Ana səhifə

Arrigo et al. American Journal of Botany 100(8): 1672-1682. 2013. Data Supplement S2-Page


Yüklə 66.5 Kb.
tarix26.06.2016
ölçüsü66.5 Kb.

Arrigo et al.—American Journal of Botany 100(8):1672-1682. 2013.—Data Supplement S2—Page

Arrigo, Nils, James Therrien, Cajsa Lisa Anderson, Michael D. Windham, Christopher H. Haufler, and Michael S. Barker. 2013. A total evidence approach to understanding phylogenetic relationships and ecological diversity in Selaginella subg. Tetragonostachys. American Journal of Botany 100(8):1672-1682.
Appendix S2. Divergence time estimations: details about the synapormorphies used for the placement of fossils and settings of the BEAST analysis.
Fossil placements
Four age constraints from the fossil record were used. The split between Selaginella and its sister group Isoetes occurred at least 370 mya, based on the isoetalean Lepidosigillaria (Bateman et al., 1992; Kenrick and Crane, 1997) found in the Upper Devonian (Late Givetian–Early Famennian). According to the latest international chronostratigraphy adopted in 2012, it may have occurred as long ago as 382 mya. However, the analyses in this study were based on ages from Gradstein et al. (2004) and done before the ICS adopted the new timescale. The age of Lepidosigillaria was used as fixed age in the PL and PATHd8 analyses (but as a minimum constraint in BEAST; see below). The other three fossils were used as minimum age constraints.

A number of heterosporous fossils from Carboniferous with dimorphic, decussate leaves are assignable to the stem group of the rhizophoric clade: Selaginella suissei from the Duckmantian (Thomas, 1997), S. gutbieri from the Westphalian, and S. primaeva from the middle Bolsovian. The rhizophoric clade is recognized by two synapomorphies: decussate sporophylls and rhizophores. Besides these two characters, microphyll dimorphism is a feature that occurs only in this clade, but it is unclear whether uniform or dimorphic leaves are the plesiomorphic condition. In the absence of rhizophores in selaginellalean fossils and scarce reports on sporophyll arrangements, dimorphic leaves were chosen as the diagnostic feature of this node. The minimum age assigned to the rhizophoric clade and, hence, crown group Selaginellaceae, was therefore set to 310 Ma, based on the first occurrence of S. suissei.



Selaginella anasazia (Ash, 1972), a fossil belonging to the stem group of the clade consisting of the species S. remotifolia, S. kraussiana, S. sericea, S. articulata, S. lingulata, S. diffusa, S. sulcata, S. suavis, and S. fragilis, has been found in the Late Triassic. Selaginella anasazia has a bistelic stem, a feature that occurs only in this clade, and the fossil provides a minimum age of 210 Ma.

Megaspore fossils of the genus Erlansonisporites have been reported from many localities and range from Triassic to Cretaceous in age (Takahashi et al., 2001). They are easily distinguished by an ordered, grid-like exospore wall structure, a feature seen only in extant species in clade A in the Selaginella phylogeny. Erlansonisporites scanicus has, in addition, a proximal pole (Takahashi et al., 2001) most closely resembling megaspores of the extant S. gracillima, S. pygmaea, S. lyallii, and S. polymorpha, where the laesurae become highly convoluted and, together with interradial sculptural elements, form a complex mass at the pole (Korall and Taylor, 2006). The combination of cross-section structure and proximal pole sculpture is found only in a small clade of three species in the extant phylogeny, including S. lyallii, S. polymorpha, and S. moratii (Korall and Taylor, 2006). Erlansonisporites scanicus thus gives a minimum age constraint for the stem group of these three species of 85 Ma.


BEAST analysis
As a test of stability, phylogenetic reconstruction and divergence time estimation were also analyzed using BEAST version 1.7.3 (Drummond and Rambaut, 2007). The heterogeneous data set caused the initial UPGMA starting tree to have zero likelihood, and the dated MrBayes/PL analysis was therefore used as a starting tree. The GTR+G substitution model and the uncorrelated lognormally distributed clock model (Drummond et al., 2006) were used, and monophyly of groups constrained by fossils was enforced. Fifty million generations were run and logged every 1000 generations. Convergence and effective sample size (ESS) for parameters were checked with Tracer version 1.5, with a burn-in of 10%. Median ages and credibility intervals (CI) were calculated using TreeAnnotator.

The prior distributions for fossil-constrained ages are always one of the main problems when setting up a BEAST analysis. With a clear synapomorphy-based approach (see fossil constraints above), we can tell what the minimum age for a node is, but assigning a distribution of possible ages is less trivial. The exponential prior distribution of ages implicitly suggests that it is most likely for the estimated ages to be close to the age of the fossil constraint, which may lead to an underestimate of the true age. The prior is, however, less prone to bias (e.g., than the widely used lognormal prior). For the root node (the split between Isoetes and Selaginella), we used a uniform prior of 370–420 Ma. For the internal fossil constraints an exponential prior was used, using the minimum age as offset and 10 as mean.


LITERATURE CITED

Ash S. 1972. Late Triassic plants from the Chinle formation in north-eastern Arizona. Palaeontology 15:598-618.

Bateman R. M., W. A. DiMichele, and D. A. Willard. 1992. Experimental cladistic analysis of anatomically preserved arborescent Lycopsids from the Carboniferous of Euramerica: An essay on paleobotanical phylogenetics. Annals of the Missouri Botanical Garden 79:500-559.

DiMichele, W. A. 1980. Paralycopodites Morey and Morey, from the Carboniferous of Euramerica - A reassessment of generic affinities and evolution of “Lepidodendronbrevifolium Williamson. American Journal of Botany 67:1466-1476.

Drummond, A. J., S. Y. W. Ho, M. J. Phillips, and A. Rambaut. 2006. Relaxed phylogenetics and dating with confidence. PLOS Biology 4.

Gradstein, F. M., J. G. Ogg, A. G. Smith, F. P. Agterberg, W. Bleeker, R. A. Cooper, V. Davydov, et al. 2004. A Geologic Time Scale 2004. Cambridge University Press.

Jermy A. 1990. Selaginellaceae. In K. Kramer and P. Green [eds.], The families and genera of vascular plants, vol. I. Pteridophytes and Gymnosperms, Springer, Berlin, Germany.

Kenrick, P., and P. R. Crane. 1997a. The origin and early evolution of plants on land. Nature 389:33-39.

Kenrick P. and P. Crane. 1997b. The origin and early diversification of land plants: A cladistic study. Smithsonian Institution Press, Washington, DC, USA.

Korall P. and P. Kenrick. 2002. Phylogenetic relationships in Selaginellaceae based on rbcL sequences. American Journal of Botany 89:506-517.

Korall P. and P. Kenrick. 2004. The phylogenetic history of Selaginellaceae based on DNA sequences from the plastid and nucleus: Extreme substitution rates and rate heterogeneity. Molecular Phylogenetics and Evolution 31:852-864.

Korall, P., P. Kenrick, and J. P. Therrien. 1999. Phylogeny of Selaginellaceae: evaluation of generic/subgeneric relationships based on rbcL gene sequences. International Journal of Plant Sciences 160:585-594.

Korall P. and W. A. Taylor. 2006. Megaspore morphology in the Selaginellaceae in a phylogenetic context: A study of the megaspore surface and wall structure using scanning electron microscopy. Grana 45:22-60.

Rowe, N. P. 1988. A herbaceous lycophyte from the Lower Carboniferous Drybrook Sandstone of the Forest of Dean, Gloucestershire. Palaeontology 31:69-83.

Takahashi M., P. Crane, and H. Ando. 2001. Fossil megaspores of Marsileales and Selaginellales from the upper Coniacian to lower Santonian (upper Cretaceous) of the Tamagawa formation (Kuji group) in northeastern Japan. International Journal of Plant Sciences 162: 431-439.

Thomas, B. A. 1992. Paleozoic herbaceous lycopsids and the beginnings of extant Lycopodium sens. lat. and Selaginella sens. lat. Annals of the Missouri Botanical Garden 79:623-631.

Thomas B. A. 1997. Upper Carboniferous herbaceous Lycopsids. Review of Palaeobotany and Palynology 95: 129-153.

Wikström, N., and P. Kenrick. 1997. Phylogeny of Lycopodiaceae (Lycopsida) and the relationships of Phylloglossum drummondii Kunze based on rbcL sequences. International Journal of Plant Sciences 158:862-871.



Wikström, N., and P. Kenrick. 2001. Evolution of Lycopodiaceae (Lycopsida): Estimating divergence times from rbcL gene sequences by use of nonparametric rate smoothing. Molecular Phylogenetics and Evolution 19:177-186.


Verilənlər bazası müəlliflik hüququ ilə müdafiə olunur ©atelim.com 2016
rəhbərliyinə müraciət