1228 S. Mali, R.M. Borges / Biochemical Systematics <strong>and</strong> Ecology 31 (2003) 1221–1246Table 3Levels of astringency, total phenolics, condensed tannins, gallotannins, ellagitannins <strong>and</strong> ADF in plant parts (mean ± SD)Category Astringency Total phenolics Condensed tannins Gallotannins Ellagitannins ADFImmature 11.37 ± 10.54 (17) 2.70 ± 3.59 (16) 16.85 ± 19.02 (16) 0.56 ± 1.00 (17) 0.24 ± 0.38 (16) 33.98 ± 15.96 (17)leavesMature 7.56 ± 7.31 (17) 1.05 ± 0.74 (16) 13.14 ± 20.29 (16) 0.28 ± 0.65 (16) 0.05 ± 0.14 (20) 33.18 ± 10.12 (17)leavesPetioles 10.42 ± 7.52 (9) 3.01 ± 4.19 (10) 20.37 ± 20.57 (10) 0.10 ± 0.16 (10) 0.09 ± 0.21 (10) 35.27 ± 2.35 (9)Flowers 9.51 ± 7.30 (14) 1.99 ± 2.04 (13) 18.82 ± 21.56 (13) 2.70 ± 5.34 (13) 0.22 ± 0.53 (13) 25.93 ± 12.39 (13)Ripe fruit 4.57 ± 5.43 (14) 1.13 ± 1.58 (15) 6.54 ± 7.76 (15) 0.05 ± 0.14 (15) 0.01 ± 0.04 (16) 27.00 ± 14.30 (14)pulpMature 4.13 ± 6.41 (21) 1.00 ± 1.25 (18) 8.78 ± 20.49 (18) 0.72 ± 1.58 (18) 0.05 ± 0.13 (17) 16.10 ± 11.90 (21)seedsTree twigs 1.95 ± 2.77 (10) 0.29 ± 0.23 (8) 3.18 ± 4.34 (8) 0.001 ± 0.001 (5) 0.001 ± 0.001 (8) 49.70 ± 20.67 (10)Inner bark 6.35 ± 7.11 (17) 0.87 ± 0.76 (16) 16.61 ± 20.65 (15) 0.06 ± 0.19 (16) 0.03 ± 0.09 (16) 38.48 ± 14.07 (17)Values are pooled for trees <strong>and</strong> lianas. Astringency <strong>and</strong> total phenolics expressed as percent tannic acid, condensed tannins as percent quebracho tannin,gallotannins as percent gallic acid <strong>and</strong> ellagitannins as percent ellagic acid in terms of dry weight. Values in parentheses are number of species (N).
S. Mali, R.M. Borges / Biochemical Systematics <strong>and</strong> Ecology 31 (2003) 1221–12461229Table 4Correlates of various measures of phenolic compounds <strong>and</strong> fibreTotal Condensed Gallotannin Ellagitannin ADFphenolics tanninAstringency 0.48 (119)∗∗∗ 0.37 (118)∗∗∗ 0.20 (116)∗∗ 0.22 (115)∗∗ 0.13 (129)∗Total phenolics 0.28 (121)∗∗∗ 0.36 (116)∗∗∗ 0.17 (114)∗ 0.08 (119)Condensed tannin 0.14 (115) 0.02 (113) 0.13 (118)∗Gallotannin 0.32 (113)∗∗∗ 0.2 (116)∗∗Ellagitannin 0.02 (114)Values are Kendall’s correlation coefficients. Values in parentheses are sample sizes (N). ∗P 0.05;∗∗P 0.01; ∗∗∗ P 0.001 (∗∗ is significant after Bonferroni’s correction for multiple tests).Although correlations were performed between these phenolic measures separatelyfor each plant part type, e.g. mature leaves, the results were not significant afterBonferroni’s correction for multiple tests, except for the positive correlations betweenastringency <strong>and</strong> total phenolics in mature leaves of trees (Kendall’s t = 0.4, N =17, P 0.01) <strong>and</strong> between astringency <strong>and</strong> total phenolics (Kendall’s t = 0.61 N= 16, P 0.01), as well as total phenolics <strong>and</strong> condensed tannins (Kendall’s t =0.58, N = 16, P 0.01) in inner bark.3.4. Co-occurrence of condensed <strong>and</strong> hydrolysable tanninsSince almost all plant parts contained condensed tannins (Table 2), we were unableto examine the segregation between hydrolysable <strong>and</strong> condensed tannins in plantparts. Within each plant part we, therefore, compared the frequency of species containingboth hydrolysable <strong>and</strong> condensed tannins to those containing condensed tanninsalone using binomial probabilities, <strong>and</strong> we found that only in ripe fruit pulpwas there a significantly higher frequency of samples that contained condensed tanninsbut also did not contain hydrolysable tannins (N = 15, P 0.02). Since therewere samples that did not contain gallotannins, we examined the independence ofoccurrence of gallotannins <strong>and</strong> ellagitannins using a 2 × 2 contingency test, withYates’ correction, <strong>and</strong> found that there was no segregation between gallotannins <strong>and</strong>ellagitannins in any plant part.3.5. Co-occurrence of <strong>alkaloids</strong>, <strong>saponins</strong> <strong>and</strong> phenolicsSince tannins occurred in almost all plant parts examined, we were unable toexamine whether the occurrence of tannins <strong>and</strong> <strong>alkaloids</strong> or tannins <strong>and</strong> <strong>saponins</strong>were independent of each other. We, therefore, examined whether significantly fewernumbers of plant parts of the different species contained both tannins <strong>and</strong> <strong>alkaloids</strong>or both tannins <strong>and</strong> <strong>saponins</strong> than those that contained tannins alone. We did thisfor each plant part category by calculating exact probabilities of the binomial since
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