GARRYALES Lindl.

Lindley in C. F. P. von Martius, Consp. Regn. Veg.: 16. Sep-Oct 1835 [‘Garryaceae’]

Eucommiales Nèmejc ex Cronq., Integr. Syst. Class. Fl. Pl.: 182. 10 Aug 1981; Eucommianae Takht. ex Reveal in Phytologia 74: 179. 25 Mar 1993; Aucubales Takht., Divers. Classif. Fl. Pl.: 385. 24 Apr 1997

Habit Dioecious, usually evergreen (rarely deciduous) trees or shrubs or perennial herbs. Young stems and branches quadrangular in cross-section.

Vegetative anatomy Phellogen ab initio superficial. Vessel elements with simple or scalariform perforation plates; lateral pits alternate, scalariform or opposite, bordered pits; vessel elements with helical thickenings. Imperforate tracheary xylem elements tracheids with simple or bordered pits, usually non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma usually apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty or banded. Sieve tube plastids S type. Nodes usually 3:3, trilacunar with three leaf traces (rarely 1:1, unilacunar with one trace). Phloem and cortex sometimes with articulated, unicellular, gutta-percha secreting laticifers. Often with calciumoxalate crystal sand.

Trichomes Hairs unicellular, simple, with ridges running counter-clock-wise (sometimes micropapillate), or absent.

Leaves Usually opposite (sometimes alternate), usually simple (occasionally somewhat pinnately compound), entire or slightly lobed, coriaceous, with conduplicate, supervolute or curved ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata laterocytic, paracytic or anomocytic. Cuticular wax crystalloids as tubuli (and sometimes platelets) or absent. Mesophyll often with sclerenchymatous idioblasts. Sometimes with laticifers containing gutta-percha. Leaf margin serrate or entire.

Inflorescence Terminal or axillary, cymose, catkin-like or dichotomously thyrsoid. Male inflorescences axillary catkin-like cymose; female flowers solitary axillary. Floral prophylls (bracteoles) sometimes absent.

Flowers Actinomorphic. Epigyny (sometimes hypogyny?). Sepals two or four with valvate aestivation, strongly reduced, free or more or less connate, caducous or persistent (sometimes absent). Petals four, with valvate aestivation, free, or absent. Nectariferous disc intrastaminal, quadrangular, bilobate or absent.

Androecium Stamens four (to twelve, rarely more than twelve), usually haplostemonous, alternisepalous or antesepalous. Filaments free from each other and from tepals. Anthers basifixed or dorsifixed, non-versatile, tetrasporangiate, introrse or latrorse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial or secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 2–3(–15)-colporate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate, or exine intectate with smooth, spinulate or verrucate surface.

Gynoecium Pistil composed of two (or three) connate carpels (gynoecium finally pseudomonomerous) or one carpel. Ovary usually inferior (sometimes superior?), unilocular. Stylodia two (or three), free, or style single, simple, short, or absent. Stigmatic surfaces elongate, decurrent, non-papillate, Dry type, or stigma one, capitate, slightly bifid. Male flowers with pistillodium or pistillodium absent.

Ovules Placentation apical. Ovules one or two (or three) per ovary, anatropous, pendulous, apotropous, unitegmic, usually crassinucellar (rarely tenuinucellar). Archespore sometimes multicellular. Large placental obturator sometimes present. Megagametophyte monosporous, Polygonum type. Fertilization often delayed. Antipodal cells sometimes early degenerating. Endosperm development nuclear or cellular. Endosperm haustoria? Embryogenesis solanad.

Fruit A one- or two-seeded and gradually drying berry with persistent calyx or style/stylodia, or a single-seeded samara.

Seeds Aril absent. Arilloid formed from placental obturator or absent. Testa usually thick, multiplicative (sometimes membranous); outer part sarcotesta, inner part with tangentially elongate cells. Exotestal cells large, palisade, fleshy. Endotesta? Perisperm not developed. Endosperm copious, fleshy, with oil, petroselinic acid, starch and hemicellulose. Suspensor sometimes very long. Embryo small or large, straight, well differentiated, with chlorophyll. Cotyledons two. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin), O-methylated flavonoids, cyanidin, Route II decarboxylated iridoids, Group I carbocyclic iridoids (e.g. scandoside, aucubin, ipolamiide, ajugol, ajugoside, and harpagide), Group II decarboxylated iridoids, oleanolic acid derivatives, ellagic acid, caffeic acid, chlorogenic acid, strongly toxic diterpene alkaloids, saponins, petroselinic (cis-6-octadecenoic) acid, eucommin A, lignans (pinoresinol, syringaresinol), gutta-percha, and acetylenes present. Gallic acid and cyanogenic compounds not found. Carbohydrates sometimes stored as inulin.

Fossils Leaves and samaras of Eucommia is frequently recorded in Cenozoic layers in North America, Europe and East Asia from the Eocene of eastern North America and Japan onwards. Eucommia is known from Central and North America in the Oligocene and the Miocene and in Europe (also by typical tricolpate pollen grains) from the Oligocene to the Pleistocene. Fossil pollen of Eucommiaceae has been found in Paleocene and younger layers.

Vegetative anatomy Phellogen ab initio superficial. Medulla lamellated. Vessel elements with simple perforation plates (earliest secondary xylem scalariform); lateral pits alternate or opposite, bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, homocellular or somewhat heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal banded. Sieve tube plastids Ss type (with approx. ten starch grains). Nodes 1:1, unilacunar with one leaf trace. Phloem fibres present. Phloem and cortex with articulated, unicellular laticifers secreting gutta percha. Crystals?

Leaves Alternate (spiral to distichous), simple, entire, with supervolute (involute?) to curved ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids absent. Laticifers with gutta-percha. Leaf margin coarsely serrate; single vein proceeding into each glandular-tipped tooth.

Inflorescence Male inflorescences axillary catkin-like cymose; female flowers solitary axillary. Floral prophylls (bracteoles) absent.

Flowers Actinomorphic, small. Hypogyny? Tepals absent. Nectary absent. Disc absent.

Androecium Stamens usually five to twelve (rarely four or more than twelve). Filaments short, free. Anthers basifixed, non-versatile, tetrasporangiate, introrse?, longicidal (dehiscing by longitudinal slits); connective prolonged at apex. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate with indistinct ora, shed as monads, bicellular at dispersal. Exine intectate. Pollen surface spinulate to verrucate.

Gynoecium Pistil composed of two connate carpels, one fertile and one aborted. Ovary superior?, unilocular due to pseudomonomery. Stylodia two, short, unequally sized, diverging. Stigmas adaxially decurrent, type? Pistillodium absent.

Ovules Placentation apical. Ovules two (one fertile) per ovary, anatropous, collateral, pendulous, apotropous, unitegmic, tenuinucellar to weakly crassinucellar. Micropyle very long. Integument five to nine cell layers thick. Parietal tissue approx. three cell layer thick. Nucellar cap present. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria? Embryogenesis solanad.

Seeds Aril absent. Testa membranous. Exotestal cells? Endotesta? Perisperm not developed. Endosperm copious, oily? Embryo large, straight, well differentiated, with chlorophyll? Cotyledons two. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin), O-methylated flavonols, cyanidin, Route II decarboxylated iridoids, Group I carbocyclic iridoids (e.g. aucubin, ipolamiide, ajugol, ajugoside, and harpagide), oleanolic acid derivatives, ellagic acid, condensed tannins (in bark), saponins, eucommin A, lignans (pinoresinol, syringaresinol), gutta percha, and acetylenes present. Alkaloids and cyanogenic compounds not found. Carbohydrates stored as inulin. Oxalate sometimes accumulated.

Systematics Eucommia (1; E. ulmoides; temperate central China).

Eucommia is sister to Garryaceae.

Aucubaceae Bercht. et J. Presl, Přir. Rostlin: 2: 91. 1825 [’Aucubeae’]

Distribution Eastern Himalayas, northern Burma, China, southern Korean Peninsula, Japan, the Ryukyu Islands, Taiwan, western North America, Mexico, Central America, the West Indies.

Habit Dioecious, evergreen trees or shrubs (Garrya); shrubs or perennial herbs (Aucuba). Young stems and branches quadrangular in cross-section.

Vegetative anatomy Phellogen ab initio superficial. Cauline pericyclic envelope absent in Aucuba. Vessel elements with simple or scalariform perforation plates; lateral pits alternate, scalariform or opposite, bordered pits. Imperforate tracheary xylem elements tracheids (vascular tracheids present in Aucuba) with simple (Aucuba) and/or bordered pits (in Garrya with spiral wall thickenings), usually non-septate (in Aucuba sometimes septate). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma usually apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty or banded. Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Often with calciumoxalate crystal sand.

Trichomes Hairs unicellular, simple, with ridges running counter-clock-wise, or absent.

Leaves Opposite, usually simple (occasionally somewhat pinnately compound), entire or slightly lobed, coriaceous, with conduplicate ptyxis. Stipules and leaf sheath absent. Petioles more or less connate at base. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata in Garrya laterocytic or paracytic, in Aucuba anomocytic. Cuticular wax crystalloids as tubuli (and sometimes platelets). Mesophyll often with sclerenchymatous idioblasts. Leaf margin serrate (Aucuba) or entire (in Garrya cartilaginous).

Inflorescence Terminal or axillary, cymose; in Garrya catkin-like, silk-hairy, with bracts more or less connate; in Aucuba dichotomously thyrsoid. Floral prophylls (bracteoles) absent? in Garrya and in male flowers of Aucuba.

Flowers Actinomorphic, small. Epigyny. Sepals four in male flowers of Garrya, small, with valvate aestivation, strongly reduced, connate at apex, early caducous, in female flowers usually two (rarely four; prophylls?), almost entirely connate, strongly reduced or absent; sepals in Aucuba four, free, persistent in fruit, or absent. Petals absent in Garrya; petals in Aucuba four, with valvate aestivation, free. Nectariferous disc intrastaminal, in Aucuba fleshy, quadrangular, on ovary apex, in male flowers of Garrya heavily reduced (bilobate), in female flowers absent.

Androecium Stamens four, alternisepalous (Garrya) or antesepalous, alternipetalous (Aucuba). Filaments free from each other and from tepals. Anthers basifixed (Garrya) or dorsifixed (Aucuba), non-versatile, tetrasporangiate, introrse or latrorse, longicidal (dehiscing by longitudinal slits). Tapetum amoeboid-periplasmodial. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 2–3(–15)-colporate, shed as monads, bicellular at dispersal. Exine semitectate with columellate infratectum, reticulate tectum and simpli- or duplicate muri (Garrya), or exine intectate with smooth surface (Aucuba).

Gynoecium Pistil composed of usually two (rarely three) connate carpels (gynoecium finally pseudomonomerous, Garrya), or one carpel (seemingly? monomerous, Aucuba). Ovary inferior, unilocular, in Aucuba nectariferous. Stylodia usually two (rarely three), subulate, free, diverging (Garrya) or style single, simple, short, thick (Aucuba). Stigmatic surfaces elongate, decurrent, non-papillate, Dry type (Garrya), or stigma one, capitate, slightly bifid, type? (Aucuba). Male flowers with pistillodium.

Ovules Placentation apical. Ovules usually one or two (sometimes three) per ovary, anatropous, pendulous, epitropous (Garrya), unitegmic, crassinucellar. Integument twelve to c. 30 cell layers thick (Garrya). Parietal tissue in Garrya four or five cell layers thick, in Aucuba approx. eight cell layers thick. Hypostaste present in Garrya. Nucellar cap sometimes two cell layers thick. Archespore sometimes multicellular (Aucuba). Megasporocytes in Aucuba sometimes several. Large placental obturator closing upper part of locule in Garrya. Megagametophyte monosporous, Polygonum type. Antipodal cells in Garrya early degenerating or persistent. Endosperm development nuclear (Garrya) or cellular (Aucuba). Endosperm haustoria? Embryogenesis solanad (Garrya).

Fruit A one- or two-seeded and gradually drying berry with persistent calyx or style (stylodia).

Seeds Aril absent. Arilloid in Garrya formed from placental obturator. Testa thick, multiplicative; outer part sarcotesta, inner part with tangentially elongate cells. Exotestal cells in Garrya large, palisade, fleshy. Endotesta not persistent (Garrya). Perisperm not developed. Endosperm copious, fleshy, with oil, petroselinic acid, starch and hemicellulose. Suspensor in Garrya very long. Embryo small, straight, well differentiated, with chlorophyll. Cotyledons two. Germination phanerocotylar (Garrya).

Phytochemistry Flavonols (kaempferol, quercetin, in Aucuba), flavonoid glycosides, Route II decarboxylated iridoids, Group I carbocyclic iridoids (e.g., scandoside and aucubin), strongly toxic diterpene alkaloids (e.g. garryfoline in Garrya), chlorogenic acids, and petroselinic acid present. Caffeic acid? Gutta percha in Aucuba? Ellagic and gallic acids, tannins, proanthocyanidins, saponins, and cyanogenic compounds not found.

Systematics Aucuba (3–10; A. chinensis, A. himalaica, A. japonica; Sikkim, northern Burma, China, southern Korean Peninsula, Japan, the Ryukyu Islands, Taiwan), Garrya (c 15; western United States, Mexico, Central America to Panamá, the Greater Antilles).

Garryaceae are sister group to Eucommia (Eucommiaceae).

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GARRYIDAE R. G Olmstead, W. S. Judd et P. D. Cantino

GARRYALES Lindl.

EUCOMMIACEAE Juss.

GARRYACEAE Lindl.