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POSITION OF THE ORANGE SUBFAMILY, AURANTIOIDEAE, IN THE PLANT FAMILY RUTACEAE The family Rutaceae belongs to the division Embryophyta Siphonogama, subdivision Angiospermae, class Dicotyledoneae, subclass Archichlamydeae (Choripetalae and Apetalae), order Geraniales, suborder 1, Geraniineae, along with eleven other plant families classed by Engler and Diels (1936, p. xl) in the following order: Oxalidaceae, Geraniaceae, Tropaeolaceae, Linaceae, Erythroxylaceae, Zygophyllaceae, Cneoraceae, Rutaceae, Simarubaceae, Burseraceae, Meliaceae, Akariaceae. The other suborders of the Geraniales are as follows: 2, Malpighiineae (3 families); 3, Polygalineae (2 families); 4, Dichapetalineae (1 family); 5, Tricoccae (2 families); 6, Callitrichineae (1 family). The order Geraniales is preceded by the order Rosales (including 17 families, among them Rosaceae and Leguminosae) and followed by the order Sapindales (including 23 families, among them Anacardiaceae and Sapindaceae). The relationship of the orange subfamily to the six other subfamilies of the Rutaceae is shown by the general key given by Engler (1931, pp. 105-11), which, translated somewhat freely from the German, reads as follows: Subfamily I. RUTOIDEAE. Carpels usually 4-5, seldom 1-3 or more, often united only by the common pistil and free below, at maturity more or less separated, opening inward by splitting the follicle (loculicidal), usually with a dehiscent endocarp, very seldom with 4-1 fleshy drupes (Pitaviinae). Leaves and bark of twigs with schizolysigenous oil glands. (5 tribes, 17 subtribes, 86 genera.) Tribe (i) Xanthoxyleae. Trees or shrubs, mostly small, greenish or greenish-white (seldom large and clear white) flowers which are always actinomorphous (radial) and often unisexual. Carpels only seldom with more than 2 ovules, Embryo [sic] mostly with flat cotyledons in endosperm (except Bosistoa and Pagetia). (5 subtribes, 30 genera.) Subtribe 1, Evodiinae (includes Xanthoxylon and Fagara) (20 g.), tropics and subtropics of Old and New Worlds; subtr. 2, Lunasiinae ( 1 g.), Monsoon region;1 subtr. 3, Decatropidinae (3 g.), Central America; subtr. 4, Choisyinae (5 g.), Central America, Pacific Islands, and Australia; subtr. 5, Pitaviinae (1 g.), Chile; Tribe (ii) Ruteae. Herbs or perennial herbs, seldom shrubs, with medium-sized, always perfect flowers, which sometimes (Dictamnus) are slightly zygomorphic. Carpels as a rule with more than 2 ovules (only in Ruta in the subgenus Haplophyllum with 2 ovules and occasionally indehiscent fruitlets). Seeds with endosperm. (2 subtribes, 6 genera.) Subtr. 6, Rutinae (5 g.), subtropical and temperate regions of Old and New Worlds (includes Cneoridium, a California shrub); subtr. 7, Dictamninae (1 g.), temperate zone of Europe and Asia; Tribe (iii) Boronieae. Perennial herbs or shrubs. Carpels with only 1 or 2 ovules. Flowers always actinomorphic, mostly perfect. Embryo usually straight, cylindrical, usually immersed in abundant fleshy endosperm. (5 subtribes, 19 genera.) Subtr. 8, Boroniinae (6 g.), Australia and New Caledonia; subtr. 9, Eriostemoninae (9 g.), Australia and New Caledonia; subtr. 10, Correinae (1 g.), Australia; subtr. 11, Nematolepidinae (2 g.), Western Australia; subtr. 12, Diplolaeninae (1 g.), Western Australia; Tribe (iv) Diosmeae. Mostly perennial herbs and shrubs, seldom trees (Calodendrum), always with simple leaves. Flowers almost always actinomorphic, mostly perfect. Seeds without endosperm. Embryo mostly straight with fleshy cotyledons. (3 subtribes, 12 genera.) Subtr. 13, Calodendrinae (1 g.) East Africa (Kenya) to Cape Province (South Africa); subtr. 14, Diosminae (9 g.), Cape Province (South Africa); subtr. 15, Empleurinae (2 g.), Cape Province (South Africa); Tribe (v) Cusparieae. Shrubs or tree. Flowers actinomorphic or the corolla and the androecium zygomorphic. Seeds with little or no endosperm. Embryo curved, the plumule lying between the cotyledons. (2 subtribes, 19 genera.) Subtr. 16, Pilocarpinae (3 g.), tropical America and subtropical South America; subtr. 17, Cuspariinae (16 g.), tropical America, mostly Brazil and northern South America. Subfamily II. DICTYOLOMATOIDEAE. Leaves with many-celled but not lysigenous oil glands. Flowers actinomorphic. Stamens isomerous and alternate with the petals, with bractlets at the base. Carpels united only at the base, with several ovules. Fruits with dehiscent endocarp, 3-4 seeded. Small trees with doubly pinnate leaves. (1 tribe, 1 genus.) Tribe (vi) Dictyolomateae. (1 genus.) Brazil and eastern Peru. Subfamily III. FLINDERSIOIDEAE. Trees or shrubs. Carpels 5-3, united, each with 2-8 ovules in 2 rows. Fruit a loculicidal or septicidal capsule, with persistent endocarp. Seed winged, without endosperm. Leaves with lysigenous oil glands. (1 tribe, 2 genera.) Tribe (vii) Flindersieae. (2 genera.) Eastern Australia, New Caledonia, East Indian Archipelago, Ceylon, and India. Subfamily IV. SPATHELIOIDEAE. Carpels 3, completely fused, each with 2 pendent ovules. Fruit a winged drupe with a 3-loculed hard pit. With oil-bearing secretory cells in the leaves, bark, and pith; lysigenous oil glands in the leaf margins. (1 tribe, 1 genus.) Tribe (viii) Spathelieae. (1 genus.) West Indies. Subfamily V. TODDALIOIDEAE. Carpels 5-2, incompletely or completely united, or else only 1, each with 1 or 2 ovules. Fruit formed out of 4-2 drupelets united only at the base, or which some occasionally abort, or else a drupe with a thick or thin mesocarp and a thick or thin endocarp, or a dry, winged, indehiscent fruit. Seeds with or without endosperm. (1 tribe, 6 subtribes, 25 genera.) Tribe (ix) Toddalieae. (6 subtribes, 25 genera.) Subtr. 18, Phellodendrinnae (2 g.), temperate and subtropical eastern Asia and tropical Africa; subtr. 19, Sohnreylinae (1 g.), Amazon Valley, Brazil; subtr. 20, Pteleinae (4 g.), tropical and temperate America; subtr. 21, Oriciinae (2 g.), tropical Africa; subtr. 22, Toddaliinae (13 g.), tropics, Old and New Worlds (include Casimiroa, a Mexican and Central American fruit tree); subtr. 23, Amyridinae (3 g.), northern South America, West Indies, Central America, Texas, Florida, tropical Africa. [The genus Amyris, having about 30 species native to Florida, Texas, Mexico, Central America, West Indies, and northern South America, is rather closely related to the tribe Clauseneae of the next subfamily, Aurantioideae.] Subfamily VI. AURANTIOIDEAE. Fruit a berry [or hesperidium] with a leathery rind or hard shell, in tribe Citreae often with pulp formed by juicy emergenzen that arise on the carpellary walls. Seeds without endosperm, sometimes with 2 or more nucellar [false] embryos. Leaves and bark [of twigs and young branches] with schizolysigenous oil glands [small or sometimes large trees, rarely shrubs.]; (2 tribes, 33 genera.) [This subfamily is given as classified by Swingle for this chapter. Engler made a single tribe with 2 subtribes and with a total of 29 genera.] Tribe (x) Clauseneae. (3 subtribes, 5 genera.) Subtr. 24, Micromelinae (1 g.), Monsoon region and western Polynesia as far as Tonga, Fiji, and Samoan Islands; subtr. 25, Clauseninae (3 g.), Monsoon region and tropical Africa; subtr. 26, Merrilliinae (1 g.), Malay Peninsula and Sumatra; Tribe (xi) Citreae. (3 subtribes, 28 genera.) Subtr. 27, Triphasiinae (8 g.), Monsoon region; subtr. 28, Citrinae (13 g.), Monsoon region and tropical Africa; subtr. 29, Balsamocitrinae (7 g.), Monsoon region and tropical Africa. Subfamily VII. RHABDODENDROIDEAE. Flowers with a bowl-shaped concave receptacle, with obliterated calyx, 5 petals, and very numerous stamens. Ovary free, ovoid, 1-locular, with 1 basal ovule. Pistil attached to the side of the ovary, with a long lateral stigma. Fruits with thin exocarp and thin endocarp. Leaves simple. (1 tribe, 1 genus.) Tribe (xii) Rhabdodendreae. (1 genus.) Amazon Valley. Total for the Rutaceae: 7 subfamilies, 12 tribes (containing 29 subtribes), with about 150 genera and 1,600 species.
THE ORANGE SUBFAMILY, AURANTIOIDEAE The subfamily Aurantioideae is defined briefly above in the outline of the Rutaceae, where all the subfamilies, tribes, and subtribes of this plant family are given. All the species of the Aurantioideae are trees or shrubs with persistent (evergreen) leaves except in the three monotypic genera, Poncirus, Aegle, and Feronia, and in three species of Clausena (C. pentaphylla, C. dentata var. dulcis, and C. suffruticosa) and one of Murraya (M. alternans). The flowers are usually white and very often fragrant. Many of the genera bear subglobose fruits with a green, yellow, or orange peel dotted with numerous oil glands that often give an agreeable aroma when the fruit is handled. The fruits of the genus Citrus are among the most beautiful, most fragrant, and most delicious known to man. This subfamily contains Citrus and thirty-two other genera related more or less to Citrus, classed in two tribes and six subtribes which contain, as treated here, 203 species. The genus Citrus and a few others closely related to it have fruits unlike any others known to botanists in being filled with curious pulp-vesicles which contain in many species a delicious juicy tissue. The subtribe Balsamocitrinae, which belongs to the tribe Citreae, has fruits as large as oranges or grapefruits but with a hard woody shell. These hard-shelled fruits do not contain juicy pulp-vesicles, although some of them are pleasantly aromatic and much liked by both natives and Europeans in India and Indo-China. Many of the remote relatives of Citrus belonging to the tribe Clauseneae have extremely small fruits very unlike those of Citrus and usually semidry and entirely inedible. Nevertheless some of these remote relatives have been found to be graft-compatible with Citrus. The native habitat of the subfamily Aurantioideae is limited to the Old World. Most of the genera are found in the Monsoon region from West Pakistan to north-central China and thence south through the East Indian Archipelago to New Guinea and Bismarck Archipelago, northeastern Australia, New Caledonia, Melanesia, and the western Polynesian islands. Of the thirty-three genera that constitute the Aurantioideae, no fewer than twenty-nine are native to the Monsoon region and twenty-seven of them are found only there. Five genera, belonging to two tribes and to three subtribes, are native to tropical Africa and four genera are found only there. Only one genus, Clausena, is native both to the Monsoon region and to tropical Africa. Many of the species of Citrus and of the genera closely related to Citrus are now found in cultivation or are grown for ornament in all the tropical and subtropical regions of the world. 153554b96e