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Aluminium Accumulation in Rubiaceae

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Aluminium Accumulation in Rubiaceae IAWA Journal, Vol. 21 (2), 2000: 197–212 ALUMINIUM ACCUMULATION IN RUBIACEAE: AN ADDITIONAL CHARACTER FOR THE DELIMITATION OF THE SUBFAMILY RUBIOIDEAE? by S. Jansen1, E. Robbrecht2, H. Beeckman3 & E. Smets1 SUMMARY The chrome azurol-S test, which is a chemical spot-test for Al accumu- lation in wood, was applied to 443 wood samples of members of the Rubiaceae. A positive reaction was found in 103 specimens. Compari- son of the results with earlier analyses of leaves of Rubiaceae shows that Al accumulation occurs more frequently in leaves than in wood. The strongest Al accumulators occur in the neotropical genera Psy- chotria subg. Heteropsychotria, Coussarea, Faramea, and Rudgea. The distribution of Al accumulators is discussed in view of recent tribal and subfamilial classification of the Rubiaceae. The major conclusion is that Al accumulation is almost limited to the subfamily Rubioideae. Within the Rubioideae, however, not all tribes show the character, es- pecially the predominantly herbaceous Anthospermeae, Paederieae, Rubieae, and Spermacoceae. Al accumulation in the Urophylleae, Pauri- diantheae, Craterispermeae, and Knoxieae supports earlier associations of these tribes with the Rubioideae. Key words: Aluminium accumulation, chrome azurol-S test, chemo- taxonomy, Rubiaceae, Rubioideae. INTRODUCTION Plants containing a high level of Al in their above-ground tissues (more than 1,000 ppm / dry weight) are termed ʻaluminium plantsʼ or ʻaluminium accumulatorsʼ (Hutch- inson & Wollack 1943; Hutchinson 1945; Robinson & Edgington 1945). They have mainly been recorded by Chenery (1946, 1948a, b, 1949), Webb (1954), and Moomaw et al. (1959). In these studies, the high Al content is detected by the ʻaluminonʼ test (based on ammonium aurine tricarboxylate) applied to leaves of living or dried speci- mens. At present, the number of known accumulating families has increased to about 45. Al accumulators are especially common in families such as Anisophylleaceae, Celastraceae, Cornaceae, Diapensiaceae, Geissolomataceae, Grossulariaceae, Melas- 1) Laboratory of Plant Systematics, Institute of Botany and Microbiology, K.U. Leuven, Kard. Mercierlaan 92, B-3001 Leuven, Belgium. 2) National Botanic Garden of Belgium, Domein van Bouchout, B-1860 Meise, Belgium. 3) Royal Museum for Central Africa, Leuvensesteenweg 13, B-3080 Tervuren, Belgium. Downloaded from Brill.com09/24/2021 04:57:17PM via free access 198 IAWA Journal, Vol. 21 (2), 2000 tomataceae, Pentaphylacaceae, Polygalaceae, Proteaceae, Rubiaceae, Symplocaceae, Theaceae, and Vochysiaceae (Chenery & Sporne 1976; Metcalfe & Chalk 1983). They are in general woody plants inhabiting tropical or subtropical regions. Above the family level, Al accumulation has been accorded very little taxonomic significance. The fami- lies listed above belong to different major groups of the dicotyledons, and it is beyond doubt that the character has arisen a number of times in plant evolution. A chemical spot-test for Al and its application for wood identification was devised by Kukachka and Miller (1980). Although these investigators used a different stain (a chrome azurol-S solution) from that employed by Chenery and others, many of their results confirm earlier findings. Almost all families which showed a positive chrome azurol-S test were represented in the list of families given in Chenery and Sporne (1976) and Metcalfe and Chalk (1983). Apart from a small number of more recent papers (e.g., Quirk 1980: Vochysiaceae; Bridgwater & Baas 1982: Xanthophyl- lum; Keating & Randrianasolo 1988: Anisophylleaceae) which briefly refer to some of the results published by Kukachka and Miller (1980), very few new studies on chrome azurol-S tests are reported in literature, despite the fact that the spot-test is included in the IAWA list of standard wood characters as feature 216 (IAWA Com- mittee 1989). Since Chenery (1948b) stated that the Rubiaceae contain the largest number of Al accumulators of any family, with 647 species in 91 genera, the present study aimed to determine the taxonomic significance of Al accumulation in this very large family. Chrome azurol-S tests were applied to 443 wood samples representing all subfamilies and tribes of the family (except for some small herbaceous tribes such as Ophiorrhizeae and Argostemmateae). In addition, the results on Al accumulation in rubiaceous taxa obtained by Chenery (1946, 1948a, b) and Webb (1954) are summarised. Thus, it is possible to compare accumulation in wood and leaves, and evaluate the information in the light of recent systematic insights. The intrafamilial classification of the Rubia- ceae has drastically changed since the publication of these early works, and still is in a state of flux (e.g., Robbrecht et al. 1996; Bremer & Thulin 1998; Andersson & Rova 1999). The discussion of our results follows Robbrechtʼs (1994) classification of the family, with reference to recently proposed modifications. MATERIALS AND METHODS The material investigated came from the xylaria of Madison (MADw-SJRw), Mont- pellier (CTFw), Tervuren (Tw), and Utrecht (Uw), from the herbaria of Brussels (BR), Kew (K), Leiden (L), Missouri (MO), Paris (P), and Wageningen (WAG), and from living collections in the greenhouses of the National Botanic Garden of Belgium. The number of specimens that show a positive Al test/total number of specimens tested is given in brackets; e.g. Coussarea (11/11) means that all 11 specimens of Coussarea tested gave a positive Al test. The presence of high Al concentrations in wood was detected by use of a 0.5% solution of chrome azurol-S as described by Kukachka and Miller (1980). This solu- tion has a yellow to orange colour. One or two drops of the solution are applied to the Downloaded from Brill.com09/24/2021 04:57:17PM via free access Jansen, Robbrecht, Beeckman & Smets — Al accumulation in Rubiaceae 199 freshly exposed end-grain surface of the wood sample, because dirt and other con- taminants may affect the test. Also, we applied the solution to the transverse side of the wood block, since uptake readily occurs in this direction. Wood samples produc- ing a blue to dark blue colour indicate a positive reaction and are regarded as strong accumulators. A light purple to bluish colour shows that the test is intermediate which is characteristic for weaker Al accumulators. Wood that contains a low Al concentra- tion does not change colour in the presence of chrome azurol-S solution; these speci- mens are considered to be negative. The Al test was repeated in the case of dubious and intermediate reactions. The chrome azurol-S solution can easily be used to test woody parts of herbarium specimens by making a fresh cut a few mm2 into the sur- face before applying the test solution. RESULTS Results of the chrome azurol-S tests are given in column I of Table 1. In 89 specimens the wood produced a (bright) blue colour in a matter of minutes. In 14 specimens the chrome azurol-S test gave an intermediate reaction. Most of these woods required five to ten minutes before a change of colour was visible. Within the Cinchonoideae sensu Robbrecht, positive specimens occurred in the tribes Pauridiantheae (Pauridiantha, Poecilocalyx, and Stelecantha) and Urophylleae (Leucolophus, Maschalocorymbus, Praravinia, and Urophyllum). No accumulators were found in the subfamily Ixoroideae. In the Antirheoideae sensu Robbrecht, posi- tive taxa were restricted to the monogeneric tribe Craterispermeae, and some genera of the Knoxieae, namely Calanda, Knoxia and Pentanisia. As for the subfamily Rubi- oideae, many wood samples reacted positively. We observed high frequencies for accumulation in the neotropical genera Psychotria subg. Heteropsychotria (formerly placed in Cephaelis) (5/6), Coussarea (11/11), Faramea (13/13), and Rudgea (9/10). Other Rubioideae proved to be marked accumulators, viz. Danais (3/4), Hedyotis (3/3), Lasianthus (7/9), Prismatomeris (4/5), Rennellia (3/3), Saprosma (2/2), Tri- chostachys (2/2), and the single specimen tested of Margaritopsis (1/1). In contrast, only few representatives of Colletoecema (2/6), Gaertnera (1/4), Morinda (1/6), Palicourea (3/10), and Psychotria p.p. reacted positively. The secondary xylem of Lasianthus acuminatus, Saprosma ceylanica, and S. ternatum was found to be nega- tive, but the outermost part near the bark gave a positive reaction. For most species investigated here, only one wood sample was tested. However, from 40 species two or more specimens were investigated. The specimens were all negative in 26 species; the two wood samples of 5 species (Faramea anisocalyx, Pentanisia renifolia, Prismatomeris beccariana, Psychotria cotejensis, Trichostachys microcarpa) proved to be positive. Positive and intermediate reactions were found in 4 species (Craterispermum cerinanthum, Faramea occidentalis, Lasianthus batangen- sis, Psychotria vasiviensis). In contrast, the wood samples of Colletoecema dewevrei, Maschalocorymbus corymbosus, Otomeria micrantha, Pentanisia schweinfurthii, and Psychotria peduncularis differed in their reaction; the tests gave negative as well as positive or intermediate results for these species. Downloaded from Brill.com09/24/2021 04:57:17PM via free access 200 IAWA Journal, Vol. 21 (2), 2000 Table 1. Summary of tests on Al accumulation in wood or leaves of Rubiaceae; genera in bold have one or more positive species; genera underlined are intermediate; genera in italic are negative; if known, the nominator between brackets gives the number of Al accumulating specimens, the denominator is the total
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