Celastraceae R.Br.

Celastraceae, N. K. Robson. Flora Zambesiaca 2:2. 1966

Morphology General Habit

Trees, shrubs, shrublets or woody climbers, without tendrils, glabrous or with simple hairs, unarmed or with axillary shoots terminating in a spine, sometimes with rubber-like latex (gutta) in various parts appearing as elastic threads when a leaf is broken

Morphology Leaves

Leaves simple, alternate or spiral to subopposite or opposite, sometimes fasciculate on short shoots, entire or with crenate or denticulate to spinose margins, penninerved; stipules small, simple or laciniate, sometimes united by a transverse ridge, usually deciduous, or absent

Morphology Reproductive morphology Flowers

Flowers bisexual or polygamous or unisexual, monoecious or dioecious, actinomorphic, often fragrant, in axillary and/ or terminal dichasial or monochasial cymes or panicles or thyrses, sometimes with accessory branches, or fasciculate or solitary, usually bracteate; pedicels often articulated

Morphology Reproductive morphology Flowers Calyx

Sepals (3)4–5(6), imbricate or rarely valvate in bud, free or united at the base, persistent

Morphology Reproductive morphology Flowers Corolla

Petals (3)4–5(6), free or rarely united at the base, imbricate or rarely valvate in bud, usually persistent, sometimes with ventral grooves or hollows or appendages

Morphology Reproductive morphology Flowers Androecium

Stamens (2)3–5(6–10), antisepalous, free or more rarely with filaments partly united to form a tube, inserted outside or on or inside the disk; anthers usually short, (1)2-thecous, extrorse or introrse, basifixed or dorsifixed or versatile, sometimes deciduous, dehiscing by longitudinal or oblique or horizontal slits; pollen simple or more rarely in tetrads or polyads

Morphology Reproductive morphology Flowers Nectaries

Disk nectariferous, annular, entire or angular or crenulate or lobed or covered with fleshy processes, concave to convex, rarely wholly or partly forming an androgynophore or discontinuous pockets, fleshy or membranous, very rarely absent

Morphology Reproductive morphology Flowers Gynoecium

Ovary free or partly or wholly immersed in the disk, sessile or on a short androgynophore, syncarpous, completely or very rarely incompletely 2–5-locular, or rarely 1-locular by abortion, with 1-? erect or rarely pendulous ovules in 2 rows or rarely superimposed in each loculus; styles as many as the loculi, free or ± united, or absent; stigmas various, free or ± united

Morphology Reproductive morphology Fruits

Fruit capsular, loculicidal, or of divergent ± dorsiventrally flattened dehiscent mericarps or baccate or drupaceous or dry, indehiscent and sometimes winged

Morphology Reproductive morphology Seeds

Seeds with a fleshy or submembranous brightly coloured aril, or winged with the funicle free from the wing (Tab. 80 figs 6–7) or united to its base (Tab. 85 fig. A8) or neither arillate nor winged (usually in indehiscent fruits), with or without endosperm; embryo erect, with cotyledons flat or fleshy, rarely united

Celastraceae, R.A. Blakelock. Flora of West Tropical Africa 1:2. 1958

Morphology General Habit

Small trees, shrubs or woody climbers without tendrils, sometimes spiny

Morphology Leaves

Leaves simple, alternate or opposite

Morphology Leaves Stipules

Stipules inconspicuous or absent

Morphology Reproductive morphology Flowers

Flowers bisexual or unisexual

Morphology Reproductive morphology Flowers Calyx

Sepals 4–5, usually free

Morphology Reproductive morphology Flowers Corolla

Petals 4–5, free

Morphology Reproductive morphology Flowers Androecium

Stamens 3–5, alternate with petals

Morphology Reproductive morphology Flowers Gynoecium

Carpels 3–5 united, free from or half-embedded in the disk Ovules 2–12 in each carpel

Morphology Reproductive morphology Fruits

Fruit a loculicidal capsule, an indehiscent fleshy or hard drupe, a berry, a 3-lobed capsule or a capsule with 3 nearly separate mericarps

Morphology Reproductive morphology Seeds

Seed sometimes arillate, sometimes winged, sometimes neither

Archer, R.H. & Lombardi, J.A. (2013). Neotropical Celastraceae.

Morphology

Description

Lianas, shrubs, or trees, sometimes treelets with slender branches ± scandent; stems with anomalous growth, with  phloem present in the fleshy -fruited genera (Salacia L.); lateral brachyblasts sometimes present and tendril -like, sometimes restarting growth after twining, rarely with stems terminating in sharp points; hairs rarely present, simple and generally restricted to inflorescences and flowers, rare on young branches, leaves and fruits; stipules present, minute and usually caducous , sometimes leaving interstipular scar. Leaves opposite or subopposite, alternate , simple ; blade margins crenulate , serrate or entire , petiolate or rarely sessile . Inflorescences axillary or on axillary brachyblasts (some may falsely appear to be terminal ), thyrsoid, cymose, corymbose or fasciculate ; bracts, and sometimes bracteoles, present, small. Flowers actinomorphic , small, bisexual ; flower buds spherical to ovate ; perianth usually persistent ; sepals 4 or 5, imbricate , free , connate at the very base, or rarely connate , not enclosing petals in bud ; petals 4 or 5, imbricate , distinct, alternate to sepals, usually yellow-green to white; extrastaminal nectariferous disc present and usually well-developed, annular, pulvinate , patelliform, short-tubular, cupuliform, cylindric, or columnar; androecium usually with 3 stamens, rarely 5 (in Hippocrateoideae), 4 or 5 (in the remaining subfamilies), minute, distinct, reflexed outward after anthesis, the filaments sometimes connate to inner disk wall, the anthers dehiscing by transverse or oblique slits; gynoecium syncarpous, the ovary superior , often triangular in shape, the carpels usually 3, 4 or 5, the locules equal to the number of carpels, style short, awl-shaped, sometimes absent, the stigmas usually 3 entire , bilobed, or obscure ; placentation axillary or subapical, the 1-10 ovules per locule , patent or somewhat pendulous. Fruits berries, or woody capsules, the capsules commonly 3- lobed , or a schizocarp with 3 strongly divergent dehiscent mericarps on swollen receptacle , or samara with single apical wing, 3-5 lateral wings or single surrounding wing; seeds usually 3-6 (in berries), 6 to ± 30 (in capsules), often 3-sided (in berries), sometimes winged at base (in capsules), wing sometimes vestigial and testa spongy , or with sarcotesta and embedded in mucilaginous pulp (in berries) exarrilate or with membraneous or fleshy , basal to completely enveloping seed ; endosperm absent or present.

General Description

Notes on delimitation

Hippocrateaceae is usually included in Celastraceae as two distinct subfamilies in most recent classifications, particularly the APG III. The Celastroideae comprise most additional genera in the Neotropics:

• Celastroideae (trees, shrubs, simplecapsule -fruited species).
• Cassinoideae (trees, drupe -fruited species).
• Hippocrateoideae (mostly lianas, dry-fruited species).
• Salacioideae (mostly lianas, fleshy -fruited species).
• Goupia (Goupiaceae) and Perrottetia (Dipentodontaceae) are placed in the order Malpighiales following molecular analysis. Parnassia and Lepuropetalon are treated in the family Parnassiaceae.

Number of genera

100 genera worldwide.

27 genera native in the Neotropics (see Distribution in the Neotropics above).

Status

• All genera in the Neotropics are native.

Distribution

Distribution in the Neotropics

• Acanthothamnus Brandegee - Endemic with 1 species, Central Mexico.
• Anthodon Ruiz & Pav. - Endemic Panama to SE Brazil.
• Celastrus L.- Mostly Northern hemisphere in Mexico to Bolivia, Brazil and West Indies
• Cheiloclinium Miers - Endemic Mesoamerica to SE Brazil.
• Crossopetalum P.Browne  - Endemic Florida, West Indies to Venezuela.
• Cuervea Triana ex Miers - Endemic Mesoamerica to SE Brazil, Caribbean islands (St Vincent, Cuba, and Jamaica).
• Euonymus L. - Mostly Northern hemisphere but with a few in Mesoamerica.
• Elachyptera A.C.Sm. - Mesoamerica to SE Brazil.
• Elaeodendron Jacq. - Mostly southern hemisphere with 1 (or possibly 2) species in Mesoamerica, Caribbean islands.
• Fraunhofera Mart. - Endemic, in NE Brazil.
• Hippocratea L. - Florida to N Argentina, and Caribbean islands.
• Hylenaea Miers - Endemic Costa Rica to Amazon basin.
• Goniodiscus Kuhlm. - Endemic, 1 species in Maués, Amazonas, Brazil.
• Gyminda Sarg. - Endemic, W. Cuba, Mexico to Panama and Venezuela.
• Haydenia M.P.Simmons - Endemic, 3 species; Veracruz, Mexico to Bolivia and Amazonian Brazil.
• Maytenus Molina - Near-endemic and widespread in Neotropics; largest group with more than 100 species.
• Peritassa Miers - Endemic Costa Rica to Paraguay.
• Plenckia Reissek - Endemic
• Prionostemma Miers - Endemic as a monotypic genus; Mesoamerica to NE Brazil and Bolivia.
• Pristimera Miers - México to N Argentina.
• Quetzalia Lundell - Endemic, Mesoamerica.
• Salacia L. - México to Paraguay, and Cuba.
• Schaefferia Jacq. - Endemic, South America, West Indies.
• Semialarium N.Hallé - Endemic Mexico to SE Brazil.
• Tetrasiphon Urb. - Endemic, Jamaica.
• Tontelea Miers - Endemic Mesoamerica to Paraguay.
• Wimmeria Schltdl & Cham. - Endemic Mexico to Panama.
• Zinowiewia Turcz. - Endemic with Mexico to Bolivia, and Venezuela.

Rzedowskia, Canotia, and Mortonia, are all small shrubby Celastraceous species with small or scaly leaves only that occur as far south as Central Mexico, thus just outside the Neotropics area and not here included.

Diagnostic

Other important characters

• Fruit particularly varied, loculicidally and/or septicidally capsular or baccate, drupaceous, samaroid or with seed often arillate or sometimes winged.
• Twining lianas with leaves opposite (Hippocrateoideae and Salacioideae).
• Leaves glabrous, rarely scabrous.
• Stamens and carpels 3 (Hippocrateoideae and Salacioideae) 4 or 5 in Celastroideae and Cassinoideae.
• Extrastaminal nectariferous disc (Hippocrateoideae and Salacioideae), intrastaminal in Celastroideae and Cassinoideae).
• Leaves often crenate/serrate.

Distinguishing characters (always present)

• Leaves simple.
• Axillary buds glabrous.
• Stipules present and minute, but usually caduceous, leaving a stipular scar.
• Nectariferous disc present.

Key differences from similar families

• Stipules present (absent in Myrtaceae).
• Axillary buds glabrous, hairs rare anyway (axillary buds pilose and hairs common in Malpighiaceae).
• Superiorovary and dialypetalous (inferior ovary and gamopetalous in Rubiaceae).

Notable genera and distinguishing features

• Hippocratea - young branches, inflorescences, pedicels, sepals and petals pilose, petals barbelate inside in the apex, fruit a dehiscentschizocarp.
• Salacia - young branches, inflorescences and pedicels rarely puberulous, petals glabrous, fruit a berry.
• Maytenus- rather indistinguishable with small flowers and loculicidally dehiscentcapsule, frequently opening in 2 valves, with 1 or 2 arillate seeds.

Useful tips for generic identification

• Acanthothamnus - shrub, stems with dark glands and terminating in sharp point, fruit a fleshy drupe.
• Anthodon - lianas, inflorescence a composite dichasium, petals serrate, disc annular, fruit a dehiscent obinfundibuliform capsule.
• Celastrus - scandent scrubs or lianes, with conspicuous lenticellate branchlets, fruit a loculicidal deshiscent capsule, seeds enclosed by fleshy red aril.
• Cheiloclinium - lianas to trees, inflorescence a composite dichasium or thyrsoid-paniculate, disc enclosing the pistil, only the 3(-5)-lobed stigmas apparent, stamens in little pockets in the outside of the disc, fruit a berry.
• Crossopetalum - small trees or shrubs, inflorescence axillary, cymose, ovary (2-) 4-locular with ovules erect, small drupes.
• Cuervea - lianas, inflorescence a monochasium, composite dichasium, or thyrsoid, flowers 8-15 mm diam. at anthesis, disc annular or short-tubular, fruit a dehiscent schizocarp.
• Elaeodendron - dioeciousevergreen trees, pistillate flowers with petaloid staminodes (Neotropics), fruit a large indehiscent drupe with a hard stone enclosing seeds.
• Euonymus - small trees or shrubs, capsule loculidically dehiscent subspheroid with aril enveloping seed.
• Elachyptera - lianas, inflorescence a glabrous composite pleochasium with branches conspicuously 4-angled, disc annular, fruit a dehiscent schizocarp.
• Fraunhofera - trees, pubescent, inflorescence racemose, fruit possibly a drupe.
• Goniodiscus - tree, fruit a large drupe, seed exalbuminous, oily?
• Gyminda - unisexual shrubs, flowers white, 2-locular, ovules pendulous, fruit 2-locular.
• Haydenia - dioecious trees, inflorescence large, cymose with 4-merous flowers, fruit a loculicidal capsule, 2-4 lobed.
• Hippocratea - lianas, inflorescence a pilose composite pleochasium, young branches and inflorescence branches pilose, discpulvinate, petals barbelate in the apex, fruit a dehiscent schizocarp.
• Hylenaea - lianas, inflorescence a glabrous composite pleochasium with flattened or cylindrical branches, disc annular, fruit a dehiscent schizocarp.
• Maytenus - shrubs or small trees, inflorescences small fasciculate, fruit a loculicidal deshiscent capsule usually separating in 2 valves.
• Peritassa - lianas to treelets, inflorescence thyrsoid or a composite dichasium, disc short-tubular, anthers dehiscence longitudinal or oblique by ± divergent slits, sometimes slits confluent in the apices, stigma punctual or rarely 3-lobed, fruit a berry.
• Plenckia - often dioecious, trees or shrubs, leaves alternate, fruit a samara with an apical wing, surrounding seed, elongate or oblong, symmetrical, and eventually septicidally dehiscent capsule with age.
• Prionostemma - lianas, scabrous, red exudate when cut, inflorescence a composite dichasium, petals fimbriate, disc patelliform, fruit a dehiscent schizocarp.
• Pristimera - lianas, inflorescence a glabrous composite dichasium, rare a pleochasium, with cylindrical to or 4-angled branches, disc annular to short-tubular, fruit a dehiscent schizocarp.
• Quetzalia - shrubs, small trees or vines, fruit capsular, septicidally dehiscent, 2-valved, seed with reddish sarcostesta.
• Salacia - lianas to trees, inflorescence fasciculate, thyrsoid or corymbose, disc annular-pulvinate, pulvinate, annular, cupuliform, cylindric, or patelliform, anthers dehiscence oblique by slits ± divergent or apical, slits confluent or not, stigma punctual, fruit a berry.
• Schaefferia - dioecious shrubs or trees, fruit small dry drupe.
• Semialarium - lianas to treelets, inflorescence a glabrous or pilose composite dichasium, with cylindrical to or 4-angled branches, discpulvinate, fruit a dehiscent obinfundibuliform capsule.
• Tontelea - lianas to trees, inflorescence thyrsoid, disc tubular, anthers dehiscence transversal, stigma 3-lobed, fruit a berry.
• Tetrasiphon -tree or shrub, 4-merous flowers with 1 pendulous ovule per locule, fruit a small flattened drupe.
• Tricerma - small trees or shrubs, inflorescences axillary, solitary, fasciculate or condensed thyrsoid, flowers bisexual or unisexual (polygamous), ovule 1 per locule, fruit a 3-locular loculicidal capsule with seeds enveloped by an aril.
• Wimmeria - trees or shrubs, fruit a samara with (2-)3(-4) wings.
• Zinowiewia - trees or shrubs, leaves opposite; fruit like Plenckia , a samara with a lateral-apical wing, oblanceolate or obovate, curved like a sable.

Literature

Important literature

Coughenour, J. M., Simmons, M. P., Lombardi, J. A., Yakobson, K., & Archer, R. H. 2011. Phylogeny of Celastraceae subfamily Hippocrateoideae inferred from morphological characters and nuclear and plastid loci. Molecular Phylogenetics and Evolution 59:   320-330.Görts-van Rijn, A. R. A. & A. M. W. Mennega. 1994. Hippocrateaceae. Fl. Guianas, Ser. A, Phanerogams, 16: 3-81.Hallé, N. 1962. Monographie des Hippocratéacées d'Afrique occidentale. Mém. Inst. Franç. Afrique Noire 64: 1-246.Hallé, N. 1983. Révision des Hippocrateae (Celastraceae): 3. Fruits, graines et structures placentaires. Bull. Mus. Natl. Hist. Nat., B, Adansonia 5: 11-25.Hedin, J. P. T. 1999. Systematic studies of the Neotropical species of Salacia L. (Hippocrateaceae) and its relatives. Thesis, Washington University, Saint Louis.Lombardi, J. A. & A. C. M. Lara. 2003 [2004]. Hippocrateaceae. Pp. 109-122 in M. G. L. Wanderley, G. J. Shepherd, T. S. Melhem, A. M. Giulietti & M. Kirizawa (eds.), Flora Fanerogâmica do Estado de São Paulo 3. FAPESP/RiMa, São Paulo.Lombardi, J. A. 2001. Hippocrateaceae. Flora del Paraguay 36: 1-36. McKenna, M. J., Simmons, M. P., Bacon, C. D., & Lombardi, J. A. 2011. Delimitation of the Segregate Genera of Maytenus s. l. (Celastraceae) Based on Morphological and Molecular Characters. Systematic Botany 36: 922-932.Mennega, A. M. W. 1997. Wood anatomy of the Hippocrateoideae (Celastraceae). IAWA J. 18: 331-368.Miers, J. 1872. On the Hippocrateaceae of South America. Trans. Linn. Soc. London 28: 319-432.Mory, B. 2001. Notes on Crossopetamum, Myginda and Gyminda (Celastraceae) from Cuba. Wildenowia 31: 129-135.Mory, B. 2010. Flora de la República de Cuba: Celastraceae, Serie A, Plantas Vasculares Fasc. 16" 1-80.Peyritch, J. 1878. Hippocrateaceae. Pp. 125-164 in C. F. P. Martius & A. G. Eichler (eds.), Flora brasiliensis 11(1). Frid. Fleischer. Lipsiae.

Simmons, M. P. 2004. Celastraceae. Pp. 29-64 in K. Kubitzki (ed.), The families and genera of vascular plants 6. Springer-Verlag, Berlin.Smith, A. C. 1940. The American species of Hippocrateaceae. Brittonia 3: 341-555.

Celastraceae, N.K.B. Robson, N. Hallé, B. Mathew and R. Blakelock†. Flora of Tropical East Africa. 1994

Morphology General Habit

Trees, shrubs, shrublets or woody climbers, without tendrils, glabrous or with simple hairs, unarmed or with axillary shoots terminating in a spine, sometimes with rubber-like latex (gutta) in various parts appearing as elastic threads when a leaf is broken

Morphology Leaves

Leaves simple, alternate or spiral to subopposite or opposite, sometimes fasciculate on short shoots, entire or with crenate or denticulate to spinose margins, penninerved; stipules small, simple or laciniate, sometimes united by a transverse ridge, usually deciduous, or absent

Morphology Reproductive morphology Flowers

Flowers bisexual, polygamous or unisexual, monoecious or dioecious, actinomorphic, often fragrant, in axillary and/or terminal dichasial or monochasial cymes or panicles or thyrses, sometimes with accessory branches (additional to the normal one in the bract axil), or fasciculate or solitary, usually bracteate; pedicels often articulated

Morphology Reproductive morphology Flowers Calyx

Sepals (3–)4–5(–6), imbricate or rarely valvate in bud, free or united at the base, persistent

Morphology Reproductive morphology Flowers Corolla

Petals (3–)4–5(–6), free or rarely united at the base, imbricate or rarely valvate in bud, usually persistent, sometimes with ventral grooves or hollows or appendages

Morphology Reproductive morphology Flowers Androecium

Stamens (2–)3–5(–10), antisepalous, free or more rarely with filaments partly united to form a tube, inserted outside or on or inside the disc (when present); anthers usually short, (1–)2-thecous, extrorse or introrse, basifixed or dorsifixed or versatile, sometimes deciduous, dehiscing by longitudinal or oblique or horizontal slits; pollen simple or more rarely in tetrads or polyads

Morphology Reproductive morphology Flowers Nectaries

Disc nectariferous, annular, entire or angular or crenulate or lobed or covered with fleshy processes, concave to convex, rarely wholly or partly incorporated in an androgynophore or formed of discontinuous pockets, fleshy or membranous, very rarely absent

Morphology Reproductive morphology Flowers Gynoecium

Ovary free or partly or wholly immersed in the disc, sessile or on a short androgynophore, syncarpous, completely or very rarely incompletely 2–5-locular, or rarely 1-locular by abortion, with 1–many erect or rarely pendulous ovules in 2 rows or rarely superimposed in each locule; styles as many as the locules, free or ± united, or absent; stigmas various, free or ± united

Morphology Reproductive morphology Fruits

Fruit capsular, loculicidal, or of divergent ± dorsally flattened dehiscent mericarps or baccate or drupaceous or dry, indehiscent and sometimes winged

Morphology Reproductive morphology Seeds

Seeds with a fleshy or submembranous brightly coloured aril, or winged with the funicle free from the wing or united to its base, or neither arillate nor winged (usually in indehiscent fruits), with or without endosperm; embryo erect, with cotyledons flat or fleshy, rarely united

Every, J.L.R. (2009). Neotropical Parnassiaceae.

Morphology

Description

Tiny (less than 2 cm broad) annuals or herbaceous perennials (Parnassia townsendii B.L. Rob), forming hemispherical tufts, caulescent or acaulescent , or rhizomatous (P. townsendii), golden brown-red tannin sacs visible in epidermis . Leaves simple , whorled in a basal rosette and/or alternate to subopposite, entire , glabrous , petiolate , margins entire or serrate (P. townsendii), venation acrodromous . Inflorescence 1-flowered (rarely 2) at ends of short, leafy, angular scape. Flowers bisexual , minute (Lepuropetalon spathulatum Elliott), appearing sessile in the basal rosette or terminal ; calyx a campanulate , 5-ribbed floral cup, persistent ; corolla (4-)5 or absent (L. spathulatum), inserted on rim of the calyx cup, linear , white, smaller than and alternating with sepal lobes, fimbricate (P. townsendii); stamens 5, antesepalous, free from perianth , staminodes 5, antepetalous or absent, anthers dehiscing longitudinally; ovary subinferior, superior (P. townsendii), syncarpous, carpels 3-4, 3-locular (1 in P. townsendii), styles 3-4 (1 in P. townsendii), short or absent. Fruit loculicidally dehiscent capsules.  Seeds numerous, minute.

General Description

Notes on delimitation

• Treated here to include the Lepuropetalaceae.
•  Zhang and Simmons asked the question 'Are Parnassiaceae the sister group of Celastraceae, or do they represent an early derived lineage within Celastraceae?' (Zhang & Simmons, 2006).

Number of genera

• Two: the monotypicLepuropetalon and a single species of Parnassia.

Status

• Parnassia townsendii is endemic to western slopes of Sierra Madre Occidental of Chihuahua, Mexico.
• Lepuropetalon spathulatum is a native.

Distribution

Distribution in the Neotropics

• Lepuropetalon spathulatum can be found by looking very carefully in disturbed areas, growing amidst grasses and other small annual herbs, mosses and liverworts, some times in damp and often sandy soil and outcrops.
• Parnassia townsendii is endemic to the western slopes of Sierra Madre Occidental of Chihuahua, Mexico.

Diagnostic

Distinguishing characters (always present)

• Herbs.
• Tannin sacs visible as red lines or dots on the leaves and calyx, especially when dry.
• Flowers hermaphroditic.
• Calyx quincunical and persisting into fruit.
• Syncarpous ovary.
• Fruit a capsule.
• Numerous tiny seeds.

Other important characters

• Staminodes opposite the petals and alternating with the fertile stamen.
• Flowers usually solitary, if not then paired.

Key differences from similar families

• Saxifragaceae do not have staminodes and posses carpels with a stylar beak, which persists into fruit.
• Droseraceae have sticky glandular hairs.
• Hypericacae generally have coloured sap, opposite leaves and numerous stamens.
• Celastraceae are a woody family.

Useful tips for generic identification

Lepuropetalon spathulatum :

• Small, annual herb ca. 2cm broad.
• Winter-growing.
• Blade 2-6mm long, spatulate.
• Flowers inconspicuous.
• Corolla small, sub -microscopic or absent.
• Ovary sub -inferior.
• 3-locular.

Parnassia townsendii :

• Rhizomatous.
• Herbaceous perennial.
• Leaves elliptic to ovate.
• Petals fimbricate.
• Ovary superior.
• Unilocular.
• Flowers September - October.

Literature

Important literature

APG 2. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants. Botanical Journal of the Linnean Society. 141: 399-436.

Berkov, A. 2004. Lepuropetalaceae. In: Smith, N., Mori, S. A., Henderson, A., Stevenson, D. W. & Heald, S. V. (eds.). Flowering Plants of the Neotropics. p. 214. The New York Botanical Garden, Princeton University Press, Princeton.

Bye, R. A. & Soltis, D. E. 1979. Parnassiatownsendii (Saxifragaceae), a Mexican Endemic. The Southwestern Naturalist 24:2 pp. 209-222

Culham, A. 2007. In: Heywood, V.H., R.K. Brummitt, A. Culham and O. Seberg (eds). Flowering plant families of the world, pp. 243-4. Kew: Royal Botanic Gardens, Kew.

Maas, P.J.M. & Westra, L.Y. Th. 2005. Neotropical Plant Families. 3rd ed. p. 166. A.R.G. Gantner Verlag K.G., Ruggell.

Simmons, M.P. 2004. Parnassiaceae. In Kubitzki, K. (ed.), amilies and genera of vascular plants vol. 6. Flowering plants. Dicotyledons. pp. 291-296.  Springer-Verlag, Berlin.

Spongberg, S. A. 1972. The genera of Saxifragaceae in the Southeastern United States. Journal of the Arnold Arboretum 53:4. pp. 409-498.

Stevens, P. F. 2008. Angiosperm Phylogeny Website. Version 9 onwards. http://www.mobot.org/MOBOT/research/APweb/.

Watson, L. & Dallwitz, M. J. 1992 onwards. The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000.

Zhang, L. & Simmons, M.P. 2006. Phylogeny and delimitation of the Celastrales inferred from nuclear and plastid genes. Systematic Botany 31:1. pp. 122-137

Lombardi, J.A. (2009). Neotropical Hippocrateaceae.

Diagnostic

Distinguishing characters (always present)

• Axillary buds glabrous.
• Stipules present and minute.
• Extrastaminal nectariferous disc present.

Other important characters

• Twining lianas.
• Leaves glabrous, rarely scabrous.
• Leaves opposite.
• Stamens and carpels 3.
• Leaves often crenate/serrate.

Key differences from similar families

• Extrastaminal nectariferous disc (intrastaminal in Celastraceae).
• Stipules present (absent in Myrtaceae).
• Axillary buds glabrous, hairs rare anyway (axillary buds pilose and hairs common in Malpighiaceae).
• Superiorovary and dialypetalous (inferior ovary and gamopetalous in Rubiaceae).

Useful tips for generic identification

• Anthodon - lianas, inflorescence a composite dichasium, petals serrate, disc annular, fruit a dehiscent obinfundibuliform capsule.
• Cheiloclinium - lianas to trees, inflorescence a composite dichasium or thyrsoid-paniculate, disc enclosing the pistil, only the 3(-5)-lobed stigmas apparent, stamens in little pockets in the outside of the disc, fruit a berry.
• Cuervea - lianas, inflorescence a monochasium, composite dichasium, or thyrsoid, flowers 8-15 mm diam. at anthesis, disc annular or short-tubular, fruit a dehiscent schizocarp.
• Elachyptera - lianas, inflorescence a glabrous composite pleochasium with branches conspicuously 4-angled, disc annular, fruit a dehiscent schizocarp.
• Hippocratea - lianas, inflorescence a pilose composite pleochasium, young branches and inflorescence branches pilose, discpulvinate, petals barbelate in the apex, fruit a dehiscent schizocarp
• Hylenaea - lianas, inflorescence a glabrous composite pleochasium with flattened or cylindrical branches, disc annular, fruit a dehiscent schizocarp.
• Peritassa - lianas to treelets, inflorescence thyrsoid or a composite dichasium, disc short-tubular, anthers dehiscence longitudinal or oblique by ± divergent slits, sometimes slits confluent in the apices, stigma punctual or rarely 3-lobed, fruit a berry.
• Prionostemma - lianas, scabrous, red exudate when cut, inflorescence a composite dichasium, petals fimbriate, disc patelliform, fruit a dehiscent schizocarp.
• Pristimera - lianas, inflorescence a glabrous composite dichasium, rare a pleochasium, with cylindrical to or 4-angled branches, disc annular to short-tubular, fruit a dehiscent schizocarp.
• Salacia - lianas to trees, inflorescence fasciculate, thyrsoid or corymbose, disc annular-pulvinate, pulvinate, annular, cupuliform, cylindric, or patelliform, anthers dehiscence oblique by slits ± divergent or apical, slits confluent or not, stigma punctual, fruit a berry.
• Semialarium - lianas to treelets, inflorescence a glabrous or pilose composite dichasium, with cylindrical to or 4-angled branches, discpulvinate, fruit a dehiscent obinfundibuliform capsule.
• Tontelea - lianas to trees, inflorescence thyrsoid, disc tubular, anthers dehiscence transversal, stigma 3-lobed, fruit a berry.

Notable genera and distinguishing features

• Hippocratea - young branches, inflorescences, pedicels, sepals and petals pilose, petals barbelate inside in the apex, fruit a dehiscentschizocarp
• Salacia - young branches, inflorescences and pedicels rarely puberulous, petals glabrous, fruit a berry

General Description

Number of genera

25 genera worldwide

12 genera native in the Neotropics;

• Anthodon
• Cheiloclinium
• Cuervea
• Elachyptera
• Hippocratea
• Hylenaea
• Peritassa
• Prionostemma
• Pristimera
• Salacia
• Semialarium
• Tontelea

Status

• All genera in the Neotropics are native

Literature

Important literature

Görts-van Rijn, A. R. A. & A. M. W. Mennega. 1994. Hippocrateaceae. Fl. Guianas, Ser. A, Phanerogams, 16: 3-81.

Hallé, N. 1962. Monographie des Hippocratéacées d'Afrique occidentale. Mém. Inst. Franç. Afrique Noire 64: 1-246.

Hallé, N. 1983. Révision des Hippocrateae (Celastraceae): 3. Fruits, graines et structures placentaires. Bull. Mus. Natl. Hist. Nat., B, Adansonia 5: 11-25.

Hedin, J. P. T. 1999. Systematic studies of the Neotropical species of Salacia L. (Hippocrateaceae) and its relatives. Thesis, Washington University, Saint Louis.

Lombardi, J. A. & A. C. M. Lara. 2003 [2004]. Hippocrateaceae. Pp. 109-122 in M. G. L. Wanderley, G. J. Shepherd, T. S. Melhem, A. M. Giulietti & M. Kirizawa (eds.), Flora Fanerogâmica do Estado de São Paulo 3. FAPESP/RiMa, São Paulo.

Lombardi, J. A. 2001. Hippocrateaceae. Flora del Paraguay 36: 1-36.

Mennega, A. M. W. 1997. Wood anatomy of the Hippocrateoideae (Celastraceae). I. A. W. A. J. 18: 331-368.

Miers, J. 1872. On the Hippocrateaceae of South America. Trans. Linn. Soc. London 28: 319-432.

Peyritisch, J. 1878. Hippocrateaceae. Pp. 125-164 in C. F. P. Martius & A. G. Eichler (eds.), Flora brasiliensis 11(1). Frid. Fleischer. Lipsiae.

Simmons, M. P. 2004. Celastraceae. Pp. 29-64 in K. Kubitzki (ed.), The families and genera of vascular plants 6. Springer-Verlag, Berlin.

Smith, A. C. 1940. The American species of Hippocrateaceae. Brittonia 3: 341-555.

Biggs, N. (2014). Neotropical Malesherbiaceae.

Morphology

Description

Habit: shrubs, subshrubs, or annual or perennial xerophytic herbs, often cyanogenic, erect or procumbent , from 15 cm to 2 m tall; indumentum of simple and glandular trichomes, often with a foetid smell.  Leaves alternate , simple or often deeply 1-10 lobed , sessile or shortly petiolate (some species with long-attenuate leaf base resembling a petiole ), margin serrate or entire ; stipules present.  Inflorescences axillary or terminal , usually racemes or leafy panicles, rarely fasciculate or flowers solitary.  Flowers scented, regular, pedicels often with 2 leaf-like bracts at the base; floral tube chartaceous , tubular, conical, campanulate or funnel-shaped, 10-nerved, 4-48 mm long; perianth and corona inserted at rim of floral tube; sepals 5, imbricate , the margins sometimes lined with glandular hairs; petals 5, white, yellow, greenish, pink, red or dark blue, clawed; a denticulate corona well-developed or absent, when present, then represented by a raised ring of tissue at the base of the perianth ; androgynophore 1.5-13.5 mm long; stamens 5, exerted, filaments inserted on the androgynophore below the ovary , anthers 2-locular, dorsifixed, dehiscing longitudinally; ovary superior , 1-locular, with 3-4 carpels, styles 3, free , filiform , stigmas generally extending beyond anthers.  Fruits stipitate capsules, enclosed by the persistent floral tube, dehiscing with 3-4 valves.  Seeds 1 to many, ovoid , exarillate, reticulately ridged, pitted, with an oily endosperm .

Distribution

Distribution in the Neotropics

Malesherbiaceae is a monogeneric family with 24 species and 5 varieties (Gengler-Nowak, 2004) endemic to the Pacific coastal desert and adjacent arid Andes of Chile, Peru and Argentina. The family is distributed throughout Chile.  Northwards from the Atacama Desert, it extends to the Andean valleys east of Lima and to the central Peruvian Andes. The species grow in mediterranean, desert and dry montane habitats of varying aridity, dry river valleys and in the coastal fog zones and lomas vegetation of Peru (Kubitzki, 2007). Most species prefer rocky, sunny areas inhabited by few other plants.

• Malesherbia lactea Phil. is a low rhizomatous perennial of very arid regions of the Atacama desert, Chile and adjacent Argentina, growing at an altitude of 2500 - 3570 m.
• The annualherbM. humilis Poepp. occurs from coastal to pre-Andean Chile and is one of the lowest altitude species. (Kubitzki 200).

General Description

Number of genera

One - Malesherbia Ruiz. & Pav.

Status

Native, endemic.

General notes

• In Peru Malesherbia scarlatiflora Gilg is a popular cure-all herb sold in Lima. A tea made from M. ardens J.F.MacBr. has been used to treat influenza. M. tubulosa (Cav.) J.St.-Hil. is sometimes used to treat bruises (Gengler-Nowak, 2004).
• Malesherbiaceae, like Turneraceae and Passifloraceae, possess poisonous cyanogenic glycosides, and several species from Chile and Argentina have been reported to be fatal to goats (Gengler-Nowak, 2004.
• Some Malesherbia species could be of ornamental value but their cultivation is difficult (Ricardi 1967).  M. linearifolia (Cav.) Poir. and M. lirana Gay have, however, been cultivated at the Royal Botanic Gardens, Kew in the past according to the Living Collections Database.
• The Argentinian vernacular names include "Aroma del campo" for M. humilis; "Margarita" and "Aroma de la montana" for M. lirana (Espinar, 1995).

Notes on delimitation

The Angiosperm Phylogeny Group (APG III website, Stevens 2001) placed Malesherbia as subfamily Malesherbioideae along with Turneroideae in the Passifloraceae in the Malpighiales. Kubitzki (2007), however, treated Malesherbiaceae as a separate family.  Takhtajan (2009) and Cronquist (1981) also consider Malesherbiaceae as a distinct family related to Passifloraceae and Turneraceae in theViolales (Passiflorales). Cronquist (1981) suggests that these three families have a common origin in or near the Flacourtiaceae.  The three families Malesherbiaceae, Turneraceae, and Passifloraceae share important characters such as an extrastaminal corona, although this is not as obvious in Malesherbiaceae as in Passiflora, exotestal seeds and cyanogenic glucosides.  Kubitzki (2007) included a key to separate the three families.

Diagnostic

Useful tips for generic identification

• For a key to the species with illustrations see Ricardi (1967). Espinar (1995) includes a key to the species in Argentina; and Gengler-Nowak (2003) contains a key to sections.

Notable genera and distinguishing features

The genus is divided into five sections by Gengler-Nowak, 2003:

• Cyanpetala - shrubby species with dark blue or purple flowers, in Central and North Chile;
• Albitomenta - perennials covered with white tomentose trichomes, with white sometimes violet tinted flowers, in the high Andes and foothills of Chile and adjacent Argentina, above 3500 m;
• Parvistella - subshrubs, leaves pinnately lobed, flowers greenish, in coastal Chile and Argentina;
• Malesherbia - shrubs with colourful flowers, yellow, red and orange and striking coronas, mostly in Peru, some in North Chile;
• Xeromontana - shrubs, densely pubescent with white, light blue or greenish flowers from montane and very arid areas of Chile and extreme western Argentina.

Distinguishing characters (always present)

• Plants usually densely pubescent, the trichomes glandular or simple.
• Flowers with floral tube, corona and androgynophore.
• Fruits loculicidal capsules.

Other important characters

• Glandular trichomes with unpleasant smell often present.

Key differences from similar families

• Malesherbiaceae - tendrils absent; corona often present but not as flamboyant as seen in Passifloraceae; androgynophore usually present; stamens 5; stylodia (stigma branch) inserted beneath the top of ovary; calyx and floral tube persistent in fruit; seeds exarillate.
• Passifloraceae - tendrils present; extrastaminal corona of threads, tubercles or scales, in 1 to many rows; androgynophore usually present; stamens(4)5-8(numerous);  stylodia inserted on top of ovary; seeds arillate.
• Turneraceae - tendrils absent; corona rarely present and then weakly developed; androgynophore absent; stamens 5; stylodia inserted on top of ovary; calyx and corolla separate from developing fruit and falling together; seeds arillate.

Literature

Important literature

Cronquist, A.J. 1981. An Integrated System of Classification of Flowering Plants, p.411-412. Columbia University Press, New York

Espinar, L.A. 1995. Malesherbiaceae. Flora Fanerogámica Argentina 17 (202): 1-4.

Gengler-Nowak, K.M. 2003. Molecular Phylogeny and Taxonomy of Malesherbiaceae. Syst. Bot. 28(2): 333-344

Gengler-Nowak, K. 2004. Malesherbiaceae. In: Smith, N., Mori, S.A., Henderson, A., Stevenson, D.W. and Heald, S.V. (eds.). Flowering Plants of the Neotropics, p. 227-228. The New York Botanical Garden, Princeton University Press, Princeton.

Kubitzki, K. 2007. Malesherbiaceae. In: Kubitzki, K. (ed.). The Families and Genera of Vascular plants vol. IX, pp. 12, 247-249. Springer-Verlag, Berlin.

Ricardi, M. 1961. Estudios en Malesherbiaceae I. Gayana Bot. 3: 5-13.

Ricardi, M. 1965. Estudios en Malesherbiaceae II. Gayana Bot. 12: 3-10.

Ricardi, M. 1967. Revision taxonomica de las Malesherbiaceas. Gayana Bot. 16: 3-139.

Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 12, July 2012. http://www.mobot.org/MOBOT/research/APweb/.

Takhtajan, A. 2009. Flowering  Plants, p. 871. Springer Science &  Business Media.

Watson, L. & Dallwitz, M.J. (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version 3rd March 2009. http://delta-intkey.com

Brexiaceae, N. K. B. Robson. Flora Zambesiaca 4. 1978

Morphology Reproductive morphology Flowers Androecium

Stamens 4–5(6?), antisepalous, free or united with the disk, inserted outside or inside (Roussea) the disk; anthers dithecous, introrse, versatile or dorsifixed, deciduous

Morphology Reproductive morphology Flowers Nectaries

Disk 4–5(6?)-lobed, with the lobes entire or bearing filaments (staminodes?), fleshy, not nectariferous

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, free, sessile, syncarpous, completely or incompletely 4–5(7)-locular, with ¥ or 2 (Ixerba), collateral ovules in each loculus; styles as many as the loculi, completely united; stigmas capitate, lobed to punctiform

Morphology Reproductive morphology Fruits

Fruit loculicidal, capsular (Ixerba) or baccate (Roussea) or drupaceous (Brexia)

Morphology Reproductive morphology Seeds

Seeds exarillate, not winged, with thin layer of endosperm; embryo erect, with cotyledons free, flat

Morphology General Habit

Trees or shrubs, rarely climbing (Roussed), glabrous or rarely with simple hairs (Roussea)

Morphology Leaves

Leaves simple, alternate to subopposite or subverticillate, entire or with crenate to glandular-serrate or spinose-dentate margins, penninerved, rarely glandular beneath (Roussea); stipules present or absent (Ixerba), sometimes united by a transverse ridge

Morphology Reproductive morphology Flowers

Flowers bisexual, actinomorphic, axillary, solitary or in cymes or false umbels or fascicles

Morphology Reproductive morphology Flowers Calyx

Sepals 4–5(6), free or adnate to the ovary base, imbricate or valvate (Roussea), persistent (Roussea) or deciduous

Morphology Reproductive morphology Flowers Corolla

Petals 4–5(6), free or united at the base (Roussea), imbricate or valvate (Roussea), deciduous or persistent (Roussea)

Brexiaceae, B. Verdcourt. Flora of Tropical East Africa. 1968

Morphology General Habit

Trees or shrubs

Morphology Leaves

Leaves coriaceous, simple, alternate or rarely subopposite or subverticillate, entire to spinously toothed; stipules present or absent

Morphology Reproductive morphology Flowers

Flowers axillary, solitary or in cymes or false umbels

Morphology Reproductive morphology Flowers Calyx

Sepals 4–6, imbricate or valvate, persistent or falling

Morphology Reproductive morphology Flowers Corolla

Petals 4–6, imbricate and clawed or valvate, deciduous or persistent

Morphology Reproductive morphology Flowers Nectaries

Disk annular or joined with the staminodes and 5-lobed

Morphology Reproductive morphology Flowers Androecium

Stamens 4–6, hypogynous to perigynous, free; anthers dithecous, opening lengthwise, usually large

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, syncarpous, 4–7-locular with 2–numerous ovules in each locule; style 1, capitate, lobed or punctiform

Morphology Reproductive morphology Fruits

Fruit a capsule, drupe or berry