Origin,Differentiation, and Geographic Distribution of the Chenopodiaceae

Numbers of species and genera,endemic genera,extant primitive genera,relationship and distribution patterns of presently living Chenopodiaceae(two subfamilies,12 tribes,and 118 genera)are analyzed and compared for eight distributional areas,namely central Asia,Europe,the Mediterranean region,Africa,North America,South America, Australia and East Asia． The Central Asia,where the number of genera and diversity of taxa are greater than in other areas,appears to be the center of distribution of extant Chenopodiaceae．North America and Australia are two secondary centers of distribution． Eurasia has 11 tribes out of the 12,a total of 70 genera of extant chenopodiaceous plants,and it contains the most primitive genera of every tribe． Archiatriplex of Atripliceae,Hablitzia of Hablitzeae,Corispermum of Corispermeae,Camphorosma of Camphorosmaea,Kalidium of Salicornieae,Polecnemum of Polycnemeae,Alexandra of Suaedeae,and Nanophyton of Salsoleae,are all found in Eurasia,The Beteae is an Eurasian endemic tribe,demonstrating the antiquity of the Chenopodiaceae flora of Eurasia．Hence,Eurasia is likely the place of origin of chenopodiaceous plants． The presence of chenopodiaceous plants is correlated with an arid climate．During the Cretaceous Period,most places of the continent of Eurasia were occupied by the ancient precursor to the Mediterranean,the Tethys Sea．At that time the area of the Tethys Sea had a dry and warm climate．Therefore,primitive Chenopodiaceae were likely present on the beaches of this ancient land．This arid climatic condition resulted in differentiation of the tribes Chenopodieae,Atripliceae,Comphorosmeae,Salicornieae,etc．,the main primitive tribes of the subfamily Cyclolobeae. Then following continental drift and the Laurasian and Gondwanan disintegration, the Chenopodiaceae were brought to every continent to propagate and develop, and experience the vicissitudes of climates, forming the main characteristics and distribution patterns of recent continental floras. The tribes Atripliceae, Chenopodieae, Camphorosmeae, and Salicornieae of recent Chenopodiaceae in Eurasia, North America, South America, southern Africa, and Australia all became strongly differentiated. However, Australia and South America, have no genera of Spirolobeae except for a few maritime Suaeda species. The Salsoleae and Suaedeae have not arrived in Australia and South America, which indicates that the subfamily Spirolobeae developed in Eurasia after Australia separated from the ancient South America-Africa continent, and South America had left Africa. The endemic tribe of North America, the tribe Sarcobateae, has a origin different from the tribes Salsoleae and Suaedeae of the subfamily Spirolobeae. Sarcobateae flowers diverged into unisexuality and absence of bractlets. Clearly they originated in North America after North America had left the Eurasian continent. North America and southern Africa have a few species of Salsola, but none of them have become very much differentiated or developed, so they must have arrived through overland migration across ancient continental connections. India has no southern African Chenopodiaceae floristic components except for a few maritime taxa, which shows that when the Indian subcontinent left Africa in the Triassic period, the Chenopodiaceae had not yet developed in Africa. Therefore, the early Cretaceous Period about 120 million years ago, when the ancient Gondwanan and Laurasian continents disintegrated, could have been the time of origin of Chenopodiaceae plants.The Chinese flora of Chenopodiaceae is a part of Chenopodiaceae flora of central Asia. Cornulaca alaschnica was discovered from Gansu, China, showing that the Chinese Chenopodiaceae flora certainly has contact with the Mediterranean Chenopodiaceae flora. The contact of southeastern China with the Australia Chenopodiaceae flora, however, is very weak.     

藜科植物的起源、分化和地理分布

全球藜科植物共约130属1500余种,广泛分布于欧亚大陆、南北美洲、非洲和大洋洲的半干旱及盐碱地区。它基本上是一个温带科,对亚热带和寒温带也有一定的适应性。本文分析了该科包含的1l族的系统位置和分布式样,以及各个属的分布区,提出中亚区是现存藜科植物的分布中心,原始的藜科植物在古地中海的东岸即华夏陆台(或中国的西南部)发生,然后向干旱的古地中海沿岸迁移、分化,产生了环胚亚科主要族的原始类群;起源的时间可能在白垩纪初,冈瓦纳古陆和劳亚古陆进一步解体的时期。文章对其迁移途径及现代分布式样形成的原因进行了讨论。     

国产藜科14种植物种皮微形态特征比较研究

应用扫描电子显微镜和光学显微镜观察了中国产藜科(Chenopodiaceae)5族10属14种植物的种子形态和种皮微形态结构。观察结果表明:供试的14种藜科植物的种子多为卵形和圆形,少数为双凸镜形;种皮表面都具有网状纹饰,包括浅网纹、负网纹和穴状网纹。种皮可分为膜质和壳质2类,其中,膜质型种皮表面不光滑,除网状纹饰外,还有瘤状或褶皱状等纹饰,网眼无规则,纹饰排列无序,猪毛菜族(Salsoleae C.A.Mey.)的松叶猪毛菜(Salsola laricifolia Turcz.ex Litv.)及樟味藜族(Camphorosmeae Moq.)的地肤〔Kochia scoparia(L.)Schrad.〕、全翅地肤(K.krylovii Litv.)、黑翅地肤(K.melanoptera Bunge)、雾冰藜〔Bassia dasyphylla(Fisch.et Mey.)O.Kuntze〕、钩刺雾冰藜〔B.hyssopifolia(Pall.)O.Kuntze〕、樟味藜(Camphorosma monspeliaca L.)、兜藜(Panderia turkestanica Iljin.)和棉藜(Kirilowia eriantha Bunge)均属此类;壳质型种皮表面近光滑,网眼规则,排列整齐有序,少数为不规则浅穴,网脊平而不明显或凹陷,滨藜族(Atripliceae C.A.Mey.)的滨藜〔Atriplex patens(Litv.)Iljin.〕、碱蓬族(Suaedeae Reich.)的角果碱蓬〔Suaeda corniculata(C.A.Mey.)Bunge〕、纵翅碱蓬〔S.pterantha(Kar.et Kir.)Bunge〕和异子蓬(Borszczowia aralocaspica Bunge)以及藜族(Chenopodieae C.A.Mey.)的刺藜(Chenopodium aristatum L.)均属此类。各族之间种皮微形态结构也有一定的区别。根据种皮微形态,地肤属(Kochia Roth)与雾冰藜属(Bassia All.)的关系较近,属的界限和部分种类的亲缘关系需要进一步界定。藜科植物种皮表面大部分凹凸不平,这种结构可能与荒漠干旱生境下吸收和保留水分有关。     

Fruit and Seed Anatomy of Chenopodium and Related Genera (Chenopodioideae,Chenopodiaceae/Amaranthaceae): Implications for Evolution and Taxonomy

A comparative carpological study of 96 species of all clades formerly considered as the tribe Chenopodieae has been conducted for the first time. The results show important differences in the anatomical structure of the pericarp and seed coat between representatives of terminal clades including Chenopodium s.str.+Chenopodiastrum and the recently recognized genera Blitum, Oxybasis and Dysphania. Within Chenopodium the most significant changes in fruit and seed structure are found in members of C. sect. Skottsbergia. The genera Rhagodia and Einadia differ insignificantly from Chenopodium. The evolution of heterospermy in Chenopodium is discussed. Almost all representatives of the tribe Dysphanieae are clearly separated from other Chenopodioideae on the basis of a diverse set of characteristics, including the small dimensions of the fruits (especially in Australian taxa), their subglobose shape (excl. Teloxys and Suckleya), and peculiarities of the pericarp indumentum. The set of fruit and seed characters evolved within the subfamily Chenopodioideae is described. A recent phylogenetic hypothesis is employed to examine the evolution of three (out of a total of 21) characters, namely seed color, testa-cell protoplast characteristics and embryo orientation.     

Wood anatomy of the subfamily Crotonoideae (Euphorbiaceae s.s.) from India: systematic implications with special reference to the taxonomic delimitation of Givotia and Vernicia

The wood anatomy of 15 representative species belonging to 12 genera of nine tribes of the subfamily Crotonoideae (Euphorbiaceae) are comprehensively described with focus on systematic implications. In addition, ecological and evolutionary aspects are evaluated. An identification key to the species based on wood anatomical features is presented. The wood microstructure of the tribes was found to be considerably heterogeneous reflecting an unnatural classification of the subfamily. However, the results confirm the generic relationship within subtribe Aleuritinae and tribe Ricinodendreae. Vernicia and Givotia may be recognized based on wood anatomical and morphological characters. The tribes Micrandreae and Adenoclineae have considerable similarity in wood anatomy. The wood structure of the monogeneric tribes Trigonostemoneae and Geloneae idicate a close relationship with the tribe Crotoneae.     

Radiation of the Australian Salicornioideae (Chenopodiaceae)--based on evidence from nuclear and chloroplast DNA sequences

In phylogenetic analyses of nuclear ITS and chloroplast trnL DNA sequences, the mostly endemic Australian genera; Halosarcia, Pachycornia, Sclerostegia, Tecticornia, and Tegicornia of the subfamily Salicornioideae (Chenopodiaceae) together form a monophyletic group, congruent with the hypothesis that they evolved from a common ancestor. However, limited genetic differentiation evident in both nrDNA and cpDNA sequences among these taxa suggests a possible rapid radiation. Based on fossil pollen records and climatic models of other authors, it is hypothesized that the expansion of the Australian endemic Salicornioideae most likely occurred during the Late Miocene to Pliocene, when increasing aridity caused the formation of extensive salt lakes along endorheic paleodrainage channels. Moreover, Australian Sarcocornia representatives were supported as monophyletic, nested within a paraphyletic Sarcocornia clade that also comprised European Salicornia in the ITS analysis. This suggests that Sarcocornia arrived in Australia subsequent to the ancestor of the Australian endemic genera most likely via long-distance dispersal.     

Molecular systematics and morphological character evolution of the Condamineeae (Rubiaceae)

? Premise of the study: The Condamineeae have in previous molecular studies been shown to be part of an early-divergent clade within the subfamily Ixoroideae, together with the tribes Calycophylleae, and Hippotideae, and genera of the former Cinchoneae and Rondeletieae. Generic relationships within this clade have, however, remained largely unresolved. ? Methods: In this study, the systematics of the Condamineeae was further examined by phylogenetic reconstruction of six cpDNA regions and one nrDNA region using parsimony and Bayesian Markov chain Monte Carlo inference. Morphological character evolution within the tribe was assessed by ancestral state reconstruction using likelihood optimization of characters onto Bayesian trees. ? Key results: Calycophylleae appears polyphyletic. "Hippotideae" is monophyletic but nested within the Condamineeae. The phylogenetic hypotheses presented support a resurrection of the genera Holtonia, Schizocalyx, and Semaphyllanthe. Furthermore, Bathysa is found to be polyphyletic, Tresanthera is found nested within Rustia, and the taxonomically disputed genus Dialypetalanthus is here shown to be sister to a Bothriospora-Wittmackanthus clade. Morphological ancestral state reconstructions indicate that protogyny have evolved at least two times within the tribe and that indehiscent fruits, loculicidal fruit dehiscence, and intrapetiolar stipules have evolved independently several times. The occurrence of calycophylls (leaf-like calyx lobes), poricidal anthers, and winged seeds also appear homoplastic within the tribe. ? Conclusions: A diagnosis and delimitation of the tribe Condamineeae is presented, with taxonomic proposals to synonymize Tresanthera and to transfer several species of Bathysa as well as Phitopis to a resurrected Schizocalyx.     

中国水鳖科植物种皮微形态特征及其系统学意义

通过扫描电镜对国产水鳖科植物(包括6属13种)的种皮微形态特征进行观察,并作了系统描述。根据种皮细胞形态、外种皮表面纹饰和内种皮内层小瘤状突起的特点将水鳖科植物的种皮微形态特征划分为3种类型,即海菜花型(海菜花属)、水鳖型(水鳖属)和苦草型(苦草属、水筛属、虾子草属和黑藻属),并作出了分属检索表。本文结果表明,种皮微形态特征可作为该科族、属以及属内种级水平分类的依据,对探讨属间关系和该科的系统发育关系亦具有重要的价值。种皮微形态特征支持Hutchinson(1959)和Eckhardt(1964)将海菜花属和水鳖属分别作为一个独立的族处理的观点。苦草属、水筛属和虾子草属种皮微形态特征的高度相似性表明它们间有密切的联系,不支持将它们置于不同亚科和族的分类处理。黑藻属虽与上述3属近缘,但其外种皮特征则较为独特,因此与水筛属放在不同族中更为合理。本文种皮微形态特征的研究结果支持iki1937)和Shaffer-Fehre(1991b)等关于水鳖科与茨藻科近缘的观点。     

Ovules and seeds in Acalyphoideae (Euphorbiaceae): structure and systematic implications

Acalyphoideae, the largest subfamily of Euphorbiaceae, are investigated with respect to ovule and seed structure on the basis of 172 species of 80 genera in all 20 tribes of Acalyphoideae sensu Webster. All species of Acalyphoideae examined have bitegmic ovules with a non-vascularized inner integument. However, noticeable differences exist among and sometimes within the genera in the thickness of the inner and outer integument, the presence or absence of vascular bundles in the outer integument, whether ovules are pachychalazal or not, the presence or absence of an aril, seed coat structure (in terms of the best-developed mechanical cell-layer), and the shape of cells constituting the exotegmen. For the latter two characters, two different types of seed coat (i.e., "exotegmic" and "exotestal") and three different types of exotegmic cell (i.e., palisadal, tracheoidal and ribbon-like) were distinguished. Comparisons showed that three tribes Clutieae, Chaetocarpeae and Pereae are distinct from the other Acalyphoideae as well as from the other Euphorbiaceae in having an exotestal seed coat with a tracheoidal exotegmen. The tribe Dicoelieae is also distinct from the other Acalyphoideae in having an exotegmic seed that is composed of ribbon-like cells of exotegmen (i.e., cells both longitudinally and radially elongated, sclerotic and pitted). The tribe Galearieae, which should be treated as a distinct family Pandaceae, is also distinct from the other Acalyphoideae in having an exotegmic seed with a tracheoidal exotegmen (i.e., cells longitudinally elongated, sclerotic and pitted). The remaining genera of Acalyphoideae always have an exotegmic seed with a palisadal exotegmen (i.e., cells radially elongated, sclerotic and pitted). The shared palisadal exotegmen supports the close affinity of Acalyphoideae (excluding five tribes) with Crotonoideae and Euphorbioideae. Within the remaining genera of Acalyphoideae, a significant diversity is found in ovule and seed morphology with respect to the thickness of the inner and outer integument, the size of chalaza, vascularization of an outer integument and an aril.     

Molecular phylogeny of Palaearctic genera of Gomphocerinae grasshoppers (Orthoptera, Acrididae)

Abstract.  Molecular phylogenetic methods were used to examine morphologically based hypotheses concerning the taxonomic structure and relationships of the grasshopper subfamily Gomphocerinae. Two mitochondrial gene (cytochrome b and cytochrome oxidase subunit I) sequences were determined for twenty-five species representing eleven Palaearctic genera. The studied Gomphocerinae species constituted a monophyletic group; furthermore, the earlier division of Gomphocerinae into tribes was supported, with each tribe monophyletic. There was no support for various systems uniting Stenobothrini and Gomphocerini into one tribe. Two separate clusters were discerned in Gomphocerini and two tribes were distinguished – Gomphocerini (genera Aeropus , Stauroderus , Chorthippus ) and Stenobothrini (genera Omocestus , Stenobothrus ).     

A classification of the Ericaceae: subfamilies and tribes

The variation shown by 60 characters (morphological, anatomical, embryological, chemical and cytological) is tabulated for the whole of the Ericaceae. The reliability of the characters and the observations on them are discussed, and it is emphasized that many characters are difficult to use because they are rarely restricted to one taxon. Six subfamilies are recognized. Within the Rhododendroideae seven tribes are recognized, the monogeneric Daboecieae being new; Epigaea is transferred to the Rhododendroideae from the Andromedeae and Diplarche from the Diapensiaceae. Calluna is placed in a new monotypic tribe within the Ericoideae. The extended Vaccinioideae includes the Arbutoideae; five tribes are recognized, of which the Enkiantheae and Cassiopeae are new. The Pyroloideae and Monotropoideae are left unchanged: a new subfamily, the Wittsteinioideae, includes the deviant monotypic genus Wittsteinia. A key to subfamilies, and to genera of the Pyroloideae, Rhododendroideae and superior-ovaried Vaccinioideae is given.     

Trifolieae and related seeds from archaeological contexts: problems in identification

Small legume seeds are commonly recovered from Epipalaeolithic and Early Neolithic sites in north Africa and south-west Asia, often as a high proportion of the total plant remains. Their role is uncertain, but it is thought that small-seeded legumes may once have been a human food resource. They are difficult to identify. The gross morphology and testa surface micromorphology have been investigated in fresh seeds of members of the tribe Trifolieae and some related tribes in the same subfamily Faboideae. The separation of the four genera Medicago, Melilotus, Trifolium, and Trigonella has been attempted by multivariate analysis of seed characters that can be seen in experimentally charred seed specimens. The characters used have also been observed in well-preserved ancient charred small legume seeds, and so can be readily used for the purpose of identification. Although identification to the level of species is rarely possible, identification to generic level could provide information useful for the interpretation of this category of ancient plant remains.     

The position of Najas within the subclass Alismatidae (Monocotyledones) in the light of new evidence from seed coat structures in the Hydrocharitoideae (Hydrocharitales)

SHAFFER-FEHRE, M., 1991. The position of Najas within the Alismatidae (Monocotyledones) in the light of new evidence from seed coat structures in the Hydrocharitoideae (Hydrocharitaceae). This paper gives details of seed coat structure in several genera of the Hydrocharitoideae (Hydrocharitaceae) and of Najas (Najadaceae). The taxonomic position of Najas L., as a genus or as the basis of a higher taxonomic category, has changed with the publication of every new system of classification in which it was considered. In the light of evidence derived from seed coat anatomy, particularly that of the testa, and from the presence of endotegmen tuberculae, Najas is placed in the enlarged tribe Stratioteae in the subfamily Hydrocharitoideae (Hydrocharitaceae).     

基于28S rDNA D2基因片段与形态特征的矛茧蜂亚科系统发育研究（膜翅目：茧蜂科）

本研究选取矛茧蜂亚科Doryctinae（昆虫纲Insecta：膜翅目Hymenoptera：茧蜂科Braconidae）的6族15属18种做内群,茧蜂科其它7亚科11属11种做外群,首次结合同源核糖体28S rDNA D2基因序列片段和100个形态学和解剖学特征对该亚科进行了系统发育学研究。利用“非圆口类"的小腹茧蜂亚科Microgastrinae为根,以PAUP*4.0和MrBayes 3.0B4软件分别应用最大简约法（MP）和贝叶斯法对矛茧蜂亚科的分子数据和分子数据与非分子数据的结合体进行了运算分析;并以PAUP*4.0对矛茧蜂亚科的28S rDNA D2基因序列片段的碱基组成与碱基替代情况进行了分析。结果表明：矛茧蜂亚科的28S rDNA D2基因序列片段的GC含量在39.33%～48.28%之间变动,而对于碱基替代情况来讲,矛茧蜂亚科各成员间序列变异位点上颠换（transversion）大于转换（transition）。不同的分析算法所产生的系统发育树都表明矛茧蜂亚科是一个界限分明的单系群;在矛茧蜂亚科内,除了吉丁茧蜂族Siragrini为单系群外,其他族（矛茧蜂族Doryctini和方头茧蜂族Hecabolini）都是并系群。对于矛茧蜂亚科内各属之间的相互亲缘关系,不同算法所得的系统发育树的拓扑结构不完全一致,表明矛茧蜂亚科内（属及族）的系统发育关系还有待于进一步研究。     

Olive embryo in vitro germination potential: role of explant configuration and embryo structure among cultivars

The in vitro germination of excised embryos can break dormancy rapidly and shorten the time required to produce seedlings, speeding up olive breeding programmes as well as rootstock production. In this study, the in vitro germination potential of four Sicilian olive cultivars was evaluated during two years of experiments, using explants with three different morphological configurations that represent three different degrees of embryo exposure: (1) intact stoneless seeds containing the embryo, the endosperm and the seed coat (Emb+En+SC), (2) seeds without the seed coat (Emb+En) and (3) naked, isolated embryos (seed coat and endosperm both removed: Emb). Differences were found in the germination percentages and timing due to both genotype and explant type. The root and shoot meristems, the radicle-hypocotyl axis, the provascular tissues and embryo storage reserves were identified as embryo anatomical structures which could influence germination capacity. Observation of these structures, however, indicated similar germination potential among cultivars, suggesting possible differences in other dormancy factors. In spite of variation in cultivar performance, after 60 days of in vitro culture all cultivars demonstrated the highest germination of naked embryos (explant type 3) and lowest for stoneless seeds (explant type 1); stoneless seeds without the seedcoat (explant type 2) showed intermediate germination percentages.     

On the structure of the aedeagus in shield bugs (Heteroptera, Pentatomidae): II. Subfamily Podopinae

This paper is the second communication in a series of papers that treats the structure of the aedeagus in the family Pentatomidae and its taxonomic applications. Inner structures of the aedeagus were examined in the completely inflated state. The structure of the aedeagus was studied on 25 species. It is impossible to offer a diagnosis of the subfamily Podopinae on the basis of the structure of the aedeagus only, because this structure is very diverse. Nevertheless, some distinguishing characters were found in each of the three tribes, currently recognized within the subfamily. On the basis of the structure of the aedeagus, the tribe Graphosomatini can be subdivided into four groups, partly corresponding to the previously recognized tribes Graphosomatini (sensu Puchkov, 1965), Ancyrosomatini, and Trigonosmatini. Three different types of the structure of the aedeagus were revealed in the tribe Podopini. Special types of the structure of the aedeagus were found in the genera Bolbocoris and Testrica with unclear taxonomic position.     

A systematic survey of leaf and stem anatomical characters in the genus Gaultheria and related genera (Ericaceae)

MIDDLETON, D. J., A systematic survey of leaf and stem anatomical characters in the genus Gaultheria and related genera (Ericaceae). Leaf and stem anatomical characters of 123 species within the Gaultheria group of genera of the tribe Andromedeae are surveyed. A number of characters show considerable variation within a species which limits their taxonomic use. Other characters such as marginal sclerenchyma and, to a lesser extent, the hypodermis, the presence of adaxial stomata, the presence of free fibres and the pith type were of more taxonomic interest. These characters are discussed in relation to the generic and infrageneric classification of the group.     

Systematic and phylogenetic significance of the seed coat in the shrubby African Iridaceae, Nivenia, Klattia and Witsenia

MANNINGJ. C. & GOLDBLATT, P., 1991. Systematic and phylogenetic significance of the seed coat in the shrubby African Iridaceae, Nivenia, Klattia and Witsenia. The seeds of Nivenia, Klattia and Witsenia, a natural alliance within Nivenioideae, are among their most distinctive features, and seed number and shape constitute at least two synapomorphies for this monophyletic lineage. In this paper we survey the structure of the seed surface of five species of Nivenia and one each of Klattia and Witsenia by scanning electron microsopy (SEM), and study the development of the seed coat in a representative species of each of the three genera. Outgroups for comparison were selected from genera inside and outside the subfamily. The six genera of Nivenioideae are united in having a 3 (– 2) seriate outer integument but the shrubby genera are unique in the family in their tangentially flattened ovules which mature into depressed, flat, scutiform seeds. They display a range of variation in seed coat characters but are specialized in the subfamily and family in having a transparent outer integument. Other genera have pigmented deposits in the outer epidermis of the outer integument. Nivenia and Witsenia are further specialized in having a partially exfoliating outer integument. Klattia is unique among the shrubby genera in having deposits in the second layer of the tegmen, in retaining an intact outer epidermis of the tegmen, and in the fusiform shape of the epidermal cells and striate surface of the seed. Our results are incorporated in a cladogram that expresses our current understanding of the phylogenetic relationships of Nivenioideae.     

One-Seeded Fruits in the Core Caryophyllales: Their Origin and Structural Diversity

The core Caryophyllales consist of approximately 30 families (12 000 species) distributed worldwide. Many members evolved one-seeded or conjoined fruits, but their origin and structural diversity have not been investigated. A comparative anatomical investigation of the one-seeded fruits within the core Caryophyllales was conducted. The origin of the one-seeded fruits and the evolutionary reconstructions of some carpological characters were traced using a tree based on rbcl and matK data in order to understand the ancestral characters and their changes. The one-seeded fruit type is inferred to be an ancestral character state in core Caryophyllales, with a subsequent increase in the seed number seen in all major clades. Most representatives of the ‘Earlier Diverging’ clade are distinguished in various carpological traits. The organization of the pericarp is diverse in many groups, although fruits with a dry, many-layered pericarp, consisting of sclerenchyma as outer layers and a thin-walled parenchyma below, with seeds occupying a vertical embryo position, are likely ancestral character states in the core Caryophyllales clade. Several carpological peculiarities in fruit and seed structure were discovered in obligate one-seeded Achatocarpaceae, Chenopodiaceae, Nyctaginaceae, Seguieriaceae and Sarcobataceae. The horizontal embryo evolved in only certain groups of Chenopodiaceae. The bar-thickening of endotegmen cells appears to be an additional character typical of core Caryophyllales. The syncarpy-to-lysicarpy paradigm in Caryophyllaceae needs to be reinterpreted.