Cytisus elongatus Waldst. & Kit., Descr. Icon. Pl. Hung. 2: 200, t. 183 (1804)

Sennikov, Alexander N. & Tikhomirov, Valery N., 2024, Atlas Florae Europaeae notes, 33. Taxonomic synopsis of East European species of the Cytisus ratisbonensis group (Fabaceae), PhytoKeys 238, pp. 157-197 : 157

5. Cytisus elongatus Waldst. & Kit., Descr. Icon. Pl. Hung. 2: 200, t. 183 (1804)

- Chamaecytisus elongatus (Waldst. & Kit.) Link, Handbuch 2: 155 (1831) - Cytisus hirsutus subsp. elongatus (Waldst. et Kit.) Briq., Etud. Cytis. Alp. Marit.: 168 (1894) - Chamaecytisus ciliatus subsp. elongatus (Waldst. & Kit.) Soó in Feddes Repert. 85: 439 (1974) - Chamaecytisus glaber var. elongatus (Waldst. & Kit.) Kuzmanov in Jordanov, Fl. Narodna Republ. Bulg. 6: 86 (1976).

= Cytisus leucotrichus Schur in Oesterr. Bot. Z. 10: 179 (1860), syn. nov. - Chamaecytisus leucotrichus (Schur) Czerep., Sosud. Rast. SSSR: 229 (1981) - Chamaecytisus triflorus subsp. leucotrichus (Schur) Holub in Bertová, Fl. Slovenska 4(4): 35 (1988). Type. Romania. "Rothen Berg bei Mühlbach [ Sebeș]”, [05].07.1853, F. Schur (lectotype LW00205768, designated by Pifkó (2009a: 153); isolectotype LW00205839).

= Cytisus lindemannii V.I.Krecz. in Bot. Zhurn. SSSR 25: 259 (1940), syn. nov. - Chamaecytisus lindemannii (V.I.Krecz.) Klásk. in Preslia 30: 214 (1958). Type. Ukraine. “Elisabethgrad” [Kropyvnytskyi], 06.05.1873, E. Lindemann (holotype LE01024081; isotype LE01024082). Fig. 6 .

= Cytisus czerniaevii V.I.Krecz. in Bot. Zhurn. SSSR 25: 261 (1940), syn. nov. - Chamaecytisus czerniaevii (V.I.Krecz.) Tzvelev, Fl. Evropeiskoi Chasti SSSR 6: 223 (1987). Type. Ukraine. Kharkov Region, Zmiev District, Hamlet of Fedorchenko, 24.04.1910, G.I. Širjaev (lectotype KW000114840, designated here). Other original material. Ukraine. Kharkov Region: Steppes near Chuguev, 19.05.1852, V.M. Cherniaev (KW). Sumy Region, Lebedin District, "prope Grun, in steppis princ. Kapnist" [near Grun’, in steppes of Count Kapnist = 'Mikhailovskaya Tselina’ Nature Reserve], 09.06.1905, G.I. Širjaev (KW000114839).

= Cytisus ponomarjovii Seredin in Novosti Sist. Vyssh. Rast. 13: 192 (1976), syn. nov. - Chamaecytisus ponomarjovii (Seredin) Czerep., Sosud. Rast. SSSR: 229 (1981). Type. Russia. Krasnodar Territory, Tuapse District, 1 km NW of Dzhubga Village, oak forest, 08.07.1973, R.M. Seredin (holotype LE).

= Chamaecytisus korabensis Pifkó & Barina in Stud. Bot. Hung. 47(1): 164 (2016), syn. nov. Type. Albania. Qarku i Dibrës: [Korab-Koritnik Nature Park,] Mali i Bardhë Mts, near peak Maja e Pelpenikut, above village Sllatinë, on evaporites, 41.78419°N, 20.45978°E, 1928 m, 17.06.2013, Z. Barina & D. Pifkó 22354 (holotype BP759110; isotype BP759111).

Type.

Romania. Historical Banat Region : "In sylvis Beregh, Banaticis et Croaticis", 1800, P. Kitaibel (lectotype W20030003241, left-hand fragment, designated here: https://w.jacq.org/W20030003241). Possibly Ukraine. [" In comitatis Bereghensis " = Bereg County, "in sylvis",] Herb. Waldstein (superseded lectotype PR155757/738a, designated by Chrtek and Skočdopolová (1982: 226)) .

Description.

Upright shrubs with erect, basally ascending stems up to 40-60(80) cm tall and long branches. Leaves with elliptic to obovate leaflets, densely hairy above, with lax hairs 0.4-0.8 mm long below, petioles rather densely covered with laxly appressed to subpatent hairs. Flowers strictly lateral, 1-4 in axils, on pedicels 2-4 mm long, yellow; calyx 11-12 mm long, with subpatent hairs 0.8-1.2 mm long; standard suborbicular, glabrous or hairy above.

Distribution.

Europe: France (along the valley of Rhône: Tison and de Foucault (2014)), Italy, Albania, Serbia, Greece, Bulgaria, Turkey ( Cristofolini 1991), Bosnia and Herzegovina (new record), Montenegro (new record), Slovenia (new record), Croatia ( Lovašen-Eberhardt 1997), North Macedonia ( Micevski 2001), Austria ( Cristofolini 1991), Hungary ( Pifkó 2009b), Slovakia ( Cristofolini 1991), Romania ( Grinţescu 1957), Moldova ( Heydemann 1986), Ukraine ( Kreczetowicz 1940; Fedoronchuk 2022), Belarus (new record), Russia (south-western part) ( Kreczetowicz 1940; Borisova 1964; Tzvelev 1987). Asia: Russia (western and central Caucasus: Grossheim (1952); Zernov (2006); Ivanov (2019)), Abkhazia ( Kolakovsky 1985), Georgia (Ajaria: Gvinianidze (1981)), Turkey (Artvin Province: Kreczetowicz (1940)). Reported for the first time from Belarus and Bosnia and Herzegovina here. New to Bryansk and Lipetsk Regions of Russia. The actual distribution in Asian Turkey and the Balkans may be more extensive, but has been obscured due to the confusion with C. hirsutus ( Gibbs 1970).

Ecology.

In the forest zone, this species occurs in sparse forest stands and on forest margins with steppe herbaceous species, mostly in xerophilous oak forests, at elevations below 500(700) m; in the forest steppe and steppe zones, it is found among sparse shrubs in dry creeks, steppe-like meadows and steppes. It also occurs in oak forests and steppe-like meadows in the mountains.

Chromosome counts.

2n = 50 ( Semerenko 1984); material collected from native populations in Kursk and Lipetsk Regions; vouchers at MSK. Dubious record: 2n = 48 ( Frahm-Leliveld (1957), as Cytisus elongatus ); cultivated material; vouchers unknown.

Notes on nomenclature.

Skalická (1986) and Cristofolini (1991) accepted Cytisus triflorus Lam. as the priority name for this species. Its lectotype actually belongs to C. hirsutus L. ( Sennikov and Tikhomirov 2024b).

Cytisus elongatus was described from present-day Romania ( Caraş-Severin, Banat) and Ukraine (former Bereg County) ( Waldstein and Kitaibel 1804). The original description of C. elongatus refers to plants with elongated branches and numerous flowers in lateral inflorescences, flowers shortly pedicellate and "slightly larger than in C. supinus ", branches with appressed hairs, leaves greyish-pubescent on both sides and calyces grey because of dense pubescence. The presumed original material ( Pifkó 2007) is apparently heterogeneous, but the original description and drawing clearly indicate the intention to describe a species of C. ratisbonensis s.l. with the calyces having long subappressed pubescence and the leaves being hairy on the upper side, which unambiguously point at the species known as C. lindemannii ( Tzvelev 1987) or C. triflorus ( Cristofolini 1991).

According to the published diaries of P. Kitaibel ( Gombocz 1945), he collected C. elongatus in Bereg County (7 July 1803, forest near Bereg, present-day Beregovo Town, mentioned as C. elongatus ) and in Banat Region (26 July - 11 August 1800, many places, mentioned, according to Pifkó (2007), as C. patens ). This means that the taxonomic concept of C. elongatus had been shaped on the basis of the Banat material prior to the travel to Bereg County. In Bereg County, besides the locality mentioned in the diary, where the plants were collected in fruits due to the late season, the species could have been collected anywhere on the route in northern and north-eastern Hungary.

After the protologue of C. elongatus was published, Kitaibel collected further specimens of this species ( Lőkös 2001). In 1805, he travelled to Banat for the second time (5 July, near “Szlatina” = Slatina-Timiș, Caraș-Severin County, Romania, as C. elongatus ). In 1815, he revisited Transcarpathia and collected in “Rhonaszegh” (6 August, Coștiui, Maramureș County, Romania, as C. elongatus ) and near Bereg (25 September, Beregovo, Ukraine). The actual collections of Kitaibel may not have been limited to the localities mentioned in the diaries, but these data may be used as guidance to shape our understanding of the collections. For example, Kitaibel had an opportunity to collect the species during his three travels to the Matra Mts. and also in other travels that included present-day Croatia and Romania.

Chrtek and Skočdopolová (1982) designated a lectotype of C. elongatus from the collections of F. de Paula von Waldstein at the National Museum in Prague. The specimens kept as C. elongatus in this collection are accompanied by a generic label written by K. Sternberg, who possessed the collections after Waldstein’s death, whereas their original label data are lacking. Four plants are kept on two sheets under a single label. Of these plants, two were designated as a lectotype.

Chrtek and Skočdopolová (1982) preferred the designated sheet because the other one was a mixed collection of two different plants. However, they failed to observe that the two lectotype plants are also apparently different. The lanceolate leaflets of the right-hand plant of the lectotype are in apparent conflict with the protologue that states "foliolis obovatis"; besides, its inflorescence looks capitate rather than elongated as stated in the protologue ("totos ramos annotinos undique dense tegentes"). The other fragment agrees with the protologue in morphology, but there is no evidence that this particular material can be associated with the protologue and was not collected in any of the numerous later travels of Kitaibel. Due to the lack of the association with the protologue, the lectotype of C. elongatus designated by Chrtek and Skočdopolová (1982) cannot be accepted and should be superseded in favour of some certain element of the original material that is in agreement with the protologue.

In search for the other original material, we examined online collections of B, BP, PRC and W. Specimens in Herbarium Willdenow at B, which are labelled “Hungaria”, are likely original material because Willdenow received manuscripts and specimens from Waldstein and Kitaibel, of which hundreds are currently kept in Berlin ( Hiepko 1972). Two of these specimens represent elongated branches, of which one (B-Willd 13622-03) has the leaves glabrous on the upper side and belongs to C. cinereus , whereas the other (B-Willd 13622-04) has the leaves hairy on the upper side and belongs to C. triflorus sensu Cristofolini. Plants collected from Bereg County are represented at PRC (PRC 454937), but their elongated branch has the leaves glabrous above and belongs to C. cinereus . Some original material collected in Banat is kept at BP ( Pifkó 2007), including a specimen with elongated branches (Hb. Kitaibel XXIV: 161) collected near “Oravicza” (Oravița, Caraș-Severin County, Romania).

The most important specimen was found at W (W 20030003241). The plants on this sheet were identified as C. elongatus with a reference to the protologue; the label of this specimen written by Kitaibel is composite and reads "In sylvis Beregh, Banaticis et Croaticis". This label reflects Kitaibel’s travels to Banat in 1800, to Croatia in 1802 and to Bereg County in 1803; it makes the specimen firmly linked to the protologue of C. elongatus . The sheet bears three fragments: a branch on the right side, densely leafy and abundantly flowering, corresponding to C. cinereus ; a small fragment in immature fruit in the middle, also belonging to C. cinereus (possibly collected in 1803 from the locality in Bereg County mentioned in Gombocz (1945)); and an elongated branch in flower on the left side, whose calyces are villous and leaves are densely hairy above. The latter fragment fully agrees with the protologue of C. elongatus . We assume that the left-hand specimen belongs to the plants collected by Kitaibel in Banat in 1800 and used for the original description of C. elongatus and, therefore, designate it as a new lectotype.

This lectotype agrees with the usage in the Hungarian exsiccata ( Kerner 1884; Anonymous 1919) and other specimens identified as C. elongatus , later usage favoured the application of this species name to C. hirsutus s.l. and the illustration was considered mismatching the original description ( Kerner 1884). The usage of C. elongatus by Skalická (1986) and Pifkó (2009b) agrees with our lectotypification (except for their inclusion of plants belonging to C. cinereus ); the placement of C. elongatus to the synonymy of " C. triflorus " by Cristofolini (1991) also agrees with our taxonomy.

The treatment of C. leucotrichus has been controversial. Schur (1859) described this plant as deviating from C. hirsutus by a denser “white” (sericeous) pubescence and smaller leaves. Tzvelev (1987) and Cristofolini (1991) placed it to the synonymy of C. hirsutus , in spite of its dense subpatent pubescence on branches and leaves (vs. sparse patent pubescence in C. hirsutus ). Holub and Bertová (1988) accepted and subordinated it to C. triflorus , which was a name for C. elongatus at that time. The type collection of C. leucotrichus is represented by large branches in fruit, which are densely covered by subappressed (partly subpatent) hairs. This type of pubescence matches the characters of " C. triflorus " ( Cristofolini 1991) and C. lindemannii ( Tzvelev 1987) and agrees with the taxonomic concept of C. elongatus accepted here.

Although Kreczetowicz (1940) already employed the type concept, he did not indicate a type of Cytisus czerniaevii . Neither did he cite any specimen in the protologue; instead, he listed two localities in Kharkov Region. We found three specimens corresponding to those localities and identified by Kreczetowicz as Cytisus lindemannii × C. ruthenicus , in agreement with the hybrid origin of C. czerniaevii indicated in its protologue. One specimen was collected by Vasily Cherniaev and formerly deposited at CWU (which was transferred to KW after the Second World War), in the Ukrainian collections of Cherniaev which were extracted from his personal herbarium and placed within the main collections of KW for the preparation of Flora of the Ukrainian SSR ( Krytzka et al. 2002). This specimen apparently provided the reason for naming the hybrid. Two specimens were collected by Grigory Širjaev in the former Kharkov Region of the Russian Empire (now Kharkov and Sumy Regions of Ukraine).

All the original material of C. czerniaevii belongs to C. elongatus . Kreczetowicz (1940) stated that his hybrid differed from the species by its subglabrous standard, which is, however, variable in C. elongatus ( Wissjulina 1954). For this reason, the name C. czerniaevii cannot be used for a hybrid between C. ruthenicus and C. elongatus , but is a synonym of the latter.

Krytzka et al. (1999: 610) believed that the holotype of C. czerniaevii is kept at LE, but cited the species provenance from the protologue instead of the label data. Fedoronchuk et al. (2003) did not mention the presence of the original material of C. czerniaevii at KW. This material was recognised as such in 2012 by M. Shevera (on herbarium labels).

Notes on taxonomy and distribution.

Ledebour (1843) distinguished between the plants with appressed and subpatent hairs on the calyces, which he called C. biflorus L’Her. and C. elongatus Waldst. & Kit., respectively. The plants with the subpatent pubescence were reported from the steppe zone of Eastern Europe for the first time by Lindemann (1867), who used the nomenclature from Ledebour (1843).

Kreczetowicz (1940) believed that C. elongatus s. str. is replaced in steppes of Eastern Europe (Ukraine) and the North Caucasus by another taxon with a hairy (vs. glabrous) standard and a denser pubescence, which he named C. lindemannii . Skalická (1986) and Tzvelev (1987) accepted C. lindemannii in the same sense. Since this widely distributed species is variable in the length and density of pubescence and Kreczetowicz (1940) himself admitted that the pubescence on standard is variable within one species, we do not consider the western and eastern plants to be taxonomically different and restore the priority name for this species, C. elongatus . Cristofolini (1991) reduced C. elongatus to a synonym of " C. triflorus ", but placed C. lindemannii in the synonymy of C. ruthenicus ; the latter decision is against the original description and type material of C. lindemannii , which has the subappressed to patent pubescence (vs. appressed in C. ruthenicus ) and the leaves hairy above (vs. glabrous above in C. ruthenicus ) ( Kreczetowicz 1940).

Kreczetowicz (1940), Grossheim (1952) and Portenier and Solodko (2002) treated C. hirsutissimus as endemic to the Caucasus, a mountainous species which reportedly differed from the steppic, lowland East European C. lindemannii (= C. elongatus ) in longer pedicels and a patent (vs. subappressed) pubescence of the whole plant. These minor and variable characters cannot be considered species-specific, and C. hirsutissimus of these authors was correctly identified with " C. triflorus " (= C. elongatus ) ( Cristofolini 1991).

Seredin (1976) described C. ponomarjovii as a local endemic of the western Caucasus and distinguished it from C. caucasicus by its denser pubescence. Cristofolini (1991) omitted this species, which was accepted in very few works ( Czerepanov 1995; Ivanov 2019). Portenier and Solodko (2002) correctly noted that C. ponomarjovii , a species of lower elevations, corresponds to ' C. hirsutissimus C. Koch’ of Russian authors ( Kreczetowicz 1940; Grossheim 1952; Portenier and Solodko 2002), which is C. triflorus in the sense of Cristofolini (1991). We place it to the synonymy of C. elongatus , accordingly.

Chamaecytisus korabensis was recently described by Pifkó and Barina (2016) as a local endemic of north-western Albania, which was considered as related to "the C. ratisbonensis and C. triflorus agg." The protologue described and illustrated a minute plant collected at higher altitudes, with ascending stems covered by subappressed pubescence, leaves appressedly pubescent on both sides, and calyces 1-1.3 mm long with abundant subpatent hairs. These characters correspond to alpine forms of C. elongatus , which may be highly reduced in size in the subalpine mountain belt, whereas the differences in plant size played a major role in identification according to Pifkó and Barina (2016).

The earlier records of C. lindemannii from Belarus ( Fedtschenko 1950) belong to C. semerenkoanus , but the presence of this species in the country is confirmed on the basis of recent collections.

Conservation status.

Although the species is not included in national or regional Red Lists, it occurs in some protected areas, for example, in the Mikhailovskaya Tselina Nature Reserve in Ukraine and in the Utrish Nature Reserve in Russia.

Figures

Figure 6. Holotype of Cytisus lindemannii V. I. Krecz.