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Wood Anatomy of Brabejum and Macadamia (Macadamiinae, Proteaceae)

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Wood Anatomy of Brabejum and Macadamia (Macadamiinae, Proteaceae) Botanica Pacifica. A journal of plant science and conservation. 2019. 8(2): 35–41 DOI: 10.17581/bp.2019.08204 Wood anatomy of Brabejum and Macadamia (Macadamiinae, Proteaceae) Anna V. Stepanova1, 2* & Alexei A. Oskolski1,2 AnnaV. Stepanova1,2* ABSTRACT e-mail: [email protected] Wood structure of Brabejum stellatifolium and Macadamia integrifolia (subtribe Maca­ Alexei A. Oskolski1,2 damiinae, tribe Macadamieae, Proteaceae) is described for the first time. These spe­ e-mail: [email protected] cies share exclusively simple perforation plates, minute to small alternate intervessel pits, non-septate libriform fibers, banded and unilaterally paratracheal axial paren­ chyma, and the rays of two distinct sizes, i.e. the suite of typical wood traits for 1 Department of Botany and Plant many of Proteaceae. Our observations together with reported data suggest that Biotechnology, University of Johannes­ the lineage comprising Brabejum, Macadamia and Panopsis is distinctive from closely burg, Johannesburg, South Africa related members of the tribe Macadamieae by the occurrence of vasicentric tra­ 2 Komarov Botanical Institute RAS, cheids combined with the lack of fiber tracheids. More abundant axial parenchyma Saint-Petersburg, Russia in Macadamia and Panopsis than in Brabejum is seemingly consistent with the global tendency to increase of amount of this tissue in the regions with warm climate. Libriform fibers is Macadamia integrifolia show one of the highest degrees of intru­ * corresponding author siveness (F/V ratio = 3.98) reported within Proteaceae to date. Keywords: Brabejum, Macadamia, Proteaceae, wood anatomy, vasicentric tracheids Manuscript received: 21.05.2019 Review completed: 17.07.2019 РЕЗЮМЕ Accepted for publication: 22.08.2019 Published online: 30.08.2019 Степанова А.В., Оскольский А.А. Анатомия древесины Brabejum и Macadamia (Macadamiinae, Proteaceae). Впервые описано анатомичес- кое стро ение древесины Brabejum stellatifolium и Macadamia integrifolia (подтри­ ба Macadamiinae, триба Macadamieae, семейство Proteaceae). Для этих видов ха рактерен набор признаков, типичных для многих протейных, таких как исключительно простые перфорационные пластинки, очень мелкие по- ры межсосудистой поровости, несептированные древесинные волокна, аки с альная паренхима односторонне-паратрахеальная и в тангентальных по лосах, две отчетливых размерных группы лучей. Наши и литературные данные указывают на то, что монофилетическая группа, включающая роды Brabejum, Macadamia и Panopsis, отличается от других представителей Mac a da- mieae наличием вазицентрических трахеид в сочетании с отсутствие волок - нистых трахеид. Более обильная аксиальная паренхима у Macadamia и Panopsis по сравнению с Brabejum соответствует глобальной тенденции к увеличению количества этой ткани у растений в регионах с тёплым климатом. Древесин­ ные волокна Macadamia integrifolia характеризуются одним из самых высоких уровней интрузивности (отношение средней длины волокон к средней дли­ не члеников сосудов равно 3,98), известным для протейных. Ключевые слова: Brabejum, Macadamia, Proteaceae, анатомия древесины, вазицент- ри ческие трахеиды Subtribe Macadamiinae L.A.S. Johnson & B.G. Briggs land and one species from Sulawesi. MonospecificNothorites belonging to the tribe Macadamieae C. Venkata Rao of P.H. Weston, that has initially been described as a species of Wes ton & Barker’s (2006) classification of Proteaceae Juss. Orites R. Br. from the tribe Roupaleae (George & Hyland com p rises five genera with 36 species of evergreen trees 1995), is also endemic of north-eastern Queensland. and shrubs with mostly verticillate leathery leaves (Wes­ Unlike these three genera, 25 species of Panopsis Salisb. are ton 2007). This small lineage is widespread across the sou- widespread in shrublands, savannas, xerophytic woodlands, thern hemisphere. The best known genus of this group is riparian forests and lowland to montane rainforests of tro- Macadamia F. Muell. with four species which are naturally pi cal South and Central America (Prance et al. 2007, Weston dist ri bu ted in subtropical regions of eastern Australia, two 2007). Finally, Brabejum stellatifolium L., the only species of of which (M. integrifolia Maiden & Betche and M. tetraphylla the genus Brabejum L., is a component of fynbos biome L.A.S. John son and their hybrids) are widely cultivated and Afrotemperate forests restricted to the Cape region for their edible seeds (macadamia nuts). The genus Lasjia in South Africa with its Mediterranean climate (Mucina & P.H. Weston & A.R. Mast, that has recently been separated Rutherford 2006, Weston 2007, Mast et al. 2008). from Macadmia (Mast et al. 2008), comprises four tro pi cal The time of radiation of the subtribe Macadamiinae is species from tropical rain forests of north-eastern Queens - estimated as the late Oligocene – early Miocene. Disjunct ©Botanical Garden-Institute FEB RAS. 2019 35 Stepanova & Oskolskii dist ri bu tion of this group is seemingly a result of long-distance dispersal of their buoyant diaspores from Australia to South America and Africa by the An- tarc tic Circumpolar Cur rent that has also developed in the Oligocene. The evo- 2.3±0.21 1.0–3.2 18 4.3±0.13 3.4–5.7 lu tio nary shifts from the early fruit and seed maturity found in Macadamia and other Australian members of this group to the tardy dehiscence at the seed ger mi na tion, which is sha red by extra-Australian taxa are well consistent with ofm); 16– number μ 1.1±0.22 0–2.1 17 3.1±0.14 1.9–4.9 this scena rio (Mast et al. 2008). The subtribe Macadamiinae is poorly explored by wood ana tomists: the –1.9 m); 12 –number ofm); 12 –number axial detailed information on wood structure has been published to date only for two 8 μ 1.3±0.14 1.0–2.1 16 species of Panopsis (Mennega 1966, Detienne & Jacquet 1983). Brief wood ana- 1.2±0.07 0. to mi cal description and microphotos are available also for Macadamia ternifolia F. Muell. (Chattaway 1948, InsideWood 2004–onwards). Quantitative data on 3266 409–5729 15 wood density, dia meter of vessel lumina and vessel wall thickness in the Brabejum 1123 229–2893 wood have been reported by Crous et al. (2012). The aim of the current study is 750 – the detailed examination of wood structure of Brabejum and Macadamia, and its 8 279 3 304 33–475 14 com parisons with related groups of Proteaceae. 34 20 – 7 / MATERIAL AND METHODS 3 – –2/7– 1 1 13 Wood sample of Brabejum stellatifolium was collected by Alexei Oskolski on m); 7 – length of fibres (mean ± standard error, min–max, 05.09.2016 in afrotemperate forest on the eastern slope of Table Mountain near μ 2–6 2–4 the Kirstenbosch Bo ta nical Garden, Cape Town, South Africa. Wood sample 12 3 m); 11 – vessels wall thickness (min–max, thickness wall m); 11 – vessels μ –5. of Macadamia integrifolia was taken by him on 10.04.2019 from the tree culti vated m); 15 – height of (mean, min–max, rays multiseriate 8 μ 2. 2.9–5.8 in Pretoria, South Africa. Both wood samples were collected from large bran- 11 ches. Herbarium vouc hers (A.A. Oskolski # 98 for B. stellatifolium and # 281 for M. integrifolia) are deposited in JRAU; wood samples are deposited in the Xyla ri- 2.8–3.7 3.8–10.5 10 um of the Komarov Botanical Institute, St. Petersburg, Russia (LEw). 43 2 Transverse, radial, and tangential microtome sections of 20–30 µm in thick- . 22 ness were stained with alcian blue/safranin (Jansen et al. 2004), dehydrated in ±0. 9 – . 1 15 24.4±0.82 18.6–33.9 8. gra dient series of alco hol, and then mounted in Euparal. Macerations were made 9 using Jeffrey’s solution (Johansen 1940). Descriptive ter mi nology followed the 3.98 2.33 re com mendations of the IAWA Com mittee (1989). The quantitative data are 8 sum ma rized in Table 1. Details of the microstructure of intervessel pits were in ves tigated by scanning electron microscopy (SEM TESCAN, soft – VegaTS) m); 10– fibre wall thickness (min–max, wall thickness m); 10– fibre at Spectrum, University of Johan nesburg. For the SEM examination, dried wood μ 1494±28.6 1080–1890 1657±46.0 1080–2060 7 samples were mounted on aluminum stubs with double-sided carbon tape and coated with gold. Evolution of selected wood anatomical features was cla ri fied by mapping dis- 376±11.4 201–523 710±30.2 299–987 crete characters on a subsample of the tree recovered from the Bayesian analyses 6 9 Macadamia integrifolia m); 2 – number ofm); 2 – number of lumina per sq.mm(mean, min–max) ; 3 – mean/ max number vessel lumina per group; vessel 4 – percent of a data set of three chloroplast DNA and four nuclear DNA sequences as well m); 6– length of min–max, elements (mean ± standard error, vessel μ –5.0 μ 2 4 5 and 4. 3. 4. 3.1–5. as mor phological data for the tribe Macadamieae (Mast et al. 2008). Character 5 opti mi za tion along the branches of the tree was illustrated using the parsimony 2 17 1 re con struction method of the Character History Tracing option in the com pu ter 4 8 package Mesquite 2.75 (Maddison & Maddison 2011). / 2 . .3/12 2 2 3 RESULTS 37 4 23. 17–30 23.1 14– Wood anatomy of Brabejum stellatifolium (AO 98-16) 2 Brabejum stellatifolium Wood diffuse-porous. Growth ring boundaries indis tinctly marked by 3–6 7 5 rows of weakly flattened fibres. Vessels few (up to 25 per mm2), medium to –99 –178 5 7 66±2. 2 110±4. 6 wide (67–178 µm in tangential diameter), solitary, in clusters and in tangential 1 mul tiples of 2–8 (up to 12), thin-walled (wall thickness 2.8–5.3 µm) (Fig. 1a). Ves sel lumina round to slightly angular.
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