Blake Vogt Number 4 Envelope

Kelly WA. 1973. Pollen morphology and relationships in the genus Calandrinia H.B.K. – Ph.D. diss., California State University, Northridge, California.

Kadereit G, Hohmann S, Kadereit JW. 2006. A synopsis of Chenopodiaceae subfam. Betoideae and notes on the taxonomy of Beta. – Willdenowia 36 (Spec. issue): 9-19. Gaskin JF. 2002. Tamaricaceae. – In: Kubitzki K, Bayer C (eds), The families and genera of vascular plants V. Flowering plants. Dicotyledons. Malvales, Capparales and non-betalain Caryophyllales, Springer, Berlin, Heidelberg, New York, pp. 363-368. Two strongly supported main clades can be discerned. One large monophyletic group comprises “the carnivorous clade” plus [[Tamaricaceae+Frankeniaceae]+[Polygonaceae+Plumbaginaceae]]. The second main clade includes [Rhabdodendraceae+[Simmondsiaceae+[Asteropeiaceae+Physenaceae]]] plus the core Caryophyllales in the strict sense.7/160–170. Pisoniella (1; P. arborescens; tropical and subtropical regions in America), Pisonia (22; tropical and subtropical regions on both hemispheres, especially America), Neea (70–80; southern Mexico, Central America, the West Indies, tropical South America; in Guapira?), Guapira (c 70; southern Mexico, Central America, the West Indies, tropical South America; incl. Neea?), Cephalotomandra (2; C. fragrans, C. panamensis; Central America)?, Grajalesia (1; G. fasciculata; Mexico)?, Neeopsis (1; N. flavifolia; Guatemala)? – Distribution as for Nyctaginaceae. Trees or (often scandent) shrubs. Leaves alternate (spiral), opposite or verticillate (sometimes anisophyllous), often glandular. Bracts two or three subtending each flower, caducous or persistent. Flowers sometimes zygomorphic. Perianth campanulate, urceolate, hypocraterimorphic or tubular, quinquelobate. Stamens (two to) five to ten (to numerous). Filaments connate at base, often adnate to pistil base. Pollen grains usually tricolpate. Stigma penicillate or papillate. Anthocarp oblong, clavate or ellipsoid, five-ribbed, coriaceous and glandular-viscid, or globose, carnose, glabrous. Testa in Pisonia multiplicative, unstructured. Embryo straight.

Butterworth CA. 2006b. Molecular phylogenetics of Cactaceae Jussieu – a review. – In: Sharma AK, Sharma A (eds), Plant genome biodiversity and evolution 1 C. Phanerogams (Angiosperm-Dicotyledons), Science Publ., Enfield, New Hampshire, pp. 489-524.Meulen-Bruijns C van der. 1976. The vascular pattern in the flower of some Mesembryanthemaceae: Aptenia cordifolia and Dorotheanthus bellidiformis. – Blumea 23: 189-201. Datson B. 2002. Samphires in Western Australia: a field guide to Chenopodiaceae tribe Salicornieae. – Department of Conservation and Land Management, Perth, Western Australia. Sharma AK, Banik M. 1965. Cytological investigation of different genera of Amaranthaceae with a view to trace their interrelationships. – Bull. Bot. Soc. Bengal 19: 40-50.

Barthlott W, Porembski S, Kluge M, Hopke J, Schmidt L. 1997. Selenicereus wittii (Cactaceae): an epiphyte adapted to Amazonian Igapó inundation forests. – Plant Syst. Evol. 206: 175-185.

Weller SG, Sakai AK, Wagner WL. 2001. Artificial and natural hybridization in Schiedea and Alsinidendron (Caryophyllaceae: Alsinoideae): the importance of phylogeny, genetic divergence, breeding system, and population size. – Syst. Bot. 26: 571-584.Hood ME, Mena-Ali JI, Gibson AK, Oxelman B, Giraud T, Yockteng R, Arroyo MTK, Conti F, Petersen AB, Gladieux P, Antonovics J. 2010. Distribution of the anther-smut pathogen Microbotryum on species of the Caryophyllaceae. – New Phytol. 187: 217-229.

Ovules Placentation basal-axile. Ovules one to three per carpel, anatropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Obturator placental. Parietal tissue? Megagametophyte monosporous, Polygonum type. Antipodal cells usually ephemeral. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?
Systematics Trigastrotheca (3; T. molluginea, T. pentaphylla, T. stricta; tropical and subtropical Asia and Australia), Mollugo (c 15; tropical to warmtemperate regions in North to South America, one species, M. disticha, in India and Sri Lanka), Glinus (c 10; tropical to warmtemperate regions on both hemispheres), Paramollugo (6; P. nudicaulis: tropical Africa, Madagascar, Yemen, Socotra, tropical Asia; P. angustifolia: Somalia; P. decandra, P. elliotii, P. simulans: Madagascar; P. digyna: New Caledonia), Hypertelis (5; H. cerviana, H. fragilis, H. spergulacea, H. umbellata, H. walteri; southwestern and southeastern Europe, tropical and subtropical regions, South Africa), Polpoda (2; P. capensis, P. stipulacea; Western Cape), Psammotropha (11; tropical and southern Africa), Adenogramma (10–11; Northern and Western Cape), Suessenguthiella (1; S. scleranthoides; Namibia, Northern and Western Cape), Coelanthum (3; C. grandiflorum, C. semiquinquefidum, C. verticillatum; southern Namibia, Northern and Western Cape), Pharnaceum (c 28; southern Africa). Olvera HF. 2003. Classification of the North American species of Atriplex section Obione (Chenopodiaceae) based on numerical taxonomic analysis. – Taxon 52: 247-260. Neumann M. 1935. Die Entwicklung des Pollens, der Samenanlage und des Embryosackes von Pereskia amapola var. argentina. – Österr. Bot. Zeitschr. 84: 1-30.

Apatesieae are sister-group to the clade [Dorotheantheae+[Delospermeae+Ruschieae]] and possess an annular holonectary. The capsule often lack hygrochastic properties and has very reduced expanding keels. The clade [Dorotheantheae+[Delospermeae+Ruschieae]] has a covering membrane. Dorotheantheae are annual herbs with semi-succulent leaves. The nectar is a wide and flat meronectary. The [Delospermeae+Ruschieae] clade has a lophomorphic nectary and a hygrochastic capsule. Moreover, the intron of the plastid gene rpoC1 is absent (lost). Delospermeae have a lophomorphic meronectary. Ruschieae have a lophomorphic holonectary and the nuclear gene ARP (a leaf developmental gene) is often duplicated (absent in some species). Fruit characters are highly homoplasious. Apatesieae are confined to the Cape Floristic Region.Simonet M. 1930. Nouvelles recherches sur le nombre des chromosomes chez les Iris et sur l’existence de mitoses didiploïdes dans ce genre. – Compt. Rend. Soc. Biol. Paris 103: 1197-1200.

Ormond WT, Cortella C de, Bezerra Pinheiro AR, Rodrigues Correia MC. 1978. Contribuição ao estudo anatômico das populações diplóide e tetraplóide de Petiveria alliacea L. – Bol. Mus. Nac. Rio de Janeiro, Bot. II, 51: 1-13. Vegetative anatomy Phellogen ab initio subepidermal. Primary vascular tissue a cylinder, without separate vascular bundles. Continuous sclerenchyma cylinder surrounding vascular cylinder in young stems. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with small simple pits or reduced bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate, homocellular or heterocellular (consisting of erect or square cells). Axial parenchyma apotracheal diffuse, paratracheal aliform, confluent, or unilateral. Sieve tube plastids? Nodes 1:1, unilacunar with one leaf trace. Pericyclic sclereids present. Secondary phloem with brachysclereids. Medullary parenchyma with sclereids. Crystals usually absent (rarely a single crystal per cell); silica grains absent. Prismatic crystals sometimes present in wood ray cells. Seeds Aril absent. Seed coat testal. Testa multiplicative, vascularized (prominent postchalazal vascular bundles reaching to micropyle). Exotestal cells palisade, with thickened walls. Mesotesta aerenchymatous. Endotesta mainly collapsed. Tegmen collapsed. Perisperm not developed. Endosperm sparse or absent. Embryo large, straight, well differentiated, chlorophyll? Cotyledons two, thick, fleshy, with simmondsin (a nitrile glycoside), aleurone grains and a liquid wax. Germination phanerocotylar?Borsch T, Flores-Olvera H, Zumaya S, Müller K. 2018. Pollen characters and DNA sequence data converge on a monophyletic genus Iresine (Amaranthaceae, Caryophyllales) and help to elucidate its species diversity. – Taxon 67: 944-976.Wood rays absent. Stipules absent. Hypanthium absent. Stamens ten or more. Nectary position. Placentation sometimes axile at least basally when ovary young. Fruit usually a septicidal and loculicidal capsule (sometimes a nutlet). Fruit with two or more seeds.Robins RJ, Juniper BE. 1980a. The secretory cycle of Dionaea muscipula Ellis I. The fine structure and the effect of stimulation on the fine structure of the digestive gland cells. – New Phytol. 86: 279-296.

Pollen grains Microsporogenesis simultaneous. Pollen grains triporate (Aldrovanda) or 10–30-porate (Dionaea, Drosera), shed as tetrads (in Drosera usually with radial discs), usually bicellular (sometimes tricellular) at dispersal. Exine usually tectate (sometimes intectate), with columellate infratectum, usually spinulate (not in Dionaea), in Drosera often operculate.Androecium Stamens usually ten (rarely five or 15), in one whorl (staminal primordia five?); antepetalous stamens larger. Filaments short, fleshy, connate at base, adnate to petal bases (epipetalous). Anthers basifixed, non-versatile, tetrasporangiate, introrse or latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.Seeds Aril tiny (present at hilum). Testa? Outer exotestal wall with stalactite-shaped outgrowths? Tegmen? Perisperm copious, mealy. Endosperm rudimentary or absent. Embryo annular, peripheral, enclosing perisperm, chlorophyll? Cotyledons two. Germination?Seeds Seed large. Aril absent. Seed coat vascularized, 16 to 20 cell layers thick, with insignificantly thickened cell walls. Exotesta? Endotesta? Tegmen? Perisperm not developed. Endosperm absent. Embryo straight, chlorophyll? Cotyledons two, unequally sized. Germination?Ihlenfeldt H-D. 1978. Monographie der Mitrophyllinae Schwantes III. Morphologie und Taxonomie der Gattung Oophytum N. E. Br. (Mesembryanthemaceae). – Bot. Jahrb. Syst. 99: 303-328.

Heimerl A. 1934b. Phytolaccaceae. – In: Engler A (†), Pax F, Harms H (eds), Die natürlichen Pflanzenfamilien, 2. Aufl., Bd. 16c, W. Engelmann, Leipzig, pp. 135-164.
Sukhorukov AP. 1999b. Some general evolutionary trends in the genus Atriplex L. s.l. (Chenopodiaceae). – In: Melikyan AP, Lotova LI (eds), Tenth Moscow Meeting in Plant Phylogeny (Materials), Moscow, pp. 166-168. [in Russia]

Barthlott W. 1983. Biogeography and evolution in neo- and palaeotropical Rhipsalinae. – In: Kubitzki K (ed), Dispersal and distribution, Sonderbd. Naturbd. Naturwiss. Ver. Hamburg 7, P. Parey, Hamburg, pp. 241-248.

Shepherd KA. 2007. Three new species of Tecticornia (formerly Halosarcia) (Chenopodiaceae: Salicornioideae) from the Ememaean Botanical Province, Western Australia. – Nuytsia 17: 353-366.
Phytochemistry Virtually unknown. Cytotoxic 16-β-[(D-xylopyranosyl)oxy]oxohexadecanyl triterpene glycosides and oxohexadecanoic acid present. Ellagic acid? Alkaloids?

Brockington SF, Yang Y, Gandia-Herrero F, Covshoff S, Hibberd JM, Sage RF, Wong GKS, Moore MJ, Smith SA. 2015. Lineage-specific gene radiations underlie the evolution of novel betalain pigmentation in Caryophyllales. – New Phytol. 207: 1170-1180.
Bortenschlager S, Auimger A, Blaha J, Simonsburger P. 1972. Pollen morphology of Achatocarpaceae (Centrospermae). – Bot. Naturwiss.-Med. Vereins Innsbruck 59: 7-13.Nepenthales Bercht. et J. Presl, Přir. Rostlin: 267. Jan-Apr 1820 [‘Nepenthaceae’]; Nepenthineae Link, Handbuch 1: 369. 4-11 Jul 1829; Nepenthanae Takht. ex Reveal in Phytologia 79: 71. 29 Apr 1996

Ovules Placentation basal. Ovule one per ovary, hemianatropous or (following fertilization) anacampylotropous, ascending, usually bitegmic (in Abronia and Boerhavia unitegmic), crassinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Parietal tissue? Megagametophyte usually monosporous, Polygonum type (in species of Mirabilis Tridax type: nuclei arranged according to the principle 1:2:1). Antipodal cells usually ephemeral (often persistent, long-lived and somewhat proliferating). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad. Polyembryony present in Boerhavia.
Sherbrooke WC, Haase EF. 1974. Jojoba: a wax-producing shrub of the Sonoran Desert. – Arid Lands Resource Information Paper 5, University of Sonora, Office of Arid Lands Studies, Tucson, Arizona.Hong S-P, Oh I-C, Ronse De Craene LP. 2005. Pollen morphology of the genera Polygonum s. str. and Polygonella (Polygoneae: Polygonaceae). – Plant Syst. Evol. 254: 13-30.

Butterworth CA, Butterworth KM, Fitz Maurice WA, Fitz Maurice B. 2007. A localized loss of the chloroplast rpl16 intron in Mammillaria series Stylothelae (Cactaceae) delineates members of the M. crinita group. – Bradleya 25: 187-192.
Grady BR, Reveal JL. 2011. New combinations and a new species of Eriogonum (Polygonaceae: Eriogonoideae) from the Great Basin Desert, United States. – Phytotaxa 24: 33-38.Trichomes Glandular hairs multicellular, insensitive, stalked or sessile, secreting a viscous mucilage with and without proteolytic enzymes, respectively. Glandular heads in Triphyophyllum consisting of two layers of secretory cells and inside these layers an endodermis; glandular stalk usually vascularized (xylem and phloem). Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpate, shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, perforate to finely punctate, spinulate. Androecium Stamens (five to) seven (to ten). Filaments connate at base, adnate to tepals (epitepalous). Anthers dorsifixed, versatile?, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.Fruit Usually a nut (often a utriculus or an achene) or an irregularly dehiscent capsule (often a pyxidium, rarely a berry, a baccaceous fruit or a drupe; adjacent ovaries sometimes fusing and forming a syncarp), often surrounded by more or less fleshy perianth forming an anthocarp; persistent and often accrescent bracts and floral prophylls sometimes forming parts of dispersal unit.

Heimerl A. 1934c. Achatocarpaceae. – In: Engler A (†), Pax F, Harms H (eds), Die natürlichen Pflanzenfamilien, 2. Aufl., Bd. 16c, W. Engelmann, Leipzig, pp. 174-178.
Stutz HC, Chu G-L, Sanderson SC. 1990. Evolutionary studies of Atriplex: phylogenetic relationships of Atriplex pleiantha. – Amer. J. Bot. 77: 364-369.

Ellis AG, Weis AE, Gaut BS. 2006. Evolutionary radiation of “stone plants” in the genus Argyroderma (Aizoaceae): unravelling the effects of landscape, habitat, and flowering time. – Evolution 60: 39-55.
Vegetative anatomy Mycorrhiza usually absent (sometimes with vesicular-arbuscular mycorrhiza). Kranz’ anatomy present in numerous species. Phellogen usually superficial or pericyclic. Stem collenchyma well developed. Cortical and/or medullary bundles usually present. Primary medullary strands usually wide. Endodermis usually significant. Secondary lateral growth usually anomalous (polycyclic, anomalous secondary vascular bundles from concentric cambia; not in Polycnemoideae) or absent. Pericyclic fibres few or absent. Vessel elements with simple perforation plates; lateral pits usually alternate (sometimes opposite), simple or bordered pits. Imperforate tracheary xylem elements libriform fibres (with cell nuclei) with simple or (reduced) bordered pits, non-septate (also vasicentric tracheids). Wood rays usually absent (sometimes uniseriate or multiseriate, homocellular or heterocellular). Axial parenchyma paratracheal scanty, aliform, winged-aliform, confluent, vasicentric, or banded. Vessel elements, fibres and/or parenchyma sometimes partially or entirely storied. Intraxylary (concentric or diffuse) phloem present. Sieve tube plastids P3cf type, without a central protein crystal, with circular peripheral protein fibrils (sometimes with starch grains). Nodes 1:1 or 1:3, unilacunar with one or three leaf traces (sometimes 1:5, unilacunar with five traces), often swollen. Heartwood sometimes with gum-like substances. Calciumoxalate usually as crystal sand, druses or prismatic crystals (raphides and styloids absent).Ovules Placentation basal and free central. Ovule one per ovary, usually orthotropous (occasionally campylotropous to anatropous), ascending, usually bitegmic (rarely unitegmic), crassinucellar. Funicle present (representing a reduced free central placenta?), or absent. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Nucellar beak present. Hypostase present or absent. Archespore usually unicellular (rarely multicellular). Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells usually uninucleate (in Rumex multinucleate). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad.

Frajman B, Oxelman B. 2007. Reticulate phylogenetics and phytogeographical structure of Heliosperma (Sileneae, Caryophyllaceae) inferred from chloroplast and nuclear DNA sequences. – Mol. Phylogen. Evol. 43: 140-155.
Fruit A one-seeded berry (Rivina, Trichostigma), a nutlet (in Petiveria with bristle-like processes) or a samara (in Seguieria and Gallesia with enlarged persistent lignified sepals; in Schindleria, Ledenbergia and Hilleria a utriculus; in Ledenbergia with wing-like sepals). Pericarp usually adnate to seed.Watson SL. 1877. Descriptions of new species of plants, with revisions of Lychnis, Eriogonum and Chorizanthe. – Proc. Amer. Acad. Arts Sci. 12: 246-278.

Ihlenfeldt H-D, Hartmann HEK. 1982. Leaf surfaces in Mesembryanthemaceae. – In: Cutler DF, Alvin KL, Price CE (eds), The plant cuticle, Academic Press, London, pp. 397-423.

Raj G, Kurup R, Hussain AA, Baby S. 2011. Distribution of naphthoquinones, plumbagin, droserone, and 5-O-methyl droserone in chitin-induced and uninduced Nepenthes khasiana: molecular events in prey capture. – J. Exp. Bot. 62: 5429-5436.
Behnke H-D, Mabry TJ, Eifert IJ, Pop L. 1975. P-type sieve-element plastids and betalains in Portulacaceae (including Ceraria, Portulacaria, Talinella). – Can. J. Bot. 53: 2103-2109.Phytochemistry Insufficiently known. Betalains (betacyanins and betaxanthins) present. Oxalic acid often accumulated. Triterpene saponins? Anthocyanins not found.

Helsen P, Browne RA, Anderson DJ, Verdyck P, Dongen S. 2009. Galapagos’ Opuntia (prickly pear) cacti: extensive morphological diversity, low genetic variability. – Biol. J. Linn. Soc. 96: 451-461.Ohsako T, Fukuoka S, Bimb HP, Baniya BK, Yasui Y, Ohnishi O. 2001. Phylogenetic analysis of the genus Fagopyrum (Polygonaceae), including the Nepali species F. megacarpum, based on nucleotide sequence of the rbcL-accD region in chloroplast DNA. – Fagopyrum 18: 9-14.

Douglas NA, Manos PS. 2007. Molecular phylogeny of Nyctaginaceae: taxonomy, biogeography, and characters associated with radiation of xerophytic genera in North America. – Amer. J. Bot. 94: 856-872.
Flowers Actinomorphic. Hypogyny. Sepals five, with valvate or plicate aestivation, persistent, often membranous or petaloid, with five or ten ridges, usually connate into a tube (rarely free). Petals five, with usually contorted (sometimes imbricate) aestivation, often persistent, connate usually only at base (sometimes entirely), in Aegialitis coriaceous; petal and staminal primordia common. Nectaries often present (sometimes as five glands alternating with stamens). Disc absent. Seeds Aril usually absent. Seed coat usually exotestal and/or endotegmic. Exotesta often tanniniferous; outer exotestal wall sometimes with stalactite-like processes. Endotesta sometimes thickened, often crystalliferous. Exotegmen? Endotegmen sometimes with rod-shaped thickenings in cell walls, often tanniniferous. Perisperm copious, starchy (with starch grains), surrounded by embryo, or not developed. Endosperm copious, sparse or absent. Embryo usually lateral-peripheral, curved around perisperm or straight (rarely cochleate or annular), with or without chlorophyll. Cotyledons usually two (rarely one or three). Germination phanerocotylar. Leaves Alternate (spiral), simple, entire, often reduced and scale-like, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation? Stomata? Cuticular waxes? Leaf margin entire.

Talavera S, Muñoz Garmendia F. 1989. Sinopsis del género Silene L. (Caryophyllaceae) en la Península Ibérica y Baleares. – An. Jard. Bot. Madrid 45: 407-460.Gynoecium Pistil composed of three connate carpels. Ovary superior, ab initio multilocular, later unilocular. Stylodia (four or) five (or six). Stigmas receptive on both surfaces, papillate, Dry type. Pistillodium absent.Kim S-T, Sultan SE, Donoghue MJ. 2008. Allopolyploid speciation in Persicaria (Polygonaceae): insights from a low-copy nuclear region. – Proc. Natl. Acad. Sci. U.S.A. 105: 12370-12375.

Lledó MD Crespo MB, Fay MF, Chase MW. 2005. Molecular phylogenetics of Limonium and related genera (Plumbaginaceae): biogeographical and systematic implications. – Amer. J. Bot. 92: 1189-1198.
Eliasson UH. 1987. 44. Amaranthaceae. – In: Harling G, Sparre B (eds), Flora of Ecuador 28, Swedish Natural Science Research Council, Stockholm, pp. 1-137.Behnke H-D, Mabry TJ, Neumann P, Barthlott W. 1983. Ultrastructural, micromorphological and phytochemical evidence for a ”central position” of Macarthuria (Molluginaceae) within the Caryophyllales. – Plant Syst. Evol. 143: 151-161. Fuertes-Aguilar J, Rossello JA, Nieto Feliner G. 1999. Nuclear ribosomal DNA (nrDNA) concerted evolution in natural and artificial hybrids of Armeria (Plumbaginaceae). – Mol. Ecol. 8: 1341-1346. Inflorescence Usually terminal, thyrsopaniculate, scorpioid or cincinnus (flowers in Aldrovanda solitary). Bracts and floral prophylls (bracteoles) sometimes absent.Shepherd KA, Lyons MN. 2009. Three new species of Tecticornia (Chenopodiaceae, subfamily Salicornioideae) identified through Salinity Action Plan surveys of the wheatbelt region, Western Australia. – Nuytsia 19: 167-180.Moreno NC, Amarilla LD, Las Peñas ML, Bernardello G. 2015. Molecular cytogenetic insights into the evolution of the epiphytic genus Lepismium (Cactaceae) and related genera. – Bot. J. Linn. Soc. 177: 263-277.

The clade [Caryophyllaceae+[Achatocarpaceae+Amaranthaceae]] has the potential synapomorphies: stamens as many as tepals, antetepalous; a single ovule; parietal tissue approx. four cell layers thick; nucellar cap two to four cell layers thick; especially outer exotestal cell walls thick, with stalactite-like processes; mitochondrial genes rps1 and rps19 absent (lost); and often presence of phytoecdysteroids.
Seeds Aril present, covering larger part of seed. Seed coat exotestal-endotegmic? Exotesta palisade, with non-lignified cell walls. Endotesta? Exotegmen? Endotegmen enlarged, persistent. Perisperm copious, nutritious. Endosperm almost absent. Embryo curved, chlorophyll? Cotyledons two. Germination?McNeill J, Majumdar NC. 1980. A new species of Arenaria subgenus Odontostemma from Tibet, with a review of the status of the genus Gooringia (Caryophyllaceae). – Bot. J. Linn Soc. 80: 371-378.

Vegetative anatomy Mycorrhiza absent. Phellogen ab initio pericyclic. Primary vascular tissue a cylinder of bundles. Medullary vascular bundles may be present; young stems with cortical bundles. Medulla, pericycle and bark tissues with large idioblasts having spiral thickenings and cap-like apices. Pith and cortex with idioblastic helical-banded fibre-sclereids. A certain amount of lateral growth may occur. Vessel elements dimorphic, with simple (sometimes vestigial scalariform) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits, usually non-septate (occasionally septate). Wood rays usually uniseriate or biseriate (rarely multiseriate), homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal banded. Sieve tube plastids S type. Nodes 5–9:5–9, multilacunar with five to nine leaf traces. Older parts of secondary xylem with gum-like deposits. Silica bodies present. Crystals?Inoue M, Ohtani K, Kasai R, Okukubo M, Andriansiferana M, Yamasaki K, Koike T. 2009. Cytotoxic 16-β-[(D-xylopyranosyl)oxy]oxohexadecanyl triterpene glycosides from a Malagasy plant, Physena sessiliflora. – Phytochemistry 70: 1195-1202.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually tri- or hexacolp(oroid)ate (sometimes hexarugate, rarely dicolp[oroid]ate), shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, echinate to spinulate.
6/c 95. Aspazoma (1; A. amplectens; Namaqualand and Richtersveld in Northern and Western Cape), Brownanthus (10; southern Angola, Namibia, Northern and Western Cape), Caulipsolon (1; C. rapaceum; Namaqualand in Northern Cape), Mesembryanthemum (c 70; southern Angola, Namibia, western and central South Africa to Eastern Cape), Psilocaulon (13; southern Angola, western Namibia, western, southern and central South Africa), Synaptophyllum (1; S. juttae; near Lüderitz in southwestern Namibia). – Southern Africa. CAM photosynthesis present. Cortical vascular bundles present. Stem sometimes with succulent persistent green cortex. Wide-band tracheids absent. Stomata usually transversely orientated. Inflorescence indistinct. Flowers tetra- or pentamerous. Half epigyny. Tepals, petaloid staminodia and stamens often more or less united at base into a tube. Nectaries usually hollow or koilomorphic (shell-shaped), discontinuous (meronectary). Pistil composed of (three or) four or five (or six) connate carpels. Placentation axile. Expanding fruit keels entirely septal. Alkaloids usually present.Zibareva L, Volodin V, Saatov Z, Savchenko T, Whiting P, Lafont R, Dinan L. 2003. Distribution of phytoecdyosteroids in the Caryophyllaceae. – Phytochemistry 64: 499-517.

Hartl WP, Barbier B, Klapper H, Müller P, Barthlott W. 2003. Dimorphism of calcium oxalate crystals in stem tissues of Rhipsalideae (Cactaceae) – a contribution to the systematics and taxonomy of the tribe. – Bot. Jahrb. Syst. 124: 287-302.
Schatz GE, Lowry PP II, Wolf A-E. 1999. Endemic families of Madagascar IV. A synoptic revision of Asteropeia (Asteropeiaceae). – Adansonia, sér. III, 21: 255-268.Carlquist SJ. 2002. Wood anatomy and successive cambia in Simmondsia (Simmondsiaceae): evidence for inclusion in Caryophyllales s.l. – Madroño 49: 158-164.

Erdtman G, Dunbar A. 1966. Notes on electron micrographs illustrating the pollen morphology in Armeria maritima and A. sibirica. – Grana Palynol. 6: 435-475.
Akhani H, Barroca J, Koteeva N, Voznesenskaya E, Franceschi V, Edwards G, Ghaffari SM, Ziegler H. 2005. Bienertia sinuspersici (Chenopodiaceae): a new species from Southwest Asia and discovery of a third terrestrial C4 plant without Kranz anatomy. – Syst. Bot. 30: 290-301.Androecium Stamens ten or 25 to 30 (in Dioncophyllum sometimes five antepetalous). Filaments free or slightly connate at base, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective somewhat prolonged. Tapetum secretory. Staminodia absent.

Loza-Cornejo S, Terrazas T. 1996. Anatomía del tallo y de la raíz de dos especies de Wilcoxia Britton y rose (Cactaceae) del noreste de México. – Bol. Soc. Bot. México 59: 13-23.
Phytochemistry Flavonols (kaempferol, quercetin), 6-7-methylene-dioxyflavonols, isoflavones, flavonol sulphates, betacyanins (e.g. amaranthin, celosianin and betamin or phyllocactin), betaxanthins, isoquinoline alkaloids and other alkaloids (particularly in Salsoleae), triterpene saponins, cyanogenic compounds, betaine, anthraquinones, and sterols present. Ferulic acid present in non-lignified cell walls. Ellagic acid and proanthocyanidins not found. Nitrate or free oxalates accumulated in many species. Monoszon MK. 1964. Pollen of halophytes and xerophytes of the Chenopodiaceae family in the periglacial zone of the Russian plain. – Pollen Spores 6: 147-155. Calvente A, Moraes EM, Lavor P, Bonatelli IAS, Nacaguma P, Versieux LM, Taylor NP, Zappi DC. 2017. Phylogenetic analyses of Pilosocereus (Cactaceae) inferred from plastid and nuclear sequences. – Bot. J. Linn. Soc. 183: 25-38.Vegetative anatomy Phellogen ab initio subepidermal. Medulla with or without diaphragms. Secondary lateral growth at least sometimes anomalous (from concentric/successive cambia). Vessel elements with simple perforation plates; lateral pits alternate, simple and/or bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple pits, usually non-septate (in Rivina?) (also vasicentric fibres). Wood rays usually multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal vasicentric scanty or banded. Intraxylary (concentric) phloem present. Sieve tube plastids P3cf type, with a central globular protein crystal surrounded by a ring of protein filaments. Nodes 1:1, unilacunar with one leaf trace. Calciumoxalate as prismatic crystals, styloids and elongate crystals (in Gallesia and Seguieria hexagonal crystals).

Hernándes-Lopes J, Oliveira-Neto MA, Melo-de-Pinna GFA. 2016. Different ways to build succulent leaves in Portulacineae (Caryophyllales). – Intern. J. Plant Sci. 177: 198-208.
Conran JG, Jaudzems G, Hallam ND. 2007. Droseraceae gland and germination patterns revisited: support for recent molecular phylogenetic studies. – Carniv. Plants Newsl. 36: 14-20.Androecium Stamens (one to) five to more than 4.000 (staminal primordia usually five), in one or more whorls or in several groups (outer stamens often initiated in pairs), antetepalous when stamens isomerous relative to tepals; staminal development usually centrifugal. Filaments free or more or less connate, sometimes adnate to sepals or petals. Anthers basifixed or dorsifixed (sometimes latrorse), versatile or non-versatile, usually tetrasporangiate (rarely disporangiate), usually introrse (sometimes extrorse), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores or pore-like slits); outer parietal cells developing directly into endothecium. Tapetum usually secretory (rarely amoeboid-periplasmodial). Female flowers often with staminodia (staminodia sometimes numerous in bisexual flowers). ‘Petaloids’ possibly in reality petaloid staminodia, developing simultaneously as or after androecium (not prior to); petaloid staminodia (’petals’) and ’antepetalous’ stamens possibly forming a developmental unit. Laundon JR. 1959. Droseraceae. – In: Hubbard CE, Milne-Redhead E (eds), Flora of tropical East Africa, Crown Agents for Oversea Governments and Administrations, London, pp. 1-6. Trichomes Hairs unicellular or multicellular, uniseriate or m
ultiseriate, simple or branched (two-branched, T-shaped, dendritic, candelabra-shaped, stellate, fasciculate, lepidote, rosulate, barbed or vesicular), or absent; glandular hairs often frequent, multicellular, stalked or sessile (occasionally peltate-lepidote; sometimes secreting viscous mucilage).

Lahondère C. 2004. Les salicornes s.l. (Salicornia L., Sarcocornia A. J. Scott et Arthrocnemum Moq.) sur le côtes françaises. – Bull. Soc. Bot. Centre-Ouest, sér. II, num. spec. 24.
Ronse De Craene L-P, Volgin SA, Smets EF. 1999. The floral development of Pleuropetalum darwinii, an anomalous member of Amaranthaceae. – Flora 194: 189-199.

The monophyletic group [Talinaceae+[Portulacaceae+[Anacampserotaceae+Cactaceae]]] is supported by the potential synapomorphies (Stevens 2001 onwards): presence of mucilaginous cells; absence of pericyclic fibres; leaves with axillary uniseriate, biseriate or multiseriate hairs, bristles or scales; stomata parallelocytic (stoma with a lateral series of at least three alternating subsidiary cells increasing in size away from guard cells; also present in Montiaceae); pericarp two-layered; fruit covered by dry tepals; exocarp completely or almost caducous. Portulacaceae and Cactaceae may be sister-groups, according to Ocampo & Columbus (2010), using data mainly from non-coding plastid DNA. Portulaca and at least Pereskia share, e.g., a c. 500 bp deletion in rbcL, and non-lignified parenchyma cells are sometimes present in the wood in Portulaca and Cactaceae. On the other hand, Nyffeler & Eggli (2010), using sequence data from matK and ndhF, identified Anacampserotaceae and Cactaceae as sister-groups (they share among morphological features the character of numerous stamens). A special arrangement of testa cells along the dorsal juncture, presence of a dry aril and a central field type of cuticular ornamentation are characteristic features in several clades of Caryophyllales such as Cactaceae and Portulacaceae (Barthlott 1984).Brysting AK, Elven R. 2000. The Cerastium alpinum-C. arcticum complex (Caryophyllaceae): numerical analyses of morphological variation and a taxonomic revision of C. arcticum Lange s.l. – Taxon 49: 189-216.

Klak C, Hanáček P, Bruyns PV. 2017a. Out of southern Africa: origin, biogeography and age of the Aizooideae (Aizoaceae). – Mol. Phylogen. Evol. 109: 203-216.
Systematics Kewa (8; K. angrae-pequenae, K. arenicola, K. bowkeriana, K. caespitosa, K. salsoloides: eastern and southern Africa; K. suffruticosa: Madagascar; K. acida: St. Helena).Seeds Seeds with funicular strophiole or (in, e.g., Calyptrotheca) aril. Testa? Tegmen? Perisperm sparse or absent. Endosperm entirely or almost entirely absent. Embryo large, curved around perisperm, well differentiated, chlorophyll? Cotyledons two, often fleshy. Germination phanerocotylar.

Mattfeld J. 1938. Über eine angebliche Drymaria Australiens nebst Bemerkungen über die Staminaldrüsen und die Petalen der Caryophyllaceae. – Feddes Repert. Beih. 100: 147-164.
Sobolevskaya KA, Vysochina GI. 1972. Ecological and geographical aspects and some problems of the chemosystematics of section Aconogonon Meisn. of genus Polygonum. – Izv. Sib. Otd. Akad. Nauk. SSSR, ser. Biol. Med. Nauk. 3: 29-39. [In Russian with English summary]
Schuster TM, Setaro SD, Kron KA. 2013. Age estimates for the buckwheat family Polygonaceae based on sequence data calibrated by fossils and with a focus on the Amphi-Pacific Muehlenbeckia. – PloS ONE 8:e61261
Sun Y-X, Wang A, Wan D, Wang Q, Liu J. 2012. Rapid radiation of Rheum (Polygonaceae) and parallel evolution of morphological traits. – Mol. Phylogen. Evol. 63: 150-158.

Seeds Aril sometimes present (surrounding seed in Sesuvioideae). Exotesta palisade or tangentially elongate. Endotesta and tegmen usually crushed. Perisperm copious, starchy. Endosperm sparse or absent. Suspensor massive, usually biseriate to multiseriate. Embryo peripheral, curved around perisperm, without chlorophyll. Cotyledons two. Radicula dorsal. Germination phanerocotylar.
Ovules Placentation usually axile (in early stage, primary; rarely basal or apical) to parietal (in later stage, secondary). Ovules one to numerous per carpel (in Acrosanthes a single basal ovule per carpel), campylotropous (hemianatropous?) to anacampylotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument two or three cell layers thick. Inner integument two cell layers thick. Obturator placental. Parietal tissue approx. three cell layer thick. Nucellar cap micropylar. Apical cells of megasporangium often radially elongate. Megagametophyte usually monosporous, Polygonum type (rarely Endymion, Penaea, Drusa, or Adoxa type). Endosperm development ab initio nuclear. Endosperm haustoria chalazal? Embryogenesis caryophyllad or solanad.Habit Usually bisexual (rarely dioecious), evergreen trees, shrubs or lianas (Ercilla), perennial or annual herbs (Anisomeria is a succulent). Anisomeria and some species of Phytolacca with napiform roots. Androecium Stamens c. 20–25, in several (four or five?) whorls. Filaments short, free from each other and from tepals, inserted at annular disc. Anthers subbasifixed, non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent. Gynoecium Pistil composed of (one or) two or three (to six) connate carpels (median carpel sometimes abaxial); when two carpels then usually transverse. Ovary usually superior (rarely semi-inferior), unilocular. Style single, simple, or stylodia two or three, long or short, more or less connate. Stigma one, capitate (simple or penicillate), or stigmas two or three (to six), narrowly elongate, papillate, Dry type, often persistent. Pistillodium usually absent (male flowers sometimes with pistillodium). Schuster TM, Reveal JL, Bayly MJ, Kron KA. 2015. An updated molecular phylogeny of Polygonoideae (Polygonaceae): relationships of Oxygonum, Pteroxygonum, and Rumex, and a new circumscription of Koenigia. – Taxon 64: 1188-1208. Fuentes-Bazán S, Uotila P, Borsch T. 2012. A novel phylogeny-based generic classification for Chenopodium sensu lato, and a tribal rearrangement of Chenopodioideae (Chenopodiaceae). – Willdenowia 42: 5-24. Sloan DB, Alverson AJ, Wu M, Palmer JD, Taylor DR. 2012. Recent acceleration of plastid sequence and structural evolution coincides with extreme mitochondrial divergence in the angiosperm genus Silene. – Genom Biol. Evol. 4: 294-306. Olvera HF, Vrijdaghs A, Ochoterena H, Smets E. 2011. The need to re-investigate the nature of homoplastic characters: an ontogenetic case study of the ‘bracteoles’ in Atripliceae (Chenopodiaceae). – Ann. Bot. 108: 847-865.Jay M, Lebreton P. 1973. Recherches chimiotaxinomiques sur les plantes vasculaires XXVI. Les flavanoides des Sarracéniacées, Nepenthacées, Droséracées, et Céphalotacées; étude critique de l’ordre des Sarracéniales. – Nat. Canadien 91: 607-613.

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