À.Ï. Ñóõîðóêîâ Карпология семейства Chenopodiaceae в связи с проблемами филогении, систематики и диагностики его представителей ББК 28.59 С 91 Сухоруков А.П. С 91 Карпология семейства Chenopodiaceae в связи с проблемами филогении, систематики и диагностики его представителей. — Тула: Гриф и К, 2014. — 400 с. Монография содержит сведения о строении покровов плода и семени у представителей одной из самых крупных клад в порядке Caryophyllales — Chenopodiaceae, охватывающей не менее 110 родов и 1600 видов. Впервые показано, что диверсификация признаков пе- рикарпия и спермодермы затронула все без исключения крупные группы (подсемейства Chenopodioideae, Salsoloideae, Betoideae, Corispermoideae, Suaedoideae, Salicornioideae), каждая из которых характеризуется определенными особенностями в строении плода и семени. Дан анализ карпологических признаков в других (внешних по отношению к Chenopodiaceae) группах Кариофиллид («core Caryophyllales»). Реконструкция эволюционной истории числа семян в плоде на основе молекулярно-филогенетического метода свидетельствует в поль- зу наличия односемянного типа плода у предков Кариофиллид и клады Chenopodiaceae. Прослежены эволюционные изменения состояний наиболее важных признаков перикар- пия и спермодермы во всем порядке. Для многих Кариофиллид свойственны лестнич- ные утолщения клеточных оболочек тегмена. Клада AAC (Achatocarpaceae, Amaranthaceae, Chenopodiaceae) имеет некоторые общие тенденции в карпологии. Среди всех представи- телей Caryophyllales ряд групп Chenopodiaceae характеризуется уникальной синапоморфи- ей — возникновением горизонтально ориентированного зародыша семени. Представлена современная таксономическая ревизия Chenopodiaceae в пределах Европейской России (38 родов и 133 вида), содержащая оригинальные диагностические ключи, важнейшую си- нонимику, подробное распространение видов по областям и республикам. Для многих так- сонов выбраны лектотипы. Книга рассчитана на специалистов широкого профиля. Издание осуществлено при финансовой поддержке Российского фонда фундаменталь- ных исследований по проекту № 14-04-07034-д. Издание РФФИ не подлежит продаже. ISBN 978-5-8125-2034-2 © А.П. Сухоруков, 2014 © Российский фонд фундаментальных исследований, 2014 The carpology of the Chenopodiaceae with reference to the phylogeny, systematics and diagnostics of its representatives Alexander P. Sukhorukov Th e book is the published dissertation which includes investigations from previously published articles with many additions and conclusions. Th e set of carpological characters is distinctive for ev- ery subfamily in the Chenopodiaceae, and these data are of particular importance for delimiting each of the subfamilies. Chenopodioideae. — Reproductive diaspores: (1) fruit enclosed in anthocarp (perianth), (2) fruit, and (3) seed arising from rupture of the pericarp. Ovary superior. Th e anthocarp not concres- cent with pericarp, 1- or several-layered, thin-walled. Th ere is a tendency to form heteromorphic seed types. Seed with vertical annular embryo. Seed-coat testa much thicker than tegmen, smooth or alveolate. Outer cell wall of the testa oft en impregnated with vertically or obliquely oriented stalac- tites. Perisperm present. Th e members of tribes Dysphanieae and Axyrideae do not deposite stalac- tites in the outer wall of the testa cells. In Axyris (tribe Axyrideae) the pericarp of one of two heteromorphic fruit types comprises two diff erent topographic zones. A many-layered pericarp (up to 12 layers) is rare and found in some Che- nopodium s.str., Oxybasis (O. macrosperma), the monotypic Holmbergia, Exomis and Manochlamys. Th e smooth amd fi nely mamillate pericarp surface is peculiar to the genera Blitum and Oxybasis. In mature fruits the papillate cells oft en fail to maintain turgidity thereby appearing crater-like, and thus the dry fruits diff er from soaked or non-abscissed ones in their surface. For this reason the crater-like or reticulate pericarp surface should not be considered as a relevant taxonomic trait. In many genera, especially Chenopodium s.str., Chenopodiastrum and part of Atriplex, the pericarp is indeed papillate. In contrast to Chenopodieae, the genus Dysphania (tribe Dysphanieae) possesses diff erent trichomes or short papillae, particularly in the majority of American species (glandular hairs) and Eurasian species (pericarp surface smooth or fi nely papillate). Th e fruit wall of Cycloloma bears two types of trichomes (simple and glandular hairs) not found else. In Atriplex, no keeled seeds are present. Th e seed heterogeneity in annual species can be visible (annual Atriplex) or cryptic (core Chenopodium, Chenopodiastrum, Oxybasis). Trispermy — the highest degree of seed heteromorphism, expressed as three seed types combining the cryptic and visible kinds of seed heterogeneity — has evolved twice independently in some Chenopodium s. str., and Atriplex. Th e hard seed coat is highly resistant to environmental degradation processes and, therefore, may be found fossilized. Betoideae. — Reproductive diaspores: (1) infructescence, (2) fruit, and (3) seed when the fruit dehisces by a lid. Ovary semi-inferior or superior. Th e anthocarp segments of neighbouring fl owers can be concrescent with each other. Pericarp free from anthocarp or fused with it, many-layered, comprising several thin-walled cells and many-layered sclerenchyma beneath. Seed with horizontal annular embryo. Seed-coat testa much thicker than tegmen. No fruit/seed heteromorphism is ob- served. Outer cell wall of the testa is oft en impregnated with tannins (stalactites) or lacking them. Perisperm present. Th e pericarp of Hablitzia and Acroglochin is homocellular or divided into parenchyma as out- er layer(s) and sclerenchyma beneath. Despite the geographical disjunction, some members like Aphanisma (North America) and Oreobliton (North Africa) as well as Hablitzia (Caucasus) and Acro- glochin (Himalaya) are carpologically closely related. 3 А.П. Сухоруков Corispermoideae. — Reproductive diaspores: fruit. Ovary superior. Fruit indehiscent or dehis- cent in special portion, fl attened. Pericarp not concrescent with hyaline perianth segments, oft en mamillate, papillate and/or with dendroid hairs, many-layered, comprising several thin-walled cells and many-layered sclerenchyma beneath. Th ere is a tendency for increasing numbers of sclerenchy- ma layers towards the fruit margins underneath the wing. Heterocarpy or heterospermy absent. Seed with vertical annular embryo. Seed coat smooth (with no papillae or mamillae), comprising two thin, tannin-fi lled (sub)equal layers, rarely the outer seedcoat layer thicker (Agriophyllum). Stalactites in the cell walls absent. Perisperm present. Th e pericarp anatomy is the most valuable character for the delimitation of Corispermum species when morphological traits do not allow precise identifi cation. Corispermum and Anthochlamys are carpologically closely related. Salsoloideae (incl. Camphorosmeae). — Reproductive diaspores: (1) fruit enclosed in antho- carp (perianth) and (2) fruit. Fruit indehiscent. Ovary superior. Pericarp not adherent to perianth, multi-layered (at least in upper third of the fruit), diff erentiated into topographical zones: (1) outer, usually one-layered epidermis, (2) parenchyma, (3) U-shaped cells with crystalliferous con- tent, and (4) inner epidermis. Seeds with horizontally, vertically or obliquely oriented, coiled or horseshoe-shaped embryo. Seed coat smooth (with no papillae or mamillae), comprising two thin, transparent, (sub)equal layers. Stalactites in the cell walls absent. Perisperm present (Salsoloideae s.str.) or absent (Camphorosmeae). Th e two- to four-cell-layered outer epidermis in the pericarp of three species of Anabasis (A. eriopoda, A. jaxartica, A. turkestanica) seems to be an apomor- phic feature in the Salsoloideae. Th e appropriate fruit term for the members with “fl eshy” pericarp (Anabasis and relatives) is ‘half-berry’ (semibacca). Th e fruit and seed covers have low resistance to environmental degradation processes and are unlikely to be found fossilized. Th e genera Sevada and Fadenia previously included in Suaedoideae are a part of Salsoloideae based on both molecular and carpological data. Suaedoideae. — Reproductive diaspores: (1) fruit enclosed in anthocarp (perianth), (2) fruit, and (3) seed arising from rupture of the pericarp. Ovary superior. Pericarp not concrescent with the anthocarp, 1-3-layered, thin-walled, without papillae or mamillae. Visible heterospermy is common in annual species. Seeds with vertical or horizontal embryo. Seed coat testa much thicker than teg- men, smooth. Th e outer cell wall of the testa in the dark seeds is impregnated with vertically (subgen. Suaeda) or obliquely oriented (subgen. Brezia excl. Suaeda australis) tannins (stalactites). Th is fea- ture is an additional character for delimiting both of these large groups. Perisperm absent or traces only (in dark seed type). Salicornioideae. — Reproductive diaspores: (1) fruit enclosed in anthocarp (perianth), (2) fruit, and (3) seed arising from rupture of the pericarp. Ovary superior. Pericarp not concrescent with the anthocarp, 1-3-layered, thin-walled. Structural heterocarpy or heterospermy not known, but termi- nal and lateral fl owers can be distinguished by morphometry. Seeds with horizontally or vertically oriented, annular or bent embryo. Seed-coat testa smooth, mamillate or papillate, much thicker than tegmen, or both testa and tegmen thin and equal in size. Outer cell wall of the testa (if thick) impreg- nated