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Wood Anatomy of Selected West African Species Of IAWA Journal, Vol. 20 (2), 1999: 115-146 WOOD ANATOMY OF SELECTED WEST AFRICAN SPECIES OF CAESALPINIOIDEAE AND MIMOSOIDEAE (LEGUMINOSAE): A COMPARATIVE STUDY by Alexa Höhn Seminar für Vor- und Frühgeschichte, Archäologie und Archäobotanik Afrikas , Johann Wolfgang Goethe-Universität, Robert-Mayer-Str. 1,60325 Frankfurt/M., Germany SUMMARY Leguminosae constitute an important proportion of the charcoal sam­ pIes recovered at archaeological sites in the West African savannas. Identification of these fragments to a level below family or subfamily was problematic, because a comparative survey was missing. There­ fore, the wood anatomy of 31 species (23 genera) of Mimosoideae and Caesalpinioideae growing in the Sudanian savannas ofWest Africa was examined. The species were grouped into 18 types according to wood anatomical structure. The types represent single species or genera (four­ teen types), two genera (three types) or three genera (one type). The following features are regarded as suitable for a reliable delimitation and identification . Heterocellular rays and storied structure allow for a first differentiation. Enlarged, non-bordered vessel-ray pitting , non­ vestured vessel-pits, silica, axial canals, septate fibres and crystals in non-eharnbered ray cells are additional features characterizing few or single types. Types without these features are delimited less easily. Parenchyma distribution and ray width are, due to variability, not as reliable, but remain necessary features for identification. Types charac­ terized by these features only may not always be recognized correctly. Quantitative features of the vessels are not regarded as helpful for the differentiation within the set of examined species. A table (Table 1) summarizes the results for easy reference . Key words: Mimosoideae, Caesalpinioideae, Fabaceae, West Africa, wood anatomy, wood identification. INTRODUCTION The present survey is intended to compile dependable data, provide wood anatomi­ cal descriptions and evaluate the possibilities and reliability of the identification of selected West African Leguminosae. It was triggered by research on archaeobotany and vegetation history. The woody Leguminosae, as a significant element of tropical floras, play an important role in the charcoal sampIes recovered at archaeological excavations undertaken in Burkina Faso and Nigeria by the joint research project "History of Culture and Language in the Natural Environment of the West African Downloaded from Brill.com10/06/2021 10:09:34PM via free access 116 IAWA Journal, Vol. 20 (2), 1999 Savanna" (Neumann in press). Woody Leguminosae in partieular eharaeterize eertain types of vegetation and/or are valued for providing wood, fruits and fodder. lsober­ linia doka Craib & Stapf, for instanee, is the eharaeteristie speeies of the "forets claires" in the Sudano-Guinean zone and is thought to have been a eonstituting ele­ ment of the aneient dry forests , rieh in woody Legumes, whieh nowadays exist in reliets only (Aubreville 1950). In addition, the "Sudanian lsoberlinia and related woodland", a vegetation-type which extends from Mali to Uganda (White 1983), is named after the genus, thus indieating its signifieanee. The adjoining vegetation to the North is dominated by other Leguminosae, such as Prosopis africana,Acacia spp. and Tamarindus indica (White 1983). In this way, a fragment of lsoberlinia doka will provide information about past vegetation and climate only if it is reeognized. The level of identifieation determines the possibilities of interpretation. When identifying wood fragments it is sometimes admissible to take the additional measure of redueing, by geographical means, the set of speeies taken into aeeount. Possible shifts in distribution over time due to ehanges in climate or to anthropogenie influenees have to be eonsidered. Correspondingly, the examined speeies were se­ leeted aeeording to geographical distribution for the survey presented here.Aubreville (1950) lists about 100 speeies ofLeguminosae for the Sahelian, Sudanian and Guinean zones ofWest Afriea - the Sahelian and Sudanian speeies being of greater importanee for the arehaeobotanieal research in Burkina Faso and Nigeria . Sinee some of the Sahelian speeies have already been examined within arehaeobotanieal frameworks (Neumann 1989; Rolando 1992; UebeI1996), the present survey foeused on speeies of Sudanian distribution. Papilionoideae of the Sudanian zone [e.g., Pericopsis laxi­ flora (Baker) van Meeuwen, Cordyla pinnata (A. Rieh.) Milne-Redh., Pterocarpus erinaceus Poir.], however, were excluded. As a group, they show a distinet wood strueture, eharaeterized by storeying of some or all elements, homoeellular rays and mainly two-eelled parenehyma strands. The type of elements that are storied, paren­ ehyma distribution, and ray width distinguish the genera from one another. While the eharaeteristie wood strueture of the Leguminosae usually allows for a se­ eure delimitation of the family, or subfamily as with the Papilionoideae of the region, a further subdivision has so far proven to be diffieult. During the survey it beeame evident that the wood anatomy of some of the examined taxa does not allow iden­ tifieation down to speeies and in some eases even to genus level. Instead, wood types were reeognized, most of them eomprising species of the same genus, but some erossing generie borders. In delimiting the types two objeetives were followed. The aim of identifieation to the lowest possible taxonomie level made eomprising as few speeies as possible the first objeetive . However, separating types with overlapping features does not render the required high level of aeeuraey of identifieation. The seeond ob­ jeetive was, therefore, to set up easily reeognizable and weIl delimited types. There is published information on the wood anatomy of all the examined genera, but some of the speeies have not yet been deseribed. Even though wood identifieation guides toparticular regions, as Lebaeq (1957) or Normand (1950) foeus on Guinean Downloaded from Brill.com10/06/2021 10:09:34PM via free access Höhn - Wood of West African Leguminosae 117 rather than Sudanian species, they contain some of the examined species . Some ofthe examined genera are included in taxonomie surveys .(e.g. Baretta-Kuipers 1981). Wood anatomical research on Leguminosae in an archaeobotanical context mainly focuses on the important, yet difficult differentiation of species within the genus Acacia (Ro­ lando 1992; Barakat 1995). MATERIALS AND METHODS In this study 155 specimens representing 31 species in 23 genera were examined. Samples could not be obtained for all genera with Sudanian distribution listed by Aubreville (1950), and only selected species were examined for large genera. 116 slides were derived from samples of more than four centimetres diameter. For the description of the wood structure, only these 'mature' specimens were taken into account, 73 specimens (19 species) belonging to Caesalpinioideae and 43 specimens (12 species) to Mimosoideae. 39 slides (20 of Caesalpinioideae, 19 of Mimosoideae) of specimens with a diameter of four centimetres and less - 'branch wood' - were examined for comparison. Most of the sampIes examined are part of the collection at the "Seminar für Vor­ und Frühgeschichte, Archäologie und Archäobotanik Afrikas" at Frankfurt/Main University. Sampies or slides received from other institutes are indicated by Stem's (1988) acronyms . The collection numbers of samples taken by members of the joint research project are preceded by the abbreviation 'SFB' . Nearly all of these samples are backed by Herbarium vouchers at the "Naturmuseum und Forschungsinstitut Senckenberg" (FR). In an appendix, the studied material is listed alphabetically, divided into 'mature wood' and 'branch wood' material. If known , it is referred to geographical origin, collector and herbarium voucher, wood collection number, and diameter of the sample , The wood samples were sectioned on a rotary microtome, bleached, stained with Safranin and mounted in Euparal for light microscopical observation. For SEM ob­ servations, sectioned wood blocks were coated with gold using an IB-2 ion coater and observed with a Hitachi S 4500 . Terminology and methodology follow the IAWA list of microscopic features for hardwood identification (IAWA Committee 1989), if not stated otherwise. Accord­ ingly, vessel diameters, for example, were measured tangentially 25 times, excluding the walls. Vessel element length , however, was only measured ten times and not in maceration. The quantitative data are given as ranges of mean values of the different samples and/or species, usually followed by upper extremes of all the samples taken together. For ray width, the most frequent ranges are given. Single mean values are cited when there is no range because only one sample was obtained for the corre­ sponding species. Furthermore, the mean number of vessels per group, as suggested by Carlqui st (1988), is given as 'vessel grouping index'. Features not explicitly men­ tioned were absent or not applicable. The description was generated with DELTA (Dallwitz 1980; Dallwitz et al. 1996), according to the character list by Richter & Trockenbrodt (1995, 1996). Downloaded from Brill.com10/06/2021 10:09:34PM via free access 118 IAWA Journal , Vol. 20 (2),1999 DESCRIPTIONS In line with the above mentioned objectives, the species were grouped into 18 weIl delimited anatomical types, comprising species or genera with similar features . The types represent single
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