MAGNOLIIDAE Novák ex Takht.

Takhtajan, Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 51. 4 Feb 1967


[[Magnoliales+Laurales]+[Canellales+Piperales]]


MAGNOLIANAE Takht.

Takhtajan, Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 51. 4 Feb 1967


[Magnoliales+Laurales]


MAGNOLIALES Juss. ex Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 223. Jan-Apr 1820 [’Magnoliae’]

Fossils Fossil flowers, which may be assigned to Magnoliales or closely related lineages, are comparatively frequent in some Turonian layers (the Sayreville Turonian angiosperm flora). A cupular receptaculum is present in several of these fossil remnants. Endressinia brasiliana, from the Upper Aptian to the Lower Albian of Brazil, consists of leafy shoots with pedicellate apocarpous flowers. The shallow receptacle have tepals along their margin, a spiral of presumed glandular staminodia inside these and central spirally arranged plicate carpels.

Habit Usually bisexual (sometimes monoecious or dioecious), evergreen or deciduous trees, shrubs or lianas (rarely dwarf-shrubs or suffrutices). Often aromatic.

Vegetative anatomy Phellogen ab initio usually superficial. Medulla septate, with sclerenchymatous diaphragmata. Primary stem with eustele or (pseudo)siphonostele with separate vascular bundles or with continuous vascular cylinder. Vessel elements with usually scalariform or simple (rarely reticulate) perforation plates; lateral pits alternate, scalariform or opposite, usually simple or reduced bordered pits. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple or bordered pits, septate or non-septate, or absent. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse, diffuse-in-aggregates or banded, or paratracheal scanty vasicentric, scalariform, reticulate, banded, or absent. Secondary phloem stratified. Sieve tube plastids Ss or Psc type (rarely Pcs or Pcsf type). Nodes 3–≥5(–11):3–≥5(–11), trilacunar, pentalacunar or multilacunar with three, five or up to eleven leaf traces (sometimes 1:1, unilacunar with one trace). Secretory cavities with resins present or absent. Wood rays sometimes with oil cells or crystals. Idioblasts with ethereal oils often present. Asterosclereids often present. Silica bodies often abundant in xylem walls. Calciumcarbonate as prismatic or acicular crystals, styloids, druses or crystal sand often present.

Trichomes Hairs unicellular or multicellular, usually uniseriate, simple or branched (sometimes stellate or dendritic, rarely peltate-lepidote, candelabra-shaped, T-shaped or fimbriate), or absent.

Leaves Alternate (spiral or distichous), simple, usually entire (rarely lobed), with conduplicate or convolute ptyxis. Stipules usually absent (sometimes early caducous, ocreate, open just opposite petiole, enclosing young leaf); leaf sheath absent. Petiole vascular bundle transection arcuate (with an adaxial plate consisting of vascular tissue) or annular (sometimes bicollateral). Palisade parenchyma present. Venation usually pinnate, eucraspedodromous or brochidodromous (rarely palmate). Stomata usually paracytic (rarely anomocytic or actinocytic). Cuticular wax crystalloids as platelets or rodlets (often transversely ridged Aristolochia type crystalloids), chemically characterized by presence of palmitone (hentriacontan-16-one) and absence of nonacosan-10-ol. Lamina often gland-dotted. Mesophyll usually with idioblasts (secretory cavities) containing ethereal oils, resin or mucilage. Sclerenchymatous idioblasts with branched sclereids of various kinds (also asterosclereids) or fibres often frequent. Leaf margin entire.

Inflorescence Terminal, axillary or supra-axillary, cymose, paniculate or fasciculate, or flowers solitary axillary, terminal or pseudo-axillary (terminal on small axillary short shoots). Floral prophylls (bracteoles) usually single (rarely paired).

Flowers Actinomorphic. Usually hypogyny (rarely half epigyny, with urceolate, campanulate or infundibuliform receptacle and without tepals). Outer tepals (two or) three (or four), with valvate or imbricate aestivation, spiral or whorled, sepaloid or petaloid, free or connate at base; inner tepals three to numerous, with valvate or imbricate aestivation, petaloid, spiral or whorled, usually free (rarely connate at base). Nectary usually absent. Disc absent.

Androecium Stamens c. 20 to more than 200, laminar (foliaceous), spiral, not differentiated into filament and anther, with separate microsporangia embedded in distal part, or differentiated into filament and anther. Filaments when present usually free from each other (rarely connate at base), free from tepals; filaments often with three vascular bundles. Anthers when present basifixed, non-versatile, tetrasporangiate, sometimes with transversely septate thecae, usually extrorse (rarely latrorse or introrse), longicidal (dehiscing by longitudinal slits) or valvicidal (dehiscing by valves), sometimes connate into synandrium; or microsporangia four, usually adaxial or lateral (sometimes abaxial), usually introrse or latrorse (sometimes extrorse), longicidal (dehiscing by longitudinal slits) or valvicidal (dehiscing by valves). Tapetum secretory. Staminodia extrastaminal, intrastaminal, or absent.

Pollen grains Microsporogenesis simultaneous, or intermediate between simultaneous and successive (cytoplasm partially divided after first meiotic nuclear division by callose furrow, developing centripetally and orthogonally relative to spindle; growth of callose furrow subsequently ceasing and later completed after second nuclear division; cell plate not formed), or sometimes probably successive. Pollen grains usually monosulc(ul)ate (anasulcate) or inaperturate (rarely trichotomosulcate, zonizonasulc[ul]ate or ulcerate), boat-shaped, shed as monads, bicellular at dispersal. Exine tectate or intectate, with granular or intermediary infratectum, microperforate, rugulate or foveolate, gemmate, clavate, echinate or psilate.

Gynoecium Carpels usually ten to numerous (rarely one to three), spiral or whorled, usually free (sometimes connate or paracarpous); carpel plicate to conduplicate (sometimes basally ascidiate and not differentiated into ovary and style), usually postgenitally partially or entirely occluded by fusion and secretion, with secretory canal often open and filled by secretions (sometimes without canal). Ovary usually superior (rarely semi-inferior), unilocular to 15-locular or more. Stylodium single, simple. Stigma capitate, often decurrent, papillate, Dry or Wet type. Nectar sometimes secreted from exposed carpel surfaces. Pistillodium absent.

Ovules Placentation laminar, marginal, parietal, basal, subbasal or lateral. Ovules one to numerous per carpel, usually anatropous (rarely orthotropous or hemiorthotropous), ascending or pendulous, apotropous, usually bitegmic (rarely tritegmic), crassinucellar. Micropyle bistomal or endostomal (rarely exostomal). Funicular obturator sometimes present. Nucellar cap absent. Megagametophyte monosporous, Polygonum type. Antipodal cells and/or synergids ephemeral or persistent. Endosperm development usually cellular (sometimes nuclear). Chalazal endosperm haustoria sometimes present. Embryogenesis onagrad (Myosurus type) or irregular.

Fruit A usually fleshy (sometimes leathery or more or less woody), dehiscent or indehiscent, apocarpous follicular fruit or a cluster of follicles (multifolliculus), or a dry or fleshy pseudosyncarp, as ripe usually dehiscing and with sarcotestal seeds pendant in their funiculi, a loculicidal capsule or an assemblage of samaras (rarely berries or dry follicles).

Seeds Aril usually absent. Seed coat mesotestal or endotestal. Testa often vascularized. Exotesta and mesotesta sometimes as a multiplicative arilloid oily sarcotesta. Endotesta multi-layered, often with crystals. Tegmen unspecialized, usually crushed (often irregularly ruminate). Perisperm usually not developed. Endosperm usually copious, oily, often irregularly ruminate. Embryo small, straight, usually undifferentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology x = 7–10, 12, 19

DNA A deletion of 30 bp (corresponding to 10 amino acids) present in PI-derived motif in nuclear gene AP3 in most Magnoliales (not in Myristicaceae).

Phytochemistry Flavonols (kaempferol, quercetin), flavones (velutin, 5-desoxyflavone, 5-desoxyisoflavone), flavanonols (taxifolin), diarylpropanes (1,3-diarylpropanes, 1,3-diaryl-2-hydroxypropanes) and other flavonoids, cyanidin, diterpenes (kauranes, clerodanes etc.), triterpenoids (tetracyclic etc.), oleanolic acid derivatives, sesquiterpenes, sesquiterpene lactones, oxyphenols and other aromatic substances, proanthocyanidins, caffeic acid, tannins, aporphine alkaloids (aporphines, oxoaporphines, etc.), benzylisoquinoline alkaloids (morphine etc.), protoberberine alkaloids (berberines etc.), C-methylated alkaloids, indole alkaloids (tryptamine), polyketide alkaloids (e.g. hallucinogenic pyridine alkaloids), quercetin glycosides, cyanogenic glycosides (triglochinin etc.), myristicin, lignans (veraguensin, dihydrocubebin), neolignans (aryltetralin, diaryltetrahydrofurans, etc.), polyketides (alkylsalicylate, acylfloroglucinol, arylalkylbutyrolactones, antibacterial acetogenins), nitrophenyl ethan, myo-inisitol, galbacin, licarin A, and amides present. Ellagic acid, gallo- and ellagitannins not found.

Systematics Myristicaceae are sister to the remaining Magnoliales. Features interpreted by Stevens (2001 onwards) and others as potential synapomorphies of Magnoliales except Myristicaceae, are, e.g., the following: primary stem containing eustele with often separate vascular bundles; wood rays wide; flowers solitary and large; floral receptacle sometimes well developed, usually with cortical vascular bundles; androecium consisting of a large number of spirally arranged stamens; filaments (when present) three-veined; thecae separate and embedded in broad foliaceous filaments; connective often prolonged; gynoecium often consisting of spirally arranged carpels; funicular obturator sometimes present; testa sometimes multiplicative; and a 30 bp (corresponding to ten amino acids) in the PI-derived motif of the nuclear AP3 gene.

Degeneria (Degeneriaceae) and Galbulimima (Himantandraceae) share the characters: axillary flowers; annular outer integument; indehiscent fruit; and x=12. Moreover, Annonaceae and Eupomatia (Eupomatiaceae) have the following characters in common: 8- to 15-seriate wood rays; stem leaves spirally arranged; arcuate petiole bundle transection; presence of inflorescence; baccate fruit; and fibrous mesotesta.

These four clades have usually: valvate anthers and intrastaminal staminodia; intectate pollen grains with psilate sexine and granular infratectum; differentiated pollen tube transmission tissue; and indehiscent fruit. However, molecular data etc. recover Magnoliaceae as sister-group to [Degeneria+Galbulimima] and this clade as sister to [Annonaceae+Eupomatia].

Phylogeny of Magnoliales based on morphological and DNA sequence data (Qiu & al. 1999; Sauquet & al. 2003).

ANNONACEAE Juss.

( Back to Magnoliales )

de Jussieu, Gen. Plant.: 283. 4 Aug 1789 [’Anonae’], nom. cons.

Annonales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 223. Jan-Apr 1820 [‘Anoneae’]; Hornschuchiaceae J. Agardh, Theoria Syst. Plant.: 65. Apr-Sep 1858 [’Hornschuchieae’]; Monodoraceae J. Agardh, Theoria Syst. Plant.: 126. Apr-Sep 1858 [’Monodoreae’]; Annonanae Doweld, Tent. Syst. Plant. Vasc.: xxiv. 23 Dec 2001

Genera/species 105–106/2.345–2.360

Distribution Pantropical, with their highest diversity in tropical parts of the Old World; Asimina also in temperate eastern North America.

Fossils Fossilized seeds (e.g. Anonaspermum) are fairly abundant in Early Cenozoic layers of Europe and the Middle East (e.g. the Early Eocene London Clay of England). Foveomorphomonocolpites is a Maastrichtian to Paleocene pollen fossil of presumed annonacean affinity. Likewise, the fossil flower of Futabanthus asamigawaensis from the Early Coniacian of Japan possibly represents an early Annonaceae.

Habit Usually bisexual (sometimes dioecious, rarely monoecious), evergreen or deciduous trees, shrubs or lianas. Often aromatic.

Vegetative anatomy Phellogen ab initio usually superficial. Primary stem usually with eustele or (pseudo)siphonostele with continuous vascular cylinder. Cortical vascular tissue present. Medulla and cambium usually stratified. Medulla often septated by diaphragmata. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits usually alternate (rarely opposite?), bordered pits. Imperforate tracheary xylem elements fibre tracheids with simple and/or bordered pits, non-septate (also vasicentric tracheids). Wood rays usually multiseriate (rarely uniseriate), homocellular or somewhat heterocellular. Axial parenchyma apotracheal, banded, or paratracheal scanty vasicentric, scalariform, reticulate, or banded. Secondary phloem usually stratified into hard fibrous and soft parenchymatous layers. Sieve tube plastids usually Pcs type (in some species of Xylopia Pcsf type), with large protein crystalloids, often with protein fibrils. Nodes usually 3:3, trilacunar with three leaf-traces (median trace split; lateral traces often originating deep inside internode; sometimes 1:1, unilacunar with a single trace). Parenchyma often storied. Wood in some species fluorescent. Medulla and cortex usually with sclerenchyma cells. Wood rays sometimes with oil cells or crystals. Secretory cavities with resins present or absent. Prismatic crystals and druses sometimes abundant (in some species also styloids, crystal sand and/or acicular crystals).

Trichomes Hairs uniseriate, multicellular, simple or branched, with usually pointed terminal cell, stellate (peltate-lepidote in, e.g., Duguetia), or absent.

Leaves Alternate (usually distichous), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation pinnate, eucraspedodromous or brochidodromous. Stomata usually paracytic (sometimes allelocytic etc.). Cuticular wax crystalloids as platelets or rodlets? Abaxial domatia (as pockets or hair tufts) present in some genera. Lamina often gland-dotted. Mesophyll usually with idioblasts (secretory cavities) with ethereal oils, resin or mucilage. Sclerenchymatous idioblasts with branched sclereids (also asterosclereids) or fibres abundant, or absent. Calciumoxalate crystals present. Leaf margin entire.

Inflorescence Terminal, axillary or supra-axillary (inserted between two nodes), cymose, or flowers solitary axillary. Floral prophyll (bracteole) usually single, median, adaxial (bracteoles rarely paired, lateral).

Flowers Actinomorphic. Hypogyny. Outer tepals (two or) three (or four), with open, valvate or imbricate aestivation, sepaloid, often thick and fleshy, in one whorl, free or connate at base; inner tepals three to six (to twelve), with open, valvate or imbricate aestivation, petaloid, often thick and fleshy, usually in one or two (rarely three) series, usually free (rarely connate at base). Adaxial side of inner tepals often with osmophores or perfume glands, often in collateral pairs, sometimes with nectaries in bands at base. Disc present.

Androecium Stamens c. 25 to several hundred, spiral, or three to six (to 15), whorled, with separate thecae embedded in distal part. Filaments usually free (rarely connate at base), free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, sometimes with transversally septate thecae, usually extrorse (rarely latrorse or introrse), longicidal (dehiscing by longitudinal slits) or valvate (dehiscing by valves); connective usually peltate-truncate, often widened. Tapetum secretory, with usually binucleate (sometimes tri- or quadrinucleate) cells. Extrastaminal staminodia present in some species.

Pollen grains Microsporogenesis at least sometimes intermediate between simultaneous and successive (similar to Magnoliaceae). Pollen grains inaperturate or monosulc(ul)ate (rarely zonizonasulculate, with two circular sulculi, or ulcerate), usually shed as monads (rarely tetrads or polyads), bicellular at dispersal. Exine tectate or intectate, with granular infratectum, microperforate, perforate, microreticulate, foveolate, gemmate, clavate, echinate or psilate; endexine sometimes lamellate.

Gynoecium Carpels usually ten to c. 100 (rarely one to three), spiral or whorled (when three, then antesepalous), usually free (sometimes partially or entirely connate or paracarpous; in some species fused when mature); carpel plicate, usually postgenitally partially closed, with secretory canal open and filled by secretions; extragynoecial compitum present (internal and partial compitum rarely present). Ovary superior, unilocular to multilocular (bilocular to 15-locular or more). Stylodia single, simple, short, thick. Stigma capitate to U-shaped, papillate, Wet type. Pistillodium absent.

Ovules Placentation parietal, (sub)basal or lateral. Ovules one to numerous per carpel (or rarely per ovary), anatropous, ascending, apotropous, usually bitegmic (rarely tritegmic, with a multiplicative seedcoat-forming integument between inner and outer integuments), crassinucellar (outer integument in Monodora relatively thin). Micropyle endostomal. Outer integument at least four cell layers thick, vascularized. Inner integument two or three cell layers thick. Nucellar cap absent. Hypostase present or absent. Chalaza perichalazal. Obturator? Megagametophyte monosporous, Polygonum type. Antipodal cells ephemeral. Endosperm development ab initio cellular. Endosperm haustoria? Embryogenesis onagrad.

Fruit A usually fleshy (sometimes partially lignified), dehiscent or indehiscent, apocarpous follicular fruit or a cluster of follicles, multifolliculus etc. (rarely berries; in Anaxagorea dry follicles).

Seeds Seeds often relatively large, usually without wings (in ‘Richella’ winged). Aril absent or present (e.g. Anaxagorea, Annona, Canangium and Xylopia with a micropylar aril). Seed coat mesotestal. Testa multiplicative, ruminate (vascularized?). Exotesta unspecialized. Mesotesta a fibrous (consisting of crushed longitudinal and/or transverse fibres) sarcotesta. Endotesta with crystals (with thin-walled, elongate fibres); endotestal plug often present. Tegmen crushed, usually regularly (in some species irregularly) ruminate. Perisperm usually not developed. Endosperm copious, ruminate with usually regularly spiniform or lamelliform transverse ruminations, hard, oily (with amyloid?). Embryo small, straight, without chlorophyll. Cotyledons two. Germination phanerocotylar or cryptocotylar.

Cytology x = 7, 8, 9 (up to n = 32) – Polyploidy occurring.

DNA Plastid gene infA lost/defunct (Annona). Mitochondrial coxI intron present in Asimina. 30 bp deletion in PI-derived motif in nuclear gene AP3?

Phytochemistry Flavonols (quercetin, myricetin), cyanidin, diterpenes (kauranes, clerodanes etc.), triterpenoids (tetracyclic etc.), tannins, proanthocyanidins, aporphine alkaloids (aporphines, oxoaporphines, etc.), benzylisoquinoline alkaloids (morphine etc.), protoberberine alkaloids (berberine), azaphenanthrene alkaloids, C-methylated alkaloids, cyanogenic compounds, polyketides (antibacterial acetogenins), and nitrophenyl ethan present. Sesquiterpenes? Lignans? Neolignans? Ellagic acid not found.

Use Ornamental plants, fruits (Annona atemaya, cherimoya, ilarma, Artabotrys, Rollinia), spices, perfumes (Cananga odorata, Mkilua fragrans, Artabotrys odoratissima), medicinal plants, timber.

Systematics The sister-group of Annonaceae is Eupomatia (Eupomatiaceae).

Anaxagoreoideae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 32. 18 Apr 2012

1/c 30. Anaxagorea (c 30; Sri Lanka, Indochina, West Malesia, the Moluccas, tropical Central and South America). – Shrubs or small trees. Hairs uniseriate, with rounded terminal cell. Petiole vascular bundle transection annular, with three separate bundles often present in petiole and with adaxial bicollateral xylem plate in midvein. Stomata often allelocytic. Receptacle without vascular bundles. Stamens spirally arranged. Connective not peltate. Inner staminodia sometimes present. Pollen grains with lamellate endexine. Intine with massive inner layer having a channelled structure. Ovules two basal. Fruit a follicle, explosively dehiscing. Aril absent or rudimentary. Endotesta aerenchymatous (with only the tegmen being involved in the ruminations) and possessing idioblastic oil globules. x = 8. – Anaxagorea is sister to the remaining Annonaceae based on combined morphological and molecular data. Scharaschkin & Doyle (2005) suggested a West Gondwanan Eocene origin for Anaxagorea.

[Ambavioideae+[Annonoideae+Malmeoideae]

Hairs stellate. Internal staminodia absent. Endosperm with amyloid staining brownish violet.

Ambavioideae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 32. 18 Apr 2012

9/54. Meiocarpidium (1; M. lepidotum; tropical West Africa); Tetrameranthus (7; T. duckei, T. globuliferus, T. guianensis, T. laomae, T. macrocarpus, T. pachycarpus, T. umbellatus; tropical South America), Mezzettia (5; M. curtisii, M. herveyana, M. leptopoda, M. parviflora, M. umbellata; the Malay Peninsula to the Moluccas), Ambavia (2; A. capuronii, A. gerrardii; Madagascar), Cleistopholis (4; C. glauca, C. myristiciflora, C. patens, C. staudtii; tropical Africa), Lettowianthus (1; L. stellatus; Kenya, Tanzania), Drepananthus (26; tropical Asia to Fiji), Cananga (2; C. brandisia, C. odorata; tropical Asia to tropical Australia), Cyathocalyx (7; C. annamensis, C. globosus, C. harmandii, C. magnifructus, C. martabanicus, C. sumatranus, C. zeylanicus; tropical Asia to islands in southwestern Pacific). – Pantropical. Anthers with tongue-shaped connective. Pollen grains heteropolar-sulcate, with granular infratectum. Carpellary stipe articulated. Ovules usually two per carpel. A third integument present between the ordinary two integuments. x = 7.

[Annonoideae+Malmeoideae]

Annonoideae Raf., Anal. Nat.: 175. Apr-Jul 1815 [‘Annonidia’]

Bocageeae Endl., Gen. Pl.: 830. Jun 1839. 8/65. Mkilua (1; M. fragrans; Kenya, Tanzania), Porcelia (7; P. macrocarpa, P. magnifructa, P. mediocris, P. nitidifolia, P. ponderosa, P. stenbachii, P. venezuelensis; Central America, tropical South America), Cymbopetalum (27; Mexico to tropical South America), Cardiopetalum (3; C. calophyllum, C. plicatum, C. surinamense; tropical South America), Hornschuchia (10; eastern Brazil), Trigynaea (c 10; northern tropical South America), Bocagea (2–3; B. longipedunculata, B. viridis; eastern Brazil, possibly extinct), Froesiodendron (3; F. amazonicum, F. longicuspe, F. urceocalyx; Amazonia in Brazil). – Xylopieae Endl., Gen. Pl.: 831. Jun 1839. 2/c 260. Artabotrys (102; tropical regions in the Old World to tropical Australia), Xylopia (c 160; pantropical). – Duguetieae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 33. 18 Apr 2012. 5/100. Pseudartabotrys (1; P. letestui; tropical West Africa), Letestudoxa (3; L. bella, L. glabrifolia, L. lanuginosa; tropical West and Central Africa), Fusaea (3; F. decurrens, F. longifolia, F. peruviana; tropical South America), Duguetia (c 90; Central America, tropical South America, tropical West Africa), Duckeanthus (1; D. grandiflorus; tropical South America). – Guatterieae Hook. f. et Thomson, Fl. Ind. 1: 92, 126. Jul 1855. 1/c 210. Guatteria (c 210; southern Mexico, Central America, the West Indies, tropical South America). – Annoneae Endl., Gen. Pl.: 833. Jun 1839. 7–8/c 340. Anonidium (5; A. brieyi, A. floribundum, A. letestui, A. mannii, A. usambarense; tropical Africa), Neostenanthera (4; N. gabonensis, N. hamata, N. myristicifolia, N. robsonii; tropical Africa, tropical South America), Goniothalamus (c 135; India, Sri Lanka, Southeast Asia, West Malesia, New Caledonia), Annona (c 165; tropical Africa, tropical America), Disepalum (9; Southeast Asia, West Malesia), Asimina (17; eastern United States; incl. Deeringothamnus?), Deeringothamnus (2; D. pulchellus, D. rugelii; Florida; in Asimina?), Diclinanona (3; D. calycina, D. matogrossensis, D. tessmannii; eastern Peru, Brazil). – Monodoreae Baill., Hist. Pl. 1: 263, 288. Aug-Dec 1868. 11/c 90. Ophrypetalum (1; O. odoratum; tropical East Africa), Sanrafaelia (1; S. ruffonammari; Tanzania), Mischogyne (2; M. elliotianum, M. michelioides; tropical Africa), Uvariodendron (15; tropical Africa), Monocyclanthus (1; M. vignei; tropical West and Central Africa), Uvariopsis (18; tropical Africa), Isolona (20; tropical Africa, Madagascar), Monodora (14–16; tropical Africa), Asteranthe (3; A. asterias, A. lutea, A. trollii; tropical East Africa), Hexalobus (5; H. bussei, H. crispiflorus, H. monopetalus, H. mossambicensis, H. salicifolius; tropical and southern Africa, Madagascar), Uvariastrum (9; tropical Africa). – Uvarieae Hook. f. et Thomson, Fl. Ind. 1: 91, 92. 1-19 Jul 1855. 14/465–470. Dielsiothamnus (1; D. divaricatus; tropical East Africa), Mitrella (9; Malesia to tropical Australia), Fissistigma (c 65; tropical regions in the Old World), Uvaria (185–190; tropical Africa, Madagascar, tropical Asia to Japan and tropical Australia), Toussaintia (4; T. congolensis, T. hallei, T. orientalis, T. patriciae; tropical Africa), Sphaerocoryne (4; S. affinis, S. blanfordiana, S. diospyrifolia, S. gracilis; tropical Africa), Monanthotaxis (c 95; tropical and subtropical Africa, Madagascar), Friesodielsia (c 38; tropical and subtropical Asia), Dasymaschalon (c 27; tropical and subtropical Asia), Desmos (25; India, Southeast Asia to tropical Australia and islands in western Pacific), Afroguatteria (2; A. bequaertii, A. globosa; tropical Africa), Cleistochlamys (1; C. kirkii; tropical East Africa), Melodorum (12; Southeast Asia, Malesia to tropical Australia), Pyramidanthe (1; P. prismatica; West Malesia).

Malmeoideae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 34. 18 Apr 2012

Piptostigmateae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 34. 18 Apr 2012. 6/35. Annickia (8; tropical Africa), Greenwayodendron (2; G. oliveri, G. suaveolens; tropical Africa), Mwasumbia (1; M. alba; coastal Tanzania), Sirdavidia (1; S. solannona; Gabon), 'Piptostigma' (14; tropical Africa; non-monophyletic), Polyceratocarpus (9; tropical Africa). – Malmeeae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 34. 18 Apr 2012. 13/180. Unonopsis (48; Central America, tropical South America), Bocageopsis (4; B. canescens, B. mattogrossensis, B. multiflora, B. pleiosperma; Central America), Onychopetalum (2; O. amazonicum, O. periquino; Peru, Brazil), Malmea (7; M. depressa, M. dielsiana, M. dimera, M. guianensis, M. manausensis, M. obovata, M. surinamensis; southern Mexico, Central America, tropical South America), Pseudoxandra (23; tropical South America), Cremastosperma (29; tropical South America), Mosannona (14; Central America, tropical South America), Ruizodendron (1; R. ovale; Colombia, Ecuador, Peru, Amazonian Brazil, Bolivia), Ephedranthus (7; E. amazonicus, E. boliviensis, E. columbianus, E. dimerus, E. guianensis, E. parviflorus, E. pisocarpus; tropical South America), ‘Oxandra’ (27; Mexico, Central America, the West Indies, tropical South America; polyphyletic), Pseudomalmea (4; P. boyacana, P. darienensis, P. diclina, P. wingfieldii; Central America, northern tropical South America), 'Klarobelia' (12; Central America, tropical South America; non-monophyletic), Pseudephedranthus (2; P. enigmaticus: Guyana, Suriname, Pará in northern Brazil, P. fragrans; southern Venezuela, northeastern Brazil). – Maasieae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 34. 18 Apr 2012. 1/6. Maasia (6; M. discolor, M. glauca, M. hypoleuca, M. multinervis, M. ovalifolia, M. sumatrana; Southeast Asia, Malesia). – Fenerivieae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 35. 18 Apr 2012. 1/9–10. Fenerivia (9–10; Madagascar). – Phoenicantheae X. Guo et R. M. K. Saunders in Sci. Rep. 7: 7323. Aug 2017. 1/2. Phoenicanthus (2; P. coriacea, P. obliqua; Sri Lanka). – Dendrokingstonieae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 35. 18 Apr 2012. 1/3. Dendrokingstonia (3; D. acuminata, D. gardneri, D. nervosa; peninsular Thailand, the Malay Peninsula, Sumatra). – Monocarpieae Chatrou, Pirie, Erkens et Couvreur in Bot. J. Linn. Soc. 169(1): 35. 18 Apr 2012. 1/6. Monocarpia (6; M. blancoi, M. euneura, M. kalimantanensis, M. maingayi, M. marginalis, M. siamensis; Southeast Asia, West Malesia). – Miliuseae Hook. f. et Thomson, Fl. Ind. 1: 147. 1-19 Jul 1855. 23/490–495. Mitrephora (47; Southeast Asia, West Malesia), Platymitra (2; P. arborea, P. macrocarpa; West Malesia), Orophea (c 50; southern India, Hainan, Southeast Asia, Malesia to Moluccas), Trivalvaria (6; T. argentea, T. casseabriae, T. costata, T. macrophylla, T. nervosa, T. rubra; Assam to West Malesia), Pseuduvaria (57; India, Sri Lanka, Southeast Asia, Malesia to New Guinea and tropical Australia), Popovia (26; tropical Asia to tropical Australia), Huberanthus (27; tropical East Africa, Madagascar, the Comoros, southern India, Sri Lanka, Southeast Asia to New Guinea, Queensland, New Caledonia, Fiji), Alphonsea (25; tropical Asia to southern China and Malesia), Miliusa (c 50; Hainan and Malesia to Queensland), Wangia (1; W. sacopetaloides; Yunnan), Phaeanthus (9; India, Sri Lanka, Southeast Asia, Malesia), Sageraea (9; India, Sri Lanka, Southeast Asia to the Philippines), Stelechocarpus (1; S. burahol; Java), Winitia (2; W. cauliflora, W. expansa; Vietnam, peninsular Thailand, the Malay Peninsula, Borneo), ’Polyalthia’ (45–50; tropical regions in the Old World to Queensland; polyphyletic), Marsypopetalum (6; M. crassum, M. littorale, M. lucidum, M. modestum, M. pallidum, M. triste; Hainan, Southeast Asia, West Malesia), Neouvaria (7; N. acuminatissima, N. foetida, N. merrillii, N. parallelivenia, N. sparsistellata, N. telopea, N. viridifolia; West Malesia), Monoon (56; India, Sri Lanka, Ryukyu Islands, Hainan, Southeast Asia, West Malesia, New Guinea, northern Australia), Meiogyne (24; southwestern India, Hainan, Southeast Asia, Malesia to New Guinea, Queensland, New Caledonia, Micronesia, Polynesia), Wuodendron (1; W. praecox; Assam and Yunnan to Vietnam and Cambodia), Tridimeris (2; T. hahniana, T. tuxtlensis; Mexico), Sapranthus (7; S. campechianus, S. isae, S. microcarpus, S. nicaraguensis, S. palanga, S. violaceus, S. viridiflorus; Central America), Desmopsis (28; Mexico, Central America, Cuba, northwestern South America). – Piptostigmateae are sister-group to the rest and Malmeeae successive sister to the remaining Malmeoideae. The clade [Desmopsis+[Tridimeris+Sapranthus]] is proposed to form the Neotropical Sapranthinae (Ortiz-Rodriguez & al. 2016).

One of numerous most parsimonious cladograms of Annonaceae based on DNA data (Chatrou & al. 2012).

Cladogram of Annonaceae based on DNA sequence data (Doyle & Le Thomas 1997b).

DEGENERIACEAE I. W. Bailey et A. C. Sm.

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Bailey et Smith in J. Arnold Arbor. 23: 357. 15 Jul 1942, nom. cons.

Degeneriales C. Y. Wu in Acta Phytotaxon. Sin. 40: 291. 2002

Genera/species 1/2

Distribution Fiji.

Fossils Unknown.

Habit Bisexual, evergreen trees. Aromatic.

Vegetative anatomy Phellogen ab initio superficial. Primary stem containing eustele with separate vascular bundles. Cortical vascular tissue? Medulla septated by diaphragmata (with sclereids). Vessels usually in groups. Vessel elements with scalariform perforation plates; lateral pits scalariform. Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma usually apotracheal diffuse, diffuse-in-aggregates or banded (paratracheal parenchyma rare or absent). Wood non-storied. Tyloses sometimes present in vessels. Secondary phloem stratified into hard fibrous and/or sclerified layers and soft parenchymatous layers. Sieve tube plastids Psc type (PI type), with a small protein crystalloid and approx. 15 starch grains. Nodes 5:5, pentalacunar with five leaf traces. Cells containing ethereal oils present in wood rays. Calciumoxalate druses frequent in phloem ray cells.

Trichomes Hairs absent.

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate?, with medullary bundles as well. Venation pinnate. Stomata paracytic. Cuticular wax crystalloids as platelets. Lamina gland-dotted. Mesophyll with idioblasts containing ethereal oils. Secretory cells absent. Leaf margin entire.

Inflorescence Flowers supra-axillary, solitary (flower inserted between two nodes). Floral prophyll (bracteole) single, median.

Flowers Actinomorphic, large. Hypogyny. Outer tepals usually three (sometimes four), with imbricate aestivation, sepaloid, in a single whorl, fleshy, caducous, free; inner tepals twelve to c. 25, with imbricate aestivation, petaloid, spiral or whorled, fleshy, caducous, free. Nectary absent. Disc absent.

Androecium Stamens c. 20 to c. 30, laminar (foliaceous), spiral, free from each other and from tepals, not differentiated into filament and anther, with separate thecae embedded in distal part. Microsporangia four, abaxial, extrorse, longicidal (dehiscing by longitudinal slits) or valvate (dehiscing by valves); connective strongly prolonged and extended. Tapetum secretory. Staminodia three to approx. ten?, intrastaminal, foliaceous, fewer than fertile stamens, distinctly cucullate (domed above carpel), glanduliferous.

Pollen grains Microsporogenesis simultaneous. Pollen grains monosulcate (anasulcate, with often very elongate sulcus), shed as monads, bicellular at dispersal. Exine tectate, with granular infratectum, psilate.

Gynoecium Pistil composed of one foliaceous carpel (monocarpellate); carpel basally ascidiate, otherwise plicate, not differentiated into ovary and style, not completely closed in bud and early anthesis, postgenitally entirely occluded by fusion alone (without canal). Carpel superior, unilocular. Stigma adaxially deeply decurrent, papillate-hairy, type? Pistillodium absent.

Ovules Placentation laminar-lateral to marginal. Ovules c. 20 to 32 per carpel, anatropous, bitegmic (with lobed integuments), crassinucellar. Micropyle endostomal. Outer integument at least four cell layers thick, vascularized, annular. Inner integument two or three cell layers thick. Funicle long or absent. Funicular obturator distinct. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio cellular. Endosperm haustoria? Embryogenesis?

Fruit A many-seeded, fleshy, ventricidal follicle with hard exocarp.

Seeds Aril absent. Seed coat endotestal. Testa multiplicative (and vascularized?). Exotesta palisade (cells radially elongate), thin-walled. Mesotesta an orange to red and oily sarcotesta. Endotesta multi-layered; endotestal cells with lignified fibrillar endoreticulum. Tegmen degenerating. Perisperm not developed. Endosperm copious, oily, ruminate. Multiseriate massive suspensor present. Embryo small, well differentiated, chlorophyll? Cotyledons usually three (sometimes four). Germination phanerocotylar.

Cytology x = 12 – Isozyme duplication not observed.

DNA Probably a 30 bp deletion in the PI-derived motif in the gene AP3?

Phytochemistry Very insufficiently known. Quercetin glycosides and cyanogenic compounds present. Sesquiterpenes? Alkaloids and proanthocyanidins not found.

Use Unknown.

Systematics Degeneria (2; D. roseiflora: Viti Levu in Fiji; D. vitiensis: Vanua Levu and Taveuni in Fiji).

Degeneria is sister to Galbulimima (Himantandraceae).

EUPOMATIACEAE Orb.

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d’Orbigny, Dict. Univ. Hist. Nat. 5: 511. 29 Mar 1845 [’Eupomatieae’], nom. cons.

Eupomatiales Takht. ex Reveal in Novon 2: 238. 13 Oct 1992

Genera/species 1/3

Distribution New Guinea, eastern and southeastern Australia.

Fossils Ambiguous.

Habit Bisexual, evergreen small trees, shrubs or dwarf-shrubs, with starch-storing tuberous roots (sometimes rhizomatous suffrutices). Often aromatic.

Vegetative anatomy Phellogen? Primary stem with eustele with separate vascular bundles. Cortical vascular tissue? Medulla non-septated, in young stems with secretory cells. Vessel elements with usually scalariform (sometimes reticulate) perforation plates; lateral pits scalariform to opposite, bordered pits. Vestured pits probably absent. Imperforate tracheary xylem elements fibre tracheids with simple and/or (reduced) bordered pits, septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma absent or very rare (apotracheal diffuse, or paratracheal scanty). Secondary phloem stratified? Sieve tube plastids Pcs type (PIb type), with rod-shaped and polygonal protein crystalloids and few starch grains. Sieve tube nuclei with non-dispersive protein bodies? Nodes (6–)7(–11):(6–)7(–11), multilacunar with six to eleven leaf traces. Secretory idioblasts with ethereal oils relatively abundant in medulla and wood rays. Pith parenchyma cells often with single druses.

Trichomes Hairs (on young leaves) simple, ferrugineous, or absent.

Leaves Alternate (distichous), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole on young stem with wings often extending to next lower node; petiole parenchyma with small lumps of crystals. Petiole vascular bundle transection arcuate. Venation pinnate, brochidodromous or eucamptodromous. Stomata usually paracytic (occasionally actinocytic). Cuticular waxes? Lamina often gland-dotted. Mesophyll usually with numerous idioblasts containing ethereal oils. Leaf margin entire.

Inflorescence Terminal or axillary, few-flowered, fasciculate, or flowers solitary axillary. Floral prophyll (bracteole) single, median.

Flowers Actinomorphic, large. Pedicel with approx. three bracts in two rows. Half epigyny. Receptacle urceolate. Tepals absent. Bud surrounded by calyptra, consisting of one or two perfoliate bracts with sclereids, dehiscing like an operculum at anthesis. Nectary absent. Disc absent.

Androecium Stamens c. 20 to more than 100, spiral, with separate thecae embedded in distal part. Filaments short, wide, connate at base, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits); connective somewhat prolonged. Tapetum secretory, parietal. Staminodia c. 40 to c. 80, intrastaminal, petaloid, glandular, curved above gynoecium, connate at base and fused with fertile stamens to a synandrium; inner staminodia with mucilaginous unicellular hairs and covered by an oily secretion.

Pollen grains Microsporogenesis simultaneous. Pollen grains zonizonasulcate (with band-shaped aperture encircling equator), shed as monads, bicellular at dispersal. Exine intectate, with granular ’infratectum’, psilate.

Gynoecium Pistil composed of 13 to c. 70 largely connate, spiral carpels, enclosed in urceolate receptacle (postgenitally fused carpels); carpel (secondarily?) ascidiate, possibly primarily plicate; partial compitum present. Ovary semi-inferior, unilocular (apocarpy). Style absent. Stigma flattened, hairy, decurrent at base, papillate, type? Pistillodium absent.

Ovules Placentation sublaminal (ventral) to marginal. Ovules two to eleven per carpel, anatropous, apotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument at least four cell layers thick. Inner integument two or three cell layers thick. Obturator? Megagametophyte monosporous, Polygonum type. Endosperm development cellular? Endosperm haustoria? Embryogenesis?

Fruit A many-seeded berry-like multifolliculus (secondarily syncarpous, sunken into receptacle). Exocarp with groups of stone cells.

Seeds Aril absent. Seed coat exotestal-mesotestal. Testa multiplicative, ruminate, non-vascularized? Exotestal cells with thickened, non-lignified walls. Mesotesta fibrous. Endotesta unspecialized, non-lignified. Tegmen unspecialized. Exotegmic cells cuboid, somewhat lignified. Endotegmic cells enlarged, crushed. Perisperm not developed. Endosperm copious, oily, ruminate. Embryo small, straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Cytology x = 10 – Isozyme duplication not observed.

DNA Probably a 30 bp deletion in the PI-derived motif in the gene AP3?

Phytochemistry Insufficiently known. Flavones (e.g. velutin), cyanidin and lignans present. Alkaloids? Sesquiterpenes? Proanthocyanidins? Flavonols, ellagic acid and cyanogenic compounds not found.

Use Ornamental plants, carpentries.

Systematics Eupomatia (3; E. barbata: northeastern Queensland, E. bennettii: northeastern New South Wales, eastern Queensland, E. laurina: eastern New Guinea, temperate parts of Victoria to subtropical regions of New South Wales and tropical Queensland).

Eupomatia is sister to Annonaceae. Staminodia and fertile stamens in Eupomatia surround the gynoecium in a perianth-like fashion, whereas the perianth itself is absent. This pattern and the calyptrate bracts surrounding the bud are similar to Galbulimima (Himantandraceae), yet is due to parallel evolution. The urceolate receptacle resembles that present in many Laurales.

HIMANTANDRACEAE Diels

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Diels in Bot. Jahrb. Syst. 55: 126. 27 Nov 1917, nom. cons.

Himantandrales Doweld et Shevyryova in Ann. Bot. (London) 81: 345. Feb 1998

Genera/species 1/2

Distribution Sulawesi to Queensland.

Fossils Unknown.

Habit Bisexual, evergreen trees. Aromatic.

Vegetative anatomy Phellogen ab initio superficial. Primary stem containing eustele with separate vascular bundles. Cortical vascular tissue? Medulla septated by diaphragmata with sclereids. Vessel elements with usually simple (sometimes scalariform) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements fibre tracheids with simple or bordered pits, non-septate. Wood rays uniseriate or multiseriate, homocellular. Axial parenchyma apotracheal diffuse, diffuse-in-aggregates, or banded. Secondary phloem usually stratified into hard fibrous and/or sclereidal layers, and soft parenchymatous layers. Sieve tube plastids S type, with approx. ten starch grains. Sieve elements with non-dispersive fusiform protein bodies. Nodes 3:3, trilacunar with three leaf traces. Heartwood with gum-like substances. Prismatic crystals abundant.

Trichomes Hairs multicellular, stellate, peltate, lepidote or fimbriate, copper-brown.

Leaves Alternate (distichous), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate? Venation pinnate, brochidodromous. Stomata paracytic. Cuticular waxes absent. Mesophyll cells often with sclereids and with idioblasts containing ethereal oils. Secretory cells abundant. Abaxial epidermis with pairs or lumps of crystals. Leaf margin entire.

Inflorescence Flowers usually axillary, solitary (rarely paired). Floral prophyll (bracteole) single, median.

Flowers Actinomorphic, large. Hypogyny. Tepals probably absent. Bud covered by two calyptrae, each consisting of a closed bract (possibly tepal?), dehiscing like an operculum at anthesis. Gynoecium surrounded by androecial receptacle. Nectary absent. Disc absent.

Androecium Fertile stamens 13 to c. 130, band-shaped, spiral, free from each other and from tepals, not differentiated into filament and anther, with separate thecae embedded in distal part. Microsporangia four, adaxial, sunken, valvate (dehiscing by longitudinal valves); connective much prolonged and widened. Tapetum secretory, with binucleate cells. Outer staminodia three to 23, extrastaminal, petaloid; inner staminodia 13 to c. 20, intrastaminal; inner staminodia and some fertile stamens glanduliferous.

Pollen grains Microsporogenesis successive. Pollen grains monosulcate (anasulcate), shed as monads, bicellular at dispersal. Exine intectate, with granular ’infratectum’, psilate.

Gynoecium Carpels six to ten (to 28), spiral, closed, ab initio almost free, in fruit more or less connate; carpel ascidiate, postgenitally fused in style, ‘ovary’ open and filled by secretions; extragynoecial compitum present. ’Ovary’ superior, multilocular. Stylodium single, simple, short. Stigma papillate, Wet type. Gynoecial apex at anthesis covered by mucilage layer (mucilaginous compitum?). Pistillodium absent.

Ovules Placentation apical to laminar-marginal. Ovule usually one (rarely two) per carpel, anatropous, pendulous, apotropous, bitegmic, crassinucellar. Micropyle endostomal. Integuments lobed? Outer integument at least four cell layers thick, vascularized, annular. Inner integument two or three cell layers thick. Obturator? Megagametophyte monosporous, Polygonum type. Antipodal nuclei ephemeral (rarely forming complete cells). Endosperm development cellular? Endosperm haustoria? Embryogenesis?

Fruit A drupaceous secondary syncarp with several usually one-seeded pyrenes.

Seeds Seeds laterally flattened, without long funicle. Seed coat mesotestal. Testa non-multiplicative. Exotesta unspecialized. Mesotesta partly fibrous, partly sarcotestal. Endotesta aerenchymatous, unspecialized. Tegmen unspecialized. Perisperm not developed. Endosperm copious, oily, not ruminate. Embryo small, straight, chlorophyll? Cotyledons two. Germination phanerocotylar.

Cytology x = 12

DNA Probably a 30 bp deletion in the PI-derived motif in the gene AP3?

Phytochemistry Insufficiently known. Sesquiterpenes, polyketide alkaloids (hallucinogenic pyridine alkaloids etc.), and neolignans (aryltetralin and diaryltetrahydrofurans) present. Flavonoids? Proanthocyanidins?

Use Medicinal plants.

Systematics Galbulimima (2; G. baccata: northeastern Queensland, G. belgraveana: Sulawesi, the Moluccas, New Guinea, New Britain, northeastern Queensland).

Galbulimima is sister to Degeneria (Degeneriaceae). Staminodia and fertile stamens in Galbulimima surround the gynoecium in a perianth-like fashion, whereas the perianth itself is either absent or transformed into bract-like calyptrae covering the bud. Hence, the flower resembles that in Eupomatia (Eupomatiaceae), although the similar patterns have evolved in parallel.

MAGNOLIACEAE Juss.

( Back to Magnoliales )

de Jussieu, Gen. Plant.: 280. 4 Aug 1789 [’Magnoliae’], nom. cons.

Liriodendraceae F. A. Barkley in Phytologia 32: 304. 29 Oct 1975

Genera/species 2/210–220

Distribution Southern India, Sri Lanka, eastern Himalayas, Assam, East Asia to the Korean Peninsula and Japan, Southeast Asia, Malesia to New Guinea, New Britain, southeastern North America, Mexico, Central America, the West Indies, scattered areas in northern and central South America from Colombia to Uruguay.

Fossils Fossil wood, leaves and seeds of Magnoliaceae are known from the mid-Cretaceous onwards and abundant in Cenozoic layers in many places in North America, Greenland, Svalbard, Europe and Asia. Fossil pollen grains are rarely found. Fossilized wood is known as Liriodendroxylon, Magnolioxylon and Magnoliaceoxylon, and fossil seeds as Liriodendroidea (winged) and Magnoliaespermum. Fossil leaves (Liriodendrites, Liriodendropsis, Liriophyllum, etc.) are known from Late Cretaceous layers of North America and Europe. Archaeanthus – bilobate leaves, flowers and many-seeded spirally arranged stipitate follicles – from the Albian to the Cenomanian of North America may be assigned to Magnoliaceae, as well as Litocarpon (fruits and seeds) from the Santonian to the Campanian of Canada.

Habit Usually bisexual (in Magnolia Section Kmeria monoecious), evergreen or deciduous trees or shrubs.

Vegetative anatomy Phellogen ab initio superficial. Primary stem with eustele with separate vascular bundles. Cortical vascular tissue present. Medulla often septated by diaphragmata (sometimes with sclereids). Vessel elements with scalariform and/or simple perforation plates; lateral pits scalariform or opposite, usually simple (rarely bordered) pits. Imperforate tracheary xylem elements tracheids or fibre tracheids with simple and/or bordered pits, non-septate, or absent. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty or banded, or absent. Wood fluorescent. Tyloses sometimes abundant. Secondary phloem often stratified into hard fibrous and/or sclereidal layers, and soft parenchymatous layers. Sieve tube plastids S type, with approx. ten starch grains, or Psc type, with a single small protein crystalloid and few starch grains. Nodes ≥5:≥5, pentalacunar or multilacunar with five or more leaf traces. Secretory cells absent. Idioblasts with ethereal oils often present. Silica often abundant in xylem walls. Calciumcarbonate as, e.g., acicular crystals present in many species.

Trichomes Hairs unicellular or multicellular, usually uniseriate, simple (sometimes stellate), or absent.

Leaves Alternate (spiral), simple, usually entire (in Liriodendron lobate), with ? ptyxis. Stipules early caducous, often ochreate, open just opposite petiole, enclosing young leaf; leaf sheath absent. Petiole vascular bundle transection arcuate?, usually with medullary bundles as well (in Liriodendron only as a cylinder). Venation usually pinnate (in Liriodendron palmate). Stomata usually paracytic (rarely anomocytic). Cuticular wax crystalloids as platelets or rodlets (or tubuli?). Lamina sometimes gland-dotted. Mesophyll often with mucilage cells, often with idioblasts with ethereal oils. Sclereids of various types (also asterosclereids) present in leaf cells of some species. Leaf margin entire.

Inflorescence Flowers terminal or pseudo-axillary (in reality terminal on small axillary short shoots), usually solitary (sometimes few together). Bud surrounded by caducous bracts, falling off at anthesis.

Flowers Actinomorphic, large. Hypogyny. Outer tepals three, with imbricate aestivation, sepaloid or petaloid, spiral or whorled, free; inner tepals three to numerous, with imbricate aestivation, petaloid, spiral or whorled, free. Sclerenchymatous diaphragms well developed in Magnolia Section Michelia. Nectary usually absent. Disc absent.

Androecium Stamens c. 50 to more than 200, spiral, laminar (foliaceous), not differentiated into filament and anther, with separate thecae embedded in distal part. Microsporangia four, usually adaxial or lateral (in Liriodendron abaxial), usually introrse or latrorse (in Liriodendron extrorse), longicidal (dehiscing by longitudinal slits) or valvate (dehiscing by valves); connective somewhat prolonged. Tapetum secretory, with multinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis intermediate between simultaneous and successive: cytoplasm after first nuclear division partly dividing by callose furrow, developing centripetally and orthogonally relative to meiotic spindle; growth of callose furrow ceasing and later completed after second nuclear division; cell plate not formed. Pollen grains usually monosulcate (anasulcate; rarely trichotomosulcate), boat-shaped, shed as monads, bicellular at dispersal. Exine tectate or intectate, with intermediary to granular infratectum, microperforate or often rugulate.

Gynoecium Carpels usually numerous (in one species of Magnolia Section Michelia a single carpel, monocarpellary), ab initio free, in some species finally more or less connate, spiral; carpel ascidiate or plicate, postgenitally completely fused, with secretory canal, often stalked (on gynophore). Ovary superior, unilocular or multilocular. Style single, simple. Stigma terminal, often elongated, papillate, Dry type. Nectar in some species of Magnolia secreted from exposed surface of carpels. Pistillodium absent.

Ovules Placentation laminar (ventral, marginal). Ovules (one or) two to twelve (to 16) per carpel, anatropous, pendulous, bitegmic, crassinucellar, with secretory canal. Micropyle usually bistomal (in Magnolia Section Michelia exostomal). Integuments lobed. Outer integument at least four cell layers thick, vascularized. Inner integument two or three cell layers thick. Funicular obturator present in at least Liriodendron. Megagametophyte monosporous, Polygonum type. Antipodal cells and synergids ephemeral. Endosperm development cellular. Endosperm haustoria usually absent (chalazal endosperm haustoria present in, e.g., Magnolia obovata). Embryogenesis onagrad (Myosurus type) or irregular.

Fruit A dry or fleshy pseudosyncarp, often almost cone-shaped, as ripe usually dehiscing abaxially (sometimes both abaxially and adaxially or transversally dehiscent, rarely indehiscent) with fleshy sarcotestal seeds pendant in funiculi by extended annular thickenings on protoxylem, a loculicidal capsule (in Magnolia Section Pachylarnax) or an assemblage of samaras (Liriodendron).

Seeds Aril absent. Seeds in dehiscing fruits pendant in spirally twisted strongly elongated funicular vascular strand. Seed coat endotestal. Testa in Magnolia with simple or tubular pore (indicating passage of vascular strand through sclerotesta). Exotesta and mesotesta usually a multiplicative arilloid oily sarcotesta (absent in Liriodendron), and situated inside scleroendotesta (sclerotic endotesta) with crystals and lignified fibrils in cells. Endotesta multilayered, sclerenchymatous. Tegmen unspecialized, mostly crushed. Perisperm not developed. Endosperm usually copious (in Liriodendron scarce), usually not ruminate. Multiseriate massive suspensor present. Embryo small, usually undifferentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology x = 19 – Polyploidy and aneuploidy occurring. Isozyme duplication indicating paleopolyploidy.

DNA The PI-derived motif of the nuclear gene AP3 has a deletion corresponding to ten amino acids.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavanonols (taxifolin), cyanidin, sesquiterpene lactones, proanthocyanidin, caffeic acid, isoquinoline alkaloids (benzylisoquinoline alkaloids, liriodenine), triglochinin (a cyanogenic glycoside), acuminatin (a bis-phenylpropide), lignans (calopiptin, galgravin, veraguensin, etc.), and neolignans present. Ellagic acid, gallo- and ellagitannins not found. Aluminium possibly accumulated in some species.

Use Ornamental plants, medicinal plants, timber, carpentries.

Systematics Magnolia (210–220; the Himalayas to Japan and West Malesia, eastern United States to tropical South America), Liriodendron (2; L. tulipifera: eastern North America; L. chinense: China).

Magnolia Section Talauma is sister to the remaining species of Magnolia.

Magnoliaceae are sister-group to [[Degeneria+Galbulimima]+[Eupomatia+Annonaceae]].

Maximum likelihood tree of Magnolia s.lat. based on DNA sequence data (Azuma & al. 2001). The neotropical clade Section Talauma is sister-group to the remainder. The former genera Elmerrillia, Kmeria, Manglietia, Michelia and Pachylarnax are nested inside Magnolia.

MYRISTICACEAE R. Br.

( Back to Magnoliales )

Brown, Prodr. Fl. Nov.-Holl.: 399. 27 Mar 1810 [’Myristiceae’], nom. cons.

Myristicales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 235. Jan-Apr 1820 [’Myristiceae’]

Genera/species 20/475–485

Distribution Pantropical, with their largest diversity in tropical Asia.

Fossils Fossil wood from Paleocene layers of Sahara has been described under the name of Myristicoxylon, although it cannot unambiguously be assigned to Myristicaceae. An Early Eocene ruminate seed is known from the London Clay in England and several other Cenozoic fossil Myristicaceae have been found.

Habit Usually dioecious (in Endocomia and some Iryanthera monoecious), usually evergreen (rarely deciduous) trees (rarely shrubs or lianas). Often aromatic.

Vegetative anatomy Phellogen ab initio superficial (also in outer cortex). Primary stem with (pseudo)siphonostele with continuous vascular cylinder. Medulla often septated by diaphragmata. Vessel elements with simple and/or scalariform(-reticulate) perforation plates; lateral pits scalariform, opposite or alternate. Imperforate tracheary xylem elements libriform fibres with usually simple (sometimes bordered) pits, septate or non-septate. Wood rays uniseriate or multiseriate, usually heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty vasicentric, banded, or absent. Tyloses often abundant (sometimes sclerotic). Secondary phloem usually stratified into hard fibrous and/or sclereidal layers, and soft parenchymatous layers. Sieve tube plastids Psc type, with small protein crystalloids and ten or fewer starch grains, or Ss type, with c. 20 starch grains. Sieve element nuclei with non-dispersive protein bodies. Nodes 3:3, trilacunar with three leaf traces. Parenchyma, wood rays and phloem with tanniniferous canals containing reddish to yellowish resinous secretions. Idioblasts with ethereal oils present in many organs. Wood cells sometimes with silica bodies. Nucleus of young sieve element with a single protein crystal (rarely two crystals), which are released into cell lumen during nuclear degeneration and persist in enucleate mature sieve element (such crystals being unique among Magnoliidae).

Trichomes Hairs uniseriate, multicellular, dendritic, stellate (sometimes candelabrous), or T-shaped.

Leaves Alternate (usually distichous), simple, entire, with conduplicate or convolute ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles bicollateral. Venation pinnate. Stomata paracytic. Cuticular wax crystalloids as platelets or rodlets. Lamina sometimes gland-dotted. Mesophyll cells and sometimes epidermal cells with acicular crystals or druses. Branched asterosclereids or fibres present in some genera. Mesophyll with idioblasts containing ethereal oils. Leaf margin entire.

Inflorescence Usually axillary (rarely terminal), panicle or fasciculate. Floral prophyll (bracteole) usually single (occasionally two).

Flowers Actinomorphic, usually small with little developed receptacle. Hypogyny. Tepals (two or) three (to five), with valvate aestivation, in one whorl, petaloid, often fleshy, connate at base and infundibuliform, campanulate or urceolate. Nectary absent. Disc absent.

Androecium Stamens two to c. 40, usually whorled (sometimes spiral). Filaments entirely or partially connate into a tubular, infundibuliform, cupular or globular structure; free from tepals. Anthers longifixed or basifixed, non-versatile, tetrasporangiate, usually extrorse (rarely latrorse), longicidal (dehiscing by longitudinal slits), usually connate into a synandrium (in Myristica with a sterile axial or staminal apex). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis probably successive. Pollen grains monosulcate (sulcoidate, spiraperturate or ulcerate) to almost inaperturate, shed as monads, bicellular at dispersal. Exine usually tectate (rarely intectate), with intermediary infratectum, microperforate (rarely psilate).

Gynoecium Pistil composed of one carpel; carpel ascoplicate (intermediate between ascidiate and plicate), postgenitally completely fused by secretory canal, sometimes shortly stipitate. Ovary superior, unilocular. Style single, simple, usually short (in Brochoneura long), or absent. Stigma simple or bifid, papillate, Dry type? Pistillodium absent.

Ovules Placentation basal or subbasal. Ovule one per ovary, usually anatropous (rarely orthotropous or hemiorthotropous), usually bitegmic (rarely tritegmic with a rudimentary intervening integument), crassinucellar. Micropyle endostomal. Outer integument at least four cell layers thick. Inner integument three to ten cell layers thick. Chalaza pachychalazal. Obturator? Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A fleshy, leathery or woody follicle, usually both adaxially and abaxially dehiscent (rarely indehiscent).

Seeds Seed large, usually with a funicular-exostomal aril (absent in Haematodendron); aril usually large, entire or lobed, hard or fleshy, red, orange or yellow, partially enclosing seed. Seed coat endotestal. Testa vascularized, usually multiplicative. Inner epidermis of outer integument transformed into lignified palisade layer with crystals. Mesotesta stout, unspecialized. Endotesta palisade, with thick-walled, prismatic cells; parts of inner integument transformed into vascularized, massive and ruminate tegmen. Exotegmen with long wood fibres and sclerotic or tracheidal cells. Chalazal end with thick and lignified counter palisade. Perisperm not developed. Endosperm copious, oily, proteinaceous and starchy, ruminate. Embryo small, straight, well differentiated, without chlorophyll. Cotyledons two, often entirely or partially connate. Hypocotyl not developed. Germination cryptocotylar.

Cytology n = 19, 21, 22, 25, 26, 50, 51, c. 140 – Isozyme duplication in Myristica indicates paleopolyploidy.

DNA Mitochondrial coxI intron present.

Phytochemistry Flavonols (kaempferol, quercetin), flavones (5-desoxyflavone, 5-desoxyisoflavone), cyanidin, diarylpropanes (1,3-diarylpropanes, 1,3-diaryl-2-hydroxypropanes), oleanolic acid derivatives, sesquiterpenes, oxyphenols and other aromatic substances, proanthocyanidin, benzylisoquinoline alkaloids, protoberberine alkaloids, indole alkaloids (tryptamine), myristicin, lignans (veraguensin, dihydrocubebin), neolignans, polyketides (alkylsalicylate, acylfloroglucinol, arylalkylbutyrolactones), myo-inisitol, galbacin, licarin A, and amides present. Gallo- and ellagitannins not found.

Use Spices (Myristica fragrans: nutmeg from seeds, mace from arils), narcotics, medicinal plants, perfumes, carpentries.

Systematics Otoba (9; Central America, tropical South America), Coelocaryon (4; C. botryoides, C. oxycarpum, C. preussii, C. sphaerocarpum; tropical Africa), Pycnanthus (4; P. angolensis, P. dinklagei, P. marchalianus, P. microcephalus; tropical Africa), Staudtia (2; S. kamarunensis, S. pterocarpa; tropical West Africa to western Uganda), Cephalosphaera (1; C. usambarensis; mountains in tropical East Africa), Doyleanthus (1; D. arillata; Madagascar), Brochoneura (5; B. acuminata, B. chapelieri, B. dardainii, B. humblotii, B. rarabe; eastern Madagascar), Mauloutchia (10; eastern Madagascar); Compsoneura (17; southern Mexico, Central America, tropical South America), Endocomia (4; E. canarioides, E. macrocoma, E. rufirachis, E. virella; India, Sri Lanka, Southeast Asia, Malesia), Virola (c 75; tropical South America), Bicuiba (1; B. oleifera; Brazil), Haematodendron (1; H. glabrum; Madagascar), Iryanthera (c 20; Panamá, tropical South America), Osteophloeum (1; O. platyspermum; Amazonia in Brazil), Gymnacranthera (7; G. bancana, G. canarica, G. contracta, G. farquhariana, G. forbesii, G. maliliensis, G. ocellata; India, Sri Lanka, Southeast Asia, Malesia to New Guinea and the Bismarck Archipelago), Horsfieldia (c 100; tropical Asia to tropical Australia), Knema (c 60; India, Sri Lanka, Southeast Asia, Malesia to New Guinea), Scyphocephalium (2; S. mannii, S. ochocoa; Cameroon, Equatorial Guinea, Gabon), Myristica (150–160; tropical Asia to tropical Australia).

The flowers are unisexual. Parenchyma, wood rays and phloem contain canals with red, orange or yellow resins. The nucleus of the immature sieve element has one or rarely two protein crystals which are released inte the cytoplasm as the nucleus degenerates and persist within the mature nucleus-free sieve element. The androecium is very variable, although usually the stamens are borne in a single whorl and connate into a synandrium surrounding a central sterile column. Mauloutchia has (secondarily) free stamens, in some species spirally arranged and numerous, a small aril and distinctly granular exine. However, it does not hold a basal position in Myristicaceae. Instead, a clade comprising Otoba, Coelocaryon, Pycnanthus, Staudtia, Cephalosphaera, Doyleanthus, Brochoneura, and Mauloutchia (the “mauloutchioids” sensu Sauquet 2003) is sister-group to the remaining Myristicaceae.

Cladogram of Myristicaceae based on morphology and DNA sequence data (Sauquet 2003).


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