Solanum dulcamara L., 1753

Knapp, Sandra, 2013, A revision of the Dulcamaroid Clade of Solanum L. (Solanaceae), PhytoKeys 22, pp. 1-432 : 96-117

14. Solanum dulcamara L. , Sp. Pl. 185. 1753 Figure 35

Solanum scandens Neck., Delic. Gallo-Belg. 1: 119. 1768. Type: Based on Solanum dulcamara L.

Dulcamara lignosa Gilib., Fl. Lit. Inch. 1: 37. 1782. Type: Based on Solanum dulcamara L.

Lycopersicon dulcamara (L.) Medik., Beobacht. 245. 1783. Type: Based on Solanum dulcamara L.

Solanum rupestre F.W.Schmidt, Fl. Boem. 2: 96, tab. 241. 1793. Type: Czech Republic. "Termas Carolinas, Dorotheenau" [Karoly Vary] (no specimens found, plates unpublished; synonymy ex descr.).

Dulcamara flexuosa Moench, Meth. 514. 1794, nom. illeg. superfl. Type: Based on Solanum dulcamara L.

Solanum ruderale Salisb., Prodr. Stirp. Chap. Allerton 133. 1796, nom. illeg. superfl. Type: Based on Solanum dulcamara L.

Solanum dulcamara L. var. villosissimum Desv., Observ. Fl. Angers 111. 1818. Type: France. Anjou: environs de Brissac, N.A. Desvaux s.n. (possible type specimen: P [P00582490]).

Solanum dulcamara L. var. rupestre (H.W.Schmidt) Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 581. 1819. Type: Based on Solanum rupestre F.W.Schmidt

Solanum persicum Willd., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 662. 1819. Type: "In Persia, Pallas "(neotype, designated here: BM [BM000942806]).

Solanum littorale Raab, Flora 2: 414. 1819. Type: Switzerland. Vaud: Lac Leman "inter Vidy et Pully; prope Ouchy et Champlande", C.W. Raab s.n. (no specimens found; synonymy ex descr.).

Solanum kieseritzkii C.A.Mey., Verz. Pfl. Cauc. 113. 1831. Type: Azerbaijan. Lenkoran, 23 May 1830, C.A. Meyer s.n. (holotype: LE; isotypes: LE (3 sheets), OXF).

Solanum assimile Friv., Flora 19: 439. 1836. Type: Greece. "Turkey in Rumelia", I. Frivaldszky s.n. (holotype: BP [n.v.]; isotype: NY [NY00169744]).

Solanum dulcamara L. var. tomentosum W.D.J.Koch, Syn. Fl. Germ. Helv. 2: 508. 1837. Type: Based on Solanum littorale Raab

Solanum dulcamara L. var. album G.Don, Gen. Hist. 4: 409. 1838. Type: "Dulcis amara flor albo", Besler, Hortus Eystettensis, fol. 46, t. ii. 1613. (lectotype, designated here: Besler, Hortus Eystettensis, fol. 46, t. ii. 1613 [illustration on p. 180 in reprints]).

Solanum dulcamara L. var. carneum G.Don, Gen. Hist. 4: 409. 1838. Type: Sweden. "Celsius, Ups. 32, Linnaeus, Fl. Suec. p. 66" (no specimens found).

Solanum dulcamara L. var. hirsutum G.Don, Gen. Hist. 4: 409. 1838. Type: "Plant hairy, or downy. Flowers violaceous. On the sea-coast." (no specimens cited).

Solanum dulcamara L. var. violaceum G.Don, Gen. Hist. 4: 409. 1838. Type: "Dulcis amara flor coeruleo vulgatica", Besler, Hortus Eystettensis, fol. 46, t. iii. (lectotype, designated here: Besler, Hortus Eystettensis, fol. 46, t. iii on folio 46 t. iii. 1613 [illustration on p. 180 in reprints]).

Solanum dulcamara L. var. cordifolium Peterm., Fl. Lips. Excurs. 173. 1838. Type: Germany. Sin. loc. (no specimens found).

Solanum dulcamara L. var. ovatum Peterm., Fl. Lips. Excurs. 173. 1838. as ovata. Type: Germany. Sin. loc. (no specimens found).

Solanum dulcamara L. var. marinum Bab., Man. Brit. Bot. 210. 1843. Type: Ireland. County Galway: Connemara, Renvyle [ “Renville”] (lectotype, designated here: CGE; isolectotype: BM [BM000941486]).

Dulcamara flexuosa Moench var. cordata Opiz, in Bercht. & Opiz, Oekon.-techn. Fl. Böhm. 3: xv. 1843. Type: Based on Solanum dulcamara L. var. cordifolium Peterm.

Dulcamara flexuosa Moench var. ovata Opiz, in Bercht. & Opiz, Oekon.-techn. Fl. Böhm. 3: xv. 1843. Type: Based on Solanum dulcamara L. var. ovatum Peterm.

Dulcamara flexuosa Moench var. hastiifolia Opiz, in Bercht. & Opiz, Oekon.-techn. Fl. Böhm. 3: xvi. 1843. as hastaefolia. Type: Czech Republic [?]. sin. loc., P.M. Opiz s.n. (holotype: PR?).

Solanum lignosum Sloboda, Rostlinnictví 358. 1852. Type: "Dulcamara Marina, Solanum lignosum Rayi" (no specimens found).

Solanum dulcamara L. subsp. cordatum (Opiz) Dunal, Prodr. [A.P. de Candolle] 13(1): 78. 1852. Type: Based on Solanum dulcamara L. var. cordifolium Peterm.

Solanum dulcamara subsp. ovatum (Peterm.) Dunal, Prodr. [A.P. de Candolle] 13(1): 78. 1852. Type: Based on Solanum dulcamara L. var. ovatum Peterm.

Solanum dulcamara L. var. rupestre (F.W.Schmidt) Dunal, Prodr. [A.P. de Candolle] 13(1): 79. 1852. Type: Based on Solanum rupestre F.W.Schmidt

Solanum dulcamara L. var. palustre Dunal, Prodr. [A.P. de Candolle] 13(1): 79. 1852. Type: Greece. “Lernacicis”, J. Sartori s.n. (no specimens found, see discussion).

Solanum dulcamara L. var. hirsutum Dunal, Prodr. [A.P. de Candolle] 13(1): 79. 1852. Type: Greece. “Lernacicis”, J. Sartori 84 (lectotype, designated here: G-DC [G00144899]).

Solanum dulcamara L. var. laciniatum Dunal, Prodr. [A.P. de Candolle] 13(1): 78. 1852. Type: United States of America.Massachusetts: Boston, 1827, Mrs. Dutton s.n. (lectotype, designated here: G-DC [G00144856]).

Solanum dulcamara L. subsp. hastiifolia (Bercht. & Opiz) Dunal, Prodr. [A.P. de Candolle] 13(1): 78. 1852. Type: Based on Dulcamara flexuosa Moench var. hastiifolia Opiz in Bercht. & Opiz

Solanum dulcamara L. var. alpinum Schur, Enum. Pl. Transsilv. 478. 1866. Type: Romania. (Transylvania) Siebenbürgen [Sibiu], "Arpaser-Kerzesor Alpen" [Arpas], Jul-Aug, J.F. Schur s.n. (holotype: GOET?, not found).

Solanum dulcamara L. var. macrocarpum Maxim., Index Seminum [St. Petersburg] suppl. 1869: 26. 1870. Type: japan. Jezo Island: Hakodate, 1861, C. Maximowicz s.n. (lectotype, designated here: LE; isolectotypes: G [G00301664], LE).

Solanum dulcamara L. var. integrifolia Willk. in Willk. & Lange, Prodr. Fl. Hispan. 2: 526. 1870. Type: Spain. Aragón: Jaca, Jun 1850, H.M. Willkomm 313 (lectotype, designated here: BM [BM000847923]; isolectotype: G [G00357859]).

Solanum dulcamara L. var. indivisum Boiss., Fl. Orient. 4: 285. 1879. Type: Based on Solanum persicum Willd.

Solanum dulcamarum St.- Lag., Ann. Soc. Bot. Lyon, 7: 135. 1880, nom. illeg. superfl. Type: Based on Solanum dulcamara L.

Solanum dulcamara L. forma subglabrum Kuntze, Trudy Imp. S.-Peterburgsk. Bot. Sada 10: 222. 1887. Type: Turkmenistan. Asgabat [ As’chabad], May 1886, O. Kuntze [Gen. Komarov] s.n. (holotype: B, destroyed; lectotype, designated here: NY [NY00172263]).

Solanum rupestre Waisb., Oesterr. Bot. Z. 45: 143. 1895, non Solanum rupestre F.W.Schmidt, 1793. Type: Austria. "ad pagum Rödlschlag solo serpentino", 700-750m, Jun-Jul 1894, A. Waisbecker s.n. (no specimens found, holotype: BP?).

Solanum dulcamara subvar. villosissimum (Desv.) Bornm., Beih. Bot. Centralbl. 33: 305. 1915. Type: Based on Solanum dulcamara L. var. villosissimum Desv.

Solanum macrocarpum (Maxim.) Kudô, in Kudô & Susaki, Hokkaido-Yakuyo-Shokub t. 84. 1922. Type: Based on Solanum dulcamara L. var. macrocarpum Maxim.

Solanum dulcamara L. var. canescens Farw., Papers Mich. Acad. Sci. 2: 39. 1923. Type: United States of America. Michigan: Rochester, 15 Aug 1909, O.A. Farwell 2105 (holotype: BLH [n.v.]).

Solanum dulcamara L. forma albiflora Farw., Papers Mich. Acad. Sci. 2: 39. 1923. Type: United States of America. Michigan: Trenton, 26 Jul 1921, O.A. Farwell 5944 (holotype: BLH [n.v.]).

Solanum dulcamara L. forma albiflorum House, Bull. New York State Mus. 254: 613. 1924. Type: United States of America. New York: Livingston County, Caledonia, F. Beckwith s.n. (holotype: NYS [n.v.]).

Solanum maximowiczii Koidz., in Mayebara, Fl. Austrohigo 50, 85. 1931. Type: Based on Solanum dulcamara L. var. ovatum Peterm.

Solanum macrocarpum (Maxim.) Koidz., Acta Phytotax. Geobot. 1: 23. 1932, nom. illeg., non Solanum macrocarpum (Maxim.) Kudô Type: Based on Solanum dulcamara L. var. macrocarpum Maxim.

Solanum megacarpum Koidz., Acta Phytotax. Geobot. 4: 159. 1935. Type: Based on Solanum dulcamara L. var. macrocarpum Maxim.

Solanum depilatum Kitag., Lin. Fl. Manshur. 390. 1939. Type: China. [Heilongjiang]: “Chéjché, Manchur. bor.", M. Kitagawa s.n. (type not located).

Solanum asiaemediae Pojark., Not. Syst. Herb. Inst. Bot. Acad. Sci. URSS 17: 330. 1955. Type: Uzbekistan. Kokaidskii region, Tamariskovii Krai, [lower Najman-Saj River], 21 Jul 1934, A.V. Prozorovskii 187 (holotype: LE).

Solanum pseudopersicum Pojark., Not. Syst. Herb. Inst. Bot. Acad. Sci. URSS 17: 328. 1955. Type: Russian Federation. Starvropolskii Krai: Pyatigorsk, Mashukh [protologue Ciscaucasia], 3 Jul 1896, D. Litvinov s.n. (holotype: LE).

Solanum marinum (Bab.) Pojark., in Komarov, Fl. URSS, 22: 16. 1955. Type: Based on Solanum dulcamara L. var. marinum Bab.

Solanum persicum Roem. & Schult. subsp. pseudopersicum (Pojark.) Schönb.-Tem., Fl. Iranica 100: 14: 1972. Type: Based on Solanum pseudopersicum Pojark.

Solanum kitagawae Schönb.-Tem., Fl. Iranica 100: 15. 1972. Type: Based on Solanum depilatum Kitag.

Solanum dulcamara L. luc. atroviolaceum Máthé, Bot. Közlem. 59(2): 129. 1972. Type: Hungary. Nógród: "ad pagum Nótines”, I. Máthé s.n. (holotype: VBI [n.v.]).

Solanum dulcamara L. forma lucidum Máthé, Bot. Közlem. 59(2): 129. 1972. Type: Hungary. Pest: "ad viam pagi Vácrátot”, I. Máthé s.n. (holotype: VBI [n.v.]).

Solanum dulcamara L. var. pusztarum Soó, Acta Bot. Acad. Sci. Hung. 18(1-2): 172. 1973. Type: Hungary. Mittelungarn, Kom. Bács-Kiskun, Bugacpuszta, R. Sóo s.n. (holotype: BP [n.v.]).

Solanum dulcamara subsp. pusztarum ( Soó) Soó, Acta Bot. Acad. Sci. Hung. 23(3-4): 382. 1978 [ “1977”]. Type: Based on Solanum dulcamara L. var. pusztarum Soó

Solanum borealisinense C.Y.Wu & S.C.Huang, Fl. Reipubl. Popul. Sin. 67(1): 84, tab. 21, Figure 1-7. 1978. Type: Based on Solanum depilatum Kitag.

Type.

“Europa”, Anon. (lectotype, designated by Knapp and Jarvis 1990, pg. 335: LINN 248.7).

Description.

Herbaceous or woody vine, above ground stems trailing to 8-10 m long and with spreading or creeping underground stems. Stems very rarely glabrous, more often pubescent with simple uniseriate or dendritic trichomes with short branches, or a mixture of the two types, these often tangled, 4-8-celled, to 1.5 mm long, usually white, pubescence density extremely variable; new growth usually white pubescent. Bark of older stems grey to yellowish grey. Sympodial units plurifoliate. Leaves simple to ternately pinnatifid, extremely variable in shape and size, even along a single stem, 2.5-7 cm long, 1.2-6 cm wide, elliptic or ovate to cordate in outline, membraneous, the upper surfaces glabrous to moderately pubescent with simple uniseriate or dendritic trichomes to 1.5 mm long on the veins and lamina, the lower surfaces sparsely to densely pubescent with trichomes like those of the upper surfaces, but usually denser; primary veins 6-9 pairs, usually pubescent; base truncate or cordate; margins entire or the leaves lobed, the lobes most commonly 2, rarely more, basal, narrowing near the sinuses; apex acute to acuminate; petiole 0.5-2 (+) cm long, pubescent like the stems. Inflorescences terminal or lateral, not leaf-opposed, (1-)4-15 cm long, many times branched, with up to 40 flowers, only a few open at a time, glabrous to moderately pubescent, the rachis often purplish in hue; peduncle (0.5-)1-7 cm long; pedicels 6-12 mm long at anthesis, ca. 1 mm in diameter, slender, spreading, often purplish green, glabrous to sparsely pubescent with simple unieriate or more rarely dendritic trichomes to 0.5 mm long, articulate at the base in a small sleeve leaving a prominent swollen peg on the axis; pedicel scars irregularly spaced 1-5 (-10) mm apart, the axis zig-zag. Buds turbinate, the corolla long-exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 1-1.5 mm long, broadly conical, the lobes <0.5 mm long, broadly triangular, glabrous or pubescent with uniseriate white trichomes, the apex pointed, the margins papillate. Corolla 1.5-2 cm in diameter, purple, violet or white, with green and white shiny spots at each lobe base, deeply stellate, lobed 3/4 of the way to the base, the lobes 6-8 mm long, 2.5-3 mm wide, strongly reflexed at anthesis, glabrous or minutely papillate on tips and margins, occasionally densely pubescent with simple uniseriate trichomes abaxially, glabrous adaxially. Filament tube minute, the free portion of the filaments to 0.5 mm long, glabrous; anthers 4.5-6 mm long, ca. 1 mm wide, fused into a single column and tightly connivent, poricidal at the tips, the pores not lengthening to slits with age. Ovary glabrous; style 5-9 mm long, glabrous; stigma minutely capitate, the surface papillose. Fruit a globose to ellipsoid berry, 0.6-1.1 cm long, 0.6-1.5 cm wide, bright red when ripe, the pericarp thin and shiny; fruiting pedicels to 1.3 cm long, 1-1.5 mm in diameter, not markedly woody, spreading. Seeds>30 per berry, ca. 3 mm long, ca. 2 mm wide, flattened reniform, pale yellow or tan, the surfaces minutely pitted, the cells of the testa pentagonal. Chromosome number: n=12 (many counts available at ICPN, http://mobot.mobot.org/W3T/Search/ipcn.html)

Distribution

( Figure 36 ). Solanum dulcamara is widely distributed across Eurasia and northern North America, where it is also common; sea level to ca. 2000 m. The North American populations are thought to be introductions, but it is possible that the species has a truly circumboreal distribution.

Ecology.

Solanum dulcamara is a weedy species and grows in a wide variety of temperate habitats, often associated with water and open places with abundant light. Although somewhat woody, it rarely reaches into the canopy but is more often found in thickets and sprawling in other low vegetation.

Common names.

Douce-amére (French); woody nightshade, bittersweet (English speaking Europe)

Conservation status.

Least Concern (LC); EOO>100,000 km2 (LC) and AOO>10,000 km2 (LC). See Moat (2007) for explanation of measurements. Solanum dulcamara has a circumboreal distribution and is very common so it is not of conservation concern. Peripheral populations such as those in eastern Russia, however, may harbour interesting genetic variation (see Knapp 2011).

Discussion.

Solanum dulcamara is one of the most variable species in the Dulcamaroid group, all of which show extremes of phenotypic plasticity. It is also the most widespread, and is found throughout the northern hemisphere, in Europe and Asia from Spain to Siberia and into extreme northern Japan, and across the northern United States (reported from Florida on the PLANTS database [http://plants.usda.gov/java/profile?symbol=SODU], from where I have seen no specimens), where it is presumed to have been introduced from Europe. I can find no documentary evidence for the introduction of Solanum dulcamara into the New World, but early collections from North America held in BM (e.g., Bartram, Clayton) do not contain specimens of this species. The earliest collections I have seen are from the Boston area in the early 19th century (e.g., Dutton s.n.); it is possible that Solanum dulcamara was introduced to North America for its medicinal properties.

Some of the variation in Solanum dulcamara appears to be related to habitat (e.g., coastal forms with succulent leaves that have been recognised as var. marinum and Solanum littorale ) but much of the variation is in leaf size and division, pubescence and flower color; I have been able to find no pattern related to either geography or habitat in this variation. Turesson (1922) suggested that differences in morphology related to habitat were genetic, despite his finding that in non-coastal environments thick-leaved coastal forms developed thinner, less pubescent leaves. Sun and shade leaves of Solanum dulcamara have been studied for physiological and some morphological properties (Clough et al. 1979a, b). Differences in morphology can occur along a single stem depending upon the shade environment. Unlike some other species in the Dulcmaroid clade (e.g., Solanum umbelliferum ) variation does easily sort into distinguishable units, but the sheer range and complexity of the variation has led to the description of many taxa within what I recognise here as a single highly polymorphic species, especially in floristic treatments from Central Asia ( Pojarkova 1955; Schönbeck-Temesy 1972), where much of the variation exists.

Chemical variation in Solanum dulcamara has also been intensively studied in eastern Europe ( Máthé and Máthé 1972, 1979); populations with pubescent individuals from the northen part of the eastern European range tended to have higher percentages of the alkaloid aglycone tomatidinol while more glabrous populations had higher concentrations of soladulcinine. The concentrations of various alkaloids were affected by the environmental conditions under which plants were grown ( Máthé et al. 1975) as well as by what were presumed to be genetic (ecotypic) factors ( Bernáth and Tétényi 1977). Clough et al. (1979a) and Pegtel (1985) both suggested that ecotypic differentiation did not explain the variation observed, but instead that the variability was the result of short-term phenotypic acclimatization to differing environments. A study using AFLP (Associated Fragment Length Polyomorphisms) data from Europe-wide accessions of Solanum dulcamara found almost no genetic variation or population structuring either geographically or in association with habitat ( Golas et al. 2010).

A few of the more extreme forms have recognised at the species level, but the variation in the diagnostic features of these forms is continuous across the range of the species. Identifications of individual specimens by botanists describing these variants were occasionally inconsistent from duplicate to duplicate.

Plants occurring in the swampy areas around the hot springs at Lenkoran in what is now Azerbaijan were described as Solanum kieseritzkii ; all of the collections from this locality are composed of small erect shoots connected with creeping stems and have very small inflorescences of only a few flowers.

Glabrous plants with simple leaves have been recognised as Solanum pseudopersicum ; glabrous plants with divided leaves as S. kitagawae.

Pubescent plants with simple leaves were described as Solanum persicum ; these tended to be from the southern part of the species range in Eurasia, but occur throughout the species’ range.

Large-fruited plants from the eastern margins of the Asian range have been called var. macrocarpum; these fruits also tend to be more ellipsoid than globose, but as with other chracters, this occurs sporadically throughout the range of the species.

White-flowered plants have been given varietal or subspecific status both in Europe and the United States; polymorphism in flower color is common in Solanum in general.

Solanum dulcamara has a long history of use in medicine in Europe and in the United States. Gerard (1597:350) says "The leaves and fruit of the Bitter-sweet are in temperature hot and dry, clensing and wasting away" and cites as its “Vertues” effects on liver and spleen, against jaundice and bruising, and of its use to help breathing difficulties and as a restorative after childbirth. The common name bittersweet (or douce-amère) comes from the Latin ‘Amaradulcis’, and refers to the fact that when chewed, the twigs taste at first bitter, then sweet ( Grieve 1931). Bittersweet had many uses ranging from inflammatory diseases of all kinds (such as asthma and rheumatism) to nymphomania and syphilis (Dunal 1813; Felter and Lloyd 1898); Grieve (1931: 590) states "there are few complaints for which it has not at some time been recommended." As ‘Dulcamara’ or ‘Dulc’ ( van Zandvoort 1996), the stems are used still in homeopathic medicine ( Bharatan et al. 2002). It is today listed as a poisonous plant in many countries and US states (see for example the FDA at http://www.accessdata.fda.gov). Michel-Félix Dunal treated the uses and superstitions associated with this species (Dunal 1813) and concluded that the many properties attributed to bittersweet were mostly not substantiated in fact. The steroidal alkaloids responsible for the active properties of Solanum dulcamara were not discovered until the late 19th century, they are principally dulcamarine and solanine, the first of which is responsible for the bittersweet taste and the second for the narcotic effects ( Grieve 1931).

Solanum dulcamara has been thought to be an intermediate host and possible source of primary infections for several agronomically important potato diseases such as late blight (caused by the oomycete Phytophthora infestans (Mont.) de Bary) and brown rot/bacterial wilt (caused by the bacterium Ralstonia solanacearum (Smith) Smith). Eradication of Solanum dulcamara has been attempted as a control of brown rot (Persson 1998), and use of water from ditches in which Solanum dulcamara grows is a known cause of brown rot spread in irrigated potato fields. The role of Solanum dulcamara in the spread of late blight, however, has been shown to be minimal (Golas et al. 2010b), as most plants are resistant, and those that are susceptible to the disease do not harbour it over winter. Even in very favourable conditions only sporadic infections occur ( Cooke et al. 2002; Flier et al. 2003; Dandurand et al. 2006). Solanum dulcamara has novel genes for resistance to late blight, and there is scope for its use in new potato breeding programs in Europe (Golas et al. 2010b). Little work has been done to investigate the mechanisms or genetics of resistance to the major diseases of potato in species outside the Potato clade, but new breeding methods have stimulated the investigation of disease in wild species such as Solanum dulcamara (Golas et al. 2010b) and Solanum nigrum L., a European hexaploid member of the Morelloid clade ( Lebecka 2008, 2009).

Typification of the many synonyms of Solanum dulcamara has been very difficult. Many of the early names are found in early 19th century floristic accounts, and while validly and effectively published, usually do not cite specimens. I suspect many of these were not necessarily based on specimens, but rather on field observations. I have not neotypified any of the synonyms for which I could not find specimens directly linked to the original description. In addition, many of the European floras coined complex series of replacement names in the early parts of the 19th century, before the codes of nomenclature became firmly established. Where I have been able to trace the identity and derivation of epithets at varying ranks through citation of specimens I recognised these names as homotypic, but for many of the early names (e.g., those coined by George Don in 1838) the links are very tenuous and I have thus recognised them as heterotypic (and often have not designated types).

The type of Solanum persicum was a Pallas specimen held in Berlin, now destroyed. A Pallas specimen in BM that matches the description perfectly has been selected as a neotype, although it may not be a duplicate of the original.

The many varietal names coined by George Don (1838) were based not on specimens, but on literature references (e.g., var. album) or personal observations (var. hirsutum). His var. hirsutum referred to plants that were "Hairy or downy. .. On the sea coast", with no specimens or plates cited. I have not neotypified this name. Varieties album, violaceum and plenum were based on one plate (plate 46) in Hortus Eystettensis ( Besler 1613) which is composed of three plants (see Figure 37 ): one white-flowered Solanum dulcamara that I have designated as the lectotype of var. album, one violet flowered Solanum dulcamara that I have designated the lectotype of var. violaceum, and a central plant, used by Don as the basis for var. plenum, that is a drawing of Nigella sativa L. ( Ranunculaceae ) (see Figure 37 ). He cannot have been looking very carefully!

Petermann (1838) coined several new infraspecific taxa in his local flora of the Leipzig region; I have found no material related to these names. Solanum maximowiczii was based on " Solanum dulcamara var. ovatum Maxim." and was coined at the specific level to represent that variety. In his discussion of Solanum dulcamara , however, Maximowicz ( Regel and Maximowicz 1870) refered explicity to Dunal’s variety ovatum and did not coin a new name. Thus, this specific epithet is homotypic with the chain of epithets linked to the variety ovatum treated by Maximowicz, ultimately traceable to the (presumably) first instance of this varietal name being used in Petermann’s floristic treatment ( Petermann 1838). Dunal (1852), cited by Maximowicz, cites Opiz ( Berchtold and Opiz 1843) and Opiz cites Petermann as the origin of the varietal name. I have found no specimens associated with these names.

Dunal’s (1852) treatment of Solanum dulcamara for the Prodromus has a broad range of variation recognised as a single taxon; he recognised 14 infraspecific entities at two levels, mostly based on literature citations, his final words in the description - "Multum variat." - couldn’t be more true. He recognised as A, B and C forms previously recognised by Opiz ( Berchtold and Opiz 1843) and Petermann (1838) - cordatum, ovatum and hastiifolia (as hastaefolia). Below them on the page he recognised a series of 11 taxa ("Hae tres formae vicissum variant") that he listed with Greek letters (α to ξ, leaving out ι, κ and λ), some of which had binomial and others polynomial epithets. Infraspecific taxa denominated with Greek letters in Dunal (1852) have always been recognised at the varietal rank, and no where else in the treatment of Solanum in the Prodromus does Dunal use an apparently nested set of ranks as he does with Solanum dulcamara . I am therefore assuming that he intended his A, B and C ranks to be hierarchically above the varietal epithets and have treated them as subspecies in the synonymy. It is possible that var. hastiifolia of Opiz is in fact an illegitimate name for var. flexuosa, as it seems from the text that Opiz ( Berchtold and Opiz 1843) treated this variety as the typical one, but I cannot be certain and so have recognised it as legitimate. The polynomial varietal epithets (e.g., " corollis violaceis ", " corollis albis ", " corollis carneis ", " flore pleno ", corollis variegatis ") are not legitimate names; I have lectotypified the rest based on either herbarium material or literature citations cited by Dunal. For var. laciniatum, Dunal cited specimens in both P and G-DC; I have found only the G-DC sheet, so have selected it as the lectotype.

The holotype of Solanum dulcamara forma subglabrum was destroyed in Berlin and a duplicate of Kuntze’s collection held in NY has been selected as the lectotype.

In describing Solanum dulcamara var. macrocarpum, Maximowicz ( Regel and Maximowicz 1870) cites as a locality "Circa Hakodate insulae Jezo pluribus locis". Several collections of Maximowicz’s from Hakodate dated 1861 are held at LE; others are in BM, GH and NY. It is very unclear whether any of these are true duplicates, so I have selected one of the flowering and fruiting sheets at LE as the lectotype and another at LE that is almost identical in morphology as a possible isolectotype.

Wasibecker’s (1895) Solanum rupestre is a later homonym of Solanum rupestre H.W.Schmidt. It is possible that the name Solanum serpentini Borbas & Waisb., apparently coined in 1897 ( Soó 1968: 153) is a replacement name for the homonym, as the epithet indicates an origin on serpentine soils (like the type of Solanum rupestre Waisb.). I have been unable to locate the place of publication of Solanum serpentini , and so treat Solanum rupestre Waisb. as a synonym of Solanum dulcamara and Solanum serpentini as a doubtful name (see Doubtful and excluded names and Names not validly published).

Máthé and Máthé (1972) undertook an exhaustive study of the variation of Solanum dulcamara in Hungary and described two new variants, one at the form level (forma lucidum) and the other (luc. atroviolaceum) at the rank ‘lucus’ that was used in by Hungarian botanists between variety and forma. He followed Soó (1968) who transfered many earlier epithets to the rank of lucus, however, none of these are validly published, as no explicit reference to the basionym’s place of publication was made as is required by the Code.

I here cite specimens I have examined; these do not cover all European countries, although Solanum dulcamara is recorded from throughout Europe. I have selectively cited material from the range edges, as that is where I anticipated taxonomic problems might lie. Solanum dulcamara is very commonly collected and the specimens cited here do not completely represent the range.

Specimens examined.

Afghanistan. Kurrum valley, at Zabardastkalla and Alikhél, Dec 1879, Aitchison 861 (BM, GH, K, LE, S); Sere Tange, south of Marshad valley, Keshan, 1250 m, 30 Jun 1969, Carter 375 (K); Parwan, 33 miles from main Salang road, by side of Gharband River, 1920 m, 13 Jun 1969, Hewer 1274 (K, LE); Gulbahar, 14 May 1937, Koelz 11428 (LE, US).

Albania. Shkoder, North Albania, environs of the city of Shkoder near the river Umri, 14 Jul 1959, Kuvaev 95 4 (LE).

Algeria. Clemcen, vers les Cascades, 800 m, 24 Aug 1932, Faure s.n. (S); Elenicens?, Aug 1849, Romain s.n. (BM).

Andorra. Santa Julia de Loria, de borda del Sabater a Tolse, 840 m, 27 Aug 2002, Aedo et al. 8428 (MA).

Armenia. Kafanskii Region, near Ang, 24 Aug 1980, Avetisyan s.n. (MO); Karassan, 1837, Koch s.n. (LE); Arpa, between Iahichevan and Yerevan River Arpa to the right of the bridge, 21 Jun 1974, Mamakli s.n. (LE); Kirovakan station, 26 May 1953, Tamamschian, S., s.n. (LE); Megra, Megrinskii Region, 21 Jul 1954, Tamamshek & Deiisova s.n. (LE); Kirovakai, 26 Aug 1953, Tamaschira s.n. (LE); Alaverdskii reg., S. Sanain, Uzunar, 25 Jul 1960, Avegisyan & Gabrielian s.n. (LE).

Austria. Tyrol: Tirolia centralis, in valle Gschnitz prope pagum Trins, Kerner 3290 (BM, LE); Vienna: Wien, erster Bezirk, an der Grundmauer des Naturhistorischen Museums auf der Seite der Ballariastrasse, 200 m, 2 Oct 1992, Wallnöfer 4212 (BM).

Azerbaijan. Distr. Schemacha, Fagraüksch, Prope st. Scharodilskaja, ad lacum Fagraüksch, 792 m, 8 Aug 1900, Alexeenko 11304 (LE); Bakinskaya Oblast’, Kusary, Kubinskii region, 20 Jul 1900, Grigoreev 20 (LE); Lenkoran, 1834, Hohenacker s.n. (G-DC, K, LE, MO); Distr. Kuba, Seput, 24 Jul 1930, Kasumova s.n. (LE); Eshakchi, 3 Jul 1909, Kirichenko s.n. (LE); Shirvan steppe, inter Kerzi-Kent et Kotoran, 24 Aug 1930, Kolakovskii s.n. (LE); Gandzha Oblast’, District Kazakh, near the road to Poyly, 13 Jun 1928, Kolakovskii s.n. (LE); Lenkoranskiy Okrug, [near village of Nipolaevna, on river Vilyazh-chai], 30 Jul 1931, Matveeva 791 (LE); Gandzha Oblast’, Bardy, Agdam District, on the Karabkh steppe, near town, by the River Terter, 17 Aug 1927, Prilipko s.n. (LE); distr. Saljany, Saljany, prope st. Dzhafarchan, 6 Jul 1927, Prilipko s.n. (LE); distr. Lenkoran, Tangovan, near village Tangovan, 26 Jun 1931, Shipchinskii 275 (LE); Talysh, Lenkoranskii, 16th km on the road from Lenkoran to Leric, left bank of the river Lenkoran-chai, 100 m, 1 Jul 1952, Vasiliev & Vasilieva s.n. (LE); 18 km from Sheka, 26 Jun 1972, Vlasov s.n. (LE).

Belgium. Brussels: garden on Chaussee Pas in Auderghem, near a forest east of Brussels area, Jul 1987, Geerinck-Coutrez 4343 (BM); West Flanders: Knokke, Le Zwin, 21 Jun 1971, Auquier et al. s.n. (LE); Knokke, Zrvin, 8 Aug 1955, Lonvalrée 6781 (BM).

Bosnia-Herzegovina. Mostar, 1886, Bornmüller 1405 (B).

Bulgaria. Varna, Aug 1886, Bornmüller s.n. (B); 8-9 km W of Velingrad, along the stream Lukovica, 980 m, 8 Jul 1982, Frost-Olsen 4401 (MA); Vitosa, 10 Jun 1927, Georgieff s.n. (LE); Malkija Kozuh, Struma valley by the hill Malkija Kozuh, 9 Jun 1980, Kuzmanov 801405 (B); Samokor, 30 Aug 1912, Post 277 (G); Sliven, 16 Jul 1907, Schneider 710 (BM); Prilap, May 1917, Stribrny s.n. (LE).

Canada. British Columbia: Vancouver, Vancouver 8: Alma. Roadside, 18 Jul 1954, Bird 43 (BM);, S. Okanagon, White Lake, Oliver road, 480 m, 6 Jul 1961, Vrugtman & Campbell 610348 (BM); New Brunswick: Kent County, Newcastle, rocky shore of Miramichi River, 17 Jul 1945, Dore & Gorham 45555 (G); St. John County, Fairville, near St. John River, 19 Sep 1926, Fernald et al. 394 (GH); Bass River, Kent County, Fowler s.n. (LE); Newfoundland: Salmonier, 1931, Ayre s.n. (GH); Nova Scotia: Annapolis County, Granville, North Mtn, 18 Jul 1921, Bartram & Long 24435 (GH, LE); Yarmouth County, Markland, Cape Forchu, 13 Jul 1921, Fernald et al. 24434 (K, LE); Shelburne County, East Jordan, 4 Aug 1921, Fernald & Long 24436 (GH); Halifax County, Halifax, 30 Sep 1924, Jack 3644 (GH); Guysborough County, Guysborough, 14 Aug 1930, Rousseau 35472 (K); Ontario: Plevna, 26 Jul 1902, Fowler s.n. (GH); Hastings County, Marmora, Concession 3-4, Twin Sister Lakes, 8.5 miles NNE of Marmora, 14 Aug 1952, Gillett & Hammond. 6922 (G); Bruce County, Tobermory, 2 Jul 1933, Krotkov 7758 (GH); Waterloo County, Westmount Woods, 1 mi. W of Kitchener, 16 Jun 1939, Montgomery 338 (GH); Brittania, 18 Jun 1921, Rolland 15680 (GH); Halton County, Great Lakes Region, Lake Medad, 25 Jun 1940, Soper & Burcher 2011 (BM, GH); Prince Edward Island: Queens County, Charlottetown, 28 Aug 1912, Fernald et al. 7789 (GH); Quebec: Iberville County, Ile aux Noix, 12 Aug 1928, Adrien 2098 (K, LE); Missisquoi, Noyan, 23 Jun 1959, Gervais & Lavigne s.n. (G); Pontiac County, Beech Grove, 19 Aug 1943, Lamarre s.n. (MA).

Croatia. Dalmatia, Spalolo, Mossor, 1886, Bornmüller s.n. (B); Golubevec coal district, 1924, Leathes s.n. (BM); island of Veglia (Crk-Venica), Sep 1924, Leathes s.n. (BM).

Denmark. Copenhagen, Lyngby Mose, 14 Jul 1941, Dahl D-45 a (BM); Odder, Sondrup Plantage, ca. 11km S of Odder, W part of the plantation, 25 Jul 1978, Frost-Olsen 1748 (LE); Lake Tissø, D42, Jul 1968, Hjorih-Olsen 10 (BM); Jutland, Silkeborg, Borre Lake, 1 Aug 1968, Jensen et al. 464 (BM, MA); North Jutland, Stenbjerg, 9 Jul 1964, Larsen & Laegaard s.n. (LE); Skagen, 9 Aug 1973, Larsen et al. S.L. 6847 (BM, MA).

Egypt. Sin. loc, Anon., s.n. (LE).

Estonia. At village Oiu, 12 Sep 1967, Aasamaa 11616 (MO); Dorpet [Dorpat = Tartu], 12 Jul 1860, Gruner s.n. (BM);Vijlandi County, Tipu, River Halliste, 14 Jul 1994, Reier s.n. (B).

Finland. Pyhäranta, Rihtniemi point and village, 21 Aug 1968, Alava et al. s.n. (LE); Nauvo, Seili, near church, 4 Aug 1976, Arkkola et al. s.n. (MA); Åland, Nafsby, Prov. Hammarland, 28 Jul 1963, Haakana s.n. (LE); Kupittaa, Turku, Varsinais-Suomi, 14 Aug 1924, Kari s.n. (LE); South Häme, Heinola, Tommola, Maitiaislahti, 7 Aug 1980, Kemppainen s.n. (MA); Kärkölä, Järvelä, middle part of the NE shore of Lake Hähkäjärvi, just NW of the man-made cape, 97 m, 29 Jun 1992, Lampinen 14764 (MA); Kuitia, Parainen, Lemlaxö, 7 Aug 1971, Kukkonen 9106 (LE); Sauvo, Karona, near the church, 13 Jul 1976, Lempiäinen & Ravanko s.n. (MA); Broby, Nylandia, par. Pyttis, 28 Jun 1963, Majander & Nordström s.n. (LE); Norra byn, par. Sottunga, 11 Jul 1973, Nordström s.n. (LE); Uusimaa, Jätkäsaari, Helsinki, harbour, 28 Jul 1968, Oinonen s.n. (LE); Versinais-Suomi, Iniö, Norrby, 13 Jul 1929, Valle s.n. (MA); Jyddo-ojen, Foglo, Ahvenanmaa ( Åland), 29 Aug 1907, Wahlberg s.n. (LE).

France. Alsace: at the edge of the river at Haguenau, 24 Jul 1846, Billot 160 (BM); Auvergne: Auliac, 26 Sep 1835, Anonymous s.n. (BM); chemin de Surat à Maringues; chemin de Bussière à St.Clement, Jun 1906, Caumel 334 (BM); Brittany: chemin de Chantenay à Rochemaurice, 12 Aug 1901, Gadeceau 5106 (BM); Corsica: zwischen Pte d’Abatesco und Mignattaja S von Ghisonaccia, 10 Aug 1933, Aellen 4601 (LE); Languedoc-Rousillon: Bouches du Rhone, Camargue, Tour du Valat, SE of Mas, 11 Jun 1968, Kendrick & Moyes 231 (BM); Bouche du Rhone, Camargue, Etang de la Dame, 2 miles SE of Salin de Giraud, 21 Jun 1968, Kendrick & Moyes 346 (BM); Loire-Atlantique: Ouest de France, environs de Vertou, Jun 1880, Gadeceau s.n. (BM); Nord Pas de Calais: Beuvrequent, ruisselet de Slack, 24 Jul 1986, Coutrez 3780 (MA); Pays-de-la Loire: Nantes, loin le Fevre, 18 Jul 1847, Genevier s.n. (BM); Picardy: Aisne, St. Quentin, Jun 1871, Moignier s.n. (BM); Poiteau Charente: Rochel, Plaisance en Oléron (Charente-Inferieure), 20 Oct 1892, Reau s.n. (BM); Rhône-Alpes: Bonneville, bords d’Arve, Depierre s.n. (BM); Haute Savoie, route d’Etrumbiern a Mornex, 14 Jun 1859, Déséglise 281 (BM); Hirault, Mont Blanc, Station Crenugua, 7 Jul 1954, Witte 14547 (MA).

Georgia. Dusheti District, south and east of Fudauri, north of Pasanauri, slopes along Georgian M3 Road (Georgian Military Highway), 1784 m, 5 Aug 2005, Atha et al. 5005 (US); Chevsurya, between Barshak and Reshka, 1 Jul 1903, Busch s.n. (LE); Akmolinsk prov., Kokczetav District, 4 May 1913, Drobov 203 (S); South of the Krestovyy Pereval, Georgian Military Highway, Belaya Aragvi valley, between Ordzhonikidze and Tbilisi, Gruzinskaya S.S.R. along walls in the village of Mleti, 4 Jul 1971, Jenkins 2946 (BM); Somchetya, Teleti, near Tiblisi, Oct 1923, Juzepczuk s.n. (LE); Klukhorskii, between villages of [illegible] and Madniekheva, 13 Aug 1946, Kemularia-Nathadze & Chinthibidze s.n. (LE); Kutaisi, Ozur, 15 Jun 1914, Kikodse s.n. (LE); Abkhasia, Teberdinskii, Protected Area [Terbeda], 1938, Komarov & Komarova s.n. (LE); Kazbek, Dushetskii region, near the 31st field, Tifliskaya Province, 1916, Krylov & Steinberg s.n. (LE); Megrelia, near village Inghuri towards city Tobashi, 14 Aug 1952, Madenova & Kuthatheladze s.n. (LE); along the Georgian Military road in the area of Krestovi Pass N of Tibilisi, 31 Jul 1984, McNeal et al. 332 (MO); Atskhur, Akhaltsikhskii region, vicinity of village Atskhur, 16 Jul 1926, Meffert 314 (LE); Kazbegi, Aug 1971, Menitskii s.n. (LE); Abkhasia, Sukhumsk, District of Tsebeld, in the Peukirskoe gorge on open slopes of the mountain Agysh near the road, 23 Aug 1937, Pobedimova 383 (LE); Abkhasia, Gagra, on the lower road towards Zhoevarskii waterfall, 4 Nov 1915, Sakharov s.n. (LE); Mtiuleti, Kasbegi Region, Dariali Gorge, Greater Caucasus Mountain Range; ca. 12 km N of Kasbegi village, on footpath to Gveleti village to local waterfalls, 1600 m, 24 Aug 1998, Schmidt et al. 2891 (G); Tiblisi, by the River Vera, 26 May 1917, Shishkin s.n. (LE); Adzharskaya, Nizovaya Natanebi, 4 Aug 1948, Sochava & Semenova s.n. (LE); Adzharskaya, gorge of the river Bakhvis-tskali, 9 May 1941, Tatishvili s.n. (LE); valley of the River Terek to the south of the Station Lars, 25 Aug 1949, Vasilchenko et al. 926 (LE); Sukhum, on the right bank of the River Besleta, near the famous factory, 29 Jul 1951, Vasiliev s.n. (LE).

Germany. Bayern: Oberbayern, München, obhere Speilbahn des Golfplatzes Thalkirchen 1 km sudlich des Asam-Schlössels, 550 m, 12 Jul 1992, Förther 48 (B); Lkr. Ostallgau, NW Trauchgau, 845 m, 24 Jul 2001, Herb. Willing [Willing & Willing] 13418 D (B); Lkr. Garmisch-Partenkirchen, SW Krün, 911 m, 5 Aug 2001, Herb. Willing [Willing & Willing] 15543 D (B); Berlin: Pichelswerder, Bez. Spandau, 35 m, 4 Jun 1998, Herb. Willing [Eisenblatter & Willing] 4624 (B); North Rhine-Westphalia: Münster in Westfalen, Wilms s.n. (BM); Rheinland-Pfalz: Westerwaldkreis, Seeburg, westerwalder Seenplatte, Haidenwalder bei Seeburg, 421 m, 2 Jul 1978, Kalheber 78-424 (B); Schleswig-Holstein: Rendsburg, Larsen et al. 106 (BM, LE, MA); Thüringen: Am Ufer des Altenberger Teiches bei Eisenach, 28 Jul 1947, Launert s.n. (BM).

Greece. Thracia, prov. Xanthi, in valle infra pagum Dhamari, 4 km infra junctionem torrentium, 500 m, 16 Aug 1978, Greuter 16361 (B); Dhrama, prope phylacion nom. Praseki, a pago Ano Vrondous, ca. 6 km boreo-occidentum versus, 1320 m, 22 Aug 1978, Greuter 16703 (B); Mt. Olympe, Thess, Jun 1929, Guiol 642 (BM); Etolia-Akarnania, Arahova (Ar 358), 900 m, 8 Jul 1996, Herb. Willing [Eisenblatter & Willing] 48029 (B); Thesprotia, Ep. Souliou, NW Paramithia (The 209), 190 m, 26 Jun 1997, Herb. Willing [Eisenblatter & Willing] 55434 (B); Trikala, Ep. Kalambakas, Notia Pindhos, NW Kalambaka (Tri 200), 330 m, 18 Jul 1997, Herb. Willing [Eisenblatter & Willing] 60744 (B); Nom. Ionnanina, Ep. Dhodhonis, Tsepelovo (Ioa 574), 1000 m, 17 Jul 1998, Herb. Willing [Eisenblatter & Willing] 67904 (B); Nom. Kastoria, Ep. Kastorias, SW Vyssinia (KAS 164), 820 m, 31 Jul 1998, Herb. Willing [Eisenblatter & Willing] 71725 (B); Pella, Ep. Edhessis, W Loutraki (Pel 115), 260 m, 27 Jul 2000, Herb. Willing [Eisenblatter & Willing] 86148 (B); Arkadhia, Elati, 1180 m, 27 Sep 2003, Herb. Willing [Willing & Willing] 118384 (B); Paranestion, Nomos Drama, Dimos Paranestion, river Nestos ca. 1 km SW of Paranestion, 100 m, Aug 1996, Liebfritz & Schuler s.n. (B); Nomos Kavala, Dimos Keramoti, mouth of river Nestos, 1 m, 31 Aug 1998, Schuler 98/447 (B); Ipiros, prob. Ioannina, distr. Konitsa, Konitsa, on both sides of the Aoos River above the bridge, 460 m, 23 Jul 1971, Stamatiadou 13382 (BM).

Hungary. Sin. loc., Jul 1877, Anonymous s.n. (BM); Kozragohid, Budapest, 26 Aug 1904, Degen s.n. (LE).

India. Jammu and Kashmir. Kashmir, Kishtwar, 1829 m, 17 Sep 1876, Clarke 31361 (BM); above Barangalla, 2000 m, 25 Jun 1902, Drummond 13886 (E); Gandarbal, Sind valley, 1585 m, 1 Jun 40, Ludlow & Sherriff 8101 (BH, BM, E); Hajipir pass, Poonch to Uri, 2134 m, 4 Jul 1934, Stewart 14000 (A); Naranag, Wangat valley, 2134 m, 8 Aug 1939, Stewart & Stewart 18104 (GH); Poonch & Azad Kashmir, below Hajiperi, 2134 m, 3 Jul 52, Stewart & Nasir 24062 (BM).

Iran. Ghilan, Rescht, 13 Jul 1902, Alexeenko 47 (LE); Fars, Tang-e-Sorkh, between Ardakan and Yasuj, 2450 m, 18 Jun 1977, Bokhari & Edmondson 2159 (G); Meshed, 1941, Mrs Donaldson s.n. (K); Amol, 25 miles S of Amol, Central Elburz N. slopes, 1219 m, 21 Aug 1966, Furse 9071 (K);. Keredj, 29 Sep 1934, Gauba 829 (B); Kurdestan, 31 km N of Sanandaj, 2000 m, 8 Jul 1964, Grant 16064 (MO); Dozein, 5 km N of Dozein, which is a village 55 km SE of Gonbad-e-Kabus, 950 m, 19 Jun 1977, Hewer 3961 (K); Khairat, Mazenderan, 2439 m, 25 Jul 1940, Koelz 16569 (US); Kuhi, near Teheran, Pidjehek, Jul, Kotschy s.n. (BM); Fars, 36 km NW Ordekam, 14 Jul 1959, Pabot 2446 (G); Qotur River,W Khvoy towards the Turkish border, 1800 m, 10 Jul 1971, Rechinger 41735 (B, G); Keredj, Elburs Mountains, Keredj River, 6 Jun 1937, Rechinger 733 (S); Hamadan, 2134 m, 7 Sep 1929, Rogers 538 (K); Joistan, Talighan Valley, 1829 m, Jul 1953, Trott 24 (K); Astava, shoreline of Caspian Sea, 21 Aug 1966, Wright 65 (K).

Iraq. Baghdad, Pig Island, 10 May 1960, Agnew et al. W-1679 (K); Tigris Plain. near Mosul, 200 m, 5 May 1936, Low 284 (BM); Penjiwin, 1200 m, 8 Jun 1948, al Rawi 12159 (K, US); Pushtashan, 15 km NE of Rania, lower slope of Qandil Range, 1150 m, 29 Jul 1957, al Rawi & Serhang 23865 (K); Erbil, Pushtashan, Montes Qandil, (Kurdistan), 1000 m, 28 Jul 1957, Rechinger 11027 (G).

Ireland: West Galway, Connemara, Renville, Babington 299 (BM); Cork, Bishopsbrook, 1849, Herb. J. Carroll s.n. (BM); South Kerry, Dooks, 15 Jul 1903, Marshall s.n. (BM); Meath, Drogheda, Aug 1956, McClintock s.n. (BM); North Kerry, Walls between Killarney and Mucross, 24 Aug 1883, Ridley s.n. (BM); South Kerry, Rossbeigh Dunes, 5 Aug 1935, Simpson 35620 (BM); South-East Galway, near Dalystown, 24 Jun 1952, Simpson 52108 (BM).

Israel. Gilead, Burmah to Gerashi, 4 May 1886, Anonymous s.n. (K).

Italy. Abruzzo: Teramo, cerca Castelli, Findo de al Salsa, 1100 m, 2 Jul 2002, Navarro et al. 4348 (MA); Apulia: Puglia, farm about 7 km NE of Taranto, 1881, Vitantonio 1675 (BM); Emilia-Romagna: Parma, Anonymous s.n. (BM); Friuli-Venezia Guilia: Trieste, Sistiana, 25 May 1980, Greuter et al. 17348 (B); Sicily: Nebroda Mtns., ad pedes montium Madoniarum prope Polizzi, 600 m, 6 Jul 1874, Strobl s.n. (BM); Catania, Randazzo, Lago de Gurrida, 820 m, 5 Jun 2000, Ávarez et al. 1490 (MA); Tuscany: Agro Lucchese, Lucca country side, Jun 1835, Puccinelli s.n. (BM).

Japan. Hokkaido: Sapporo, 4 Jul 1903, Arimoto s.n. (GH, MO); Iburi, Tomakomai, 30 Jul 1930, Akiyama s.n. (S); between Kotoni and Sapporo, 22 Aug 1929, Dorsett & Morse 1128 (US); Yezo, prov. Nemuro, to Attoko-too from Attoko-Nemuro-gun, 8 Jul 1959, Furuse s.n. (A, S); on the coasts vicinity and Oikamanai-numa Bansei, Taiki-choo, Hirowo-gun, Province Tokachi, 19 Jul 1974, Furuse 6434 (K); Wakkanai-Park. Wakkanai-shi. Province Kitami. Hokkaido ((Yezo)), 18 Aug 1975, Furuse 9465 (K); Jyuusam-mura. Nishi-tsugaru-gun. Province Mutsu. Hondo, 20 Sep 1955, Furuse 29993 (K); Watsakinai. Toyotomi-choo. Teshio-gun. Province Teshio. Hokkaido (Yezo), 2 Jul 1975, Furuse 8980 (K); prov. Iburi, between Tomakomai-shi and Lake Shikotsu, 15 Aug 1981, Takahashi 1650 -A (NY); Honshu: Shimura, Itabashi-ku, Tokyo Prefecture, Jun 1907, Makino 75314 (LE); Shiga, Yotsugawa, Adogawa Cho, Takashima gun, Lake Biwa, 86 m, 17 Sep 1991, Kanda & Kuribayashi 1 (A); Hondo, Oze-ga-hara, prov. Kozuke, 26 Aug 1950, Mizushima 417 (A); Aomori-shi, Arakawa, below Jogakura Spa, 810 m, 8 Aug 2000, Yonekura 5960 (A).

Kazakhstan. West Kazakhstan Prov., Right side of river Ural, 2 km downstream of Kopseharov, 15 Feb 1955, Grubov & Lynbarskii 32 (MO); South Kazakhstan Oblast, Shymkent, prov. Syr-Darja, Distr. Tschimkent, Tchiluk, 1908, Knorring 604 (S); Tselinogradskaya Oblast’, near village of Kurgaljinskii, on the lower eastern shore of the Lake Kuraljin, in the surroundings of the settlement of Karazhar, 20 Jun 1980, Sidorova 155 (LE).

Lebanon. Ain Zahatallah, Mt. Lebanon, Aug 1912, Letchworth s.n. (K).

Lithuania. Holonety, dist. Wilkomierz, 30 Jul 1895, Rudiminowna s.n. (B, BM).

Luxembourg. Jardin Botanique, 20 Aug 1837, Gay s.n. (K).

Macedonia. Popovo, NW foot of Krusa Balkan, 3 m E of Lake Doiran [Doirani], 1918, Gooding s.n. (BM); Kojnsko, distr. Duditza-et Saharupa-planina, 550 m, Jun 1917, Schultze-Jena 138 (B).

Mongolia. southwestern Mongolia, Khovdinskii Aymak, southern slope of the Mongolian Altai, 45 km to the E of the villageg of Bulgan, along the River Bulgan, on the border with KNR [China], 1979, Gubanof 7321 (LE); "ad Mongolia borealis, Altai australis ad Totyask nigrum", 1876, Potanin s.n. (K).

Morocco. Xauen, 650 m, 28 May 1928, Font Quer 354 (BM); Tazzeka, 13 km from Taza on minor road, Ras-El-Ma, 990 m, 5 Jul 1993, Jury et al. 11718 (K).

Norway. Oslo, 24 km hinter, am fjord, 8 Sep 1980, Meyer s.n. (B); Stjordal, near Hagra, 1926, Trethewy s.n. (BM); Astre Biarum, 6 Sep 1902, Triatz s.n. (BM).

Poland. Wolka wysoka prope Sanniki, distr. Gostyn, 22 Jul 1895, Drymmer s.n. (BM); Orawicza, in Banat, 6 Aug 1840, Wierzbicki s.n. (GOET).

Portugal. Salvador, as margenes do rio Sever, Sera de S. Maccede, 12 Jun 1959, Beliz & Raimundo 5046 (MA); inter Garvas et Panvies, Jun 1875, Daneau 1184 (BM); Alto Alentejo, Elvas, Frequisia de Varche, Horto do Carlos Ranita, margem da Ribereira do Cancão, 30 Sep 1984, Guerra 1518 (MA); Beira Alta, Vizeu, Santa Comba-Dão, Pinheiro de Azer, descida para a capela de Sra. do Pranto, 11 Jun 1982, Marques 2087 (MA); Rangel prope a Coimbra, Jun 1889, Moller s.n. (BM); Oliveira, M.P. de, s.n. (BM); Entremadura, Obidos nas margens dos vales na La Goa de Obidos, 14 Jun 1917, Rainha 377 (MA); Idanha a Nova, Ribeira do Poasul, Jul 1883, Ricardo da Cunha s.n. (BM); Alijó, Pinhão, beira-Doura, foz do Rio Pinhão, 30 Jun 1070, Rozeira et al. s.n. (MA); Douro Litoral, near Oporto, 2 Jul 1887, Murray s.n. (BM); Madeira, sin. loc, 1804, Turner s.n. (K); Tagum, 1848, Welwitsch 202 (LE).

Romania. Comm. Balcani, prope pagum Frumoasa, 550 m, 12 Jul 1972, Barabas 487 (BM).

Russian Federation. Central Federal District: Moscow, 30 Aug 1992, Croat 73991 (MO); Far Eastern Federal District: Amurskaya Oblast’, Radde, around the village, on the River Amur, 25 Sep 1926, Selivanova s.n. (LE); Amurskaya Oblast’, Zea, Zeyskii region, around the town of Zea, 30 Jul 2000, Shaligin s.n. (LE); Dalnevostochnii Krai, near Valdivostock, 12 Jul 1929, Transhel 284 (LE); Kurilskii Islands, Iturup [illegible], 914 m, 29 Aug 1946, Vorobeev s.n. (LE); Archangelsk Oblast, Archangelsk, Beresnik, 1919, Grantham s.n. (BM); around Vladivostock, on the bank of Lake Khanka, 10 Aug 1929, Shishkin 1306 (LE); island of Sakhalin, village of Vladimirovskoe, 26 Jun 1899, Shestunov 53 (LE); Primorskaya Oblast’, around 25 verst [measure of distance] to the west of Vladivostock, on the slope of a mountain, 29 Jun 1926, Ivanova s.n. (LE); Primorskaya Oblast’, Khabarovsk, left bank of the River Amur, near Khabarovsk, 25 Jul 1926, Solokhin s.n. (LE); Primorskaya Oblast’, Petrovolikova Bay, island of Furugelma, 21 Sep 1922, Kozlov 483 (LE); Primorskaya Oblast’, the village of Chernshovekoe [Chernyshevka], 30 Jul 1902, Paltzevskii s.n. (LE); Sakhalin Oblast, on the sea slope of southern Sakhalin, between the settlements of Khvostovo and Altasovo, 13 Aug 1959, Pobedinova & Konovalova 1347 (LE); North Caucasian Federal District: Tegenekli, basin of the River Baksan, right side of the gorge Prielbrusye, 23 Aug 2000, Menitskii & Popova s.n. (LE); Nalchik, to the south of the settlement of the village Belaya Rechka, 2 Aug 2000, Menitskii & Popova s.n. (LE); [Chechnya] Itun-kale, Argun, left side of river Argun, above the village, 23 Aug 1988, Menitskii & Nikolaev 38 (LE); [Dagestan] Machachkala, by the river Terstagay, 26 Sep 1926, Orlov s.n. (LE); [Dagestan] Gergebilskii region, between villages of Arakani and Mogok, 14 Jun 1961, Tsvelev et al. 914 (LE); [Dagestan] Levashinskii region, between villages of Gersibil and Levashy, 27 Jul 1961, Tsvelev et al. 4175 (LE); Chechyna, Terekaya Oblast’, Beslan, May 1901, Alexeenko 14374 (LE); Kabardino-Balkar, Bezeniyskoe gorge, on the slope of the mountain Ushtensk, 12 Aug 1995, Shkhagapsoev & Korchevskaya 675-2000 (LE); Ordzhonikidzevskii, “Chechnya-Ingushetia”, 22 Jul 1966, Vlasov & Kuvas s.n. (LE); slope of the left bank of the River Bizhgon 10 km S of the station Storozhevaya, 31 Aug 1947, Borisova 369 (LE); Stavropolskii Krai, bank of River Podkushok in the region of Pyatigorsk, 8 Aug 1990, Yagushevskii s.n. (LE); Terekaya Oblast’, Nalchik, near town, 30 Jun 1917, Paltseva s.n. (LE); Northwestern Federal District: Ladozhskoe Lake, town Lakhdempokhye, 6 Jul 1961, Pobedimova & Gladkova 97 (LE); near St. Petersburg, pag. Olgino, in litore sinus Fennici, Jun 1919, Czernjakovskaja s.n. (BM); Siberian Federal District: Siberia, regio transbaicalensis, Utotschkina, Selenga River, 500 m, Jul 1900, Ehnberg s.n. (G); Irkutsk Oblast, Irkutsk, Besser s.n. (K); Krasnoyarsk Krai, Yeniseysk, "Siberia, Jenisei, Jeniseisk", 23 Jun 1876, Arnell s.n. (S); Central Siberia, region of Novosibirsk, near railway station Seyatel’ ca. 3 km Nw of Akademgorodok near river Ob’, 2-3 km below hydroelectric dam, ca. 20 km SSE of Novosibirsk, 140 m, 24 Jul 1979, Iltis et al. 1115 (G, S); Podkova, Yekaterinava, 21 Jun 1906, Yálovaya s.n. (LE); Southern Federal District: Mineralnie Vody, eastern slope of the mountain Zmeyka, 7 Aug 1996, Menitskii & Popova 122 (LE); River Mzymty [Myzmyta] near the mouth of River Agipse, 7 Sep 1928, Leskov s.n. (LE); Tuapsinsk, Black Sea Region, 8 Jul 1915, Litvinov s.n. (LE); [Adygea] Kubanskii Oblast’, Maykopsk, between the stations of Dondukovkskaya and Sergieskaya, in the valley of the River Fars, May 1915, Kozo-Polyanski & Preobrasheiski s.n. (LE); Krasnodavskii Krai, Novorossika, Black Sea region, near the road on the high shore of the sea, 25 Jun 1923, Pojarkova 156 (LE); Urals Federal District: Goygol Rayon, Goygol, territory of Elizabethapolis [Yekaterinburg], 8 Jun 1844, Kolenati 1753 (LE).

Serbia. mountain above Jviato Petka, 22 Aug 1921, Yermoloff s.n. (BM).

Slovenia. within 6 miles of Bled (in Slovenja, at the borders with Austria), which includes the lower slopes of Mt.Triglav, 304 m, 1923, Leathes s.n. (BM).

Spain. Alcabete: Reolid, 800 m, 19 Aug 1985, Figuerola s.n. (MA); Andalucia: Granada, Castril, pr. Cortijo del Nacimiento, 1100 m, 21 Jul 1978, Charpin et al. 15109 (MA Asturias: Villaviciosa, playa de España, 5 m, 5 Oct 1992, Aedo 2469 (MA); Avila: Conabeltrán (?), Jul 1918, Cogolludo s.n. (MA); Baleares: Mahón, San Sancho, 18 Apr 1899, Pons, Gereau s.n. (MA); Burgos: Carazo, junto a la Fuente de la Mora, Peñas de Cervera y aledaños, 1200 m, 24 Jul 1979, Pons Sorolla & Susanna 616 (MA); Caceres: Baños de Montemayor, 15 May 1944, Cabellero s.n. (MA); Cadiz: collected in a farm named ‘Marcellino’ in a rough country between Los Barrios and Algeciras, 60 m, 14 Jun 1973, Brinton-Lee 1346 (BM); Cantabria: Santander, Cueto, El Panteón del Inglés, 22 May 1992, Busqué s.n. (MA); Castellón: Villafarnés, on same sheet several specimens, also from Castielfavib and Vallanca (Valencia, 1792, 1793), and Jardin de la Chevette (France, 1781), 1792, Anonymous s.n. (MA); Castilla: Bujedo, 28 Jul 1920, Elías s.n. (BM); Catalunya: Castello, Castillo de Villamalefa, Río Villahermosa, 540 m, 22 Oct 1998, Riera et al. 19505 (MA); Cerdagne, Llivia, val de l’Estahuja, 1300 m, 10 Aug 1918, Sennen s.n. (BM); Cuenca: Hoz de Cañizares, 10 Jul 1932, Cabellero s.n. (MA); Tarancón, 7 Apr 1977, López 8 (MA); Cuidad Real: Frencaliente, margenes del Río Pradillo, 600 m, 7 Jul 1997, García Río s.n. (MA); Galicia: Lugo, Riberas de Lea, 25 Jul 1956, Carreira s.n. (MA); Guadalajara: Zaorejas, Hoz del Tajo, piscifactoria del Campillo, 900 m, 18 Jul 1981, Muñoz 512 (MA); Huelva: Almonte, Coto de Donaña, Reserva de La Rocina, 28 Jun 1977, Castroviejo & Porta 733 (MA); Madrid: San Agustín de Guadalquix, riberas del Río Guadalquix, 3 Aug 1982, Moreno s.n. (MA); Navarro: Pamplona, 16 Jun 1971, Moriyóu s.n. (MA); Salamanca: alrededores de Bejar, Aug 1914, Cogolludo s.n. (MA); Segovia: Granja de San Ildefonso, desde la finca La Sauca hacia la carretera a Torrecabelleros, 30TVL1529, 1150 m, 9 Sep 1985, García Adá & Egido 1816 (MA); Soria: Cañon del Río Lobos, 14 Jul 1980, Buades s.n. (MA); Tarragona: 20 km SW of Tarragona, between Cambrils and Hospitalet, 29 May 1962, Brummitt et al. 298 (BM); Toledo: ribera del Río Tajo, 500 m, 18 Jun 1983, Egido 947 (MA); Valencia: Bco. Bocairente, 510 m, 7 Jul 1980, Palasí s.n. (MA); Valladolid: Encinos de Esqueva, en Arroyo de los Frios, 11 Jul 1983, Fernández Alonso 2071 (MA); Zamora: Parque Natural Lago de Sanabria, Forcadura, 1100 m, 1 Jul 1987, Anonymous s.n. (MA).

Sweden. Uppland, ad sopes Upsalia passim, Aug 1870, Ahlberg s.n. (BM); Uppland, Funbo Parish, southern shore of Lake Goren, 6 Jul 1944, Alm & Smith 272 (BM); Bohnslam, Väderöarna, 10 Jul 1909, Almquist s.n. (BM); Brohuslan, Fjalibacha, 3 Jul 1910, Almquist s.n. (BM); Uppsala, Andersson s.n. (MA); Skafto, Gaso, 20 Jun 1948, Ankarsward s.n. (BM); Södermanland, Toiosocken, Stora Grasskar, 6 Aug 1922, Asplund s.n. (BM); Västergötland, Goteborg, 14 Aug 1959, Blom s.n. (BM); Skåne, Buutofha, Jul 1893, Johansson s.n. (BM); Scania, Kivik, Aug 1911, Johansson s.n. (BM); Malmo, Aug 1886, Johansson s.n. (BM); Östergötland, Linköping, Sep 1912, Johansson s.n. (BM); Sm. Oscasthaven, 4 Aug 1909, Köhler s.n. (BM); Yonstrup, Jul 1861, Mortensen s.n. (BM); Södermanland, Paroecia Strängnäs, Sundby, 27 Jun 1925, Samuelsson 1371 (BM); Bohuslän, Högås Parish, the innermost part of the bay Svältekilen, 14 Aug 1960, Santesson 13962 (BM).

Switzerland. Vaud: Montherod pres d’Aubonne, Anonymous s.n. (BM); inter Nyon et Lausanne ad lacum Lemanni, Jul 1838, Sander s.n. (BM); Zürich: Rümlang, Gemeinde Rümlang, Näherer Einschlag, 420 m, 20 Jun 1947, Bührer s.n. (MA).

Syrian Arab Republic. Monts Arnanis, 762 m, Aug 1906, Haradjian 481 (K); Zahleh, Cacle [coll. Prof. Post, ex. Herb. J.T. Rothrock], Post s.n. (F).

Tajikistan. Giesar, Pamiro-alay, near the village of Giesar on the river Khanaka, 31 Aug 1942, Grigoriev 757 (LE).

Turkey. Ouchak, Phrygie, Bords du ruisseau d’Ouchak (Phrygie), 910 m, 9 Jul 1858, Balansa 1137 (A, B, BM, K); Cumasia, 24 Apr 1889, Bornmüller 344 (B); Prov. Konya distr., Ermeneh (Cilican tracks), Sorivandt, 1200 m, 15 Aug 1949, Davis 16221 (K); Ankara prov., Hacikacum Valley, 11 Jun 1952, Davis & Dodds 18783 (BM, K); Cankiri-Kalecik, 9 Jun 1954, Davis & Polunin 21737 (BM); B9 Agri, Murst Valley, 3-5 km from Tutak to Hamur, 1600 m, 2 Jun 1966, Davis 44044 (K); Suluçem, (Musum) S edge of Balik G, 2300 m, 23 Jul 1966, Davis 47185 (K); C10 Hakkari, Nehil çayi, 48-55 km from Hakkari to Yuksekova, 1600 m, 14 Jun 1966, Davis 44905 (K); Cankiri-Kalecik, 9 Jun 1954, Davis 21737 (K); Tokat-Artova, 4 Sep 1954, Davis & Polunin 24854 (K); Abant Gol, Prov. Bolu, 1400 m, 11 Jul 1962, Davis & Coode D-37291 (K); Prov. Zonguldak, Bartin-Amasra, 25 Aug 1960, Khan et al. 796 (K); Çibuktal, bei Ankara, 10 Jun 1932, Kotte s.n. (K); Adapazari, Kuntay 8735 (MO); Dorukhan geçidi, Nord-seite (A4 Zonguldak), 800 m, 8 Aug 1982, Raus 6964 (B); Takiansi, 304 m, 29 Aug 1878, Regel s.n. (LE); Yagan, Erzerumskii sandzhak, river Arakas above Yagan, 9 Jul 1916, Sapozhnikov & Shishkin s.n. (LE); River Fol-chai, Trapezundskii sandzhak, 28 Jun 1917, Shishkin s.n. (LE);Vitse, Lazistan, 12 Aug 1912, Shishkin s.n. (LE); Pamphylia, Antalya, 23 Jun 1936, Tengwall 48 (GH, K, S); Bolu, Koru Motel, 850 m, 30 Aug 1972, Uotila 20137 (G); Ismid, (Nicaea), 1834, Wiedemann s.n. (LE); vicinity of Tortum Lake. East Turkey, 30 Jul 1961, Winter 278 (BM).

Turkmenistan. Germad, 8 Jun 1899, Antonov s.n. (LE); gorge Verkhanyaya Firyuza, central Kopet Dag, 31 Aug 1945, Blinoskii s.n. (LE); going up the river Dyuynee near the spring Autet, 1200 m, 25 May 1928, Bobrov & Yarmelenko 168 (LE); Karasashinskii region, river Gebe-Zvud, 14 Jul 1931, Borieova 472 (LE); Zakaspiiskaya Oblast, Shorkala [Shor-Kala], Alta-Yaba, aul Shorkala, 23 Jun 1924, Bukinich 45 (LE); Kopet Dag, road Utsch-Kuyu Nukhur, 28 Jun 1925, Fedchenko et al. 790 (LE); Germad, on the banks of the river Saki[l]yad, near village of Germad, 13 Jul 1934, Gnezdillo 97 (LE); Zakaspiiskaya Oblast’, Askabadskii area, mountains of Kopet-Dag, gorge of Chuli, 4 May 1912, Lipskii 1807 (LE); Zakaspiiskaya Oblast’, Askabadskii area, mountains of Kopet-dag, Chuli, 17 Jun 1911, Mickelson 487 (LE); Zakaspiiskaya Oblast’, [Finshchi], Jun 1895, Minkevich s.n. (LE); Tilutschi, 4000 m, 23 Jul 1879, Regel s.n. (K); Zakaspiiskaya Oblast’, Chuli Mountains, 26 Jun 1912, Samokish 1373 (LE); Zakaspiiskaya Oblast’, Chuli Mountains, near Askabad, 9 Jun 1911, Sevdturabov s.n. (LE); Kisil-arwat, Karakala, by a little river the valley of [Foltere], 10 Jul 1901, Sintenis 2035 (B, LE).

Ukraine. Kiev, 28 Aug 1906, Lonaczewsky s.n. [7272a](BM, MO); Zakarpatskaya Oblast’, Uzhgorodsky Raion, road to Imina (Transcarpatia Province, Uzhgorod District), 26 Jul 1967, Apentyeva s.n. (K).

United Kingdom. Channel Isles: Guernsey, Petit Bo, Jul 1883, Fawcett s.n. (BM); Jetou, 26 Jun 1932, Meinertzhagen s.n. (BM); Sark, 8 Jul 1953, Sowerby 4 (BM); England: Huntingdonshire, Woodwalton, 7 Jun 1914, Adamson s.n. (BM); Leicestershire, Scraptoft, 15 Aug 1949, Bangerter s.n. (BM); Warwickshire, Between Pillerton Priors and Eatington; Ettington parish, 9 Jul 1961, Bangerter 1058 (BM); North Wiltshire, Calne, 27 Aug 1914, Barton s.n. (BM); Nottinghamshire, Cresswell Gorge, east of Cresswell, 60 m, 22 Jul 1963, Bowden & Hillman 137 (BM); Yorkshire, Spern Point, 18 Aug 1070, Castroviejo s.n. (MA); Buckinghamshire, Aston Clinton parish, Dancers End Nature Reserve, 16 Aug 1942, Dandy, J.E., 943 (BM); Herefordshire, Startop’s End Reservoir. Tring Rural parish, 20 Jun 1943, Dandy 974 (BM); East Suffolk, Corporation Marshes, Walberswick; Walberswick parish, 6 Jun 1961, Elword 19 (BM); Kent, Wrotham, 21 Aug 1913, Fox 1200 (BM); West Cornwall & Scillies, Tresco harbour, Isles of Scilly, 9 Jun 1963, Gerrans 1098 (BM); Cornwall, Daymer Bay, Trebetharick, near Weybridge, 8 Jul 1967, Halliday 163 /67 (LE); East Gloucestershire, Clapton, 9 Aug 1884, Hanbury s.n. (BM); West Kent, Woolwich Arsenal, 23 Aug 1894, Herb. A.H. Wolley-Dod s.n. (BM); Oxfordshire, Banbury, 3 Aug 1863, Herb. Alfred French s.n. (BM); South Essex, Romford, Jun 1914, Herb. H. Stanley Redgrove s.n. (BM); East Cornwall, Tintagel, 26 Jun 1907, Herb. L.B. Hall 306-1134 (BM); West Cornwall & Scillies, Lizard, 27 May 1886, Herb. R.P. Murray s.n. (BM); West Gloucestershire, over Sapperton Tunnel, 18 Jun 1908, Herb. W.C. Barton s.n. (BM); Lancashire, road to Newland, Hodgson 738 (BM); Berkshire, Speen Moors near Newbury, 8 Sep 1894, Jackson s.n. (BM); East Norfolk, Lopham Middle Fen, 3 Jul 1973, Jones & Vickery 20 (BM); Hampshire, Southampton, edge of The Avenue, Southampton Common, 17 Jul 1964, Kerr 220 (LE); South Lancashire, Nr. Freshfield, Ainsdale Dunes Reserve, 26 Jul 1966, Kendrick 30 (BM); Suffolk, Brome, 19 Jul 1832, Kirby s.n. (BM); Durham, Hartlepool, North sands, Sep 1872, Lees s.n. (BM); West Lancashire, Lancaster, Shingle at Bare, Morecambe Bay, 27 May 1912, Lees s.n. (BM); Lincolnshire, Skegness, 24 Aug 1904, Linton s.n. (BM); North Essex, Frinton to Clacton, 14 Sep 1889, Linton s.n. (BM); South Hampshire, Lymington, Pilewell, 24 Aug 1893, Linton s.n. (BM); East Kent, Wittersham, May 1927, Meinertzhagen s.n. (BM); Norfolk, Sutton Broad, Sep 1928, Meinertzhagen s.n. (BM); North Devon, Woolacombe, 14 Sep 1931, Meinertzhagen s.n. (BM); South Somerset, Knowle, Jul 1881, Painter s.n. (BM); Staffordshire, Knypersley, Jul 1886, Painter s.n. (BM); Hertfordshire, Hadley Wood, 16 Jul 1882, Parker s.n. (BM); West Norfolk, Thompson, 12 Aug 1917, Robinson s.n. (BM); South Devon, Dartmouth, 30 Jul 1952, Robson 350 (BM); Dorset, Pentridge, Cranborne Chase, 19 Jun 1934, Simpson 34161 (BM); West Sussex, Chichester, Kingley Vale, West Stoke, 12 Aug 1905, Standen s.n. (BM); Cheshire, Parkgate, 12 Oct 1965, Valdés s.n. (MA); Surrey, Farncombe near Godalming, edge of R. Wey, 14 Aug 1952, Welch 4613 (BM); North Somerset, Bristol, Clifton Down, 15 Jul 1929, White s.n. (BM); Derbyshire, Monsal Dale, near the station, 21 Jun 1916, Wilmott 708 (BM); Middlesex, Buckingham Palace garden, 9 Jun 1995, Wiltshire s.n. (BM); Isle of Man: Andreas, 22 Jun 1929, Paton s.n. (BM); Scotland: East Lothian, Anonymous s.n. (BM); Fife, Burntisland, 12 Jun 1847, Anonymous s.n. (BM); East Sutherland, Creich, road between Invershin and Bonarbridge, 20 Aug 1958, Groves 2838 (BM); East Lothian, Haddington, Longniddry, 1850, Herb. J.T.I. Boswell-Syme s.n. (BM); West Perthshire, Clackmannanshire, Dollar [v.c.87], 1851, Herb. J.T.I. Boswell-Syme s.n. (BM); Mid Ebudes, Mull, near Nun’s Pass, west of Carsaig, 24 Sep 1967, Kenneth & Stirling s.n. (BM); Renfrewshire, Crossmyloof, 14 Aug 1953, Mackechnie s.n. (BM); Easterness, Beauly, 5 Aug 1892, Marshall s.n. (BM); near Mull of Kintyre, 2 Sep 1961, McClintock s.n. (BM); South Aberdeenshire, Jul 1947, Robson 349 (BM); Wales: Pembrokeshire, Pembroke, Lydstep, 28 Aug 1907, Bickham 1134 (BM); Radnorshire, Gladestry, Jul 1951, Edwards 5 (BM); Glamorgan, Sully Island, Jul 1905, Herb. H.J. Riddelsdell s.n. (BM); Merionethshire, Dolgelley, [v.c.48], 15 Aug 1923, Herb. W.C. Barton s.n. (BM); Radnorshire, Between Erwood and Aberedw, 17 Aug 1885, Ley s.n. (BM); Holyhead, Anglesey, 4 Aug 1955, Robertson D-11 (LE).

United States of America. California: Shasta County, Fall River Hills, 3 miles SW, 4 Jul 1934, Howell 12411 (GH); District of Columbia: Washington DC, National Arboretum, 15 Jul 1965, Mazzeo & Meyer 1090 (G); Idaho: Boise, 18 Jul 1911, Clark 345 (G, GH); Power County, Meader’s Trout Farm, 28 May 1961, Thiel s.n. (LE); Illinois: Kane County, Elgin, north part, 24 Jul 1928, Benke 4878 (GH); McHenry County, Fox River Grove, 18 Jun 1934, Benke 5661 (GH); Lake County, Lake Villa, 3 Aug 1906, Gleason & Shore 123 (GHDu Page County, Morton Arboretum, near Lisle, 26 Oct 1922, Palmer 22347 (A); Indiana: Wells County, Harrison TP, 31 May 1903, Deam s.n. (GH); Noble County, Kendalle, Martin Farm about 2 miles southeast of Kendalle, 23 Aug 1928, Dean 46131 (GH); Berrien County, Niles, 17 Jun 1933, Palmer 40391 (A); Putnam County, Greencastle, East Seminary Street, 1 Sep 1940, Welch 6120 (GH); Maine: Washington County, Lake View Campground, Moose Island, along western passage into Passamaquoddy Bay, N of East Port, 10 m, 30 Aug 2000, Nee & Atha 50937 (BM); Maryland: Cumberland, Jul 1849, Prior s.n. (K); Massachusetts: Hampden County, Knightville Dam, vicinity of overlook on Hwy. 112, about 1.3 mi N of Knightville, 23 Jun 1984, Bates & Elsik 83 (LE); Berkshire County, Tyringham Valley, 10 Sep 1926, Meredith s.n. (LE); Suffolk County, Franklin Park, 7 Jul 1883, Seymour s.n. (USM); Michigan: Mason County, Hamlin Lake, Ludington, 25 Jul 1910, Chaney 91 (GH); Flint, Aug 1877, Clarke 2195 (K); Cheboygan County, Douglas Lake, northern end of Lower Peninsula, Gleason 269 (K); Clinton County, East Lansing, 7 miles northeast near route 78, 9 Sep 1955, Perdue 2016 (GH); Montana: Powell County, Grant Kohrs Ranch National Historic Site, Deerlodge on US Hwy 10, 1373 m, 12 Aug 1983, Ray 194 (GH); Nevada: Washoe County, Truckee River, 3-7 miles W of Sparks, 1219 m, 6 Aug 1937, Moore & Franklin 929 (GH); New Jersey: Union County, Summit, Passiac River, 5 Jun 1969, Hellquist s.n. (GH); Cape May County, Five-mile Beach, 3 Oct 1899, MacElwee 1448 (GH); New York: Tompkins County, Ithaca, Dryden Road, near Dwyer’s Bridge, 12 Jun 1916, Bechtel 7126 (GH); Washington County, Lake George, N. Beaver Creek, Vaughn’s, north of Hudson Lake, 12 Jun 1913, Burnham s.n. (GH); Westchester County, Yonkers, Oatskill Aqueduct, 10 Jul 1934, Gleason et al. 1404 (LE, MA); Tompkins County, in creekbed of Taughannock Creek upstream from the falls, 24 Sep 1983, Kearney 132 (BM); Bronx, New York Botanical Garden, in the Swale, 28 m, 9 Jun 2006, Nee & McClelland 54448 (G); Cayuga County, Lake Como, 15 Jun 1919, Randolph & Wiegand 12864 (GH); Orange County, Aleck Meadow Reservoir, Black Rock Forest, 6 Jul 1936, Raup 7502 (GH); Yates County, Canandaigua Lake, south end, 19 Aug 1933, Swartley 66 (A); Delaware County, North Harpersfield, 11 Jun 1906, Topping 137 (A); North Carolina: Haywood County, Waynesville, US 276 near city limits, W side of highway, 823 m, 16 Aug 1969, Pittillo & Hensley 3102 (GH); Ohio: Butler County, Oxford, south of Oxford, 13 Jul 1958, Cobbe m-52 (G); Portage County, Garrettsville, 15 Jun 1895, Webb 68 (GH); Oregon: Multnomah County, St. Johns, near slough below St. Johns, 23 Jun 1915, Gorman 3575 (K); Polk County, vacant lot back of Taylor’s, 30 May 1960, Murata 71 (GH); Willamette County, Salem, 16 Jul 1917, Nelson 1684 (GH); Washington County, Hillsboro, 23 Aug 1911, Smith 4093 (GH); Pennsylvania: Northumberland County, Turbotville, 4 mi NNW of town, 13 Aug 1940, Adams 5416 (A); Berks County, McKnight’s Gap, 3/4 mile SW of town, 164 m, 12 Jun 1941, Berkheimer 2565 (GH); Buchs County, Riegelsville, 3 mi below (Kintnerville), 27 Jun 1925, Dreisbach 3573 (LE); Pike County, Kimble, along Lackawanna River, 24 Jun 1936, Fogg 10784 (A); Montgomery County, Jenkintown, 28 Jun 1938, Frazier s.n. (A); Somerset County, Tire Hill, beside Pa. Highway 53, 0.9 km SSE of Tire Hill, 427 m, 9 Aug 1956, Shetler 266 (LE); Fulton County, Needmore, 15 Jun 1947, Wahl 2514 (A); Utah: Tooele County, Open Reventment Area, zone 12, 23 Jul 2000, Long 1090 (GH); Vermont: Bellamy Falls, Anonymous s.n. (K); Leicester, 18 Aug 1907, Dutton s.n. (G); Virginia: Arlington County, Clarendon, Allard’s garden, 10 May 1938, Allard 4653 (GH); Frederick County, Belle Grove, near Middletown, 23 May 1943, Hunnewell 17854 (GH); Rappahannock County, Piney River, 27 Jun 1949, Hunnewell 19218 (GH); Washington: Yakima County, Silah Valley, 29 Aug 1902, Cotton 882 (GH); Skamania County, White Salmon, along Hwy 830 just W of White Salmon,, 10 Aug 1961, Dennis 2323 (A, G, GH); King County, Seattle, 9 Aug 1935, Eyerdam s.n. (BM, G); Spokane County, Chattaroy, 12 Jul 1923, Lackey s.n. (G); Thurston County, Mima Prairie, along the Black River 3 miles south of Littlerock, 22 Jun 1939, Meyer 1650 (GH); Whatcom County, Northwood Swamp, 26 Jun 1937, Muenscher 8394 (GH); Whitman County, Bonnie Lake, north of Rock Lake, 29 May 1938, Sharsmith 3570 (GH); West Virginia: Pendleteon County, Snowy Mountains, east slope, 13 Aug 1931, Core s.n. (GH); Smyth County, Marion, 29 Jun 1892, Small s.n. (K); Sweet Spring, 8 Sep 1903, Steele & Steele 246 (GH); Wisconsin: Dane County, Rice Lake, 19 Jun 1926, Fassett 2783 (GH); Iron County, Mercer, near outlet of Grand Portage Lake, 17 Sep 2000, Nee & Atha 50993 (BM); Brown County, Kellogg’s Work Farm, 6 Aug 1902, Schuette 96 (GH); Wyoming: Fremont County, Lander, along the Popo Agie River just south of city limits; overflow channel of river, 1615 m, 2 Aug 1965, Scott 588 (GH).

Uzbekistan. on the right bank of the Amu-dari [Amu-Darya River] between two rocks, near [Kumksntau], 15 Jul 1873, Karolkov & Krause s.n. (LE); Karakalpakiya, near Amu-dari [Amu-Darya River], 22 Aug 1874, Smirnov s.n. (LE).

Figures

Figure 35. Solanum dulcamara L. (All drawn from live plants in Battleboro, Vermont, USA). Illustration by Bobbi Angell.

Figure 36. Distribution of Solanum dulcamara L.

Figure 37. " Dulcis amara " from Hortus Eystettensis (Besler 1613). The centre plant is clearly Nigella (Ranunculaceae), but was described as a variety of Solanum dulcamara by David Don. Reproduced with permission of the Natural History Museum Botany Library.