OPCE-STR.vp Acta Bot. Croat. 71 (1), 115–123, 2012 CODEN: ABCRA 25 ISSN 0365–0588 eISSN 1847-8476 Fruit and seed traits of Berberis croatica Horvat and Berberis vulgaris L. DARIO KREMER1, RENATA JURI[I] GRUBE[I]1, ZVJEZDANA POPOVI]2, KSENIJA KARLOVI]3* 1 Faculty of Pharmacy and Biochemistry, University of Zagreb, Schrottova 39, HR-10000 Zagreb, Croatia. 2 Public institution »Nature Park Biokovo«, Trg Tina Ujevi}a 1, HR-21300 Makarska, Croatia. 3 Faculty of Agriculture, University of Zagreb, Sveto{imunska 25, HR-10000 Zagreb, Croatia. Abstract – The three year variability of fruits and seeds was investigated in Berberis croatica (Vo{ac, Mt Biokovo, Rakov Potok near Zagreb) and in B. vulgaris (Fran Ku{an Pharmaceutical Botanical Garden in Zagreb). Berberis croatica had the following dimen- sions of fruits (seeds): length 7.28–7.88 (4.57–5.03) mm; width 3.85–3.99 (width 1: 2.06–2.20; width 2: 1.44–1.63) mm; weight 0.065–0.078 (0.0116–0.0134) g. Dimensions of B. vulgaris fruits (seeds) were: length 10.20–11.29 (5.71–6.24) mm; width 5.29–5.83 (width 1: 2.40–2.71; width 2: 1.60–1.98) mm; weight 0.1602–0.2199 (0.0146–0.0235) g. The fruit shape of both species was similar and the length/width ratio was 1.91–2.04 in B. croatica and 1.77–2.07 in B. vulgaris. The number of seeds per fruit was 1.23–1.58 in B. croatica and 1.36–1.54 in B. vulgaris. Generally, fruits and seeds of B. vulgaris were sig- nificantly longer, wider and heavier than fruits and seeds of B. croatica. ANOVA showed significant statistical differences between populations for all analyzed fruit and seed traits while the species significantly differed in all traits, except in the fruit shape and number of seeds in fruit. Keywords: Berberis croatica, Berberis vulgaris, fruit, seed Introduction Inter- and intraspecific variability in fruit and/or seed traits has been in the focus of in- terest of many authors (JORDANO 1984, HENDRIX and SUN 1989, WILLSON et al. 1990, HERRERA 1992, OBESO and HERRERA 1994, ERIKSSON 1999) but only few of them address the genus Berberis. At the same time, when dealing with the fruit and seed properties of ACTA BOT. CROAT. 71 (1), 2012 115 * Corresponding author, e-mail: karlovic@agr.hr Copyright® 2012 by Acta Botanica Croatica, the Faculty of Science, University of Zagreb. All rights reserved. U:\ACTA BOTANICA\Acta-Botan 1-12\432 Kremer.vp 26. o ujak 2012 11:25:00 Color profile: Disabled Composite 150 lpi at 45 degrees barberry species in relation with interannual variation (HERRERA 1998, ALLEN and WILSON 1992), papers are mainly concentrated on the seed and/or fruit production. Even though data regarding basic fruit and seed related traits for some of the European species of the ge- nus Berberis, namely B. vulgaris (RUDOLF 1974, AKBULUT et al. 2009) and B. hispanica (OBESO and HERRERA 1994) could be found, those concerning B. croatica are lacking. B. croatica is a species with a still unresolved taxonomic status and is still not recognized as a distinctive species by the leading authorities on European flora. Several attempts have been made to crystallize the morphological (KARLOVI] et al. 2009) and chemical difference (ZOVKO KON^I] et al. 2010) between B. croatica and B. vulgaris. However, fruit and seed characters have not been included in the analysis to date yet they could help in the charac- terization of the distinctive morphological traits of these species. Material and methods Fruits of Berberis croatica Horvat and B. vulgaris L. were randomly collected during three years (from 2008 to 2010). B. croatica fruits were collected in one natural habitat (Vo{ac, Mt Biokovo, in text as abbreviation Vo; 43°18’ N; 17°02’ E; 1422 m a.s.l.), while B. vulgaris fruits were collected in one natural (Rakov Potok near Zagreb, in text as abbrevia- tion RP; 45°47’ N; 15°42’ E; 160 m a.s.l.) and one cultivated habitat (Fran Ku{an Pharma- ceutical Botanical Garden, Zagreb, in text as abbreviation PBG; 45°50’ N; 15°59’ E; 195 m a.s.l.). Twenty plants were harvested in the locality Vo{ac, ten in the locality Rakov Potok and six in the cultivated habitat. According to our permit from the Ministry of Culture of the Republic of Croatia, we were able to analyse 100 fruits for the locality Vo{ac and so the same number of B. vulgaris fruits (100) was collected for each population. Length, width and weight of fruits, as well as seeds, were measured. Since the seeds of the researched Berberis species have irregular shapes, the width of the seed was measured on two axes (width 1 and width 2). The ratio between length and width of fruits and length and width (width 1, width 2) of seeds was obtained through mathematical calculation, as well as their mutual (fruit-seed) ratio, which included length, width and weight. A greater ratio indicates narrower fruits (or seeds), while a smaller ratio indicates more orbicular fruits (or seeds). The differences in fruit and seed traits between species and populations were analyzed by the General Linear Models (GLM) procedure using repeated measures ANOVA and Tuckey’s post-hoc test at the p £0.05 level. The statistical analysis was performed using Statistica 7 software (StatSoft Inc., Tulsa, OK, USA). Results Table 1 presents descriptive statistics for the analyzed traits of fruits and seeds of Berberis croatica and B. vulgaris. The average length of B. croatica fruits ranged from 7.28 mm (Vo 2010) to 7.88 mm (Vo 2009) while the width ranged from 3.85 mm (Vo 2010) to 3.99 mm (Vo 2008). Weight of B. croatica fruits ranged from 0.065 g (Vo 2008) to 0.078 g (Vo 2010). Fruits of B. vulgaris were the largest and their length ranged from 10.20 mm (PBG 2009) to 11.29 mm (RP 2010), while the width ranged from 5.29 mm (RP 2008) to 5.83 mm (PBG 2009). Weight of B. vulgaris fruits ranged from 0.1602 g (RP 2009) to 0.2199 g (PBG 2009). The shapes of the two species were similar and the ratio between 116 ACTA BOT. CROAT. 71 (1), 2012 KREMER D., JURI[I] GRUBE[I] R., POPOVI] Z., KARLOVI] K. U:\ACTA BOTANICA\Acta-Botan 1-12\432 Kremer.vp 26. o ujak 2012 11:25:00 Color profile: Disabled Composite 150 lpi at 45 degrees length and width of fruits was from 1.91 (Vo 2010) to 2.04 (Vo 2009) in B. croatica and from 1.77 (PBG 2009) to 2.07 (RP 2010) in B. vulgaris. The most variable trait was fruit weight, ranging from 25.61 % (Vo 2010) to 28.55 % (Vo 2008) and from 22.84 % (RP 2010) to 36.54 % (PBG 2008) in B. croatica and B. vulgaris, respectively. The least vari- able trait was fruit length, which ranged from 5.84 % (Vo 2009) to 9.62 % (Vo 2010) and from 6.37 % (PBG 2009) to 9.83 % (RP 2008) in B. croatica and B. vulgaris, respectively (Tab. 1). The number of seeds per fruit ranged from 1.23 (Vo 2010) to 1.58 (Vo 2009) in B. croatica and from 1.36 (RP 2008, RP 2010) to 1.54 (PBG 2010) in B. vulgaris. ACTA BOT. CROAT. 71 (1), 2012 117 BERBERIS CROATICA AND B. VULGARIS FRUIT AND SEED TRAITS Tab. 1. Statistics for analyzed traits of fruits and seeds of Berberis croatica – Vo{ac (Vo) and B. vulgaris – Rakov Potok (RP) and Pharmaceutical Botanical Garden »Fran Ku{an« (PBG). Min, Max and Std. dev. are in mm (length, width) and g (weight); CV (coefficient of variabil- ity) is in %. Minimum and maximum are bolded. Statistics B. croatica B. vulgaris Vo 2008 Vo 2009 Vo 2010 RP 2008 RP 2009 RP 2010 PBG 2008 PBG 2009 PBG 2010 Fruit traits Length N Mean Min Max Std. dev. CV 100 7.83 7.00 9.04 0.50 6.39 100 7.88 6.89 9.07 0.46 5.84 100 7.28 5.26 9.43 0.70 9.62 100 10.89 5.34 12.74 1.07 9.83 100 10.39 8.64 12.10 0.79 7.60 100 11.29 5.84 13.26 0.91 8.06 100 10.42 8.94 12.54 0.74 7.10 100 10.20 9.01 12.47 0.65 6.37 100 10.41 7.33 12.12 0.67 6.44 Width N Mean Min Max Std. dev. CV 100 3.99 3.03 5.03 0.42 10.53 100 3.91 2.86 5.06 0.44 11.25 100 3.85 2.23 5.01 0.51 13.25 100 5.29 3.91 6.90 0.69 13.04 100 5.30 4.22 6.90 0.62 11.70 100 5.55 4.16 7.04 0.54 9.73 100 5.42 3.76 7.76 0.94 17.34 100 5.83 4.59 7.58 0.68 11.66 100 5.71 3.50 7.43 0.73 12.78 Length/ Width N Mean Min Max Std. dev. CV 100 1.98 1.46 2.88 0.25 12.63 100 2.04 1.52 2.88 0.25 12.25 100 1.91 1.40 3.05 0.22 11.52 100 2.05 1.22 2.77 0.19 9.27 100 1.98 1.55 2.78 0.22 11.11 100 2.07 1.30 2.40 0.19 9.18 100 1.97 1.49 2.60 0.30 15.23 100 1.77 1.47 2.07 0.16 9.04 100 1.85 1.12 2.66 0.25 13.51 Weight N Mean Min Max Std. dev. CV 100 0.065 0.034 0.119 0.018 27.06 100 0.071 0.032 0.113 0.020 28.55 100 0.078 0.036 0.132 0.020 25.61 100 0.182 0.066 0.307 0.050 27.48 100 0.160 0.082 0.250 0.037 22.85 100 0.209 0.107 0.328 0.048 22.84 100 0.180 0.062 0.362 0.066 36.54 100 0.220 0.102 0.376 0.0531 24.15 100 0.206 0.051 0.348 0.062 30.05 U:\ACTA BOTANICA\Acta-Botan 1-12\432 Kremer.vp 26. o ujak 2012 11:25:01 Color profile: Disabled Composite 150 lpi at 45 degrees 118 ACTA BOT. CROAT. 71 (1), 2012 KREMER D., JURI[I] GRUBE[I] R., POPOVI] Z., KARLOVI] K. Statistics B. croatica B. vulgaris Vo 2008 Vo 2009 Vo 2010 RP 2008 RP 2009 RP 2010 PBG 2008 PBG 2009 PBG 2010 Seed traits Length N Mean Min Max Std. dev. CV 145 5.03 4.13 5.72 0.31 6.16 158 4.92 3.97 5.84 0.30 6.10 123 4.57 3.58 5.74 0.38 8.32 136 5.88 4.30 6.94 0.47 7.99 138 5.71 4.23 6.88 0.52 9.11 136 5.99 4.29 6.99 0.47 7.85 142 6.14 4.81 7.29 0.45 7.33 149 5.94 5.14 6.64 0.36 6.06 154 6.24 4.51 7.75 0.49 7.85 Width 1 N Mean Min Max Std. dev. CV 145 2.10 1.28 2.90 0.26 12.38 158 2.06 1.22 2.51 0.23 11.17 123 2.20 1.30 2.60 0.18 8.18 136 2.57 1.43 3.09 0.27 10.51 138 2.55 1.54 3.16 0.35 13.73 136 2.71 2.03 3.14 0.23 8.49 142 2.46 1.78 3.28 0.33 13.41 149 2.40 1.87 3.14 0.32 13.33 154 2.50 1.68 3.32 0.31 12.40 Width 2 N Mean Min Max Std. dev. CV 145 1.49 0.82 2.27 0.26 17.45 158 1.44 0.92 1.97 0.21 14.58 123 1.63 0.85 1.96 0.22 13.50 136 1.94 0.87 2.48 0.30 15.46 138 1.82 1.09 2.75 0.25 13.74 136 1.98 0.96 2.57 0.30 15.15 142 1.61 0.50 2.36 0.34 21.12 149 1.60 0.79 2.24 0.34 21.25 154 1.66 0.83 2.66 0.36 21.69 Length/ Width 1 N Mean Min Max Std. dev. CV 145 2.43 1.87 3.62 0.30 12.35 158 2.41 1.93 3.53 0.26 10.79 123 2.09 1.66 2.83 0.20 9.57 136 2.30 1.88 3.30 0.23 10.00 138 2.26 1.87 2.96 0.22 9.73 136 2.22 1.71 2.67 0.21 9.46 142 2.53 1.88 3.40 0.31 12.25 149 2.50 2.01 3.07 0.24 9.60 154 2.52 1.90 3.96 0.30 11.90 Length/ Width 2 N Mean Min Max Std. dev. CV 145 3.48 2.04 5.09 0.57 16.38 158 3.48 2.45 4.95 0.51 14.66 123 2.86 2.17 5.01 0.45 15.73 136 3.12 2.22 5.65 0.59 18.91 138 3.18 2.15 4.71 0.42 13.21 136 3.10 2.15 5.46 0.59 19.03 142 3.95 2.62 6.11 0.80 20.25 149 3.88 2.58 6.67 0.84 21.65 154 3.89 2.44 6.72 0.83 21.34 Width 1/ Width 2 N Mean Min Max Std. dev. CV 145 1.45 0.56 2.12 0.29 20.00 158 1.46 0.92 1.91 0.22 15.07 123 1.37 1.01 1.89 0.16 11.68 136 1.35 1.07 2.40 0.19 14.07 138 1.42 0.76 2.20 0.20 14.08 136 1.40 0.98 2.29 0.21 15.00 142 1.58 1.02 2.51 0.32 20.25 149 1.56 1.03 2.96 0.34 21.79 154 1.55 0.93 2.81 0.30 19.35 Tab. 1. – continued U:\ACTA BOTANICA\Acta-Botan 1-12\432 Kremer.vp 26. o ujak 2012 11:25:01 Color profile: Disabled Composite 150 lpi at 45 degrees Seeds of B. vulgaris were also larger than seeds of B. croatica. The length of B. croatica seeds ranged from 4.57 mm (Vo 2010) to 5.03 mm (Vo 2008), width 1 from 2.06 mm (Vo 2009) to 2.20 mm (Vo 2010), width 2 from 1.44 mm (Vo 2009) to 1.63 mm (Vo 2010) and the weight from 0.0116 g (Vo 2008) to 0.0134 (Vo 2010) g. In B. vulgaris seeds the length ranged from 5.71 mm (RP 2009) to 6.24 mm (PBG 2010), width 1 from 2.40 mm (PBG 2009) to 2.71 mm (RP 2010), width 2 from 1.60 mm (PBG 20) to 1.98 mm (RP 2010) and the weight from 0.0146 g (PBG 2010) to 0.0235 g (RP 2010). The most variable trait was seed weight, ranging from 20.15 % (Vo 2010) to 24.14 % (Vo 2008) and from 22.13 % (RP 2010) to 57.72 % (RP 2009) in B. croatica and B. vulgaris, respectively. The least variable trait was seed length and it ranged from 6.10 % (Vo 2009) to 8.32 % (Vo 2010) and from 6.06 % (PBG 2009) to 9.11 % (RP 2009) in B. croatica and B. vulgaris, respectively (Tab. 1). ANOVA showed significant statistical differences between species for most analyzed fruit and seed traits (Tab. 2). The only exception was the ratio between fruit length and width as well as the number of seeds in the fruit. The analysis of fruit traits between species showed that fruits of B. vulgaris were significantly longer, wider and heavier than fruits of B. croatica (Tab. 3). Nevertheless, fruit shape (based on ratio between length and wide) was similar in the two species. ACTA BOT. CROAT. 71 (1), 2012 119 BERBERIS CROATICA AND B. VULGARIS FRUIT AND SEED TRAITS Statistics B. croatica B. vulgaris Vo 2008 Vo 2009 Vo 2010 RP 2008 RP 2009 RP 2010 PBG 2008 PBG 2009 PBG 2010 Weight N Mean Min Max Std. dev. CV 145 0.012 0.001 0.018 0.003 24.14 158 0.012 0.001 0.021 0.003 23.97 123 0.013 0.005 0.019 0.003 20.15 136 0.022 0.004 0.035 0.007 31.78 138 0.022 0.007 0.034 0.013 57.72 136 0.024 0.010 0.035 0.005 22.13 142 0.018 0.003 0.038 0.008 44.32 149 0.017 0.003 0.046 0.007 43.79 154 0.015 0.002 0.033 0.006 42.47 Seed and fruit traits Seeds in Fruit N Mean Min Max Std. dev. CV 100 1.45 1.00 2.00 0.50 34.49 100 1.58 1.00 3.00 0.52 32.91 100 1.23 1.00 2.00 0.42 34.15 100 1.36 1.00 2.00 0.48 35.29 100 1.38 1.00 2.00 0.49 35.51 100 1.36 1.00 2.00 0.48 35.29 100 1.42 1.00 2.00 0.50 35.21 100 1.49 1.00 2.00 0.50 33.56 100 1.54 1.00 2.00 0.50 32.47 Seed/Fruit Length 0.64 0.62 0.63 0.54 0.55 0.53 0.59 0.60 0.60 Seed width 1/Fruit Width 0.61 0.62 0.54 0.43 0.43 0.40 0.47 0.43 0.44 Seed width 2/Fruit Width 0.87 0.89 0.74 0.59 0.60 0.56 0.73 0.67 0.68 Seed/Fruit Weight 0.177 0.170 0.172 0.119 0.137 0.113 0.098 0.077 0.071 Tab. 1. – continued U:\ACTA BOTANICA\Acta-Botan 1-12\432 Kremer.vp 26. o ujak 2012 11:25:01 Color profile: Disabled Composite 150 lpi at 45 degrees 120 ACTA BOT. CROAT. 71 (1), 2012 KREMER D., JURI[I] GRUBE[I] R., POPOVI] Z., KARLOVI] K. Tab. 2. Results of ANOVA for analysed traits of fruit and seed. Asterisks indicate significant p val- ues at p£0.05. Effect Sum of Squares Degr. of freedom Mean Square F p-level Fruit Length Population 1765.34 2 882.67 1608.8 <0.0001* Species 1725.57 1 1725.57 2536.49 <0.0001* Fruit Width Population 523.84 2 261.92 643.30 <0.0001* Species 513.08 1 513.08 1161.04 <0.0001* Fruit Length/Width Population 4.504 2 2.252 40.52 <0.0001* Species 0.202 1 0.202 2.89 0.0904 Fruit Weight Population 2.99660 2 1.49830 832.78 <0.0001* Species 2.94489 1 2.94489 1497.422 <0.0001* Seed Length Population 347.93 2 173.97 977.2 <0.0001* Species 335.54 1 335.54 1608.2 <0.0001* Seed Width 1 Population 49.319 2 24.659 300.74 <0.0001* Species 43.625 1 43.625 454.14 <0.0001* Seed Width 2 Population 31.795 2 15.898 189.02 <0.0001* Species 15.041 1 15.041 119.48 <0.0001* Seed Length/Width 1 Population 16.161 2 8.081 126.2 <0.0001* Species 1.598 1 1.598 15.92 0.0001* Seed Length/Width 2 Population 142.38 2 71.19 165.92 <0.0001* Species 18.93 1 18.93 25.67 <0.0001* Seed Width 1/Width 2 Population 6.560 2 3.280 48.44 <0.0001* Species 0.826 1 0.826 10.08 0.0016* Seed Weight Population 0.023150 2 0.011575 44.116 <0.0001* Species 0.016446 1 0.016446 59.0474 <0.0001* Seeds in fruit Population 2.047 2 1.023 4.241 0.0153* Species 0.005 1 0.005 0.020 0.8870 U:\ACTA BOTANICA\Acta-Botan 1-12\432 Kremer.vp 26. o ujak 2012 11:25:01 Color profile: Disabled Composite 150 lpi at 45 degrees ANOVA of seed traits showed similar results (Tab. 4). In general, seeds of B. vulgaris were significantly longer, wider and heavier than fruits of B. croatica. Interspecies and ACTA BOT. CROAT. 71 (1), 2012 121 BERBERIS CROATICA AND B. VULGARIS FRUIT AND SEED TRAITS Tab. 3. p values from Tukey post-hoc test for variability of fruits of Berberis croatica in Vo{ac and B. vulgaris in Rakov Potok and Pharmaceutical botanical garden »Fran Ku{an« (PBG) pop- ulations. Asterisks indicate significant p values at p£0.05. Population Vo{ac Rakov Potok Population Vo{ac Rakov Potok Length Length/Width Vo{ac Vo{ac Rakov Potok <0.0001* Rakov Potok 0.1867 PBG <0.0001* 0.0001* PBG 0.9058 0.0757 Width Weight Vo{ac Vo{ac Rakov Potok <0.0001* Rakov Potok <0.0001* PBG <0.0001* 0.4388 PBG <0.0001* 0.9738 Tab. 4. p values from Tukey post-hoc test for variability of seeds of Berberis croatica in Vo{ac and B. vulgaris in Rakov Potok and Pharmaceutical botanical garden »Fran Ku{an« (PBG) pop- ulations. Asterisks indicate significant p values at p£0.05. Population Vo{ac Rakov Potok Length Vo{ac Rakov Potok <0.0001* PBG <0.0001* <0.0001* Width 1 Vo{ac Rakov Potok <0.0001* PBG <0.0001* 0.0058* Width 2 Vo{ac Rakov Potok <0.0001* PBG 0.0031* <0.0001* Length/Width 1 Vo{ac Rakov Potok 0.0008* PBG 0.0163* <0.0001* Population Vo{ac Rakov Potok Length/Width 2 Vo{ac Rakov Potok 0.0002* PBG <0.0001* <0.0001* Width 1/Width 2 Vo{ac Rakov Potok 0.0198* PBG 0.0004* <0.0001* Weight Vo{ac Rakov Potok <0.0001* PBG <0.0001* <0.0001* Seeds in fruit Vo{ac Rakov Potok 0.4001 PBG 0.9030 0.6651 U:\ACTA BOTANICA\Acta-Botan 1-12\432 Kremer.vp 26. o ujak 2012 11:25:01 Color profile: Disabled Composite 150 lpi at 45 degrees intraspecies variability for seed traits was something higher than for fruit traits. The least inter- and intraspecies variability was recorded for the number of seeds in fruit. Discussion The least variable traits in this analysis were fruit and seed length while the most vari- able traits were fruit and seed weight. That was expected since dimensionality consider- ations predict that, all else being equal, the coefficients of variability (CVs) of mass-related traits will be larger than those of linear or surface measurements (LANDE 1977), such as length and width of fruits and seeds. The largest variability for fruit and seed weight was also reported by OBESO and HERRERA (1994) which state CVs for these traits ranging gener- ally from 20–30 % for eight investigated species, including Berberis hispanica. The same authors also reported fruit length and width variability ranging generally between 5 and 12 % (7.6 % and 7.4 % in B. hispanica for fruit length and fruit width, respectively). These re- sults are in keeping with the results reported in this paper. Regarding fruit traits of B. vulgaris, AKBULUT et al. (2009) reported much lower values for the wild population of B. vulgaris growing in Turkey. For example, average fruit length, width and mass are reported to be 7.69 mm, 3.32 mm, and 0.07 g, (compared with the aver- age value range for the same traits in this investigation, that is, 10.20–11.29 mm, 5.29–5.83 mm and 0.1602–0.2199 g). This could be explained by environmental factors but also by great intraspecific variation in Berberis already reported by LANDRUM (1999). Generally, fruits and seeds of B. vulgaris were significantly longer, wider and heavier than fruits and seeds of B. croatica. This is in congruence with the results of KARLOVI] et al. (2009) who, when analyzing distinguishing morphological traits between these two spe- cies, stated that B. croatica is characterized by reduced growth/size traits compared with B. vulgaris. Our results support this thesis. Results regarding fruit and seed traits show signifi- cant differences between species for most analyzed traits (10 out of 12). Nevertheless, whether the observed differences between B. croatica and B. vulgaris are merely non-he- reditary modifications caused by environmental factors or species-related differences, should be further tested by means of genetic analysis. Acknowledgements This work was supported by the Ministry of Science, Education and Sports of the Re- public of Croatia (project »Micromorphological and chemotaxonomic researches on some species of the family Lamiaceae«). References AKBULUT, M., CALISIR, S., MARAKOGLU, T., COKLAR, H., 2009: Some physicomechanical and nutritional properties of barberry (Berberis vulgaris L.) fruits. Journal of Food Pro- cess Engineering 32, 497–511. ALLEN, R. B., WILSON, J. B., 1992: Fruit and seed production in Berberis darwinii Hook., a shrub recently naturalised in New Zealand. New Zealand Journal of Botany 30, 45–55. 122 ACTA BOT. CROAT. 71 (1), 2012 KREMER D., JURI[I] GRUBE[I] R., POPOVI] Z., KARLOVI] K. 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Food and Chemical Toxicology 48, 2176–2180. WILLSON, M. F., MICHAELS, H. J., BERTIN, R. I, BENNER, B., RICE, S., LEE, T. D., HARTGERINK, A. P., 1990: Intraspecific variation in seed packaging. American Midland Naturalist 123, 179–185. ACTA BOT. CROAT. 71 (1), 2012 123 BERBERIS CROATICA AND B. VULGARIS FRUIT AND SEED TRAITS U:\ACTA BOTANICA\Acta-Botan 1-12\432 Kremer.vp 26. o ujak 2012 11:25:01 Color profile: Disabled Composite 150 lpi at 45 degrees