Diphelypaea tournefortii (Desf.) Nicolson (1975: 654)

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 : 18-22

2. Diphelypaea tournefortii (Desf.) Nicolson (1975: 654) . Type:— TURKEY (lectotype designated here):— Phelypaea Orientalis , flore coccineo [ Arménie, nowadays Erzurum Province, Turkey], Tournefort (1703: t. 479!). TURKEY (epitype designated here):— Armenia, im valle Kapuclu, inter Baiburt [Bayburt] et Erzeroum [Erzurum], Maio 1853, Huet de Pavillon [Plantae Orientales exsiccatae] (P-02970941!). Basionym:— Phelypaea tournefortii Desfontaines (1807: 298) . Homotypic synonyms:— Anoplon tournefortii (Desf.) G. Don (1837 –1838: 633). Anoplanthus tournefortii (Desf.) Reuter (1847: 42) . Heterotypic synonyms:— Anoplanthus coccineus var. peduncularis Boissier (1879: 494) . Fig. 9 .

Taxonomic Notes: —Here, we formally designated the Tournefort illustration of his Phelypaea Orientalis , flore coccineo (t. 479), which is cited in the protologue of Desfontaines (1807: 298), as the lectotype, as was suggested by Cullen (2010: 380). This is because no specimen of the original material could be found in the Tournefort herbarium (P). In addition, we chose an epitype because the lectotype lacks a complete stem and scales which show important diagnostic features for this species.

Distribution: —Mainly in the central and southern part of Armenia, in the Aragatsotn, Ararat, Gegharkunik, Kotayk, Yerevan and Vayots Dzor provinces. Usually, S of Pambaki mountain range, W and SW of Geghama mountain range, E slopes of Mt Aragats and in the Vayots Dzor mountain range, and rarely in the NW part of the Zangezur mountain range ( Fig. 10).

General distribution: —Caucasus ( Armenia, Azerbaijan, Georgia, N Iran) and Turkey.

Habitat: —Rocky slopes, hills, steppe, edges of forests, near paths around ruins and monasteries; usually (1200)1800–2100(2500) m.

Hosts: — Tanacetum sp. ( Asteraceae ), mainly T. polycephalum subsp. argyrophyllum (K. Koch) Podlech [syn. T. argyrophyllum (C. Koch) Tzvel. ], T. chilophyllum (Fisch. & Mey.) , T. canescens DC. [ Pyrethrum canescens (DC.) Boiss. ]. Need confirmation: Anthemis L., Pyrethrum myriophyllum (Willd.) C.A.M. [ Tanacetum polycephalum subsp. duderanum (Boiss.) Podlech ] ( Novopokrovskij & Tzvelev 1958).

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

Conservation status: —Least Concern (LC). The species has rather wide distribution in Armenia (6 floristic regions). One subpopulation grows in the Khosrov Forest State Reserve. Populations consist usually of ca. 10–50 shoots, rarely even a few hundred (e.g. Karmrashen, Fantan, Sers, Garni).

Notes: — Diphelypaea tournefortii differs from D. coccinea mainly by corolla with short cup-shaped tube, broad lobes which overlap, hairy calyx and stem, densely hairy anthers, leaves located on the base of stem, and host plant ( Figs. 7 , 9 ).

Specimens examined: — ARMENIA. Aragatsotn prov.: Ashtarak distr., Antarut, forest, with Tanacetum sp. , 3 June 1961, Sh. Aslanyan et al. (ERE) [as Phelipea baierni by I. Mulkidjanjan]; Antarut, forest, 2 June 1961, Sh. Aslanyan, R. Karapetyan & A. Pogosyan (ERCB); Byurakan, 28 May 1969, E. Matusova (ERE); Byurakan, 28 May 1969, K. Avakyan (ERCB); Ashtarak distr., between Koshabulah [Antarut] village and Aragats Mt., Amberd river gorge, [40°22’52”N, 44°14’26”E], 3 June 1961, Aslanyan, Karapetyan & Pogosyan (ERCB); Ararat prov.: Ashtarak distr., massif of Eranos Mt., S slope, with Tanacetum sp. , 27 June 1972, Mulkidjanjan et al. (ERE); mountis Eranos, in regione montana media, 18 May 1940, A. Fedorov (ERE); Azizkend x Daynag, left bank of the Vedi river, 1800 m, 1200–2000 m, with Tanacetum sp. , 27 May 1960, V. Manyakan & A. Bobrov (ERE, LE) [as Phaelipea bayernii by Mulkidjanjan]; Azizkend x Dagnaz, left bank of the Vedi river, 27 May 1960, A. Takhtajan, Ja. Mulkidjanian, E. Gabrielyan, A. Bobrov & V. Manakyan (ERCB); Ararat distr., Dagnak mountain range, Dagnak Mt., buffer zone of Khosrov reserve, mountain steppe, with host Anthemis tinctoria , 39°55’17,2”N, 44°54’14,47”E, 2068 m, 6 June 2012, E. Gabrielyan, B. Siebert & M. Aghababyan (ERE); NE macroslope Ketuz Mt, 16 km N of Dashtakar, 1500‒2100 m, 15 June 1977, G. Torosyan (ERE); Khosrov reserve, road to And, 3 June 1977, E. Gabrielyan & G. Torosyan (ERE); NNE of Zangakatun, mountain slope ca. 1.4 km E of Gortun, 39°51’44”N, 45°03’41”E, 2130 m, 14 June 2009, K. Kugler & E. Vitek (BP, ERE, W); 3 km S from Lusashoh village, mountain steppe, 39º49’72N, 44º59’195E, 1760 m, 28 May 2005, K. Tamanyan 10-2005 & G. Fayvush (P, NY); Ararat distr., 5‒6 km to North-East from ruins of the Gortun [ruins of the Hortun, 8 km to North-East from Zangakatun village] village, subalpine meadows, 26 June 1973, T. Popova & V. Manakyan (ERCB); Vedi distr., Khosrov reserve, near river Khosrov, [39°58’32”N, 45°00’15”E], 9 June 1972, O. Lovelius (ERCB, LE); Vedi rayon [Ararat rayon], 2 km to the south of the village of Kyarki [Tigranashen], 1400 m, 29 March 1973, N. Shvedchikova (MW) [on Tanacetum sp. ]; Vedi rayon, surroundings of the Council (Vedi), 30 May 1973, N. Shvedchikova (MW); ArmSSR, Vedi distr., between villages Kjarki [Tigranashen] and Sovetashen [Zangakatun], traganth steppe, [39°48’14”N, 44°58’37”E], 29 May 1960, A. Takhtajan, E. Gabrielyan & Ja. Mulkidjanyan (ERCB); Artashat distr., between Gelajsor and Gilanlar [Khosrov reserve], [40°06’01”N, 44°48’20”E], 26 May 1959, Takhtajan, Mulkidjanyan & Gabrielyan (ERCB); ArmSSR, Dorakh [in Khosrov reserve], on dry grassy slopes], 19 June 1981, M. Grigoryan (ERCB); Vedi distr., Khosrov forest, 29 May 1977, S. Grigoryan (ERCB); Khosrov forest, SE slope, 20 May 1960, Avakyan (ERCB); Armenian SSR, Ararat marz, Vedi distr., Urts range, Karmir sar Mt., vil. Asni, [39°51’26”N, 44°55’01”E], 9 June 1963, E. Gabrielyan (ERCB) [as P. bayernii by V. Manakyan]; Saray-Bulakh pass [Urts range, pass], 10 June 1930, A. Schelkovnikov (ERCB); Vedi distr., south slopes above the village Shugaib [now it is destroyed, even ruins don’t exist], 1600‒1700 m, phryganoid vegetation, [40°00’18”N, 44°46’29”E], Sh. Aslanyan (ERCB); Vedi distr., south slopes above the village Shugaib [now the village is destroyed], 1700‒1800 m, phryganoid vegetation, [40°00’18”N, 44°46’29”E], 22 May 1955, Sh. Aslanyan & R. Karapetyan (ERCB); Artashat distr., vicinity of Gelaysor village [now this village is destroyed and abandoned, it is in the territory of Khosrov Forest reserve, 40°02’04”N, 44°52’22”E], 26 May 1959, Takhtajan, Mulkidjanyan & Gabrielyan (ERCB); Vedi distr., Keshish-dag Mt., SW stony slope, steppe, 1850 m, 4 June 1965, M. Grigoryan (ERCB); Khosrov reserve, under the “jagdhaus”, [40°01’29”N, 44°54’35”E], 3 June 1977, L. Mnatsakanyan & M. Oganesyan (ERCB); Gegharkunik prov.: Martuni distr., on xerophilous slopes, 15 July 1939, S. Narinian (ERCB); “ Diphelypaea tournefortii , filmed in the Tarachichaki [Tsaghkunk/ Varazhnunik] (town) ...”, photo 85148 by V. Martirosyan, 2011, (in vahemart.livejournal.com; accessed Jan 2015); Kotayk prov.: Kotayk distr., near monastery Geghard, opposite bank of the river Azat, 22 May 1961, A. Takhtajan et al. (ERE) [as Phelipaea bayernii by Mulkidjanjan]; valley of river Goght, below of monastery Geghard, SW of Goght, 40°8’14”N, 44°48’22”E, 1640 m, 23 May 2006, K. Tamanyan & E. Vitek (ERE, W, G); between Kaghsi and Fantan, 6 June 1969, T. Tsaturyan (ERE); prope pagum Garni, 14 May 1939, A. Fedorov (ERE); near Garni, Milli dara, semi-desert, parasitise Achillea , 19 May 1957, E. Gabrielyan (ERE); Garni, rocky slopes above old bridge in direction to Khosrov reserve, 40°06’54,9”N, 44°44’34,3”E, 1370 m, 24 May 2016, R. Piwowarczyk (ERCB, KTC); slopes between Garni and Havuts Tar monastery, 40°07’13,3”N, 44°45’51,4”E, 1560 m, 25 May 2016, R. Piwowarczyk (KTC); SW of Goght, S and W of Havuts Tar monastery, 40°07’21”N, 44°46’08”E, ca. 1600 m, 25 May 2016, R. Piwowarczyk (KTC); Abovyan distr., near Geghard, on the rocks below monastery, right bank of the river Azat, 29 May 1967, G.Babayan (ERE); Erivan, Inter Arzni and Eilar,in „Pyrethretrum”, 31 May 1931, S. Yamaschian (ERE); Hrazdan distr., valley of river Hrazdan, up stream of village Bjni, mountain slope, 40°28’32”N, 44°40’37”E, 1932 m, 17 June 2004, E. Vitek, K. Tamanyan, G. Fayvush, M. Oganesian & A. Ter-Voskanian (ERE); Zar, 29 May 1951, B. Bramyan (ERE); NNE of Fantan, small hills, 40°24’24”N, 44°42’21”N, 1830 m, 6 June 2016, R. Piwowarczyk (KTC); Fantan, 15 June 1967, A. Melikyan (ERCB); NE of Aghveran, slope near the road, 40°30’54,7”N, 44°33’57,3”E, 1951 m, 6 June 2016, R. Piwowarczyk (KTC); Arzni, Zangu [Hrasdan] river gorge, slopes, [40°17’37”N, 44°35’23”E], 30 May 1933, A. Takhtajan (ERCB); in steppe inter Eilar [Abovian] et Arzni, [40°16’59”N, 44°33’54”E], 30 May 1931, S. Tamamschian & V. Maleev (ERCB); Tsagkhadzor, edges of forest, near monastery, 24 June 1968, T. Tsaturyan (ERCB); Vicinity of Gokcha [Sevan] Lake, Darachichag [Tsahkadzor], gravely slope, 2200 m [40°31’53”N, 44°42’44”E], 19 June 1929, P. Smirnov. N 7013 (MW) [as Anoplanthus coccineus Walp. ]; Tsagkhadzor, 20 May 1982, Agasaryan (ERCB); Hrazdan rayon, Tsaghkunyats Mountains, Marmarik, Tsaghkadzor, 5 June 1977, M.G. Pimenov, E.V. Klyuykov, L.P. Tomkovich & N.A. Cyrillin (MW) [on Tanacetum sp. ]; Tsagkhadzor, NE slope of the Teghenis range, 30 May 1975, S. Avakyan (ERCB); vicinity of Kaghsi village, [40°28’08”N, 44°44’09”E], 15 June 1959, T. Tsaturyan (ERCB); Kaghsi, 2 June 1971, I. Hrimilyan (ERCB); Kaghsi, gorge of the river Hrazdan, 2 June 1958, R. Gusyan (ERCB); Agveran, 28 June 1972, A. Melikyan (ERCB); Arzni, right slope of forest, 25 May 1952, T. Koharyan (ERCB); Arzni, dry, rocky slopes, 18 June 1952, T. Tsaturyan (ERCB); Arzni Plateau, 30 May 1933, N. Troitskiy (ERCB); Nairi distr., Eghvard vil., south slope of Arailer Mt., [40°23’04”N, 44°26’45”E], 28 May 1966, Ja. Mulkidjanjan (ERCB) [as P. bayernii ]; Arailer Mt., in the crater under steep walls among rocks, on the roots of Prangos ferulacea , [40°24’10”N, 44°27’25”E], 4 June 2010, E. Gabrielyan (ERCB); Vayots Dzor prov.: Azizbekov distr. [now it is Syunik marz, Sisian distr.], vicinity of Gushchi village [now Tasik vil.], [39°26’58”N, 45°58’00”E], 25 May 1945, Sh. Aslanyan & R. Karapetyan (ERCB); area ca. 5.8 km ENE of Yeghegnadzor, around monastery Thanahat, 39°46’48”N, 45°24’18”E, 1630 m, 16 May 2006, G. Fayvush, K. Tamanyan & E. Vitek (ERE, W); Daralagez [Vayots Dzor reg.], Ortakend vil. [Yeghegnadzor distr., Gladzor vil.], 8 April 1933, A. Movsesyan (ERE); Armenian SSR, Daralaghez, Mt. Teke-Dolduran, locality Khasma, mountain meadow [39°39’27”N, 45°22’30”E], 10 July, 1922, Ant . & I. Schchukiny (MW); Yeghegnadzor distr., between Khachik and Gnishik, grassy slopes, left of the road, 2400‒2500 m, 9 July 1976, B. Arevshatyan (ERE); Arazdayan [Yeraskh] – Mikoyan (Yeghegnadzor], 16–17 km right of road, 29 May 1960, A. Takhtajan, Ja. Mulkidjanjan & E. Gabrielyan (ERCB, ERE); Djermuk, Kechut village, near gorge with mineral fountain, slopes up to mountain, 39°49’21”N, 45°38’14”E, 2119 m, 19 June 2004, E. Vitek, K. Tamanyan, G. Fayvush, M. Oganesian & A. Ter-Voskanian (ERE, W); Yeghegnadzor distr., rocky slopes, left of the road between Areni and Amaghu, 31 May 1977, A. Takhtajan & E. Gabrielyan (ERE); NE Artabuynk, track to monastery Ts’agats’ Khor, below monastery, 39°53’25”N, 45°21’28”E, 2065 m, 31 May 2009, G. Fayvush & E. Vitek (ERE, W); ca. 5.7 km NNE of center of Yeghegnadzor, track from Vernashen to monastery Spitakavor Astvatsatsin, 1955 m, 39°48’31”N, 45°21’40”E, 1 June 2009, E. Vitek (ERE, W); area c. 14.5 km S of Yeghegnadzor, Gnishik, ca. 3.1 km SE of Gnishik, besides old road to Khachik, 2285 m, 39°38’07”N, 45°19’17”E, 16 June 2009, K. Kugler & E. Vitek (ERE, W); SW of Yelpin, slopes near road, 39°48’04,6”N, 45°05’56,6”E, 1576 m, R. Piwowarczyk (KTC); between Goghtanik and Karmrashen, W slope, 39°50’31,3”N, 45°29’08,3”E, 2100 m, 28 May 2016, R. Piwowarczyk (KTC); N of Khachik, W slope above road, 39°39’14,6”N, 45°12’23,6”E, 29 May 2016, R. Piwowarczyk (KTC); N of Sers, E slope, 39°34’22,1”N, 45°29’03,9”E, 2010–2060 m, 31 May 2016, R. Piwowarczyk (KTC); gorge SSW of Zaritap, 39°36’37,8”N, 45°29’16,4”E, 1730 m, 31 May 2016, R. Piwowarczyk (KTC); NNE of Gnishik, 39°40’25,5”N, 45°18’24,9”E, 2145 m, slope near the road, 31 May 2016, R. Piwowarczyk (KTC); Yeghegnadzor distr., between villages Gyulliduz [Vardahovit] and Karakaja [Getikvank], west stony slope, [39°55’14”N, 45°30’21”E], 15 June 1957, E. Gabrielyan, V. Avetisyan, Sh. Aslanyan & R. Karapetyan (ERCB) [as P. bayernii by Ja. Mulkidjanjan]; between villages Gyulliduz [Vardahovit] and Karakaja [Getikvank], west stony slope, 2100‒2200 m, [39°55’14”N, 45°30’21”E], 15 June 1957, V. Avetisyan, E. Gabrielyan, R. Karapetyan & Sh. Aslanyan (ERCB) [as P. bayernii by Ja. Mulkidjanjan]; Mikoyan distr. [Vayots Dzor marz, Yeghegnadzor distr.] v. Gjulliduz [Vardahovit, 39°53’41”N, 45°28’59”E], 15 June 1959, Ja. Mulkidjanjan (ERCB); Elegis river gorge, between Kavushugh and Gjulliduz [Hermon and Vardahovit], oak forest. 1700 m, [39°53’15”N, 45°28’12”E], 11 July 1968, Ja. Mulkidjanjan, K.K. Browicz, T.N. Popova & B. Arevshatyan (ERCB) [as P. bayernii by Ja. Mulkidjanjan]; between villages Kavushug [Hermon] and Gjulliduz [Vardahovit], [39°53’02”N, 45°28’05”E], 15 June 2003, K. Tamanyan & G. Fayvush (ERCB); Vayk distr., vicinity of Martiros village, 10 km to Khndzorut village, 1900 m, traganth communities, [39°34’59”N, 45°27’32”E], 8 June 1978, E. Gabrielyan (ERCB); Yeghegnadzor distr., just under Elpin’s electro substantion, on the right side of the road from Yerevan to Yeghegnadzor, [39°46’30”N, 45°05’12”E], 11 May 1995, K. Tamanyan (ERCB); Yerevan distr.: Yerevan, vicinity of Botanical Garden (on Pyrethrum ), 5 May 1937, A. Oganesyan (ERCB); Hortus botanicus Erivanensis [Yerevan, area of the Botanical Garden], 15 May 1940, R. Karapetyan (ERE), Sovietashen [Nubarashen], 14 July 1973, K. Tamanyan (ERE); Armeniae, Prope Erivan, Sh. Aslanyan (ERE, LE); Locus Armenia, Erivan, prope Kanaker [now it is a part of Yerevan, 40°13’50”N, 44°32’11”E], 13 May 1922, Tempus (ERCB); Hilar prope Erivan Bayerni Herb. Caucasi, Phelipaea tournefortii Dr. G. de Beck.

Identification key for Diphelypaea species

1. Stems usually covered with dense, long, multicellular white hairs. Scales 2(‒3) concentrated towards the base of the stems. Calyx clearly bilabiate (upper lip subentire with three short teeth and lower lip with two lanceolate lobes), with conspicuous white hairs (similar to stem). Corolla-lobes broadly obovate-orbicular to orbicular, overlapping. Anthers hairy. On Tanacetum ( Asteraceae )... ..................................................................................................................................................................................... D. tournefortii

+ Stems covered with slightly or ± dense, short, glandular brownish hairs. Scales 2‒5 evenly distributed along the stems. Calyx weakly bilabiate, 5-lobed (lobes deep, lanceolate and acute, more or less equal), with glandular brownish hairs (similar to stem). Corolla-lobes obovate to orbicular, not or clearly overlapping. Anthers hairy or glabrous. On Centaurea [s.l.] ( Asteraceae ) ........ 2

2. Stems up to 30 cm (rarely more). Corolla-tube conspicuous and with ± parallel-sides; corolla-lobes obovate to obovate-orbicular, not or scarcely overlapping. Anthers glabrous or slightly hairy. Usually on Psephellus [ Centaurea sect. Psephellus ] ( Asteraceae ). ......................................................................................................................................................................................... D. coccinea

+ Stems up to 15 cm (rarely more). Corolla-tube short and cup-shaped; corolla-lobes broadly obovate-orbicular to orbicular, overlapping. Anthers hairy. Usually on Centaurea [s.str.] ( Asteraceae ) ...................................................................................... D. boissieri

Genus Phelipanche Pomel (1874: 102)

Herbs annual, biennial, or perennial, holoparasitic, achlorophyllous. Stem simple or branched, with scattered scales. Flowers usually numerous, sessile or short-pedicellate, with one bract and two bracteoles on sides of calyx. Calyx usually gamosepalous, campanulate, with 4 subequal teeth, rarely with a fifth, very small, rudimentary. Corolla tubular, campanulate or infundibuliforum, bilabiate, upper lip 2-lobed, lower lip 3-lobed, usually blue, violet to rarely whitish or yellowish, with two, usually white, hairy folds. Stamens 4 inserted in the lower parts of corolla tube, anthers glabrous or hairy. Ovary 1-locular, ovate or cylindrical, placentae 4. Style elongated, usually not persistent, top opening, stigma 2-4-lobed, usually white or more rarely yellowish, and, sometimes, ± stained with violet. Pollen tricolpate with microreticulate sculpture of the exine. Capsule ovoid-globose, or ellipsoid, dehiscing by three valves. Seeds numerous, small, oblongoid, elipsoid, to globose, with reticulate testa with fibrillar ornamentation of the periclinal wall.

About 50–62 species (according to concept of species of the different authors) occur mostly in temperate regions of Europe, especially in the Mediterranean Basin, western and central Asia, north Africa, less represented in America and Oceania (alien). Parasitise roots of many different species.

Figures

FIGURE 7. Diphelypaea coccinea. A. General habit with host-Centaurea. B. Leaves on the upper part of stem. C. Flower with acute lobes, front view. D. Flower, side view, glabrous calyx and stem. E. Flower, rear view, glabrous calyx and stem. F. Flower, bottom view. Photos by Renata Piwowarczyk.

FIGURE 9. Diphelypaea tournefortii. A. General habit. B. Leaves at the base of the stem. C. Flower, front view, rounded lobes. D. Flower, side view, hairy calyx and stem. E. Flower, rear view, hairy calyx and stem. F. Flower, bottom view. Photos by Renata Piwowarczyk.