ROSANAE Takht.

Takhtajan, Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 264. 4 Feb 1967

Rosopsida Batsch, Tab. Regni Veg.: 1. 1802 [’Rosaceae’]; Hamamelididae Takht, Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 461. 4 Feb 1967, pro parte; Rosidae Takht., Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 264. 4 Feb 1967 pro parte; Dilleniidae Takht. ex Reveal et Takht. in Phytologia 74: 171. 25 Mar 1993, pro parte


[Fabidae+Malvidae]


FABIDAE W. S. Judd, D. E. Soltis et P. S. Soltis

Judd, Soltis et Soltis in Taxon 56: E29. Aug 2007

(eurosids I)


[Zygophyllales+COM clade+Nitrogen Fixing clade]


ZYGOPHYLLALES Link

Link, Handbuch 2: 228. 4-11 Jul 1829 [’Zygophylleae’]

Zygophyllanae Doweld, Tent. Syst. Plant. Vasc.: xxxviii. 23 Dec 2001

Habit Usually bisexual (rarely dioecious), usually evergreen shrubs, suffrutices, or perennial or annual herbs (rarely trees), sometimes with axillary, simple or branched, spines. Often xerophytes or halophytes. C4-photosynthesis present.

Vegetative anatomy Mycorrhiza probably absent. Phellogen ab initio superficially or deeply seated (pericyclic to deep cortical). Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary elements tracheids or fibre tracheids with simple or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma usually apotracheal diffuse, diffuse-in-aggregates or banded (rarely paratracheal scanty vasicentric). Sieve tube plastids usually S type (rarely Pcs type, with two unequally sized protein crystalloids). Nodes 1:1 or 3:3, unilacunar with one leaf trace or trilacunar with three traces. Parenchyma often with sclerenchymatous idioblasts. Tanniniferous secretory cells sometimes present. Dark-brown substances present in heartwood in some species (e.g. Guaiacum officinale). Calciumoxalate present as druses or styloids, prismatic or acicular crystals, often in idioblasts.

Trichomes Hairs usually unicellular (sometimes multicellular), simple or furcate (rarely peltate or capitate); glandular hairs rarely present.

Leaves Usually alternate (spiral) or opposite, usually pinnately compound (rarely unifoliolate, bifoliolate or trifoliolate), leaflets entire, usually persistent (rarely caducous), often coriaceous, sometimes succulent or modified into spines, with ? ptyxis. Stipules cauline or interpetiolar (sometimes modified into spines) or absent; leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation usually pinnate (rarely palmate, leaves sometimes uninerved), eucamptodromous or brochidodromous. Stomata usually anomocytic or paracytic (rarely cyclocytic or actinocytic), sometimes transversely orientated. Cuticular wax crystalloids at least sometimes as platelets. Epidermis and mesophyll sometimes with mucilaginous idioblast. Tannins and crystals often frequent. Lamina rarely gland-dotted. Leaves or leaflet margins entire.

Inflorescence Usually axillary (sometimes terminal or leaf-opposite), usually few-flowered cymose or racemose of different shape, or flowers solitary axillary. Floral prophylls (bracteoles) small or absent.

Flowers Usually actinomorphic or zygomorphic. Usually hypogyny (rarely half epigyny). Sepals (four or) five (or six), with imbricate or valvate aestivation, sometimes caducous, usually free (rarely connate at base). Petals (four or) five (or six), usually with imbricate aestivation, often clawed, usually free (sometimes partially connate). Disc annular or intrastaminal and lobate, or absent, sometimes also with extrastaminal and/or intrastaminal nectariferous glands.

Androecium Stamens usually four, eight or ten (sometimes three, five or twelve), in one or two whorls, antesepalous and/or alternisepalous (antesepalous whorl sometimes consisting of staminodia or absent). Filaments sometimes winged, sometimes with scales or other appendages at base, free or connate at base, usually free from tepals. Anthers dorsifixed or basifixed, versatile or non-versatile, tetrasporangiate, introrse to latrorse, longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical pores or short slits). Tapetum secretory. Staminodia usually absent (extrastaminal antesepalous staminal whorl or median abaxial stamen rarely staminodial).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes tricolpate, tricolporoidate, tetracolporate, triporate, synorate, or polypantoporate, rarely hexarugorate), shed as monads, bicellular or tricellular at dispersal. Exine tectate or semitectate, with columellate infratectum, usually reticulate (sometimes striate, rarely rugulate, columellate or retipilate).

Gynoecium Pistil composed of two to five (or six) connate antepetalous carpels. Ovary usually superior (rarely semi-inferior), bilocular or (quadrilocular or) quinquelocular (to duodecemlocular); locules sometimes divided by secondary septa. Style single, simple (sometimes gynobasic). Stigma punctate, capitate, clavate or as commissural ridges down style, papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation axile(-marginal; sometimes apical). Ovules usually one to ten (sometimes more than ten) per carpel, usually anatropous (rarely campylotropous, hemianatropous or orthotropous), pendulous, epitropous, bitegmic, weakly crassinucellar. Micropyle bistomal or endostomal. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis usually solanad (rarely caryophyllad).

Fruit Usually a loculicidal and/or septicidal capsule (sometimes a nutlike capsule or a schizocarp with two or five nutlike mericarps; rarely a drupe).

Seeds Aril usually absent. Seed coat usually testal or exotestal (rarely endotegmic). Exotesta often palisade, sometimes with enlarged tanniniferous cells. Endotesta with crystals, often lignified. Exotegmen? Endotegmic cells periclinally elongate, lignified. Perisperm not developed. Endosperm usually absent (sometimes oily). Embryo straight or somewhat curved, well differentiated, at least sometimes with chlorophyll. Cotyledons two. Germination phanerocotylar or cryptocotylar.

Cytology x = 6, 8–13 (14)

DNA Plastid gene infA present. Mitochondrial intron coxII.i3 lost.

Phytochemistry Flavonols (kaempferol, quercetin), methylated flavonoids, ethereal oils (sesquiterpenes etc., in wood), oleanolic acid derivatives, tannins, phlobaphene, indole and quinazoline alkaloids (harman alkaloids, e.g. harmane, harmine, harmol), steroidal and triterpene saponins, anthraquinones, lignans, neolignans, nor-neolignans, pinitol, apiitol, and N-methyltyrosine present. Ellagic acid, tannins, proanthocyanidins, and cyanogenic compounds not found.

Systematics Zygophyllales may be sister-group to [‘Nitrogen fixing clade’+’COM clade’] (Soltis & al. 2011).

KRAMERIACEAE Dumort.

( Back to Zygophyllales )

Dumortier, Anal. Fam. Plant.: 20, 23. 1829, nom. cons.

Krameriales Kunth in C. F. P. von Martius, Consp. Regn. Veg.: 44. Sep-Oct 1835 [’Krameriaceae’]

Genera/species 1/18

Distribution Southern United States and southwards to Chile and Argentina.

Fossils Unknown.

Habit Bisexual, usually shrubs or suffrutices (Krameria lanceolata is a perennial herb, usually with woody rhizome). Root hemiparasites.

Vegetative anatomy Mycorrhiza probably absent. Phellogen ab initio deeply seated (sometimes in pericycle). Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Non-vestured pits present? Imperforate tracheary xylem elements (fibre?)tracheids with bordered pits, non-septate (also vasicentric tracheids). Wood rays usually uniseriate (rarely biseriate), homocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, or banded. Wood elements sometimes (axial parenchyma) partially storied. Wood fluorescent? Phloem non-lignified. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Parenchyma with tanniniferous secretory cells. Rhomboidal calciumoxalate crystals present especially in secondary phloem.

Trichomes Hairs unicellular, simple.

Leaves Leaves alternate (spiral), usually simple? (in reality unifoliolate?, rarely trifoliolate), entire, small, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate (hippocrepomorphic, sometimes almost circular). Venation pinnate? Stomata usually paracytic (sometimes anomocytic). Cuticular wax crystalloids as narrow platelets. Mesophyll with sclerenchymatous idioblasts with sclereids and calciumoxalate druses. Tannins and crystals abundant. Leaf(let) margin entire.

Inflorescence Terminal or axillary, raceme-like or botryoid-paniculate, or flowers solitary axillary. Floral prophylls (bracteoles) foliaceous, small.

Flowers Zygomorphic, probably inverted. Pedicel articulated. Hypogyny. Sepals (four or) five, with imbricate aestivation, petaloid inside, free; median sepal abaxial, larger than remainder; three outer sepals larger than two inner sepals, often almost enclosing remaining parts of flower. Petals (four or) five, with imbricate aestivation; median petal adaxial; (two or) three adaxial petals long-clawed, usually connate at base into velum; two abaxial petals often modified into sessile glands, excreting bee-attracting lipids (free β-acetoxy fatty acids) from elaiophores in epidermis. Nectary absent. Disc absent.

Androecium Stamens (three or) four, adaxial, antesepalous, alternipetalous. Filaments stout, usually connate at base, usually free from petals (sometimes adnate to claws of adaxial petals). Anthers basifixed, curved, non-versatile, tetrasporangiate, introrse, poricidal (dehiscing by one or two apical pores or short slits). Tapetum? Staminodia usually absent (median abaxial stamen rarely staminodial).

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, tetracolporate, triporate or synorate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, striate.

Gynoecium Pistil composed of two connate carpels; adaxial carpel early degenerating. Ovary superior, unilocular (pseudomonomerous, first bilocular). Style single, simple, stout, curved. Stigma small, punctate, sunken, type? Pistillodium absent.

Ovules Placentation apical. Ovules two per carpel, collateral, anatropous, pendulous, bitegmic, weakly crassinucellar. Micropyle endostomal. Outer integument approx. six cell layers thick. Inner integument four or five cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A one-seeded nutlike capsule with hairs and barbed spines; thin pericarp irregularly dehiscing.

Seeds Aril absent. Seed coat exotestal. Exotestal cells enlarged, tanniniferous. Endotesta? Tegmen up to seven cell layers thick above, mostly degenerating. Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, chlorophyll? Haustoria formed from young adventitious roots. Cotyledons two, large, ventrally flattened, cordate. Germination? Seedling without root hairs.

Cytology n = 6

DNA Plastid gene rps16 absent (lost). Plastid gene infA?

Phytochemistry Tannins of catechin type, phlobaphene (red root pigment), neolignans, nor-neolignans, apiitol, and N-methyltyrosine present. Saponins not found. Harman-alkaloids?

Use Medicinal plants, dyeing substances (roots), cosmetics.

Systematics Krameria (18; southwestern and southern United States, Mexico, Central America, the West Indies, northern and central South America to Chile and Argentina).

Krameria is sister to Zygophyllaceae.

ZYGOPHYLLACEAE R. Br.

( Back to Zygophyllales )

Brown in Flinders, Voy. Terra Austral. 2: 545. 19 Jul 1814 [’Zygophylleae’], nom. cons.

Balanitaceae M. Roem., Fam. Nat. Syn. Monogr. 1:26. 14 Sept-15 Oct 1846 [‘Balaniteae’], nom. cons.; Zygophyllineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 265, 266. 1846 [‘Zygophylleae’]; Tribulaceae Trautv., Estestv. Istorija Gub. Kievsk. Uchebn. Okr. Bot. Sist.: 28. 1853; Balanitales C. Y. Wu in Acta Phytotaxon. Sin. 40: 314. 2002

Genera/species 18/150–175

Distribution Mainly tropical and subtropical regions on both hemispheres, with their largest diversity in arid and subarid areas; some species in warm-temperate regions.

Fossils Unknown.

Habit Usually bisexual (rarely dioecious), evergreen shrubs, suffrutices, or perennial or annual herbs (rarely trees), sometimes with axillary, simple or branched, spines. Nodes often swollen and/or articulated. Bark often bitter. Some species are succulent. Many species are xerophytes or halophytes. C4 photosynthesis present (almost all species in Kallstroemia have C4 photosynthesis; nearly all in Zygophyllum are C3 photosynthetic).

Vegetative anatomy Mycorrhiza usually absent (arbuscular mycorrhiza sometimes present in Larrea). Phellogen ab initio usually superficially (sometimes deeply) seated. Primary vascular tissue cylinder without separate vascular bundles. Primary medullary rays usually narrow (in Balanites wide). Secondary lateral growth normal or anomalous. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary elements tracheids or fibre tracheids (or libriform fibres?) with simple and/or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, one or two (to four) cells wide (in Balanites up to c. 35 cells wide), homocellular or heterocellular. Axial parenchyma usually apotracheal diffuse, diffuse-in-aggregates or in uniseriate bands (rarely paratracheal scanty vasicentric). Wood elements usually storied. Wood often fluorescent. Sieve tube plastids usually S type (rarely Pcs type, with two unequally sized protein crystalloids). Nodes usually 1:1, unilacunar with one leaf trace, often also with split laterals (trilacunar nodes present in Viscainoa). Cortex and parenchyma often with fibres and sclerenchymatous idioblasts. Secretory cavities absent. Dark-brown substances present in heartwood in some species (e.g. Guaiacum officinale). Calciumoxalate present as druses and styloid, prismatic and acicular crystals, often in idioblasts.

Trichomes Hairs usually unicellular (sometimes multicellular), simple or furcate (rarely peltate or capitate); glandular hairs present in Fagonia.

Leaves Usually opposite (in Morkillia and Viscainoa alternate, spiral), usually pinnately compound (rarely bifoliolate or trifoliolate or probably seemingly simple: unifoliolate?), leaflets entire, usually persistent (rarely caducous), often coriaceous, sometimes succulent or modified into spines, with usually flat ptyxis. Stipules cauline or one interpetiolar (sometimes modified into spines; rarely absent); leaf sheath absent. Petiole vascular bundle transection annular; petiole with wing bundles. Venation usually pinnate (rarely palmate, leaves sometimes one-veined), eucamptodromous or brochidodromous. Stomata usually anomocytic (rarely paracytic, cyclocytic or actinocytic). Cuticular wax crystalloids? Epidermis sometimes with mucilaginous idioblasts. Mesophyll with or without sclerenchymatous idioblasts. Lamina rarely gland-dotted. Leaflet margins usually entire (sometimes serrate).

Inflorescence Usually axillary (sometimes terminal or leaf-opposite), few-flowered cymose or racemose of different appearance, or flowers solitary. Floral prophylls (bracteoles) absent.

Flowers Usually actinomorphic (rarely zygomorphic). Usually hypogyny (rarely partially perigynous). Sepals (four or) five (or six), with imbricate or valvate aestivation, sometimes caducous, usually free (rarely connate at base). Petals (four or) five (or six), usually with imbricate (or contorted?) aestivation, often clawed, free (absent in Seetzenia and one species of Zygophyllum). Disc annular or as intrastaminal lobes or absent, sometimes also with extrastaminal and/or intrastaminal nectariferous glands.

Androecium Stamens usually eight or ten (sometimes five or twelve), in one or two whorls, antesepalous and/or alternisepalous (antesepalous whorl staminodial or absent), often obdiplostemonous. Filaments sometimes winged, sometimes with scales or other appendages at base, free from each other and from tepals. Anthers dorsifixed, versatile, tetrasporangiate, introrse or latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia usually absent (extrastaminal antesepalous staminal whorl rarely staminodial).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes tricolpate, tricolporoidate or polypantoporate, rarely hexarugorate), shed as monads, bicellular or tricellular at dispersal. Exine semitectate, with columellate infratectum, usually reticulate (rarely rugulate, columellate or retipilate).

Gynoecium Pistil composed of (two to) five (or six) connate antepetalous carpels. Ovary usually superior (rarely semi-inferior), (quadrilocular or) quinquelocular (to duodecemlocular), often angular or winged (sometimes stipitate, with gynophore); locules sometimes divided by secondary septa. Style single, simple, short to long, narrow (in Zygophyllum gynobasic). Stigma capitate, punctate, clavate or as commissural ridges down style, papillate, Dry or Wet type. Pistillodium absent?

Ovules Placentation axile(-marginal; rarely apical). Ovules usually one to ten (sometimes more than ten) per carpel, usually anatropous (rarely campylotropous, hemianatropous or orthotropous), pendulous, epitropous (sometimes apotropous?), bitegmic, weakly crassinucellar. Micropyle endostomal or bistomal, usually not Z-shaped (in Balanites zig-zag). Outer integument two to six (to eight) cell layers thick. Inner integument two to four (to six) cell layers thick. Obturator present. Hypostase present. Endothelium usually present (absent in Seetzenia). Parietal tissue one or two (to four) cell layers thick. Nucellar cap sometimes present. Archespore sometimes multicellular. Megagametophyte monosporous, Polygonum type, elongated. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis usually solanad (rarely caryophyllad).

Fruit Usually a loculicidal and/or septicidal capsule (sometimes a schizocarp with two or five nutlike mericarps; in Balanites a one-seeded drupe with oily mesocarp).

Seeds Aril usually absent. Seed coat usually testal (rarely endotegmic). Exotesta often palisade. Endotesta usually crystalliferous, often lignified. Exotegmen? Endotegmic cells periclinally elongate, lignified. Perisperm not developed. Endosperm usually absent (sometimes present, oily). Embryo straight or somewhat curved, at least sometimes with chlorophyll. Cotyledons two. Germination phanerocotylar or cryptocotylar.

Cytology x = 6, 8–13 (14) – Polyploidy occurring.

DNA The plastid gene infA is present.

Phytochemistry Flavonols (kaempferol, quercetin), methylated flavonoids, ethereal oils (sesquiterpenes etc., in wood), oleanolic acid derivatives, quinazoline alkaloids (harmane, harmin, harmol and other harmala alkaloids), indole alkaloids, steroidal and triterpene saponins, anthraquinones, lignans, neolignans, and pinitol present. Ellagic acid, tannins, proanthocyanidins, and cyanogenic compounds not found. Mustard oils?

Use Ornamental plants, timber and carpentries (extremely hard wood, Lignum vitae, from Guaiacum), fruits (Balanites aegyptiaca), medicinal plants, edible flower buds.

Systematics Zygophyllaceae are sister-group to Krameria (Krameriaceae).

[[Morkillioideae+Balanitoideae]+[Seetzenioideae+[Larreoideae+Zygophylloideae]]] is a plausible topology (Sheahan & Case 2000).

[Morkillioideae+Balanitoideae]

Morkillioideae Thorne et Reveal in Bot. Rev. (Lancaster) 73: 106. 29 Jun 2007

3/4. Morkillia (2; Mexico), Viscainoa (1; V. geniculata; Baja California in northwestern Mexico), Sericodes (1; S. greggii; northern Mexico). – Mexico. Leaves usually alternate (spiral).

Balanitoideae Engl. in Engler et Prantl, Nat. Pflanzenfam. III, 4: 354, 355. Jul 1896

6/38–56. Tribulus (7–25; tropical and subtropical regions on both hemispheres), Kallstroemia (17; tropical and subtropical America), Kelleronia (3; northeastern Africa, southern Arabian Peninsula), Sisyndite (1; S. spartea; southern Namibia, Northern and northwestern Western Cape), Neoluederitzia (1; N. sericeocarpa; southern Namibia), Balanites (9; tropical Africa, the Arabian Peninsula and eastwards to India and Burma). – Tropical to warm-temperate regions on both hemispheres. C4 photosynthesis frequent in Kallstroemia. Fruit usually a schizocarp with four or five or up to ten single-seeded nutlike mericarps. – Balanites are spiny shrubs or trees with bitter bark. Medullary and wood rays very wide. Stipules tiny (absent?). Petiole anatomy complicated. Stem stomata transversely orientated relative to longitudinal axis. Leaves alternate (spiral) and pinnately compound with one pair of leaflets. Corolla enclosing floral bud. Ovule one per carpel, pendulous. Micropyle bistomal, Z-shaped (zig-zag). Outer integument four to eight cell layers thick. Inner integument three to six cell layers thick. Fruit a single-seeded drupe. Endosperm absent. Testa multiplicative, vascularized.

[Seetzenioideae+[Larreoideae+Zygophylloideae]]

Seetzenioideae M. C. Sheahan et M. W. Chase in Bot. J. Linn. Soc. 122: 299. 18 Dec 1996

1/1. Seetzenia (1; S. lanata; North Africa and eastwards to Afghanistan and India, Northern and Eastern Cape). – Prostrate perennial herb. Leaves opposite, trifoliolate. Petals absent. Stamens five. Filaments without scale-like processes. Carpels five. Stylidia five. Ovule one per carpel, epitropous. Micropyle bistomal. Outer integument six or seven cell layers thick. Endothelium absent. Fruit a septicidal capsule, dehiscing into five single-seeded cocci with fleshy exocarp and bony endocarp. Endosperm present.

[Larreoideae+Zygophylloideae]

Larreoideae M. C. Sheahan et M. W. Chase in Bot. J. Linn. Soc. 122: 299. 18 Dec 1996

7/29. Bulnesia (8–10; South America), Guaiacum (6; tropical and subtropical America), Porlieria (3–4; Mexico, the Andes), Larrea (5–6; southwestern United States, Mexico, South America), Pintoa (1; P. chilensis; Chile), Plectrocarpa (3; temperate South America), Metharme (1; M. lanata; northern Chile). – Southwestern United States, Mexico, Central and South America to Chile and Argentina. Sieve tube plastids in Larrea with protein and starch. Filaments often with scale-like processes. Ovary stipitate or sessile. Capsule often winged. Seed one per locule. Endosperm present.

Zygophylloideae Arn., Botany: 104. 9 Mar 1832 [‘Zygophylleae’]

1/80–85. Zygophyllum (80–85; Macaronesia, the Mediterranean, southeastern Europe, northern and northeastern Africa and eastwards to Iran, Central Asia and northwestern India, China, tropical and subtropical to southern Africa, Australia, southwestern United States, Mexico). – Mainly drier regions of the Old World, southwestern United States, Chile. Nectariferous disc octa- or decemangulate. Stamens eight or ten. Filaments with basal scale-like processes. Style in Zygophyllum simple, gynobasic. Stigma punctate. Outer and inner integuments in Zygophyllum approx. two cell layers thick. Fruit an angular to winged capsule or schizocarp.

Unplaced Zygophyllaceae: Izozogia (1; I. nellii Bolivia).

Cladogram (simplified) of Zygophyllaceae based on DNA sequence data (Sheahan & Chase 2000).


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