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RESEARCH ARTICLE
Contents Vol 21(6)

Fruit anatomy of the genus Anabasis (Salsoloideae, Chenopodiaceae)

Alexander P. Sukhorukov
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Department of Higher Plants, Biological Faculty, Moscow Lomonosov State University, 119992, Vorobyovy Gory, Moscow, Russia. Email: suchor@mail.ru

Australian Systematic Botany 21(6) 431-442 https://doi.org/10.1071/SB08013
Submitted: 19 February 2008  Accepted: 30 October 2008   Published: 23 December 2008

Abstract

The fruit anatomy and morphology of 22 representatives of the genus Anabasis L. were studied, with the aim of clarifying the taxomomic importance of carpological characters in the genus. As shown in cross-sections, the pericarp of Anabasis is differentiated into the following four zones: (i) outer epidermis, (ii) subepidermal hydrated parenchyma, (iii) crystalliferous layer with lignified U-shaped cell walls and (iv) inner epidermis. Anatomical differences mainly relate to the outer epidermal structure. Fruit anatomy does not confirm the separation of the genera Brachylepis and Esfandiaria. A combination of carpological characteristics separates A. annua and A. setifera from the other species studied. Also, characters of reproductive organs in representatives of Anabasis are shown. When vegetative and reproductive features are considered, the genus Fredolia appears rather distant from Anabasis s.l. The pericarp histology of almost all the Salsoloideae (incl. Anabasis) is fully presented in the upper third of the fruit. In the lower parts of the fruit, some histological layers are either reduced or absent altogether. On the basis of the anatomical structures in the upper third of the fruit, the common carpological features of the Salsoloideae can be defined. These include a pericarp consisting of several, usually well-differentiated layers and the presence of crystalliferous cells with U-shaped walls. The two- to four-cell layered outer epidermis of three Anabasis representatives (A. eriopoda, A. jaxartica, A. turkestanica) seems to be an apomorphic feature in the Salsoloideae. The seed coat is thin (two cell layers thick) and non-differentiated. Owing to the pericarp and seedcoat structure, the fruit and seed covers have low resistance to environmental degradation processes and, therefore, are unlikely to be found among fossil remnants.


Acknowledgements

I thank Professor A. P. Melikyan, Professor A. Danin, Dr A. G. Devyatov, Dr A. I. Konstantinova, Dr M. V. Nilova, Dr E. S. Zaitseva, Dr E. Yu. Yembaturova, Dr L. Klimeš, T. Makhamov and anonymous referees for valuable comments and discussion. The work was supported by the RFFR (project 08-04-00393).


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Appendix 1.  Origin of the material of the Salsoloideae species investigated

Agathophora alopecuroides Bunge: Israel, 5 km S of Jericho, A. Sukhorukov I-39, Dec. 2007 (MW);

Anabasis africana Murb. ex H. Lindb.: Marocco, Dar-Drius, H. Mauricio s.n., Sept. 1930 (LE);

A. annua Bunge: Persia borealis, A. Bunge 68 (LE);

A. aphylla L.: Kazakhstan, ptov. Turgay, Kop-Mulla, N. Androssov, exs. 3086a, Sept. 1908 (MW);

A. articulata (Forsk.) Moq.: (1) Hispania, Baetica, Gros 18, Jul. 1925, (LE) sub A. hispanica Pau; (2) Hispania, Murcia, Lorca, H. Jeronimo 6447, Nov. 1927 (LE); sub A. hispanica; (3) [Egypt], C Sinai, Gebel El-Tih, N. Tadmor, S-729, Nov. 1969 (LE); (4) Hispania, prov. Murcia, Cartagena, S. Castroviejo & al. 9572, Nov. 1984 (MHA);

A. brachiata Fisch. & C.A.Mey. ex Kar. & Kir.: (1) Turkmenistan, prov. Tashauz, Ust-Urt, A.A. Mesheryakov s.n., Oct. 1954 (LE); (2) SW Kazakhstan, Karynzharyk, D.D. Vyshivkin s.n., Sept. 1956 (MW);

A. brevifolia C.A.Mey.: [Russia], Montain Altai region, Kosh-Agach, M. Danilov & O. Tur s.n., Aug. 1982 (MW);

A. calcarea (Charif & Aellen) Bokhari & Wendelbo: Iran, between Kerman and Bam, E.S. Brown 3428, Nov. 1960 (LE);

A. cretacea Pall.: (1) [Kazakhstan], Semipalatinsk, city surroundings, M. Iljin & A. Heinrichsson s.n., Sept. 1928, (LE); (2) Russia, prov. Saratov, Ozinki, A. Sukhorukov, s.n., Sept. 2006 (MW);

A. ebracteolata Korov. ex Botsch.: (1) Kazakhstan, Ust-Urt, Ash-Orpa, F.N. Rusanov s.n., Sept. 1926 (LE); (2) Kazakhstan, Ust-Urt, Beyneu, A. Sukhorukov s.n., Sept. 2001 (MW);

A. elatior (C.A. Mey.) Schischk.: East Kazakhstan, Aktogai, M. Lomonosova & A. Sukhorukov s.n., Sept. 2000 (MW);

A. eriopoda (Schrenk) G.Volkens: (1) [Uzbekistan], prov. Bukhara, Kuyu-Mazar, N. Androsov 1889b, Sept. 1905 (MW); (2) Kazakhstan, prov. Kzyl-Orda, N. Pavlov s.n., Sept. 1929 (MW);

A. ferganica Drobov: (1) Uzbekistan, Alai range, distr. Khalmion, Lopotin & Tinkhasov 467, Oct. 1940 (TASH); (2) Uzbekistan, Fergana valley, between Kadamzhay and Vuadil, U. Pratov 104, Nov. 1963 (LE);

A. haussknechtii Bunge ex Boiss.: (LE): N. Iran, prov. Damgan, M.P. Petrov s.n., Dec. 1942 (LE);

A. jaxartica (Bunge) Benth. ex Volkens: 1) Kazakhstan, Karatau, N. Pavlov 1302, Sept. 1931, (MW); 2) Uzbekistan, prov. Andizhan, T. Makhamov s.n., Nov. 2006 (MW);

A. oropediorum Maire: Algeria, Chellala, Ras Nokra, V.P. Bochantsev 236, Nov. 1967, (LE);

A. pelliotii Danguy: Kirghizia, Alai, Kyzyl-su, Koman, I. Tyshenko 1037, Sept. 1933 (LE);

A. prostrata Pomel: (1) Algeria, A.Pomel s.n., Dec. 1861 (P); (2) Dept. d’Oran, Arzew, R.Cesve 5293, Dec. 1926 (B);

A. salsa (C.A.Mey.) Benth. ex Volkens: (1) Kazakhstan, distr. Irghiz, Androsov & Bubyr 3087, Dec. 1910 (MW) sub A. ramosissima Minkw.; (2) Russia, prov. Volgograd, distr. Pallasovka, Elton, A. Seregin & A. Sukhorukov R-71, Sept. 2002 (MW);

A. setifera Moq.: (1) Israel, Southern shore of the Dead Sea, M. Zohary s.n., Dec. 1938 (MHA); (2) Iraq, between Shithatha and Hindiya, K.H. Rechinger 140, Nov. 1956 (LE);

A. tianschanica Botsch.: Kirghizia, C. Tien-Shan, Akche-tau, P. Gomolitsky 1094, Aug. 1932 (LE);

A. truncata Bunge: Kazakhstan, prov. Almaty, Kungey-Alatau, P. Polyakov 393, Oct. 1954 (LE);

A. turkestanica Korovin: Uzbekistan, 15 km NE from Dzhizak, P. Gomolitsky 433, Sept. 1934 (LE);

Arthrophytum lehmannianum Bunge: [Kazakhstan], distr. Karsakpay, Arys-Kul, N.V. Pavlov 2266, Sept. 1929 (MW);

A. subulifolium Schrenk: E Kazakhstan, Ili valley, between Chunzha and Tas-Karasu, L. Rodin 1655, Oct. 1931 (MW);

Fredolia aretioides Coss. & Durieu: (1) Algeria, Colomb-Bechar, ex herb. Bochantsev s.n. (LE); (2) Algeria, N Sahara, anonym s.n., Nov. 1965 (P);

Girgensohnia bungeana Sukhor.: Uzbekistan, prov. Samarkand, Karakchita, A. Sukhorukov 250, Oct. 2006 (MW);

Halothamnus glaucus (M.Bieb.) Botsch.: [Tadzhikistan], Zeravshan valley, P. Gordienko & L. Chilikina 531, Sept. 1930 (MW);

H. hispidus (Bunge) Botsch.: W Tien-Shan, Talass valley, N.V. Pavlov 102, Aug. 1966 (MW);

Haloxylon persicum Bunge ex Boiss. & Buhse: Uzbekistan, Kzyl-Dzhar, T.T. Trofimov s.n., Oct. 1954 (MW);

H. thomsonii Bunge ex Boiss.: NW India, Jammu & Kashmir, Ladakh, L.Klimesh 5064, Sept. 2004 (Pr);

Hammada eriantha Botsch.: Uzbekistan, between Shirabad and Zarabag, V.P. Bochantsev 18, Oct. 1970, (LE);

H. leptoclada (Popov) Iljin: Uzbekistan, prov. Surkhandarya, Kuhitang, Aktash, V.P. Bochantsev 53, Oct. 1970 (LE);

H. wakhanica (Pauls.) Iljin: Tadzhikistan, W Pamir, Abdusalyamova 4902, Aug. 1963 (LE);

Iljinia regelii (Bunge) Korovin: [Kazakhstan], distr. Lepsy, Boin-Nor, S. Lipshitz 1260, Sept. 1928 (MW);

Lagenantha gillettii (Botsch.) M.G. Gilbert & Friis: Kenia, Thompson s.n., 1964 (B);

Nanophyton erinaceum (Pall.) Bunge: Kazakhstan, prov. Aktyubinsk, Ust-Urt, Kurusay, Vostokova s.n., Sept. 1948 (MW);

Noaea mucronata (Forsk.) Asch. & Schweinf.: Cypern, Larnaca Bay, A. Sukhorukov s.n., Oct. 2006 (MW);

Petrosimonia brachiata (Pall.) Moq.: Russia, prov. Astrakhan, A. Sukhorukov s.n., Sept. 2002 (MW);

Salsola inermis Forsk.: Cypern, Larnaca Bay, A. Sukhorukov s.n., Oct. 2006, (MW; B);

S. incanescens C.A.Mey.: Uzbekistan, prov. Syr-Darya, road to Tashkent, A. Sukhorukov s.n., Oct. 2006 (MW);

S. komarovii Iljin: Russia, Vladivostok, Russkiy island, S. Petrova s.n., Sept. 2007 (MW);

S. tamariscina Pall.: Russia, prov. Saratov, Ozinki, A. Sukhorukov s.n., Sept. 2006 (MW);

S. verticillata Schousb.: Spain, Costa del Sol, T. Konovalova & N. Shevyreva s.n., Oct. 1996 (MHA);

Seidlitzia rosmarinus Bunge: [Turkmenistan], SE Karakum deserts, E.V. Korovin s.n., Oct. 1926 (MW)