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Atlas of Leaf Venation and Oil Gland Patterns in the Eucalypts

Atlas of Leaf Venation and Oil Gland Patterns in the Eucalypts

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Atlas of Leaf Venation and Oil Gland Patterns in the Eucalypts

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431 pages
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Jun 12, 2013


Atlas of Leaf Venation and Oil Gland Patterns in the Eucalypts is an aid to the identification of eucalypts in the field and a confirmation of the natural affinities between species and higher-level taxa on the basis of their comparative morphology. Its purpose is to standardise leaf venation and oil gland terminology and to demonstrate the taxonomic value of leaf venation and oil gland patterns within the eucalypts.

The work discusses the visible features of the adult leaves of eucalypts as seen with reflected and transmitted light. Because venation and oil glands become obscure in dried specimens this work relies entirely on the comprehensive sampling and observation of fresh leaves.

High quality, scaled, leaf venation images of vouchered specimens are used to compare all taxonomic groups in the eucalypts. All genera, sections, series and subseries are represented.

Jun 12, 2013

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Atlas of Leaf Venation and Oil Gland Patterns in the Eucalypts - Ian Brooker



In this study we discuss the visible features of the adult leaves of eucalypts (and those of the neotenous species) as seen with reflected and transmitted light (Figure 1). We intend this work to be an aid in field identification and for confirmation of natural affinities between species and higher-level taxa on the basis of their comparative morphology. It is desirable that the illustrations, descriptions and findings in this book be complemented later by anatomical studies to resolve the internal structure and possible functional implications behind the images shown.

There are no comprehensive comparative studies of the internal structure and appearance of the leaves in the eucalypts. Descriptive Flora treatments have conventionally been confined to the size, shape and colour of the leaves, observations on the principle veins and the apparent ‘density’ of the oil glands. The venation and oil glands in the leaf become obscure in dried specimens, and this work has only been possible by the comprehensive observation and sampling of fresh leaves.

Figure 1. Part of a leaf of a) C. citriodora and b) E. gregsoniana, photographed with reflected light and with transmitted light. The secondary and tertiary venation and oil glands are only readily visible when viewed with transmitted light. As such, all other images in this book have been taken using mostly or entirely transmitted light with little or no reflected light.

Background – leaf venation

Maiden (1922–1923) summarised in considerable detail venation patterns given by various earlier authors. He placed many eucalypt species in groups with similar venation but made no overall classification. He emphasised that ‘real genetic relationships take cognisance of all the characters’. Maiden’s work is not illustrated.

Blakely (1934) made some references to leaf venation, but his terminology is not very informative. For example, for the bloodwoods (now Corymbid), he states, ‘almost transverse’ in his summary for E. ser. Corymbosae, then within the species descriptions, he uses many other terms such as ‘finely veined’, ‘very fine and obscure’, ‘very fine and inconspicuous’, ‘fine and semitransverse’, ‘fine and subtransverse’, ‘close subparallel, ‘close and semitransverse’ and ‘almost parallel’.

For later taxonomic descriptions, Blakely does not in every case give venation patterns although he did for some species, e.g. the monocalypt E. rossii, in which he described the venation as ‘penninerved’. The venation of E. rossii (p. 205) could in no way be described as ‘penninerved’ as generally understood. It must be said that what various authors meant by many terms such as ‘transverse’, ‘penninerved’, ‘pennivenation’, ‘pinnate’, ‘parallel’, etc., is not always clear, particularly without illustration. We prefer the use of the terms ‘pinnate’ and ‘pinnation’, when referring to relevant species, in particular Angophora, Corymbia and Eucalyptus sect. Latoangulatae. More contemporarily the use by various eucalypt authors of numerous terms describing the position of the intramarginal vein, the density of the tertiary venation, and the frequency of visible oil glands, is likewise unclear, and we have attempted to clarify the terminology by clearly defining some of these terms.

An attempt was made to characterise leaf venation by Blake (1953) who gave for each species of bloodwood (Corymbia) the number of pairs of secondary (side) veins and the angle at which they left the primary vein (midrib). Blake’s scheme has not taken on acceptance although the secondary vein angle has sometimes been given in descriptions by later authors. Here we provide the measured secondary venation angle for all 363 samples illustrated in the Atlas, as well as providing descriptive terminology for the secondary venation angle.

Baker and Smith (1920) discussed the evolution of the eucalypt leaf from the multiple secondary-veined (pinnate) Angophora and Corymbia, through the discolorous-leaved, eastern blue gums (E. sect. Latoangulatae) to some of the peppermints (monocalypts) which they imply are highly modified taxa. As such, in the peppermints they refer to the ‘subordination’ of the reticulation and the ‘room given for the formation of oil glands’ – island oil glands as we discuss later. The principal purpose of their publication is, however, eucalypt oils.

In protologues for new eucalypt species, the discussion of leaf venation is not a requirement and in any case is not diagnostic for some species. L.A.S. Johnson’s earliest descriptions in 1962 were provided in Latin only, not illustrated, and were therefore not readily accessible. For E. corticosa of New South Wales, Johnson (1962) stated, ‘venatione modice regulari venis angulum circiter 30–40° cum costa media formantibus’ (venation moderately-regular with the angle of the veins at 30–40° to the midrib). For E. cypellocarpa of New South Wales and Victoria in the same publication, he provided no description of the venation at all.

In Johnson’s later work co-authored with K.D. Hill, the diagnosis in Latin is followed by a comprehensive description in English. The authors make reference to the venation, but the use of words like ‘regular’, ‘not regularly pinnate’, ‘spaced’, etc., are of little value without illustrations or definitions. For their recircumsciption of E. spathulata (E. ser. Erectae), the description was ‘reticulation sparse, incomplete; oil glands crowded, spherical’ (Hill & Johnson 1992). The term ‘spherical’ is three-dimensional which may be correct and based on Baker and Smith (1920), but cannot be judged with certainty from transmitted light. E. spathulata has similar venation and gland patterns to several related species of Western Australia, e.g. E. utilis and E. tenera (p. 81).

Carr et al. (1986) in a detailed study mainly focusing on ontogeny, demonstrated that the intramarginal vein was formed by the linking of the ends of the secondary veins only in juvenile leaves. They distinguished this feature by the introduction of the term ‘paramarginal vein’. In the true adult leaves, they found that the veins linking the secondary veins (intramarginals) were independently formed. We refer to both forms as the ‘intramarginal vein’, although most illustrations included in the Atlas are from adult leaves. In their work on leaf venation and ovule patterns, Carr and Carr (1986) studied internal and therefore protected structures, thereby shifting the emphasis from the length, breadth, shape and colour of the organs, all of which are variable and of only moderate reliability in identification.

Chippendale (1988) gave lateral vein angles and a comment on the intramarginal veins for all 513 species in the Eucalyptus Flora treatments. His descriptions included ‘lateral veins faint’ for many species. It is probable that his observations were made from dried herbarium specimens only. Two of the most widespread Corymbia species (C. polycarpa and C. terminalis), whose distributions broadly overlap in northern Australia, have superficially similar leaves. For C. polycarpa, Chippendale stated that the intramarginal vein was confluent with the leaf margin. Under C. terminalis, he stated that the intramarginal vein was very close to the leaf margin. Our observations are that the situation is the reverse for these two species. Blakely (1934) recognised that for some bloodwood (Corymbia) species the intramarginal veins are ‘confluent and indistinguishable from the thick nerve-like margin’. The secondary veins in these species ‘run’ direct to the leaf edge.

In the Canberra foothills there are two common white gums, viz. the scribbly gum (E. rossii of E. subg. Monocalyptus) and the brittle gum (E. mannifera of E. subg. Symphyomyrtus). These two species are similar in stature, bark, and leaf shape and size, and they may be difficult to distinguish without flowers if the Ogmograptis moths, whose larvae cause the prominent black ‘scribbles’ on the outside of the bark of E. rossii, are absent (probably through isolation of the trees as outliers from natural populations where the moths are abundant or when the trees are cultivated). Familiarity with the leaf venation patterns for these two species allows instant identification, i.e. by the very sparse tertiary venation in E. rossii and the moderate tertiary venation seen in E. mannifera (Figure 2). This phenomenon is a common pattern in parallel situations elsewhere in the eucalypts.

Figure 2. a) E. rossii and b) E. mannifera. These two superficially very similar but distantly-related species can be immediately differentiated by the venation pattern in the adult leaf.

Background – leaf oil glands

Welch (1920) made a study of the leaf oil glands in the eucalypts, particularly with respect to their formation. Basically they are formed by ‘the breaking down of the secretory cells’, or ‘separating apart of the secretory cells’, or a combination of both. He discussed the glands of seven species but for the oil contents only.

Baker and Smith (1920) made anatomical studies of some eucalypt leaves. Their transverse sections show that the oil glands are more or less spherical and not distributed throughout the leaf tissue, i.e. there is a row beneath the upper epidermis and another above the lower epidermis. The larger upper glands often extend to the centre of the leaf. The upper glands are usually more numerous than the lower (see Figure 3).

Oil glands in the leaves can be apparently absent (not visible). With fresh leaves of C. ficifolia, this provides an easy distinction from the co-occurring bloodwood, C. calophylla, both of southern Western Australia.

Within the leaf tissue the glands must be three-dimensional (Baker

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