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10. Rutaceae

芸香科 yun xiang ke

Authors: Prof. Dianxiang Zhang, Thomas G. Hartley & David J. Mabberley

Shrubs, trees, or sometimes herbs, sometimes scrambling or scandent, sometimes armed, with aromatic volatile oils contained in glands visible at surface of at least leaves, young branchlets, inflorescences, flower parts, fruit, or cotyledons in seed. Stipules absent [or stipular excrescences rarely present]. Leaves alternate, opposite [or whorled], simple (petiole neither apically swollen nor articulate with leaf blade), 1-foliolate (in individual specimens at least some 1-foliolate leaves with petiole apically swollen and/or articulate with leaf blade), or variously compound. Flowers bisexual or unisexual, usually 3-5-merous, actinomorphic or rarely zygomorphic, hypogynous [or rarely perigynous]. Perianth in 2 series, with clearly differentiated calyx and corolla or sometimes in 2 irregular series or 1 series, with ± undifferentiated tepals. Sepals distinct or connate to their full length. Petals distinct [or rarely coherent or connate for part of their length]. Stamens usually as many as or 2 × as many as petals or sometimes more numerous; filaments distinct or sometimes coherent or connate for at least part of their length; anthers introrse or sometimes latrorse, longitudinally dehiscent. Disk [rarely lacking] within androecium, nectariferous, flattened, annular, cup-shaped, pulvinate, or sometimes columnar, bell-shaped, conic, or hourglass-shaped. Gynoecium of 1-5 distinct 1-loculed carpels or 2 to many partially to completely connate carpels; placentation axile [very rarely becoming parietal]; ovules 1 to many per locule. Fruit of 2-5 follicles [drupes or samaras] or a single follicle, capsule, or berry [or samara]. Seeds with relatively large embryo; endosperm present and fleshy or lacking.

About 155 genera and ca. 1600 species: nearly cosmopolitan but mainly tropical and subtropical; 22 genera (one endemic, one introduced) and 126 species and hybrid species (49 endemic, at least two introduced) in China.

Oil glands of Rutaceae, when viewed from the surface of plant parts they occupy, are usually pellucid. They also appear to be ± isodiametric and to have ± definite patterns of distribution. In blades of leaves, for example, where they are most commonly observed, they are usually ± evenly scattered throughout, or sometimes they are restricted to the margins. Rarely they are alleged to occur only along the secondary veins of the blades.

In a cladistic analysis of selected genera of Rutaceae and related families based on rbcL and atpB molecular data (but only on rbcL data for Harrisonia), M. W. Chase, C. M. Morton, and J. A. Kallunki (Amer. J. Bot. 86: 1191-1199. 1999) recommended the placement of Harrisonia (traditionally Simaroubaceae) in Rutaceae. We do not agree with this classification, particularly because Harrisonia appears to lack oil glands, and suggest that the genus is most correctly placed in Cneoraceae, in which it is treated herein.

Several taxa of cultivated Rutaceae treated by C. C. Huang (Fl. Reipubl. Popularis Sin. 43(3). 1997) are not treated here: Casimiroa edulis La Llave, which is native to Mexico, has been introduced as a cultivated plant to the Xishuangbanna Botanical Garden in Yunnan; Limonia acidissima Linnaeus (Feronia limonia (Linnaeus) Swingle), which is native to India and Sri Lanka, has been reported as cultivated in Taiwan by C. C. Huang (Fl. Reipubl. Popularis Sin. 44(3): 212. 1997) but was not reported as a cultivated plant by T. C. Huang in A Checklist of the Vascular Plants of Taiwan (Fl. Taiwan 6: 81-83. 1979); Flindersia amboinensis Poiret, which is native to Indonesia and New Guinea, is cultivated in S China; Ptelea trifoliata Linnaeus, which is native to North America, is cultivated in Beijing and Liaoning; and Ruta graveolens Linnaeus, which is native to the Mediterranean region, is widely cultivated in China.

Huang Chengchiu. 1997. Rutaceae. In: Huang Chengchiu, ed., Fl. Reipubl. Popularis Sin. 43(2): ii-viii, 1-250.


1 Leaves opposite   (2)
+ Leaves alternate   (6)
       
2 (1) Leaves mostly odd-pinnate; inflorescences terminal or terminal and axillary; plants dioecious   (3)
+ Leaves digitately 3-foliolate or 1-foliolate; inflorescences axillary or basal to leaves; plants usually monoclinous or dioecious   (4)
       
3 (2) Axillary buds exposed; ovary in female flowers with carpels connate at base, otherwise contiguous; fruit of 1-5 follicles.   3 Tetradium
+ Axillary buds concealed in excavated base of petioles; ovary in female flowers syncarpous; fruit a drupaceous berry.   10 Phellodendron
       
4 (2) Leaves digitately 3-foliolate or 1-foliolate; ovary in bisexual and female flowers with carpels connate at base, otherwise contiguous; fruit of 1-4 basally connate follicles.   4 Melicope
+ Leaves 1-foliolate; ovary in bisexual and female flowers syncarpous or rarely (Acronychia) separated apically by septicidal fissures; fruit a drupaceous berry   (5)
       
5 (4) Flowers male or female; stamens glabrous, similar in male and female flowers but lacking pollen in latter; gynoecium rudimentary and minute in male flowers, ± as long as petals in female flowers; seed coat with thick inner layer of dense black sclerenchyma and spongy outer layer bounded externally by a shiny black pellicle.   12 Maclurodendron
+ Flowers bisexual; staminal filaments with an indumentum; gynoecium ± half as long as petals; seed coat with thick inner layer of dense black sclerenchyma and an outer layer of compact dull to ± shiny parenchymatous tissue.   11 Acronychia
       
6 (1) Plants herbaceous perennials   (7)
+ Plants woody   (10)
       
7 (6) Leaves mostly simple.   6 Haplophyllum
+ Leaves mostly compound   (8)
       
8 (7) Flowers zygomorphic, 5-merous; stamens 10; fruit of 5 basally connate follicles, 1-2 cm.   8 Dictamnus
+ Flowers actinomorphic, 4-merous; stamens usually 8; fruit follicular or capsular, to 0.5 cm   (9)
       
9 (8) Leaves pinnately to ternately decompound; fruit of 4 distinct follicles.   5 Boenninghausenia
+ Leaves mostly digitately 3-foliolate; fruit a capsule.   7 Psilopeganum
       
10 (6) Fruit follicular or drupaceous; endocarp cartilaginous or leathery; seeds with endosperm; flowers male, female, or sometimes bisexual; functional stamens as many as petals except in plants of Zanthoxylum with undifferentiated perianth which have 5-9 tepals and 3-8(-10) functional stamens; functional gynoecium 1-7-loculed, syncarpous or with distinct or only basally connate carpels   (11)
+ Fruit baccate; endocarp membranous or fleshy; seeds without endosperm; flowers bisexual or bisexual and male; stamens at least 2 × as many as petals or rarely fewer; functional gynoecium 2- to many loculed, syncarpous   (14)
       
11 (10) Leaves mostly compound; plants usually armed   (12)
+ Leaves simple; plants unarmed   (13)
       
12 (11) Leaves odd-pinnately 3- to many foliolate or sometimes digitately 3-foliolate (occasional leaves even-pinnate, 2-foliolate, or 1-foliolate); functional gynoecium 1-5-loculed, with distinct carpels or carpels basally connate; fruit of 1-5 distinct or basally connate follicles.   1 Zanthoxylum
+ Leaves digitately 3-foliolate (occasional leaves 1- or 2-foliolate); functional gynoecium 4-7-loculed, syncarpous; fruit a 4-7-loculed drupaceous berry.   9 Toddalia
       
13 (11) Inflorescences axillary, between leaves, or basal to leaves; flowers male or female; functional gynoecium 4-loculed; ovaries basally connate; fruit of 1-4 basally connate follicles.   2 Orixa
+ Inflorescences terminal; flowers male, female, or sometimes bisexual; functional gynoecium 2-5-loculed, syncarpous; fruit drupaceous berry, with 1-5 1-seeded leathery pyrenes.   13 Skimmia
       
14 (10) Flowers bisexual or bisexual and male; stamens more than 2 × as many as petals; gynoecium 2- to many loculed; ovules 2 to many per locule; fruit with stalked or rarely sessile pulp vesicles; leaves odd-pinnately 3(or 5)-foliolate, digitately 3-foliolate, 1-foliolate, or simple   (15)
+ Flowers bisexual; stamens 2 × as many as petals or rarely fewer; gynoecium 2-5-loculed; ovules 1 or 2 per locule; fruit either without pulp vesicles or with sessile pulp vesicles; leaves odd-pinnately 3- to many foliolate, digitately 3-foliolate, 1-foliolate, or simple (occasional leaves 2-foliolate)   (16)
       
15 (14) Plants evergreen, rarely deciduous (Citrus trifoliata); fruit with leathery (or rarely soft) exocarp and spongy mesocarp; seeds embedded in pulp vesicles; leaves 1-foliolate, simple, or rarely digitately 3-foliolate.   21 Citrus
+ Plants deciduous; fruit with thin, parenchymatous exocarp and woody mesocarp; seeds embedded in a clear glutinous substance; leaves odd-pinnately 3(or 5)-foliolate.   22 Aegle
       
16 (14) Radial walls of locules in gynoecium becoming curved after anthesis; cotyledons in seed thin and flat, convolute, and/or folded.   14 Micromelum
+ Radial walls of locules in gynoecium straight; cotyledons in seed plano-convex, neither convolute nor folded   (17)
       
17 (16) Terminal and axillary buds and usually young inflorescences with a rust-colored villosulous indumentum (sometimes partly bleached); style persistent in fruit.   15 Glycosmis
+ Terminal and axillary buds and young inflorescences without a rust-colored villosulous indumentum; style or at least its distal portion usually deciduous in fruit   (18)
       
18 (17) Leaves 1-foliolate or simple   (19)
+ Leaves, or most of them, compound   (20)
       
19 (18) Fruit without pulp vesicles; stamens distinct; flowers axillary, solitary or in few-flowered fascicles; petiole 0.4-2.5 cm, usually bent and/or twisted and ± apically swollen; erect or scrambling shrubs or woody climbers, usually armed.   19 Paramignya
+ Fruit usually with pulp vesicles; stamens distinct or with filaments variously connate or coherent; flowers in terminal and/or axillary inflorescences, in fascicles, racemes, or panicles; petiole 0.2-1.3 cm, usually not bent, twisted, or swollen; shrubs or trees, armed or unarmed.   20 Atalantia
       
20 (18) Leaves mostly digitately 3-foliolate; woody climbers, armed.   18 Luvunga
+ Leaves odd-pinnate; shrubs or trees, unarmed   (21)
       
21 (20) Flower buds globose, pyriform, or rarely ovoid; staminal filaments ± abruptly dilated toward base, ± straight or geniculate; style 0.5-2.5 × as long as ovary; disk columnar, conic, bell-shaped, or ± hourglass-shaped; seeds glabrous.   16 Clausena
+ Flower buds ellipsoid to obovoid or narrowly so or subcylindric; staminal filaments linear or sublinear, ± straight; style 3-7 × as long as ovary; disk annular, pulvinate, or columnar; seeds glabrous or villous.   17 Murraya

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