Biogeography and Chromosome Number Evolution of Linaceae
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Songyi Xu, Will A. Freyman, Adam C. Schneider, Bruce G. Baldwin Department of Integrative Biology, University of California Berkeley [email protected] Results: IntroductionIntroduction Results Figure 2: Maximum likelihood estimate of biogeographic range Linaceae is a diverse family of flowering Figure 1: Maximum a posteriori estimates of ancestral evolution for Linaceae inferred using the Dispersal-Extinction- DEC 8b on Linaceae plants, distributed throughout the temperate chromosome numbers for Linaceae inferred using the Cladogenesis (DEC+J) model.ancstates: global optim, 5 areas max. d=0.0068; e=0.0093; j=0; LnL=−200.18 ChromoSSE model. Ixonanthes_icosandra G G and tropical regions of the world. It contains Ixonanthes_reticulata EFG EFG G G EFG EFG G EG Ixonanthes_chinensis G BEFG B B Ochthocosmus_longipedicellatus B B B B Cyrillopsis_paraensis B B Hebepetalum_neblinae 13 genera and approximately 260 species B B B B Hebepetalum_humiriifolium 18 18 Linum prostratum B B Roucheria_schomburgkii 18 18 B BDGHB B B B Roucheria_monsalveae B B 183618 36 Linum macraei B B Roucheria_calophylla including 3 genera and 16 species native to Linum oligophyllum H H Hugonia_jenkinsii 18 1818 18 BDGHBDGH 18 36 Linum littorale H H Durandea_pentagyna 9 18 G H H H H Hugonia_penicillanthemum H H 18 18 Cliococca selaginoides DGH DGH H H Durandea_neocaledonica 9 9 F F Hugonia_mystax California. Using a fossil calibrated phylogeny, 15 15 Linum comptonii D DF 15 15 D D Hugonia_longipes 15 15 Linum gracile H H D D Hugonia_orientalis 10 15 DD Linum sect. Linopsis D D D D Hugonia_castaneifolia 9 9 15 15 Linum africanum DD D D Hugonia_gabunensis 10 15 Linum sulcatum D D we examined the historical biogeography and 9 9 D D D D Hugonia_busseana 9 18 Linum rupestre DG D D Hugonia_obtusifolia 9 9 D DDD Hugonia_spicata 9 9 DD 15 15 Linum vernale Hugonia_platysepala 10 15 C BCE DD G G Philbornea_magnifolia 15 15 Linum berlandieri G G chromosome evolution of Linaceae and tested 9 18 Linum striatum G G Indorouchera_griffithiana 9 9 E EFG Reinwardtia_indica 9 13 Linum neomexicanum E E Tirpitzia_sinensis E E E E E E Tirpitzia_ovoidea 18 18 Hesperolinon congestum E E previously published biogeographic and 18 18 EFG EFG Anisadenia_pubescens 18 18 Hesperolinon californicum E EFG EFGEFG Anisadenia_saxatilis 18 18 E EFG 9 9 18 18 Hesperolinon breweri EEF Anisadenia_khasyana Chromosome Number 18 17 C C Linum_pubescens 18 18 Hesperolinon clevelandii C C Linum_viscosum 17 17 C C ancestral chromosome number hypotheses. Hesperolinon disjunctum C C C C Linum_hypericifolium 10 17 17 C C 17 18 C C Linum_hirsutum 17 17 Hesperolinon didymocarpum Hesperolinon E E Linum_stelleroides 17 17 C C Hesperolinon bicarpellatum C C Linum_grandiflorum 18171717 C CE C CE E E Linum_mesostylum Hesperolinon adenophyllum CE CE 18 18 C C Linum_leonii 20 9 181718 C C 18 18 Hesperolinon spergulinum C CE C C Linum_narbonense 18 18 Linum_decumbens 18 18 Hesperolinon micranthum C C C C H H Linum_monogynum C C H H 17 17 Hesperolinon tehamense H H Linum_marginale C C C CH 8 9 ABCEABCE Linum_usitatissimum 30 15 15 Linum flavum C ABCE 14 14 Linum campanulatum CC Linum_bienne Materials and Methods 14 141414 C CE Linum_nervosum Materials and Methods C Linum_alpinum 14 14 Linum arboreum C C C 14 14 C C A A Linum_pratense 14 14 Linum mucronatum C C E AE 14 14 E E Linum_pallescens 9 13 C CE CECE Linum_austriacum 14 14 Linum album E CE 40 EE Linum_altaicum • 13 13 Linum nodiflorum CE ACEF We acquired information about the 9 9 A A Linum_lewisii Linum trigynum ACEFACEFACEFACEF Linum_perenne 10 10 ACEFACEF 10 10 Linum_komarovii 8 8 10 10 Linum tenue Linum sect. Syllinum EE 9 10 C CD Radiola_linoides biogeographic origins of the different 10 10 Linum maritimum C CE Linum_catharticum 9 9 D D Linum_volkensii 9 9 Linum tenuifolium C CD Posterior Probability CD CD Linum_strictum 9 9 C C 18 36 Linum suffruticosum C C Linum_setaceum 9 9 C C 8 8 C C C C Linum_tenuifolium 0.00 9 9 Linum setaceum C C species from the Global Biodiversity 9 9 C C Linum_suffruticosum 9 27 Linum volkensii C C Linum_maritimum 9 9 C C C C C C Linum_trigynum 9 9 Linum strictum C C C C C C Linum_tenue 0.25 C C 8 8 Linum catharticum C C Linum_nodiflorum Information Facility and Tropicos, and Linum_mucronatum C C 8 9 Radiola linoides C C C C C C Linum_album 43 43 Linum monogynum C C 0.50 14 42 C CE Linum_arboreum 42 42 Linum marginale C CC CE Linum_flavum 9 14 CC chromosome numbers from the 15 15 Linum usitatissimum C C Linum_campanulatum A A Linum_striatum 15 15 C AC 0.75 9 9 15 15 Linum bienne A A A A Linum_neomexicanum A A 9 9 9 9 Linum decumbens A A Linum_kingii A A Linum_sulcatum A A Chromosome Counts Database 1.00 9 14 Linum narbonense E A A A A Linum_rupestre A A 9 9 A A Linum_vernale 9 9 Linum mesostylum A A 8 9 A A Linum_berlandieri 9 9 9 Linum leonii A 9 8 AABD D D Linum_africanum • 8 8 Linum grandiflorum C D DD D Linum_comptonii We used multiple maximum likelihood DD D D Linum_gracile 9 9 Linum perenne D D 9 9 Linum sect. Linum BDBD D D Linum_esterhuysenae 9 99 9 Linum komarovii F A A B B Cliococca_selaginoides 9 9 9 9 Linum lewisii B B B B Linum_littorale models of geographic range evolution that 9 9 B B B B Linum_oligophyllum 9 9 Linum austriacum G B B B B Linum_prostratum 9 9 B B 9 9 9 9 Linum altaicum B B Linum_macraei D A A Sclerolinon_digynum 9 9 Linum pratense A A Hesperolinon_drymarioides 8 9 A A represented different biogeographic 9 99 9 Hesperolinon_tehamense 9 9 Linum pallescens B A A A A AA Hesperolinon_clevelandii 9 9 Linum alpinum AA 9 9 H AA Hesperolinon_sharsmithiae AA 9 15 Linum nervosum A AA A AA Hesperolinon_disjunctum scenarios in the R package BioGeoBears. 8 8 AA Hesperolinon_breweri 9 9 Linum stelleroides AAA Hesperolinon_congestum AA 8 12 Linum hypericifolium AA A Hesperolinon_californicum 8 8 A A Hesperolinon_spergulinum Linum hirsutum A A 8 8 8 8 A A Hesperolinon_micranthum • Linum sect. Dasylinum AA Hesperolinon_adenophyllum Linum pollen fossil observations were used 8 8 8 8 Linum viscosum A A A AAA Hesperolinon_didymocarpum 8 8 Linum pubescens AA to cross-validate our biogeographic range AA Hesperolinon_bicarpellatum 38 35 30 25 20 15 10 5 0 evolution results (Cavagnetto, 1996). 38 35 30 25 20 15 10 5 0 80 60 40 20 80 60 40 20 00 Millions of years ago • We used Bayesian models of anagenetic and Million of years ago Million of years ago cladogenetic chromosome number evolution to estimate both the mode of chromosome Discussion and ConclusionDiscussion and Conclusion evolution and ancestral chromosome numbers in the software RevBayes. Biogeography Findings: The Likelihood Ratio Test and Akaike information criterion indicate vicariance and long distance dispersal. We found support for a dispersal origin of Hugonoidaeae in South America approximately 45 million years ago and a North America founder speciation event for Linum sect. Linopsis from Acknowledgements Acknowledgements Europe about 35-40 million years ago. We found evidence that all yellow-flowered flax in North America results from a single colonization from Europe or Funding: Undergraduate Research Apprenticeship Program, UC the Mediterranean region approximately 11.8 million years ago. The ancestor of the blue-flowered flax originated from Europe or the Mediterranean Berkeley region approximately 24 million years ago. ReferencesReferences A. C. Schneider, W. A. Freyman, C. M. G., Y. P. S., B. G. Baldwin. 2016. Chromosome Number Evolution: Pleistocene radiation of the serpentine-adapted genus Hesperolinon and other The estimated ancestral chromosome numbers for Linum sect. Linopsis is n=9, Linum sect. Linum is n=9, and Linum sect. Dasylinum is n=8, supporting divergence times in Linaceae (Malpighiales), American Journal of Botany 103(2):221-232. previously proposed hypotheses of ancestral chromosome numbers (McDill, 2009). The ancestral chromosome number for all of Linum sensu lato is 8. J. McDill, R. M., B. B. Simpson, J. W. Kadereit. 2009. The Phylogeny of Linum and Linaceae Subfamily Linoideae, with Implications for Their Systematics, Though repeated polyploidization events are reconstructed, particularly in Linum sect. Linopsis, all polyploidization events occurred relatively recently (<5 Biogeography, and Evolution of Heterostyly. Systematic Botany 34(2):386-405. Ma). 21% of all speciation events in Linoideae are associated with dysploid chromosome gains, while only 7% of all speciation events are associated with C. Cavagnetto, and P. Anadon. 1996. Preliminary palynological data on floristic and climatic changes during the Middle Eocene–Early Oligocene of the polyploidy. These results suggest that cladogenetic dysploidization played a greater role than cladogenetic polyploidization during the diversification of eastern Ebro Basin, northeast Spain. Review of Palaeobotany and Palynology 92 : 281-305 . Linaceae. .