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Anthocyanins of the Sterculiaceae: Flavonoid scores and Hennigian phylogenetic systematics
8iochemistrySysrernaticsandEcology, Printed in Great Britain.
Vol. 10, No. 2, pp. 197-199, 1982.
0306-1978/82/Q20197-(}3 $O3.0010 C) 1982PergamonPressLtd.
Letter to the Editors Anthocyanins of the Sterculiaceae: Flavonoid Scores and Hennigian Phylogenetic Systematics P. MICK RICHARDSON Harding Laboratory, New York Botanical Garden, Bronx, NY 10458, USA Key Word Index - Sterculiaceae; anthocyanins; flavonoid scores; cladistics; chemosystematics.
/Ig~r==Ct - Published results of the distribution of anthocyanins in the Sterculiaceae have been re-interpreted on the basis of the phylogenetic status of the compounds present. A flavonoid score system was less useful than a cladistic interpretation based on Hennigian arguments.
Introduotion The anthocyanins of the Sterculiaceae have recently been investigated by Scogin [1] who carried out a limited survey of 13 taxa in 7 genera in the 5 sections of the family. He concluded "floral pigments appear on this limited survey to be of little systematic utility in the Sterculiaceae." In my opinion, Scogin has underestimated the value of his data. The use of a ftavonoid score system and Hennigian phylogenetic systematics both reveal the phylogenetic content of Scogin's data and suggest that this limited survey has produced much useful information.
Results The phylogenetic status of anthocyanin pigments has been discussed in the literature for many years, most recently in a review by Gornall and Bohm [2]. The consensus of opinion is that cyanidin represents the primitive state, and delphinidin and pelargonidin represent advanced states. Similarly, as regards O-glycosylation, glucose is primitive while galactose and arabinose are advanced [2, 3]. Using this information, a quick examination of Scogin's data [1] suggests that the section Dombeyeae contains only primitive compounds (i.e. cyanidin-3-O-glucoside), while all the other sections have some advanced features; e.g. the presence of delphinidin, pelargonidin, galactose or arabinose. We can express this more meaningfully by the use of a flavonoid score system. Such systems were originally independently devised by Bate-Smith [4, 5] and da (Received 1 November 1981)
Mata Rezende and Gottlieb [6], and subsequently used by Harborne [7-9] and Campbell et al. [10]. Briefly, the method is as follows: the various flavonoid features are allocated points depending on their presumed relative advancement and each taxon is then assigned a total score dependent on the flavonoids it contains. In the case of the Sterculiaceae, it is appropriate that cyanidin and Oglucosylation are given the score 0, while pelargonidin, delphinidin, O-arabinosylation and O-galactosylation each score one point (Table 1). Thus, each section of the family accumulates the following scores: Dombeyeae 0; Fremontieae 1; Lasiopetaleae 1; Buttnerieae 2 and Sterculieae 3 (Table 2). However, the accumulated totals obscure the fact that the score for sections Frernontieae and Lasiopetaleae is due to the presence of advanced anthocyanidins, while Buttnefieae has points only for advanced OL glycosylation features; Sterculieae scoring points for both features. This information can be revealed by using a second method which is relatively little used by botanists but more widely used by zoologists. The method is Hennigian phylogenetic TABLE 1. CHARACTERSTATESOF ANTHOCYANIN FEATURESOF THE STERCULIACEAE Character state
Characters
Primitive
1 Cyanidin 2 Delphinidin 3 Pelargonidin 40-Glucosylation 50-Arsbinosylation 60-Galactosylstion
Present Absent Absent Present Absent Absent
197
Dedved (advanced) Absent Present Prtment
Absent
Present
Pr~,gt
196
P MICK RICHARDSON
TABLE 2. DATA MATRIX OF OCCURRENCE OF ADVANCED ANTHOCYANIN FEATURES IN THE SECTIONS OF THE STERCULIACEAE Character" Taxon Del Pel Ara Gal Fremont/eee Las~oetaleee Stercu#eee Buttner/eae Oomtxweee
+ + + .
+ +
-
+ .
.
+
.
"See Table1: + = present; - = absent.
systematics and produces cladograms (branched, rooted diagrams) based on the presence of shared, advanced characters. For botanical examples, see refs. [11-13], and for examples using flavonoid studies see ref. [14]. In the case of the Sterculiaceae the ctadogram is decided by the advanced anthocyanin features which are present in one or more sections of the family. The method is as follows: decide the phylogenetic status of the anthocyanin features (Table 1), produce a data matrix of the occurrence of the advanced anthocyanin features in each taxon (Table 2), and finally produce the most parsimonious cladogram from the data matrix (Fig. 1). The primitive and advanced features of the anthocyanin characters (Table 1) were decided by traditional interpretations as discussed earlier. It is more preferable to decide the primitive and advanced features by an out-group comparison, i.e. on the presence or absence of these features in the groups most closely related to the one under study. However, we do not know the anthocyanins which occur in the sister group to the Sterculiaceae, or even which family to use as a sister group. At the present time we must assume the character polarity states based on traditional interpretations are correct. In the future they can
be changed, if necessary, and a new cladogram produced. We must also assume that the sections and the family itself are monophyletic. In the cladogram (Fig. 1), sections Fremontieae and Lasiopetaleae cannot be separated because they produce the same anthocyanins. These two sections and the Sterculieaeshare the advanced feature of delphinidin production and form a monophyletic grouping. Section Sterculieae has the unique derived feature of pelargonidin synthesis and of O-arabinosylation. The latter also occurs in the Buttnerieae and is interpreted as being independently derived (a parallelism) in both sections. Section Buttnerieae has the unique derived feature of O-galactosylation while section Dombeyeaehas no advanced features, so there is a basal trichotomy. Fig. 1 is the most parsimonious ctadogram that can be derived from the data matrix: any other arrangement of the five sections would result in the production of more parallelisms, or even reversals in characters. Dkmu=don The cladogram in Fig. 1 shows which sections of the Sterculiaceae are most closely related in terms of recent common ancestry based on shared advanced anthocyanin features. The cladogram would be greatly improved by the addition of further characters, such as the distribution of other flavonoids. It could then be compared to other cladograms based on morphological or cytological features to give further insights into the phylogeny of the Sterculiaceae. Cladistic interpretations of flavonoid data such as the one presented here can readily be performed and they transform raw data into a form which can be readily understood by other taxonomists. Other chemosystematists are urged to consider the possibilities of presenting their data in a Hennigian fashion. A c k n o w l e d ~ m t e n t s - I am grateful to Chris Humphries and David Young, who criticized earlier drafts of a paper similar to this one. The study was largely supported by NSF grant DEB 79-08525 awarded to D. S. Seigter and D. A. Young, and its completion was undertaken at NYBG.
Referene~
FIG. 1. CLAOOGRAM SHOWING PHYLOGENETICRELATIONSHIPS OF SECTIONS IN THE STERCULIACEAE RASED ON ANTHOCYANIN CHARACTERS: Key: S=Stercu/~ee; F==Frem(~l~ee; L=/.ai~o~eee; B = B u t ~ ; D=Dombeyel~, 2= Delphinidin; 3 = Pe~-O~,-~din;5 -- O-Atabino~t~rt; 6 = O-Galectosylation.
1. Scogin, R. (1979)Biochem. Syst. Ecol. 7, 35. 2. Gornall, R. J. and Bohm, B. A. (1978) Syst. Bot. 3. Wallace, J. W. and Markham, K. R. (1978) Amer. 86,965. 4. Bate-Smith, E. C. (1973) Bot. J. Linn. Soc. 87, 5. Bate-Smith, E. C. and Richens, R. H. (1973)
3, 353. J. B o t a n y
347.
Biochem. Syst. 1 , 1 4 1 . 6. da M a t a Rezende, C. M. A. and Gottlieb, O. R. (1973) B i o c h e m . Syst. 1, 111.
ANTHOCYANINS OF THE STERCULIACEAE 7. Harborne, J. B. (1976) Nova Acta Leopoldina Suppl. 7, 563. 8. Harborne, J. B. (1977) Biochem. Syst. Ecol. 5, 7. 9. Smith, D. M., Glennie, C. W., Harborne, J. B. and Williams, C. A. (1977) B~ochem. Syst. Ecol. 5, 107. 10. Campbell, E. O., Markham, K. R., Moore, N. A., Porter, L. J. and Wallace, J. W. (1979) J. Hattori Bot. Lab. 45, 185. 11. Bremer, K. and Wanntorp, H.-E. (1978) Taxon27, 317.
199 12. Humphries, C. J. (1979) Bull. Br. Mus. Nat. Hist. (Bot.) 7,83. 13. Humphries, C. J. and Richardson, P. M. (1980) Chemosystematics. Principles and Practice (Bisby, F. A., Vaughan, J. G. and Wright, C. A., eds.) p. 353. Academic Press, London. 14. Richardson, P. M. (1962) Advancesin Cladistics (Funk, V. and Platnick, N., eds.) in press.
Vol. 10, No. 2, pp. 197-199, 1982.
0306-1978/82/Q20197-(}3 $O3.0010 C) 1982PergamonPressLtd.
Letter to the Editors Anthocyanins of the Sterculiaceae: Flavonoid Scores and Hennigian Phylogenetic Systematics P. MICK RICHARDSON Harding Laboratory, New York Botanical Garden, Bronx, NY 10458, USA Key Word Index - Sterculiaceae; anthocyanins; flavonoid scores; cladistics; chemosystematics.
/Ig~r==Ct - Published results of the distribution of anthocyanins in the Sterculiaceae have been re-interpreted on the basis of the phylogenetic status of the compounds present. A flavonoid score system was less useful than a cladistic interpretation based on Hennigian arguments.
Introduotion The anthocyanins of the Sterculiaceae have recently been investigated by Scogin [1] who carried out a limited survey of 13 taxa in 7 genera in the 5 sections of the family. He concluded "floral pigments appear on this limited survey to be of little systematic utility in the Sterculiaceae." In my opinion, Scogin has underestimated the value of his data. The use of a ftavonoid score system and Hennigian phylogenetic systematics both reveal the phylogenetic content of Scogin's data and suggest that this limited survey has produced much useful information.
Results The phylogenetic status of anthocyanin pigments has been discussed in the literature for many years, most recently in a review by Gornall and Bohm [2]. The consensus of opinion is that cyanidin represents the primitive state, and delphinidin and pelargonidin represent advanced states. Similarly, as regards O-glycosylation, glucose is primitive while galactose and arabinose are advanced [2, 3]. Using this information, a quick examination of Scogin's data [1] suggests that the section Dombeyeae contains only primitive compounds (i.e. cyanidin-3-O-glucoside), while all the other sections have some advanced features; e.g. the presence of delphinidin, pelargonidin, galactose or arabinose. We can express this more meaningfully by the use of a flavonoid score system. Such systems were originally independently devised by Bate-Smith [4, 5] and da (Received 1 November 1981)
Mata Rezende and Gottlieb [6], and subsequently used by Harborne [7-9] and Campbell et al. [10]. Briefly, the method is as follows: the various flavonoid features are allocated points depending on their presumed relative advancement and each taxon is then assigned a total score dependent on the flavonoids it contains. In the case of the Sterculiaceae, it is appropriate that cyanidin and Oglucosylation are given the score 0, while pelargonidin, delphinidin, O-arabinosylation and O-galactosylation each score one point (Table 1). Thus, each section of the family accumulates the following scores: Dombeyeae 0; Fremontieae 1; Lasiopetaleae 1; Buttnerieae 2 and Sterculieae 3 (Table 2). However, the accumulated totals obscure the fact that the score for sections Frernontieae and Lasiopetaleae is due to the presence of advanced anthocyanidins, while Buttnefieae has points only for advanced OL glycosylation features; Sterculieae scoring points for both features. This information can be revealed by using a second method which is relatively little used by botanists but more widely used by zoologists. The method is Hennigian phylogenetic TABLE 1. CHARACTERSTATESOF ANTHOCYANIN FEATURESOF THE STERCULIACEAE Character state
Characters
Primitive
1 Cyanidin 2 Delphinidin 3 Pelargonidin 40-Glucosylation 50-Arsbinosylation 60-Galactosylstion
Present Absent Absent Present Absent Absent
197
Dedved (advanced) Absent Present Prtment
Absent
Present
Pr~,gt
196
P MICK RICHARDSON
TABLE 2. DATA MATRIX OF OCCURRENCE OF ADVANCED ANTHOCYANIN FEATURES IN THE SECTIONS OF THE STERCULIACEAE Character" Taxon Del Pel Ara Gal Fremont/eee Las~oetaleee Stercu#eee Buttner/eae Oomtxweee
+ + + .
+ +
-
+ .
.
+
.
"See Table1: + = present; - = absent.
systematics and produces cladograms (branched, rooted diagrams) based on the presence of shared, advanced characters. For botanical examples, see refs. [11-13], and for examples using flavonoid studies see ref. [14]. In the case of the Sterculiaceae the ctadogram is decided by the advanced anthocyanin features which are present in one or more sections of the family. The method is as follows: decide the phylogenetic status of the anthocyanin features (Table 1), produce a data matrix of the occurrence of the advanced anthocyanin features in each taxon (Table 2), and finally produce the most parsimonious cladogram from the data matrix (Fig. 1). The primitive and advanced features of the anthocyanin characters (Table 1) were decided by traditional interpretations as discussed earlier. It is more preferable to decide the primitive and advanced features by an out-group comparison, i.e. on the presence or absence of these features in the groups most closely related to the one under study. However, we do not know the anthocyanins which occur in the sister group to the Sterculiaceae, or even which family to use as a sister group. At the present time we must assume the character polarity states based on traditional interpretations are correct. In the future they can
be changed, if necessary, and a new cladogram produced. We must also assume that the sections and the family itself are monophyletic. In the cladogram (Fig. 1), sections Fremontieae and Lasiopetaleae cannot be separated because they produce the same anthocyanins. These two sections and the Sterculieaeshare the advanced feature of delphinidin production and form a monophyletic grouping. Section Sterculieae has the unique derived feature of pelargonidin synthesis and of O-arabinosylation. The latter also occurs in the Buttnerieae and is interpreted as being independently derived (a parallelism) in both sections. Section Buttnerieae has the unique derived feature of O-galactosylation while section Dombeyeaehas no advanced features, so there is a basal trichotomy. Fig. 1 is the most parsimonious ctadogram that can be derived from the data matrix: any other arrangement of the five sections would result in the production of more parallelisms, or even reversals in characters. Dkmu=don The cladogram in Fig. 1 shows which sections of the Sterculiaceae are most closely related in terms of recent common ancestry based on shared advanced anthocyanin features. The cladogram would be greatly improved by the addition of further characters, such as the distribution of other flavonoids. It could then be compared to other cladograms based on morphological or cytological features to give further insights into the phylogeny of the Sterculiaceae. Cladistic interpretations of flavonoid data such as the one presented here can readily be performed and they transform raw data into a form which can be readily understood by other taxonomists. Other chemosystematists are urged to consider the possibilities of presenting their data in a Hennigian fashion. A c k n o w l e d ~ m t e n t s - I am grateful to Chris Humphries and David Young, who criticized earlier drafts of a paper similar to this one. The study was largely supported by NSF grant DEB 79-08525 awarded to D. S. Seigter and D. A. Young, and its completion was undertaken at NYBG.
Referene~
FIG. 1. CLAOOGRAM SHOWING PHYLOGENETICRELATIONSHIPS OF SECTIONS IN THE STERCULIACEAE RASED ON ANTHOCYANIN CHARACTERS: Key: S=Stercu/~ee; F==Frem(~l~ee; L=/.ai~o~eee; B = B u t ~ ; D=Dombeyel~, 2= Delphinidin; 3 = Pe~-O~,-~din;5 -- O-Atabino~t~rt; 6 = O-Galectosylation.
1. Scogin, R. (1979)Biochem. Syst. Ecol. 7, 35. 2. Gornall, R. J. and Bohm, B. A. (1978) Syst. Bot. 3. Wallace, J. W. and Markham, K. R. (1978) Amer. 86,965. 4. Bate-Smith, E. C. (1973) Bot. J. Linn. Soc. 87, 5. Bate-Smith, E. C. and Richens, R. H. (1973)
3, 353. J. B o t a n y
347.
Biochem. Syst. 1 , 1 4 1 . 6. da M a t a Rezende, C. M. A. and Gottlieb, O. R. (1973) B i o c h e m . Syst. 1, 111.
ANTHOCYANINS OF THE STERCULIACEAE 7. Harborne, J. B. (1976) Nova Acta Leopoldina Suppl. 7, 563. 8. Harborne, J. B. (1977) Biochem. Syst. Ecol. 5, 7. 9. Smith, D. M., Glennie, C. W., Harborne, J. B. and Williams, C. A. (1977) B~ochem. Syst. Ecol. 5, 107. 10. Campbell, E. O., Markham, K. R., Moore, N. A., Porter, L. J. and Wallace, J. W. (1979) J. Hattori Bot. Lab. 45, 185. 11. Bremer, K. and Wanntorp, H.-E. (1978) Taxon27, 317.
199 12. Humphries, C. J. (1979) Bull. Br. Mus. Nat. Hist. (Bot.) 7,83. 13. Humphries, C. J. and Richardson, P. M. (1980) Chemosystematics. Principles and Practice (Bisby, F. A., Vaughan, J. G. and Wright, C. A., eds.) p. 353. Academic Press, London. 14. Richardson, P. M. (1962) Advancesin Cladistics (Funk, V. and Platnick, N., eds.) in press.