She-oaks
This small but distinctive family of shrubs and trees is widely distributed in Australia and extends to South-east Asia and the Pacific Islands. Species have also become naturalised in Africa and the Americas, particulary in coastal habitats. The trees commonly have a drooping habit with slender, green, grooved and jointed branchlets. At the joints are whorls of small, dark-coloured, scale-like leaves; the number of scale-leaves per whorl (between four and twenty) is used in the identification of species.
Flowers are unisexual, and plants are either monoecious or dioecious. The male flower has two scale-like perianth parts and one stamen. Each flower is subtended by a bract, and the flowers are borne in elongated spikes (Pl. 17d, e). The female flower has no perianth parts, and the two fused carpels, with two long reddish styles, are subtended by a bract and two tiny bracteoles. There are two loculi but only one is fertile, and one seed produced. The flowers form small globular heads (Pl. 17g) and, at maturity, the bracts become woody to form the characteristic woody cone (Pl. 17f). The flowers are wind-pollinated.
There are two genera in southern Australia, Casuarina and Allocasuarina. The members of the latter genus were formerly included in Casuarina.
Flora of Australia volume 3 provides keys and descriptions of Australian species.
ILLUSTRATIONS Plate 17d–g.
SPOTTING CHARACTERS
Shrubs or trees, leaves very reduced, scale-like, forming a ring of small teeth at each node of the dark green branchlets.
22 CELASTRACEAE (including STACKHOUSIACEAE)
Bittersweet and Spindle-tree Family
Celastraceae form a medium-sized family, widely distributed in tropical and subtropical regions, with several genera extending into temperate zones. Recent reviews have resulted in the limits of the family being extended to include several smaller ones previously recognised such as Hippocrateaceae and Stackhousiaceae.
Species of several genera, for example Euonymus (Spindle-tree), are occasionally seen in gardens.
FLORAL STRUCTURE
| Flowers | Actinomorphic, usually small and bisexual, often with a conspicuous fleshy disc, sometimes with a short floral tube. Often borne in cymes, sometimes racemes, or solitary. |
| Calyx | Sepals usually 4 or 5, sometimes fewer, free or slightly united. |
| Corolla | Petals usually 4 or 5, sometimes fewer, free or united. |
| Androecium | Stamens usually 3–5, sometimes fewer, free or united. Staminodes sometimes present. |
| Gynoecium | Carpels 2–5, united. Ovary superior to partly inferior. |
| Fruit | Capsule, schizocarp, drupe or berry. Seeds often with a coloured aril. |
Plants are usually glabrous trees or shrubs, some are climbers, a few are herbaceous. Stipules are usually small, or sometimes absent.
The genus Stackhousia and two smaller genera of annual and perennial herbaceous species, were formerly included in Stackhousiaceae, a family of about 19 species almost entirely restricted to Australia. In the flowers of Stackhousia the corolla is tubular in the centre with 5 lobes at the top, as well as splitting into 5 parts at the base. The stamens are of unequal length. The flowers are particularly fragrant at night, and some are pollinated by moths.
Celastraceae and Stackhousiaceae, as separate families, are treated in Flora of Australia volume 22.
ILLUSTRATIONS Figure 75.
SPOTTING CHARACTERS
Flowers often small, greenish or pale-coloured, a fleshy disc often prominent.
Fig. 75 Stackhousia monogyna (Creamy Stackhousia)K5 C(5) A5 G(3) floral tube present
Small, erect perennial herb about 40 cm tall with a few branches from near the base; corolla creamy-white; stamens of unequal length. Widespread in southern and SE Australia. Flowering in spring and early summer. (a ×0.6, b–c ×12, d ×24)
23 ELAEOCARPACEAE (including TREMANDRACEAE)
Oliveberries, Aristotelias and Pinkbells
This is a rather small family of trees and shrubs distributed in eastern Australia, New Zealand, South-east Asia and Japan, southern South America, and the West Indies. DNA analysis has seen the inclusion of the former Tremandraceae, adding about 43 species of small shrubs, all confined to Australia.
Elaeocarpus (Oliveberry) is the largest genus with nearly 300 species. Some of these, as well as species of Aristotelia and Crinodendron, are sometimes seen in gardens. Some species produce edible fruits, and others yield useful timber.
Tetratheca (Pinkbells), from southern Australia, is one of the three genera formerly included in Tremandraceae. The bright pink flowers are often conspicuous in springtime. Flowers of species in the eastern states are 4-partite with 4 sepals, 4 petals and 8 stamens (Fig. 77). The gynoecium consists of 2 united carpels (Fig, 77, Pl. 4l). Most of the Western Australian species and one from Kangaroo Island are 5-partite but have a similar ovary structure.
Fig. 76 Tetratheca ciliata (Pinkbells)
Variable small shrub to c. 1 m tall, with vivid purplish-pink flowers; leaves very variable in size, alternate or opposite, or more often in whorls of 3 or 4; anthers dehisce through a terminal pore (Fig. 77c). Common in heaths and forests across southern Vic. and adjacent parts of SA and NSW, and occasionally grown in gardens. Flowering in spring and early summer. See also Pl. 4l. (×0.7)
| Flowers | Actinomorphic, usually bisexual, often drooping in racemose or cymose inflorescences. |
| Calyx | Sepals usually 4–5, free or partly united. |
| Corolla | Petals usually 4–5, sometimes absent, usually free, often noticeably fringed or torn at the apex. |
| Androecium | Stamens few to numerous, free, often borne on the disc. Sometimes in 5 groups. Connective often prolonged. Anthers often longer than filaments, opening by short terminal slits or terminal pores. |
| Gynoecium | Carpels usually 2–9, united. Ovary superior. |
| Fruit | Usually a capsule or drupe. |
Leaves are simple and alternate (rarely opposite). Stipules are usually present but often readily falling.
ILLUSTRATIONS Figures 76, 77; Plate 4l.
SPOTTING CHARACTERS
Flowers often drooping.
Petals often fringed/torn. Anthers usually longer than filaments, opening by apical pores or short slits; the connective often prolonged. Style simple.
Fig. 77 Tetratheca ciliata (Pinkbells, a–b ×3, c–d ×12)K4 C4 A8 G(2)
St John’s Wort and allies
This is a rather small but widely distributed group, well developed in temperate regions. In recent classifications it has found extra support from molecular studies but is sometimes seen included at the rank of subfamily within the closely related family Clusiaceae. The largest genus, of some 360 species, is Hypericum (St John’s Wort) which includes both weedy and ornamental species as well as some valued for medicinal purposes.
FLORAL STRUCTURE
| Flowers | Actinomorphic, bisexual. |
| Calyx | Sepals usually 4–5, free or slightly united. |
| Corolla | Petals usually 4–5, free, often colourful. |
| Androecium | Stamens 5–many, sometimes in bundles. Anthers dorsifixed. |
| Gynoecium | Carpels 3–5, united; ovary superior; styles often free, each with a very small stigma; placentation axile or parietal (Pl. 4k). |
| Fruit | Often a capsule, sometimes a berry or drupe. |
Plants are trees, shrubs or herbs. Leaves are usually opposite or sometimes alternate or whorled, usually with small translucent or blackish secretory cavities visible as tiny dots. There are no stipules.
ILLUSTRATION Plate 4j, k.
SPOTTING CHARACTERS
Plants shrubby or herbaceous. Leaves often opposite and dotted. Flowers showy, often yellow, the stamens numerous and often in bundles.
25 EUPHORBIACEAE
Spurges
The Spurge Family is a large one, and dominated by the distinctive genus Euphorbia which includes about one-third of the species. The family is very widely distributed but shows greatest diversity in subtropical and tropical regions. Species of several genera are grown as ornamentals including numerous Euphorbia spp., notably E. characias (Dalmatian Spurge, Pl. 18a, b), E. milii (Crown-of-thorns) and E.pulcherrima (Poinsettia). The important food staple cassava is the root of Manihot esculenta, and castor oil is extracted from Ricinus communis. Other species provide timber and medicines. Hevia brasiliensis provides the raw material for most of the world’s natural rubber. Euphorbia paralias (Sea Spurge), well established on coastal sands in south-eastern Australia, is one of a number of weedy species.
FLORAL STRUCTURE
| Flowers | Unisexual, usually actinomorphic but often reduced and borne in highly modified inflorescences. |
| Calyx | Sepals up to 6, sometimes more or absent, free or united. |
| Corolla | Often absent or petals up to 6, free or united, sometimes showy. |
| Androecium | Stamens 1–numerous, free or united. |
| Gynoecium | Carpels usually 3, sometimes more, united; ovary superior, often 3-lobed; styles usually 3, each sometimes (deeply) forked and appearing as 6 (Fig. 79a), or further branched. Loculi usually 3 with one ovule in each (Fig. 78d; Pl 18g). |
| Fruit | Usually a capsule, sometimes breaking up into 1-seeded segments, sometimes a drupe or berry. |
The family includes trees, shrubs, herbs, and vines, usually dioecious, sometimes succulent (resembling a cactus), and often with poisonous milky sap. Variation is large but leaves of most species are alternate and simple. Stipules are often large, sometimes small or absent.
The specialised inflorescence of Euphorbia, called a cyathium, consists of several male flowers and one female flower enclosed in a cup-shaped involucre of 5 fused bracts (Pl. 18b). In the small, weedy, herbaceous annual E. peplus (Petty Spurge, Fig. 78) there is a crescent-shaped gland at four of the five junctions between the bracts. Within the involucre, the female flower consists only of the gynoecium and is exserted over the part of the involucre that has no gland. The male flowers consist of 1 stamen only, with a jointed stalk (Fig. 78c). The part below the joint is interpreted as a pedicel, with the filament above. There are no perianth parts. The cyathium of E. characias (Dalmatian Spurge, Pl. 18b) is very similar.
Another distinctive Australian member of the family, Amperea xiphoclada (Broom Spurge, Pl. 18c–g), is an almost leafless, small, tufted shrub with triangular stems, found in heathlands and poor forest country.
ILLUSTRATIONS Figures 78, 79; Plate 18.
SPOTTING CHARACTERS Milky sap often present. Flowers unisexual, the petals absent or 4–6. Ovary superior, trilocular, and sometimes stalked.
female flowers: P0 A0 G(3)
male flowers: P0 A1 G0
Fig. 78 Euphorbia peplus (Petty Spurge)
Small, herbaceous annual usually up to c. 30 cm tall. Native to Europe but very widely introduced elsewhere, and a common weed of gardens. Flowering winter to summer. (a ×0.7, b ×7, c–d ×30)
female flowers: K(5) C5 A0 G(3)
male flowers: K(5) C5 A(∞) G0
Fig. 79 Ricinocarpos pinifolius (Wedding Bush)
Bushy dioecious or monoecious shrub to c. 1.5 m tall with opposite, narrow-linear leaves 1–3.5 cm long with revolute margins; flowers unisexual, showy with white petals (perianth parts vary from 4 to 6) in small clusters at the ends of the branches; male flowers have filaments united in a central column; ovary of female flowers covered with short, stout hairs, each of the three styles deeply divided. Native to near-coastal heaths of Vic., Tas., NSW and Qld. Flowering in spring. (a–b ×3, c ×12, d ×3, e ×8)
Evening Primroses, Fuchsias and Willowherbs
This is a relatively small family with greatest diversity in the Americas. It is probably best known through the genus Fuchsia, widely cultivated in gardens, but be aware that many ornamental Fuchsias have been modified and no longer show the standard floral structure illustrated in Figure 80 or Plate 19d. Various species of Clarkia, Epilobium (Willowherb), Oenothera (Evening Primrose, Butterfly Bush) and others are also grown for ornament.
The family is included with Myrtaceae and several others in the order Myrtales, and the symmetrical flowers with an inferior ovary, floral tube and often prominent stamens may appear quite similar. Members of Onagraceae however do not possess aromatic oil glands.
FLORAL STRUCTURE
| Flowers | Actinomorphic, bisexual, and nearly always 4-partite. Floral tube often developed, sometimes prominent (Pl. 19d), often producing nectar (Fig. 80b). Flowers borne in indeterminate inflorescences, often terminal, or solitary in leaf axils (Fig. 80). |
| Calyx | Sepals usually 4, usually regarded as free, sometimes united to a small degree at the base as in Fuchsia magellanica (Fig. 80), or united in pairs (at least initially) as in Oenothera lindheimeri (Butterfly Bush, Pl. 19b). |
| Corolla | Petals usually 4, free. |
| Androecium | Stamens usually 8 (in two whorls of 4), sometimes fewer, the whorls often of different lengths. |
| Gynoecium | Carpels usually 4, united. Ovary inferior. Placentation usually axile. |
| Fruit | Usually a capsule or a berry, sometimes a nut. |
Plants are usually shrubs or herbs, often with simple, opposite leaves, sometimes with stipules.
See Flora of Australia volume 18 for an account of this family.
ILLUSTRATIONS Figure 80; Plate 19a–e.
SPOTTING CHARACTERS
Plants often herbaceous as in Epilobium (Willowherb) or shrubby. Flowers actinomorphic, 4-partite, floral tube present (often quite pronounced, Pl. 19d).
Fig. 80 Fuchsia magellanicaK(4) C4 A4+4 G(4) floral tube present
Erect shrub to 2 m, branchlets drooping; leaves opposite, ovate-lanceolate, to 8 cm long, margins serrate; flowers red and purple, in the upper axils; sepals united to a small degree; nectary present at the base of the floral tube. Native to South America, commonly grown, and sometimes escaping from gardens and becoming naturalised. Flowering mostly in summer and autumn. (a ×0.7, b ×2, c ×7, d ×12)
Myrtles, Eucalypts, Bottlebrushes, Tea-trees
This large family is mainly of the southern hemisphere and the tropics, with an extension to South-east Asia. Most genera occur in Australia and South America, but only a few in Africa. The genus Myrtus (Myrtle), from which the family name is derived, is found in the Mediterranean region.
The family has been very important to indigenous Australians. Water was collected from the roots of many mallee eucalypts, and edible grubs and insects found in the foliage and under the bark of numerous species. The wood and bark of eucalypts and melaleucas had a variety of uses.
The family has considerable economic importance. Many species of Eucalyptus are utilised for timber, chipboard, paper-making or extraction of aromatic oils, and they are also important to bee-keepers. Essential oils are also derived from species of Backhousia, Corymbia, Leptospermum (Tea-tree) and Melaleuca. Syzygium smithii (Lilly Pilly, Pl. 19h, i), Lophostemon (Brush Box, Pl. 20a–d) and Angophora (Apple Box) are eastern Australian trees, sometimes used to provide timber for cabinet work and, like Eucalyptus, are widely planted as street trees and ornamentals. Acca sellowiana (formerly Feijowa, Pineapple Guava, Pl. 19f, g) and Psidium guajava (Guava), introduced from South and Central America respectively, are among species that produce edible fleshy fruits. The dried unripe berries of Pimenta dioica of the West Indies are sold as allspice and the dried flower buds of Syzygium aromaticum from the Moluccas are marketed as cloves.
Many well-known ornamentals are among the genera Agonis (Willowmyrtle), Callistemon (Bottlebrush), Chamelaucium (Waxflower), Leptospermum (Tea-tree), Melaleuca (Paperbark or Honeymyrtle) and Thryptomene. Countries in South America, Africa and the Mediterranean region, and also California and New Zealand have successfully introduced eucalypts for use in plantations and as street trees. In some situations eucalypts have become serious weed issues.
In the past, Myrtaceae has been divided into two subfamilies; the predominantly Australian Leptospermoideae, most of which have dry capsular fruits, and the mainly South American Myrtoideae, with succulent fruits. Most southern Australian members of the family belonged to the Leptospermoideae, one of the exceptions being Syzygium smithii (Lilly Pilly, Pl. 19h, i). Research in the last decade or so, particularly molecular studies, has challenged this classification. Two subfamilies are still recognised but Myrtoideae has been expanded to include virtually all of the genera, accommodated in 15 tribes. The other subfamily, Psiloxyloideae, includes just two very small genera of limited distribution, one in the islands of Réunion and Mauritius, the other endemic in south-eastern Africa.
FLORAL STRUCTURE
| Flowers | Nearly always actinomorphic and bisexual, floral tube present. Inflorescences varied. |
| Calyx | Sepals usually 4–5, usually free, united in Eucalyptus. |
| Corolla | Petals usually 4–5, usually free, united in Eucalyptus. |
| Androecium | Stamens usually numerous and free, occasionally 5 (rarely fewer) or 10, sometimes united in bundles, for example Melaleuca (Fig. 91) and Lophostemon (Pl. 20c), often with an apical gland (Fig. 90b). |
| Gynoecium | Carpels commonly 5, sometimes 2–10 or more, united. Usually the number of loculi equals the number of carpels, but occasionally there is only one loculus as in Calytrix (Fig. 83), Thryptomene, and several other genera. Ovary usually inferior, sometimes semi-inferior or superior. A floral tube, the length of which varies with species, is fused with and may extend above the ovary and bears the other whorls of parts. The inside of the tube is often lined with nectar-producing tissue (dark red in Pls. 20f, i). Placentation usually axile. Ovules 1 or 2 to many per cell. |
| Fruit | Often a woody capsule, opening by valves at the top, sometimes dry and indehiscent, or a berry. |
Differences in the proportions of petals and stamens can disguise the real similarities in structure. For example, in Leptospermum (Tea-tree, Fig. 90), the petals form the showy part of the flower and the stamens are relatively short. In the bottlebrushes, the long showy stamens are conspicuous and often obscure the petals (Callistemon, Fig. 82).
Members of the family are shrubs and trees. Leaves are alternate or opposite, simple and mostly entire. Leaves of species from drier areas are often rather leathery (for example in many eucalypts) compared to those of more benign habitats such as rainforests. Stipules, if present, are rudimentary. The leaves, and often other parts of the plant, are dotted with glands containing aromatic oil.
Volume 19 of Flora of Australia deals with the genera Eucalyptus and Angophora. Volumes 20 and 21, as yet unpublished and likely to be available in electronic form only, will complete the account of the family. See also Brophy et al. (2013), George (2002), Govaerts et al. (2008), Holliday (1989, 1997), Wrigley & Fagg (1993).
ILLUSTRATIONS Figures 81–91; Plates 19f–j, 20
SPOTTING CHARACTERS
Plants woody. Flowers regular, stamens often numerous, ovary often inferior. Fruit usually dry, often a woody capsule. Oil glands present in the leaves, usually distinctively aromatic when crushed.
THE GENUS EUCALYPTUS
Eucalyptus is one of the largest genera in the Myrtaceae, with many hundreds of species, almost all of which are restricted to Australia. They are spread throughout the continent in a wide range of habitats and climatic conditions, and are the chief components of the forests of southwestern, south-eastern and eastern Australia. Many species, such as E. pauciflora (Snow Gum), survive freezing temperatures, while others, like the mallees, are adapted to high temperatures and drought.
Classification
The first Eucalyptus species, E. obliqua (Messmate) was described in 1789, from specimens collected on Bruny Island, southern Tasmania, during Cook’s third expedition. By 1866 George Bentham, in his Flora Australiensis, accepted 135 species, grouping them into five series based on features of the stamens. Later botanists also relied on the stamens (particularly the anthers) in devising systems of classification although the weighting of a particular set of characteristics such as this is now considered unsatisfactory. In practice, anther types can be difficult to observe even in young flowers. The three main anther types are:
• macrantherous—anther lobes parallel, dehiscence by parallel slits
• renantherous—anthers reniform (kidney-shaped), dehiscence by slits
• porantherous—anthers dehisce by terminal pores (Fig. 88d).
During the twentieth century, many botanists contributed to the understanding of the genus, which had come to be regarded as a taxonomically difficult group including over 700 species. In 1995 botanists from the NSW Herbarium published a proposal to remove the ‘bloodwoods’ from the genus Eucalyptus and place them in a new genus, Corymbia (Hill & Johnson, 1995). This was part of a formal move to reclassify eucalypts, and followed many years of study, discussions, and proposals, some of which, although published, were nevertheless intended only as informal reviews (e.g. Pryor & Johnson, 1971). At various times up to nearly a dozen genera have been suggested as warranting recognition alongside a more restricted interpretation of Eucalyptus.
Most bloodwoods are native to northern Australia but examples common in cultivation in the south-east include Corymbia ficifolia (syn. Eucalyptus ficifolia, Western Australian Flowering Gum) and C. citriodora (syn. E. citriodora, Lemon-scented Gum). The natural ranges of C. gummifera (Bloodwood) and C. maculata (Spotted Gum) extend south into Victoria in the far east.
The genus Corymbia, with more than 90 species, is characterised by the combination of a number of rather technical features including well developed tertiary venation in the leaves, and the presence of elongated bristle-glands. Several microscopic and anatomical features are also important. These features are shared by the closely related genus Angophora from which Corymbia is distinguished by its possession of an operculum, and ‘alternate’ adult leaves. In addition, corymbias bear their flowers in ‘terminal panicles’ which often results in the canopies being conspicuous at flowering time. They also produce urn-shaped fruits, their leaves have secondary veins at a large angle to the midrib, and the bark is often distinctively rough and flaky (tessellated). However, as happens in many plant groups, not all species possess all of these features (i.e. exceptions occur), and further, some of these characteristics occur in other eucalypts. This situation (overlap of ‘distinctive’ characters) occurs occasionally in classification, and it is up to the botanist, having weighed all the evidence, to make the decision as to where to draw the lines between the various groups.
It should be remembered that plant classification is continually refined over time with the benefit of more information and increasing sophistication in techniques. As well, interpretation of the evidence is sometimes not unanimous—opinions differ, and botanists may hold different views particularly over the ranks to be assigned to various groups. With respect to the genus Eucalyptus, an example of this situation is the proposal to broaden the generic concept (to include the closely related genus Angophora) rather than to subdivide it into a number of smaller genera (Brooker, 2000). In this case those species now in Angophora would change their names to Eucalyptus, and of course those species assigned to Corymbia would remain in Eucalyptus.
For the non-expert, perhaps the main question now becomes ‘Well, what proper name do I apply to the Western Australian Flowering Gum?’ Whether one uses Corymbia ficifolia or Eucalyptus ficifolia is really a matter of personal choice, realising that behind the use of either of these names is the acknowledgement of one alternative view of the classification of the group. In time, one view of the classification of eucalypts will probably prevail, and the use of one or other of the scientific names for the Western Australian Flowering Gum will come to be accepted. At the time of writing of this third edition, Corymbia appears to be withstanding the test of time.
For a summary of evolutionary relationships within Eucalyptus (in a broad taxonomic sense) see West (2006).
Plant structure
The structure of the open flowers of Eucalyptus is relatively uniform, and the identification of species is largely based on characters of the buds, fruits, leaves and bark. The following notes refer to Eucalyptus as it is traditionally known, that is, including Corymbia but excluding Angophora.
Buds
The eucalypt bud has three main parts: a pedicel (which may be absent), floral tube (variously called a calyx tube, hypanthium, thalamus tube or torus) and an operculum (Fig. 87b). The operculum, which is deciduous, is formed by one or two whorls of united perianth parts. Some species with a double operculum shed the outer or sepaline one as the bud develops, leaving a circular scar on the outside of the bud. A scar is not seen when the opercula fall together or when the operculum is single.
Bud characters used in identification include:
• pedicel—presence or absence; shape in T.S. (pedicels are often flattened)
• length of the floral tube measured from the top of the pedicel, relative to the operculum
• shape of the operculum—conical, rounded, beaked etc.
• surface characters—whether smooth, glaucous, warty, ribbed etc.
• inflorescence type—buds may be solitary or arranged in umbel-like clusters (usually simply called umbels, Fig. 85) or panicles; the number of buds per umbel, and characters of the peduncle are also used.
The eucalypt flower has numerous stamens borne on the floral tube. The appearance of the zone of attachment of the stamens varies (cf. Figs. 86a and 88b) and is variously called the staminal ring or the staminophore. In bud, the stamens are either all inflexed towards the style, all erect, or inflexed but with the anthers lying irregularly around the style; at anthesis the filaments more or less straighten and spread out. In some species the outermost stamens are replaced by staminodes (Fig. 88b).
Fruit
The fruit is a woody capsule with usually 3–5 valves through which the seeds are released. The number of valves may vary even in fruits from the same tree. At the fruiting stage the opercular scar is known as the rim, and the scar left by the stamens may also be distinct (or simply considered part of the rim by some authors). Sometimes the staminophore is relatively large and may persist as a withered brown ring on the developing fruit as in E. leucoxylon (Yellow Gum, Fig. 88) and E. tricarpa (Red Ironbark). The tissue between the rim and the base of the valves is the disc, and this may be flat, ascending (Fig. 85b) or descending (Fig. 87c). The valves open outwards and their position in relation to the rim is described as level, exserted (Fig. 85b) or enclosed (Fig. 87c).
Leaves
The leaves of most Eucalypts change as the plant grows from seedling to adult form (Fig. 84). Juvenile leaf characters are sometimes used to distinguish between closely related species such as E. rubida (Candlebark) and E. viminalis (Manna Gum). The adult leaves of most species are tough and leathery and tend to hang vertically from the branches. Leaves form in opposite pairs but in the majority of species adult leaves finally appear alternate due to unequal elongation of the axis between the leaf bases. A notable exception is Eucalyptus crenulata (Buxton Gum), quite common in cultivation, which retains the distinctive juvenile leaf phase on the mature tree. Most of the aromatic oils found in the oil glands of the leaves have a ‘eucalyptus’ odour, even though the constituent oils vary with species. The smell of crushed leaves of the peppermint group (e.g. E. radiata, Narrow-leaf Peppermint) is often quite distinctive. Many eucalypt leaves are slightly sickle-shaped, others are asymmetrical, with the two halves of the lamina joining the petiole at different points. This feature of its leaves led to the name E. obliqua (Messmate), a widespread forest tree in southern Australia.
Leaf venation is another useful feature for identification. Secondary veins are sometimes quite widely separated and almost parallel to the midvein (e.g. E.pauciflora, Snow Gum), and range to being closely spaced at a wide angle to the midvein as in Corymbia maculata (Spotted Gum) in the bloodwood group, and E. botryoides (Southern Mahogany) in the mahogany group. The distance from the leaf margin to a vein running roughly parallel to the margin (the intramarginal vein) may also be useful.
Bark
The type of bark is a considerable help in the identification of species, particularly in the field. The main categories are nonpersistent and persistent. In the non-persistent type, the outer layer of bark is shed each season, leaving the trunk and branches smooth and light-coloured. This type is often referred to as ‘gum bark’ and may be shed completely, in irregular patches, or, for a time, hang from the branches in long ribbons.
In the persistent type, the outer layer is not shed regularly but tends to flake off gradually, so the surface of the trunk is rough and mostly dark-coloured. Several variations are recognised:
• stringybark—thick, often furrowed, with long coarse fibres; the bark can be pulled off in strips
• peppermint—fibrous but relatively compact; fibres of medium length, not easily pulled off
• box—thin, compact, short-fibred, sometimes breaking horizontally into irregular flakes
• ironbark—deeply furrowed, dark and hard due to gum deposition
• bloodwood—tessellated (that is, cracking vertically and horizontally to form small plates).
Form classes
The majority of eucalypts can be divided into three general form classes:
• Forest trees have a tall single stem with small branches high above the ground forming a relatively small crown.
• Woodland trees have a relatively short, single stem, before branching to form a spreading crown.
• Mallees have a number of thin stems arising at ground level. The stems are usually less than 10 m high, with a small crown.
At or below ground level, most eucalypts develop a swollen woody structure called a lignotuber, which contains food reserves and many dormant vegetative buds. These buds rapidly develop into new shoots when the aerial parts of the plant are destroyed, for example, by fire or harvesting for oil distillation. Mallee eucalypts have particularly well-developed lignotubers, which are collected from cleared land and sold for fuel as mallee roots. Eucalypts also recover from fire in other ways. When only the crown and small branches are killed, concealed buds just under the bark of the trunk and branches sprout to form a mass of epicormic shoots. When the crown has been re-established most of these shoots die off. This type of regeneration is typical of the stringybarks, though not restricted to them. Species with thin bark and no lignotuber, such as E. regnans (Mountain Ash), are readily killed by fire and must regenerate from seed. The capsules, which protect the seeds from all but very severe fires, release them soon after the tree has died, and when conditions are suitable a thick growth of seedlings will appear.
Flora of Australia volume 19 includes numerous line drawings, keys, and descriptions for all eucalypt species recognised at the time of publication (1988), but there has been much research into the genus since then and numerous new species have been described. Brooker & Kleinig’s Field Guide to Eucalypts (volumes 1–3) probably remains the most comprehensive and useful identification text, but see also CPBR (2006) in the reference list for a DVD.
Native trees and shrubs of south-eastern Australia (Costermans, 2009) is a valuable introduction to the eucalypt species of the region, and the family in general. Trees of Victoria and adjoining areas (Costermans, 2006) is an excellent starting point for learning eucalypts and a most useful field guide. Other works include Brooker & Kleinig (1996), and Nicolle (1997), (2006).
Fig. 81 Callistemon citrinus (Crimson Bottlebrush, ×0.5)
Fig. 82 Callistemon citrinus (Crimson Bottlebrush)K5 C5 A∞ G(3) floral tube present
Shrub 1–3 m tall, branches often arching over; young stems and leaves covered with deciduous, silky hairs; leaves lanceolate, up to 8 cm long; flowers red, borne in terminal spikes, the apical bud remaining vegetative and the shoot continuing to grow beyond the inflorescence; fruit a capsule, often retained on the plant for several years. Common in swampy heaths of eastern Vic., NSW and Qld. Widely cultivated, with many cultivars and hybrids. Flowering in spring to early summer, sometimes autumn. (a–b ×4, c ×7)
Fig. 83 Calytrix tetragona (Common Fringe-myrtle)K5 C5 A∞ G(loculus 1) floral tube present
Erect shrub 1–2 m tall; leaves variable, usually less than 1 mm long, usually hairy, blunt or with a small mucro; flowers white to pink, numerous and forming corymbose clusters; sepals often tapering into long awns; ovary structure and placentation unusual (the two ovules, surrounded by a loose membrane, are attached laterally to a thread-like placenta extending from top to bottom of the loculus); fruit one-seeded, crowned by the persistent calyx. Widespread in temperate areas of all states except NT. Flowering in spring. (a–b ×6, c ×24)
Fig. 84 Eucalyptus globulus ssp. globulus (Tasmanian Blue-gum, Southern Blue-gum)
Medium to very tall tree to 60 m; trunk stout, branches relatively small, crown spreading when open-grown; bark of the gum type, peeling in strips so the trunk is streaked with dark and pale grey, blue-grey, cream and brown; adult leaves alternate, lanceolate, to 30 cm long, dark green; juvenile leaves opposite, sessile, ovate, glaucous; intermediate leaves very long, up to 60 cm; flowers white, solitary, sessile, axillary; fruit a capsule. Widespread in Tas., restricted to the Otways and South Gippsland in southern Vic. Commonly grown in parks and larger gardens, and valued for timber. Flowering winter to spring. (a–b ×0.5)
Fig. 85 Eucalyptus camaldulensis (River Red-gum)
(a–b ×4)
Fig. 86 Eucalyptus camaldulensis (River Red-gum)P operculum A∞ G(4) floral tube present
Variable, medium to tall tree to 45 m high; trunk thick, main branches heavy, crown spreading; forest forms more upright, with a smaller crown; the gum bark is variable in colour, grey, brown, or pinkish or white, and often patchy; adult leaves lanceolate, usually 10–15 cm, sometimes to 25 cm long, dull green; juvenile leaves opposite at first, then alternate; broad lanceolate; flowers white or creamy, 7–11 in axillary umbels; fruit a capsule with exserted valves. Widespread along the river systems of all states except Tas. (a–b ×7)
Fig. 87 Eucalyptus leucoxylon ssp. megalocarpa (Large-fruited Yellow-gum)
(a ×0.6, b–c ×1.2)
Fig. 88 Eucalyptus leucoxylon ssp. megalocarpa (Large-fruited Yellow-gum)P operculum A∞ G(5) floral tube present
Small tree to about 20 m; bark rough, box-like for about 2 m, then gum-type, whitish, cream or yellowish-grey; adult leaves lanceolate, dull green, to about 10 cm long; juvenile leaves opposite, sessile, ovate, dull green; flowers usually red or pink, and usually in threes in axillary umbels; the ring of tissue to which the stamens and staminodes are attached often persists in the fruiting stage, and may be called a staminophore or staminal ring; fruit a capsule, valves enclosed. Mainly restricted to coastal calcareous regions of western Vic. and south-eastern SA. Commonly cultivated. Flowering mainly in autumn. (a–c ×2.5, d ×25)
Fig. 89 Kunzea ambigua (White Kunzea)K5 C5 A∞ G(3) floral tube present
Erect shrub to 3 m; branchlets hairy; leaves crowded, lanceolate-oblanceolate, up to 1 cm long, concave above; flowers white or creamy, in dense clusters on very short lateral branches, each flower, at least when young, with bracts at the base (some authors state bracts are absent); capsules non-woody, with persistent calyx. Wilsons Promontory and coastal in eastern Vic., NSW and Tas., sometimes becoming naturalised outside its original range. Flowering in spring to summer. Cf. Pl. 20e–g. (a–c ×7)
Fig. 90 Leptospermum myrsinoides (Silky Tea-tree)K5 C5 A∞ G(5) floral tube present
Shrub to 2 m tall; leaves oblanceolate, up to 1 cm long, concave, dull green, and glabrous; flowers white or pink, terminal on short lateral branches; fruit a nonwoody capsule. Widespread in heathlands and sandy forests, mainly near-coastal in Vic., SA and NSW. Flowering in spring. Cf. Pl. 20h, i. (a–c ×7)
Fig. 91 Melaleuca linariifolia (Snow-in-summer)K5 C5 A(∞) G(3) floral tube present
Shrub or small tree to 10 m; leaves lanceolate and slightly concave, 2–3 cm long; flowers white, borne in spikes near the shoot apex which continues to grow; stamens united by filaments into 5 bundles (in some species the union is not so obvious); fruit a capsule that persists unopened for some years. Coastal and damp localities in NSW and Qld. Widely planted as an ornamental and street tree, sometimes becoming naturalised. Flowering in summer. Cf. Pl. 19j. (a ×5, b–c ×7)
Rue, Citrus, Boronias and Correas
This is a medium-sized family, widely distributed in temperate and tropical regions, especially in Australia and South Africa. The Australian representatives total about 43 genera and over 480 species. The name Rutaceae is derived from the genus Ruta, which includes R. graveolens (Rue), a European herb supposedly possessed of many healing properties.
Many species are scented due to the presence of oil glands, most readily seen in their leaves and flowers, and some, including boronias, are grown commercially to provide oils for use in perfumes. Several genera have edible fruits, the most important being the citrus group which includes grapefruit, kumquats, lemons, limes, mandarins and oranges. Some Australian natives, Citrus australasica (Finger Lime, formerly Microcitrus) and C. glauca (Desert Lime, formerly Eremocitrus), bear fruits that are similar to the cultivated citrus and these were used by Indigenous people and colonists. With increasing interest in Australian bush foods these species are slowly returning to favour.
Numerous native and introduced genera are grown as ornamentals, for example Boronia, Correa, Philotheca (previously Eriostemon), Choisya (Mexican Orange), Coleonema (Diosma) and Murraya. Calodendrum capense (Cape Chestnut) forms a handsome, floriferous tree occasionally seen in parks and larger gardens.
FLORAL STRUCTURE
| Flowers | Usually actinomorphic and bisexual, sometimes unisexual. Inflorescences various. |
| Calyx | Sepals 4–5, sometimes fewer, free or united, inconspicuous in Asterolasia. |
| Corolla | Petals 4–5, usually free, sometimes united as in some species of Correa (Fig. 93). |
| Androecium | Stamens usually 8 or 10, in 2 whorls of 4 or 5, sometimes united at the base. In Citrus there are 15 or more. In Crowea each anther is tipped with a prominent bearded appendage (Fig. 94). Staminodes are quite common. |
| Gynoecium | Carpels usually 4–5, sometimes free, commonly fully united, sometimes free at the base and then united at the top to form a single style and stigma as in Boronia (Fig. 92; Pl. 4i) and Philotheca (Fig. 96). Ovary superior. Placentation usually axile with 1–2 ovules per loculus. A disc is often present at the base of the ovary (Figs. 92, 96; Pl. 4h, i). |
| Fruit | Of various types. Commonly dry and leathery, splitting into segments at maturity, or a berry as in the citrus group, sometimes a capsule or samara. |
The majority of plants are shrubs or trees, sometimes thorny or spiny, and most have alternate leaves, often compound, that have no stipules. Boronia, Correa and Zieria (Fig. 97) have opposite leaves, which is a useful spotting character for these genera.
Volume 26 of Flora of Australia provides keys and descriptions for Australian species.
ILLUSTRATIONS Figures 92–8; Plate 4h, i
SPOTTING CHARACTERS
Plants usually woody. Leaves contain oil glands (leaf blades often dotted) and may be pleasantly aromatic when crushed. Other surface features include stellate hairs and peltate scales (Fig. 95d). Flowers actinomorphic, 4- or 5-partite, stamens often double the number of petals. Ovary superior, disc often present.
Fig. 92 Boronia mollis (Soft Boronia)K4 C4 A4+4 G(4)
a flower, side view; b half flower (the plane of section has passed between the loculi); c L.S. carpels (loculi and ovules sectioned); d T.S. ovaries, disc and bases of filaments, placentation axile. (a–b ×7, c–d ×12)
Small shrub to 2 m tall; branches densely hairy; leaves opposite, pinnate with 3–7 leaflets; flowers pink, in small axillary or terminal inflorescences; pedicels, calyx and filaments bearing stellate hairs; outer whorl of stamens longer than the inner; disc prominent surrounding the ovary and filament bases; carpels united by the single style, the ovaries free. Native in sandstone gullies inland and south of Sydney, NSW. Sometimes cultivated. Flowering in spring. Cf. Pl. 4h, i.
Fig. 93 Correa reflexa (Common Correa, Native Fuchsia)K(4) C(4) A4+4 G(4)
Variable, small shrub to 2 m tall; young branches, leaves, and flowers with stellate hairs; leaves opposite, 2–5 cm long, lanceolate to ovate; flowers usually pendulous, one or few at the ends of small lateral branches; flower colour variable, red, red with green tips, or all green; bracteoles borne on the pedicel, sometimes falling early; inner whorl of stamens with broad bases to the filaments; carpels united, ovaries separating at maturity. Native and common in all states. Flowering late autumn to spring. (a–b ×2.5, c ×13)
Fig. 94 Crowea
Crowea is a small Australian genus of pink-flowered shrubs, including three species and several cultivars that are commonly grown. It is closely related to Philotheca and has the same floral formula. One of the key differences is the hairy anthers: note the prominent, bearded terminal appendages. (a–b ×12)
Fig. 95 Phebalium squamulosum (Forest Phebalium)K(5) C5 A5+5 G(5)
Erect shrub of variable form, most parts of the plant covered with silvery or brownish, overlapping scales; leaves alternate, narrow-oblong to elliptical, 1–7 cm long; flowers cream to yellow, borne in small terminal umbel-like clusters; each anther bears a small, rounded, apical gland; ovaries densely scaly, united at the base and by the single style. Native in forests of eastern Vic., SA, NSW and Qld. Sometimes grown for ornament. Flowering in spring. (a–b ×10, c–d ×20)
Fig. 96 Philotheca myoporoides (Long-leaf Waxflower)K5 C5 A5+5 G(5)
Variable shrub to 5 m tall; stems glabrous, rough due to prominent oil glands; leaves alternate, lanceolate, aromatic; buds pink, flowers white in stalked axillary umbels; filaments hairy but anthers glabrous, tipped by a small point; ovaries united by the base of the single style, each ovary with a prominent terminal lobe; disc prominent around the base of the ovaries. Occurs naturally in eastern Vic., NSW and Qld., and often grown for ornament. Flowering in spring and summer. Formerly known as Eriostemon myoporoides. (a–b ×5, c–d ×10)
Fig. 97 Zieria arborescens (Stinkwood)
(×0.7)
Fig. 98 Zieria arborescens (Stinkwood)K4 C4 A4 G(4)
Tall shrub or small tree to 5 m; leaves opposite, trifoliolate, leaflets 3–10 cm long with an unpleasant odour; branchlets and undersides of leaves covered with minute stellate hairs; flowers white, small, borne in axillary cymose panicles; disc 4-lobed; stamens with warty filaments. Occurs in damp forests and gullies of Tas., Vic., NSW and Qld. Flowering late winter to early summer. Occasionally cultivated. (a–c ×10)
Mallows, Baobabs, Lindens and Bottle Trees
As defined in recent classifications supported by molecular research, Malvaceae is now a large, cosmopolitan family. The traditional boundaries of the family have been broadened to take in Bombacaceae (Baobabs and Balsa Trees), Sterculiaceae (Kurrajongs, Bottle Trees and Paperflowers) and Tiliaceae (Lindens). Kept separate, these four families are considered not to be monophyletic, and the boundaries between them to be rather arbitrary. Nine subfamilies have been recognised.
The following paragraphs introduce these four former groups:
Bombacaceae: a small group, mostly trees from tropical South America, including Adansonia (Baobab) and others with distinctive swollen trunks, Ochroma pyramidale the source of balsa wood, and Ceiba and Bombax spp. from which the fibre kapok is derived.
Sterculiaceae: quite a large group, mainly tropical and subtropical trees and shrubs, with several genera extending into more temperate zones. The type genus Sterculia, of some 150 species, includes some native in northern Australia. Various species are valued for timber, oil, and edible seeds among other uses. Brachychiton rupestris (Queensland Bottle Tree) with a distinctive swollen trunk, and B. acerifolius (Flame Tree, Pl. 22a–e) with striking red flowers, are grown for ornament. Lasiopetalum (Pl. 22f, g) and Thomasia (Fig. 100), both endemic Australian genera of small shrubs, are seen in native gardens. Theobroma cacao, originally from South America but now much cultivated, is the source of chocolate.
Tiliaceae: a small, widely distributed group of trees and shrubs, mostly tropical, some extending to temperate areas. Corchorus spp. provide the fibre jute, and Tilia (Linden) trees are valued for timber and ornament.
Malvaceae: a large, cosmopolitan group of trees, shrubs and herbs. Many ornamentals are seen in gardens including Alcea (Hollyhock), Abutilon (Lantern Flower), Alyogyne (Native Hibiscus), Hibiscus (Pl. 21h), Malva (Mallow), and Malvaviscus (Wax Mallow, Fig. 99; Pl. 21i). Gossypium sturtianum (Sturt’s Desert Rose), a widely distributed native species in inland mainland Australia, is the floral emblem of the Northern Territory. Other Gossypium species are of considerable economic value as the source of cotton. Common weeds include Lavatera spp., Malva nicaeensis (Mallow of Nice, Pl. 21f, g), M. parviflora (Small-flowered Mallow, Pl. 21e) and other Malva spp., Modiola caroliniana (Red-flowered Mallow), and Sida rhombifolia (Paddy’s Lucerne).
Fig. 99 Malvaviscus arboreus (Wax Mallow, Turk’s Cap)
Spreading shrub to 2 m; leaves broadly ovate, to 15 cm long, hairy, becoming 3-lobed at the apex; flowers bright red (Pl. 21i), solitary in the axils, epicalyx of bracts surrounding the tubular calyx; petals joined to the filament tube which surrounds, but is free from, the style. Native to South America, sometimes grown for ornament. Flowering mostly in summer to autumn. (a–b ×2, c ×2.5, d ×2, e ×7)
| Flowers | Usually actinomorphic, bisexual or unisexual, often with extra segments forming an epicalyx (Fig. 99a; Pl. 21h, i). Inflorescences diverse. |
| Calyx | Sepals usually 5, commonly united (Fig. 99a; Pl. 21b, h), sometimes free. |
| Corolla | Petals usually 5, free, sometimes absent (as in Brachychiton, Pl. 22b–d), often joined to the base of a column of united filaments (Fig. 99b; Pl. 21c). |
| Androecium | Stamens 5 to numerous, the filaments often united into a tube (filament column, staminal column, staminal tube) around the style (Fig. 99b; Pl. 21c, h). Staminodes often present. |
| Gynoecium | Carpels few to many, united (sometimes regarded as free, e.g. in Brachychiton Pl. 22b, d). Ovary superior. Placentation usually axile (Fig. 99e; Pl. 21d). |
| Fruit | Numerous types. Often breaking into segments (mericarps) each derived from 1 carpel. |
Plants are mostly trees or shrubs, sometimes lianes or herbaceous. Leaves are usually alternate, the blades often quite broad and lobed or toothed, the main veins palmate and ending in the marginal teeth. Stipules usually present.
ILLUSTRATIONS Figures 99, 100; Plates 21, 22.
SPOTTING CHARACTERS
Plants often with stellate hairs (Pl. 22g), sometimes peltate scales. Stipules usually present. Leaf blades often with palmate venation. Flowers actinomorphic, the sepals often united and an epicalyx often present. Stamens numerous, the filaments often united into a column around the style. The weedy annual or biennial Mallows have distinctive, broad, orbicular leaf blades (Pl. 21f).
Fig. 100 Thomasia petalocalyx (Paper Flower)K(5) C0 A5 G(3)
Small shrub to c. 1 m, with a covering of stellate hairs; stipules large, leaf-like; sepals mauve-lilac, with a prominent midrib; petals absent or very small, dark red and opposite the stamens which dehisce through terminal pores; fruit a capsule. Native to WA, SA and Vic. often coastal, sometimes grown in gardens. Flowering in spring and summer. Cf. Pl. 22f, g. (a ×2, b ×6, c ×20)
Daphnes and Riceflowers
The Thymelaeaceae form a small family with greatest diversity in temperate and tropical regions of the southern hemisphere, particularly Africa. Most species are trees and shrubs. The genus Thymelaea, from which the family name is derived, is native to Asia and the Mediterranean, and one species has been recorded as (sparingly) naturalised in South Australia and Victoria. Daphne, with highly fragrant flowers, is a common garden shrub.
In south-eastern Australia, an interesting member of the family is Kelleria dieffenbachii (formerly Drapetes tasmanica), one of the so-called cushion plants. It is also found in New Guinea and New Zealand. Plants are small-leaved and densely branched, forming a low compact mound usually only a few centimetres high.
Of about eight genera in Australia, all but Pimelea (Riceflower) are very small. Pimelea is widespread, with some 90 endemic species found in a range of habitats. A small number are cultivated in native gardens and for revegetation programs. Several other species extend to New Zealand, Lord Howe Island and the Philippines.
The flowers of Pimelea (Fig. 101) are small, bisexual or unisexual, and commonly clustered in terminal heads surrounded by 4 or more involucral bracts. The four white, yellow or pink sepals extend from the top of the slender floral tube. Petals are absent, and the 2 stamens are inserted near the top of the tube. Sometimes this arrangement is interpreted as a calyx tube with episepalous stamens, and in other genera extra small ‘scales’ are variously described as petals or staminodes. The ovary is superior, with a single loculus containing 1 pendulous ovule. (There are two carpels but one is sterile and fails to develop.) The fruit is a drupe or nut that is enclosed in the persistent lower part of the floral tube.
The leaves of Pimelea are entire, often small, opposite or alternate and without stipules. The bark on the stems and branches of some species such as P. axiflora (Bootlace Bush) is very strong, and colonists often used strips of it in place of twine.
Flora of Australia volume 18 includes keys and descriptions of Australian species.
Fig. 101 Pimelea glauca (Smooth Riceflower)K4 C0 A2 G loculus 1 floral tube present
Small bushy shrub, 0.5 m tall, with opposite leaves; flowers creamy-white, sometimes unisexual, in terminal heads surrounded by 4 bracts (the inner 2 with ciliate margins). Widespread in SA, Vic., Tas., NSW and Qld. Flowering in spring. (a ×1, b–c ×7, d ×20)
Cabbages, Mustards and Wallflowers
Brassicaceae form quite a large cosmopolitan family, mostly of herbaceous species, widely distributed in cold to temperate regions (particularly in the northern hemisphere), as well as drier parts. The older name Cruciferae, still acceptable under the nomenclatural rules, is based on the Latin word for a cross and alludes to the shape of the four-petalled flowers.
The family is important for food plants and sources of condiments and vegetable oils such as canola. Numerous species are grown for ornament and stock feed, and many are weeds of waysides and agricultural cropping. Isatis tinctoria provides the traditional blue dye known as woad. Edible examples include Brassica rapa (Chinese Cabbage, Turnip), B. oleracea (Cabbage) including all its related forms such as broccoli, brussels sprouts, cauliflower, kale, and kohlrabi, Eruca vesicaria ssp. sativa (Rocket) and Raphanus sativus (Radish). A number of genera provide different types of salad cress. Mustard is prepared from the seeds of several Brassica spp. as well as Sinapis alba (White Mustard). Horse-radish comes from the roots of Armoracia rusticana, a native of south-east Europe.
Some common garden species include Erysimum cheiri (Wallflower), Iberis sempervirens (Candytuft), Lobularia maritima (Sweet Alyssum), Lunaria annua (Honesty) and Matthiola incana (Common Stock).
Numerous species have been introduced to Australia and are now weeds. Many have yellow flowers and are conspicuous along roadsides. Nearly 100 species are recorded in the Victorian flora but more than half of these are introduced.
FLORAL STRUCTURE
Plants are usually annual, biennial or perennial herbs, sometimes shrubby, rarely small trees or vines. Leaves are usually simple, the margins often slightly to deeply pinnately dissected, often varying on the one plant from a basal rosette to those on the stem, and from the lower to the upper stem. Stipules are absent. When crushed, the leaves may release a sharp or unpleasant taste or odour due to the production of mustard oils, a characteristic of the family.
For keys and descriptions of Australian species see Flora of Australia volume 8. Rich (1991) describes and illustrates species of Britain and Ireland, many of which are introduced elsewhere.
ILLUSTRATIONS Plates 23, 24a–e.
SPOTTING CHARACTERS
Plants herbaceous, leaves often characteristically malodorous when crushed. Inflorescence often a raceme. Flowers (sepals and petals) 4-partite, stamens 6. Fruit structure distinctive (Pls. 23g, 24e).
Mistletoes
The Mistletoe Family, Loranthaceae, is closely related to Santalaceae (Sandalwoods) and Viscaceae. In the latest APG IV classification, Viscaceae has been merged into an enlarged Santalaceae. Members of these families are root or stem parasites. In Loranthaceae most species are stem parasites, and some are found on several hosts, including eucalypts, wattles, she-oaks and introduced trees such as oaks, birches, Liquidambar, planes and fruit trees. Others are specific to one host genus only. The Western Australian Christmas Tree, Nuytsia, is a root parasite.
Loranthaceae is mostly found in the tropics and subtropics. The type genus, Loranthus, at one time including about 600 species, is no longer considered to occur in Australia. Those Australian species once thought to belong there are now assigned to Amyema and some other genera.
The seeds of Loranthaceae are surrounded by a sticky layer rich in glucose and are an important food source for birds, especially the Mistletoe Bird. The seeds pass through the bird’s gut in 3–12 minutes and, as the bird twists when defecating, the seeds usually drop on a branch. The sticky layer is still present and the seed adheres to the branch, where it germinates readily (Pl. 24g). The parasite is attached to the host stem by one or more structures called haustoria, through which the respective vascular systems are linked.
Amyema (Pl. 24f, g) is the largest genus of mistletoes in southern Australia. The bisexual flowers are borne in axillary umbel-like groups of 2 or 3. Each flower is usually subtended by a bract. The calyx is reduced to a rim of tissue on top of the inferior ovary. Petals are 4–6 in 1 whorl, free or united into a tube, which is often split down one side. Stamens are equal in number to the petals and epipetalous with basifixed anthers. Ovary structure is obscure, and the ovules are undifferentiated. The fruit is berrylike and contains 1 seed. The stems are rather brittle, usually with thick, simple, opposite leaves, with parallel venation. There are no stipules. In many cases the foliage of the parasite is visually similar to that of the host.
For descriptions and keys to Australian species, see Flora of Australia volume 22. See also Watson (2011).
Sundews
Even though forming a very small family, the Sundews have a world-wide distribution. There are only three genera, two of which contain just one species. Plants are annual or perennial carnivorous herbs. The leaves are modified for trapping insects which are then digested, supplementing the nutritional intake from photosynthesis and from rather poorly developed root systems.
The leaf blades of Dionaea muscipula (Venus Flytrap) from south-east North America are broad, ‘hinged’ along the centreline and able to fold inwards. Marginal teeth intermesh as the leaf folds over to ensnare prey insects attracted by secretions from the leaf margins.
Aldrovanda vesiculosa (Waterwheel) is a free-floating, rootless, aquatic herb, widely distributed through Europe and Asia, and extending to Africa and north-eastern Australia. Leaves are borne in whorls of 5–9 or more, with the blades forming traps in a similar way to those of Dionea.
The genus Drosera (Sundew, Pl. 24h) includes about 100 species, most of which are found in Australia. Sundews are annual or perennial herbs, commonly growing each season from a tuberous structure which lies dormant underground during the summer. They vary in habit from rosettes to small erect plants or straggly scramblers with stems up to 1 m or more long. The leaves may be more or less round, or strap-shaped or spoon-shaped, or sometimes forked, and the upper surfaces are covered with sensitive glandular hairs. Small animals, such as ants or mosquitoes are trapped by a sticky substance exuded by the hairs, which then fold over to prevent escape. Other hairs secrete enzymes that digest the prey.
The flowers of Drosera are regular and bisexual, mostly with 5 sepals, 5 white or coloured petals, and usually 5 stamens. The ovary is superior, unilocular with 2–5 styles (which may be further divided) and 3–many ovules on parietal placentas. The fruit is a capsule.
Droseraceae is covered in volume 8 of Flora of Australia. See also Carniverous plants of Australia by Lowrie (the earlier volumes are in a small field guide style; those published in 2013 are in large format).
Campions, Catchflies, Chickweeds and Pinks
The Chickweed Family is quite large, mostly herbaceous, and widely distributed in temperate regions, with a centre of diversity in the Mediterranean and adjacent Europe and Asia. Traditional classifications have divided the family into three subfamilies. These groups have not been well supported by recent research, and a current proposal sees the subfamilies abandoned in favour of a larger number of smaller groups recognised at tribal level. Research continues at the generic level also, and changes in circumscription can be expected.
Many species are garden favourites, and possibly a similar number are common weeds. Ornamentals include numerous cultivars of Dianthus (Carnation, Pink, Sweet William), as well as Cerastium (Mouse-ear Chickweed), Saponaria (Soapwort) and Silene (Campion, Catchfly, and now encompassing Lychnis). The multi-branched inflorescences of Gypsophila (Baby’s Breath) are valued in floristry. Weedy species can be found among genera such as Cerastium, Moenchia, Polycarpon, Sagina, Silene, Spergula, Spergularia, and Stellaria.
Colobanthus, a southern temperate genus, is the only ‘Dicot’ to venture into the Antarctic Circle. Arenaria bryophylla reaches the highest altitude of any plant on Mt Everest.
FLORAL STRUCTURE
| Flowers | Usually actinomorphic and bisexual, and often arranged in clearly cymose inflorescences (Pl. 25a, f), sometimes solitary. |
| Calyx | Sepals usually 4–5, free (Pl. 25h) or united (Pl. 25e). |
| Corolla | Petals usually 4–5, free, often deeply notched (Pl. 25g, k) or markedly differentiated into a narrow basal claw and broader upper limb (Pl. 25d), sometimes absent. |
| Androecium | Stamens often 5–10, sometimes fewer, usually free (Pl. 25d). |
| Gynoecium | Carpels 2–5, united, but the styles often free (Pl. 25d) or partly free. Ovary superior. Placentation often free central (Pl. 25i). Stamens and gynoecium sometimes raised on a stalk (androgynophore) above the level of attachment of the petals. |
| Fruit | Usually a capsule, often opening by as many, or twice as many, apical teeth or valves as styles. Rarely indehiscent. |
The above description of floral structure accords with that most often adopted in floras. Some texts regard true petals as absent and interpret the apparent petals as modified stamens from an outer whorl.
Plants are nearly always herbaceous, sometimes shrubby, with opposite leaves that are simple, entire and often (sub-)sessile. Stipules, when present, are often scarious, and sometimes joined across the node.
An annotated list of genera is provided by Hernández-Ledesma et al. (2015).
ILLUSTRATIONS Plate 25.
SPOTTING CHARACTERS
Plants herbaceous, the stems often with swollen nodes. Leaves often opposite. Inflorescence a monochasium or dichasium (Figs. 11a, b). Flowers actinomorphic, the stamens often twice as many as petals. Fruit often capsular.
Saltbushes and Samphires
Members of this medium-sized family are distributed world-wide, except for the tropics, and most common in saline habitats and arid environments. They have a significant presence in inland Australia.
A few species are cultivated for food, for example varieties of Beta (Beetroot and Sugar Beet), and Spinacia (Spinach). The fleshy fruits of Enchylaena (Pl. 26f) and Rhagodia (Pl. 26d) are edible and were used by indigenous Australians, and the seeds of Atriplex and Tecticornia were made into flour. As a nutritious seed crop, Chenopodium quinoa (Quinoa) from South America is becoming increasingly popular.
Some Atriplex species (Saltbushes), particularly A. nummularia (Old Man Saltbush), and Maireana spp. (Bluebushes, Pl. 27a, b) provide fodder in pastoral areas of the Australian arid zone. A number of members of the family are common weeds of waste ground, roadsides and cultivation, for example Chenopodium album (Fat Hen, Pl. 26a, b).
Classification within the family has received attention in recent years. Some new genera have been proposed, accepted for a time, and subsequently merged. Species of Sarcocornia have been returned to the older genus Salicornia (Fig. 102). Others remain under evaluation. At the family level, a proposal to merge the family with the Amaranthaceae has not yet received universal acceptance.
FLORAL STRUCTURE
| Flowers | Generally small, usually actinomorphic, bisexual or unisexual. A short floral tube may be present but this is usually ignored. Solitary (Pl. 26f) or few together or borne in dense clusters (Pl. 26a), or as in the samphires, immersed in the stem tissue (Fig. 102; Pl. 26g, h). If flowers are unisexual, the species may be monoecious or dioecious. |
Fig. 102 Salicornia quinqueflora (Beaded Glasswort)P(4) A2 G(2)
Decumbent or shortly erect or tufted perennial with apparently leafless stems made up of fleshy cylindrical segments; inflorescences spike-like with the flowers in small groups immersed in the stem tissue towards the tips of the branchlets; perianth segments 3 or 4, succulent, with the upper lobe overlapping the lateral ones. Locally common in saltmarsh communities of all states except NT, mainly coastal, but occasionally fringing saline lakes inland. Flowering mainly in autumn. Cf. Pl. 26g, h. (a ×0.6, b ×3, c–f ×24)
| Perianth | Of one whorl, usually referred to as tepals, sometimes regarded as sepals. Tepals usually 5 (sometimes less), free, or often united at the base or more completely, often greenish and herbaceous to somewhat fleshy, usually persistent and enlarging in fruit. In female flowers of Atriplex (Saltbush), the perianth has usually been regarded as absent, the ovary enclosed by a pair of bracteoles (Pl. 27g). Detailed research observing the development of the bracteoles from inception to maturity has prompted suggestions they could be interpreted as tepals. |
| Androecium | Stamens 1–5, opposite the perianth parts (Pl. 26b, e), usually free, sometimes united at the base, sometimes alternating with extra outgrowths or lobes (which may be referred to as staminodes). |
| Gynoecium | Carpels usually 2, sometimes 3–5, united. Ovary nearly always superior, with 1 loculus and one basally attached ovule. The style is usually bifid (Fig. 102e, f). |
| Fruit | Generally a berry, achene or nut. The perianth is generally persistent and in many genera enlarges in fruit, often developing appendages such as wings (Maireana, Pl. 27b), spines (Sclerolaena), or tubercles (Malococera). The fruiting perianth is succulent in Enchylaena (Pl. 26f). In Atriplex, the bracteoles enlarge (Pl. 27g) and often develop appendages, or become bladdery. |
The family is mostly made up of annual and perennial herbs, and shrubs. Leaves are generally small, often succulent, and crowded on the branches. There are no stipules. Young plants are often covered with tiny bladder-like hairs (Pl. 26b, e), which later deflate to give the plant a greyish or mealy appearance. Fruit characters are used extensively in the identification process.
Flora of Australia volume 4 covers this family. A more up-to-date, annotated list of genera for this and other familes in the order Caryophyllales is provided by Hernández-Ledesma et al. (2015). Shepherd & Wilson (2007) merged several Australian genera.
ILLUSTRATIONS Figure 102; Plates 26, 27a–g.
SPOTTING CHARACTERS
Small shrubs or herbs mostly with fleshy leaves, often mealy, without stipules. Flowers small, often closely clustered. Perianth single. Style usually two-branched (and/or stigmas usually two).
Amaranths
The Amaranths are mostly herbs and shrubs, and members of the family are widely distributed in warmer regions. Some weedy species venture into cooler temperate zones.
Several species are well known in gardens including Celosia cristata (Cockscomb), Amaranthus caudatus (Love-lies-bleeding) and A. tricolor (Joseph’s Coat), and Gomphrena globosa (Globe Amaranth). The genus Ptilotus with about 90 species, all but one Australian, provides a number of attractive subjects for native gardens. Various Amaranthus spp. are harvested for edible leaves and seeds. The family also includes a number of weedy species, in many cases opportunistic inhabitants of disturbed sites, but sometimes seriously invasive such as Alternanthera philoxeroides (Alligator Weed) and Gomphrena celosioides (Gomphrena Weed).
Amaranthaceae and Chenopodiaceae have long been recognised as closely related groups. Suggestions to merge the two into an enlarged Amaranthaceae have not yet received universal acceptance.
FLORAL STRUCTURE
| Flowers | Individually often small, actinomorphic. Solitary or in clusters, these often dense and spike-like or head-like (Pl. 27h, l). Usually bisexual. If flowers are unisexual, the species may be monoecious or dioecious. Often subtended by rather dry, papery bracts and bracteoles. |
| Perianth | Of one whorl, the segments referred to as tepals or sepals. |
| Calyx | Sepals usually 4–5 (sometimes fewer), usually free, the segments often rather dry and papery, often whitish to pink or reddish (Pl. 27 i, j, m) |
| Corolla | Petals absent. |
| Androecium | Stamens 1–5 (Pl. 27i, m) or sometimes more, filaments often united, the tube sometimes bearing petal-like or fringed lobes or outgrowths between each anther. |
| Gynoecium | Carpels often 2, sometimes 3, united (Pl. 27j). Ovary superior, with 1 loculus and usually a solitary ovule. The style is usually bifid. |
| Fruit | Usually dry and dehiscent (sometimes by an apical lid, Pl. 27j), the fruit wall often thin and breaking irregularly. Sometimes indehiscent. Sometimes referred to as a utricle. |
Plants are generally annual or perennial herbs or sometimes shrubs, with simple, usually entire leaves that bear no stipules. In cultivated plants, the leaves are often variously coloured.
An annotated list of genera is provided by Hernández-Ledesma et al. (2015).
ILLUSTRATIONS Plate 27h–m.
SPOTTING CHARACTERS
Plants herbaceous or shrubby. Leaves exstipulate. Flowers small, often with a dry papery perianth (of a single whorl) and associated bracts aggregated in dense inflorescences.
37 ERICACEAE
Heaths and Heathers
A close relationship has long been recognised between the traditional Ericaceae, a large virtually cosmopolitan family, and the smaller Epacridaceae (Heath Family) mostly from Australia. Now, in recent classifications, the Ericaceae has been expanded to encompass not only Epacridaceae but several other smaller families such as Vacciniaceae (Blueberries) and Pyrolaceae (Wintergreens). About eight subfamilies have been established within this enlarged family; subfamily Epacridoideae accommodates those genera formerly included within the family Epacridaceae.
Among well-known ornamentals are Arbutus (Strawberry Tree), Erica (Pl. 28a–e) and Rhododendron (which includes azaleas). Although Epacris (Heath) is also grown as an ornamental, other members of the subfamily are not common in cultivation, apparently due to difficulties with propagation. Epacris impressa (Common Heath, Figs. 103, 104; Pl. 28h), the floral emblem of Victoria, is common in heathlands and open forests of south-eastern Australia.
The succulent fruits of some genera were eaten by early settlers and Aboriginal people, who also valued the plentiful nectar produced by many species. Blueberries and cranberries harvested from Vaccinium spp. have become more popular in recent times, valued for their nutritional content.
In its older sense, Ericaceae was poorly represented in Australia by only four or five small genera including Gaultheria (Wax-berry), found in alpine areas of the south-east. Rhododendron lochiae (Pl. 28i–k) from north Qld is the sole native member of this horticulturally important genus.
The term heath is also applied to a plant community that is dominated by shrubs varying in height from about 0.5 to 2.0 metres. These communities include many members of the Epacridoideae, and are found on shallow and/or poorly drained soils that are low in nutrients.
FLORAL STRUCTURE
| Flowers | Usually actinomorphic and bisexual, often small; larger and showy in genera such as in Rhododendron. |
| Calyx | Sepals usually 4 or 5, free or slightly united. On the pedicel there are often additional bracts which may resemble the sepals and grade into them (Figs. 104a, 111a). |
| Corolla | Petals usually 4 or 5, usually united, rarely free. The corolla is often tubular, with 5 lobes at the top that may be densely hairy as in Leucopogon (Beard-heath, Figs. 105, 106). In the genus Richea (Fig. 108a) the corolla remains closed, and as the anthers mature it splits transversely near the base, the upper part (the operculum or cap) falling leaving a cup-like rim. |
| Androecium | Stamens usually 5–10, sometimes more, often free, sometimes united by filaments or anthers (Pl. 28c, d), often epipetalous (e.g. in many Australian genera, Figs. 104b, 107b), but also often hypogynous as in Dracophyllum, Prionotes, Richea (Fig. 108b), Sprengelia (Fig. 110b) and Woollsia from Australia, and many overseas genera such as Erica (Pl. 21c) and Rhododendron. Anthers often 1-celled at maturity in Australian genera, opening by a longitudinal slit. |
| Gynoecium | Carpels usually 4–5. Ovary often superior, sometimes inferior, with 1–10 loculi (mostly 4–5, Pl. 28e, k). Ovules 1–many per loculus. Placentation apical or axile (Pl. 28e), sometimes parietal. The style is simple and may be terminal (Figs. 105b, 106b), or inserted in a depression in the top of the ovary (Fig. 104b). A nectary disc or nectary glands are usually present at the base of the ovary (e.g. Fig. 106b). The nectary is absent in Sprengelia (Swamp Heath). |
| Fruit | A capsule, berry or drupe. |
Members of the family are mostly evergreen shrubs, sometimes trees, lianes or epiphytic, rarely herbaceous. Leaves are commonly alternate and simple. There are no stipules. Australian species are usually small woody shrubs.
‘Ericoid’ leaves, small and more or less linear with the blade margins recurved such that the underside is obscured, are mostly restricted to the subfamily Ericoideae, and are particularly common in the genus Erica. Australian genera often possess small, tough, sharply pointed leaves with the main veins running more or less parallel (Pl. 28f). A few genera, such as Sprengelia (Fig. 109), Dracophyllum, and Richea have leaves with broad sheathing bases. Some of the Tasmanian richeas resemble small palm trees, and specimens of R. pandanifolia up to 18 m high are found in the dense rainforests.
ILLUSTRATIONS Figures 103–11; Plate 28.
SPOTTING CHARACTERS
Woody shrubs. Leaves often small, tough and pointed, with main veins more or less parallel and more easily seen on the lower surface (Pl. 28f). Flowers actinomorphic, 4- or 5-partite. Sepals often grading into bracts of similar size and texture. Corolla often tubular.
Fig. 103 Epacris impressa (Common Heath) (×0.7)
Fig. 104 Epacris impressa (Common Heath)
Erect to spreading shrub to about 1 m high; leaves lanceolate, to 1.5 cm long, pungent, the midrib prominent on the underside; flowers white, pink or red, axillary; corolla lobes overlapping in bud; fruit a capsule, opening by 5 valves. Widespread in heaths and drier forests with a heathy understorey, in Vic., Tas., NSW and SA. Flowering autumn to spring. See also Pl. 28h. (a–b ×7, c ×12)
Fig. 105 Leucopogon ericoides (Pink Beard-heath)
Slender, small, wiry shrub to 2 m; branches pubescent; leaves oblong, less than 1 cm long, mucronate, the margins recurved; flowers white to pink, 2–4 in short axillary spikes, each flower subtended by a bract and two bracteoles; corolla lobes valvate in bud; fruit a drupe. Widespread in heathlands, Vic., Tas., NSW, SA and Qld. Flowering in spring. (a–b ×15, c ×30)
Fig. 106 Leucopogon virgatus (Common Beard-heath)
Wiry small shrub to 1 m; leaves to 1.5 cm long, lanceolate, incurved, sharply pointed, lower surface with conspicuous parallel veins; flowers white in short, dense, axillary spikes, each flower subtended by a bract and two bracteoles; corolla lobes valvate in bud; anthers with sterile tips; number of loculi in the ovary variable, usually 4 or 5; fruit a drupe. Widespread in heathlands, Vic., Tas., NSW, SA and Qld. Flowering in spring. (a–b ×15, c ×40)
Fig. 107 Lissanthe strigosa (Peach Heath)
Small, rigid shrub up to about 0.5 m high; leaves linear, pungent, to 1.5 cm long, the margins recurved; flowers pale pink in small axillary clusters; corolla lobes valvate in bud; fruit a drupe. Widespread in all states except WA. Flowering late winter to spring. (a–b ×12, c ×25)
Fig. 108 Richea procera (Lowland Richea)K5 C(5) A5 G(5)
Sparingly branched shrub to 3 m; leaves imbricate, recurved, ovate-lanceolate and tapering to a point, 1–3.5 cm long, bases sheathing; flowers creamy-yellow, in short spikes or heads terminating main or lateral shoots; corolla splitting transversely, the upper part shed as a cap or operculum; filaments thickened and papillose in the upper part. Endemic in Tas. where locally common, occurrences scattered through central and southern parts. Flowering late spring. (a–b ×10, c–d ×20)
Fig. 109 Sprengelia incarnata (Swamp Heath) (×0.7)
Fig. 110 Sprengelia incarnata (Swamp Heath)K5 C(5) A5 G(5)
Slender shrub to 2 m, with rigid erect stems; leaves with sheathing bases, to 2 cm long, tapering to a sharp point; flowers pink in leafy terminal clusters; fruit a capsule. Occurs in wet heaths and swamps in southern Vic., SA, Tas., and NSW. Flowering in spring. (a–c ×7, d ×25)
Fig. 111 Woollsia pungens (Snow Wreath)K5 C(5) A5 G(5)
a flower with enveloping bracts and subtending leaf; b half flower—the dotted lines indicate the limits of the bracts and sepals, to distinguish them from the corolla tube; c T.S. ovary, placentation axile (a–b ×7, c ×25)
Erect shrub to 2 m high; leaves crowded, sessile, ovate, acuminate, about 1 cm long; flowers white or reddish, sessile and axillary; bracts around the base of the flower grade into the 5 sepals; corolla lobes crumpled in bud; anthers borne on very fine filaments that in this sample were free from the corolla tube (some texts report filaments (partly) fused to the corolla); fruit a capsule. Common in coastal heaths and dry forests of NSW and Qld. Occasionally grown. Flowering in winter or early spring to summer.
Borages and Forget-me-nots
Boraginaceae is a medium-sized family widely distributed in temperate and tropical regions, particularly in western North America, and east from the Mediterranean Sea. In many existing floras the family is treated in a broader sense (i.e. including more genera) than the most recent classifications accept; various groups of genera have now been moved into separate families. One such family, Heliotropiaceae, includes several hundred species of Heliotropium (Heliotrope) quite common in gardens and as wayside weeds. A characteristic feature of this latter small family is the distinctive style-end which, in H. europaeum (Pl. 29f), is conical with the apex 2-lobed and the stigmatic zone a ring around the base.
Boraginaceae is well known in gardens through genera such as Borago (Borage) and Myosotis (Forget-me-not). Borago officinalis (Borage) and Symphytum officinale (Comfrey) are familiar herbs.
Weedy species are found in a number of genera including Amsinckia (Fiddle-neck), Buglossoides (Bastard Alkanet, Sheepweed) and Echium. In southern Australia, Echium plantagineum (Vipers Bugloss, Pl. 29 a–c) is a major weed of grazing areas, although eaten by sheep in times of drought. Two alternative common names, Patersons Curse and Salvation Jane, appear to reflect differing attitudes towards the species’ presence.
FLORAL STRUCTURE
Plants are mostly coarsely hairy herbs, sometimes shrubs or trees, with alternate, simple, exstipulate leaves. Basal leaves often form a short-lived or persistent rosette.
ILLUSTRATIONS Plate 29a–c. (See also Pl. 29 d–f.)
SPOTTING CHARACTERS
Plants usually herbaceous, often covered with coarse hairs (thus rough to the touch). Inflorescences distinctively curved, straightening as they mature (Pl. 29b).
39 SOLANACEAE
Nightshades and Kangaroo Apples
The Solanaceae is a medium-sized family, widely distributed in tropical and temperate regions, particularly in Central and South America. It is also well represented in Australia and Africa.
Alkaloids are found in the foliage, flowers and immature fruits of most genera. Many are toxins, and some are important in the pharmaceutical industry. Solandine, used in contraceptive pills, is extracted from Solanum aviculare and S. laciniatum (Kangaroo Apple, Pl. 30m). Aboriginal people are known to have chewed the leaves of native members of Nicotiana (Pl. 30j), and the American species N. tabaccum is the source of commercial tobacco. The leaves of Duboisia hopwoodii (Pituri, Pl. 30h) contain nicotine and nornicotine, and were also chewed by Aboriginal people as well as being used as a source of animal poison.
Family members cultivated for food include Capsicum spp. (peppers and chillies), Solanum lycopersicum (syn. Lycopersicon esculentum, Tomato, Pl. 30k), Solanum betaceum (Tree Tomato), S. melongena (Eggplant) and S. tuberosum (Potato). Potatoes are underground storage organs produced on the ends of rhizomes arising from the plant axis. Well-known ornamentals are Cestrum spp. (Pl. 30g), Brugmansia spp. (Angel’s Trumpet, Pl. 29g–i), Lycianthes rantonnei (Blue Potato Bush, Pl. 30e, f), Nicotiana spp., Petunia hybrids, Physalis spp., and Solanum laxum (syn. S. jasminoides, Potato Vine). S. aviculare and S. laciniatum (Kangaroo Apple, Pl. 30m) are commonly grown for revegetation work.
A number of introduced species have become troublesome weeds including Datura spp. (Thornapple), some Nicotiana and Physalis spp. (Pl. 30i), Lycium ferocissimum (Boxthorn), Solanum nigrum (Black Nightshade, Pl. 30n), and Salpichroa origanifolia (Pampas Lily of the Valley, Pl. 30a–d).
FLORAL STRUCTURE
| Flowers | Actinomorphic or slightly zygomorphic, mostly bisexual. Inflorescence various, often terminal, or flowers solitary. |
| Calyx | Sepals usually 5, united. Calyx tubular to campanulate, usually five-lobed, persistent and sometimes developing to enclose the fruit (Physalis, Pl. 30i). |
| Corolla | Petals usually 5, united. The corolla may be tubular, campanulate, urceolate or funnel-shaped (Fig. 2g, j, k). The limb is often rotate or stellate (Fig. 2h). |
| Androecium | Stamens usually 5, sometimes 4 or 6, free or united, epipetalous and alternating with the corolla lobes. Anthers sometimes coherent, dehiscing by vertical slits or apical pores, sometimes not all fertile. |
| Gynoecium | Carpels usually 2, sometimes more, united. Ovary superior, with a nectary disc usually present (Pl. 30c). Loculi usually 2 but sometimes more due to development of false septa, these sometimes incomplete. Placentation axile, the ovules usually numerous (Pls. 29i, 30k). |
| Fruit | A berry, sometimes rather dry, or a capsule. |
Members of the family are mostly herbs and shrubs, as well as some small trees and climbers. Stems are glabrous or hairy, or with prickles. Leaves are alternate, occasionally almost opposite, simple, entire, deeply lobed or pinnate, exstipulate, and sometimes prickly.
For descriptions and keys to Australian species see Flora of Australia volume 29. Hunziker (2001) provides an overview of genera.
ILLUSTRATIONS Plates 29g–i and 30.
SPOTTING CHARACTERS
Flowers usually actinomorphic, with 5 united sepals, and 5 united petals. Corolla often rotate (Fig. 2h). Stamens 5, epipetalous. Fruit a berry or capsule, with many seeds.
Figworts, Myoporums and Emubushes
The Scrophulariaceae form a medium-sized family of herbs, shrubs and small trees widely distributed in temperate and tropical parts of the world.
The limits of the family have changed significantly in recent classifications with numerous genera being assigned to other families, and other genera previously placed elsewhere being included. Many familiar herbaceous garden plants such as Antirrhinum (Snapdragon), Digitalis (Foxglove), Linaria (Toadflax) and Penstemon have now moved to the family Plantaginaceae. The commonly grown shrubby Hebes, now included in the genus Veronica, have also moved to Plantaginaceae.
Garden subjects retained or now placed in Scrophulariaceae include Nemesia, Buddleja, and Verbascum (Mullein). The latter genus also includes a number of weedy species. Scrophularia auriculata (Water Figwort, Pl. 31a–e) is sparingly naturalised in South Australia and Victoria.
Also now in Scrophulariaceae is the Australian endemic genus Eremophila (Emubush), as well as Myoporum (Boobialla) from Southeast Asia, Australasia and Hawaii, and the monotypic Bontia from the West Indies and tropical America. These three genera have in the past constituted the small family Myoporaeae.
There are more than 200 species of Eremophila, most of which grow in the Australian arid zone. Usually shrubs or small trees with colourful flowers, many are in cultivation and make attractive garden plants.
There are 30 species of Myoporum with about half of these found mostly in southern and temperate parts of Australia. M. insulare (Common Boobialla) has been widely planted in windbreaks in South Australia and Victoria, and M. floribundum and M. parvifolium (Creeping Boobialla) make attractive ornamentals. M. platycarpum (False Sandalwood, Sugarwood) is locally common in low woodlands in the arid zone.
FLORAL STRUCTURE
Members of the family are herbs or shrubs to small trees, a few are vines.
For further reading on Eremophila see Brown & Buirchell (2011). Chinnock (2007) is a major text covering the former family Myoporaceae.
ILLUSTRATIONS Plate 31.
SPOTTING CHARACTERS
Herbs, shrubs or small trees with alternate, exstipulate leaves. In Eremophila and Myoporum plants are often resinous, the leaves with scattered small secretory cavities appearing as pale dots. Flowers often zygomorphic, the corolla often 2-lipped. Stamens epipetalous, the anther lobes joined, opening by one slit.
Mints and Mintbushes
This large family, found throughout the world, is particularly well represented in the Mediterranean region. About 40 genera are native in Australia. The older name, Labiatae, still acceptable under nomenclatural rules, is derived from the character of the corolla, in which one or more petals form a distinct lip. The alternative name is based on the Mediterranean genus Lamium.
The family includes many aromatic culinary herbs such as Mentha (Mint), Ocimum (Basil), Origanum (Marjoram, Oregano), Rosmarinus (Rosemary), Salvia (Sage) and Thymus (Thyme), all of which were early introductions to Australia. Lavandula dentata (Lavender) is cultivated in Tasmania and Victoria for the commercial extraction of the oil, which is used in perfumes. The many introduced genera grown as ornamentals include Coleus (Painted Nettle), Lavandula, Leonotis (Lion’s Ear), Nepeta (Cat Mint), Plectranthus, Salvia and Stachys (Lamb’s Ears). Two well-known native genera widely cultivated are Prostanthera (Mintbush, Fig. 112) and Westringia. Several European species are weedy in Australia, including Marrubium vulgare (Horehound), Lavandula stoechas (Topped or Spanish Lavender) and Lamium amplexicaule (Henbit Dead-nettle).
The limits of the family have been expanded in recent classifications to include many genera formerly placed in Verbenaceae (Lantana and Verbena family). Inflorescences in Verbenaceae as now defined are indeterminate racemes, spikes or heads.
FLORAL STRUCTURE
| Flowers | Usually zygomorphic, bisexual. Usually arranged in cymose clusters on an indeterminate main axis, sometimes appearing whorled around the stem, rarely solitary. |
| Calyx | Sepals usually 4–5, rarely more, united. Calyx often 2-lipped, persistent around the fruit. |
| Corolla | Petals usually 4–5, united. Corolla tubular but usually deeply lobed, often 2-lipped. |
| Androecium | Stamens usually 4 or 2, epipetalous (Fig. 112b, c), sometimes arranged in 2 pairs of unequal length. |
| Gynoecium Fruit | Carpels 2. Ovary superior, the apex merging into the style, or often deeply divided into 4 lobes (each with 1 loculus) and the style then gynobasic (Fig. 113c), stigma usually bifid (forked). Ovules 1 per loculus. Disc usually present. Often dry, splitting into 4 achene-like nutlets, sometimes a drupe. |
The majority of mints are aromatic shrubs or herbs, although some are forest trees. The stems are often quadrangular with simple, opposite, or occasionally whorled leaves. Plants are often hairy, the vestiture including glandular hairs. Stipules are absent.
ILLUSTRATIONS Figures 112, 113.
SPOTTING CHARACTERS
Stems square (in T.S.), often hairy, and leaves opposite and aromatic. Flowers zygomorphic, stamens 2 or 4.
Fig. 112 Prostanthera rotundifolia (Round-leaf Mintbush)
Bushy shrub to 4 m; leaves opposite, aromatic, more or less orbicular, about 1 cm long; flowers mauve-purple, profuse, axillary or in short bracteate racemes; calyx with bracteoles, and dotted with oil glands; anthers with appendages. Occurs on dry hills and along watercourses in Vic., Tas., and NSW. Many members of the genus are cultivated. Flowering in late spring. (a–b ×7, c ×2.5, d ×10, e–f ×20)
Fig. 113 Salvia verbenaca (Wild Sage)
Hairy herbaceous perennial to 60 cm high; leaves opposite, ovate-oblong, wrinkled on the surface, the margins crenate, often lobed; flowers purplish, rarely white, in apparent whorls of about 6, in the axils of orbicular bracts; calyx and corolla both 2-lipped; anthers each with a very broad connective that separates the fertile lobe from a sterile one. The sterile lobe and connective act as a lever to move the fertile lobe. A foraging insect pushing against the lever may deposit pollen on its back. Introduced from Europe, a weed widely distributed in Australia. Flowering in summer. (a–b ×6, c–d ×24)
Triggerplants and Styleworts
While predominantly Australian, this small family has a few representatives in South-east Asia, New Zealand and the tip of South America.
The largest genus, Stylidium (Fig. 114, Pl. 32a) is widespread in Australia. Most members are annual or perennial herbs, exstipulate, the larger species with a cluster of basal leaves from which the stalk or scape arises bearing the inflorescence. Flowers are bisexual and zygomorphic. There are 5 sepals and the calyx is often 2-lipped. The corolla, of 5 united, white to deep pink petals, is deeply lobed with a short tube. One lobe is comparatively short, a different shape, often reflexed, and is called the labellum. At the throat of the corolla there may be 6 or 8 prominent erect appendages (Pl. 32a). The filaments of the 2 stamens are united with the style to form a column. The anthers are attached at the top of the column, with the stigma between them. The ovary is inferior, 1- to 2-celled, with many ovules attached to a central placenta. The fruit is a capsule. The calyx, the outer wall of the ovary, and the scape are often glandular-hairy.
The column of Stylidium is sensitive to touch, and at first is reflexed over the labellum. When triggered by an insect collecting nectar, the column springs up and the pollen is showered over the back of the insect. When the pollen has dispersed, the column again reflexes and the stigma becomes receptive. If the next insect to trigger the column has pollen on its back, this may then be transferred to the receptive stigma.
The small, informative, well-illustrated text by Erickson (1958) was comprehensive in its time.
Fig. 114 Stylidium armeria
Variable tufted perennial with stems to c. 1 m high (but often much less); flowers pale to dark pink, in simple, upright racemes. Widespread in south-eastern and eastern Australia. Flowering in late winter to summer. See also Pl. 32a. (×0.4)
Fanflowers and Goodenias
This small family is almost entirely confined to the southern hemisphere, with all of the 10 genera found in Australia. A few species extend to South-east Asia. In recent classifications the limits of the family have been expanded to include Brunoniaceae which formerly accommodated the single species Brunonia australis (Blue Pincushion). The type genus Goodenia is the largest and is widespread in south-eastern Australia. The Western Australian genus Lechenaultia is widely cultivated, and the striking blue flowers of Dampiera are also favoured.
Indigenous Australians utilised a few species, such as Scaevola spinescens (Prickly Fanflower), the drupes of which were much prized by tribes of the Flinders Ranges in South Australia.
FLORAL STRUCTURE
| Flowers | Usually zygomorphic and bisexual, in inflorescences of various types. |
| Calyx | Sepals usually 5, free or sometimes united or reduced to a rim. |
| Corolla | Petals 5, united but, except in Brunonia, the corolla split open on one side (Pl. 5e), often 2-lipped, sometimes spurred as in Velleia paradoxa (Spur Velleia). The corolla lobes are sometimes spread like a fan as in Scaevola (Fanflower, Pl. 5d, e), and may be described as winged when a central band along the midrib differs in colour or texture from the thinner marginal wings that are folded under in bud (Fig. 115c). |
| Androecium | Stamens 5, free or with anthers joined in a tube around the style. Stamens sometimes epipetalous at the base of the corolla. |
| Gynoecium | Carpels united, often cited as 2, but this not always readily confirmed in T.S. ovary as septum development is often incomplete and there may be 1 loculus or 2 (sometimes incomplete) loculi. Ovules 1–many. Ovary usually inferior to semi-inferior, rarely superior. At the top of the style is a 2-lipped or cup-shaped structure, usually with hairy margins, called an indusium (Fig. 115a, b; Pl. 5e, i). Before the flower opens the pollen is shed into the indusium which is carried upwards by the lengthening style. Later, the pollen is pushed out by the growth of the stigma, which is usually bilobed. |
| Fruit | Mostly a capsule, sometimes a drupe or nut. |
Fig. 115 Goodenia ovata (Hop Goodenia)K5 C(5) A5 G(loculi 2)
Spreading shrub to 2 m high; leaves ovate, thin, 2–5 cm long, with finely toothed margins; flowers yellow, 1–3 together in the leaf axils; corolla lobes conspicuously winged; indusium present on the end of the style; within the ovary, the septum often incompletely divides the loculi; fruit a cylindrical capsule about 1 cm long. Widespread near the coast and in mountain forests, Vic., SA, Tas., NSW and Qld. Flowering mainly spring and summer. (a–c ×6, d ×8)
Members of the family are mostly annual or perennial herbs or low-growing shrubs. The leaves are simple and usually alternate or radical. There are no stipules, but an axillary tuft of hairs is often present.
For descriptions, and keys to Australian species see Flora of Australia volume 35.
ILLUSTRATIONS Figure 115; Plate 5d–i.
SPOTTING CHARACTERS
Flowers usually zygomorphic. Corolla lobes often winged, usually either all directed towards one side of the flower as in Scaevola (Pl. 5d), or forming 2 lips as in Goodenia (Fig. 115c). The indusium at the end of the style (Fig. 115a, b; Pl. 5e, i) is very characteristic of the family.
Daisies
The Asteraceae, also known as Compositae, is a very large family of worldwide distribution. They are common in grasslands and lightly forested areas as well as montane vegetation, but less well represented in humid lowland tropical forests. About 290 genera and more than 1400 species are found in Australia, and include all the daisies, thistles and everlastings. The family name is derived from the genus Aster, which is known for some familiar garden plants such as the Michaelmas and Easter Daisies. The family is usually readily recognised because the apparent ‘flowers’ are condensed inflorescences of many small flowers clustered together. It is this composite nature of the ‘flower’ that gave rise to the older name Compositae, which is still widely used.
The family is a highly successful one due to its adaptability and high reproductive rate. Many species introduced into Australia have become common weeds, some seriously invasive. Local examples include the thistles, as well as Arctotheca calendula (Capeweed), Chrysanthemoides monolifera (Boneseed, Pl. 33f–h) and Jacobaea vulgaris (formerly Senecio jacobaea, Ragwort). Members of the family are not planted in pastures although fields of blue-flowered Cichorium intybus (Chicory) are sometimes grown for forage. Some are important as food plants such as Lactuca sativa (Lettuce), Cichorium endivia (Endive), and Cynara scolymus (Artichoke, Pl. 33m). Safflower oil is extracted from the seeds of Carthamus, sunflower oil from the seeds of Helianthus, and the culinary herb tarragon is Artemisia dracunculus. Members of many genera are grown as ornamentals, including Aster, Calendula (English Marigold), Chrysanthemum, Dahlia, Helichrysum and Zinnia. Numerous other species are important commercially, in products such as insecticides, varnishes and paints, soaps and detergents, cosmetics and perfumes, and medicines.
FLORAL STRUCTURE
| Flowers | Regular or zygomorphic, usually bisexual or female, arranged in heads (capitula), and usually referred to as florets (Figs. 117a, b, 118c, d; Pl. 32d, e, etc.). |
| Calyx | If present, usually represented by scales (Fig. 117b, c; Pl. 33d, e), hairs (Pl. 32e), bristles (Fig. 123d; Pl. 32k) or awns, and known as a pappus. |
| Corolla | Petals often 5, sometimes 3 or 4, united. |
| Androecium | Stamens usually 5 and epipetalous, united by their anthers, which form a tube around the style (Fig. 117d; Pl. 32e, i). |
| Gynoecium | Carpels nearly always 2 (indicated by two style-arms), united. Ovary inferior, unilocular with 1 basal ovule. |
| Fruit | Usually dry and indehiscent and then historically and still commonly called an achene (some authors prefer the term cypsela—an achene-like fruit derived from an inferior ovary Pls. 32k, 33e). Occasionally a drupe, as in Chrysanthemoides monilifera (Boneseed, Pl. 33h). |
Within the family there is considerable variation in floral structure. The following notes amplify the brief summary given above. Most of the features discussed are illustrated in Figures 116–27 and Plates 32 and 33.
The inflorescence is called a capitulum or head (Pl. 32c, h), and the individual flowers are usually known as florets. The florets, which are sessile (stalkless), are borne on the receptacle, the expanded end of the inflorescence stalk, the peduncle. The receptacle is usually more or less flat, but in some genera is elongated to form a club-shaped or cone-shaped structure. Its surface may be smooth or pitted, and sometimes there are hairs or small scales at the base of each floret (Pls. 32j, 33b). If hairs or scales are absent, the receptacle is said to be naked.
Each head is surrounded by a series of bracts, which are usually referred to collectively as the involucre, and the component parts as involucral bracts or phyllaries (Pls. 32h, 33m).
Two types of floret, tubular and ligulate, are most commonly observed. Tubular florets, also called disc florets, (Figs. 117b, 118d; Pls. 32e, 33d) have a tubular, actinomorphic corolla of 5 (sometimes 4) united petals with 5 (4) lobes at the top. Filiform florets are a form of tubular floret in which the corolla tube is very slender, and these are usually unisexual.
Ligulate florets, often also called ray florets (Figs. 117a, 118c; Pls. 32d, i, 33c), have a short corolla tube at the base, which extends on one side into a flat strap-shaped part called the ligule or ray. There are either 3 or 5 lobes or teeth at the end of the ligule, said to represent the number of petals.
At anthesis the pollen is shed to the inside of the anther tube and pushed out of the corolla by the upward growth of the style, to be gathered by an insect, or otherwise dispersed. Finally, the style-arms separate and the stigmatic surfaces become receptive.
Depending on the arrangement of the floret types, three kinds of head are common: radiate, discoid, and ligulate.
In the radiate head (Figs. 116, 118a; Pls. 32b, c, 33a, g, i) both tubular and ligulate florets are present. The tubular florets are usually bisexual, occupy the centre of the receptacle, and may be referred to collectively as the disc. The ligulate florets are arranged in one or more rows around the edge of the receptacle and collectively may be called the ray. They commonly have a 3-toothed ligule and are often either female or neuter.
The discoid head (Fig. 121a) has only tubular florets, which are usually all bisexual. Sometimes the outer florets (next to the involucral bracts) are female (Fig. 123a) or neuter and in addition their corollas may be filiform (Fig. 123a) or absent. These latter cases are sometimes distinguished as disciform heads (Fig. 122b; Pl. 33j).
The ligulate head (Fig. 125a; Pl. 32g, h) contains ligulate florets only, which are nearly always all bisexual, but occasionally may be all male or all female. The florets usually have a 5-toothed ligule (Pl. 32i).
With reference to florets, some authors restrict the term ligulate to refer only to florets found in a ligulate head, and restrict ligule to the strap-shaped part of their corollas. The term ray then refers only to the one-lipped florets around the margin of a radiate head, and the term lamina may be used for the extended part of their corollas.
Florets in the head vary in number with species (from very few to several hundred) and they may be of similar or different sex. The head is said to be homogamous if all florets are of the same sex, and heterogamous if a mixture of sexes is present. The most common combination is bisexual and female, with the bisexual florets in the centre and one or two rows of female florets around the outside of the head (Fig. 116b; Pls. 32c, 33b). As the head is a racemose inflorescence, the youngest florets are in the centre and are the last to mature (Pl. 33g, i). Rarely all florets are unisexual, and species are then monoecious or dioecious.
Heads may be solitary or arranged in similar ways to other flowers, that is, in racemes or panicles etc. Some species have many small heads closely packed together at the end of a single stalk resulting in a compound head (Fig. 126; Pl. 33l). Each small head within the compound head will have a partial involucre and is called a partial head (Fig. 127a). Bracts at the base of a compound head are referred to as the general involucre.
As the family is so large, fine detail is often required for identification of species. The following paragraphs emphasise some further features of flower structure used in classification.
The shape of the base of the anther lobes and the presence or absence of apical and/or basal appendages is of importance. If a short basal appendage is present, the anther lobes are described as acute; if the appendage is longer, it is called a tail and the lobes are said to be tailed or caudate (Fig. 123c). Because the anthers start to wither after the pollen is shed, examination of the lobes for appendages is best carried out on buds.
Style-arms, sometimes called style-branches, vary in form (Pls. 32 d, e, i, 33c). They may be flattened or rounded in cross-section, and their apices may be acute, obtuse or truncate. The stigmatic areas are often papillose and may cover the inner surface of each arm or be more restricted.
The pappus may be of hairs (Pl. 32d, e), bristles, scales or awns. The bristles are said to be simple if they are smooth, barbellate if they have short rough appendages, and plumose if they bear long fine hairs (Pl. 32k). Scales are usually flat bract-like structures, although the distinction between a narrow scale and broad bristle is sometimes tenuous. They may be glabrous or pubescent, and are said to be fringed if the hairs are only marginal (Pl. 33e).
Involucral bracts are usually numerous, in one or more rows, and may be free from one another or united (Fig. 116). They vary considerably in structure and texture and may be herbaceous (leaf-like and green, Pl. 32h), leathery, scaly, stiff or spiny. The margins of herbaceous bracts are often dry, colourless or brownish and membranous, and are then said to be scarious.
The fruit is usually dry, 1-seeded and indehiscent and is correctly referred to as a cypsela. However, it is commonly called an achene (the term ‘achene’ is usefully restricted to refer to a dry, 1-seeded fruit derived from a superior ovary of a single carpel, but some authors omit ovary position and carpel number from its definition). If a pappus is present in the flower, it usually persists at the fruiting stage and is often an aid to fruit dispersal (Figs. 117c, 121h; Pls. 32k, 33e). A dry, hairy, persistent pappus enables the fruit to be blown away by the wind, and many thistles are dispersed in this way. Sometimes the tissue between the pappus and the ovary grows as the fruit develops, and the pappus is then connected to the top of the fruit by a small stalk called the beak (Fig. 124c; Pl. 32k). The whole structure is often then referred to as a beaked achene.
Young vegetative parts of some species, when damaged, exude a white sticky juice known as latex. This is typical of the tribe Cichorieae, to which dandelion, lettuce, and some of the thistles belong.
Most members of the Asteraceae are annual, biennial or perennial herbs. Some are exceedingly small while others, like the sunflower are large with spectacular inflorescences. A few genera include woody shrubs or small trees. Leaves lack stipules and are usually simple, but shape and margins vary enormously; blades are often deeply dissected.
Flora of Australia volume 37 deals with about half of the family. Volume 38 is yet to be published. See also Salkin et al. (1995) and ADSG (2002).
SELECTED EXAMPLES
The illustrations (Figs. 116–27; Pls. 32b–k, 33) show examples of species with radiate, discoid, ligulifloral and compound heads. Floral formulae have been omitted from individual captions. They all take the form K pappus 
except where corollas are 4-partite or florets unisexual.
Fig. 116 Tagetes ‘Cinnabar’ (Marigold)
Herbaceous annual to about 30 cm high; leaves mostly opposite, pinnatisect, up to 10 cm or more long, strongly aromatic; flowerheads radiate, surrounded by one row of united involucral bracts; ligulate florets female, deep red with a yellow border; tubular florets bisexual, yellow; pappus of white scales; ovary darkening with development, becoming almost black in fruit. One of many cultivated marigolds, most of which are double and do not show the basic structure. Flowering summer to autumn. The genus is native to Central and South America and adjacent North America. (a ×1.5, b ×3)
Fig, 117 Tagetes ‘Cinnabar’ (Marigold)
a ligulate floret; b tubular floret; c fruit; d young tubular floret—upper part of corolla opened out to show the anther tube around the style and emerging stylearms; e T.S mature ovary—the ovule completely fills the loculus, placentation basal. (a–c ×3, d ×12, e ×6)
Tagetes minuta (Stinking Roger), an aromatic annual to 1 m or more tall, is widely naturalised in mainland Australia, (but not commonly collected in Vic.).
Fig. 118 Olearia argophylla (Musk Daisybush)
Small tree to 10 m high; leaves broad-lanceolate to ovate, to 15 cm long, with a musky odour, margins slightly toothed, the upper surface green, lower surface covered with silvery hairs; flowerheads radiate, creamy white, numerous in large panicles; pappus of numerous capillary bristles. Common in cool gullies and sheltered forests of Vic., Tas. and NSW. Flowering in spring and summer. (a–d ×6, e–f ×12)
Fig. 119 Xerochrysum bracteatum ‘Dargan Hill Monarch’
Spreading perennial to 50 cm; leaves lanceolate, silvery-grey, hairy; flowerheads golden yellow, large, up to 7 cm diameter; florets all tubular, surrounded by spreading, papery involucral bracts which remain showy when the central florets have matured and fallen, hence the name ‘everlasting’. Previously placed in the genus Helichrysum, and later transferred to Bracteantha, but Xerochrysum found to have priority. The species is variable and widespread, this form native to Qld (Cunningham Gap). Flowering spring and summer. A selected cultivar, commonly grown, the cultivar name written in single quotation marks. (a ×0.6, b ×2)
Fig. 120 Senecio vulgaris (Common Groundsel)
Erect annual herb to 30 cm high; stems glabrous or with scattered hairs; leaves variably pinnatifid with widely-spaced and sinuate-toothed lobes, the upper leaves tending to be stem-clasping (amplexicaul); involucral bracts usually described as being in one row in the genus, but in this case with a number of additional, smaller bracts at the base of the involucre, often conspicuously tipped black; florets all tubular, yellow; anther bases obtuse; pappus of numerous capillary bristles. Widespread naturalised weed, common in gardens, introduced from Europe. Flowering mainly winter to spring. (×0.5)
Fig. 121 Senecio vulgaris (Common Groundsel)
(a–c, f–h ×6, d, e ×12)
Fig. 122 Leptorhynchos squamatus (Scaly Buttons)
Fig. 123 Leptorhynchos squamatus (Scaly Buttons)
Spreading herbaceous perennial; stems numerous, to 25 cm tall including the long scaly peduncles; leaves linear-lanceolate, 1–4 cm long, the upper surface with scattered hairs, the lower with dense, white woolly hairs; involucral bracts numerous, fringed, grading down into small scaly bracts on the peduncle; florets all tubular, yellow, the inner florets bisexual, a few outer florets often filiform and female; anthers tailed; pappus of barbellate bristles becoming plumose towards the tips. Widespread in all eastern states. Flowering spring and summer. (a–c ×12, d–e ×20)
Fig. 124 Taraxacum officinale (Dandelion)(a ×0.6, b–c ×6, d–e ×12)
Fig. 125 Taraxacum officinale (Dandelion)
Herbaceous perennial with a well-developed tap root; leaves in a basal rosette, pinnatifid and often described as runcinate, 3–15 cm long, with pointed lobes; flowerheads with yellow ligulate florets only, borne on single unbranched scapes that exude latex when broken; involucral bracts numerous, herbaceous; pappus of numerous capillary bristles; cypselas (sometimes called achenes) beaked. Widespread naturalised weed native to Europe and Asia, now virtually cosmopolitan. Some headway has been made in distinguishing the various dandelions naturalised in Australia, all of which have previously been included in T. officinale in what is known as a species aggregate (see Flora of Australia volume 37). Flowering mostly spring and summer. (a–c ×3)
Fig. 126 Calocephalus citreus (Lemon Beauty-heads)
(a ×0.7, b ×5)
Fig. 127 Calocephalus citreus (Lemon Beauty-heads)
Perennial herb to 40 cm high; stems branching, silvery-grey and densely hairy; leaves mostly opposite, linear, to 8 cm long, covered with fine white hairs; compound flowerheads yellow; partial heads small, with few tubular florets, immersed in woolly hairs; pappus of fine scales with plumose tips, united at the base; cypselas (sometimes called achenes) papillose. Widespread in lowland areas of all states except WA. Flowering late spring and summer. (a–b ×25)
Pittosporums
Most of the nine genera belonging to this small family are restricted to Australia. Pittosporum, from which the family name is derived, is the largest genus with about 140 species, and it extends into New Zealand, South-east Asia and across to Africa.
The berries of Billardiera scandens (Apple-berry, Fig. 128) are edible and were used by indigenous Australians. Pittosporum angustifolium (Weeping Pittosporum, Native Willow, Fig. 130), P. undulatum (Sweet Pittosporum, Fig. 131) and Bursaria spinosa (Sweet Bursaria) have been used as hedge plants and as ornamentals. A number of other Pittosporum spp., with numerous cultivars, are grown in gardens. Hymenosporum flavum (Native Frangipani) and the climbers Sollya heterophylla (Bluebell Creeper), Marianthus bignoniaceus (Orange Bell-climber, Fig. 129) and other Billardiera species are common in native gardens. Pittosporum undulatum, which is native to the wetter forests of eastern Victoria, has extended its range, becoming a weed in some southern coastal and forest regions, as well as in a number of other countries. Similarly, Sollya heterophylla has become established outside its natural range.
FLORAL STRUCTURE
| Flowers | Actinomorphic, usually bisexual. Inflorescences often paniculate or corymbose, or flowers solitary. |
| Calyx | Sepals 5, free or weakly to strongly united, falling early in some Pittosporum species. |
| Corolla | Petals 5, free or sometimes wholly or partially coherent, or united. |
| Androecium | Stamens 5, usually free. |
| Gynoecium | Carpels 2–5, united, ovary superior, with 1–5 loculi, and 2–5 parietal or axile placentas. A unilocular ovary is sometimes incompletely divided by the inward growth of parietal placentas (Fig. 131c). |
| Fruit | A capsule or berry. |
The family consists of trees, shrubs and climbers, often with a distinctive resinous odour. The leaves are simple, usually alternate, sometimes whorled. There are no stipules.
ILLUSTRATIONS Figures 128–31.
Leaves often aromatic. Flowers actinomorphic, 5-partite, the corolla often appearing tubular, often with a spreading limb. Ovary superior, often of 2 united carpels. Seeds often immersed in sticky, coloured pulp.
Fig. 128 Billardiera scandens var. scandens (Common Apple-berry)K5 C5 A5 G(2)
Twining creeper; leaves lanceolate to linear, usually with undulate margins, 1.5–5 cm long; flowers pendulous, pale yellow, solitary and axillary; corolla appearing tubular but petals free; fruit an elongated greenish berry. Widespread in forests of Vic., SA., Tas., NSW and Qld. Flowering spring to early summer. (a–b ×4, c ×12)
Fig. 129 Marianthus bignoniaceus (Orange Bell-climber)K5 C(5) A5 G(2)
Twining creeper, the stems becoming glabrous with age; leaves hairy, scattered, ovate or cordate, 2–4 cm long; flowers pendulous on slender stalks, 1–3 in the leaf axils; corolla greenish at the base, orange to salmon-coloured above; fruit a pubescent capsule. Grows in shady, damp places in the Grampians, Vic., and in SA on Kangaroo Is. and Mt Lofty Ranges. Flowering mainly summer. Sometimes included in the genus Billardiera as B. bignoniacea. (a ×1.2, b ×4, c ×16)
Fig. 130 Pittosporum angustifolium (Weeping Pittosporum)K5 C(5) A5 G(2)
Shrub or small tree to 7 m high; branches pendulous; leaves glabrous, narrowlinear, 4–10 cm long; flowers yellow, solitary or in small axillary cymose clusters; petals often virtually united when young, becoming free with age; fruit an ovoid orange capsule 1–2 cm long, the seeds red and sticky. Occurs in dry inland areas of all states except Tasmania. Flowering winter to spring. (a–b ×5, c ×10)
Fig. 131 Pittosporum undulatum (Sweet Pittosporum)K(5) C5 A5 G(2)
Bushy tree to 15 m high; leaves glabrous, lanceolate, 6–12 cm long sometimes appearing whorled near the ends of the branches, margins undulate; flowers cream, fragrant, in leafy clusters; fruit a globular orange capsule, the seeds brown and sticky. In cultivation, plants with only female flowers are quite common. In Victoria originally restricted to the wet gullies of the east, but now widely planted in hedges and parks, and escaped to become an invasive weed in some areas, particularly near-coastal. Also in NSW and Qld, rare in north-western Tas. (a–b ×5, c ×20)
Celery, Carrot, Parsley and allies
This is a widely distributed, medium-sized family important for numerous food and spice plants including Anethum (Dill), Apium (Celery), Carum (Caraway), Coriandrum (Coriander), Cuminum (Cumin), Daucus (Carrot), Foeniculum (Fennel), Pastinaca (Parsnip) and Petroselinum (Parsley). A number of species, such as members of the genus Conium (Hemlock), are seriously poisonous.
The older name, Umbelliferae, is still acceptable, and alludes to the inflorescence type common in the family.
The family is closely related to Araliaceae, and the two have at times been united. In current classifications they are kept separate and, together with Pittosporaceae and several other families, are placed in the order Apiales.
FLORAL STRUCTURE
| Flowers | Usually actinomorphic, usually bisexual, nearly always 5-partite, mostly in simple or compound umbels. |
| Calyx | Sepals usually 5, free, sometimes very small, sometimes absent. |
| Corolla | Petals usually 5, free. |
| Androecium | Stamens 5, free. |
| Gynoecium | Carpels 2, united. A nectar-secreting disc (Pl. 5b) covers the top of the ovary (called a stylopodium in some texts). Ovary inferior. Placentation usually regarded as axile but sometimes effectively apical. |
| Fruit | Dry, indehiscent, splitting into two halves (each a mericarp) from the central stalk (the carpophore, Pl. 34g, h). |
Plants are usually herbaceous, less often shrubby, often with compound or dissected leaves.
Tutin (1980) describes and illustrates British species; many are introduced elsewhere.
ILLUSTRATIONS Figure 132; Plates 5a–c, 34.
SPOTTING CHARACTERS
Plants usually aromatic herbs. Leaves often compound. Inflorescence often a simple or compound umbel. Flowers bisexual, with inflexed petals, the ovary inferior, and the fruit splitting into two halves.
Fig. 132 Foeniculum vulgare (Fennel) ApiaceaeK0 C5 A5 G(2)
Glabrous, upright, herbaceous perennial to 2 m high; leaves compound, dissected into fine segments; flowers in compound umbels; stamens mature and drop off in sequence before the styles and stigmas develop; the top of the ovary is covered by a yellow disc; fruit dry, splitting into 2 segments. Native to southern Europe, widely naturalised and often weedy. Flowering late spring to early autumn. (a ×0.5, b–e ×12)
