Orobanche colorata K. Koch (1844: 291)

Piwowarczyk, Renata, Pedraja, Óscar Sánchez, Moral, Gonzalo Moreno, Fayvush, George, Zakaryan, Narine, Kartashyan, Nune & Aleksanyan, Alla, 2019, Holoparasitic Orobanchaceae (Cistanche, Diphelypaea, Orobanche, Phelipanche) in Armenia: distribution, habitats, host range and taxonomic problems, Phytotaxa 386 (1), pp. 1-106 : 95-98

18. Orobanche colorata K. Koch (1844: 291) (TL-2: 194 n. 10744, 196). Type:—ARMENIA-GEORGIA: „In Armenia [Georgiam Armeniamque]”. (B [destroyed]) ( Novopokrovskij & Tzvelev 1958: 113). Misapplied names:— O. anatolica auct., non O. anatolica Boiss. & Reut. ex Reut. in Candolle (1847: 17). Fig. 45A .

Taxonomic Note: —For some authors (e.g., Novopokrovskij & Tzvelev 1958: 113; Domina & Raab-Straube 2010 – Present), O. anatolica is a closely related species that can be distinguished by the whitish-woolly indument that covers a large part of the plant, especially the inflorescence. According to these authors, O. colorata only grows in the countries of the Caucasus area, being replaced in Turkey, Iran and Iraq by the O. anatolica s. str., although taxonomic and chorological limits corresponding to each taxon are not entirely clear. Nevertheless, in the sense of other authors ( Reuter 1847: 17, Boissier 1879: 504, Beck 1930: 275), this name includes both the species that we call O. colorata (glabrecescente) and O. anatolica (lanate). The reason for this mixture is that Reuter, when he described the new species, mentions materials from Anatolia (e.g., Caria [leg. Pinard 1843]) and from the Caucasus (e.g., Caucasian Georgia [leg. Hohenacker 1838]). The factor that prevents these two names from being synonymised is the typification of the Reuter name and this needs to be done on Anatolian material because it is the one that best fits Reuter’s description ( Reuter 1847: 17 “pilis niveis araneosis adpressis vestitus”) and preserves the current use of this name. In addition, because we think that the type of O. colorata was destroyed during World War II, we have chosen a neotype of the same region that serves as nomenclatural type according to the Koch protologue.

Orobanche anatolica Boiss. & Reut. ex Reut. in Candolle (1847: 17). Type :— TURKEY (lectotype designated here): „ Orobanche anatolica B. et R., Caria 1843, leg. C. Pinard (G-00383282! [the sheet contains two similar specimens clearly belonging to the same species and gathering]); isolectotype: (P-02982386! [the specimen on the right side of the sheet next to its label]).

Orobanche colorata K. Koch (1844: 291) . Type :— AZERBAIJAN (neotype designated here): In pratis ad radices montis Sarial [Gora Ziyarat, Khojavend District, formerly Elisabethpol Governorate], Georgia caucasica, Jun. , leg. Hohenacker [Unio itiner. 1838] (P-02982394!); isoneotype (P-02982393!).

Distribution: —Mainly in the southeastern (Syunik prov.) and northeastern parts (Tavush prov.) ( Fig. 64).

General distribution: —The Caucasus and surrounding areas of Turkey, Iran and Iraq.

Habitat: —Rocky, sandy, loamy slopes, roadsides, rarely meadows, edges of shrubs and forests, (800)1500‒ 1900(2200) m.

Hosts: — Salvia sp. ( Lamiaceae ), usually S. verticillata L., occasionally S. glutinosa L. (in herbarium materials).

Phenology: —Flowering May to June (July), fruiting (June) July–August.

Conservation status: —Least Concern (LC). The species has rather wide distribution in Armenia (4 floristic regions). Populations are varied from a few to few dozens of individuals.

Specimens examined: — ARMENIA. Gegharkunik prov.: Krasnoselsk distr. [Gegharkunik reg., Chambarak distr.], gorge above Kara-top vil. [now it is a part of Chambarak town], SE slope, 1600 m, 20 May 1962, N . Karapetyan ( ERCB); betweem Aygut and Martuni , rocky, sandy slope, 40°40’12”N, 45°13’52”E, 1520 m, 29 July 2017, R . Piwowarczyk ( KTC) ; Syunik prov.: Kapan distr., Sananlu [it is ancient settlement near Tsav village], Mtnadzor river , 23 May 1960, M . Grigoryan ( ERCB); Sananlu, left bank of the Mtnadzor river , 23 May 1960, M . Grigoryan ( ERCB, ERE); W of Tsav , sandy slope, 39°03’18”N, 46°24’54”E, 1305 m, 18 June 2017, R . Piwowarczyk ( ERCB, KTC); Goris – Khndzoresk , 18 June 1957, B . M . Bramyan ( ERCB); between Tandzatap and Aghvani , 1855 m, 1 June 2016, R . Piwowarczyk ( KTC); S of Bardzravan , 1610 m, 2 June 2016, R . Piwowarczyk ( KTC); few scattered populations, slopes and roadsides, between Kajaran and Tashtun pass, 2018 m, 2140 m, 3 June 2016, R . Piwowarczyk ( KTC); slopes and roadsides, E of Lichk , 1775 m, 3 June 2016, R . Piwowarczyk ( KTC); slopes and roadsides, NE of Tashtun , 1815 m, 3 June 2016, R . Piwowarczyk ( KTC); S of Meghri , slopes and roadsides, 800 m, 3 June 2016, R . Piwowarczyk ( KTC); SW of Tsav , roadsides and slopes, 1730 m, 4 June 2016, R . Piwowarczyk ( KTC); NE of Shvanidzor , roadsides, 1080 m, 4 June 2016, R . Piwowarczyk ( KTC); S of Karut , rocky and sandy slope, 39°14’13”N, 46°08’34”E, 1650 m, 39°15’27”N, 46°07’44”E, 1835 m, 15 June 2017, R . Piwowarczyk ( KTC); between Lernadzor and Baghajur , sandy slope near the road, 39°09’52”N, 46°12’31”E, 1546 m, 16 June 2017, R . Piwowarczyk ( ERC, KTC); NNW of Pukhrut , sandy, rocky slope, 39°08’14”N, 46°12’59”E, 1920 m, 16 June 2017, R . Piwowarczyk ( KTC); NEE of Tashtun , slopes near sideroad, 39°04’20”N, 46°11’19”E, 1830 m, 17 June 2017, R . Piwowarczyk ( KTC); rocky and sandy slope W of Vank , 39°02’23”N, 46°13’41”E, 1720 m, 39°02’34”N, 46°13’59”E, 1810 m, 39°02’33”N, 46°13’55”E, 1820 m, 17 June 2017, R . Piwowarczyk ( KTC); NW of Vank , rocky and sandy slope, 39°03’02”N, 46°14’47”E, 1930 m, 17 June 2017, R . Piwowarczyk ( KTC); between Vank and Kaler , rocky and sandy slopes, 39°03’28”N, 46°15’34”E, 2000 m, 17 June 2017, R . Piwowarczyk ( ERCB, KTC); NNW of Kaler , sandy, rocky slope, 39°03’58”N, 46°16’24”E, 2165 m, 17 June 2017, R . Piwowarczyk ( KTC); SEE of Shiskert , sandy, loamy slope, 39°04’10”N, 46°21’32”E, 1723 m, 18 June 2017, R . Piwowarczyk ( KTC); N of Verishen , grassy slope, 39°32’49”N, 46°18’42”E, 1725 m, 19 June 2017, R . Piwowarczyk ( ERCB, KTC) ; Tavush prov.: Shamshadin distr. [Tavush region], Norashen, meadow, on Salvia glutinosa , 12 June 1971, G . Sahakyan ( ERCB); Idjevan distr., Kholarker vil. [now it is a part of Idjevan town], 1954, N . Gevorkyan ( ERCB); area E of Berd, road from Berd to Aygedzor, between Ghrhi and Karmir aghbyur, 1220 m, 40°51’57”N, 45°26’41”E, 29 May 2006, M . Oganesian, H . Ter-Voskanian & E . Vitek ( ERE) .

Identification key for Orobanche species of Armenia

1. Flowers usually blue-violet, bluish-brown, rarely whitish or yellowish. Corolla-tube cylindrical, inflated below insertion of the stames. Lower lip of corolla not ciliate. Stamens inserted at the middle or slightly below middle part of the corolla-tube. Stigma whitish or yellowish-white ................................................................................................................................................................ 2

+. Flowers usually brownish, orangish, pinkish, reddish, whitish-yellow or yellow. Corolla-tube broadly cylindrical to campanulate [bell-shaped] or infundibuliformis [funnel-shaped], expanded above insertion of the stames. Lower lip of corolla ciliate or not. Stamens inserted at lower third of the corolla-tube. Stigma (pinkish, reddish, yellow or yellowish-brown) not whitish or yellowish-white .................................................................................................................................................................................................. 3

2. Leaves 10‒17(‒25) mm, ovate-lanceolate to lanceolate. Inflorescence ± densely white arachnoid-villous. Calyx usually (8‒) 15‒17 mm, calyx-segments entire or divided to half with lobes narrowly lanceolate and ± densely white arachnoid-villous. Corolla arachnoid-villous ....................................................................................................................................................... O. coerulescens

+. Leaves (5‒) 8‒13 mm, ovate to lanceolate. Inflorescence subglabrous to glandular-pubescent. Calyx usually (5‒)7‒12(‒15) mm, calyx-segments entire or ± deeply bidentate and subglabrous to glandular-pubescent. Corolla subglabrous to glandular-pubescent ............................................................................................................................................................................................................ 4

4. Leaves lanceolate to narrowly lanceolate. Bracts oval-lanceolate to lanceolate, acuminate. Calyx segments 2-toothed or entire, teeth acuminate and ± filiform to apex. Dorsal line of the corolla is geniculate in the lower part and ± almost straight in the rest. Parasitic on Lactuca sp ...................................................................................................................................................... O. grenieri

+. Leaves ovate to ovate-lanceolate. Bracts ovate to ovate-lanceolate, acute or subulate. Calyx segments 2-toothed or entire, teeth acute or subulate. Dorsal line of the corolla is geniculate in the lower part and ± uniformly curved in the rest. Parasitic on wild and cultivated plants................................................................................................................................................................................. 5

5. Plant usually stout. Inflorescences usually dense, rarely lax at the base. Flowers at the distal part usually blue-violet, bluish-brown. Corolla tube cylindrical and relatively wide in relation to its length. Parasitic mainly on Artemisia sp ........................... O. cernua

+.

3.

+.

7.

+.

8.

+.

10.

+.

9. +. 11.

+.

13.

+.

14.

+.

15.

+.

12. +.

16.

+.

17.

+.

18.

+.

Plant usually slender. Inflorescences usually lax, rarely dense at the apex. Flowers at the distal part usually bluish or pale-violet. Corolla tube cylindrical and relatively narrow in relation to its length. Parasitic on cultivated plants ( Nicotiana tabacum ), more rarely on Artemisia sp ........................................................................................................................................................ O. cumana Calyx-segments connate [cup-shaped], rarely not in adaxial side. Corolla galeate [helmet-shaped], usually dark-red or rarely brownish-red (black or nearly in dry state). Parasitic on Geranium sp ........................................................................... O. gamosepala Calyx-segments free or slightly connate at the base (abaxial side). Corolla galeate or not, usually brownish, orangish, pinkish, reddish, whitish-yellow, yellowish or yellowish-brown (± clear brown or reddish in dry state) colour........................................... 7 Corolla galeate [helmet-shaped], ± infundibuliformis [funnel-shaped] ............................................................................................ 8 Corolla not galeate, tubular-campanulate, campanulate or infundibuliformis .................................................................................. 9 Plant (10–)15–17(–24) cm high (small). Calyx-segments narrow, entire. Corolla (11–)13–15(–17) mm (small size). Parasitic on Lamiaceae ( Scutellaria sp. )......................................................................................................................................... O. zajaciorum Plant (15–)20–55(–60) cm high (medium or large). Calyx-segments wide, bifid or entire. Corolla (15‒) 20‒36 mm (medium or large size). Parasitic on Fabaceae or Rubiaceae .............................................................................................................................. 10 Inflorescence usually dense and many-flowered. Corolla brownish-yellow, yellow or reddish; lower lip of corolla not ciliate. Stamens inserted 3‒5(‒6) mm from corolla base. Stigma yellow, rarely reddish. Parasitic on Fabaceae (mainly Medicago sp. )...... ................................................................................................................................................................................................. O. lutea Inflorescence usually lax and few flowered. Corolla pinkish or pale brownish, more rarely yellow; lower lip of corolla glandular-ciliate. Stamens inserted less than 3(‒4) mm above corolla base. Stigma red, rarely yellow. Parasitic on Rubiaceae ( Galium sp. and Asperula sp. ).......................................................................................................................................................... O. caryophyllacea Stames inserted 1‒3 mm above corolla base................................................................................................................................... 11 Stames inserted 3‒6 mm above corolla base................................................................................................................................... 12 Corolla with scattered more or less sessile dark glands (punctuated appearance), lower lip glandular-ciliate. Parasitic only on Lamiaceae (mainly Thymus sp. ).............................................................................................................................................. O. alba Corolla without sessile dark glands (without punctuated appearance), lower lip glandular-ciliate or not. Parasitic on other families ............................................................................................................................................................................................ 13 Stem thick, entirely orangish to dark orangish-red and covered with numerous scales. Inflorescences always dense, many-flowered. Calyx-segments bidentate. Corolla orangish to dark orangish-red and ± concolorous (outside and inside) with ± dark red callous on folds. Stigma always yellow. Parasitic on Salvia sp. ( Lamiaceae ).................................................................................... O. colorata Stem ± slender, brownish, pinkish, reddish, yellowish or yellowish-brown and covered with a few scales. Inflorescences dense or lax, many-flowered or relatively few-flowered. Calyx-segments entire or bidentate. Corolla color ± similar to stem, without ± dark red callous on folds. Stigma dark pink, purple, reddish or yellow. Parasitic on other families ...................................................... 14 Inflorescence cylindrical to ovate, ± dense. Corolla 12‒18 mm long (medium size) with dorsal line evenly curved at the proximal and distal part, almost straight at the middle. Stigma yellow. Parasitic on Seseli sp. ( Apiaceae ) .................................. O. bartlingii Inflorescence cylindrical to short-cylindrical or ovate, ± lax. Corolla 15‒30 mm long (medium or large size) with dorsal line only slightly curved at the proximal part and almost straight at the rest. Stigma dark pink, purple, reddish, violet or more rarely orangish to yellowish. Parasitic on other families.......................................................................................................................................... 15 Inflorescence usually long to short cylindrical, rather lax, usually many-flowered. Calyx-segments entire or bidentate, rarely with 4 teeth. Corolla tubular-bell-shaped; purple, dark or light pink, rarely dirty yellow, light brown, inside more lightly coloured (yellowish). Stigma dark pink, purple, violet, rarely orange, pale to dark yellow. Parasitic on Betulaceae , Oleaceae , Fagaceae (e.g., Fraxinus , Fagus , Carpinus ) ........................................................................................................................................... O. laxissima Inflorescence usually short-cylindrical to ovate, ± lax, usually few-flowered. Calyx-segments usually entire. Corolla bell-shaped; pale violet to purple, rarely cream, inside more lightly coloured (whitish or yellowish). Stigma reddish. Parasitic on Fabaceae ( Trifolium sp. and Vicia sp. )............................................................................................................................................... O. owerini Plant always yellowish or whitish; upper lip ± porrect (continuing the dorsal line of the corolla tube) or flexed forwards........... 16 Plant usually more strongly pigmented, rarely yellowish-white; upper lip erect or recurved ......................................................... 17 Calyx (11–)13–14(–15) mm long, (2.5–)3.0–4.0 mm wide at the widest point. Corolla (22–)24–26(–28) mm long, (8–)9–10(–11) mm in diameter in the central part, pale ± pinkish-yellow, pinkish-white or yellowish-pink and usually evenly colored (not ± reddish at the apex). Parasitic on Psephellus pulcherrimus ( Asteraceae ) ............................................................................ O. arpica Calyx (7–)8–9(–11) mm long, (1.0–)2.0–2.5(–3.0) mm wide at the widest point. Corolla (15–)18–19(–21) mm long, (5–)6–7(–8) mm in diameter in the central part, yellowish and frequently ± reddish at the apex (lobes). Parasitic on Lomelosia caucasica ( Dipsacaceae ) and Stachys macrantha ( Lamiaceae ) .................................................................................................... O. javakhetica Stem with relatively few scales. Inflorescence usually shorter than the remaining stem (rarely equal to or longer). Calyx segments appreciably connate on abaxial side. Corolla yellowish-brown. Parasitic on Cirsium sp. ( Asteraceae ) .............. O. schelkovnikovii Stem with very numerous scales. Inflorescence usually equal to or longer than the remaining stem (rarely shorter). Calyx segments free. Corolla pale pinkish-yellow, pale purplish, ± reddish or yellowish. Parasitic on other families ............................................. 18 Inflorescence usually ± dense (rarely ± lax). Bracts equal or longer, rarely shorter than the corolla. Corollas evenly colored in dry state. Stames inserted (2–) 3–4 mm above corolla base. On Cephalaria sp. ( Dipsacaceae )........................................ O. grossheimii Inflorescence usually ± lax (rarely ± dense). Bracts equal or shorter, rarely longer to the corolla. Corolla lighter colored at the base in dry state. Stames inserted 4–6 mm above corolla base. On Prangos sp. ( Apiaceae )..................................................... O. kurdica

Mistakenly listed or unclear species from Armenia

Cistanche salsa (C.A. Mey.) Beck , C. ambigua (Bunge) Beck in Engler (1930: 40), C. fissa (C.A. Mey.) Beck in Engler & Prantl (1895: 129) – confused with C. armena ;

Diphelypaea helenae (Popl.) Tzvelev in Fedorov (1981: 319) – it is probably only a color form of D. coccinea ;

Phelipanche kelleri (Novopokr.) Soják (1972: 130) – confused with P. cilicica ;

P. serratocalyx (Beck) Soják (1972: 130) – confused with O. alba ;

P. hirtiflora (Reut.) Soják (1972: 130) – confused mainly with P. cilicica ;

P. aegyptiaca (Pers.) Pomel (1874: 107) – confused with P. ramosa , some need further research;

P. hohenackeri Reut. in Candolle (1847: 10) – synonym of P. heldreichii ;

P. oxyloba (Reut.) Soják (1972: 130) – confused with P. nana , P. caesia , etc.;

P. pulchella (C.A. Mey.) Soják (1972: 130) – confused mainly with P. coelestis ;

Orobanche camptolepis Boiss. & Reut. in Boissier (1879: 515) – confused with badly preserved specimen of O. kurdica ;

O. vitellina Novopokrovskij (1950: 288) – confused with O. laxissima / O. transcaucasica ;

O. hymenocalyx Reut. in Candolle (1847: 20) – probably doubtful species;

O. reticulata Wallroth (1825: 42) – confused with O. alba ;

O. grigorjevii Novopokrovskij (1950: 326) – doubtful species, confused with O. alba ;

O. pubescens d’Urville (1822: 332) – data mistake;

O. connata C. Koch (1849: 669) – doubtful species ( Novopokrovskij & Tzvelev 1958);

O. rosea Tzvelev in Komarov (1958: 687) – synonym of O. kurdica ;

O. elatior Sutton (1798: 178) – mistakenly listed, confused with O. kurdica etc.;

O. flava Mart. ex F.W. Schultz (1829) – confused with O. laxissima / O. transcaucasica ;

O. alsatica Kirschleger (1836: 109) – we find O. bartlingii in O. alsatica aggr.;

O. longibracteata Schiman-Czeika in Rechinger (1964: 17) – confused with O. kurdica . Tsaturyan & Grigoryan (1987), Tsaturyan & Gevorgyan (2009), and Zare & Yildirmli (2012) reported O. longibracteata from Armenia: Aparanskij Rajon [Kotayk province], Fontan [Fantan], but in our opinion this is O. kurdica ; additionally in Fantan we observed in the years 2015‒2017 numerous populations of O. kurdica . Moreover, O. colorata ( O. anatolica auct.) does not grow in this part of Armenia, which is confirmed by our field research as well as herbarium materials. This species is completely different from O. colorata and species from subsect. Arcuatae;

O. gracilis Smith (1798: 172) – confused with O. alba ;

O. crenata Forsskål (1775: 113) – confused with O. owerini and O. laxissima / O. transcaucasica ;

O. picridis F.W. Schultz (1830: 504) – probably confused with O. laxissima / O. transcaucasica .

Figures

FIGURE 45. General habit of: A. Orobanche colorata, B. O. bartlingii, C. O. alba, D. O. javakhetica, E. O. owerini, F. O. arpica. Photos by Renata Piwowarczyk.