Flora Malesiana,Series I, Volume 13 (1997) 43-144 Boraginaceae 1 H. Riedl Vienna) Boraginaceae Juss., Gen. Pl. (1789) 128 (‘Borragineae ’); Brand in Engl., Pflanzenr., fam. IV.252 (1921) 1-183 (Cynoglosseae ); ibid. (1931) 1-236 (Cryptantheae); Heine in Fl. Nouv.-Caléd. 7 (1976) 95-118; I.M.Johnston, Contr. Gray Herb. 73 (1924) 42-73; Ridl., Fl. Malay Penins. 2 (1923) 438-442; C.B.Rob., Philipp. J. Sc., Bot. 4 (1909) 687-698; Van Royen, Pac. Sc. 29 (1975) 79-98. Trees, shrubs, subshrubs, woody climbers, perennial or annual herbs usually covered by hairs or bristles on the herbaceous parts, woody species sometimes entirely glabrous. Leaves alternate, very rarely oppsite (in Tournefortia), exstipulate, undivided, usual- in few with reticulate venation of which ly entire, a very species serrate, only the main detectable in most either nerves are clearly cases. Inflorescence a simple cyme or com- pound, cymes arranged dichotomously or in racemes or panicles, with or without bracts, terminal or lateral, sometimes single flowers in the axils of upper leaves. Flowers her- maphroditic, rarely unisexual and plants monoecious, composed of calyx and corolla, pentamerous, rarely tetramerous, actinomorphous, in some genera slightly zygomor- phous. Calyx campanulate to cup-shaped, lobes entirely free or more or less coherent, sometimes accrescent or spreading after flowering, sessile or distinctly stalked. Corolla coherent in lower part (tube), lobes free, erect or spreading in the upper part (limb), with an intermediate, gradually widening zone (throat), tubular, campanulate to funnel- shaped or rotate, lobes usually imbricate in bud, rarely valvate, bent into the throat (in some Heliotropium spp.) or contorted (in Myosotis). In many genera, scale-like in- in vaginations (fornices) present the throat. In some genera, a basal nectary ring, scale- like appendages (that are not hollow invaginations) or variously distributed hairs inside the corolla present. Stamens of the same numberas corolla lobes, alternating with them, included in or exserted from the corolla; anthers sessile or on distinct filaments, some- times with a sterile tip of connective tissue or produced to an awn-like structure, awns twisted of one flower or not, anthers bilocular, dorsally fixed to the corolla or the fila- often fixed either disc-like ment, opening longitudinally. Ovary superior, to a base or base produced to a conical or pyramidal structure (gynobasis), bicarpellate as a rule, un- four divided, entire or with deep furrows, breaking into two biovulatehalves or into four uni-ovulate sometimes of the locules mericarps (‘nutlets’), part aborted, rarely a greater number of carpels developed (e.g., in Trigonotis procumbens); placentation axillary; ovules erect or nearly horizontal, rarely pendent. Style single, either terminal on the un- divided free at last of the fixed ovary or on top 4-partite ovary, or basally on the disc- like base between the nutlets or apically on the conical to pyramidate gynobasis, undi- 1) With contributions by: P. Baas, Leiden (vegetative anatomy), R.W.J.M. van der Ham, Leiden (pa- lynology), R. Hegnauer, Leiden (phytochemistry and chemotaxonomy). Most drawings are by Ms. N. Spitteler; origin of photographs is as indicated. 43 44 Flora Malesiana, Ser. I, Vol. 13 (1997) vided or once to twice forked, sometimes cleft nearly to the base; stigma absent or disc- the to cushion-shaped terminating style, or (in Heliotropium and Tournefortia) forming a ring round the style at various levels (for convenience’ sake the ring together with the terminal part of the style above the ring is called stigma as a rule); heterostyly rather outside frequent (in groups Malesia). Fruit drupaceous or a berry or divided into 2-4 surrounded hard sometimes adherent mericarps by a outer wall, to the style and separat- ing with an awn; surface smooth, warty, or rugulose, often tuberculate or spiny, some- times divided into a disc-like outer and a convex inner side, the outer disc surrounded by a thickened margin or not, sometimes glochids present on the whole or certainparts of the surface or nutlets winged. Seeds usually without albumen. DISTRIBUTION of About 100 to 115 genera with a total about 2,400 species in all climatic regions of the earth. The of subfamilies Ehretioideae woody genera Cordioideae, and Heliotropioideae in and is in are predominant tropical subtropical regions, Heliotropium widespread trop- ical and temperateregions, while herbs and subshmbs belonging to Boraginoideae are most numerous in temperate regions, in the tropics mainly confined to higher altitudes or growing as weeds and also represented in the arctic with a few species (involves only Boraginoideae). In the Malesian region, 12 genera are indigenous, two more introduced. Of these, Crucicaryum is of rather doubtful value and only known from one collection, which has been lost. Pteleocarpa, sometimesregarded as a member of Boraginaceae, is not includ- in the number ed this family by present author. The of species acknowledged in this ac- is of which 64 65 the other 12 count, 77, or are indigenous, or 13 having been intro- ducedeither for economical and/or ornamental uses or as weeds. HABITAT Boraginaceae are widespread in a great number of different habitats from sandy sea shores mountain desertic semidesertic and to forests, wet meadows, or regions stony slopes. In tropical countries, they are absent from lowland rain forests as a rule as well as from vegetation types derived from them. Subfamily Boraginoideae is confined to mountainousareas at higher altitudes in the tropics and subtropics as mentionedabove. Several species are introduced into new areas of distribution as weeds. ECOLOGY Pollinationis performed by insects of various groups in a great majority of cases though special adaptations to particular pollinators are lacking. Faucal appendages prevent in- with short from the flowers in sects a proboscis pollinating many genera, however. These the in the basal of the is observ- appendages serve as guide to nectar part corolla. Selling ed in a number of taxa, but is prevented by heterostyly in others. Heterostyly is combin- ed with in self-incompatibility as a rule, some genera also with pollen dimorphism. In Cordia bats act large-flowered species may as pollinators. Riedl Boraginaceae 45 several of found within the Nutlets There are ways dispersal family. bearing glochids are well adapted to epizoochorous dispersal. Comparatively small, hard nutlets are eaten their shorter while by birds and transported in digestive tract over longer or distances, the fruits of Cordia are partly digested by mammals and the endocarp is excreted in a viable state. Species of the sea coast such as Tournefortia argentea with a spongy peri- the wind. The carp are dispersed by seawater. Winged nutlets are transported by same is true sometimes in plants with single surviving nutlets that are shed together with the enlarged calyx. FOSSILS There are only a few records of pollen of Cordia and Tournefortia from the Oligocene and Miocene from Mexico, Puerto Rico and the Marshall Islands. Nutlets of still extant and of extinct genera have been reported from the Miocene. Among the latter, epecially Prolithospermum is of interest for the phylogeny ofpresent-day Boraginaceae as demon- strated by Johnston (1954). Literature: Gabel, M.L., Amer. J. Bot. 74 (1987) 1690-1693. — Graham, A. & D.M. Jarzen, Ann. Missouri Bot. Gard. 56 (1969) 308-357; ibid. 63 (1976) 787-842. — Johnston,I.M., J. Arnold Arbor. 35(1954) 1-81. MORPHOLOGY AND ANATOMY Hairs — Hairs are among the most generally distributed features in the family. Unicel- but the of in lular hairs are always present, they are not only type hairs many species. Typical for unicellularhairs are agglomerations of calciumcarbonate or cystoliths in their lumen.To maintaintheirflexibility, there are various devices, most often a specific dis- tributionof thin and thickened segments of the cell wall. In many cases, most character- istically in members of tribe Lithospermeae, the hairs are surrounded by one to several calcified cell that different circles of so called 'accessory cells' with walls, may also be in the ratio of length to width from other epidermal cells. Widespread are bristle-like series of cells the terminal cell of which be horizontal hairs with one longitudinal may or found fruits in order them the fur of oblique. These are glochids mainly on to attach to of hairs less hairs which characterized animals. Other types common are capitate are by undivided heads in Boraginaceae, stellate hairs and hairs composed of more than one series ofcells. Literature: Hummel, K. & K. Staesche, Verbreitung Haartypen. In: Handb. Pflanzenanat., ed. 2, IV, 5 (1962). Inflorescence — The most peculiar morphological featureof Boraginaceae is their inflo- sometimes been called It is with rescence, which has 'boragoid'. sympodial a straighten- ed axis. The main axis is terminated by a single flower. From the axil of the uppermost in terminated leaves one or two side branches originate, which turn are by a flower. In Boraginaceae, either single bracts are developed or bracts are utterly suppressed. The composite axis of the inflorescence is continued by a branch originating from the axil of 46 Flora Malesiana, Ser. I, Vol. 13 (1997) the bract if present. This is not obvious in most cases, however, as the true origin of the that the in side branch is obscured by concaulescence, means, two axes are connate part. In most members of Heliotropioideae and Boraginoideae at least,