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
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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).
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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 species or genera (14 types) , two genera (three types) or three genera (one type) of leguminous woods of Sudanian West Africa. The type including three genera serves as a 'catch-all' type for the identification of Caesalpinioideae and Mimosoideae. These genera may be identified as not being papilionoid, while it is impossible to decide whether they belong to Mimosoideae or Caesalpinioideae. The description starts with this basic Caesalpinioideae /Mimosoideae type, the others fol 10w in alphabeticalorder. Some diagnostic features of individual types are printed in bold.
The following features are found in all of the species examined: vessel pits alternate and perforation plates simple, fibre pits mainly restricted to radial walls and simple to minutely bordered.
Caesalpinioideae/Mimosoideae type - Fig. 1 & 2 Description based on 19 specimens, 5 of Amblygonocarpus andongensis (Oliv.) Exell & Torre, 4 of Burkea africana Hook., 4 of Erythrophleum africanum (Benth.) Harms and 6 of Erythrophleum suaveolens (Guill. & Perr.) Brenan. Growth ring boundaries absent or inconspicuous, marked by interrupted marginal parenchyma bands, by radially flattened, sometimes thick-walled fibres, and accom panied by small vessels or vessel groups (in addition conspicuously bulging rays in Burkea africana and weakly so in Amblygonocarpus andongensis). Vessels diffuse, 3-17 per mm- , solitary (40-80%), in radial multiples of 2-4 and in clusters, vessel grouping index 1.1-2.0. Tangential diameter 110-210 um, maxi mum 320 um. Vessel element length 240-400 um. Intervessel pits vestured, 6-10 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders, similar to intervessel pits. Rarely thin-walled tyloses and locally brown, gum-like deposits . Fibres medium thick-walled to thick-walled, gelatinous layers often present. Axial parenchyma aliform to confluent, rarely to banded, in strands of 2-4 cells, next to vessels up to 6 cells. Parenchyma sheath sometimes irregular and almost uni lateral (in Burkea africana up to unilateral). Apotracheal diffuse parenchyma spo radic, often crystalliferous. Rays 6-12 per mm, 2-4(-5) cells wide, homocellular, composed of procumbent cells. Large rays 150-450 um high, maximum 1.4 mm. Crystals solitary, prismatic, located in chambered axial parenchyma cells.
Note: This type probably contains more species than the above mentioned because it represents a basic type of the subfamilies. Burkea africana is described by Banks & Kromhout (1966) and Gottwald & Noack (1966). Erythrophleum africanum and E. suaveolens (as E. guineense G. Don) are described by Lebacq (1957). Normand (1950) included E. suaveolens (as E. guineense G. Don) in his characterization of the genus.
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Fig. 1-4. - 1 & 2: Caesalpinioideae/Mimosoideae type. - 1: Amblygonocarpus andongensis (CTFw 21725) , TS, confluent parenchyma. - 2: A. andongensis (RBHw 11363), TLS , mostly 3-seriate rays. - 3 & 4: Acacia type . - 3: Acacia seyal (CTFw 27058), TS, paratracheal confluent paren chyma, crystals in apotracheal diffuse parenchyma, wide rays. - 4: Acacia sieberiana (SFB 594.0 ex Bw) , TLS , 6-7-seriate rays, mostly 2-celled parenchyma strands. - Scale bars = 1 mm.
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Fig. 5 - 8. - 5 & 6: Afzelia/Tamarindus type, Afz elia africana (CTFw 19960). - 5: TS, alifonn pa renchyma and marginal bands, radial fibre pattern. - 6: TLS, 3- or 4-seriate rays. - 7 & 8: Alb izia type, Albizia cf. malacophylla (SFB 571.1). - 7: TS, paratracheal alifonn to confluent and apotracheal diffuse parenchyma. - 8: TLS, paratracheal parenchyma in strands and apotracheal parenchyma fusifonn . - Scale bars = I mm.
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Acacia type - Fig. 3 & 4 Description based on 14 specimens , 4 of Acacia senegal (L.) Willd., 5 of A. seyal DeI. and 5 of A. sieberiana DC. Growth ring boundaries distinct, marked by marginal parenchyma bands, 1-3 (-5) cells wide, sporadically to continuously crystalliferous. Vessels diffuse, 5-19 per mm-, solitary (20-80%), in radial multiples of 2-4 and in clusters, sporadically in groups of more than 10 vessels, vessel grouping index 1.2-2.7. Tangential diameter 90-150 um, maximum 250 um. Vessel element length 150-280 um. Intervessel pits vestured, 7-9 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders, similar to intervessel pits. Locally brown, gum like deposits. Fibres medium thick-walled to thick-walled, gelatinous layers often present. Axial parenchyma (aliform to) confluent and in discontinuous and irregular bands, 5-20(-30) cells wide. Apotracheal parenchyma diffuse, mostly with large crystals. Parenchyma in strands of2(-4) cells and occasionally fusiform. Parenchyma sheaths sometimes irregular to almost unilateral. Rays 4-9 per mm, 4-10(-15) cells wide (widest in A. seyal with up to 10, but reg ularly up to 8 in the other two species), homocellular, composed of procumbent cells. Large rays 250-800 ).Ut1 high (250-400 um in A. senegal, 350-700 um in A. sieber iana, 500-800 um in A. seyal), maximum 2 mm. Crystals solitary, prismatic , located in chambered axial parenchyma cells (the lat ter conspicuously enlarged in A. seyal). Rarely crystals in chambered ray cells.
Note: Lebacq (1957) characterizes the rays of A. sieberiana as heterogeneous, but they are described as homogeneous by Gottwald & Noack (1966). Abbate (1963) examined A. seyal and A. sieberiana (in Edlmann Abbate 1970). Descriptions of the examined Acacia spp. can also be found in papers on charred wood, in Neumann (1989) as weIl as in Rolando (1992) and in Barakat (1995). The latter ascribes hetero cellular rays to A. seyal.
Afzelia/Tamarindus type - Fig. 5 & 6 Description based on 10 specirnens, 6 of Afzelia africana Pers. and 4 of Tamarindus indica L. Growth ring boundaries distinct or occasionally indistinct, marked by marginal parenchyma bands, 1-3(-4) cells wide, sporadically crystalliferous, and accompa nied by small vessels or vessel groups. Vessels diffuse, 2-10 per mm? (2-4/mm2 in Afzelia africana, 7-1O/mm2 in Tamarindus indicai, solitary (50-90%), in radial multiples of 2-4 and rarely in clus ters, vessel grouping index 1.l-1.6. Tangential diameter 100-230 um (100-130 um in Tamarindus indicai , maximum 310 um. Vessel element length 200-350 um. Intervessel pits vestured, 6-8 mm in diameter, apertures sometimes coalescent. Ves sel-ray pits with distinct borders, similar to intervessel pits. Locally brown, gum-like deposits.
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Fibres medium thiek-walled, oeeasionally with a distinet radial pattern due to regular intrusive growth, gelatinous layers often present. Axial parenehyma aliform to shortly eonfluent (2-3(- 5) vessels/v essel group s), in strands of 2- 4 eells. Diffuse apotraeheal parenehyma sporadie, often erystalliferou s. Rays 6-10 per mm (6- 7 in Afzelia africana, 9-10 in Tamarindus indica) , 2-4 eells wide (up to 8 eells in Afzelia af ricana SFB 1171.0, higher portion of uniseriate rays in Tamarindus indica) , homoeellular, eomposed ofproeumbent eells. Large rays 150-400 um high, maximum 700 1JIll. Crystals solitary, prismatie, loeated in ehambered axial parenehyma eells, at the margins of aliform parenehyma eonspieuous.
Note: Chalk et al. (1932, 1933) deseribe tendeneies of storied strueture in Afzelia afri cana Sm. Aceording to Gill et al. (1983) Afzelia africana Harms (!) and Tamarindus indica show heterogeneous rays. Further deseriptions on Af zelia africana Sm. are published by Gottwald & Noack (1966), and on Tamarindus indica by Abbate (1963), by Jagiella & Kürschner (1987) as weil as by Detienne & Ter Welle (1989). Normand (1950) included A. africana Sm. in his description of the genus.
Albizia type - Fig. 7 & 8 Deseription based on 2 speeimens, I of Albizia chevalieri Harms and I of A lbizia ef. malacophylla (A.Rieh.) Walp. Growth ring boundaries indistinet, marked by interrupted marginal parench yma bands, sporadieally erystalliferous, by radially flatten ed fibres, and aeeompanied by small vessels or vessel groups. Vessels diffuse, 3- 6 per mm-, solitary (80- 90%) and in radial multiples of 2- 4, vessel grouping index 1.1. Tangential diameter 130-150 1JIll, maximum 220 1JIll. Ves seI element length 270-280 mm. Inter vessel pits vestured, 4- 7 mm in diameter, aper tures not or faintly eoaleseent. Vessel-ray pits with distinct borders, similar to intervessel pits. Loeally brown, gum-like deposits. Fibres medium thick-walled , exclusively septate, oceasionally with a distinet pat tern due to regular intrusive growth, gelatinous layers often present. Axial parenchyma aliform to eonfluent, in strands of 2-4 eells. Apotracheal pa renchyma diffuse and fusiform, predominantly without erystals. Rays 6-8 per mrn, 1-3 eells wide, homoeellular, eomposed of proeumbent eells. Large rays 150-200 um high, maximum 350 1JIll. Crystals solitary, prismatie, loeated in ehambered axial parenchyma eells.
Note: Other speeies of the genus were surveyed for fusiform apotracheal parenchyma. The feature was also found in Albizia zygia (DC.) J.F. Maebr. (SFB 504.0) but not in A. amara (Roxb.) Boivin (SFB 122.0, 123.1) and A. aylmeri Huteh. (SFB 426.0). Published deseriptions on the examined species were not found, but the feature ean also be seen in a photograph of Albizia zyg ia by Gottwald & Noaek (1966), even though it is not menti oned in the description . Metealfe & Chalk (1950) mention apotraeheal diffuse parenehyma as weil as fusiform parenehyma as features of Albizia spp. The two features are not, however, put into relation.
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BauhinialPiliostigma type - Fig. 9-12 Description based on 11 specimens, 2 of Bauhinia rufescens Lam., 4 of Piliostig ma reticulatum (DC.) Hochst. and 5 of Piliostigma thonningii (Schum .) Milne-Redh. Growth ring boundaries inconspicuous to distinct, marked by interrupted marginal parenchyma bands. Vessels diffuse, 3-40 per mm- [20-40/mm2 in Bauhinia rufescens, 3-9(-20) in Piliostigma spp.], solitary [(20-)40-90%], in radial multiples of 2-4 and rarely in clusters, vessel grouping index 1.1-1.7(-3.2). Tangential diameter 60-150 um(60 90 um in Bauhinia rufescens , 100-150 um in Piliostigma spp.), maximum 230 um. Vessel element length 190-280~. Intervessel pits not vestured, 7-10 um in diam eter, apertures coalescent. Vessel-ray pits with reduced borders or simple, rounded and elongated (rarely horizontally elongated in Bauhinia rufescens, often diagonally to vertically in Piliostigma spp.), Locally thin-walled tyloses and brown, gum-like deposits. Fibres medium thick-walled to thick-walled, gelatinous layers often present, occa sionally some fibres septate (only in Bauhinia rufescens). Axial parenchyma confluent to discontinuously and irregularly banded, 3-10 (-20) cells wide. Parenchyma in strands of 2-4 cells. Parenchyma sheaths sometimes irregular to almost unilateral. Diffuse apotracheal parenchyma sporadic, often crystalliferous. Rays 16-20 per mm, 1-2(-4) cells wide, heterocellular, square and upright cells in one, occasionally up to two marginal rows. Rays 200-400 um high, maximum 1.3mm. Storied structure regular to irregular (irregular in Bauhinia rufescen s and some specimens of Piliostigma reticulatum), 4-6 ray tiers per axial mm. Low rays, axial ' parenchyma, vessel elements, and fibres storied, large rays 2-3(-6) tiers high. Crystals solitary, prismatic, located in chambered axial parenchyma cells and ty loses, very rarely in chambered upright ray cells .
Note: Septate fibres as found in the examined samples of Bauhinia rufeseens are de scribed by Watson & Dallwitz (1983) for the genus Bauhinia s.1. (including Pilio stigma).
BerlinialIsoberlinia type - Fig. 13 Description based on 15 specimens, 4 of Berlinia grandiflora (Vahl) Huteh. & Dalziel, 8 of Isoberlinia doka Craib & Stapf and 2 of Isoberlinia tomentosa (Harms) Craib & Stapf. Growth ring boundaries distinct, marked by marginal parenchyma bands, 2-4 (-6) cells wide, sporadically crystalliferous, and accompanied by small vessels and vessel groups . Vessels diffuse, 2-5(-7) per mm-, solitary (50-80%), in radial multiples of 2-4 and rarely in clusters, vessel grouping index 1.0-1.5. Tangential diameter (l00-)140 230~, maximum 300~. Vessel element length 270-41O~ . Intervessel pits ves tured, 6-7 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders,
Downloaded from Brill.com10/06/2021 10:09:34PM via free access 124 IAWA Journal , Val. 20 (2),1999 similar to intervessel pits, sometimes unilaterally compound (very rarely in Berlinia grandiflora). Locally brown, gum-like deposits. Fibres medium thick-walled, occasionally ina distinct pattern due to regular intru sive growth, gelatinous layers often present.
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Axial parenchyma paratracheal aliform to confluent, in strands of 2-4 cells, next to vessels up to 6. Diffuse apotracheal parenchyma sporadic, often crystalliferous. Rays 8-11 per mm, 1-2(-3) cells wide (a slightly higher portion of uniseriate rays in Berlinia grandiflora), homocellular and heterocellular with one irregular row of square to upright cells. Large rays 150-350 um high, maximum 570~. Crystals solitary, prismatic, located in chambered axial parenchyma cells, very rarely in chambered procumbent or upright ray cells.
Note: Unilaterally compound pits were only discemed within this group, though not in every cross-field. The feature is not mentioned in the descriptions of Berlinia gran diflora by Gill et al. (1983) or Normand (1950), nor in those of B. grandiflora and Isoberlinia tomentosa by Lebacq (1957).
Cassia sieberiana DC. - Fig. 14 & 15 Description based on 3 specimens . Growth ring boundaries indistinct, marked by interrupted marginal parenchyma bands, with sporadically crystalliferous cells, and by radially flattened fibres. Vessels diffuse, 5 or 6 per mm- , solitary (40-80%), in radial multiples of2-4 and rarely in clusters, vessel grouping index 1.2-1.8. Tangential diameter 120-190 um, maximum 280 um. Vessel element length 240-330 urn. Intervessel pits vestured, 9-10 mm in diameter, apertures coalescent. Vessel-ray pits with distinct borders, sim ilar to intervessel pits. Rarely thin-walled tyloses and locally brown, gum-like de posits. Fibres medium thick-walled to thick-walled, few to many septate, gelatinous lay ers often present. Axial parenchyma paratracheal, aliform to confluent, rarely discontinuously and irregularly banded, in strands of2-4 cells, next to vessels up to 6. Parenchyma sheath sometimes irregular to almost unilateral. Little apotracheal diffuse parenchyma, of ten crystalliferous. Rays 7-11 per mm, 2 or 3 cells wide, homocellular, composed of procumbent cells. Large rays 150-350 um high, maximum 450~. Crystals solitary, prismatic, located in chambered axial parenchyma cells.
Note: The species is included in Normand's (1950) description of the genus. He does not, however, refer to septate fibres.
~ Fig. 9-13. - 9-12: Bauhinia/Piliostigma type. - 9 & 10: Piliostigma thonningii (CTFw 27027). 9: TS, paratracheal confluent to banded parenchyma and marginal bands, tyloses. - 10: TLS, storied structure and uniseriate rays. - 11: Bauhinia rufescen s (RBHw 17397 ex CTFw 27066), RLS, ves sel-ray pits rounded and simple. - 12: Piliostigma thonningii (CTFw 27027), RLS, vessel-ray pits elongated and simple. - 13: Berliniallsoberlinia type, lsoberlinia doka (CTFw 19972), TS, paratracheal aliform to confluent parenchyma and marginal bands. - Scale bars = I mm in Fig. 9, 13; 250 um in Fig. 10; 50 um in Fig. 11, 12.
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Fig. 14-17. - 14 & 15: Cassia sieberiana (RBHw 8056 ex CTFw 19960). - 14: TS, paratracheal confluent parenchyrna. - 15: RLS, septate fibres. - 16 & 17: Cynometra vogelii (CTFw 6608) . 16: TS, banded parenchyrna. - 17: RLS, crystals in non-ehambered ray cells . - Scale bars = I rnrn in Fig . 14, 16; 250 /lI1I in Fig . 15, 17.
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Cynometra vogelii Hook. f. - Fig. 16 & 17 Description based on 1 specimen. Growth ring boundaries indistinct, marked by interrupted marginal parenchyma bands, by radially flattened fibres, and accompanied by vessels or vessel groups. Vessels diffuse, 13 per mrn-, solitary (70%), in radial multiples of2-4 and rarely in clusters, vessel grouping index 1.4. Tangential diameter 110 um, maximum 150 um, Vessel element length 340 um. Intervessel pits vestured, 4 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders, similar to intervessel pits. Locally brown, gum-like deposits . Fibres thick-walled, gelatinous layers often present. Axial parenchyma in discontinuous and irregular bands, 3-7 cells wide, in strands of 2-4 cells, next to vessels up to 7. Parenchyma sheaths sometimes irregular to al most unilateral. Rays 9 per mm, 3 or 4 cells wide, heterocellular, square and upright cells in one, rarely up to 3 marginal rows. Large rays about 600 um high, maximum 970 um. Crystals solitary, prismatic, located in square/upright or procumbent, non-cham bered ray cells, very rarely in non-ehambered cells ofaxial parenchyma.
Daniellia oliveri (Rolfe) Huteh . & Dalziel- Fig. 18 & 19 Description based on 4 specimens. Growth ring boundaries indistinct or absent. Vessels diffuse, 2 or 3 per mm-, solitary (60-80%), in radial multiples of 2-4 and in clusters, vessel grouping index 1.2-1.4. Tangential diameter 140-210 um, maxi mum 330 um,Vessel element length 330-460 um. Intervessel pits vestured, 8-10 um in diameter, apertures not or faintly coalescent. Vessel-ray pits with distinct borders, similar to intervessel pits. Rarely thin-walled tyloses and locally brown, gum-like deposits. Fibres medium thick-walled, with a distinct pattern due to regular intrusive growth, gelatinous layers often present. Axial parenchyma paratracheal aliform, rarely vasicentric or shortly (2 or 3 ves sels/vessel groups) confluent, in strands of2-4 cells, next to vessels up to 6. Banded parenchyma in connection with axial intercellular canals . Rays 6-8 per mm, 3-6(-8) cells wide, heterocellular, square and upright cells in one, rarely up to 3 marginal rows. Rays 300-500 um high, maximum 1.3 mm. Storied structure regular, 2 or 3 ray tiers per axial mm. Low rays, axial parenchy ma, vessel elements, and fibres storied, large rays 2-3(-4) tiers high. Intercellular canals axial, diffusely arranged and in short to long tangentiallines. Lumina ofaxial canals often smaller than vessellumina. Bands of surrounding paren chyma 5-10 cells wide. Crystals solitary, prismatic, few located in chambered axial parenchyma cells, rarely in chambered ray cells, sometimes missing altogether.
Note: Lebacq (1957) has described the wood of Daniellia oliveri.
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Fig. 18-21. - 18 & 19: Daniellia oliveri. - 18: D. oliveri (SFB 1188.1), TS, paratracheal alifonn parenchyma, heterocellular rays, axial canals diffuse and in bands. - 19: D. oliveri (SFB 915.1), TLS, storied heterocellular rays. - 20: Detarium type, Detarium microcarpum (RBHw 17411 ex CTFw 27041), TS, paratracheal alifonn parenchyma, axial canals in bands, radial fibre pattern. 21: Dichrostachys cinerea (CTFw 20966), TS, paratracheal vasicentric parenchyma, sometimes unilateral, and marginal bands. - Scale bars = I mm.
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Detarium type - Fig. 20 Description based on 10 specimens, 4 of Detarium microcarpum Guill. & Perr. and 6 of Detarium senegalense J.F. Gmel. Growth ring boundaries indistinct, marked by interrupted marginal parenchyma bands, and accompanied by small vessels and vessel groups. Vessels diffuse, 2-15 per mm-, solitary (30-90%), in radial multiples of 2-4 and in clusters, sporadically groups of more than 10 vessels (especially in D. microcar pum), vessel grouping index 1.1-2.4. Tangential diameter 90-260 um, maximum 360 um, Vessel element length 230-350 um. Intervessel pits vestured, 7-10 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders, similar to inter vessel pits. Rarely thin-walled tyloses and locally brown, gum-like deposits. Fibres medium thick-walled, in a distinct pattern due to regular intrusive growth, gelatinous layers often present. Axial parenchyma paratracheal aliforrn to weakly confluent (2-5 vessels/vessel groups) (to banded in D. senegalense SFB 440.1), in strands of 2-4 cells, next to vessels up to 8. Little apotracheal diffuse parenchyma, often crystalliferous. Bands in connection with axial intercellular canals. Rays 5-7 per mm, 3-5(-6) cells wide, homocellular to heterocellular with one irregular row of square to upright cells (higher portion of and more distinct heterocel lular rays in D. senegalense). Large rays 400-1000 umhigh, maximum 1.7 mm. Intercellular canals axial (missing in D. senegalense CTFw 6606 and RBHw 12714), in long tangentiallines, lumina often up to the size ofvessellumina. Bands of surrounding parenchyma 5-15 cells wide. Crystals solitary, prismatic, located in chambered axial parenchyma cells, very rarely in chambered ray cells. Note: The wood anatomy of Detarium senegalense is described by Normand (1950) . and Lebacq (1957).
Dialium guineense Willd. - Fig. 22 & 23 Description based on 3 specimens . Growth ring boundaries absent or indistinct, marked by interrupted and irregular marginal parenchyma bands, 1 or 2 cells wide. Vessels diffuse, 5-12 per mm- , solitary (70-80%), in radial multiples of2-4 and in clusters, vessel grouping index 1.2-1.3. Tangential diameter 100-150 um, maxi mum 190 um. Vessel element length 280-300 um. Intervessel pits not vestured, 8-9 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders, simi lar to intervessel pits. Locally brown, gum-like deposits . Fibres thick-walled, gelatinous layers often present. Axial parenchyma discontinuously and irregularly banded, 2-3(-5) cells wide, in strands of 2-4 cells, next to vessels up to 6. Diffuse apotracheal parenchyma scarce, often crystalliferous. Rays 9-11 per mrn, 2-3(-4) cells wide, homocellular, composed ofprocumbent cells. Rays about 250 um high, maximum 700 um.
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Fig. 22-25. - 22 & 23: Dialium guineense (SFB 482.1). - 22: TS, parenchyrna in narrow bands. 23: TLS , silica grains in axial parenchyrna. - 24 & 25: Entada type , Entada abyssinica (SFB 1075.0). - 24: TS, banded parenchyrna, fibre band poor in parenchyrna. - 25: TLS, septate fibres. - Scale bars = I rnrn in Fig. 22, 24; 250 J.UIl in Fig. 23, 25.
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Storied structure regular, 4 ray tiers per axial mm. Low rays, axial parenchyma, vessel elements, and fibres storied, large rays 2 or 3 tiers high . Silica as grains in (almost) all cells ofthe axial parenchyma. Crystals few, solitary, prismatic, located in chambered axial parenchyma cells. Note : Dialium guineense is included in Normand's (1950) description of the genus.
Dichrostachys cinerea (L.) Wight & Am. - Fig . 21 Description based on 4 specimens. Growth ring boundaries indistinct, marked by interrupted marginal parenchyma bands, sporadically crystalliferous, and accompanied by small vessels and vessel groups. Vessels diffuse, 6-20 per mm-, solitary (40-90%), in radial multiples of2-4 and rarely in clusters, vessel grouping index 1.1-2.0. Tangential diameter 110-140 J1Ill, maximum 250 J1Ill . Vessel element length 250-260 J1Ill . Intervessel pits vestured, 6-7 J1Ill in diameter, apertures coalescent. Vessel-ray pits with distinct borders, simi lar to intervessel pits. Locally brown, gum-like deposits. Fibres medium thick-walled to thick-walled, gelatinous layers often present. Axial parenchyma paratracheal, vasicentric to aliforrn and confluent, in strands of 2-4 cells. Parenchyma sheaths sometimes irregular to unilateral. Little apotracheal diffuse parenchyma, sporadically crystalliferous. Rays 6 per mm, 2-5(-8) cells wide, homocellular, composed of procumbent cells. Large rays 400-650 um high, maximum 1.6 mm. Crystals solitary, prismatic, located in axial chambered parenchyma cells .
Entada type - Fig. 24 & 25 Description based on 6 specimens, 4 of Entada abyssinica A. Rich . and 2 of Entada africana Guill. & Perr. Growth ring boundaries absent or inconspicuous, marked by fibre bands poor in parenchyma. Vessels diffuse, 1 or 2 per mm-, solitary (40-90%), in radial multiples of2-4 and rarely in clusters, vessel grouping index 1.1-1.9. Tangential diameter 190-250 J1Ill, maximum 420 J1Ill . Vessel element length 250-320 J1Ill. Intervessel pits vestured, 9 J1Ill in diameter, apertures coalescent. Vessel-ray pits with distinct borders, similar to intervessel pits . Locally brown, gum-like deposits. Fibres medium thick-walled to thick-walled, sporadically septate, gelatinous lay ers often present. Axial parenchyma paratracheal (aliforrn to) confluent and banded, bands discon tinuous, 3-10(-15) cells wide . Narrow vasicentric or narrow aliform parenchyma sheaths within zones poor in parenchyma. Parenchyma in strands of 2-4 cells , occa sionally fusiform. Rays 4 or 5 per mm, 3-5(-8) cells wide , homocellular, composed of procumbent cells. Large rays 600-1000 um high, maximum 2.4 mm. Crystals, solitary, prismatic, located in chambered axial parenchyma cells, con spicuous at the margins of aliforrn parenchyma.
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Fig. 26-29. - 26 & 27: Faidherbia albida (SFB 1195.1). - 26: TS, parenchyma in broad bands. 27: TLS, storied structure, uniseriate rays. - 28 & 29: Gilletiodendron glandul osum (CTFw 4932). - 28: TS, scarce parenchyma, paratracheal scanty to aliform, sheath sometimes incomplete, growth ring boundaries marked by thick-walled fibres. - 29: RLS, heterocellular rays. - Scale bars = I mm in Fig. 26-28; 250 J.I111 in Fig. 29.
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Faidherbia albida (DeI.) A. Chev. - Fig. 26 & 27 Description based on 3 specimens. Growth ring boundaries distinct, marked by marginal parenchyma bands, 1-3(-4) cells wide, occasionally to continuously crystalliferous. Vessels diffuse, 3-7 per mm-, solitary (60-70%), in radial multiples of 2-4 and rarely in clusters, vessel grouping index 1.3-1.4. Tangential diameter 200-230 um, maximum 350 um, narrow vessels very numerous. Vessel element length 190-240 um. Intervessel pits vestured, 9-11 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders, similar to intervessel pits. Fibres medium thick-walled, gelatinous layers often present. Axial parenchyma in discontinuous and irregular bands, 8-20(-30) cells wide, often wider than fibre zones in between . Parenchyma in strands of 2(-4) cells. Rays 15 or 16 per mm, uniseriate, with only small portions up to 2 cells wide, homocellular, composed of procumbent cells. Rays 100-250 um high, maximum 750~ . Storied structure regular, 5-7 ray tiers per axial mm. Low rays, axial parenchyma, vessel elements, and fibres storied, large rays 2-3(-4) tiers high. Crystals, solitary, prismatic, located in chambered axial parenchyma cells, con spicuous at the margins of aliform parenchyma.
Note: Descriptions of Faidherbia albida are given by Lebacq (1957), Ciuffi Cellai (1967), Cutler (1969), Fahn et al. (1986) and Jagiella & Kürschner (1987). Lebacq (1957) probably examined a misidentified sampie, for the described ray structure does not match Faidherbia albida (Cutler 1969). Neumann (1989) and Rolando (1992) describe charred wood of the species.
Gilletiodendron glandulosum (Porteres) J. Leonard - Fig. 28 & 29 Description based on 1 specimen. Growth ring boundaries indistinct, marked by radially flattened, thick-walled fibres, and accompanied by small vessels and vessel groups. Vessels diffuse, 50 per mm-, solitary (70%), in radial multiples of 2-4 and rarely in clusters, vessel grouping index 1.3. Tangential diameter 60 um, maximum 80 um. Vessel element length 330 um. Intervessel pits vestured, 5 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders, similar to intervessel pits. Locally brown, gum-like deposits . Fibres medium thick-walled. Axial parenchyma scarce, paratracheal, scanty to aliform, in strands of 2-4 cells, next to vessels up to 6. Parenchyma sheaths narrow and often incomplete. Rays 13 per mm, exclusively uniseriate, mostly heterocellular with one marginal row of square and upright cells, few homocellular. Large rays about 350 um high, maximum 620 um. Crystals solitary, prismatic, located in chambered axial parenchyma cells.
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Note: The wood anatomy ofthe same sample (CTFw 4932) is described by Nonnand & Chatelet (1955). They consider the rays as being homocellular.
Guibourtia copallijera Benn. - Fig. 30 & 31 Description based on 1 specimen. Growth ring boundaries indistinct, marked by radially flattened and thick-walled fibres, scattered parenchyma cells, and accompanied by small vessels and vessel groups. Vessels diffuse, 18 per mm-, solitary (70%), in radial multiples of 2-4 and rarely in clusters, vessel grouping index 1.2. Tangential diameter 80 um, maximum 110 um. Vessel element length 380 um. Intervessel pits vestured, 7 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders, similar to intervessel pits. Locally brown, gum-like deposits . Fibres medium thick-walled to thick-walled. Axial parenchyma scarce, paratracheal, scanty to aliform, in strands of 2-4 cells. Parenchyma sheaths narrow and often incomplete. Rays 8 per mm, 2-3(-4) cells wide, heterocellular, with 1 or 2 marginal rows of square and upright cells. Large rays about 600 um high, maximum 1 mm. Crystals solitary, prismatic, located in chambered and not chambered axial paren chyma cells.
Parkia biglobosa (Jacq.) G. Don - Fig. 32 & 33 Description based on 4 specimens. Growth ring boundaries indistinct to distinct, marked by marginal parenchyma bands, 1-3(-5) cells wide, sporadically crystalliferous, and accompanied by small vessels and vessel groups. Vessels diffuse, 2-7 per mm-, solitary (60-90%), in radial multiples of 2-4 and rarely in clusters, vessel grouping index 1.1-1.5. Tangential diameter (100-)130 190 um, maximum 290 um, Vessel element length 170-260 um. Intervessel pits ves tured, 8-10 um in diameter, apertures coalescent. Vessel-ray pits with distinct bor ders, similar to intervessel pits. Locally brown, gum-like deposits . Fibres medium thick-walled, occasionally with a distinct pattern due to regular intrusive growth, gelatinous layers often present. Axial parenchyma paratracheal aliform to confluent and in discontinuous and irregular bands, 4-10(-20) cells wide, in strands of 2-4 cells, next to vessels up to 6. Little apotracheal parenchyma, often crystalliferous. Rays 6-10 per mm, 3-5(-6) cells wide, homocellular, composed ofprocumbent cells. Large rays 150-350 um high, maximum 560 um, Crystals solitary, prismatic, located in chambered axial parenchyma cells, occa sionally in chambered procumbent ray cells.
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Fig. 30-33. - 30 & 31: Guibourtia copallifera (CTFw 7603) . - 30: TS, parenchyma scarce, para tracheal scanty to aliform, sheath sometimes incomplete, growth ring boundaries marked by thick walled fibres. - 31: RLS, heterocellular rays . - 32 & 33: Parkia biglobosa. - 32: P. biglobosa (SFB 848.0), TS, confluent parenchyma and marginal bands . - 33: P. biglobosa (SFB 1174.1). TLS . 4- or 5-seriate rays. - Scale bars = I mm.
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Fig. 34 & 35: Prosopis africana . - 34: P. africana (RBHw 17406 ex CTFw 27026), TS, vasicentric to alifonn parenchyrna. - 35: P. africana (RBHw 11524), TLS, septate fibres. - Scale bars =1 rnrn in Fig. 34, 100 um in Fig. 35.
Prosopis africana (Guill. & Perr.) Taub. - Fig. 34 & 35 Description based on 5 specimens. Growth ring boundaries indistinct, marked by radially flattened and thick-walled fibres, in places with parenchyma cells, and accompanied by small vessels and vessel groups. Vessels diffuse, 5-7 per mm-, solitary (70-80%), in radial multiples of 2-4 and rarely in clusters, vessel grouping index 1.2-1.3 . Tangential diameter 140-180 um, maximum 280 um. Vessel element length 180-300 um, Intervessel pits vestured, 7-9 um in diameter, apertures coalescent. Vessel-ray pits with distinct borders, simi lar to intervessel pits. Locally red-brown, gum-like deposits. Fibres medium thick-walled to thick-walled, some septate (missing in some speci mens), gelatinous layers often present, mostly remarkably thick. Axial parenchyma paratracheal, vasicentric to aliform and shortly (2 or 3 vessels/ vessel groups) confluent, in strands of 2-4 cells. Parenchyma sheath sometimes irregular to unilateral. Rays 6-8 per mm, 2-3(-5) cells wide, homocellular, composed of procumbent cells . Large rays 200-250 um high, maximum 740~ . Crystals solitary, prismatic, located in chambered axial parenchyma cells.
Note : Gottwald & Noack (1966) also describe few septate fibres for the species .
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DISCUSSION
Table I summarizes the results of the survey. It shows that some of the examined species have a unique combination of features and can easily be differentiated. These weIl delimited types are placed to the left of the table. To the right of the table follow types of species or even genera that are only weakly delimited. The lack of differ entiating features places the Caesalpinioideae /Mimosoideae type at the end of the table.
Caesalpinioideae/Mimosoideae type This type, which combines four species of both subfamilies, demonstrates the dif ferent evaluation of characters in identification of archaeological, usually smaIl, wood fragments in contrast to taxonomy. TaxonomicaIly, Papilionoideae and Caesalpinioi deae are closely linked via wood anatomical features and intergrade, while Mimosoi deae are a distinct group (Baretta-Kuipers 1981). Heterocellular rays, für instance, are shared on1y by Caesalpinioideae and Papilionoideae (Baretta-Kuipers 1981). In identifying Sudanian leguminous woods the Papi1ionoideaecan easily be distinguished from the taxonomically closely linked caesalpinioid taxa. On the other hand, some of the mimosoid and caesalpinioid taxa cannot be separated from each other, and conse quently the joint Caesalpinioideae/Mimosoideae type was recognized. The taxa of this type are characterized by typical features of Mimosoideae, as lack of storeying and homocellular rays (Baretta-Kuipers 1981), that of course are distributed among caesalpinioid taxa as weIl. In identifying archaeological sampies from the area, the Caesalpinioideae/Mimosoideae type also functions as a 'garbage bin' for those frag ments that can be identified as leguminous and non-papilionoid, which, however, do not allow further determination due to the lack of differentiating features.
Weil delimited types The Faidherbia albida, Dialium guineense, Bauhinia/Piliostigma, Daniellia oliveri, Detarium, and Albizia types differ distinctly from the Caesalpinioideae /Mimosoideae type. Within the limitations of single sampies, the Cynometra vogelii, Guibourtia copallifera and Gilletiodendron glandulosum types are weIl delimited, too. All of these types are characterized by qualitative features such as storied struc ture, heterocellular rays, axial canals, silica, septate fibres and crystals in non-cham bered ray cells. These features are most useful for the delimitation of the types for several reasons . First, they are exceptions to common combinations within the group in question . They distinguish the wood types from the basic Caesalpinioideae/Mimos oideae type. Furthermore, the qualitative features are not variable; their development is not, or only slightly, influenced by external factors (Carlquist 1988; Iqbal & Ghouse 1990; Lev-Yadun & Aloni 1995). Moreover, they are clearly marked and their pres ence or absence alone is decisive for identification purposes . Some of the above fea tures are also used in the keys for leguminous woods by Quirk (1983), Watson & Dallwitz (1983) and Detienne & Ter Welle (1989). However, these qualitative fea tures may still cause problems as far as identification of unknown fragments is con cerned, as will be discussed below.
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~f;x (>l ( 1 I >< 1 >< Faidherbia albida - Acacieae >< I 1 1 1 I" 1 I >< " Diali um guineense . Cassieae ~ >< I 1 I ~ >< >< ~ Bauhinia rufescen s - Cercideae 1 "" " ~ a~ ~.§.. "I 1 1 I >< >< Pilostigma reticulatum - Cercideae 3: s " """" ~ Cl " '< ~ " 8 " >< " >< " >< Pilestigma thonningii - Cercideae il 18 " " ~~ 8 "" I" " Daniellia olive r; - Detarieae (:l f8 " "" 8 " ~ Detarium microcarpum • Detarieae -<~ ~ ~ , """ 8 " ~ Detarium senegolense - Detariea e 3 8 x " " " Cynometra vogelii - Detarieae >< " " " Guibourtia copollifera - Detarieae >< " " Gilletiodendron glandulosum - Detar. 8 Berlinia grandiflora - Amherstieae ~ T8 "" ~~ 8 " " 8 lsoberlinia doka - Amherstieae ...~~ -<~ 8 " " ~ Isoberlinia tomentosa - Amherstieae il ~ ~ Albfzia chevalieri - Ingeae ,. " " .:
"I 1 1 1 I 1 I >< I I 1 >< III Afzelia africana - Detarieae I ~ ~ .:< ~ ~ ]1:: '~ i l "" Tamarindu s indica - Amherstieae ~- 8 1 "" Parkia biglobo sa - Parkieae """"""" Aea cia seyal - Acacieae ~2 Acacia siebe riana - Acacieae ""8 """ "& B' 8 8 """ Acacia senegal - Acacieae " I" " 8 " 8 Entada africana - Mimoseae 1.- g. I ~01!-~ >< I>< " ~ " 8 Entada abyssinica - Mimoseae 8 1 8 Amb /ygonocarpus andongen sis - Mim, ~ Q I I I" 3 g >< Erythroph/eum suaveo/ ens - Caesalpin, ~e. 8 ~~ ' g §: ~ Erythroph/eum africanu m - Caesalpin. ';g- "0 0 o ~ Burkea afncana - Caesalp meae 1
6661 '(Z:) OZ: '\01\ ' \UUlnOf YAW l 8n Höhn - Wood of West African Leguminosae 139
Storeying of elements is characteristic for some of the examined species only (Table 1). Since the radial-longitudinal divisions that lead to storeying ofthe cambium are genetically fixed and independent of exogenous factors (Iqbal & Ghouse 1990), it is also very reliable. Nevertheless, some intra-specific variability occurs. For this reason, it does not seem possible to use the regularity of storeying as a differentiating feature. A difference in regularity for Bauhinia and Piliostigma (Baretta-Kuipers 1981), for instance, cannot be confirmed . Bauhinia does show irregular storeying, but while Piliostigma thonningii has regular tiers, P. reticulatum does exhibit regular as well as irregular storeying. Heterocellular rays were, in accordance with Baretta-Kuipers' (1981) results, found in Caesalpinioideae only (Table 1). Though this is an important feature for the separation of the types, it sometimes proves to be problematic to evaluate whether rays are hetero- or homocellular in the small, often poorly preserved frag ments, obtained from archaeological excavations. Procumbent cells are often higher and shorter at the margins of rays or where they cross axial parenchyma. This might lead to the impression of heterocellularity where there is none. On the other hand, the apparently heterocellular rays of Gilletiodendron glandulosum have been described as homocellular by Normand & Chatelet (1955) . Such differences in evaluating ray composition may especially arise when the heterocellular rays are few or not distinct, as is the case with the Detarium and BerlinialIsoberlinia types. Septate fibres occur in five types. But they are a distinct and typical feature of the Albizia type only, which is delimited also by the occurrence of distinct fusiform, diffuse apotracheal paren chyma (Fig. 7, 8). In other types, e.g. the BauhinialPiliostigma type, septate fibres can be few and hardly discernible, especially within thick-walled or gelatinous fibres. They are very inconspicuous in B. rufescens and cannot be excluded for the examined Piliostigma spp. On the other hand, detached fragments of wall material, in particular within a ground-rnass of gelatinous fibres, may give the impression of septa in spe eies with non-septate fibres. Non-vestured pits and vessel-ray pits with reduced bor ders, silica, axial canals and crystals in ray cells characterize only one or two types each (Table 1). The lack of vestures can be well recognized in bleached slides. In charred fragments, however, which cannot be bleached, other incrustations may mimic vestured pits. Vessel-ray pitting with reduced or missing borders is easily discerned, especially when combined with an enlargement of the pits. The unique occurrence of silica in axial parenchyma marks Dialium guineense (Fig. 22, 23). Since there are few Leguminosae with silica and Dialium is the only genus with silica in axial paren chyma (Koeppen 1980), this character allows for easy delimitation of the type and identification of the taxon. Unfortunately silica grains are difficult to discern in charr ed woods. Axial canals are a reliable feature in Daniellia oliveri, but they may be missing in a given fragment of the Detarium type. Moreover, it may be difficult to distinguish axial canals from vessels in charred and distorted fragments.
Weakly delimited types The BerlinialIsoberlinia, Cassia sieberiana, Prosopis africana, Dichrostachys ein erea, AfzelialTamarindus, Parkia biglobosa, Acacia and Entada types are not very well separated from the Mimosoideae/Caesalpinioideae type or from one another,
Downloaded from Brill.com10/06/2021 10:09:34PM via free access 140 IAWA Journal, Val. 20 (2), 1999 because the features discussed above are either missing or not distinctly developed. In order to distinguish between these types, fewer and less reliable feature s are recog nized . Features such as parenchyma distribution, ray width and height are more vari able. Consequently, the delimitation is weaker. The chance that characters are not developed typically in a given fragment is much higher with these types. Erroneous identification may result. Cassia sieberiana, Prosopis africana, Entada, and BerlinialIsoberlinia types are only weakl y delimited, even though they possess the qualitative feature s of septate fibres, respectively heterocellular rays. However, the features are indistinct and/or developed in a few sampIes only. The additional and unique feature of the Berlinial Isoberlinia type, unilaterally compound pits, is too rare and too inconspicuous to be considered as reliable for identification purposes. Parenchyma distribution is crucial for the distinction of the weakly delimited types. Müller-Stoll & Mädel (1967 ) even count it as the most conspicuous feature for iden tifying Leguminosae. While it is helpful for the taxonomie distinction of tribes and genera (Baretta-Kuipers 1981), as far as reliability for identification is concerned, its variability may lead to erroneous identifications. Ray width too, is a variable but neces sary quantitative feature for the distinction between the examined species. Extreme values such as uniseriate or extremely wide rays are reliable in mature wood . However, variability and overlapping ranges do not allow a secure delimitation of types. And , as was the case before, while differences in ray width may be interesting in taxonomie contexts, they are often not big enough to allow for aseparation of unknown fragments. The example of Berlinia and Isober linia shows how variability leads to different conclusions. Lebacq (1957 ) ascribes uniseri ate rays to Berlinia grandiflo ra and mostly biseriate rays to Isoberlinia tomentosa. The examined material confirms the tendency towards a higher portion of biseriate rays with both Isoberlinia species. However, Baretta-Kuipers (1981) dif ferentiates between Isoberlinia with exclusively or predominantly uniseriate rays, and Berlinia with occasionally biseriate rays. Quantitative characters of the vessels, such as diameter and density, may be of assistance for the diagnosis of familie s, if they are extremely small or extremely large. Within the examined taxa, the means are too close together to allow identification. Vessel density and vessel diameter as weil as maximum height of rays are quite vari able and most often show some deviation from , or do not match, other published descriptions of the examined taxa.
Brauch wood Since differences in wood structure between mature and juvenile wood were ob served, the branch wood specimens were not considered for the recognition of the types. Characters that are reliable in mature wood - storied structure and ray struc ture - are not weIl developed yet. Due to the fact that storied structure only develops with the radial divisions of the cambial cells (Iqbal & Ghou se 1990), storeying is in distinct in the areas adjacent to the pith. Ray composition may differ as well between specimens from close to the pith and from mature wood . For example, next to the pith, the rays of all branch wood specimens from Bauhinia rufescens consist almost
Downloaded from Brill.com10/06/2021 10:09:34PM via free access Höhn - Wood of West African Legumin osae 141 exclusively of upright cells, gradually changing to rays of mixed composition, and reaching heterocellular composition only with some millimetres distance to the pith. Within the millimetre next to the pith, both juvenile specimens of Burkea africana show heterocellular rays as opposed to homocellular rays in mature wood. Tamarindus indica, Cassia sieberiana and Prosopis africana show heterocellu larity in their juve nile wood as weIl. On the other hand Dos Santos & Miller (1997) did not find significant differences in ray structure between twig and stern sampies. However, the above men tioned differences are limited to a small area of the specimen. With small fragments of twigs, as found in archaeological contexts, these differences, nonetheless, may lead to erroneous identifications.
Typical for all specimens of juvenile wood (Fig. 36, 37) is a higher variability in parenchyma distribution within a growth zone and from one to another. This high range of variation increases the difficulties of identifying branch wood, especially if the taxa lack other qualitative features such as septate fibres or axial canals . Quantita tive feature s differ from mature wood as weIl. Vessels are smaller and often more numerous in the branch wood specimens. A higher percentage of vessels in branches in comparison to stern and root material is also reported by Stokke & Manwiller (1994). Conceming ray width and ray height no clear tendencies could be detected from the material. Fahn et al. (1986) report higher rays close to the pith. Inthe exam ined specimens, however, both mean width and height, fall within the ranges set by
Fig. 36 & 37: Branch wood. - 36: Acacia sieberiana (SFB 754.0), parenchyrna variation between different growth zones. - 37: Parkia biglobosa (SFB 1183.2), parenchyrna distribution close to the pith. - Scale bars = 1 rnrn.
Downloaded from Brill.com10/06/2021 10:09:34PM via free access 142 IAWA Journal, Vol. 20 (2), 1999 the mature material, only maximum height is higher in a few specimens. On the other hand, some specimens of branch wood show rays that are more narrow and less tall than in the stern wood material. Such an increase in length with distance from the pith is reported by Iqbal & Ghouse (1990) and Lev-Yadun & Aloni (1995).
CONCLUSION
This survey has shown the possibilities and limits for identification of leguminous woods from the Sudanian savannas ofWestAfrica. The geographicallimitation was a prerequisite for this survey and it has to be kept in mind that additional taxa may belong to the recognized types. A significant number of types are weakly differenti ated from one another and the collective Caesalpinioideae/Mimosoideae type. Due to variation in parenchyma distribution, in width or height of rays, it might not be pos sible to recognize these types correctly or at all, when identifying unknown frag ments. Therefore, working on the identification of archaeological specimens of legu minous woods requires caution and restraint. At times one is forced to conclude that identifying a certain fragment beyond the taxonomie level of genus, tribe, subfamiliy or even family is impossible. On the other hand, there are a number of taxa with reliable qualitative characters, such as Faidherbia albida, Dialium guineense or Detarium spp., which facilitate easy and reliable identification. Moreover, the appealing wood anatomy of many Leguminosae pays off whatever difficulties they may impose on the wood anatomist.
ACKNOWLEDGEMENTS
This work was supported by the Special Research Project "History of Culture and Language in the Natural Environment of the West African Savanna" funded by the German Research Foundation (DFG). I am indebted to Dr. K. Neumann for suggesting and facilitating the survey, for fruitful dis cussions, lots of freedom and valuable comments on the manuscript, as weil as to Dr. P. Baas and the reviewers for critical remarks.The technical assistance of Ms. G. Schenk in preparing the slides, and of Mr. M. Ruppel in making SEM observations is greatly appreciated. Thanks are due to Mr. A. Jansen who critically reviewed the English text. The survey would not have been possible with out the sampies generously supplied by Dr. H.G. Richter of RBHw and Dr. P. Detienne of CTFw.
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APPENDIX
Mature wood material studied: Acacia senegal (L.) Willd ., Mimosoideae: K-Jw 2498 ; Senegal, RBHw 17393; Senegal, RBHw 17394 ; Sudan, Nubia, Schweinfurth s.n., SFB 116.0 ex Bw, 100 mm diam. -Acacia seyal Del ., Mimosoideae: Senegal, Mariaux 1258, CTFw 27058 ; Senegal, RBHw 17412 ; Sudan, Jebel Marra, Miehe s.n., RBHw 18062 ,80 mm diarn. ; Sudan, Wadi Karnoi, Neumann 25, SFB 117.1; Sudan, Forest Department Karthoum, SFB 245.0.- Acacia sieberiana DC., Mimosoi deae: Sudan, RBHw 10674 ; Sudan, Hawata, RBHw 10676 ; Senegal, RBHw 17395 ; Sudan, Khartoum, Soba-Arboreturn, SFB 119.0; Egypt, Schweinfurth s.n., SFB 594 .0 ex Bw, 80 mm. - Afzelia africana Pers ., Caesalpinioideae: Senegal, Mariaux 1204, CTFw 19960 ; Cöte d'Ivoire, Mariaux s.n., RBHw 4700 ex CTFw 20484; Senegal, Kolda, Mariaux & Diaite 1234, RBHw 17396 ex CTFw 27034; Sudan, Gelte, RBHw 11519; Sudan, Gelte, RBHw 11520; Burkina Faso, Diapaga, Küppers 1087, SFB 1171.0. -Albizia chevalieri Harms, Mimosoideae: Nigeria, FHI Nr. 15695, K-Jw 7859 .-Albizia cf. malacophylla (A. Rich.) Walp., Mimosoideae: Burkina Faso, Logoniegue, Neumann 981 , SFB 571 .1, 55 mm diam. - Amblygonocarpus andongensis (Oliv.) Exell & Torre , Mimo soideae: Angola, Dechamps 1454, CTFw 21725 ; Zirnbabwe, RBHw 9836 ; Mozambique, RBHw 10632 ; Mozambique, RBHw 11363; Mozam bique, RBHw 14578. - Bauhinia rufescens Lam., Caesalpinioideae: Senegal, Bakel Mouderi, Mariaux & Diaite 1266, RBHw 17397 ex CTFw 27066; Sudan, Wadi Karnoi, Neumann 54, SFB 130.1. - Berlinia grandijlora (Vahl) Huteh . & Dalziel , Caesalpinioideae: Liberia, Cooper 217, RBHw 16714ex MAD-SJRw 15144;Togo, Kersting 33, RBHw 6759; Congo, Bas Kouilou, Sargos 873, RBHw 8017; Congo, Bas Kouilou, Sargos 874, RBHw 8079 ; Benin, Cascades de Tanougou, Neumann 1489, SFB 1229.1. - Burkea africana Hook., Caesalpinioideae: Togo,
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Kersting 8, RBHw 18128 ; Nigeria, Forest Herbarium Ibadan 47986, Tw 14493; Cöted'lvoire, Pare National de 1a Cornoe, Neumann 1042, SFB 274 .1; Burkina Faso, To, Neumann 1472 , SFB 1185.1,200 mm diam.- Cassia sieberiana De., Caesalpinioideae: Congo, Bas Koui1ou, Sargos 917 , RBHw 8056 ex CTFw 19960; Senegal, Tambaeounda ouest, Mariaux & Diaite 1240 , RBHw 17398 ex CTFw 27040; Burkina Faso, Logoniegue, Neumann 982 , SFB 551.1. - Cynometra vogelii Hook. f., Cae salpinioideae: Guinea, Pobeguin 858 , CTFw 6608. Daniellia oliveri (Ro1fe) Huteh. & Dalziel, Cae salpinioideae: Cöte d'lvoire, CTFw 6693 ; Su dan, Gette, RBHw 11525 ; Burk ina Faso, Niangoloko, Rohde 12, SFB 915.1 , 160 mm diam.; Burkina Faso, To, Neumann 1479, SFB 1188.1 , 120 mm diam. - Detarium microcarpum Guill. & PeIT., Caesalpinioideae: Senegal, Tambacounda oue st, Mariaux 1241, RBHw 17411 ex CTFw 27041 ; Togo, Kersting 13, RBHw 6740; Burkina Faso, Niangoloko, Rohde 27, SFB 544 .1,50 mm diam.; Burkina Faso, Fada N'Gourma, Neumann 564 , SFB 1054.0,45 mm diam . -Detarium senegalense J.F. Gmel., Caesalpinioideae: Guinea, Pobeguin 689 , CTFw 6606 ; Liberia, Cooper-Record s. n., RBHw 12714 ; C öte d'lvoire,Danzer s.n., RBHw 12968 ; Cöte d'lvoire, Danzer s.n., RBHw 12970 ; Nigeria, Forest Herbarium Ibadan 22983 , SFB 331.0; C öte d'lvoire, Parc National de la Comoe, Neumann 1070, SFB 440 .1. - Dialium guineense Willd ., Caesalpinioideae: Guinea, Pobeguin 584, CTFw 6603; Congo, Bas Kouilou, Sargos 894 , RBHw 8036; Cöte d'Ivoire, Parc National de la Comoe, Neumann 1036, SFB 482 .1, 220 mm diam.-Dichrostachys cinerea (L.) Wight &Arn., Mimosoideae: C öte d'lvoire, Detienne 240, CTFw 20966; Senegal, Niayes Mboro, Mariaux & Diaite 1278, RBHw 17401 ex CTFw 27078; EastAfrica, Schlieben 3082, RBHw 1619; Togo , Kersting 100, RBHw 6808 . -Entada abyssinica A.Rich., Mimosoideae: Angola, Dechamps 1423 , CTFw 21496; East Africa, Schlieben 3186 , RBHw 1621,65 mm diam .; Burkina Faso, Banfora, Neumann 602, SFB 1074.0, 43 mm diam .; Burkina Faso, Tiefora, Neumann 609 , SFB 1075.0, 65 mm diam. - Entada ajricana Guill. & PeIT., Mim osoideae: Senegal, Boundou, Koussan, Mariaux 1257 , RBHw 17 410 ex CTFw 27057; Burkina Faso, To, Neumann 1474 , SFB 1186.1, 120 mm diam. Erythrophleum ajricanum (Benth.) Harrn s, Caesalpinioideae: Togo , Leving ston 26, CTFw 18880 ; East Africa,Schlieben 5258, RBHw 1771; Mozambique, RBHw 10642 ; South Africa (cult.), RBHw 15167. -Erythrophleum suaveolens (Guill. & Perr.) Brenan, Caesalp inioideae: Cameroon, Hedin 1644, CTFw 3870; Liberia, Cooper L 17, 148386, RBHw 17937 ex MAD SJRw 13791 ; East Afriea, Schli eben 5528, RBHw 1884; C öte d'lvoire, RBHw 2233 ; Togo , Kersting 31, RBHw 6757 ; Cameroon, Mildbraed 3956, RBHw 17934 . - Faidherbia albida (DeI.) A. Chev., Mimosoideae: Sud an, RBHw 18061 ; Togo, Kersting s.n., SFB 105.0 ex Bw, 200 mm diam.; Burkina Faso, Dori , Neumann s.n., SFB 1195.1 . - Gilletiodendron glandulo sum (Porteres) J. Leonard, Cae salpinioideae: Senegal, Kita , Lucchini 0, CTFw 4932. Guibourtia copallijera Benn., Caesalpinioideae: Guinea, ENITEF 129, CTFw 7603. Isoberlinia doka Craib & Stapf, Caesalpinioideae: Mali , Service Forestier 211 ar, CTFw 19972; Sudan, Loka, RBHw 9864; Sudan, Loka, RBHw 9865 ; Sudan, Yei/Juba Road Khor Kimbi, RBHw 12022; Sudan, YeilJuba Road Khor Kimbi, RBHw 12023; Sudan, Yei/Juba Road Khor Kimbi, RBHw 12024; Burkina Faso, Boulon, Neumann 1011, SFB 211.0; Burkina Faso, To, Neumann 1472, SFB 1184.1,55 mm diam.- Isoberlinia tomentosa (Harms) Craib & Stapf, Caesalpinioideae: SouthAfrica (cult.), RBHw 15179 ex PFPw ; Burkina Faso, Boulon, Neumann 1010, SFB 190.0 ,280 mm diam. - Parkia biglobosa (Jacq .) G. Don, Mimosoideae: Guinea, Pobeguin 130, CTFw 6594; Senegal, Kolda, Mariaux & Diaite 1232, RBHw 17403 ex CTFw 27032; Burkina Faso, Fada N'Gourma,Neumann s.n., SFB 848.0;Nigeria, Biu, Salzmann 89, SFB 1174.1. - Piliostigma reticulatum (DC.) Hochst., Caesalpinioideae: Sudan, Baumer 64, CTFw 15547; EastAfrica, Schlieben s.n., RBHw 1539,60 mm diam.; Burkina Faso, Gorom Gorom, Maydell s.n., RBHw 16330 ; Burkina Faso, Dori , Neumann s. n., SFB 1193.1 , 140 mm diam. - Piliostigma thonningii (Schum.) Milne-Redh., Caesalpinioideae:Senegal, Ziguinchor, Mariaux & Diaite 1227 , RBHw 17405 ex CTFw 270 ; Senegal, Mariaux 1227, CTFw 27027;
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Mozambique, RBHw 10625; Angola, Murta 1075, RBHw 15213 ex Tw 28222; Burkina Faso, Logoniegue, Neumann s.n., SFB 869.1, 120 mm diam. - Prosopis africana (GuilI. & Perr.) Taub., Mimosoideae: Senegal, Ziguinchor, Mariaux & Diaite 1226, RBHw 17406 ex CTFw 27026 ; Burkina Faso, Croise 30, CTFw 29678 ; Sudan, Gette, RBHw 11523; Sudan, Gette, RBHw 11524; Nigeria, Konduga, Salzmann 88, SFB 1173.1. - Tamarindus indica L., Caesalpinioideae: Senegal , Mariaux 1252, CTFw 27052; Senegal, RBHw 8560; Togo, Kersting s.n., SFB 640.0, 200 mm diarn.; Burkina Faso, Logoniegue, Neumann s.n., SFB 865.1,80 mm diameter.
Branch wood material studied: Acacia senegal (L.) Willd., Mimosoideae: K-Jw; Sudan, Zalingei, Neumann 85, SFB 115.0, 20 mm diarn.; Sudan , EI Fasher, Neumann s.n. , SFB 822.0, 30 mm diam. -Acacia seyal DeI., Mimosoideae: K-Jw, 20 mm diarn.; Sudan, Wadi Karnoi, Neumann 25, SFB 117.2, 13 mm diarn.; Sudan, Wadi Karnoi, Neumann 62, SFB 733.0, 20 mm diam. - Aeacia sieberiana DC., Mimosoideae: Sudan, Zalingei , Neumann 79, SFB 744.0, 15 mm diarn.; Sudan, Wadi Azurn, Neumann s.n., SFB 754.0, 25 mm diam. -Albizia cf. malaeophylla (A. Rieh.) Walp., Mimosoideae : Burkina Faso, Logoniegue, Neumann 981, SFB 571.2, 20 mm diam. - Bauhinia rufeseens Lam., Caesalpinioideae: K-Jw; Burkina Faso, Gorom-Gorom , Maydell s.n., RBHw 16329; Sudan, Wadi Karnoi, Neumann 54, SFB 130.2,25 mm diarn.; Sudan, Zalingei, Neumann 123, SFB 779.0, 15 mm diam. - Burkea afrieana Hook., Caesalpinioideae: Cöte d'Ivoire, Parc National de la Comoe, Neumann 1042, SFB 274.2, 20 mm diarn.; Burkina Faso, To, Neumann 1472, SFB 1185.3,20 mm diam. - CassiasieberianaDC., Caesalpinioideae: Burkina Faso, Logoniegue, Neumann 982, SFB 551.2, 20 mm diarn.; Burkina Faso, Fada N'Gourma, Neumann 528, SFB 1051.0,30 mm diam. -Daniellia oliveri (Rolfe) Hutch . & Dalziel, Caes alpinioideae: Burkina Faso, Niangoloko, Rohde 12, SFB 915.2,40 mm diarn.; Burkina Faso, To, Neumann 1479, SFB 1188.2, 30 mm diam. - Detarium mieroearpum GuilI. & Perr., Caesalpinioideae : Burkina Faso, Niangoloko, Rohde 27, SFB 544.2, 20 mm diam. - Detarium senegalense J.F. Grnel., Caesalpinioideae: Cöte d'Ivoire, Parc National de la Cornoe, Neumann 1070, SFB 440.2, 20 mm diam. - Dialium guineense Willd., Caesalpinioideae: Cöte d' Ivoire, Parc National de la Cornoe, Neumann 1036, SFB 482.2, 20 mm diam. - Diehrostaehys ein erea (L.) Wight & Arn., Mimosoideae: Gamble s.n., K-Jw, 40 mm diarn.; Sudan, Mortagello, Neumann 111, SFB 126.0, 18 mm diarn.; Swaziland, Prior s.n., SFB 1136.0,30 mm diam. Entada afrieana GuilI. & Perr., Mimosoideae: Burkina Faso, Fada N'Gourma, Neumann 537, SFB 1077.0,40 mm diarn.; Burkina Faso, To, Neumann 1474, SFB 1186.2,20 mm diam. Faidherbiaalbida (DeI.) A. Chev.,Mimosoideae: Sudan, Wadi Karnoi,Neumann 52, SFB 729.0, 17 mm diarn.; Burkina Faso, Dori, Neumann s.n., SFB 1195.2, 15 mm diam. - Isoberlinia doka Craib & Stapf, Caesalp inioideae: Burkina Faso, To, Neumann 1472, SFB 1184.2,20 mm diam. - Parkia biglobosa (Jacq.) G.Don, Mimosoideae: Nigeria, Biu, Salzmann 89, SFB 1174.2, 15 mm diarn.; Burkina Faso, To, Neumann 1470, SFB 1183.2,20 mm diam . Piliostigma retieulatum (DC.) Hochst., Caesalpinioideae : Sudan, Zalingei, Neumann 156, SFB 807.0,25 mm diarn.; Burkina Faso, Dori, Neumann s.n., SFB 1193.2,30 mm diam. Piliostigma thonningii (Schum.) Milne-Redh., Caesalpinioideae: Burkina Faso, Logoniegue, Neumann s.n., SFB 869.2, 20 mm diarn.; Burkina Faso, To, Neumann 1467, SFB 1180.2, 20 mm diam . - Prosopis afrieana (GuilI. & Perr.) Taub., Mimosoideae: Burkina Faso, Tansarga, Neumann 671, SFB 499.0 , 40 mm diarn.; Nigeria, Konduga, Salzmann 88, SFB 1173.2,9 mm diam. - Tamarindus indiea L., Caesalpinioideae: K-Jw, 10 mm diarn.; Tanza nia, Tabora, v.Trotha s.n., SFB 639.2, 35 mm diarn.;. Burkina Faso, Logoniegue, Neumann s.n., SFB 865.2, 40 mm diam.
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