Scorzonera comprises 180–190 species and belongs to the subtribe Scorzonerinae. Its circumscription has long been the subject of debate and available molecular phylogenetic analyses affirmed the polyphyly of Scorzonera in its wide sense. We provide a re-evaluation of Scorzonera and other related genera, based on carpological (including anatomical) and extended molecular phylogenetic analyses. We present, for the first time, a comprehensive sampling, including Scorzonera in its widest sense and all other genera recognised in the Scorzonerinae. We conducted phylogenetic analyses using Maximum Parsimony, Maximum Likelihood and Bayesian analyses, based on sequences of the nuclear ribosomal ITS and of two plastid markers (partial rbcL and matK) and Maximum Parsimony for reconstructing the carpological character states at ancestral nodes. Achene characters, especially related to pericarp anatomy, such as general topography of the tissue types, disposition of the mechanical tissue and direction of its fibres, presence or absence of air cavities, provide, in certain cases, support for the phylogenetic lineages revealed. Confirming the polyphyly of Scorzonera, we propose a revised classification of the subtribe, accepting the genera Scorzonera (including four major clades: Scorzonera s. str., S. purpurea, S. albicaulis and Podospermum), Gelasia, Lipschitziagen. nov. (for the Scorzonera divaricata clade), Pseudopodospermum, Pterachaenia (also including Scorzonera codringtonii), Ramaliellagen. nov. (for the S. polyclada clade) and Takhtajaniantha. A key to the revised genera and a characterisation of the genera and major clades are provided.

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.

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The taxonomy of perennial Sesuvium species in Africa has been poorly investigated until now. Previously five perennial species of Sesuvium were recognised in Africa (S. congense, S. crithmoides, S. mesembryanthemoides, S. portulacastrum, and S. sesuvioides). Based on the differing number of stamens, S. ayresii is accepted here as being distinct from S. portulacastrum. Field observations in Angola also led the authors to conclude that S. crystallinum and S. mesembryanthemoides are conspecific with S. crithmoides. A new subspecies, Sesuvium portulacastrum subsp. persoonii, is described from West Africa (Cape Verde, Gambia, Guinea-Bissau, Mauritania, Senegal). The molecular phylogeny indicates the position of S. portulacastrum subsp. persoonii within the “American lineage” as a part of the Sesuvium portulacastrum complex which needs further studies. A diagnostic key and taxonomic notes are provided for the six perennial species of Sesuvium found in Africa and recognised by the authors (S. ayresii, S. congense, S. crithmoides, S. portulacastrum subsp. portulacastrum, S. portulacastrum subsp. persoonii, S. verrucosum and the facultatively short-lived S. sesuvioides). The distribution of S. crithmoides, previously considered to be endemic to Angola, is now confirmed for the seashores of Republic of Congo and DR Congo. The American species S. verrucosum is reported for the first time for Africa (the Macaronesian islands: Cape Verde and the Canaries). It is locally naturalised in Gran Canaria, being a potentially invasive species. These findings as well as new records of S. verrucosum from Asia and the Pacific Islands confirm its proneness to transcontinental introduction. Lectotypes of S. brevifolium, S. crithmoides, S. crystallinum and S. mesembryanthemoides are selected. The seed micromorphology and anatomy of the perennial African species is studied. Compared to the seeds of some annual African Sesuvium investigated earlier, those of perennial species are smooth or slightly alveolate. The aril is one-layered and parenchymatous in all species and usually tightly covers the seed. The aril detachments from the seed coat that form a white stripe near the cotyledon area easily distinguish S. verrucosum from other species under study.

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The former Chenopodiumsubgen.Blitum and the genus Monolepis (Chenopodioideae) are characterised in part by a reduced (0–4) number of perianth segments. According to recent molecular phylogenetic studies, these groups belong to the reinstated genera Blitum incl. Monolepis (tribe Anserineae) and Oxybasis (tribe Chenopodieae). However, key taxa such as C.antarcticum, C.exsuccum, C.litwinowii, C.foliosumsubsp.montanum and Monolepisspathulata were not included and so their phylogenetic position within the Chenopodioideae remained equivocal. These species and additional samples of Blitumasiaticum and B.nuttallianum were incorporated into an expanded phylogenetic study based on nrDNA (ITS region) and cpDNA (trnL-trnF and atpB-rbcL intergenic spacers and rbcL gene). Our analyses confirm the placement of C.exsuccum, C.litwinowii and C.foliosumsubsp.montanum within Blitum (currently recognised as Blitumpetiolare, B.litwinowii and B.virgatumsubsp.montanum, respectively); additionally, C.antarcticum, currently known as Oxybasisantarctica, is also placed within Blitum (reinstated here as B.antarcticum). Congruent with previous studies, two of the three accepted species of Monolepis – the type species M.trifida (= M.nuttalliana) as well as M.asiatica – are included in Blitum. The monotypic genus Carocarpidium described recently with the type C.californicum is not accepted as it is placed within Blitum (reinstated here as B.californicum). To date, few reliable morphological characters have been proposed that consistently distinguish Blitum (incl. two Monolepis species) from morphologically similar Oxybasis; however, two key differences are evident: (1) the presence of long-petiolate rosulate leaves in Blitum vs. their absence in Oxybasis and (2) a seed coat structure with the outer wall of the testa cells lacking stalactites (‘non-stalactite seed coat’) but with an obvious protoplast in Blitum vs. seed coat with the outer walls of the testa cells having stalactites (‘stalactite seed coat’) and a reduced protoplast in Oxybasis. Surprisingly, the newly sequenced North American Monolepisspathulata nested within the tribe Dysphanieae (based on ITS and trnL-trnF + rbcL + atpB-rbcL analyses).The phylogenetic results, as well as presence of the stalactites in the outer cell walls of the testa and lack of the rosulate leaves, confirm the distinctive nature of Monolepisspathulata from all Blitum and, therefore, the recent combination Blitumspathulatum cannot be accepted. Indeed, the morphological and molecular distinctive nature of this species from all Dysphanieae supports its recognition as a new monotypic genus, named herein as Neomonolepis (type species: N.spathulata). The basionym name Monolepisspathulata is also lectotypified on a specimen currently lodged at GH. Finally, while Micromonolepispusilla is confirmed as belonging to the tribe Chenopodieae, its position is not fully resolved. As this monotypic genus is morphologically divergent from Chenopodium, it is retained as distinct but it is acknowledged that further work is requ...

Corispermum iljinii from Qinghai and Ningxia provinces and C. nanum from Xizang (Tibet) are described as new species. The new variety C. dutreuilii var. montanum is described. Lectotypes of C. declinatum, C. elongatum and C. macrocarpum have been designated. The fruit anatomy of 16 Chinese taxa has been investigated for the first time. Both molecular and carpological data support the specific status of C. ellipsocarpum. The general fruit structure of Corispermoideae is specified and discussed.

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