Takhtajan Armen. Flowering Plants

Подождите немного. Документ загружается.

142 Subclass IV. CARYOPHYLLIDAE

Cyatho phylla, Bolanthus, Ankyropetalum, Phrynella,

Dianthus, Kohlrauschia, Petrorhagia, Velezia; dryp-

ideae: Drypis; sileneae: Silene, Melandrium,

Pleconax, Ixoca, Oberna, Petrocoma, Lychnis, Steris,

Gastrolychnis, Petrocoptis, Cucubalus, Agrostemma,

Uebelinia.

The Caryophyllaceae are closely related to the

Molluginaceae and probably have a common origin

with them. They are probably an early branch from the

protophytolaccaceous ancestor, which had no betalain

yet. A special variety of P-type sieve- element plastids

with polygonal central protein crystal is characteristic

of the Caryophyllaceae as well as of Limeum and

Macarthuria in the Molluginaceae (Behnke 1969,

1975b, c, 1976b, c, 1993). Besides, the micromorpho-

logy of epicuticular waxes in the Caryophyllaceae is

very much similar to that of Molluginaceae by the fre-

quent occurrence of irregular platelets and rodlets (Engel

and Barthlott 1988; Barthlott in Behnke and Mabry

1994). According to Eichler (1875) the Caryo phyllaceae-

type of ﬂ ower originated from the Phytolacca-type as a

result of transformation of the outer whorl of stamens

into petals.

19. SIMMONDSIACEAE

van Tieghem 1899. 1/1. Southern California, western

Arizona, northern Baja California.

Simmondsia.

Simmondsiaceae are similar to Caryophyllales in

having normal secondary growth, wood anatomy,

pollen morphology, and seeds without endosperm.

20. AMARANTHACEAE

A. L. de Jussieu 1789 (including Achyranthaceae

Raﬁ nesque 1837, Celosiaceae Martynov 1820, Deerin-

giaceae J. Agardh 1858, Gomphrenaceae Raﬁ nesque

1837, Polycnemaceae Menge 1839). 69/1000 or more.

Widely distributed in tropical, subtropical and warm-

temperate regions. Concentrated in Africa south of the

Sahara, southwestern North America, and Central and

South America.

20.1 POLYCNEMOIDEAE

Embryo annular. Stem with normal secondary growth.

Flowers bisexual, solitary, with bracteoles. —

Polycnemum, Hemichroa, Nitrophila.

20.2 AMARANTHOIDEAE.

Anthers tetrasporangiate and 2-locular. Ovules several or

solitary on a basal or short, free-central placenta. –

celosieae: Deeringia (including Dendroportulaca),

Pleuropetalum, Henonia, Celosia, Hermbstaedtia; ama-

rantheae: Bosea, Chamissoa, Herbstia, Siamosia,

Allmania, Charpentiera, Indobanalia, Lagrezia,

Amaranthus, Digera, Neocentema, Pleuropterantha,

Saltia, Sericostachys, Sericoco mopsis, Sericocoma,

Pseudosericocoma, Kyphocarpa, Centemo psis, Nelsia,

Sericorema, Centema, Erio stylos, Lopriorea, Rosifax,

Leucosphaera, Cyathula, Allmaniopsis, Pupalia,

Marcelliopsis, Daysphaera, Volkensinia, Arthraerua,

Aerva, Polyrhabda, Trichuriella, Nothosaerva,

Nototrichium, Calicorema, Chionothrix, Stilbanthus,

Mechowia, Nyssanthes, Ptilotus, Psilotrichum,

Psilotrichopsis, Achyranthes, Centrostachys, Achyropsis,

Pandiaka.

20.3 GOMPHRENOIDEAE

Anthers disporangiate and 1-locular. Ovules usually

solitary and basal, rarely apical and pendulous. – pseu-

doplantageae: Pseudoplantago; gomphreneae:

Guilleminea, Tidestromia, Froelichia, Froelichiella,

Pfafﬁ a, Alternanthera, Woehleria, Gomphrena,

Pseudogomphrena, Iresine, Orenella, Blutaparon,

Lithophila.

The Amaranthaceae reveal some links with the

Petiveriaceae and Achatocarpaceae. In the relatively

most archaic members of the family, such as

Deeringia and Pleuropetalum, the gynoecium is

even more primitive than in Achatocarpaceae and

Petiveriaceae.

21. CHENOPODIACEAE

Ventenat 1799 (including Atriplicaceae A.L. de Jussieu

1789, Betaceae Burnett 1835, Blitaceae Post et Kuntze

1903, Dysphaniaceae Pax 1927, Salicorniaceae

Martynov 1820, Salsolaceae Menge 1839, Spinaciaceae

Menge 1839). 103/1600. Widely distributed in temper-

ate and subtropical regions, mainly in saline habitats;

particularly well represented around the Mediterranean

and Red Sea, in deserts and semideserts of Irano-

Turanian region, in the edge of the Sahara, and in the

Karoo of South Africa, the deserts of Australia, and the

alkaline areas of the United States of America and

Argentina.

Superorder CARYOPHYLLANAE 143

21.1 CHENOPODIOIDEAE

Embryo annular or less often curved. Leaves well

developed, mostly broad, sometimes narrow and ﬂ eshy

or scalelike. Fruits surrounded by the persistent

perianth or by bracteoles. Perisperm usually present.

– beteae: Acroglochin, Hablitzia, Oreobllton,

Aphanisma, Beta; chenopodieae: Rhagodia,

Holmbergia, Baolia, Chenopodium, Teloxys,

Scleroblitum, Dysphania, Monolepis; atripliceae:

Axyris, Endolepis, Atriplex, Exomis, Archiatriplex,

Microgynoecium, Suckleya, Krascheninnikovia,

Grayia, Ceratocarpus, Spinacia, Zuckia; camphoros-

meae: Kirilowia, Panderia, Camphorosma, Cycloloma,

Chenolea, Bassia, Londesia; sclerolaeneae:

Maireana, Eremophea, Enchylaena, Didymanthus,

Neobassia, Malacocera, Dissocarpus, Babbagia,

Threlkeldia, Sclerolaena, Roycea; corispermeae:

Corispermum, Agriophyllum, Anthochlamys.

21.2 MICROTEOIDEAE

Embryo annular. Annual herbs, sometimes suffrutes-

cent at the base. Flowers in spike or spike-like racemes,

bisexual, actinomorphic. Anthers globose, pollen

grains pantoporate. Style absent or nearly so. Fruit an

achene, muricate to spiny. – Microtea.

21.3 SALICORNIOIDEAE

Embryo annular or curved, rarely straight. Perisperm

present. Stems succulent, articulated, with leaves

reduced to tubercles or scales. Flowers usually in groups

of three sunken into cavities in the axis of spiciform

inﬂ orescences. – halopeplideae: Halopeplis,

Kalidium; salicornieae: Allenrolfea, Halocnemum,

Halostachys, Heterostachys, Pachycornia, Sclerostegia,

Arthrocnemum, Microcnemum, Halosarcia, Tecticornia,

Tegicornia, Sarcocornia, Salicornia.

21.4 SALSOLOIDEAE

Embryo spirally coiled. Perisperm usually lacking.

Leaves mostly linear to ﬁ liform, terete, sometimes

scalelike. Flowers usually 1–3, in the axils of the bracts,

usually bracteolate. – suaedeae: Suaeda, Alexandra,

Bienertia, Borsczowia; salsoleae: Haloxylon,

Sympegma, Noaea, Salsola, Climacoptera, Halothamnus,

Cyathobasis, Anabasis, Halogeton, Micropeplis,

Seldlitzia, Arthrophytum, Iljinia, Nanophyton,

Girgensohnia, Halocharis, Halanthium, Fadenia, Sevada,

Choriptera, Rhaphidophyton, Ofaiston, Cornulaca,

Traganum, Traganopsis, Nucularia,Horaninowia,

Piptoptera, Gamanthus, Halarchon, Petrosimonia,

Halimocnemis, Halotis, Lagenantha.

The Chenopodiaceae stand very close to the

Amaranthaceae with which they have a common

origin.

The American genus Microtea, which is usually

included in the Phytolaccaceae s. 1., has many features

in common with the Chenopodiaceae and often has

been considered a link between these two families.

According to Nowicke (Nowicke in Behnke and Mabry

1994:215), the transfer of Microtea to either

Amaranthaceae or Chenopodiaceae is supported, for

the most part, by pollen data. The grains are pantoporate,

have a thin endexine, and a thin foot layer but thick

tecta, characters common to the Amaranthaceae and

Chenopodiaceae. Moreover, the pollen of Microtea

lacks the unusual endexine (ﬁ brous and netlike under

the aperture) found in many Phytolaccaceae. Recently

Behnke (1993) has shown that the sieve-element plas-

tids of Microtea are characterized by a very broad

peripheral ring of protein ﬁ laments and a single tiny

starch grain. The average diameter of the sieve-element

plastids of Microtea also corresponds to that of the

Chenopo diaceae and negates closer relationships to

the Phytolaccaceae.

Bibliography

Aleshina LA. 1963. Morphology of pollen grains of the genus

Claytonia Cronov. and related genera. Bot. Zhurn. 48: 1191–

1196 (in Russian).

Al-Turku TA, K Swarupanandan and PG Wilson. 2003. Primary

vasculature in Chenopodiaceae: a re-interpretation and

implications for systematics and evolution. Bot. J. Linn. Soc.

143: 337–374.

Anderson EF. 2001. The Cactus Family. Timber Press, Portland.

Applequist WL and DB Pratt. 2005. The Malagasy endemic

Dendroportulaca (Portulacaceae) is referable to Deeringia

(Amaranthaceae): molecular and morphological evidence.

Taxon 54: 681–687.

Applequist WL, WL Wagner, EA Zimmer, and M Nepokroeff.

2006. Molecular evidence resolving the systematic position

of Hectorella (Portulacaceae). Syst. Bot. 31: 310–319.

Applequist WL and RS Wallace. 2000. Phylogeny of the

Madagascarian endemic family Didiereaceae. Plant Syst.

Evol. 221: 157–166.

Applequist WL and RS Wallace. 2001. Phylogeny of the

Portulacaceous cohort based on ndhF sequence data. Syst.

Bot. 26: 406–419.

Applequist WL and RS Wallace. 2002. Deletions in the plastid

trnT-trnL intergenic spacer deﬁ ne clades within Cactaceae

subfamily Cactoideae. Plant Syst. Evol. 231(1–4): 153–162.

144 Subclass IV. CARYOPHYLLIDAE

Applequist WL and RS Wallace. 2003. Expanded circumscrip-

tion of Didiereaceae and its division into three subfamilies.

Adansonia, sér. 3, 25: 13–16.

Bailey DC. 1980. Anomalous growth and vegetative anatomy of

Simmondsia chinensis. Am. J. Bot. 67: 147–161.

Balfour E. 1965. Anomalous secondary thickening in

Chenopodiaceae, Nyctaginaceae, and Amaranthaceae.

Phytomorphology 15: 111–122.

Baranova M. 1980. Comparative-stomatographical study of the

families Buxaceae and Simmondsiaceae. In: SG Zhilin, ed.

Systematics and evolution of the higher plants, pp. 68–75.

Nauka, Leningrad (in Russian).

Barthlott W. 1988. Über die systematischen Gliederungen der

Cactaceae. Beitr. Biol. Pﬂ . 63: 17–40.

Barthlott W. 1994. Epicuticular wax ultrastructure and systemat-

ics. In: H-D Behnke and TJ Mabry, eds. Caryophyllales.

Evolution and systematics, pp.75–86. Springer, Berlin

Barthlott W and DR Hunt. 1993. Cactaceae. In: K Kubitzki, ed.

The families and genera of vascular plants, vol. 2, pp. 161–

197. Springer, Berlin/Heidelberg/New York.

Barthlott W and DR Hunt. 2000. Seed-diversity in the Cactaceae,

subfamily Cactoideae. Succ. Plant Res. 5: 1–173

Barthlott W and G Voit. 1979. Mikromorphologie der

Samenschalen und Taxonomie der Cactaceae. Plant Syst.

Evol. 132:205–229.

Bedell HG. 1980. A taxonomic and morphological reevaluation of

Stegnospermataceae (Caryophyllales). Syst. Bot. 5: 419–431.

Behnke H-D. 1969. Über Siebröhren-Plastiden und

Plastidenﬁ lamente der Caryophyllales. Untersuchungen zum

Feinbau und zur Verbreitun eines weiteren speziﬁ schen

Plastidentyps. Planta 89: 275–283.

Behnke H-D. 1974. Elektronenmikroskopische Unter-suchungen

an Siebrohren-Plastiden und ihre Aussage über die system-

atische Stellung von Lophiocarpus. Bot. Jahrb. Syst.

94:114–119.

Behnke H-D. 1975. Hectorella caespitosa: Ultrastructural evi-

dence against its inclusion into Caryophyllaceae. Plant Syst.

Evol. 124: 31–34.

Behnke H-D. 1976a. A tabulated survey of some characters of

systematic importance in centrospermous families. Plant

Syst. Evol. 125: 95–98.

Behnke H-D. 1976b. Ultrastructure of sieve-element plastids in

Caryophyllales (Centrospermae), evidence for the delimitation

and classiﬁ cation of the order. Plant Syst. Evol. 126: 31–54.

Behnke H-D. 1976c. Sieve element plastids of Fouquieria,

Frankenia (Tamaricales), and Rhabdodendron (Rutaceae),

taxa sometimes allied with Centrospermae (Caryophylales).

Taxon 25: 265–268.

Behnke H-D. 1976d. Die Siebelement-Plastiden der

Caryophyllaceae: Eine weitere speziﬁ sche Form der P-Typ-

Plastiden bei Centrospermen. Bot. Jahrb. Syst. 95: 327–333.

Behnke H-D. 1977a. Regular occurring massive deposits of

phytoferritin in the phloem of succulent Centrospermae.

Z. Pﬂ anzenphysiol. 85: 89–92.

Behnke H-D. 1977b. Zur Skulptur der Pollen-Exine bei drei

Centrospermen (Gisekia, Limeum, Hectorella) bei

Gyrostemonaceen, und Rhabdodendraceen. Plant Syst. Evol.

128: 227–235.

Behnke H-D. 1977c. Transmission electron microscopy and sys-

tematics of ﬂ owering plants. Plant Syst. Evol., Suppl. 1:

155–178.

Behnke H-D. 1978. Elektronenoptische Untersuchungen am

Phloem sukkulenter Centrospermen (incl. Didiereaceen).

Bot. Jahrb. Syst. 99: 341–352.

Behnke H-D. 1981. Sieve-element characters. Nord. J. Bot. 1:

381–400.

Behnke H-D. 1982. Geocarpon minimum: Sieve-element plas-

tids as additional evidence for its inclusion in the

Caryophyllaceae. Taxon 31: 45–47.

Behnke H-D. 1993. Further studies of the sieve-element plastids

of the Caryophyllales including Barbeuia, Corrigiola,

Lyallia, Microtea, Sarcobatus, and Telephium. Plant Syst.

Evol. 186: 231–243.

Behnke H-D. 1994. Sieve-element plastids: their signiﬁ cance

for the evolution and systematics of the order. In: H-D

Behnke and TJ Mabry, eds. Evolution and systematics

of the Caryophyllales, pp. 87–121. Springer, Berlin/

New York.

Behnke H-D. 1997. Sarcobataceae — a new family of

Caryophyllales. Taxon 46: 495–507.

Behnke H-D and W Barthlott. 1983. New evidence from ultra-

structural and micromorphological ﬁ elds in angiosperm clas-

siﬁ cation. Nord. J. Bot. 3: 43–66.

Behnke H-D and BL Turner. 1971. On speciﬁ c sieve-tube plas-

tids in Caryophyllales: Further investigation with special ref-

erence to the Bataceae. Taxon 20: 731–737.

Behnke H-D, C Chang, IJ Eifert, and TJ Mabry. 1974. Betalains

and P-type sieve-tube plastids in Petiveria and Agdestis

(Phytolaccaceae). Taxon 23: 541–542.

Behnke H-D, TJ Mabry, IJ Eifert, and L Pop. 1975. P-type sieve-

element plastids and betalains in Portulacaceae (including

Ceraria, Portulacaria, Talinella). Canad. J. Bot. 53:

2103–2109.

Behnke H-D, TJ Mabry, P Neuman, and W Barthlott. 1983a.

Ultrastructural, micromorphological, and phytochemical

evidence for a “central position” of Macarthuria

(Molluginaceae) within the Caryophyllales. Plant Syst. Evol.

143: 151–161.

Behnke H-D, L Pop, and VV Sivarajan. 1983b. Sieve-element

plastids of Caryophyllales: Additional investigations with

special reference to the Caryophyl-laceae and Molluginaceae.

Plant Syst. Evol. 142: 109–115.

Bisalputra T. 1960. Anatomical and morphological studies in the

Chenopodiaceae: I. Inﬂ orescence of Atriplex and Bassia.

Austral. J. Bot. 8: 226–242.

Bisalputra T. 1961. Anatomical and morphological studies in the

Chenopodiaceae: II. Vascularization of the seedlings.

Austral. J. Bot. 9: 1–19.

Bisalputra T. 1962. Anatomical and morphological studies in the

Chenopodiaceae: III. The primary vascular system and nodal

anatomy. Austral. J. Bot. 10: 13–24.

Bittrich V. 1990. Systematic studies in Aizoaceae. Mitt. Inst.

Allg. Bot. Hamb. 23: 491–507.

Bittrich V. 1993. Achatocarpaceae, Caryophyllaceae,

Halophytaceae. In: K Kubitzki, ed. The families and genera

of vascular plants, vol. 2, pp. 35–36, 206–236, 320–321.

Springer, Berlin/Heidelberg/New York.

Bittrich V and HEK Hartmann. 1988. The Aizoaceae: A new

approach. Bot. J. Linn. Soc. 97: 239–254.

Bittrich V and U Kühn. 1993. Nyctaginaceae. In: K Kubitzki,

ed. The families and genera of vascular plants, vol. 2,

pp. 473–486. Springer, Berlin/Heidelberg/New York.

Superorder CARYOPHYLLANAE 145

Bittrich V and M Struck. 1989. What is primitive in

Mesembryanthemaceae? An analysis of evolutionary polar-

ity of character states. South Afr. J. Bot. 55: 321–331.

Blackwell WH. 1977. The subfamilies of the Chenopodiaceae.

Taxon 26: 395–397.

Blunden G, MH Yang, G Janicsak, I Mathe, and A Carabot-

Guervo. 1999. Betaine distribution in the Amaranthaceae.

Biochem. Syst. Evol. 27: 87–92.

Bocquet G. 1959 (1960). The structure of the placental column in

the genus Melandrium (Caryophyllaceae). Phytomorphology

9: 217–221.

Bogle AL. 1969. The genera of Portulacaceae and Ba-sellaceae

in the southeastern United States. J. Arnold Arbor. 50:

566–598.

Bogle AL. 1970. The genera of Molluginaceae and Aizoaceae in

the southeastern United States. J. Arnold Arbor. 51: 431–462.

Bogle AL. 1974. The genera of Nyctaginaceae in the southeast-

ern United States. J. Arnold Arbor. 55: 1–37.

Bogle AL et al. 1971. Geocarpon: Aizoaceae or Caryophyl-

laceae? Taxon 20: 473—477.

Boke NH. 1963. Anatomy and development of the ﬂ ower and

fruit of Pereskia pititache. Am. J. Bot. 50: 843–858.

Boke NH. 1964. The cactus gynoecium: A new interpretation.

Am. J. Bot. 51: 598–610.

Boke NH. 1966. Ontogeny and structure of the ﬂ ower and fruit

of Pereskia aculeata. Am. J. Bot. 53: 534–542.

Boke NH. 1980. Developmental morphology and anatomy in

Cactaceae. Bioscience 30: 605–610.

Borsch T. 1998. Pollen types in the Amaranthaceae: morphol-

ogy and evolutionary signiﬁ cance. Grana 37(3): 129–142.

Bortenschlager S. 1973. Morphologic pollinique des

Phytolaccaceae. Pollen Spores 15: 227–253.

Bortenschlager S, A Auimger, J Blaha, and P Simons-burger.

1972. Pollen morphology of Achatocarpaceae (Centrospermae).

Bot. Naturwiss.-Med. Vereins Innsbruck 59: 7–13.

Brown GK and GS Varadarajan. 1985. Studies in Caryophyllales:

I. Reevaluation of classiﬁ cation of Phytolaccaceae s. 1. Syst.

Bot. 10: 49–63.

Burret F, Z Rabesa, P Zandonella, and B Viorin. 1981.

Contribution biochimique a la systemadque de 1’ordre des

Centrospermales. Biochem. Syst. Ecol. 9: 257–262.

Butnik AA. 1991. Chenopodiaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 3, pp. 77–82. Nauka,

Leningrad (in Russian).

Butnik AA and UN Zhapakova. 1991. Amaranthaceae. In:

A Takhtajan, ed. Comparative seed anatomy, vol. 3, pp.

74–77. Nauka, Leningrad (in Russian).

Butterworth CA. 2006. Molecular phylogenetics of Cactaceae

Jussieu – a review. In: AK Sharma and A Sharma, eds. Plant

Genome Biodiversity and Evolution, vol. 1, pt C: 489–524.

Science Publishers, Enﬁ eld, NH.

Butterworth CA and RS Wallace. 2005. Molecular phylogenet-

ics of the leafy cactus genus Pereskia (Cactaceae). Syst. Bot.

30: 800–808.

Buxbaum F. 1944. Untersuchungen zur Morphologic der

Kakteenblüte: I. Das Gynoeceum. Bot. Arch. 45:190–247.

Buxbaum F. 1948. Zur Klärung der phylogenetischen Stellung

der Aizoaceae und Cactaceae im Pﬂ anzenreich. Jahrb.

Schweiz. Kakt. Ges. 2: 3–16.

Buxbaum F. 1949. Vorläufer des Kakteen-Habitus bei den

Phytolaccaceen. Oesterr. Bot. Z. 96: 5–14.

Buxbaum F. 1950, 1954, 1955. Morphology of cacti., vols. 1–3.

Abbey Garden Press, Pasadena, CA.

Buxbaum F. 1958. The phylogenetic division of the subfamily

Cereoideae, Cactaceae. Madroño 14: 177–206.

Buxbaum F. 1961. Vorläuﬁ ge Untersuchungen über Um-fang,

systematische Stellung, und Gliederung der Caryophyllales

(Centrospermae). Beitr. Biol. Pﬂ . 36: 1–56.

Buxbaum F. 1962. Stellung der Kakteen im Pﬂ anzenreich. Kakt.

und Sukk. 13: 194–197.

Buchmann SL. 1987. Floral biology of jojoba (Simmondsia chin-

ensis), an anemophilous plant. Desert Plants 8: 111–124.

Carlquist S. 1995. Wood anatomy of Caryophyllaceae: ecologi-

cal, habitual, systematic, and phylogenetic implications.

Aliso 14: 1–17.

Carlquist S. 1998a. Wood anatomy of Portulacaceae and

Hectorellaceae: ecological, habitual, and systematic implica-

tions. Aliso 16: 137–153.

Carlquist S. 1998b. Wood and stem anatomy of Petiveria and

Rivina (Caryophyllales): systematic implications. IAWA J.

19: 383–391.

Carlquist S. 1999a. Wood, stem, and root anatomy of Basellaceae,

with a relation to systematics and cambial variants. Flora

194: 1–12.

Carlquist S. 1999b. Wood anatomy of Agdestis (Caryophyllales):

systematic position and nature of the successive cambia.

Aliso 18: 35–43.

Carlquist S. 1999c. Wood and stem anatomy of Stegnosperma

(Caryophyllales): phylogenetic relationships; nature of lat-

eral meristems and successive cambial activity. IAWA J. 20:

149–163.

Carlquist S. 1999d. Wood anatomy, stem anatomy, and cambial

activity of Barbeuia (Caryophyllales). IAWA J. 20: 431–440.

Carlquist S. 2000a. Wood and stem anatomy of Sarcobatus

(Caryophyllales): Systematic and ecological implications.

Taxon 49: 27–34.

Carlquist S. 2000b. Wood and stem anatomy of phytolaccoid

and rivinoid Phytolaccaceae (Caryophyllales): ecology, sys-

tematics, nature of successive cambia. Aliso 19: 13–29.

Carlquist S. 2000c. Wood and bark anatomy of Achatocarpaceae.

SIDA 19: 71–78.

Carlquist S. 2001. Wood and stem anatomy of Rhabdodendraceae

is consistent with placement in Caryophyllales sensu lato.

IAWA J. 22: 171–18.

Carlquist S. 2002. Wood anatomy and successive cambia in

Simmondsia (Simmondsiaceae): Evidence for inclusion in

Caryophyllales s.l. Madroño 49: 158–164.

Carlquist S. 2003. Wood and stem anatomy of woody

Amaranthaceae s.s.: ecology, systematics and the problems of

deﬁ ning rays in dicotyledons. Bot. J. Linn. Soc. 143: 1–19.

Carlquist S. 2005. Lateral meristems, successive cambia and

their products: a reinterpretation based on roots and stems of

Nyctaginaceae. Bot. J. Linn. Soc. 146: 129–143.

Carlquist S. 2007. Successive cambia in Aizoaceae: products

and process. Bot. J. Linn. Soc. 153: 141–155.

Carolin RC. 1983. The trichomes of the Chenopodiaceae and

Amaranthaceae. Bot. Jahrb. Syst. 103: 451–466.

Carolin RC. 1987. A review of the family Portulacaceae. Austral.

J. Bot. 35: 383–412.

Carolin RC. 1993. Portulacaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 2, pp. 544–555.

Springer, Berlin/Heidelberg/New York.

146 Subclass IV. CARYOPHYLLIDAE

Carolin RC, SWL Jacobs, and M Vesk. 1975. Leaf structure in

Chenopodiaceae. Bot. Jahrb. Syst. 95: 226–255.

Chesselet P, Mössmer, and GF Smith. 1995. Research priorities

in the succulent plant family Mesembryanthaceae Fenzl.

South Afr. J. Sci. 91: 192–209.

Chesselet P, GF Smith, PM Burgoyne, C Klak, SA Hammer,

HEK Hartmann, H Kurzweil, EJ van Jaarsveld, BE van Wyk

and OA Leistner. 2000. Mesembryanthemaceae. In: Leistner

OA eds. Seed plants of South Africa: families and genera.

Srelitzia 10: 360–410.

Chesselet P, GF Smith and AE van Wyk. 2002. A new tribal

classiﬁ cation of Mesebryanthemaceae: evidence from ﬂ oral

nectaries. Taxon 51: 295–308.

Chesselet P, AE van Wyk, and GF Smith. 2004.

Mesembryanthemaceae. A new tribe and adjustments to inf-

rafamilial classiﬁ cation. Bothalia 31: 47–51.

Chorinsky F. 1931. Vergleichende morphologische

Untersuchungen der Haargebilde bei Portulacaceae and

Cactaceae. Osterr. Bot. Z. 80: 308–327.

Choux P. 1934. Les Didiereacees: Xerophytes de Madagascar.

Mem. Acad. Malgache 18: 1–69.

Chuong SDX, VR Franceschi, and GE Edwards. 2006. The

cytoskeleton maintains organelle partitioning required for

single cell C

photosynthesis in Chenopodiaceae species.

Plant Cell 18: 2207–2223.

Clement JS and TJ Mabry. 1996. Pigment evolution in the

Caryophyllales: a systematic overview. Bot. Acta 109:

360–367.

Costea M and DA DeMason. 2001. Stem morphology and ana-

tomy in Amaranthus L. (Amaranthaceae) – taxonomic sig-

niﬁ cance. J. Torr. Bot. Soc. 128: 254–281.

Cranwell LM. 1963. The Hectorellaceae: Pollen type and taxo-

nomic speculation. Grana Palynol. 4: 195–202.

Cronquist A and RE Thorne. 1994. Nomenclatural and taxo-

nomic history. In: H-D Behnke and TJ Mabry, eds.

Caryophyllales. Evolution and systematics, pp. 5–25.

Springer, Berlin/Heidelberg.

Cuénoud Ph. 2002. Introduction to expanded Caryophyllales. In:

K Kubitzki, ed. The Families and genera of vascular plants,

vol. 5, pp. 1–4. Springer, Berlin/Heidelberg/New York.

Cuénoud P. 2006. Phylogeny, evolution and diversiﬁ cation of

Caryophyllales. In: AK Sharma and A Sharma, eds. Plant

genome biodiversity and evolution, vol. 1, pp. 187–218.

Science Publishers, Enﬁ eld, NH.

Cuénoud Ph, V Savolainen, LW Chatrou, M Powell, RJ Grayer,

and M Chase. W. 2002. Molecular phylogenetics of

Caryophyllales based on nuclear 18S rDNA and plastid

rbcL, atpB, and matK DNA sequences. Am. J. Bot. 89:

132–144.

Daxenbichler M, GF Spencer, DG Carlson, GB Rose, AM

Brinker, and RG Powell. 1991. Glucosilonate composition of

seeds from 297 species of wild plants. Phytochemistry 30:

2623–2638.

Dickie SL. 1996. Phylogeny and evolution in the subfamily

Opuntioideae (Cactaceae): insights from rp116 intron

sequence variation. M.S. Thesis. Iowa State University.

Di Fulvio TE. 1975. Estomatogenesis en Halophytum ameghi-

noi (Halophytaceae). Kurtziana 8: 17–29.

Dinan L, P Whiting, and AJ Scott. 1998. Taxonomic distribution

of phytoecdysteroids in seed of members of the

Chenopodiaceae. Biochem. Syst. Ecol. 26: 553–576.

Douglas NA and PS Manos. 2007. Molecular phylogeny of

Nyctaginaceae: taxonomy, biogeography, and characters

associated with a radiation of xerophytic genera in North

America. Am. J. Bot. 94: 856–872.

Doweld AB. 2001. Conspectus systematicus tribus Cacteae

(Cactaceae). Genera et taxa supragenerica, 2. Novosti Sist.

Vyssh. Rast. 33: 163–171 (in Russian).

Downie SR and JD Palmer. 1994. A chloroplast DNA phylogeny

of the Caryophyllales based on structural and inverted repeat

restriction site variation. Syst. Bot. 19: 236–252.

Downie SR, DS Katz-Downie, and KJ Cho. 1997. Relationships

in the Caryophyllales as suggested by phylogenetic analyses

of partial chloroplast DNA ORF2280 homolog sequences.

Am. J. Bot. 84: 253–273.

Eckardt Th. 1954. Morphologische und systematische

Auswertung der Placentation von Phytolaccaceen. Ber.

Deutsch. Bot. Ges. 67: 113–128.

Eckardt Th. 1967a. Vergleich von Dysphania mit Chenopodium

und mit Illecebraceae. Bauhinia 3: 327–344.

Eckardt Th. 1967b. Blütenbau und Blültenentwicklung von

Dysphania myriocephala Benth. Bot. Jahrb. Syst. 86: 20–37.

Eckardt Th. 1967c (1968). Blütenmorphologie von Dysphania

planta ginella F. v. M. Phytomorphology 17: 165–172.

Eckardt Th. 1974. Vom Blütenbau der Centrospermen-Gattung

Lophiocarpus Turcz. Phyton (Horn) 16: 13–27.

Eckardt Th. 1976. Classical morphological features of centro-

spermous families. Plant Syst. Evol. 126: 5–25.

Edwards EJ. 2006. Correlated evolution of stem and leaf hydrau-

lic traits in Pereskia (Cactaceae). New Phytol. 171.

Edwards EJ and M Diaz. 2006. Ecological physiology of

Persekia guamacho, a cactus with leaves. Plant, Cell Environ.

29: 247–256.

Edwards EJ and MJ Donoghue. 2006. Pereskia and the origin of

the cactus life form. Am. Naturalist 167: 777–793.

Edwards EJ, R Nyffeler, and MJ Donoghue. 2005. Basal cacus

phylogeny: implications of Pereskia (Cactaceae) paraphyly

for the transition to the cactus life form. Am. J. Bot. 92:

1177–1188.

Ehrendorfer F. 1976a. Chromosome numbers and differentiation

of centrospermous families. Plant Syst. Evol. 126: 27–30.

Ehrendorfer F. 1976b. Closing remarks: Systematics and evolu-

tion of centrospermous families. Plant Syst. Evol. 126:

99–105.

Eliasson UH. 1988. Floral morphology and taxonomic relations

among the genera of Amaranthaceae in the New World and

the Hawaiian Islands. Bot. J. Linn. Soc. 96: 235–283.

Endress ME and V Bittrich. 1993. Molluginaceae. In:

K Kubitzki, ed. The families and genera of vascular plants,

vol. 2, pp. 419–426. Springer, Berlin/Heidelberg/New York.

Engel T and W Barthlott. 1988. Micromorphology of epicuticu-

lar waxes in Centrosperms. Plant Syst. Evol. 161: 71–85.

Erbar C. 1994. Putative origin and relationships of the order

from the viewpoint of developmental ﬂ ower morphology. In:

H-D Behnke, TJ Mabry, eds. Caryophyllales. Evolution and

systematics, pp. 3–316. Springer, Berlin/New York.

Erbar C and P Leins. 2006. Floral ontogeny and systematic posi-

tion of the Didiereaceae. Plant Syst. Evol. 261: 165–185.

Erdtman G. 1948. Pollen morphology and plant taxonomy: VIII.

Didiereaceae. Bull. Mus. Hist. Nat., Paris, ser. 2, 20: 387–394.

Esau K and VI Cheadle. 1969. Secondary growth in

Bougainvillea. Ann. Bot. 33: 807–819.

Superorder CARYOPHYLLANAE 147

Fahn A and T Arzee. 1959. Vascularization of articulated

Chenopodiaceae and the nature of their ﬂ eshy cortex. Am. J.

Bot. 46: 330–338.

Fahn A and Y Shchori. 1967 (1968). The organization of the

secondary conducting tissues in some species of the

Chenopodiaceae. Phytomorphology 17: 147–154.

Fay MF, KM Cameron, GT Prance, MD Lledo, and MW Chase.

1997. Familial relationships of Rhabdodendron

(Rhabdodendraceae): plastid rbcL sequences indicate a cary-

ophyllid placement. Kew Bull. 52: 923–932.

Fedotova TA. 1991. Hectorellaceae, Basellaceae, Halophytaceae.

In: A Takhtajan, ed. Comparative seed anatomy, vol. 3,

pp. 33–41. Nauka, Leningrad (in Russian).

Fior S, PO Karis, G Casazza, L Minuto, and F Sala. 2006.

Molecular phylogeny of the Caryophyllaceae (Caryophyllales)

inferred from chloroplast matK and nuclear rDNA ITS

sequences. Am. J. Bot. 93: 399–411.

Franz E. 1908. Beiträge zur Kenntnis der Portulacaceen und

Basellaceen. Bot. Jahrb. 42, Beibl. 97: 1–46.

Gail PA. 1964. Simmondsia chinensis (Link) Schneider: ana-

tomy and morphology of ﬂ owers. M.A. Thesis. Clermont.

Garcia M, M Lapp, M Castro, and P Torrecilla. 2000. Anatomia

foliar comparada de cuatro especies del genero Pereskia

(Plum.) Miller (Cactaceae). Ernstia 10: 27–41.

Gentry HS. 1958. The natural history of jojoba (Simmondsia

chinensis) and its cultural aspects. Econ. Bot. 12: 261–295.

Gibson AC. 1977. Comparative anatomy of secondary xylem in

Cactoideae (Cactaceae). Biotropica 5 (I): 29–65.

Gibson AC. 1977b. Vegetative anatomy of Maihuenia

(Cactaceae) with some theoretical discussions of ontogenetic

changes in xylem cell types. Bull. Torrey Bot. Club 104:

35–48.

Gibson AC. 1978. Rayless secondary xylem of Halophytum.

Bull. Torrey Bot. Club 105: 39–44.

Gibson AC and KE Horak. 1979. Systematic anatomy and phy-

logeny of Mexican columnar cacti. Ann. Missouri Bot. Gard.

65: 999–1057.

Gibson AC and PS Nobel. 1986. The cactus primer. Harvard

University Press, Cambridge, MA.

Gilbert MG. 1987. The taxonomic position of the genera

Telephium and Corrigiola. Taxon 36: 47–49.

Gilbert MG. 1993. A review of Gisekia (Gisekiaceae). Kew

Bull. 48: 343–356.

Gowik U, S Engelmann, OE Bläsing, AS Raghavendra, and

P Westhoff. 2006. Evolution of C

phosphoenolpyruvate

carboxylase in the genus Alternanthera: Gene families and

the enzymatic characteristics of the C

isozyme and its

orthologues in C

and C

alternanthera. Planta 359–368.

Grifﬁ th MP. 2002. Phylogenetic relationships in the Opuntioideae

(Cactaceae) based on nrITS sequences. In Botany 2002:

Botany in the Curriculum, p.126. Madison, WisconSinica

(Abstracts).

Guaralnick LJ and MD Jackson. 2001. The occurrence and phy-

logenetics of Crassulacean Acid Metabolism in the

Portulacaceae. Int. J. Plant Sci. 162: 257–262.

Gvinianidze ZI and TA Fedotova. 1991. Caryophyllaceae. In:

A Takhtajan, ed. Comparative seed anatomy, vol. 3,

pp. 59–74. Nauka, Leningrad (in Russian).

Hartmann HEK. 1993. Aizoaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 2, pp. 37–69. Springer,

Berlin/Heidelberg/New York.

Hassan NMS, U Meve, and S Liede-Schumann. 2005. Seed coat

morphology of Aizoaceae-Sesuvioideae, Gisekiaceae and

Molluginaceae and its systematic signiﬁ cance. Bot. J. Linn.

Soc. 148: 189–206.

Hassan NMS, J Thiede, and S Liede-Schumann. 2005.

Phylogenetic analysis of Sesuvioideae (Aizoaceae) inferred

from nrDNA internal transcribed spacer (ITS) sequences and

morphological data. Plant Syst. Evol. 255: 121–143.

Haug I, M Weiß, J Homeier, F Oberwinkler, and I Kottke. 2005.

Russulaceae and Thelephoraceae form ectomycorrhizas with

members of Nyctaginaceae (Caryophyllales) in the tropical

mountain rain forest of southern Ecuador. New Phytol. 165:

923–936.

Hershkovitz MA. 1990. Phylogenetic and morphological studies

in Portulacaceae. Ph.D. dissertration, University of

California, Davis.

Hershkovitz MA. 1993. Revised circumscriptions and subge-

neric taxonomies of Calandrinia and Montiopsis

(Portulacaceae) with notes on phylogeny of the portulaca-

ceous alliance. Ann. Missouri Bot. Gard. 80: 333–365.

Hershkovitz MA. 2006. Ribosomal and chloroplast DNA evi-

dence for diversiﬁ cation of western American Portulacaceae

in the Andean region. Gayana Bot. 63: 13–74.

Hershkovitz MA and EA Zimmer. 1997. On the evolutionary

origins of the cacti. Taxon 46: 217–232.

Heubl GR, G Bringmann, and H Meimberg. 2006. Molecular

phylogeny and character evolution of carnivorous plant

families in Caryophyllales – revisited. Plant Biol. 8: 821–830.

Hofmann U. 1973. Centrospermen-Studien: 6. Morphologische

Untersuchungen zur Umgrenzung und Gliederung der

Aizoaceen. Bot. Jahrb. Syst. 93: 247–324.

Hofmann U. 1977. Centrospermen-Studien: 9. Die Stellung von

Stegnosperma innerhalb der Centrospermen. Ber. Deutsch.

Bot. Ges. 90: 39–52.

Horak KE. 1981. Anomalous secondary thickening in

Stegnosperma (Phytolaccaceae). Bull. Torrey Bot. Club

108:189–197.

Hu Z-H and P-J Yang. 1944. Comparative anatomy of anoma-

lous structures in the axes of 44 species of Chenopodiaceae.

Cathaya 6: 145–162.

Hunt D and N Taylor, eds. 1990. The genera of Cactaceae:

Progress towards consensus. Report of the IOS Working

Party, 1987–1990. Bradleya 8: 85–107.

Hunziker JH, H-D Behnke, IJ Eifert, and TJ Mabry. 1974.

Halophytum ameghinoi: A betalain-containing and P-type

sieve-tube species. Taxon 23: 537–539.

Hunziker JH, R Pozner, and A Escobar. 2000. Chromosome

number in Halophytum ameghinoi (Halophytaceae). Plant

Syst. Evol. 221: 125–127.

Ihlenfeldt H-D. 1975. Some trends in the evolution of the

Mesembryanthemaceae. Boissiera 24: 249–254.

Ihlenfeldt H-D, G Schwantes, and H Straka. 1962. Die hoheren

Taxa der Mesembryanthemaceae. Taxon 11: 52–56.

Ihlenfeldt H-D and H Straka. 1961 (1962). Über die system-

atische Stellung und Gliederung der Mesembryanthemen.

Ber. Deutsch. Bot. Ges. 74: 485–492.

Inamdar JA. 1968. Epidermal structure and ontogeny of stomata

in some Nyctaginaceae. Flora 158: 159–166.

Inamdar JA, M Gagadhara, PG Morge, and RM Patel. 1977.

Epidermal structure and ontogeny of stomata in some

Centrospermae. Feddes Repert. 88: 465–475.

148 Subclass IV. CARYOPHYLLIDAE

Jacobs SWL. 2001. Review of leaf anatomy and ultrastructure in

the Chenopodiaceae (Caryophyllales). J. Torrey Bot. Soc.

128: 236–253.

Jansen S, LP Ronse Decraene, and E Smets. 2000. On the wood

and stem anatomy of Monococcus echinophorus

(Phytolaccaceae s.l.). Syst. Geogr. Plants 70: 171–179.

Jensen U. 1965. Serologische Untersuchungen zur Frageder sys-

tematischen Einordnung der Didiereaceae. Bot. Jahrb. Syst.

84: 233–253.

Jia A-Q, N-H Tan, Y-P Yang, S-G Wu, L-Q Wang, and J Zhou.

2004. Cyclopeptides from three arctic Caryophyllaceae

plants, chemotaxonomy and distribution signiﬁ cance of

Caryophyllaceae cyclopeptides. Acta Bot. Sinica 46: 625–630.

Joshi AC. 1937. Some salient points in the evolution of the

secondary vascular cylinder of Amaranthaceae and

Chenopodiaceae. Am. J. Bot. 24: 3–9.

Joshi AC and VSR Rao. 1934. Vascular anatomy of ﬂ owers of

four Nyctaginaceae. J. Indian Bot. Soc. 13: 169–186.

Joshi AC and VSR Rao. 1936. The embryology of Gisekia phar-

naceoides L. Proc. Indian Acad. Sci. 3B: 71–92.

Judd WS and IK Ferguson. 1999. The genera of Chenopodiaceae

in southeastern United States. Harward Pap. Bot. 4: 365–416.

Kadereit G, T Borsch, K Weising, and H Freitag. 2003. Phylogeny

of Amaranthaceae and Chenopodiaceae and the evolution of

photosynthesis. Int. J. Plant Sci. 164: 959–986.

Kadereit G, D Gotzek, S Jacobs, and H Freitag. 2005. Origin and

age of Australian Chenopodiaceae. Organisms Divers. Evol.

5: 59–80.

Kadereit G, L Mucina, and H Freitag. 2006. Phylogeny of

Salicornioideae (Chenopodiaceae): Diversiﬁ cation, biogeo-

graphy, and evolutionary trends in ﬂ oral morphology. Taxon

55: 617–642.

Kajale LB. 1954. Contribution to the embryology of the

Phytolaccaceae: II. Fertilization and development of embryo,

seed, and fruit in Rivina humilis Linn. And Phytolacca dio-

ica Linn. J. Indian Bot. Soc. 33: 206–225.

Kakkar L. 1973. Leaf vasculature in Simmondsia californica.

Indian Sci. Congr. Assoc., Chandigarh. Proc. 60th Sess. Part

III: 293–294 (Abstract).

Kamelina OP. 2001. Development of embryonic structures

of Hablitzia (Chenopodiaceae). Bot. Zhurn. 86(10): 1–9

(in Russian with English summary).

Kamelina OP and OB Proskurina. 1985. On the embryology of

Stegnosperma halimifolium (Stegnospermataceae). Bot.

Zhurn. 70: 721–730 (in Russian with English summary).

Kapralov MV, H Akhani, EV Voznesenskaya, G Edwards, V

Franceschi, and EH Ropalson. 2006. Phylogenetic relation-

ships in the Salicornioideae/Suaedoideae/Salsoloideae s.l.

(Chenopodiaceae) clade and a clariﬁ cation of the phyloge-

netic position of Bienertia and Alexandra using multiple

DNA sequence datasets. Syst. Bot. 31: 572–585.

Kapranova NN. 1991. Aizoaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 3, pp. 17–22. Nauka,

Leningrad (in Russian).

Klak C, A Khunou, G Reeves and T Hedderson. 2003. A phylo-

genetic hypothesis for the Aizoaceae (Caryophyllales) based

on four plastid DNA regions. Am. J. Bot. 90: 1433–1445.

Köhler E. 1993. Blattnervatur-Muster der Buxaceae Dumortier

und Simmondsiaceae van Tieghem. Feddes Repert. 104:

145–167.

Köhler E. 2003. Simmondsiaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 5, pp. 355–358.

Springer, Berlin/Heidelberg/New York.

Köhler E and R Brückner. 1983. Zur Pollenmorphologie und

systematischen Stellung der Gattung Simmondsia Nutt.

Wiss. Z. Friedrich-Schiller-Univ. Jena, Math. Naturwiss.

Reihe, 32: 945–955.

Komar GL. 1991. Nyctaginaceae. In: A. Takhtajan, ed.

Comparative seed anatomy, vol. 3, pp. 12–17. Nauka,

Leningrad (in Russian).

Kondorskaya VR. 1983. On the inﬂ orescences of the genus

Chenopodium L. Bull. Moscow Soc. Nat., Biology, 88:

78–86 (in Russian).

Kondorskaya VR. 1984. Peculiarities in the structure of inﬂ ores-

cences of the tribe Atripliceae C. A. Mey. of the family

Chenopodiaceae. Bull. Moscow Soc. Nat., Biology, 89: 104–

114 (in Russian).

Kowal T. 1961. Studia nad morfologia i anatomia nasion

Portulacaceae Rchb. Monogr. Bot. 12: 1–48.

Kozhanchikov VI. 1967. Morphological characters of the seeds

of the family Caryophyllaceae and probable ways of their

evolution. Bot. Zhurn. 52: 1277–1280.

Kozhanchikov VI. 1975. Variation of morphological features of

seeds of the representatives of the family Caryophyllaceae

Juss. In: LY Budantsev, ed. Questions of comparative

morphology of seed plants, pp. 108–128. Nauka, Leningrad

(in Russian).

Kravtsova TI. 1991. Didiereaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 3, pp. 57–58. Nauka,

Leningrad (in Russian).

Kubitzki K. 1993. Didiereaceae. In: K Kubitzki, ed. The fami-

lies and genera of vascular plants, vol. 2, pp. 292–295.

Springer, Berlin/Heidelberg/New York.

Kühn U, V Bittrich, R Carolin, H Freitag, IC Hedge, P Uotila,

and PG Wilson. 1993. Chenopodiaceae. In: K Kubitzki, ed.

The families and genera of vascular plants, vol. 2, pp. 253–

281. Springer, Berlin/Heidelberg/New York.

Landrum JV. 2001. Wide-band tracheids in leaves of genera in

Aizoaceae: the systematic occurrence of a novel cell type

and its implications for the monophyly of the subfamily

Ruschioideae. Plant Syst. Evol. 227: 49–61.

Landrum JV and JD Mauseth. 1995. Systematic and evolution-

ary implications of stem and leaf anatomy in portulacaceous

genera of South America, North America, and Africa. Am.

J. Bot. 82(6): 176 (Abstract).

Landrum JV and JD Mauseth. 1996. Wide-band tracheids in

genera of the Portulacaceae: Systematic and evolutionary

implications. Am. J. Bot. 83(6): 170 (Abstract).

Larsen K. 2002. Caryophyllaceae. In: HP Nooteboom, ed. Flora

Malesiana, ser.I, 16: 1–51. Leiden.

Lee CW and RA Sherman. 1985. Meiosis in jojoba, Simmondsia

chinensis. Israel J. Bot. 34: 1–6.

Leins P. 1988. Placentation in Cactaceae. In: P Leins, SC Tucker,

and PK Endress, eds. Aspects of Floral Development,

pp. 191–204. Cramer, Berlin.

Leins P and C Erbar. 1994. Putative origin and relationships of

the order Caryophyllales from the viewpoint of developmental

ﬂ ower morphology. In: H-D Behnke and TJ Mabry, eds.

Caryophyllales. Evolution and systematics, pp. 303–316.

Springer, Berlin/New York.

Superorder CARYOPHYLLANAE 149

Leins P and S Schwitalla. 1986. Studien an Cactaceen-Blüten I.

Einige Bemerkungen zur Blütenentwicklung von Pereskia.

Beitr. Biol. Pﬂ anzen 60: 313–323.

Leins P and S Schwitalla. 1988. Placentation in Cactaceae. In: P

Leins, SC Tucker, and PK Endress, eds. Aspects of ﬂ oral

development, pp. 57–68. Cramer, Berlin.

Leins P, A Walter und C Erbar. 2001. Eine morphogenetische

Interpretation der Caryophyllaceen-Kronblätter. Bot. Jahrb.

Syst. 123(3): 355–367.

Leuenberger BE. 1976. Die Pollenmorphologie der Cactaceae

und ihre Bedeutung für die Systematik. Diis. Lot. 31. Vaduz.

Leuenberger BE. 1986. Pereskia (Cactaceae). Mem. New York

Bot. Gard. 41: 1–141.

Leuenberger BE. 1997. Maihuenia: Monograph of a Patagonian

genus of Cactaceae. Bot. Jahrb. Syst. 119: 1–92.

Levin RA. 2000. Phylogenetic relationships within Nyctag-

inaceae tribe Nyctagineae: evidence from nuclear and chlo-

roplast genomes. Syst. Bot. 25: 738–750.

Liden M, Popp M, and B Oxelman. 2001. A revised generic

classiﬁ cation of the tribe Sileneae (Caryophyllaceae). Nord.

J. Bot. 20: 513–518.

Livingstone DA, M Tomlinson, G Friedman, and R Broome.

1973. Stellate pore ornamentation in pollen grains of the

Amaranthaceae. Pollen et Spores 15: 345–351.

Mabry TJ. 1977. The order Centrospermae. Ann. Missouri Bot.

Gard. 64: 210–220.

Mabry TJ, H-D Behnke, and IJ Eifert. 1976. Betalains and

P-type sieve-element plastids in Gisekia L. (Centrospermae).

Taxon 25: 112–114.

Mabry TJ, L Kimler, and C Chang. 1972. The betalains:

Structure, function, and biogenesis and the plant order

Centrospermae. In: VC Runeckles and TC Tso, eds. Recent

Adv. Phytochem. 4: 105–134. New York.

Mabry TJ, P Neumann, and WP Philipson. 1978. Hectorella:

A member of the betalain-suborder Chenopodiineae of the

order Centrospermae. Plant Syst. Evol. 130: 163–165.

Mauseth JD. 2004. Wide-band tracheids are present in almost all

species of Cactaceae. J. Plant Res. 117: 69–76.

Mauseth JD, Y Uozumi, BJ Plemons, and JK Landrum. 1995.

Structural and systematic study of an unusual tracheid type

in cacti. J. Plant Res. 108: 517–526.

McNeil J. 1974. Synopsis of a revised classiﬁ cation of the

Portulacaceae. Taxon 23: 725–728.

McNeil J. 1975. A generic revision of Portulacaceae tribe

Montieae using techniques of numerical taxonomy. Canad. J.

Bot. 53: 789–809.

Maheshwari P and RN Chopra. 1955. The structure and devel-

opment of the ovule and seed of Opuntia dillenii Haw.

Phytomorphology 5: 112–122.

Mauritzon J. 1934. Ein Beitrag zur Embryologie der

Phytolaccaceen und Cactaceen. Bot. Not. 1934: 111–135.

Mauseth JD. 1982–1984. Introduction to cactus anatomy, parts

1–11. Cact. Succ. (U.S.) 54: 263–266; 55: 18–21, 42, 84–89,

113–118, 171–175, 272–276; 56: 33–37,131–135, 181–184,

212–216, 250–255.

Melikian AP. 1968. Position of the families Buxaceae and

Simmondsiaceae in the system. Bot. Zhurn. 53: 1043–1047

(in Russian).

Melikian AP. 1991. Phytolaccaceae, Achatocarpaceae,

Barbeuiaceae, Stegnospermataceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 3, pp. 8–11. Nauka,

Leningrad (in Russian).

Melikian AP and AG Devyatov. 2000. Exocarp surface structure

as a taxonomically signiﬁ cant feature in Caryophyllaceae.

Byull. Mosk. Obshch. Ispyt. Prir., Biol. 105(1): 54–58

(in Russian with English summary).

Mikesell JE. 1979. Anomalous secondary thickening in

Phytolacca americana L. (Phytolaccaceae). Am. J. Bot. 66:

997–1005.

Moeliono BM. 1966. De caulomatische Oorsprong van

Zaadknoppen bij Caryophyllaceën en Primulaceën (een

histogenetische en anatomische Analyse). Rotoype,

Amsterdam.

Müller K and T Borsch. 2005a. Phylogenetics of Amaranthaceae

based on matK/trnK sequence data – evidence from parsi-

mony, likelihood, and Bayesian analyses. Ann. Missouri

Bot. Gard. 92: 66–102.

Müller K and T Borsch. 2005b. Multiple origins of a unique pol-

len feature: Stellate pore ornamentation in Amaranthaceae.

Grana 44: 266–281.

Nageshwar G and M Radhakrishnaiah. 1993. Afﬁ nities of

Basella L. Feddes Repert. 104(3–4): 241–244.

Nair NC and VJ Nair. 1961. Studies on the morphology of some

members of the Nyctaginaceae: I. Nodal anatomy of

Boerhavia. Proc. Indian Acad. Sci. 54B: 281–294.

Narayana PS. 1990. The bearing of amino acids on the status

and systematic position of Basellaceae. Indian J. Bot. 13:

129–135.

Narayana PS and LL Narayana. 1986. The embryology of

Stegnospermataceae, with a discussion on its status, afﬁ ni-

ties, and systematic position. Plant Syst. Evol. 154:

137–145.

Narayana PS and LL Narayana. 1988. Systematic position of

Gisekia

L.: A numerical assessment. Feddes Repert. 99

(5–6): 189–193.

Nepokroeff M, WL Wagner, RK Rabeler, EA Zimmer, SG

Weller, and AK Sakai. 2002. Relationships within

Caryophyllaceae inferred from molecular sequence data. In

Botany 2002: Botany in the Curriculum, p.105. Madison,

WisconSinica (Abstracts).

Ng Siew Yoong, WR Philipson, and JRL Walker. 1975.

Hectorellaceae: A member of the Centrospermae. New

Zealand J. Bot. 13: 567–570.

Nowicke JW. 1968. Palynotaxonomic study of the Phytolaccaceae.

Ann. Missouri Bot. Gard. 55: 294–364.

Nowicke JW. 1975. Pollen morphology in the order

Centrospermae. Grana 15: 51–77.

Nowicke JW. 1994. Pollen morphology and exine ultrastructure.

In: H-D Behnke and TJ Mabry, eds. Caryophyllales.

Evolution and systematics, pp. 167–221. Springer, Berlin/

New York.

Nowicke JW. 1996. Pollen morphology, exine structure and the

relationships of Basellaceae and Didiereaceae to

Portulacaceae. Syst. Bot. 21: 187–208.

Nowicke JW and JJ Skvarla. 1984. Pollen morphology and rela-

tionships of Simmondsia chinensis to the order Euphorbiales.

Am.J. Bot. 71: 210–215.

Nyananyo BL. 1988. The systematic signiﬁ cance of seed mor-

phology and anatomy in the Portulacaceae (Centrospermae).

Folia Geobot. Phytotax. 23 (3): 275–279.

150 Subclass IV. CARYOPHYLLIDAE

Nyananyo BL. 1990. Tribal and generic relationship in the

Portulacaceae (Centrospermae). Feddes Repert. 101 (5–6):

237–241.

Nyffeler R. 2002. Phylogenetic relationships in the cactus fam-

ily (Cactaceae) based on evidence from trnK/matK and trnL-

trnF sequences. Am. J. Bot. 89: 312–326.

O’Quinn R and L Hufford. 2005. Molecular systematics of

Montieae (Portulacaceae): Implications for taxonomy, bio-

geography and ecology. Syst. Bot. 30: 314–331.

Oxelman B, M Lidén, RK Rabeler, and M Popp. 2000. A revised

generic classiﬁ cation of the tribe Sileneae (Caryophyllaceae).

Nord. J. Bot. 20: 743–748.

Oxelman B, M Liden, K Rabeler, and M Popp. 2001. A revised

generic classiﬁ cation of the tribe Sileneae (Caryophyllaceae).

Nord. J. Bot. 20: 743–748.

Paliwal GS. 1965. The development of stomata in Basella rubra

Linn. Phytomorphology 15:50–53.

Pant DD and PF Kidwai. 1968. Structure and ontogeny of stomata

in some Caryophyllaceae. Bot. J. Linn. Soc. 60: 309–314.

Patterson GW and S Xu. 1990. Sterol composition in ﬁ ve fami-

lies of the order Caryophyllales. Phytochemistry 29:

3539–3541.

Pax F. 1927. Zur Phylogenie der Caryophyllaceae. Bot. Jahrb.

Syst. 61: 223–241.

Payne MA. 1933. The morphology and anatomy of Mollugo ver-

tuillata L. Univ. Kansas Sci. Bull. 21: 399–419.

Philipson WR. 1993. Hectorellaceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 2, pp. 331–334.

Springer, Berlin/Heidelberg/New York.

Philipson WR and JP Skipworth. 1961. Hectorellaceae: A new

family of dicotyledons. Trans. Roy. Soc. New Zealand Bot.

1: 31.

Plisko MA. 1991. Tetragoniaceae, Molluginaceae. In:

A Takhtajan, ed. Comparative seed anatomy, 3: 22–27.

Nauka, Leningrad (in Russian).

Pozner R. and A Cocucci. 2006. Floral structure, anther devel-

opment, and pollen dispersal of Halophytum ameghinoi

(Halophytaceae). Int. J. Plant Sci. 167: 1091–1098.

Prakash N. 1967a. Aizoaceae – a study of its embryology ans

systematics. Bot. Notis. 120: 305–323.

Prakash N. 1967b. Life history of Tetragonia tetragonioides

(Pal.) O. Kuntze. Austral. J. Bot. 15: 413–424.

Prance GT. 1968. The systematic position of Rhabdodendron

Gilg and Pilg. Bull. Jard. Bot. Nat. Belg. 38: 127–146.

Prance GT. 1972. Monograph of Rhabdodendraceae. Flora

Neotropica No.11: 1–22.

Prance GT. 2003. Rhabdodendraceae. In: K Kubitzki, ed. The

families and genera of vascular plants, vol. 5, pp. 339–341.

Springer, Berlin/Heidelberg/New York.

Pratt DB 2003. Phylogeny and morphological evolution of the

Chenopodiaceae-Amaranthaceae alliance. Ph.D. Thesis,

Iowa State University. Ames.

Pratt DB, LG Clark, and RS Wallace. 2001. A tale of two fami-

lies: Phylogeny of the Chenopodiaceae-Amaranthaceae. In

Botany 2001: Plants and People, p. 135 (Abstracts).

Puff C and A Weber. 1976. Contributions to the morphology,

anatomy, and karyology of Rhabdodendron and a reconsid-

eration of the systematic position of the Rhabdodendraceae.

Plant Syst. Evol. 125: 195–222.

Rabeler RK and V Bittrich. 1993. Suprageneric nomenclature in

the Caryophyllaceae. Taxon 42: 857–863.

Raghavan TS and VK Srinivasan. 1940. Studies in the Indian

Aizoaceae. Ann. Bot. 4: 651–661.

Raiput KS. 2002. Stem anatomy of Amaranthaceae: rayless

nature of xylem. Flora 197: 224–232.

Rajput KS. 2002. Stem anatomy of Amaranthaceae: Rayless

nature of xylem. Flora 197: 224–232.

Raipyt KS and KS Rao. 1999. Structural and developmental

studies on cambial variant in Pupalia lappaeae

(Amaranthaceae). Ann. Bot. Fenn. 36: 137–141.

Rajput KS and KS Rao. 2000. Secondary growth in the stem of

some species of Alternanthera and Achyranthes aspera

(Amaranthaceae). IAWA J. 21: 417–424.

Rao BS. 1975. Embryo development in ﬁ ve species of Mollugo.

Current Sci. 44: 712–713.

Rauh W. 1956. Morphologische, entwicklungsgeschich- tliche,

histogenetische, und anatomische Untersu-chungen an

Didiereaceen. Sitzungsber. Abh. Akad.Wiss. Lit., Math.-Nat.

Kl, 6: 1–104.

Rauh W. 1961. Weitere Untersuchungen an Didiereaceen.

Part 1. Sitzungsber. Heidelb. Akad. Wiss., Math.-Nat. Kl, 7:

1–118.

Rauh W. 1965. Weitere Untersuchungen an Didieraceen. II. Teil:

Inﬂ orescenz, blütenmorphologische und embryologische

Untersuchungen mit Ausblick auf die systematische Stellung

der Didieraceen. Sitzungsber. Heidelberger Akad. Wiss.

Jahresh. 1965: 221–443.

Rauh W and K Ditmar. 1970. Weitere Untersuchungen an

Didiereaceen. Part 3. Sitzungsber. Heidelb. Akad.Wiss.,

Math.-Nat. Kl, 4: 1–88.

Rauh W and H Reznik. 1961. Zur Frage der systematischen

Stellung der Didiereaceen. Bot. Jahrb. Syst. 81: 94–105.

Rauh W and HR Schölch. 1965. Weitere Untersuchungen an

Didiereaceen. Part 2. Sitzungsber. Heidelb. Akad. Wiss.,

Math.-Nat. Kl, 3: 1–218.

Record SJ. 1933. The woods of Rhabdodendron and

Duckeodendron. Trop. Woods 33: 6–10.

Rettig JH, HD Wilson, and JR Manhart. 1992. Phylogeny of the

Caryophyllales: Gene sequence data. Taxon 41: 201–223.

Reznik H. 1955. Die Pigmente der Centrospermen alssystem-

atisches Element. Z. Bot. 43: 499–530.

Reznik H. 1975. Betalaine. Ber. Deutsch. Bot. Ges. 88:

179–190.

Richardson M. 1978. Flavonols and C-glycosylﬂ avonoids of the

Caryophyllales. Biochem. Syst. Ecol. 6: 283–286.

Richardson M. 1981. Flavonoids of some controversialmembers

of the Caryophyllales (Centrospermae). Plant Syst. Evol.

138: 227–233.

Robertson KR. 1981. The genera of the Amaranthaceae in the

southeastern United States. J. Arnold Arbor. 62: 267–314.

Rodman JE, MK Oliver, RR Nakamura, JU McClammer, Jr., and

AH Bledsoe. 1984. A taxonomic analysis and revised clas-

siﬁ cation of Centrospermae. Syst. Bot. 9: 297–323.

Rogers GK. 1985. The genera of Phytolaccaceae in the south-

eastern United States. J. Arnold Arbor. 66: 1–37.

Rohweder O. 1965. Centrospermen-Studien: 1. Der Blütenbau

bei Uebelinia kiwuensis T. C. E. Fries (Caryophyllaceae).

2. Entwicklung und morphologische Deutung der Gynöciums

bei Phytolacca. Bot. Jahrb. Syst. 83: 406–418; 84: 509–526.

Rohweder O. 1967. Centrospermen-Studien; 3. Blüten-

entwicklung und Blütenbau bei Silenoideen (Caryophyllaceae).

Bot. Jahrb. Syst. 86: 130–185.

Superorder CARYOPHYLLANAE 151

Rohweder O. 1970. Centrospermen-Studien: 4. Morphologie

und Anatomic der Blüten, Früchte, und Sa-men bei

Alsinoideen und Paronychioideen s.lat. (Caryophyllaceae).

Bot. Jahrb. Syst. 90: 201–271.

Rohweder O and K Huber. 1974. Centrospermen-Studien: 7.

Beobachtungen und Anmerkungen zur Morphologie und

Entwicklungsgeschichte einiger Nyctaginaceen. Bot. Jahrb.

Syst. 94: 327–359.

Rohweder O and K Konig. 1971. Centrospermen-Studien: 5.

Bau der Blüten, Früchte, und Samen von Pteranthus dichot-

omus Forsk. (Caryophyllaceae). Bot. Jahrb. Syst. 90:

447–468.

Rohweder O and K Urmi-König. 1975. Centrospermen-Studien: 8.

Beiträge zur Morphologic, Anatomic, und systematischen

Stellung von Gymnocarpos Forsk. und Paronychia argentea

Lam. (Caryophyllaceae). Bot. Jahrb. Syst. 96: 375–409.

Rohwer JG. 1993. Phytolaccaceae, Stegnospermataceae. In:

K Kubitzki, ed. The families and genera of vascular plants,

vol. 2, pp. 506–515, 592–594. Springer, Berlin/Heidelberg/

New York.

Ronse Decraene LP and EF Smets. 1991. The ﬂ oral ontogeny of

some members of the Phytolaccaceae (subfamily Rivinoideae)

with a discussion on the evolution of the androecium in the

Rivinoideae. Biol. Jahrb. Doidonaea 39: 77–99.

Ronse Decraene LP, FF Smets, and P Vanvinckenroye. 1998.

Pseudodiplostemony, and its implications for the evolution

of the androecium in the Caryophyllaceae. J. Plant Res. 111:

25–43.

Ronse Decraene LP, P Vanvinckenroye, and EF Smets. 1997.

A study of ﬂ oral morphological diversity in Phytolacca

(Phytolaccaceae) based on early ﬂ oral ontogeny. Int. J. Plant

Sci. 158: 57–72.

Ronse Decraene LP, SA Volgin, EF Smets. 1999. The ﬂ ora deve-

lopment of Pleuropetalum darwinii, an anomalous member

of the Amaranthaceae. Flora (Germany) 194(2): 189–199.

Ross R. 1981. Chromosome counts, cytology, and reproduction

in the Cactaceae. Am. J. Bot. 68: 463–470.

Rost TL, AD Samper, P Schell, and S Alien. 1977. Anatomy of

jojoba (Simmondsia chinensis) seed and the utilization of liq-

uid wax during germination. Econ. Bot. 31: 140–147.

Sanderson SC. et al. 1988. Evolutionary loss of ﬂ avonoids and

other chemical characters in the Chenopodiaceae. Biochem.

Syst. Ecol. 16: 143–149.

Satyanarayana P and LL Narayana. 1985. Embryology of

Stegnospermataceae. Curr. Sci. 54: 882–884.

Schaeppi H. 1936. Zur Morphologic des Gynoeceums der

Phytolaccaceen. Flora 131: 41–59.

Schölch H-F. 1963. Die systematische Stellung der Di-diereaceen

im Lichte neuer Untersuchungen über ihren Blütenbereich.

Ber. Deutsch. Bot. Ges. 76: 49–55.

Schütze P, H Freitag, and K Weising. 2003. An integrated mole-

cular and morphological study of the subfamily Suaedoideae

Ulbr. (Chenopodiaceae). Plant Syst. Evol. 239: 257–286.

Scogin R. 1980. Serotaxonomy of Simmondsia chinensis

(Simmondsiaceae). Aliso 9: 555–559.

Scogin R and S Brown. 1979. Leaf ﬂ avonoids of Simmondsia

chinensis (Simmondsiaceae). Aliso 9: 475–477.

Scott AJ. 1977. Reinstatement and revision of Salicor-niaceae

J. Agardh (Caryophyllales). Bot. J. Linn. Soc. 75: 357–374.

Scott AJ. 1978. A revision of the Camphorosmoideae

(Chenopodiaceae). Feddes Repert. 89: 101–119.

Sharma HP. 1961. Contribution to the morphology and anatomy

of Basella rubra Linn. Bull. Bot. Soc. Bengal. 15: 43–48.

Sharma HP. 1962. Contributions to the morphology of the

Nyctaginaceae: I. Anatomy of the node and inﬂ orescence of

some species. Proc. Indian Acad. Sci. 56B: 35–50.

Sharma HP. 1963. Contributions to the morphology of the

Nyctaginaceae: II. Floral anatomy of some species. Proc.

Indian Acad. Sci. 57B: 149–163.

Sharma HP. 1962. Studies in the order Centrospermales: III.

Vascular anatomy of the ﬂ ower of some species of the family

Ficoidaceae. Proc. Indian Acad. Sci. 56B: 269–285.

Sharma HP. 1963. Studies in the order Centrospermales: II.

Vascular anatomy of the ﬂ ower of certain species of the

Molluginaceae, Nyctaginaceae, and Portulacaceae. J. Indian

Bot. Soc. 42: 19–32; 637–645.

Shepherd A, TD Macfarlane and TD Colmer. 2005.

Morphology, anatomy and histochemistry of fruits and

seeds of the Salicornioideae (Chenopodiaceae). Ann. Bot.

95: 917–933.

Shepherd A, TD Macfarlane, and M Waycott. 2005. Phylogeneic

analysis of the Australian Salicornioideae (Chenopodiaceae)

based on morphology and nuclear DNA. Austral. Syst. Bot.

18: 89–115.

Shepherd A, M Waycott, and A Calladine. 2004. Radiation of

the Australian Salicornioideae (Chenopodiaceae) – based on

evidence from nuclear and chloroplast DNA sequences. Am.

J. Bot. 91: 1387–1397.

Shilkina IA. 1953. Anatomical peculiarities of the family

Chenopodiaceae. Bot. Zhurn. 38: 590–598 (in Russian).

Shipunov AB. 2003. The system of ﬂ owering plants from a syn-

thetic point of view. J. Gen. Biol. 64: 501–510 (in Russian).

Singh BP. 1989. The status of Aizoaceae and Molluginaceae in

Centrospermae. J. Econ. Taxon. Bot. 13: 311–313.

Singh BP. 1992. Chemotaxonomy of Aizoaceae and

Molluginaceae. A study of free amino and organic acids. J.

Econ. Taxon. Bot. 16: 103–106.

Sivarajan VV. 1988. A preliminary taxonomic revision of Indian

Molluginaceae. J. Taiwan Mus. 41(2): 79–93.

Skipworth JP. 1961. The taxonomic position of Hectorella

caespitosa. Trans. Roy. Soc. New Zealand Bot. 1: 17–30.

Skvaria JJ and JW Nowicke. 1976. Ultrastructure of pollen exine

in centrospermous families. Plant Syst. Evol. 126: 55–78.

Skvaria JJ and JW Nowicke. 1982. Pollen ﬁ ne structure and rela-

tionships of Achatocarpus triana and Phaulothamnus

A. Gray. Taxon 31: 224–249.

Smissen RD, JC Clement, PJ Garnock-Jones, and GK Chambers.

2002. Subfamilial relationships within Caryophyllaceae as

inferred from 5’ndhF sequences. Am. J. Bot. 89: 1336–1341.

Soriano A. 1946. Halophytaceae. Nueva familia del orden

Centrospermae. Notas Prelim. Mus. La Plata, Bot. 11:

161–175.

Sperling CR and V Bittrich. 1993. Basellaceae. In: K Kubitzki,

ed. The families and genera of vascular plants, vol. 2,

pp. 143–152. Springer, Berlin/Heidelberg/New York.

Straka H. 1965. Die Pollenmorphologie der Didiereaceen.

Sitzungsber. Heidelb. Akad. Wiss., Math.-Nat. Kl., 3: 219–227.

Straka H. 1975. Palinologie et différentiation systématique

d’une famille endémique de Madagascar: Didiereacees.

Boissiera 24: 245–248.

Terrazas T and S Arias. 2002. Comparative stem anatomy in the

subfamily Cactoiodeae. Bot. Rev. 68: 444–473.