Oenanthe millefolia Janka Oesterr. Bot. Z. 22: 177-178, 1872
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|Oenanthe millefolia Janka Oesterr. Bot. Z. 22: 177-178, 1872|
= Oenanthe bulgarica Velen. Fl. Bulg. Suppl. 1: 127, 1898.
BULGARIA. in pratis inter Kalofer et Karlova ad ped. austral. m. Balkan Thraciae, ubi specimina nondum bene efflorata legi d. 2. Junii 1871 (holotype GOET!); In graminosis pagum Susam. July 1910, V. Stribrny s.n. (W!) ( Oenanthe bulgarica ); Kreis Sliven, Stara Planina (Balkan-Gebirge), Kotlenska Planina (Kotel-10 Str. -km S Kotel Richtung Gradec, Balkan), Rastplatz mit Brunnen, Quercus - Mischwald ca. 450 m s.m., 09 August 1978 Kalk., F. Ehrendorfer, F. Sorger, D. Fürnkranz, M.A. Fischer, A. Öztürk s.n. (WU!); Stara Zagora district, around Quercus robur forest, in Kilimite area, 26.06.1958, Ivan Ganchev (photo SOM!); South Black Sea coastline, oak forest between Ahtopol and Sinemorets, 09.05.2004, A. Petrova & B. Assyov (photo SOM!); Strandzha mountain , oak forest, by the roadside, Tsarevo-Malko Tarnovo, at fountain, west of Izgrev village , 09.05.2004, A. Petrova & B. Assyov (photo SOM!); Thracian Lowland, pasture, northeast of Malevo village , Haskovo district, 11.06.2004, A. Petrova (photo SOM!); Eastern Rhodopi , above Potocharka village , in forest of Carpinus orientalis , Fraxinus ornus and Paliurus spina-christi , 08.06.2006, D. Dimitrov (photo SOM!); Stara Zagora district, near Sarnevo village , in oak forest, 08.06.1960, Iv. Ganchev, St. Denchev (photo SOM!); St. Iliya hills, 22.07.1964, Iv. Ganchev, St. Denchev (photo SOM!) .
TURKEY. Kırklareli: Demirköy-Iğneada, to 5-10 km Iğneada, under Pinus sylvestris forest, 273 m, 18 July 2014, E. Doğan Güner 2044 & B. Bani (GAZI!); ibid., 03 August 2014, E. Doğan Güner 2075 & B. Bani (GAZI!) ; Kırklareli: Sarpdere-Armutveren , under Pinus sylvestris forest, 33 m, 19 June 2014, E. Doğan Güner 2046 & B. Bani (GAZI!) ; Tekirdağ: Saray, Kiyikoey district , under Quercus forest, 247 m, 16 June 2015, E. Doğan Güner 2101 & B. Bani (GAZI!) .
Perennial, 40-70 cm tall, herb, with thickened, fusiform or oblong tubers, tubers generally at stem base, rarely far away. Stem erect, simple or 3 times branched above, hollow, furrowed, minutely scabrid below, glabrous above. Basal leaves lanceolate or oblong in outline, 2-3 pinnate, 17-45 × 5-9 cm, leaves lamina longer than petiole; segments of lamina opposite at rachis, deeply pinnatisect, ultimate segments linear or elliptic up to 6 × 1 mm, excurrent into a setaceaus tip. Upper leaves similar to basal one, but only a few which reduce upwards. Umbel with 12-18 slender rays of sub-equal length, up to 2 cm, becoming slightly thickened in fruit. Umbels 5 cm diam. at flowers and 3 cm diam. at fruit. Bracts 8-9, lanceolate, 8-10 × 1.5-2 mm. Umbellules conical with unequal thickened pedicels in fruit, 20-30 flowered, about 1 -1.5 mm diam., pedicels of sterile flowers longer than fertile ones. Bracteoles 9-13, elliptic-linear, 2-3.5 × 0.5-1.5 mm. Petals radiating, white, cordate, to 2.5 mm long. Sepals ovate, 0.3-0.4 mm, acuminate at apex. Filaments at least two times longer than petals. Stylopodium is conical and not exceeding calyx teeth. Styles about as long as the body of the fruit (ca. 3 mm), erect. Fruit ovate to cylindrical, 3 × 2 mm, striate, laterally and base of fruit slightly spongiose margin.
Distribution, habitat and ecology.
Oenanthe millefolia is distributed in Bulgaria, North-eastern Greece and European Turkey (Fig. 1 View Figure 1 ). No threat factor was observed against the habitat of the species. The populations are represented by many healthy individuals. The flowering time is between June and July, the fruiting time is August. It grows on clearings of Pinus and Quercus forest between the altitudes of 240-450 m and shares the same habitat with the species of Oenanthe pimpinelloides L., Helianthemum racemosum (L.) Pau, Trachystemon orientalis (L.) G. Don., Crupina vulgaris Cass, Pinus nigra Arn. subsp. pallasiana (Lamb.) Holmboe, Pulicaria dysenterica (L.) Bernh., and Anthemis altissima L.
Mericarp macromorphology and micromorphology.
Mericarps have three dorsal and two lateral primary ribs. The lateral ridges are more prominent and broader than the dorsal ones. The lateral ridges extend towards the base and cover the base of mericarp. Sepals are generally distinctive and persistent in Oenanthe species. Stylopodium is conical and not exceeding calyx teeth. Stylopodium ending with style is almost as long as the fruit. The surface ornamentation of the pericarp is longitudinally striate. The pattern is formed by rectangular cells. Stomatal cavities are observed on the pericarp surface and the density increases towards the calyx (Fig. 3 View Figure 3 ). The style surface is ribbed.
Stem anatomy: Stems are circular and slightly 7-8 ribbed in cross sections. There is a thin layer of cuticle on the top surface and a single-line epidermis composed of rectangular cells underneath. Collenchyma cells are grouped at the edges. Cortex parenchyma cells are located around disordered collenchyma cells. Collenchyma and parenchyma cells are identified as two layers between edges. Collenchyma has 4-5 rowed, small, and circular cells. Parenchyma has 1-2 rowed, large and circular cells. Secondary slight edges are located between the edges with collenchyma cells. Secretion canals exist in cortex under the collenchyma layer. 5-6 cells in one line surround the canals. Endoderm cell walls are slightly thick one-line oval cells. Vascular bundles are embedded between the cortex and the pith. There are 6-7 rows of sclerenchymatic cell layers between the vascular bundles. Peripheral vascular bundles which are collateral, are large against the edges and small in between the edges. Central vascular bundles are connected with peripherals by sclerenchymatic tissue. Secretion canals are also found under vascular bundles. Slight thickening is seen in pith parenchyma cells. Pith cells have various sizes with large inter-cellular spaces (Fig. 4A-B View Figure 4 ).
Petiole anatomy: Petiole is almost straight in cross section, ovoid in outline and slightly-canaliculate on the lower surface. Scabrid hairs rarely occur between cubic epidermis cells. Cuticle, epidermis and collenchyma structures are designed in almost the same manner as the stem. Secretion canals between collenchyma and concentric vascular bundles are significant. Xylem elements are dominantly distributed. The pith is composed of large circular cells with a hollow centre (Fig. 4C-D View Figure 4 ).
Leaf anatomy: The epidermal layer consists of rectangular or circular cells in both adaxial and abaxial directions. Stoma exist on both surfaces. There are large respiratory spaces under the stomata. Mesophyll is composed of two-row palisade and one or two-row sponge parenchyma cells. Large ventilation spaces exist between sponge parenchyma cells. Xylem elements are located through the abaxial side and phloem elements are located through the adaxial side (Fig. 4E-F View Figure 4 ).
Fruit anatomy: The fruit is schizocarp with two mericarps. The pericarp forms a thin layer around the endocarp and seed. There are single-line and horizontally located epidermal cells on the surface. The mesocarp is formed by 2-3-row small cells. Both epidermis and mesocarp cells have significant thickness. Five ridges are seen on each mericarp. Vascular bundles are located on these ridges. They are reduced. Secretion canals are located on vascular bundles. Pericarp is surrounded with sclerenchymatic tissue which makes a continuous ring up to the carpophore. The sclerenchyma layer is composed of irregular cells with thick walls. There are 4 dorsally and 2 ventrally vittae. Oval-shaped vittae are located in the vallecular region. Endoderm is located as one line under the vittae and seems to be integrated with the testa. The seed is composed of endosperm and testa with a thickened cell wall. Endosperm contains large quantities of lipid and protein. There are many druse crystals in the endosperm. When the section is taken from the middle of the mericarp, the embryo cannot be observed because it is small and close to the tip (Fig. 4G-H View Figure 4 ).
The pollen grains are isopolar symmetric, the aperture is tricolporate type. The pollen shape is prolate with an elliptic equatorial outline, polar axis 29.5-33.5 µm, equatorial axis 15-18 µm. The ornamentation is rugulate. The colpus length is 18-27 µm and width is 0.5-2 µm. The pore length is 4-6 µm and width is 4-6 µm. The exine subtectate is 0.75-1 µm (on equator and polar), the intin is 0.75-1.25 µm (on equator and polar) (Fig. 5 View Figure 5 ).
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