particular. One of the first works in this field was ITS-based phylogenetic analysis which included 14 Potentilla s. l. species and members of 19 related genera (each represented by a single species), except for Fragaria L. Relevant tree of 34 Rosaceae taxa demonstrated lack of monophyly of Potentilla. In a subsequent, ITS- and trnLF-based study of 44 Rosaceae (including 18 Potentilla s. l.) species (Eriksson et al., 2003), monophyly of Potentilleae Juss. has been confirmed.
Potter et al. (2007) constructed a family-wide phylogeny based on a multigene (6 nuclear and 4 cp markers) analysis of 88 Rosaceae genera. Results of this analysis have been utilized for elaboration of a new system of the family. Two subtribes were recognized within Potentilleae: 1) Potentillinae J. Presl with the single genus Potentilla L. (incl. Argentina Hill, Comarella Rydb., Duchesnea Sm., Horkelia Cham. et Schltdl., Horkeliella (Rydb.) Rydb., Ivesia Torr. et A. Gray, Purpusia Brandegee, Stellariop-sis (Baill.) Rydb.); and 2) Fragariinae Torr. et A. Gray which comprised Alchemilla L. (incl. Aphanes L., Lachemilla (Focke) Lagerh., and Zygalchemi-lla Rydb.), Chamaerhodos Bunge, Comarum L. (incl. Farinopsis Chrtek et Sojak), Dasiphora Raf. (=Pentaphylloides Duhamel), Drymocallis Fourr., Fragaria L., Potaninia Maxim., Sibbaldianthe Juz. (incl. Schistophyllidium (Juz. ex Fed.) Ikonn.), Sib-baldia L., and Sibbaldiopsis Rydb. (Potter et al., 2007). The authors emphasized the lack of reliable non-molecular features for delimitation of the above subtribes.
Such a character has been reported soon by Sojak (2008) who discovered that representatives of Potentillinae have anthers with two thecae while members of Fragariinae possess anthers with one theca. Thus, results of molecular phylogeny have been supported by morphological data.
More recently, Faghir et al. (2014) reconstructed the phylogeny of Iranian species of Potentilla and some related genera (over 60 spp. in total) using ITS and trnLF markers. This study confirmed the mono-phyly of Potentilla and pored light on evolution of certain taxonomically important morphological characters. In particular, independent evolvement of anther structure, style shape and position has been demonstrated while parallel evolution was revealed for leaf morphology.
Some other phylogenetic studies puplished in between of Potter et al. (2007) and Faghir et al. (2014) also confirmed monophyly of Potentilla and division of Potentilleae onto two subtribes (e. g., Dobes, Paule, 2010; Topel et al., 2011).
Results and discussion
The current paper presents the global system of subtribe Potentillinae comprising 12 genera possessing anthers with two thecae. Majority of genera of Potentillinae are endemic to North America and characterized by exclusively pinnate leaves (often with 4-15 pairs of deeply incised and whorled leaflets) and small, usually rather narrow petals. One of the most diverse New World Potentillinae genera, Horkelia, is distributed in the west of North America. Species of Horkelia have 10 stamens with flat broadened filaments predominantly attached to the hypanthium mouth. Carpels often numerous (20-50 and more), usually attached to the hypanthium base; inflorescence laxly racemose. The closest to Horkelia is Horkeliella represented by three species also distributed in the west of North America (Rydberg, 1908). The latter genus differs from the former by having 20 filaments and sepals recurved at anthesis.
Ivesia is another relatively big North American genus of Potentillinae which is characterized by 15-20 (rarely 10 or 5) stament with filiform filaments situated in the upper part of hypanthium, 4-5 to 10-15 carpels, often dense, subcapitate inflorescence and small flowers. Monotypic genus Purpusia (arid mountains of Nevada) is represented by cespi-tose-cusion plant lacking "outer calyx" and having 5 stamens attached to the hypanthium mouth and 5-7 carpels. Another monotypic American genus of Potentillinae is Stellariopsis endemic to the mountains of California and characterized by 15 stamens attached closer to the hypanthium base. It also has anthers opening by subterminal pores to short lateral slits and a single 1-seeded carpel in an open hypanthium. Leaves compound, pinnate, with numerous pairs of tightly sitting and densely pubescent small leaves.
The above six exclusively North American genera of Potentillinae demonstrate unique patterns of leaf morphology and generative organs not characteristic to the rest of subtribe including Potentilla. Original morphology within Potentillinae is demonstrated by Duchesnea (two species and a hybrid: Sojak, 2012). It is separated from Potentilla and other Potentillinae members by big, apically 3-5-dentate epicalyx scales ("outer sepals") and glabrous torus
with numerous ovaries, considerably enlarging and becoming globose, fleshy or spongy, red but, unlike in Fragaria, not detaching from the hypanthium. Phylogenetic studies show that Duchesnea (D. indica) falls into one, almost 100 % supported, clade with P. reptans L. and P. erecta (L.) Raeusch. (Kechaykin, 2016). R. V. Kamelin (2001) and Th. Wolf (1908) merged Duchesnea with Potentilla. The prior author placed P. indica into the section Duchesnea (J. E. Smith) B. K. Dikshit et G. Panigrahi, as Pani-grahi and Dikshit (1985) have done while the latter treated the same species as a member of subsection Gomphostylae Th. Wolf and group Tormentillae, along with P. erecta, P. reptans, P. flagellaris Willd. ex D. F. K. Schltdl., etc. (Wolf, 1908). Indeed, Duch-esnea species share some characters with members of Potentilla subgen. Tormentilla (L.) Kechaykin, e. g., single flowers on long stalks. Besides, representatives of both groups are perennial herbs with short rhizome and long, creeping shoots rooting at nodes (except for P. erecta having erect stem). This morphological similarity apparently reflects common origin of Duchesnea and Tormentilla. At the same time, having in mind resemblance of fruits of Duchesnea and Fragaria and inadequate information on phylogenetic position of D. chrysantha (Zoll. et Moritzi) Miq. and D. hara-kurowae Naruh. et Sugim. it seems reasonable to treat Duchesnea for the time being as a distinct genus.
Rather separate position in the system of Poten-tillinae is occupied by Dryadanthe Endl. (Endlicher, 1840). The only species of this genus, D. tetrandra (Bunge) Juz., occurs predominantly in high mountains of Central and Middle Asia. It is a densely cespitose to cusion-forming perennial with woody rhizomes and stems to 15 cm tall. The genus Dryadanthe differs from Potentilla s. str. by having 4 stamens, 4 pistils, and lateral styles. In contrast to the other genera of Potentillinae, flowers in Dryadanthe are unisexual.
The second biggest genus of Potentillinae is Argentina. The centre of its diversity and speciation (and, probably, origin) is situated in mountains of Palaeotropics (South-East Asia). 20 species of Argentina (over 25% of the whole number) are endemic to New Guinea (Sojak, 2012). Majority of Argentina species occur at the elevations above 3000 m a. s. l., have leaves with many (in average 10-20) pairs of leaflets; 10 species produce terrestrial creeping stolons rooting at nodes, others develop erect to ascending stems. One of the characters separating Argentina from Potentilla and other genera of Po-tentillinae is narrow style attached to the ventral
side of achene. This feature is, however, not reliable enough as in more than 10 Asian species the styles are subterminal (Soják, 2010). Instead, the difference in stipule structure detected by Soják (2010) appears to be more sufficient for sepatarion of Argentina from Potentilla. While stipule auricles in Potentilla s. str. are lateral (stretching along the leaf petiole margins), in Argentina they are ventral, and this difference is consistent (Soják, 2010). Molecular studies (Kechaykin, 2016; Feng et al., 2014) also support distinctness of Argentina from other genera of Potentillinae. Based on these findings, we follow Hill (1756), Rydberg (1898, 1908) and Soják (2004, 2010, 2012) in accepting Argentina as a separate genus.
Fragariastrum Heist. ex Fabr. and Tormentilla L. are also accepted here as distinct genera and the need of a new typification of Potentilla L. with subsequent conservation of the latter name is stressed for which the following agruments can be put forward. First, majority of Potentilla species are taprooted polycar-pic herbs (Serebryakov, 1964) with erect stems and many-flowered terminal inflorescences. It was mentioned in the literature (e. g., Kamelin, 2001) that Rydberg's (1908) lectotypification of Potentilla by P. reptans L. was not a good decision, an opinion with which we fully agree. Potentilla reptans is a stoloniferous polycarpic herb (Serebryakov, 1964) without developed inflorescence (represented by single axillary flowers on long pedicels). This life form is, with few exceptions, untypical for Poten-tilla. R. V. Kamelin (2001) noted that P reptans is similar to some species of Duchesnea Sm.
Second, and most important, retention of P reptans as the lectotype of Potentilla and acceptance of the genus Tormentilla (which contains P reptans), well-sepatated both morphologically and molecu-larly, would bring severe instability in nomenclature. Namely, it would request several hundreds of transfers of species from Potentilla to Tormentilla and drastic change of the concept of Potentilla. The same action is inevitbale if the above American genera are accepted unless the paraphyletic concept of Potentilla is adopted (Ertter, Reveal, 2014).
Hence, a new typification of Potentilla is needed. The best potential type would be P. argentea L., another, along with P reptans, original member of the genus (Linneaus, 1753). Potentilla argentea exhibits basic features typical for most Potentilla s. str. species. It is a taprooted polycarpic herb with erect stems, apical many-flowered inflorescence, and terminal styles (subterminal in Tormentilla). Unlike P reptans, P. argentea readily hybridyze with members of variuos sections of P. subgen. Potentilla.
Traditional concept of Potentilla is adopted here which includes P. argentea and excludes its current lectotype, P. reptans. Proposal to conserve Potentilla with a new type is in preparation.
Potentilla students emphasized that among the species of the genus intensive hybridization takes place which complicates systematics and identification of individual specimens. However, no natural hybrids between representatives of Potentilla s. str. with species of Drymocallis or Schistophyllidium from subtribe Fragariinae (where P rupestris L. and P. bifurca L., respectively, belong) are known to the date. Furthermore, no native hybrids between the genera of Potentillinae and active hybridization (often intersectional) is restricted to the genus Po-tentilla. This is an additional argument in favor of acceptance of Argentina, Drymocallis, Fragarias-trum, Schistophyllidium, Tormentilla, etc.
A key for the genera of Potentillinae as well as the system of the subtribe with some comments are given below.
Key to the genera of the subtribe Potentillinae
1. Stipule auricles ventral (stretching along the ventral side of leaf petiole) ...................................2
+ Stipule auricles lateral (stretching along the leaf petiole margins)............................................. 3
2. Subshrubs. Leaf petioles and pedicels articulate. Carpels pubescent .............. Tylosperma
+ Herbs. Leaf petioles and pedicels not articulate.
Carpels glabrour, occasionally pubescent...............
..................................................................Argentina
3. Elements of epicalyx ("outer sepals") 2-3 as wide as sepals, apically 3-5-dentate. Torus in fruit considerably enlarged, globose, fleshy, red or pink (reminding fruits of Fragaria L.).........Duchesnea
+ Elements of epicalyx ("outer sepals") narrower than or, rarely, as wide as sepals, apically entire or rarely 2-3-dentate. Torus not enlarged in fruit, not fleshy .................................................................... 4
4. Flowers unisexual, plants dioecious. Filaments 4 (staminoid in male flowers) .............Dryadanthe
+ Flowers bisexual. Filaments usually 5 to 30 .. ............................................................................... 5
5. Anthers opening by subterminal pores to short
lateral slits; carpels single .......................................
....................... Stellariopsis (endemic to California)
+ Anthers opening by lateral slits; carpels usually two or more .......................................................... 6
6. Carpels surrounded by hypanthium walls. Stamens attached to hypanthium; N America...... 7
+ Hypanthium absent. Stamens attached to torus ............................................................................. 10
7. Epicalyx ("outer sepals") absent .... Purpusia
+ Epicalyx ("outer sepals") present ................ 8
8. Filaments filiform (except for Ivesia argyrocoma (Rydb.) Rydb. var. argyrocoma), attached to the upper part of hypanthium, not
forming a tube. Hypanthia not flat-bottomed..........
........................................................................Ivesia
+ Filaments flat, dilated at base, attached predominantly to hypanthium mouth, often forming a tube. Hypanthia often flat-bottomed ................. 9
9. Stamens 10. Epicalyx ("outer sepals") not recurved at anthesis. Leaves usually with 2-20 pairs of leaflets ..................................................Horkelia
+ Stamens 20. Epicalyx ("outer sepals") recurved at anthesis. Leaves usually with 15-35 pairs of leaflets ..................................................Horkeliella
10. Inflorescense open (indeterminate)..............
...............................................................Tormentilla
+ Inflorescense closed (determinate) .............11
11. Styles filiform throughout, about twice longer than ripen achenes. Achenes usually pubescent with simple hairs, rarely glabrous .......... Fragariastrum
+ Styles not filiform throughout, as long as or shorter, rarely 2-3 times longer than ripen achenes (sect. Persicae) but then dilated and verrucose at
base. Achenes glabrous or, rarely, glandular...........
................................................................. Potentilla
System of the subtribe Potentillinae
Subtribus Potentillinae J. Presl, 1846, Wsobecny Rostl. 1: 491.
Type: Potentilla L.
12 genera: Argentina Hill, Dryadanthe Endl., Duchesnea Sm., Fragariastrum Heist. ex Fabr., Horkelia Cham. et Schltdl., Horkeliella (Rydb.) Rydb., Ivesia Torr. etA. Gray (incl. Comarella Rydb.), Potentilla L., Purpusia Brandegee, Stellariopsis (Baill.) Rydb., Tormentilla L., Tylosperma Botsch.
Genus I. Argentina Hill, 1756, Brit. Herb.: 6; Lam. 1778, Fl. Fr., ed. 2, 3: 118, s. str. - Potentilla L. subgen. Argentina (Hill) Jeps. 1936, Fl. Calif. 2(2): 175.
Lectotype (Rydberg, 1908: 352): Argentina an-serina (L.) Rydb. [=Potentilla anserina L.].
73 species, predominantly in Palaeotropics and Holarctics.
1. Argentina sect. Argentina. - Potentilla L. sect. Pentaphylloides Tausch, 1823, Hort. Canal. 1: (unpaged). - P sect. Anserina (Gaudin) Pfeiff. 1873, Nomencl. Bot. 1(1): 205. - P. ser. Anserina (Gaudin) Lehm. 1856, Revis. Potentill.: 8 (as "ser.
Anserinae"). - P [unranked] Anserina Gaudin, 1828, Fl. Helv. 3: 370, 405. - P. subgen. Chenopo-tentilla (Focke) Juz. 1941, Fl. URSS, 10: 80, 220. -P. sect. Chenopotentilla Focke, 1889, Abh. Natur-wiss. Vereins Bremen 10(3): 415. - P. sect. Lepto-stylae (Th. Wolf) Gu§ul. 1956, Fl. Republ. Populare Rom. 4: 656. - P. subsect. Leptostylae (Th. Wolf) Th. Wolf, 1908, Biblioth. Bot. 16(71): 52. - P. [unranked] Leptostylae Th. Wolf, 1904, Syn. Mitteleur. Fl. 6(16): 671.
Lectotype: Argentina anserina (L.) Rydb. [=Po-tentilla anserina L.].
Six species in mountains and plains of Holarc-tics, Palaeo- and Neotropics and Australia.
2. Argentina sect. Brassiae Kechaykin et Shma-kov, sect. nov.
Dwarf compact herbs, cespitose or forming cutions to 60 cm diam. Leaves to 1.5 cm long, very short-petiolate. Lateral leaflets entire, more rarily dissected onto two segments.
Type: Argentina brassii (Merr. et L. M. Perry) Sojak [=Potentilla brassii Merr. et L. M. Perry]. No. 10156. L. J. Brass & E. Myer-Drees, Sept. 1938. Abundant in alpine bogs. 4120 m. alt. Northern slope of Mt. Wilhelmina (A: 00003012).
Two species in mountains of New Guinea.
3. Argentina sect. Cristatae Kechaykin et Shma-kov, sect. nov.
Densely to loosely cespitose plants, sometimes forming cusions to 40 cm diam. Leave blades usually more than 1.5 cm long, with (3)4-6(7) pairs of leaflets. Leaves incised, 2-5-toothed on each side.
Type: Argentina cristata (H. R. Fletcher) Kechaykin et Shmakov, comb. nov. = P. cristata H. R. Fletcher, 1950, Notes Roy. Bot. Gard. Edinburgh, 20: 218. [P stenophylla Diels var. cristata (H. R. Fletcher) H. Ikeda et H. Ohba.]. 4 Aug. 1920. Altd. 11200 feet. Moka-ji Pass. No. 1803. Coll. R. F. (E: E00012085).
Six species in mountains of South and SouthEast Asia including New Guinea.
4. Argentina sect. Habbemanae Kechaykin et Shmakov, sect. nov.
Non-cespitose plants. Leaves usually with 6-15 (sometimes with 20 and more on the same plant) pairs of leaflets. Leaves serrate- or pseudoserrate-pinnatisect or dissected onto two unequal elliptic lobes. Central costa of lateral leaflet less than 3 mm long.
Type: Argentina habbemana (Merr. et L. M. Perry) Sojak [=Potentilla habbemana Merr. et L.
M. Perry]. No. 9594. L. J. Brass, Aug. 1938. Alpine grassland; abundant on marshy ground. Lake Habbema, 3225 m. camp (A: 00003039). Three species in mountains of New Guinea.
5. Argentina sect. Leuconotae (Dikshit et Pani-grahi) Kechaykin et Shmakov, comb. nov. = Poten-tilla L. sect. Leuconotae Dikshit et Panigrahi, 1981, J. Orissa Bot. Soc. 3(1): 33. - Potentilla L. sect. Leptostylae (Th. Wolf) Gu§ul. subsect. Microphyl-lae (T. T. Yu et C. L. Li) H. Ikeda et H. Ohba ser. Leuconotae (Dikshit & Panigrahi) H. Ikeda et H. Ohba, 1999, Himalayan Pl., 3 (Univ. Mus. Univ. Tokyo Bull., 39): 76.
Type: Argentina leuconota (D. Don) Sojak [=Po-tentilla leuconota D. Don].
Twelve species in mountains of South and SouthEast Asia including New Guinea.
6. Argentina sect. Lineatae (H. Ikeda et H. Ohba) Kechaykin et Shmakov, comb. et stat. nov. = Potentilla L. sect. Leptostylae (Th. Wolf) Gu§ul. subsect. Microphyllae (T. T. Yu et C. L. Li) H. Ikeda et H. Ohba ser. Lineatae H. Ikeda et H. Ohba, 1999, Himalayan Pl. 3 (Univ. Mus. Univ. Tokyo Bull., 39): 43.
Type: Argentina lineata (Trevir.) Sojak [=Potentilla lineata Trevir.].
Seven species in mountains of South and SouthEast Asia including New Guinea.
7. Argentina sect. Microphyllae (T. T. Yu et C. L. Li) Kechaykin et Shmakov, comb. et stat. nov.
= Potentilla L. sect. Leptostylae (Th. Wolf) Gu§ul. ser. Microphyllae T. T. Yu et C. L. Li, 1980, Acta Phytotax. Sin. 18: 5.
Type: Argentina microphylla (D. Don) Sojak [=Potentilla microphylla D. Don].
Ten species in mountains of South and SouthEast Asia including New Guinea.
8. Argentina sect. Parvulae Kechaykin et Shmakov, sect. nov.
Usually non-cespitose plants. Leaves with more than 5 pairs of leaflets. Lateral leaflets round in outline, often with 2-6 teeth on each side. Central costa of lateral leaflet more than 3 mm long.
Type: Argentina parvula (Hook. f. ex Stapf) Sojak [=Potentilla parvula
Hook. f. ex Stapf]. 1057K, G. D. Haviland, Aug. 1892. Kinabalu, 11000 ft. (K: K000762536).
17 species in mountains of South and South-East Asia including New Guinea.
9. Argentina sect. Piletophyllum (Sojak) Ke-chaykin et Shmakov, comb. nov. = Sibbaldia L. sect. Piletophyllum Sojak, 1970, Preslia, 42: 185. [Piletophyllum (Sojak) Sojak, 2008, Bot. Jahrb. Syst. 127(3): 356].
Type: Argentina micropetala (D. Don) Sojak [=Potentilla micropetala D. Don]. Potentilla sp. E. Gos. Than. 1809. Wallich (BM: BM000521790). Two species in mountains of South Asia.
10. Argentina sect. Vittatae Kechaykin et Shma-kov, sect. nov.
Non-cespitose plants. Leaves usually with more than 5 pairs of leaflets. Lateral leaflets entire or api-cally 2-4-dentate. Central costa of lateral leaflet more than 3 mm long.
Type: Argentina vittata (Sojak) Sojak [=Poten-tilla vittata Sojak]. Kongbo, SE Tibet. No. 13966. Ba La, Pasum Chu. 22. 6. [19]47. Coll. F. Ludlow, G. Sherriff & H. H. Elliot (BM: BM000551381).
Eight species in mountains of South and SouthEast Asia including New Guinea.
Genus II. Dryadanthe Endl. 1840, Gen. Pl.: 1242.
Type: Dryananthe tetrandra (Bunge) Juz. [=Sib-baldia tetrandra Bunge, Potentilla tetrandra (Bunge) Hook. f.].
Monotypic genus distributed in mountains of Asia between 55o and 26o N.
Genus III. Duchesnea Sm. 1811, Trans. Linn. Soc. London, 10(2): 372. - Potentilla L. subgen. Duchesnea (Sm.) Shah et Wilcock, 1993, Edinb. J. Bot. 50(2): 176. - Potentilla L. sect. Duchesnea (Sm.) Panigr. et Dikshit, 1985, Bull. Bot. Surv. Ind. 27(1-4): 181.
Lectotype (Rydberg, 1908: 356): Duchesnea indica (Andrews) Teschem. [=Fragaria indica Andrews].
Two species in Holarctics, Palaeo- and Neo-tropics and Australia.
Genus IV. Fragariastrum Heist. ex Fabr. 1759, Enum. Meth. Pl.: 64. - Potentilla subgen. Fragariastrum (Heist. ex Fabr.) Reich. 1828, Consp. Regn. Veget. 167. - P. subgen. Micropogon Juz. 1941, Fl. URSS, 10: 84. - Micropogon Bunge, 1835, Mem. Acad. Imp. Sci. Saint Petersbourg, 2(7): 552, nom. provis. - P. sect. Nematostylae (Th. Wolf) Gu§uleac, 1956, in Suvulevscu, Fl. Reipubl. Popularis Roman. 4: 602. - P. subsect. Nematostylae Th. Wolf, 1908, in Biblioth. Bot. 16 (71): 42, 45 p. p.
Lectotype (Panigrahi, Diskhit, 1986: 350): Fragariastrum sterile (L.) Schur [=Fragaria sterilis L, = Potentilla sterilis (L.) Garcke].
About 40 species in Northern Hemisphere.
1. Fragariastrum sect. Ancistrifoliae (Soják) Kechaykin et Shmakov, comb nov. = Potentilla sect. Ancistrifoliae Soják, 1987, Candollea, 42(2): 495.
Type: Fragariastrum ancistrifolia (Bunge) Kechaykin et Shmakov
Three species in East Asia (Soják, 1987).
Fragariastrum ancistrifolium (Bunge) Kechaykin et Shmakov, comb nov. = Potentilla ancistrifolia Bunge, 1833, Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6. Sci. Math. 2: 99.
Fragariastrum dickinsii (Franch. et Sav.) Kechaykin et Shmakov, comb nov. = Potentilla dickinsii Franch. et Sav. 1878, Enum. Pl. Jap. 2: 337.
Fragariastrum rugulosum (Kitag.) Kechaykin et Shmakov, comb nov. = Potentilla rugulosa Kitag. 1937, Rep. Inst. Sci. Res. Manchoukuo, 1: 260.
Note. This group is intermediate between Fragariastrum and Potentilla. Woody base and pubescent fruits are characteristic to Fragariastrum, though in leaf morphology and inflorescence structure it is closer to sect. Fragarioides of Potentilla subgen. Potentilla. Available molecular data demonstrate relationship of representarives of F. sect. Ancistrifoliae with either P. sect. Fragarioides (trnLF-based phylogeny) or Fragariastrum (ITS-based phylogeny) (Kechaykin, 2016). We keep sect. Ancistrifoliae within Fragariastrum for the moment but its placement is to be elucidated.
2. Fragariastrum sect. Bhutanicae (Soják) Kechaykin et Shmakov, comb nov. = Potentilla sect. Bhutanicae Soják, 1987, Candollea 42(2): 495, 497.
Type: Fragariastrum bhutanicum (Ludlow) Kechaykin et Shmakov.
One species in Bhutan.
Fragariastrum bhutanicum (Ludlow) Kechaykin et Shmakov, comb nov. = Potentilla bhutanica Ludlow, 1956, Bull. Brit. Mus. (Nat. Hist.), Bot. 2: 68.
3. Fragariastrum sect. Biflorae (Rydb.) Ke-chaykin et Shmakov, comb et stat. nov. = Poten-tilla L. [unranked] Biflorae Rydb. 1897, Bull. Tor-rey Bot. Club 24(1): 13. - P. sect. Biflorae (Rydb.) O. Stevens, 1959, in North Amer. Fl. 22 (7): 11, comb. inval. (sine cit. bas.). - P. sect. Biflorae (Th.
Wolf) Schiman-Czeika, 1969, in K. H. Reichinger, Fl. Iran. 66: 85, comb. inval. (sine cit. bas.). -P. [unranked] Biflorae (Rydb.) Th. Wolf, 1908, Bib-lioth. Bot. 16(71): 46, 70. - P. sect. Nematostylae (Th. Wolf) Gu§uleac ser. Biflorae (Th. Wolf) T.T. Yu & C.L. Li, 1985, Fl. Reipubl. Popularis Sin. 37: 235, 256, comb. inval. - P. sect. Eriocarpae (Th. Wolf) Juz. ser. Biflorae (Th. Wolf) Sojak, 1987, Candollea, 42(2): 495, comb. inval. - P. sect. Eriocarpae (Th. Wolf) Sojak ser. Articulatae (Th. Wolf) Sojak, 1987, Candollea, 42(2): 495.
Type: Fragariastrum biflorum (Willd. ex Schl-tdl.) Kechaykin et Shmakov
Two species; Asia and North America.
Fragariastrum biflorum (Willd. ex Schltdl.) Kechaykin et Shmakov, comb. nov. = Potentilla biflora Willd. ex Schltdl. 1816, Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin, 7: 295.
Fragariastrum articulatum (Franch.) Kechaykin et Shmakov, comb. nov. = Potentilla articulata Franch. 1890, Pl. Delavay. 3: 210.
4. Fragariastrum sect. Camillae (Kapeller) Kechaykin et Shmakov, comb. nov. = Potentilla sect. Camilla Kapeller, 1956, Not. Syst. Georg. Inst. Bot. Thibiliss. 19: 18.
Type: Fragariastrum camillae (Kolak.) Kechaykin et Shmakov
One species (Caucasus: Russia, Abkhasia).
Fragariastrum camillae (Kolak.) Kechaykin et Shmakov, comb. nov. = Potentilla camillae Kolak. 1936, Bot. Zhurn. S.S.S.R. 21: 553.
5. Fragariastrum sect. Curvisetae (Th. Wolf) Kechaykin et Shmakov, comb. nov. = Potentilla L. [unranked] Curvisetae Th. Wolf, 1908, Biblioth. Bot. 16(71): 47, 94. [Potentilla sect. Curvisetae (Th. Wolf) Schiman-Czeika, 1969, Fl. Iran. 66: 87].
Type: Fragariastrum curvisetum (Hook. f.) Kechaykin et Shmakov
Three species distributed in Afghanistan, Pakistan, and India.
Fragariastrum curvisetum (Hook. f.) Kechaykin et Shmakov, comb. nov. = Potentilla curviseta Hook.f. 1878, Fl. Brit. India, 2(5): 358.
Fragariastrum collettianum (Aitch. et Hemsl.) Kechaykin et Shmakov, comb. nov. = Potentilla collettiana Aitch. et Hemsl. 1880, in J. Linn. Soc. 18: 53.
Fragariastrum pteropodum (Royle) Kechaykin et Shmakov, comb. nov. = Potentilla pteropoda Royle, 1834, Ill. Bot. Him. 2: t. 40 f. 2.
6. Fragariastrum sect. Elatiores (Kapeller) Kechaykin et Shmakov, comb. nov. = Potentilla sect. Elatiores Kapeller, 1956, Not. Syst. Georg. Inst. Bot. Thibiliss. 19: 21.
Type: Fragariastrum elatior (Willd. ex Schltdl.) Kechaykin et Shmakov
One species (Caucasus and Turkey).
Fragariastrum elatior (Willd. ex Schltdl.) Kechaykin et Shmakov, comb. nov. = Potentilla elatior Willd. ex Schltdl. 1816, Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin, 7: 295.
7. Fragariastrum sect. Eriocarpae (Th. Wolf) Kechaykin et Shmakov, comb. nov. = Potentilla L. [unranked] Eriocarpae Th. Wolf, 1908, Biblioth. Bot. 16(71): 46, 79. [Potentilla L. sect. Eriocarpae (Th. Wolf) Juz. 1941, Fl. URSS, 10: 86].
Type: Fragariastrum eriocarpum (Wall. ex Lehm.) Kechaykin et Shmakov
One species (India, Nepal, Bhutan, China).
Fragariastrum eriocarpum (Wall. ex Lehm.) Kechaykin et Shmakov, comb. nov. = Potentilla eriocarpa Wall. ex Lehm. 1831, Nov. Stirp. Pug. 3: 35.
8. Fragariastrum sect. Fragariastrum = Potentilla sect. Fragariastrum (Heist. ex Fabr.) Ser. 1825, in DC., Prodr. 2: 583. - P. sect. Micranthae Soják, 1987, Candollea 42(2): 495, 498.
Type: Fragariastrum sterile (L.) Schur [=Fragaria sterilis L.].
Six species in Eurasia and northern North America.
Fragariastrum davisii (H. Duman) Kechaykin et Shmakov, comb. nov. = Potentilla davisii H. Duman, 1999, in H. Duman et R.R. Mill, Edinburgh J. Bot. 56: 351.
Fragariastrum carniolicum (Kern.) Kechaykin et Shmakov, comb. nov. = Potentilla carniolica Kern. 1870, Oesterr. Bot. Z. 20: 44.
Fragariastrum imerethicum (Gagnigze et Sokhadze) Kechaykin et Shmakov, comb. nov. = Potentilla imerethica Gagnigze et Sokhadze, 1982, Zametki Sist. Georg. Rast. 38: 4.
Fragariastrum micranthum (Ramond ex DC.) Schur, 1853, in Verh. Mittheil. Siebenburg. Ver. Na-turw. Hermannst. iv. Anhang (Sert. Fl.Transsilv.): 23. - Potentilla micrantha Ramond ex DC. 1805, in Lam. et DC., Fl. Fran?., ed. 3, 4: 468.
Fragariastrum nerimaniae (H. Duman) Kechay-kin et Shmakov, comb. nov. = Potentilla nerima-niae H. Duman, 1999, in H. Duman et R. R. Mill, Edinburgh J. Bot. 56: 349.
Fragariastrum sterile (L.) Schur, 1853, in Verh. Mittheil. Siebenburg. Ver. Naturw. Hermannst. iv. Anhang (Sert. Fl.Transsilv.): 23. - Fragaria sterilis L. 1753, Sp. Pl.: 495. - Potentilla sterilis (L.) Gark-ke, 1856, Fl. Halle, 2: 200.
Fragariastrum ulrichii (Parolly et Nordt) Kechaykin et Shmakov, comb. nov. = Potentilla ulrichii Parolly et Nordt, 2002, Willdenowia, 32: 75.
9. Fragariastrum sect. Laterales (Döll) Kechaykin et Shmakov, comb. nov. = Potentilla L. [un-ranked] Laterales Döll, 1843, Rhein. Fl.: 769, p. p. = Potentilla L. sect. Laterales (Döll) Gren. et Godr. 1848, Fl. France, 1: 522. - P. sect. Crassinerviae (Th. Wolf) Juz. 1941, Fl. URSS, 10: 90. - P. [un-ranked] Crassinerviae Th. Wolf, 1908, Biblioth. Bot. 16(71): 47, 97. - P. sect. Umbellatae Sojak, 1987, Candollea 42(2): 495, 497.
Type: Fragariastrum caulescens (L.) Schur
About 14 species in mountains of Europe and North Africa.
Fragariastrum alchimilloides (Lapeyr.) Kechaykin et Shmakov, comb. nov. = Potentilla alchimilloides Lapeyr. 1782, Mem. Acad. Sci. Toulouse, 1: 212, t. 17.
Fragariastrum brachypetalum (Fisch. et C. A. Mey. ex Lehm.) Kechaykin et Shmakov, comb. nov. = Potentilla brachypetala Fisch. et C. A. Mey. ex Lehm. 1849, Del. Sem. Hort. Hamburg.: 9.
Fragariastrum caulescens (L.) Schur, 1853, in Verh. Mittheil. Siebenburg. Ver. Naturw. Hermannst. iv. Anhang (Sert. Fl. Transsilv.): 23. - Potentilla caulescens L. 1756, Cent. Pl. 2: 19.
Fragariastrum clusianum (Jacq.) Schur, 1853, in Verh. Mittheil. Siebenburg. Ver. Naturw. Hermannst. iv. Anhang (Sert. Fl. Transsilv.): 23. - Potentilla clusiana Jacq. 1774, Fl. Austriac. (Jacquin), 2: 10, t. 116.
Fragariastrum crassinervium (Viv.) Kechaykin et Shmakov, comb. nov. = Potentilla crassinervia Viv. 1825, App. Fl. Cors. Prodr.: 2.
Fragariastrum doddsii (P. H. Davis) Kechaykin et Shmakov, comb. nov. = Potentilla doddsii P. H. Davis, 1957, Notes Roy. Bot. Gard. Edinburgh, 22: 169.
Fragariastrum doerfleri (Wettst.) Kechaykin et Shmakov, comb. nov. = Potentilla doerfleri Wettst. 1892, in Luerss. Bibl. Bot. Heft, 26: 39.
Fragariastrum grammopetalum (Moretti) Kechaykin et Shmakov, comb. nov. = Potentilla gram-mopetala Moretti, 1826, Giorn. Fis. Chim. Storia Nat. Med. Arti Dec. 2: 9: 67.
Fragariastrum haynaldianum (Janka) Kechaykin et Shmakov, comb. nov. = Potentilla haynaldiana Janka, 1872, Oesterr. Bot. Z. 22: 176.
Fragariastrum nitidum (L.) Kechaykin et Shmakov, comb. nov. = Potentilla nitida L. 1756, Cent. Pl. 2: 18.
Fragariastrum nivalis (Lapeyr.) Kechaykin et Shmakov, comb. nov. = Potentilla nivalis Lapeyr. 1782, Mem. Acad. Sci. Toulouse, 1: 210, t. 16.
Fragariastrum savvalensis (Pawl.) Kechaykin et Shmakov, comb. nov. = Potentilla savvalensis Pawl., Fragm. Florist. Geobot. 11: 69. 1965.
Fragariastrum saxifragum (Ardoino ex De Not.) Kechaykin et Shmakov, comb. nov. = Potentilla saxifraga Ardoino ex De Not. 1848, Index Seminum [Genoa]: 25.
Fragariastrum valderium (L.) Kechaykin et Shmakov, comb. nov. = Potentilla valderia L. 1759, Syst. Nat., ed. 10. 2: 1064.
10. Fragariastrum sect. Lupinoides (Tausch) Kechaykin et Shmakov, comb. nov. = Potentilla sect. Lupinoides Tausch, 1823, Hort. Canal., sine pag. - P. sect. Campestres (Poeverl.) Juz. 1941, USSR, 10: 91. - P. [unranked] Campestres Poeverl. 1898, Denkschr. Bayer. Bot. Ges. Regensburg, 7(1): 220.
Lectotype (Sojak, 1987: 495): Fragariastrum album (L.) Schur
Two species in Europe.
Fragariastrum album (L.) Schur, 1853, in Verh. Mittheil. Siebenburg. Ver. Naturw.
Hermannst. iv. Anhang (Sert. Fl.Transsilv.): 23. -Potentilla alba L. 1753, Sp. Pl.: 498.
Fragariastrum montanum (Brot.) Kechaykin et Shmakov, comb. nov. = Potentilla montana Brot. 1805, Fl. Lusit. 2: 350.
11. Fragariastrum sect. Plumosistylae (Pawlowski) Kechaykin et Shmakov, comb. nov.
= Potentilla sect. Plumosistylae Pawlowski, 1965, Fragm. Fl. Geobot. 11: 79.
Type: Fragariastrum libanotica (Boiss.) Ke-chaykin et Shmakov
Three species in Greece, Turkey, Lebanon and Syria.
Fragariastrum arcadiensis (Latrou) Kechaykin et Shmakov, comb. nov. = Potentilla arcadiensis Latrou, 1985, Candollea, 40(1): 121.
Fragariastrum isauricum (P. H. Davis) Kechaykin et Shmakov, comb. nov. = Potentilla libanotica Boiss. var. isaurica P. H. Davis, 1957, Notes Roy. Bot. Gard. Edinburgh, 22: 171. = Potentilla isaurica
(P. H. Davis) Pawl., 1965, Fragm. Florist. Geobot. 11: 81.
Fragariastrum libanoticum (Boiss.) Kechaykin et Shmakov, comb. nov. = Potentilla libanotica Boiss. 1843, Diagn. Pl. Nov. Orient., ser. 1, 3: 4.
12. Fragariastrum sect. Speciosae (Th. Wolf) Kechaykin et Shmakov, comb. nov. = Potentilla L. [unranked] Speciosae Th. Wolf, 1908, Biblioth. Bot. 16(71): 46, 85. = Potentilla sect. Speciosae (Th. Wolf) Juz. 1941, Fl. URSS, 10: 89.
Type: Fragariastrum speciosa (Willd.) Kechay-kin et Shmakov
Seven species in Europe (Balkans), Turkey, Iraq, and Lebanon.
Fragariastrum apenninum (Ten.) Kechaykin et Shmakov, comb. nov. = Potentilla apennina Ten. 1815, Cat. Hort. Neap.: 50.
Fragariastrum deorum (Boiss. et Heldr.) Kechaykin et Shmakov, comb. nov. = Potentilla deorum Boiss. et Heldr. 1856, Diagn. Pl. Orient. ser. 2, 2: 51.
Fragariastrum discipulorum (P. H. Davis) Ke-chaykin et Shmakov, comb. nov. = Potentilla dis-cipulorum P. H. Davis, 1957, Notes Roy. Bot. Gard. Edinburgh, 22: 168.
Fragariastrum divinum (Albov) Kechaykin et Shmakov, comb. nov. = Potentilla divina Albov, 1891, Otchet Trudy Odessk. Otdel. Ross. Obshch. Sadovod., Prilozh.: 103.
Fragariastrum kionaeum (Halacsy) Kechaykin et Shmakov, comb. nov. = Potentilla kionaea Halacsy, 1888, Verh. K.K. Zool.-Bot. Ges. Wien, 38: 751.
Fragariastrum owerinianum (Rupr. ex Boiss.) Kechaykin et Shmakov, comb. nov. = Potentilla oweriniana Rupr. ex Boiss. 1872, Fl. Orient. 2: 705.
Fragariastrum speciosa (Willd.) Kechaykin et Shmakov, comb. nov. = Potentilla speciosa Willd. 1799, Sp. Pl., ed. 4, 2(2): 1110.
Genus V. Horkelia Cham. et Schltdl. 1827, Linnea, 2(1): 26. - Potentilla L. sect. Horkelia (Cham. et Schltdl.) Baill. 1869, Hist. Pl. 1(6): 372. -Potentilla L. subgen. Horkelia (Cham. et Schltdl.) Jeps. 1925, Man. Fl. Pl. Calif.: 484. - Potentilla L. subgen. Horkelia (Cham. et Schltdl.) J. T. Howell, 1945, Leafl. W. Bot. 4(6): 176, comb. inval.
Type: Horkelia californica Cham. et Schltdl.
Ca. 20 species in western North America (west of USA and north-west of Mexico).
1. Horkelia sect. Capitatae (Rydb.) Ertter et Reveal, 2007, Novon, 17(3): 316. - H. sect.
Capitatae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 7, comb. inval. (sine cit. bas.). - H. [unranked] Capitatae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120, 122. - Potentilla L. sect. Capitatae (Rydb.) Crum, 1934, Leafl. W. Bot. 1(10): 100.
Type: Horkelia capitata Lindl. [=H. fusca Lindl. var. capitata (Lindl.) M. Peck].
Note. O. A. Stevens (1959) proposed some combinations at the rank of a section which, however, do not meet the requirements of ICN Art. 41.5 (McNeill et al., 2012) and are thus invalid.
2. Horkelia sect. Hispidulae Ertter et Reveal, 2007, Novon, 17(3): 316.
Type: Horkelia hispidula Rydb.
3. Horkelia sect. Horkelia. - H. sect. Californi-cae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 7, comb. inval. (sine cit. bas.). - H. [unranked] Califor-nicae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120, 121. - H. sect. Cuneatae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 7, comb. inval. (sine cit. bas.). - H. [unranked] Cuneatae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120, 121. -H. sect. Tenuilobae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 7, comb. inval. (sine cit. bas.). -H. [unranked] Tenuilobae Rydb. 1908, N. Amer. Fl. 22(3): 269, 275.
Type: Horkelia californica Cham. et Schlecht.
4. Horkelia sect. Parryae Ertter & Reveal, 2007, Novon, 17(3): 316.
Type: Horkeliaparryi Greene
5. Horkelia sect. Tridentatae (Rydb.) Kechaykin et Shmakov, comb. et stat. nov. = Horkelia Cham. et Schltdl. [unranked] Tridentatae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120, 122. - H. sect. Tridentatae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 7, comb. inval. (sine cit. bas.).
Type: Horkelia tridentata Torr.
Syn.: H. [unranked] Ambiguae A. Gray, 1865, Proc. Amer. Acad. Arts, 6: 530. - H. sect. Hirsutae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 7, comb. inval. (sine cit. bas.). - H. [unranked] Hirsutae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120, 123. - H. sect. Sericatae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 7, comb. inval. (sine cit. bas.). - H. [unranked] Sericatae Rydb.1898, Mem. Dept. Bot. Columbia Coll. 2: 120, 123.
Genus VI. Horkeliella (Rydb.) Rydb. 1908, N. Amer. Fl. 22(3): 269, 282. - Horkelia Cham.
et Schltdl. subgen. Horkeliella Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120.
Type: Horkeliella purpurascens (S. Watson) Rydb. [=Horkeliapurpurascens S. Watson].
Three species distributed in mountains of California.
Genus VII. Ivesia Torr. et A. Gray, 1858, Pacif. Railr. Rep. 6(3): 72. - Horkelia Cham. et Schltdl. subgen. Ivesia (Torr. et A. Gray) Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120. - Potentilla L. sect. Ivesia (Torr. et A. Gray) Baill. 1869, Hist. Pl. 1(6): 372. - Potentilla L. subgen. Ivesia (Torr. et A. Gray) Jeps. 1925, Man. Fl. Pl. Calif.: 484.
Lectotype (Rydberg, 1908: 283): Ivesia gordonii (Hook.) Torr. et A. Gray ('gordoni) [=Horkelia gordonii Hook.].
26 species distributed in the west of North America (west of USA and north-west of Mexico).
1. Ivesia sect. Comarella (Rydb.) Ertter et Reveal, 2007, Novon, 17(3): 316. - Comarella Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 19, 156.
Lectotype (Rydberg, 1908: 291): Ivesia multifoliolata (Torr.) D. D. Keck. [=Horkelia multifoliolata Torr., Comarella multifoliolata (Torr.) Rydb.].
2. Ivesia sect. Ivesia. - I. [unranked] Lycopodioides (Rydb.) Rydb. 1908, N. Amer. Fl. 22(3): 283, 286. - I. sect. Lycopodioides (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 8, comb. inval. (sine cit. bas.). - Horkelia Cham. et Schltdl. subgen. Ivesia (Torr. et A. Gray) Rydb. [unranked] Lycopodioides Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 121, 124.
Lectotype: Ivesia gordonii (Hook.) Torr. et A. Gray
3. Ivesia sect. Saxosae (Rydb.) Ertter et Reveal, 2007, Novon, 17(3): 317. - Horkelia Cham. et Schltdl. subgen. Ivesia (Torr. et A. Gray) Rydb. [unranked] Saxosae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120, 125. - Potentilla L. sect. Saxosae (Rydb.) B.C. Johnst. 1985, Phytologia, 57(4): 299. - P. sect. Saxosae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - I. sect. Setosae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 8, comb. inval. (sine cit. bas.). - I. [unranked] Setosae Rydb. 1908, N. Amer. Fl. 22(3): 283, 290.
Type: Ivesia saxosa (Lemmon ex Greene) Ertter [=Potentilla saxosa Lemmon ex Greene].
4. Ivesia sect. Unguiculatae (Rydb.) Ertter et Reveal, 2007, Novon 17(3): 317. - I. sect. Unguiculatae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 8, comb. inval. (sine cit. bas.). - Horkelia Cham. et Schltdl. subgen. Ivesia (Torr. et A. Gray) Rydb. [unranked] Unguiculatae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120, 124. - I. [unranked] Unguiculatae (Rydb.) Rydb. 1908, N. Amer. Fl. 22(3): 283, 284. - I. sect. Eremicae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 8. -Horkelia Cham. et Schltdl. subgen. Ivesia (Torr. et A. Gray) Rydb. [unranked] Eremicae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 120, 124. - I. [unranked] Eremicae (Rydb.) Rydb. 1908, N. Amer. Fl. 22(3): 283, 286.
Type: Ivesia unguiculata A. Gray.
Genus VIII. Potentilla L. 1753, Sp. Pl.: 495; id. 1754, Gen. Pl., ed. 5: 219.
Lectotype (Rydberg, 1908: 293): P reptens L., but see comment above; P. argentea L. is to be proposed as a conserved type.
About 300 species in Holarctics (Arctic and temperate zones) as well as mountain regions of Palaeo-and Neotropics.
1. Potentilla sect. Arenicolae (Rydb.) B. C. Johnst. 1985, Phytologia 57(4): 297. - P sect. Arenicolae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11 comb. inval. (sine cit. bas.). - P. [unranked] Arenicolae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 22, 31.
Lectotype (Johnston, 1985: 297): Potentilla newberryi A. Gray.
2. Potentilla sect. Asperrimae (Chevtaeva) Kurbatski, 2008, Sist. Zametki Mater. Gerb. Krylova Tomsk. Gosud. Univ. 99: 6. - P. ser. Asperrimae Chevtaeva, 1983, Izv. Akad. Nauk Tadzh. SSR, Biol. Nauk, 1(90): 16.
Type: Potentilla asperrima Turcz.
3. Potentilla sect. Aureae (Lehm.) Juz. 1941, Fl. URSS, 10: 197. - P. subtrib. Aureae Lehm. 1856, Revisio Potentillarum: 6. - P. sect. Aureae (Lehm.) O. Stevens, 1959, in North Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - P. [unranked] Aureae (Lehm.) Zimm. 1884, in Jahresber. k. k. Staats-Ober-Realschule Steyr, 14: 17, sine dignit. - P. [unranked] Aureae (Lehm.) Rydb. 1896, in Bull. Torrey Bot. Club 23(10): 394, sine dignit. - P. [unranked] Aureae vernales Th. Wolf, 1901, Potentillen-Studi-en, 1: 50, 123. - P. [unranked] Aureae (Lehm.) Th. Wolf, 1904, in Aschers. et Graebn., Syn. Mitteleur. Fl. 6(1): 671, 786.
Type: Potentilla aurea L.
4. Potentilla sect. Brevifoliae (Rydb.) B. C. Johnst. 1985, Phytologia 57(4): 295, 299. - P. sect. Brevifoliae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - P. [un-ranked] Brevifoliae Rydb. 1896, Bull. Torrey Bot. Club, 23: 306. - P. [unranked] Brevifoliae (Rydb.) Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 30.
Type: Potentilla brevifolia Nutt.
5. Potentilla sect. Chrysanthae (Lehm.) Juz. 1941, Fl. URSS, 10: 180. - P. subtrib. Chrysanthae Lehm. 1856, Revisio Potentillarum: 5. - P. [unranked] Chrysanthae (Lehm.) Zimm. 1884, in Jahres-Bericht der k. k. Staats-Ober-Realschule in Steyr, 14: 13, sine dignit. - P. [unranked] Aureae chrysanthae Th. Wolf, 1901, Potentillen-Studien, 1: 50, 123. - P. [unranked] Chrysanthae (Lehm.) Th. Wolf, 1904, in Aschers. et Graebn., Syn. Mitteleur. Fl. 6 (1): 671, 775.
Type: Potentilla chrysantha Trevir.
6. Potentilla sect. Concinnae (Rydb.) A. Nelson, 1909, New Man. Bot. Centr. Rocky Mt.: 255. - P. [unranked] Concinnae Rydb. 1896, Bull. Torrey Bot. Club, 23: 431. - P. sect. Concinnae (Rydb.) B. C. Johnst. 1985, Phytologia, 57(4): 299, stat. inval.
Type: Potentilla concinna Richardson.
7. Potentilla sect. Desertae (Chevtaeva) Kechaykin et Shmakov, comb. et stat. nov. = Potentilla L. ser. Desertae Chevtaeva, 1981, Izv. Akad. Nauk Tadzh. SSR, Biol. Nauk, 3(84): 22. -P. sect. Desertae (Chevtaeva) Chevtaeva, 1983, Izv. Akad. Nauk Tadzh. SSR, Biol. Nauk, 1(90): 17, comb. inval. (sine cit. bas.). - P. sect. Desertoria Kamelin, 1979, Komaroviana annua, 31: 116, nom. nud. et provis. - P. sect. Vvedenskyi M. M. Mirachmedova, 1998, Uzb. Biol. Zhurn. 3: 33, nom. inval. (sine descr. lat.).
Type: Potentilla desertorum Bunge
8. Potentilla sect. Dumosae Sojak, 1969, Preslia, 41: 349.
Type: Potentilla dumosa (Franch.) Hand.-Mazz.
9. Potentilla sect. Fasciculato-pilosae Kamelin, 2001, Fl. Vostochnoi Evropy, 10: 397, 437.
Type: Potentilla acaulis L.
10. Potentilla sect. Fragarioides (Lehm.) Juz. 1941, Fl. URSS, 10: 210. - P. 'trib.' Fragarioides Lehm. 1856, Revisio Potentillarum: 4. -
P. [unranked] Fragarioides (Lehm.) Th. Wolf, 1908, in Biblioth. Bot. 16(71): 52, 635.
Type: Potentilla fragarioides L.
11. Potentilla sect. Graciles (Rydb.) A. Nelson, 1899, Bull. Torrey Bot. Club, 26(9): 480. - P. [unranked] Graciles Rydb. 1897, Bull. Torrey Bot. Club, 24(1): 4. - P. sect. Graciles (Th. Wolf) Sojak, 1983, Cas. Nar. Muz. Praze, Rada Pnr. 152(1): 21, comb. inval. (sine cit. bas.). - P. sect. Candidae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - P. sect. Heptaphyl-lae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - P. sect. Longipe-dunculatae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - P. sect. Nuttallianae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - P. sect. Pectinisectae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - P. sect. Permolles (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.).
Type: Potentilla gracilis Douglas ex Hook.
12. Potentilla sect. Grandiflorae (Th. Wolf) Juz. 1941, Fl. URSS, 10: 179. - P. [unranked] Grandiflorae Th. Wolf, 1904, in Aschers. et Graebn., Syn. Mitteleur. Fl. 6(1): 671, 771.
Type: Potentilla grandiflora L.
13. Potentilla sect. Haematochroae (Th. Wolf) Schiman-Czeika, 1969, Fl. Iran. 66: 91. - P. [unranked] Haematochroae Th. Wolf, 1904, in Aschers. et Graebn., Syn. Mitteleur. Fl. 6(1): 670, 702. -P. sect. Haematochri (Rydb.) B.C. Johnst. 1985, Phytologia, 57(4): 296, 300. - P. [unranked] Haematochri Rydb. 1897, Bull. Torrey Bot. Club, 24(1): 10. - P. sect. Rubrae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). -P. [unranked] Rubrae Rydb. 1908, N. Amer. Fl. 22(4): 293, 323.
Type: Potentilla hamaetochrus Lehm.
14. Potentilla sect. Heterosepalae (Rydb.) B. C. Johnst. 1985, Phytologia, 57(4): 293, 297. - P. sect. Heterosepalae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - P. [unranked] Heterosepalae Rydb. 1898, Mem. Dep. Bot. Columbia Univ. 2: 21, 23.
Type: Potentilla heterosepala Fritsch
15. Potentilla sect. Leucophyllae (Rydb.) A. Nelson, 1909, New Man. Bot. Centr. Rocky Mt.:
255, 256. - P. [unranked] Leucophyllae Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 22, 31. - P sect. Hippianae (Rydb.) Ertter et Reveal, Novon, 17(3): 317. - P. [unranked] Hippianae Rydb. 1897, Bull. Torrey Bot. Club, 24(1): 1.
Type: Potentilla leucophylla Torr. 1827, not Pallas, 1776 [=P hippiana Lehm.].
16. Potentilla sect. Multifidae (Lehm.) A. Nelson, 1909, in Coulter et Nelson, New Man. Bot. Centr. Rocky Mount.: 255. - P. trib. Multifidae Lehm. 1856, Revisio Potentillarum: 6. - P. [unranked] Multifidae (Lehm.) Rydb. 1896, in Bull. Torrey Bot. Club 23 (7): 262. - P. [unranked] Multifidae (Lehm.) Th. Wolf, 1904, in Aschers. et Graebn., Syn. Mitteleur. Fl. 6(1): 670, 698. - P. [unranked] Pensylvanicae Poeverl. 1904, in Aschers. et Graebn., Syn. Mitteleur. Fl. 6(1): 669 (as "Pennsyl-vanicae"). - P. sect. Adenocarpae Kurbatsky, 1985, Sist. Zametki Mater. Gerb. Krylova Tomsk. Gosud. Univ. Kuybysheva, 87: 1.
Type: Potentilla multifida L.
17. Potentilla sect. Multijugae (Rydb.)A. Nelson, 1909, New Man. Bot. Centr. Rocky Mt.: 255, 226. -P. [unranked] Multijugae Rydb. 1896, Bull. Torrey Bot. Club, 23: 432. - P. sect. Candicantes (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - P. [unranked] Candicantes Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 22, 32.
Type: Potentilla multijuga Lehm.
18. Potentilla sect. Niveae (Lehm.) A. Nelson, 1909, in Coulter et Nelson, New Man. Bot. Centr. Rocky Mount.: 255. - P. 'subtrib.' Niveae Lehm. 1856, Revisio Potentillarum: 8. - P. [unranked] Niveae (Lehm.) Rydb. 1896, Bull. Torrey Bot. Club 23 (8): 301. - P. [unranked] Canescentes boreales Th. Wolf, 1903, Potentillen-Studien, 2: 13, 68. -P. [unranked] Niveae (Lehm.) Th. Wolf, 1904, in Ascherson et Graebner, Syn. Mitteleur. Fl. 6(1): 670, 703.
Type: Potentilla nivea L.
19. Potentilla sect. Persicae (Th. Wolf) Juz. 1941, Fl. URSS, 10: 175. - P. 'trib.' Persicae Th. Wolf, 1906, in Bornm., Bull. Herb. Boissier, 6, ser. 7(8): 612. - P. sect. Butkovii M. M. Mirachmedova, 1998, Uzb. Biol. Zhurn. 3: 33, nom. inval. (sine de-scr. lat.).
Type: Potentilla persica Boiss. et Hausskn.
20. Potentilla sect. Potentilla. - P. sect. Terminales (Döll) Gren. et Gord. 1848, Fl. France, 1: 522,
532. - P. [unranked] Terminales Döll, 1843, Rhein. Fl.: 772, sine dignit. - P. sect. Argenteae (Lehm.) Juz. 1941, Fl. URSS, 10: 142. - P. 'subtrib.' Argenteae Lehm. 1856, Revisio Potentillarum: 6. - P. [unranked] Argenteae (Lehm.) Zimm. 1884, in JahresBericht der k. k. Staats-Ober-Realschule in Steyr, 14: 9, sine dignit. - P. [unranked] Argenteae (Lehm.) Rydb. 1898, in Mem. Dep. Bot. Columbia Univ. 2: 21, 24. - P. [unranked] Canescentes Th. Wolf, 1901, Potentillen-Studien, 1: 21, 122. - P. [unranked] Argenteae (Lehm.) Th. Wolf, 1908, in Biblioth. Bot. 16 (71): 49, 252. - P. sect. Fedtschenkoana M. M. Mirachmedova, 1998, Uzb. Biol. Zhurn. 3: 32, nom. inval. (sine descr. lat.).
Lectotype: Potentilla reptans L.
21. Potentilla sect. Ranunculoides (Th. Wolf) Juz. 1941, Fl. URSS, 10: 193. - P. [unranked] Ranunculoides Th. Wolf, 1908, Biblioth. Bot. 16(71): 51, 503.
Type: Potentilla ranunculoides Humb. et Bonpl. ex Nestl.
22. Potentilla sect. Rectae (Lehm.) Juz. 1941, Fl. URSS, 10: 160. - P. 'subtrib.' Rectae Lehm. 1856, Revisio Potentillarum: 5. - P. sect. Rectae (Lehm.) O. Stevens, 1959, in North Amer. Fl. 22 (7): 11, comb. inval. (sine cit. bas.). - P. [unranked] Rectae (Lehm.) Th. Wolf, 1901, Potentillen-Studien, 1: 39, 122.
Type: Potentilla recta L.
23. Potentilla sect. Rubricaules (Rydb.) A. Nelson, 1909, New Man. Bot. Centr. Rocky Mt.: 255, 256. - P. sect. Rubricaules (Rydb.) O. Stevens, 1959, in North Amer. Fl. 22 (7): 11, comb. inval. (sine cit. bas.). - P. [unranked] Rubricaules Rydb. 1898, Mem. Dep. Bot. Columbia Univ. 2: 21, 30.
Type: Potentilla rubricaulis Lehm.
24. Potentilla sect. Subjugae (Rydb.) A. Nelson, 1909, New Man. Bot. Centr. Rocky Mt.: 255. -P. sect. Subjugae (Rydb.) O. Stevens, 1959, in North Amer. Fl. 22 (7): 11, comb. inval. (sine cit. bas.). - P. sect. Subjugae (Rydb.) B.C. Johnst. 1985, Phytologia, 57(4): 296, 299, comb. inval. -P. [unranked] Subjugae Rydb. 1896, Bull. Torrey Bot. Club, 23: 397.
Type: Potentilla subjuga Rydb.
25. Potentilla sect. Subviscosae (Rydb.) B. C. Johnst. 1985, Phytologia 57(4): 295, 299. (Rydb.). -P. sect. Subviscosae (Rydb.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). -
P [unranked] Subviscosae Rydb. 1896, Bull. Torrey Bot. Club 23: 429.
Type: Potentilla subviscosa Greene.
26. Potentilla sect. Supinae (Lehm.) A. Nelson, 1909, in Coulter et Nelson, New Man. Bot. Centr. Rocky Mount.: 255. - P. ser. Supinae Lehm. 1856, Revisio Potentillarum: 8. - P. [unranked] Supinae (Lehm.) Rydb. 1898, Mem. Dep. Bot. Columbia Univ. 2: 21, 23. - P. sect. Rivales (Th. Wolf) Juz. 1941, Fl. URSS, 10: 165. - P. [unranked] Rivales Th. Wolf, 1903, Potentillen-Studien, 2: 11, 68.
Type: Potentilla supina L.
27. Potentilla sect. Tanacetifoliae (Lehm.) Juz. 1941, Fl. URSS, 10: 152. - P. 'trib.' Tanacetifoliae Lehm. 1856, Revisio Potentillarum: 6. - P. [unranked] Tanacetifoliae (Lehm.) Th. Wolf, 1904, in Ascherson et Graebner, Syn. Mitteleur. Fl. 6 (1): 670,743.
Type: Potentilla tanacetifolia Willd. ex D. F. K. Schltdl.
Genus IX. Purpusia Brandegee, 1899, Bot. Gaz. 27: 446. - Potentilla L. subgen. Purpusia (Brandegee) J. T. Howell, 1945, Leafl. W. Bot. 4(6): 172.
Lectotype (Rydberg, 1908: 291): Purpusia saxosa Brandegee
Three species distributed in arid regions of southwest of USA.
Genus X. Stellariopsis (Baill.) Rydb. 1898, Mem. Dept. Bot. Columbia Coll. 2: 19, 155. -Potentilla L. sect. Stellariopsis Baill. 1869, Hist. Pl. 1: 372. - P. sect. Stellariopsis (Baill.) O. Stevens, 1959, N. Amer. Fl. 22(7): 11, comb. inval. (sine cit. bas.). - Ivesia Torr. et A. Gray sect. Stellariopsis (Baill.) Ertter et Reveal, 2007, Novon 17(3): 317.
Type: Stellariopsis santolinoides (A. Gray) Rydb. [=Ivesia santolinoides A. Gray].
Monotypic genus occurring in mountains of California.
Genus XI. Tormentilla L. 1753, Sp. Pl.: 500.
Lectotype (Green, in Hitchcock et Green, 1929: 159): Tormentilla erecta L.
Eight species native to Northern and introduced to Southern Hemisphere.
1. Tormentilla L. sect. Reptantes (Lehm.) Kechaykin et Shmakov, comb. et stat. nov. = P. ser. Reptantes Lehm. 1856, Revisio Potentillarum: 8.
Type: Tormentilla linnaeana Kechaykin et Shmakov [=Potentilla reptans L.] (Herb. Linn. No. 655.38, LINN).
Six species in Northern Hemisphere:
T. canadensis (L.) Kechaykin et Shmakov, comb. nov. = Potentilla canadensis L. 1753, Sp. Pl.: 498.
T. flagellaris (Willd ex D. F. K. Schltdl.) Kechaykin et Shmakov, comb. nov. = Potentilla flagellaris Willd ex D. F. K. Schltdl. 1816 in Ges. Naturf. Freunde Berlin (Mag.) 7: 291.
T. hemsleyana (Th. Wolf) Kechaykin et Shmakov, comb. nov. = Potentilla hemsleyana Th. Wolf, 1908, Biblioth. Bot. 16(71): 667.
T. linnaeana Kechaykin et Shmakov, nom. nov. = Potentilla reptans L. 1753, Sp. Pl.: 499, non T. reptans L. (1753: 500).
T. simplex (Michx.) Kechaykin et Shmakov, comb. nov. = Potentilla simplex Michx. 1803, Fl. Bor.-Amer. 1: 303.
T. simulatrix (Th. Wolf) Kechaykin et Shmakov, comb. nov. = Potentilla simulatrix Th. Wolf, 1908, Biblioth. Bot. 16(71): 663.
2. Tormentilla L. sect. Tormentilla. - Potentilla L. sect. Tormentilla (L.) Tausch, 1823, in Hortus Canal. 1: t. 10 (unpaged). - P. ser. Tormentillae Lehm. 1856, Revis. Potentill.: 8. - P. [unranked] Tormentillae Rydb. 1898, Mem. Dep. Bot. Columbia Univ. 2: 21, 22, nom. inval. - P. [unranked] Tormentillae Th. Wolf, 1901, Potentillen-Studien, 1: 99, 123, nom. inval.
Lectotype: Tormentilla erecta L. (Herb. Linn. No. 656.1, LINN).
Two species native to Northern and introduced to Southern Hemisphere.
T. erecta L. 1753, Sp. Pl.: 500. - Potentilla tormentilla Neck. 1770, Hist. et Commentat. Acad. Elect. Sci. Theod.-Palat. 2: 491.
T. reptans L. 1753, Sp. Pl.: 500. - Potentilla anglica Laichrd. 1790, Veg. Europ., ed. 1: 475. -P. procumbens Sibth. 1794, Fl. Oxon.: 162.
Genus XII. Tylosperma Botsch. 1952, Not. Syst. Herb. Inst. Bot. Acad. Sci. Uzbekistan,
13: 17.
Type: Tylosperma lignosa (Willd. ex D. F. K. Schltdl.) Botsch. [=Potentilla lignosa Willd. ex D. F. K. Schltdl.]
Two species distributed in Turkey, Iran, Afghanistan, Pakistan, India, and Middle Asia.
Acknowledgements
This study was supported by the Russian Foun-datin for Basic Research (project 14-04-00721). We are grateful to colleagues for translation of the text to English and valuable comments on the contents.
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