Orobanche colorata K. Koch (1844: 291)

Piwowarczyk, Renata, Pedraja, Óscar Sánchez, Khutsishvili, Manana & Kharazishvili, Davit, 2023, Holoparasitic Orobanchaceae in Georgia (Caucasus): taxonomic revision, diversity, distribution, habitats and host range, Phytotaxa 604 (1), pp. 1-103 : 90-94

23. Orobanche colorata K. Koch (1844: 291)

(TL-2: 194 n. 10744, 196). Type:—ARMENIA-GEORGIA: “In Armenia [Georgiam Armeniamque]” (B [destroyed]) ( Novopokrovsky & Tzvelev 1958: 113) (neotype designated by Piwowarczyk et al. 2019: 95): In pratis ad radices montis Sarial [ Azerbaijan, Gora Ziyarat, Khojavend District, formerly Elisabethpol Governorate], Georgia caucasica, Jun. , leg. Hohenacker [Unio itiner. 1838] (P02982394!); isoneotype (P02982393!). Misapplied names:— O. anatolica auct., non O. anatolica Boiss. & Reut. ex Reut. in Candolle (1847: 17). Fig. 64 .

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

Distribution: —Scattered single localities in Abkhazia, Adjara, South Ossetia, and Samegrelo-Zemo Svaneti, Guria, Samtskhe-Javakheti, Mtskheta-Mtianeti, Tbilisi, Kakheti, Kvemo Kartli provinces ( Fig. 65 ).

Habitat: —Rocky, sandy, loamy or steppe slopes, roadsides, subalpine meadows, edges of shrublands and forests, rarely in anthropogenic habitats, near cemeteries, usually (500)700‒1700(2100) m.

Hosts: — Salvia sp. (Lamiaceae) , here usually S. verticillata L., S. nemorosa L., rarely S. glutinosa L.

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

Conservation status: —Least Concern (LC). The species has wide distribution in Georgia, but populations are fragmented and usually not numerous. EOO is ca. 50,000 km 2 and AOO is less than 1000 km 2. The main threats are loss/degradation of habitats, especially through overgrazing, development of fields, housing and road infrastructure, landslides, excessive overgrowing.

Taxonomic note: —See Piwowarczyk et al. 2019.

S pecimens examined: — GEORGIA. Abkhazia: Tsebelda, on Salvia glutinosa , April 1899, Woronow (LE); Tsebelda, on Salvia glutinosa , 19 April, 2 May, 1902, G. Woronow (LE); “ Abchasien: bei Gerjewskoje [Jurjewskoje / Jurjewskoye / Jurjevskoje / Smjewoskoje / Serjevskoje (Tsebelda)] im Juni 1902 auf Salvia glutinosa L. (G. Woronin [leg. G. Woronow]!)” ( Beck 1930: 260, sub O. kurdica [rev. Novopokrovsky & Tzvelev 1958: 106, sub O. colorata ]) [https://www.mobot.org/MOBOT/Research/LEguide/collections/119/index.html]; Adjara prov.: riv. Kintrishi, vil. Zeraboseli, Adjara, river side, shrublands, 21 May 1963, V. Memiadze (BATU); “Batum [Batumi, Adjaria] im Distr. Artwin [Artvin Province, today Turkey] bei Usot im Mai (G. Woronow!)” ( Beck 1930: 276, sub O. anatolica ); Guria prov.: distr. Chokhatauri, on Salvia sp. , 12 May 2015, G. Bolkvadze & N. Memiadze (BATU); Kakheti prov.: SE of Kvemo Bodbe, steppe slopes in the hills, 41°32.21’N, 45°54’E, 760 m, 9 May 2019, R. Piwowarczyk (KTC); between Kvemo Bodbe and Gamarjveba, slopes, 41°31.4’N, 45°55.5’E, 760 m, 9 May 2019, R. Piwowarczyk (KTC). Kvemo Kartli prov.: Kvemo Kartli Region, Minor Caucasus, Mt. Didgori [Trialeti Range, Lesser Caucasus], surroundings of the small village Der, 1200–1300 m, on Salvia sp. (Lamiaceae) , 29 May 2002, G. Schneeweiss (WU32436); Mtskheta-Mtianeti prov.: Pasanauri, along the Georgian Military Road, 3 June 1889, V. Li…? (LE); NE of Lutkhubi, Gudamakri range, slopes of Lutkhubi river, 42°22’55.191”N, 44°48’10.045”E, 24 May 2022, V. Kalníková [phot.]; N of Khomi, Gudamakri range, slopes of Aragvi river, 42°23’45.864”N, 44°55’4.287”E, 31 May 2022, V. Kalníková [phot.]; Mtiuleti, SE of Gudauri, 1600 m, Fischer et al. (2018) [phot.]; Mtiuleti, Stepancminda, Fischer et al. (2018) [phot.]; Samegrelo-Zemo Svaneti prov.: “ Mingrelien und Oberswanetien, near Mazeri, 43°05’03.6”N, 42°36’31.5”E, [ca. 1730 m], 28 May 2018, Bernd-J. Seitz (https://www.inaturalist.org/observations/107418824, accessed Jul 2022); Tvebishi, 43°04’15.0”N, 42°35’47.4”E, [ca. 1700 m], 9 June 2022, J. Pietruszewski [phot., https://www.inaturalist. org/observations/121049470]; Adishi [slopes of Adishchala river], 42°59’45.5”N, 42°54’57.6”E, [ca. 2070 m], 42°59’46.2”N, 42°54’56.9”E, 12Jun2022, J.Pietruszewski (phot.,https://www.inaturalist.org/observations/121591657, https://www.inaturalist.org/observations/121591589 [yellow form]); Samtskhe-Javakheti prov.: Borjomi distr., NE of Timotesubani, slopes, roadsides, around the cemetery, 41°48’52”N, 43°31’20”E, ca. 1200 m, 2 June 2014, R. Piwowarczyk (KTC); Timotesubani, slope near monastery, 41°48’35”N, 43°31’03”E, 1150 m, 2 June 2014, R. Piwowarczyk (KTC); Tbilisi prov.: Transcaucasia, Tiflis, in familus fl. Dabachanca [part of r. Dabakhanka gorge in Botanical Garden], 6 June 1920, A. Grossheim (ERE) [as O. caryophyllacea ]; South Ossetia: Ermani, Khudisen, subalpine meadow opposite Britata gorge, 2000 m, 16 June 1948, V. Surova (MW0723365).

Difficult to locate: Iberia, m. ill., 1797, Steven (LE); “Transcaucasia: prope Tiflis [Tbilisi] leg. Bayern!; in Iberia

[Caucasian Iberia / Iveria] leg. Wilhelms!” ( Beck 1890: 193, sub O. anatolica ).

Identification key of Orobanche species of Georgia (after Piwowarczyk et al. 2019, modified)

1. Flowers usually blue-violet, bluish-brown, rarely whitish or yellowish. Corolla-tube cylindrical, inflated below insertion of the stamens. 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 stamens. 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 [scales] 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 usually on Lactuca , rarely Tragopogon .............................................................................................................. 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. Parasitic on cultivated plants (e.g., Nicotiana tabacum ), rarely on Artemisia [in Georgia mainly on Artemisia and occasionally on Bassia ] .................................................................................................................................................... O. cumana

3. 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 .......................................................................... 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........................................... 5

5. Corolla galeate [helmet-shaped], ± infundibuliformis [funnel-shaped] ............................................................................................ 6

+. Corolla not galeate, tubular-campanulate, campanulate or infundibuliformis .................................................................................. 7

6. Plant (10–)15–17(–24) cm (small). Calyx-segments narrow, entire. Corolla (11–)13–15(–17) mm (small size). Parasitic on Lamiaceae (Scutellaria) ................................................................................................................................................ O. zajaciorum

+. Plant (15–)20–55(–60) cm (medium or large). Calyx-segments wide, bifid or entire. Corolla (15‒) 20‒36 mm (medium or large size). Parasitic on Fabaceae or Rubiaceae ......................................................................................................................................... 8

8. 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 )............ ................................................................................................................................................................................................ 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 and Asperula ) ................................................................................................................................................................ O. caryophyllacea

7. Corolla tubular (contracted at the throat). Stigma usually yellow (very rarely purplish). Parasitic only on Araliaceae species (usually on Hedera )......................................................................................................................................................................... O. hederae

+. Corolla tubular (not contracted at the throat), tubular-campanulate, campanulate or infundibuliformis. Stigma yellow or with another colour (purple, red, orange). Never parasitic on Araliaceae species .................................................................................... 9

9. Stamens inserted 1‒3 mm above corolla base................................................................................................................................. 10

+. Stamens inserted 3‒6 mm above corolla base................................................................................................................................. 11

10. Corolla shiny dark red [sanguineus] inside. Stigma yellow. Parasitic only on Fabaceae (mainly Cytisus ) species .......... O. gracilis

+. Corolla not shiny dark red inside. Stigma yellow or purple. Parasitic on Fabaceae and other families ......................................... 12

12. Corolla with scattered more or less sessile dark glands (punctuated appearance) or ± completely dark-red (outside), lower lip glandular-ciliate. Calyx-segments usually entire. Parasitic only on Lamiaceae (mainly Thymus ) or Campanulaceae (mainly Campanula ) ..................................................................................................................................................................................... 13

+. Corolla without sessile dark glands (without punctuated appearance) or never dark-red (outside), lower lip glandular-ciliate or not. Calyx-segments entire or bidentate. Parasitic on Lamiaceae (only on Salvia sp. ) and other families (never Campanulaceae ) ...... 14

13. Corolla campanulate, with scattered more or less sessile dark glands (punctuated appearance), sometimes ± dark-red (outside), dorsal line ± almost straight at the middle part. Stamens ± (not densely) hairy at the base. Stigma dark-red, red, purple or yellow. Parasitic only on Lamiaceae (mainly Thymus )........................................................................................................................ O. alba

+. Corolla tubular to tubular-campanulate, ± dark-red to reddish (outside), dorsal line ± regularly curved at the middle part. Stamens densely hairy at the base. Stigma purple or dark-red. Parasitic only on Campanulaceae (mainly Campanula )............. O. raddeana

14. 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 ± concolourous (outside and inside) with ± dark red callous on folds. Stigma always yellow. Parasitic on Salvia (Lamiaceae) ................................................................................... O. colorata

+. Stem ± slender, brownish, pinkish, reddish, yellow 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 colour ± similar to stem, without ± dark red callous on folds. Stigma dark pink, purple, reddish or yellow. Parasitic on other families (never on Salvia )........................... 15

15. Corolla 10‒18(‒19) mm long (small to medium size), tubular ...................................................................................................... 16

+. Corolla 15‒30 mm long (medium or large size), campanulate [bell-shaped], tubular-infudibuliform [funnel-shaped] or campanulateinfundibuliform [bell-shaped to funnel-shaped].............................................................................................................................. 17

16. Inflorescence cylindrical to ovate, short, ± dense. Corolla 12‒18 mm long, usually yellowish-brown or yellowish-pink or reddishpurple to reddish-brown, with dorsal line evenly curved at the proximal and distal part, almost straight at the middle, upper lip with ± erect lobes. Stamens inserted 1‒3 mm above corolla base, ± densely pubescent from the base to the middle. Stigma yellow. Parasitic on Seseli rarely Cachrys (Apiaceae) ................................................................................................................ O. bartlingii

+. Inflorescence cylindrical, long to short, ± lax. Corolla 10‒17(‒19) mm long, usually yellowish-white and ± tinged with purple on the veins and at the upper lip, with dorsal line ± evenly curved, upper lip with ± patent lobes. Stamens inserted 2‒3 mm above corolla base, sparsely pubescent at the base. Stigma purple (rarely yellow). Parasitic on many families ( Apiaceae , Asteraceae , Fabaceae , etc.) ...................................................................................................................................................................... O. minor

17. Stem (35–) 60–100 cm. Corolla (20–)23–24(–25) mm long, campanulate [bell-shaped]. Stamens inserted 1(–2) mm above the corolla base. Parasitic on large Apiaceae (mainly Heracleum ) ........................................................................................... O. ingens

+. Stem 13–65(–70) cm. Corolla 15‒25(‒30) mm long, tubular-infudibuliform [tubular to funnel-shaped] or campanulateinfundibuliform [bell-shaped to funnel-shaped]. Stamens inserted (1.5–)2–4(–5) mm above the corolla base. Parasitic on other families (never Apiaceae ) ................................................................................................................................................................ 18

18. Corolla yellow or brownish-yellow, dorsal line regularly arched-curved. Stamens inserted (1.5–) 2–3 mm above the corolla base. Parasitic on Asteraceae (mainly Inula )................................................................................................................................. O. inulae

+. Corolla yellow or other colours. Stamens inserted 2–4(–5) mm above the corolla base. Parasitic on other families (never on Inula ) .............................................................................................................................................................................................. 19

19. Inflorescence usually long to short cylindrical, rather lax, usually many-flowered. Calyx-segments entire or bidentate, rarely with 4 teeth. Corolla tubular-infudibuliform [tubular to funnel-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. Forests. Parasitic on trees and shrublands (woody plants) of e.g., 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 campanulateinfundibuliform [bell-shaped to funnel-shaped]; pale violet to purple, rarely cream, inside more lightly coloured (whitish or yellowish). Stigma reddish. Usually subalpine and alpine meadows. Parasitic on herbs (non-woody plants) of Fabaceae (mainly Trifolium ) and rarely Asteraceae (Leontodon) .................................................................................................................. O. owerinii

11. Plant always pale, yellowish or whitish; upper lip ± porrect (continuing the dorsal line of the corolla tube) or flexed forwards. 20

+. Plant usually more strongly pigmented, rarely yellowish-white; upper lip erect or recurved ........................................................ 21

20. Corolla tubular to narrowly campanulate (slightly widening towards the mouth); dorsal line ± flexed forwards or straight at the apex. Parasitic on Stachys macrantha (Lamiaceae) and also on Lomelosia (Dipsacaceae) ( Armenia))...................... O. javakhetica

+. Corolla broadly tubular-campanulate to campanulate (conspicuously widening towards the mouth); dorsal line ± porrect at the apex. Parasitic on other species (never on Lamiaceae or Dipsacaceae ).......................................................................................... 22

22. Corolla (17–)20–23(–25) × (7–)10–12(–14) mm, usually campanulate or more rarely tubular-campanulate, dorsal line slightly curved (but straight at the apex); lobes oval or rectangular to triangular, with irregularly dentate margin. Parasitic on Aconitum (Ranunculaceae) ..................................................................................................................................................... O. mlokosiewiczii

+. Corolla (21–)22–23(–25) × (6.8–)7.0–7.8(–8.3) mm, tubular, strongly curved (but ± deflexed at the apex); lobes spathulate rounded, with finely and irregularly denticulate and glandular margin. Parasitic on Caucasalia , Pojarkovia , Senecio , Prenanthes , ( Asteraceae ) .................................................................................................................................................................... O. cicerbitae

21. Stem with scales scattered not overlapping each other. Inflorescence usually ± lax (rarely ± dense). Flower erect-patent to patent. Corolla yellow, orange or reddish-brown, ± concolourous (outside and inside). Stamens inserted 3–6 mm above corolla base. Parasitic only on Asteraceae (mainly on Petasites )................................................................................................................ O. flava

+. Stem with scales very numerous, almost overlapping with each other. Inflorescence usually ± dense (rarely ± lax). Flower erect to erect-patent. Corolla yellowish, yellowish-brown, whitish ± tinged with purple, clearer inside (light yellow or whitish). Stamens inserted (2–) 3–4 mm above corolla base. Parasitic only on Dipsacaceae (Cephalaria) .............................................. O. grossheimii

Mistakenly listed or unclear species from Georgia

Cistanche salsa (C.A. Mey.) Beck (1895) – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018); confused with C. fissa , see also taxonomic note in C. fissa .

Phelypaea helenae (Popl.) Tzvelev in Fedorov (1981: 319) – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018), taxon described from Crimea (LE!) is included within P. coccinea , and was probably only a colour variation (corolla and stem orange yellow), as both have the same host.

P. tournefortii Desf. – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018), confused with P. coccinea ; herbarium revision and field studies did not confirm this species.

Phelipanche kelleri (Novopokr.) Soják (1972: 130) – species described from eastern Kazakhstan and cited from Georgia by Rätzel et al. (2018a): Georgia: Greater Caucasus, Mzcheta-Mtianeti, near Karsani, Sarkinetis Kedi, 41.835519°N, 44.714053°E, 640 m, Sexposed slope, stony, open sunny site, collected together with fragment of host [ Bassia prostrata (L.) Beck, but root attachment was not verified], 11 May 2017, Gr̂ger & Shewardnadze 2206 (M); ibid., Caucasus, Gouv. Tiflis, 14 Jun 1889, Lipsky (LE). Georgian distribution needs to be revised. In May 2019 , the primary author (R. Piwowarczyk) made an attempt to find the cited locality in the field, near Karsani. Despite intensive searches in this area, only numerous populations of P. nana parasitizing mainly Erodium cicutarium were found. It was also possible to find the slope (see photo by A. Gr̂ger in Rätzel et al. 2018a) with B. prostrata , but only P. nana was found in the vicinity.

P. mutelii (F.W. Schultz) Pomel (1874: 106) – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018); confused mainly with P. nana , herbarium revision and field studies did not confirm this species.

P. oxyloba (Reut.) Soják (1972: 130) – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018); confused with P. nana , P. pulchella , etc., see also comments in P. pulchella and P. libanotica .

P. septemloba (Beck) Soják (1972: 130) – synonym of P. pulchella , see comments in P. pulchella .

Orobanche crenata Forsskål (1775: 113) – confused with O. owerinii and O. laxissima .

O. elatior Sutton (1798: 178) – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018, Fischer et al. 2018); mistakenly listed; herbarium revision (confusion mainly with O. grossheimii ) and field studies did not confirm this species.

O. kurdica Boiss. & Hausskn. in Boissier (1879: 505) – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018); herbarium revision and field studies did not confirm this species.

O. pubescens d’Urville (1822: 332) – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018); data probably mistaken; herbarium revision and field studies did not confirm this species, the closest known place of the species is located on the north coast of the Black Sea (Krasnodar Krai, Caucasian Russia). [Samtskhe-Javakheti: “Transkaukasia: Abas-Tuman [Abastumani, Adigeni Municipality] (leg. Owerin)!” ( Beck 1890: 240, sub O. versicolor ); “Transkaukasien: bei Abas Tuman (Owerin!)” ( Beck 1930: 185, sub O. versicolor ); “Caucasus: Western Transcaucasia” ( Novopokrovsky & Tzvelev 1958 [2000]: 104[98], sub O. versicolor ); Georgia, Artvinskij Rajon, Berg Artvine, [ Turkey territory], 7 June 1914, Turkevitsch (LE s. n.) (in Teryokhin et al. 1993: 103). Orobanche squalida Steven in Bull. Soc. Nat. Mosc. 30: 354 (1857) ( Novopokrovsky & Tzvelev 1958 [2000]: 97[125]). Ind. loc.: “In Tauria montosa [Crimea] hinc inde; etiam in Iberia [ Iberia, Georgia]”]. At a distance of about 20–30 km in a straight line from Abastumani, there are several very numerous localities of O. minor (e.g., from Dviri to Kortaneti) with which this species may have been confused. Besides, in the near vicinity of Abastumani, there are numerous populations of O. laxissima , as well as O. owerinii , which, together with O. minor , belong to the species with which there is possible confusion, as they belong to the same subsect. Minores/ Speciosae .

O. quadrifida K. Koch (1849: 665) – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018); synonym of O. caryophyllacea , see also taxonomic note in O. caryophyllacea .

O. reticulata Wallroth (1825: 42) – ( Mtzchvetadze 1999, Gagnidze 2005, Nakhutsrishvili 2018); herbarium revision and field studies did not confirm this species.

O. teucrii Holandre (1829: 322) – cited with photos by Fischer et al. (2018), but photos illustrate O. alba with attached host – Thymus .

Figures

FIGURE 64. Orobanche colorata. A–C. General habit. D, E, G. Flower, front view. F. Flower, side view. H, J. Representative habitats (H. Samtskhe-Javakheti prov., Timotesubani. I. Kakheti prov., Kvemo Bodbe). I. Representative host, Salvia nemorosa. Photos by R. Piwowarczyk.

FIGURE 65. Distribution of Orobanche colorata in Georgia.