Phylogenetic analysis of the Triticeae (Poaceae) Frederiksen, Signe Elisabeth; Seberg, Ole Published in: Hereditas DOI: 10.1111/j.1601-5223.1992.tb00198.x Publication date: 1992 Document version Publisher's PDF, also known as Version of record Citation for published version (APA): Frederiksen, S. E., & Seberg, O. (1992). Phylogenetic analysis of the Triticeae (Poaceae). Hereditas, 116(1-2), 15-19. https://doi.org/10.1111/j.1601-5223.1992.tb00198.x Download date: 04. okt.. 2021 Hereditas 116: 15-19 (1992) Phylogenetic analysis of the Triticeae ( Poaceae) SIGNE FREDERIKSEN and OLE SEBERG Botanical Laboratory, Copenhagen University, Copenhagen, Denmark FKEDERIKSEN,S. and SEBERG,0. 1992. Phylogenetic analysis of the Triticeae (Poaceae). - Hereditas 116 15-19. Lund, Sweden. ISSN 0018-0661. Received August 2, 1991. Accepted December 24, 1991 A phylogenetic analysis, based on morphology, was made of the monogenomic genera of the Triticeae. Bruchypodiurn P. B. and Bromus L. were used as outgroups. No less than 1335 equally parsimonious trees were found (length 113, consistency index 0.33). The consensus tree does not support the monophyly of the Triticeae. Two well defined clades were found, a Hordeurn-clade and an Aegilops-clade. As the structure of the tree is only weakly supported, no taxonomic consequences are proposed. Signe Frederiksen and Ole Seberg, Botanical Luboratory, Copenhagen Unruersiiy, Gothersgade 140, DK- I123 Copenhagen K, Denmark A fundamental drawback of genome analysis is its Material and methods inability to reveal the phylogenetic relationships of the monogenomic groups, e.g., the genera defined Material from the following herbaria was studied: by having only a single genome. Even accepting BM, C, E, and K. The aim was to study 10-15 the basic principles of genome analysis, it is not different collections of each diploid species, but in possible to make a phylogenetic hypothesis taking a few cases less material was available. A few the diploid taxa into account (SEBERG1989). diploid species were, however, not available at all. Thus, the basic “building blocks” appear to be Data from Hordeum s. lat., Psathyrostachys, and largely unrelated, despite the general agreement Secale were obtained from N. Jacobsen, C. Baden, that the Triticeae is monophyletic ( MACFARLANEand G. Petersen, respectively. and WATSON 1982; WATSONet al. 1985; KEL- Diploid species representing genomically defined LOGG 1989; SEBERGet al. 1991). Unlike the genera (Table 2) were used as terminal taxa as the monogenomic taxa, the heterogenomic, polyploid genera were a priori supposed to be monophyletic taxa have a “known” history, as they are all pos- (KELLOGG1989). In cases where genera were tulated to be of allopolyploid origin. found to be morphologically heterogeneous and, BAUMet al. (1987) and KELLOGG(1989) have hence, probably non-monophyletic, they were split previously made phylogenetic hypotheses for the into smaller units. Triticeae. BAUM et al. (1987) used both the The Bromeae and the Brachypodieae were both monogenomic and the heterogenomic taxa in the used as outgroups, as they most probably are the cladistic analysis, and thus confounded the analy- closest relatives of the Triticeae ( MACFARLANE sis by including polyphyletic terminal taxa. KEL- and WATSON1982; WATSONet al. 1985; KEL- LOGG ( 1989) excluded the heterogenomic taxa LOGG 1989, 1992; SEBERGet al. 1991). Character from the phylogenetic analysis and added them as states found to be autapomorphies of terminal taxa reticulations on the final tree. Though KELLOGG’S were excluded, but a total of 31 morphological data matrix is a dear improvement in respect to characters were scored (Table 1). Most of them that of BAUMet al. (1987), both matrices suffer were binary, but a few have three or four states. from relying too heavily on the literature. All characters were treated as non-additive. 7 he present study follows the same basic Most character states were easily observed, but a scheme as KELLOGG’S(1989), but differs in taking few character states caused problems. On account more taxa into account and in a much more thor- of the occasionally incomplete material, the data ough study of herbarium specimens and live for each terminal taxon were compiled by using all plants. accessible collections. 16 s. FREDERIKSEN AND 0.SEBERG Hereditas 116 (1992) Table 1. Characters and character states Brachypodium (-) Pseudoroegneria (S) 1. Longevity perennial/annual Lophopyrum (E) 2. Life form caespitose/stoloniferous 3. Shoots extravaginal/intravaginal Australopyrum (W) 4. Leaves auricles presentiauricles absent Erernopyrum (F) 5. Sheaths margins joined/not joined on vegetative Thinopyrum (J) shoots Agropyron (P) 6. Sheaths margins joinedinot joined on culm leaves Peridictyon (?) 7. Leaf blades folded/rolled Dasypyrum (V) 8. Spikelets homospiculate/heterospiculate (Hordeum- type)/heterospiculate (Hereronfhelium- Crithodium (A) type)/heterospiculate (Kiharopyrum-type) Festucopsis (G) 9. Spikelets embedded in rachis/more or less H et her ant hel iu m (a) protruding 10. Rachis disarticulating/tough Secale (R) Bromus (-) 11. Rachis disarticulating at every nodeinot so i 12. Rachis disarticulating above/below spikelets Taeniatherum (T) 13. Inflorescence short ( < 1/3 the spikelet length)/ Crithopsis (K) internodes long ( > 1/2 the spikelet length) Psathyrostachys (N) 14. Spikelets spikelets solitary/in pair/in triplets Critesion 1 (H) 15. Spikelets sessileilong pedicellate (infl. panicle)/lat- era1 subsessile, central sessile (Hordeum- Critesion 2 (H) type)/all subsessile (infl. spikelike raceme) Hordeurn (I) 16. Rachilla disatticulating/tough Amblyopyrum (Z) 17. Glumes ventricose/keeled/setaceous ( awnlike) Sitopsis 1 (6) 18. Glumes abaxial/lateral with respect to rachis Henrardia (0) 19. Glume-apex with 2-more lobesiwith I or without dents Patropyrum (D) 20. Glumes broadest in lower half-middleiin upper half Sitopsis 2 (B) 21. Glumes at least some awnediall awnless Orrhopygium (C) 22. Glume-apex nerves divergentinerves confluent Comopyrum (M) or nearly parallel Kiharapyrum (U) 23. Glumes 2-keeledil -keeled Chennapyrum (L) 24. Florets max. 2 per spikeletimore than 2 25. Lemma-apex with 2-5 lobes/with 1 or without dents Fig. 1. Consensus tree based on 1335 equally parsimo- nious trees. Genera and genome designations as in Table 26. Lemmas at least some awnediall awnless 1, except that Critesion 1 = C. murinum, Critesion 2 = 27. Lemmas keeled/ventricose C. bogdanii and C. brachyantherum, Sitopsis 1 = S. 28. Paled-apex bifidientire sharonensis, S. bicornis, and S. speltoides, and Sitopsis 29. Caryopsis adherentifree 2 = S. searsii and S. longissima. 30. Ovary with/without appendage 3 I. Chromosomes large, some telocentrics/large. only meta- NIG86. Accordingly, different permutations of the centrics/small input matrix were run and the strict consensus trees (Nelsen) made for each. In all runs, the consensus trees were similar. The data-matrix shown in Table 2, based on the characters shown in Table 1, was used as input to Results HENNIG86 (FARRIS1988). Due to the size of the data-matrix only the heuristic algorithm (bb*) The individual trees were 113 steps long with a could be used in an attempt to find the shortest consistency index of 0.33. The consensus tree was, tree(s). The number (1335) of equally parsimo- as mentioned above, based on several different nious trees exceeded the storage ability of HEN- subsets of the 1335 equally parsimonious trees. It Herrdrtas 116 (1992) PHYLOGENY OF THE TRITICEAE 17 Table 2. OTU-Character matrix used as input to Hennig86. Genera, species and genome designations according to LOVE (1984). Concerning the characters 29 and 31 the following sources were consulted: KIMBERand FELDMAN( 1987); SCHULTZ-SCHAEFFER( 1956); CHENNAEVEERAIAH(1960); SCHULTZ-SCHAEFFERand JURASITS(1982); ENDOand GILL(1984); FREDERIKSEN (1986, 1991a,b); and SEBERG el al. (1991) Taxon Character No. ~- 1-5 6-10 11-15 16-20 21-25 26-30 31 Brachypodium (-) pinnatum 12122 22122 --214 11221 22-22 I2222 3 silvaricum 11122 22122 --214 11221 22-22 12222 3 Bromus (-) porleri I1 121 12122 --212 11221 22-22 12221 ? kalmii I1 121 l2l22 --212 12221 22222 12221 ? gracillimus 21221 12122 --212 12221 22222 12221 ? japonicum 21221 12122 --212 11221 22-21 12221 2 remot @orus ?2121 1?122 --212 12221 22222 12221 ? Psarhyrostachys (N) ‘caespitose’ spp. 11211 2?121 11?31 1312- 1--12 12112 2 ‘stolonif.’ spp. 12?11 2?121 11?31 1312- 1--12 121 12 2 Critesion (H) murinum 21212 22222 21134 -1121 1--l2 12112 2 bogdanii 1222 2222? ?I 133 -312- 1--12 121 12 2 brachyanrherum 1222 22221 11133 -312- 1--12 12112 2 Hordeurn (I) uulgare ssp. spontaneum 1212 22221 11133 -1121 1--12 12112 2 Lophopyrum (E) elongatum 1212 22122 --21 I 11221 22-22 21212 2 Pseudoroegneria (S) libanotica -212 22 I22 --211 11221 22-22 222?2 2 tauri -212 2? I22 --21 I 11221 22-22 12222 ? sripifolia 1121? 2?122 --211 11221 22 - 22 222?2 2 srrigosa IIII? 22122 --211 11221 22 - 22 I22?2 ? spkata 11112 21122 --211 11221 12-22 122?2 2 Thinopyrum (J) bessarahieum 1212? 2?121 1121 I 11221 22-22 22222 2 Agropyron (P) pectiniforme 11211 21122 --Ill 12221 I2222 Illl2 2 Ausrralopyrum (W) uelurinum 11211 22122 --Ill 12221 22222 11212 2 retrofracrum 1 I21 I 22122 --Ill 12221 22222 12212 2 pectinatum 11211 22122 --I14 I222 1 22222 11212 2 Eremopyrum (F) disrans 2121 I 22121 11111 22221 12222 121 12 1 rririceum 2121 1 22 I22 --Ill 1222 1 12222 121 I2 1 Peridictyon (?) sanctum 11211 21122 --214 11221 22 - 22 12212 2 Festucopsis (G) serpentini 11221 22122 --214 11221