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Convergent fruit evolution in Athysanus and Thysanocarpus (Brassicaceae)

Convergent fruit evolution in Athysanus and Thysanocarpus (Brassicaceae). P.J. Alexander (1) , I.A. Al-Shehbaz (2) , G. Rajanikanth (1) , C.D. Bailey (1) , and M.D. Windham (3) (1) New Mexico State University; (2) Missouri Botanical Garden; (3) University of Utah. Introduction

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Convergent fruit evolution in Athysanus and Thysanocarpus (Brassicaceae)

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  1. Convergent fruit evolution in Athysanus and Thysanocarpus (Brassicaceae) P.J. Alexander (1) , I.A. Al-Shehbaz (2), G. Rajanikanth (1), C.D. Bailey (1), and M.D. Windham (3) (1) New Mexico State University; (2) Missouri Botanical Garden; (3) University of Utah

  2. Introduction In the mustard genus Athysanus, fruit and trichome morphology give conflicting evidence of generic affinities. Its unusual fruit morphology is shared with Thysanocarpus, which is similar in other respects as well. However, trichome morphology differs between the two genera, and suggests a closer affinity of Athysanus to genera such as Heterodraba,Draba, or Alyssum. We generated sequence data from ITS and trnL to discover the generic affinities of Athysanus, Heterodraba, and Thysanocarpus and better understand evolution in the group.

  3. Fruits & trichomes

  4. Thysanocarpus: • six species, Thysanocarpus curvipes & T. laciniatus widespread • white-flowered desert annuals in the SW US & adjacent Mexico • fruits with 1 carpel, 1 seed, indehiscent, winged • glabrous or with sparse, simple trichomes

  5. Athysanus: • monotypic, Athysanus pusillus • small, white-flowered desert annual in western North America from southern British Columbia to Baja California • fruits with one carpel, 1 seed, indehiscent • pubescent with simple and rayed trichomes

  6. Heterodraba: • monotypic, Heterodraba unilateralis • small, white-flowered desert annual in California and S Oregon • fruits with 2 carpels, 6-12 seeds, +/- indehiscent • pubescent with simple and rayed trichomes

  7. Athysanus affinities? • Thysanocarpus is placed in Schizopetaleae with weak support in ITS trees (Bailey et al., in review; Al-Shehbaz et al., 2006) • Athysanus & Heterodraba not previously investigated with genetic data; both placed in Alysseae (Schulz, 1936) or Arabideae (Al-Shehbaz, 2006) based on morphology • So: is Athysanus related to Thysanocarpus, as suggested by fruit morphology, or within one of the branched-trichome clades like Alysseae or Arabideae?

  8. Methods We generated ITS and trnL sequence data for these three genera and several outgroups; We downloaded ITS and trnL sequences from GenBank; We aligned sequences using with ClustalW (Thompson et al., 1994); We scored gap characters with GapCoder (Simmons et al., 2001; Young & Healy, 2003); 5. We produced maximum parsimony trees with NONA (Goloboff, 2000) via WinClada (Nixon, 1999-2002).

  9. Aethionema arabicum Biscutella laevigata Phylogeny of Brassicaceae from ITSdata. Consensus of 58 most parsimonious trees. WinClada settings: hold= 25001 mult= 2500 hold/= 10 Support estimated from 100 bootstrap replicates with 100 mults each. 9 6 Alyssum alyssoides Aurinia saxatile Neotorularia torulosa 4 8 Lobularia maritima 3 6 1 0 0 Noccaea montana Microthlaspi perfoliatum 3 4 Notothlaspi australe Sisymbrium loesellii 9 8 Sisymbrium irio Thellungiella salsuginea 9 2 9 8 Eutrema wasabi Pseudoturritis turrita 6 4 Scolixon mexicanus Arabis hirsuta 4 0 6 0 Arabideae 70 Arabis alpina 94 Draba mogollonica 98 Draba tomentosa 2 0 94 Heterodraba unilateralis Athysanus pusillus Sinapidendron angustifolium Eruca pinnatifida 9 8 Zilla macroptera Brassica nigra 6 8 6 2 Raphanus sativus Hemicrambe fruticulosa 1 0 0 Matthiola incana 4 8 Hesperis matronalis 2 0 4 0 Heliophila arenaria 7 6 Iberis amara Chamira circaeoides Sibara rosulata Sisymbrium andinum Fourraea alpina 32 Thelypodiopsis vaseyi Schizopetaleae Isatis tinctoria Stanleya pinnata 7 0 Caulanthus inflatus 8 4 Streptanthus cordatus 82 Thysanocarpus laciniatus 94 Thysanocarpus radians Thysanocarpus curvipes Barbarea vulgaris 1 0 0 Cardamine hirsuta 8 2 Nasturtium officinale 1 0 0 Rorippa officinale Ianhedgea minutiflora 7 8 1 0 0 1 0 0 Descurainia californica Descurainia pinnata Smelowskia johnsonii 1 0 0 6 0 Smelowskia americana 7 2 Lepidium oleraceum 1 0 0 Lepidium virginianum 1 0 0 Thlaspi arvense 4 2 Capsella bursa-pastoris 9 2 Camelina microcarpa Olimarabidopsis pumila 7 2 Physaria fendleri 1 0 0 Nerisyrenia linearifolia 1 0 0 2 8 Lyrocarpa coulteri Arabidopsis halleri ssp. halleri 1 0 0 Arabidopsis thaliana 2 8 Transberingia bursifolia Boechera perennans 7 0 4 4 Nevada holmgrenii 6 2 Boechera laevigata Sphaerocardamum macropetalum 9 8 Mancoa pubens 1 0 0 Halimolobos diffusa

  10. Streptanthella longirostris Phylogeny of Thysanocarpus from combined ITSand trnL data. Consensus of 149 most parsimonious trees. WinClada settings: hold= 25001 mult= 2500 hold/= 10 Support estimated from 100 bootstrap replicates with 100 mults each. Thysanocarpus conchuliferus Thysanocarpus laciniatus 93 Thysanocarpus laciniatus var. crenatus 93 T. laciniatus Thysanocarpus laciniatus var. laciniatus 1 Thysanocarpus laciniatus var. laciniatus 2 82 Thysanocarpus laciniatus var. laciniatus 3 Thysanocarpus radians 1 100 T. radians Thysanocarpus radians 2 Thysanocarpus radians 3 Thysanocarpus curvipes var. eradiatus 1 Thysanocarpus curvipes var. eradiatus 3 89 Thysanocarpus curvipes 3 Thysanocarpus curvipes var. eradiatus 2 Thysanocarpus curvipes 5 Thysanocarpus curvipes 4 96 Thysanocarpus curvipes 1 T. curvipes Thysanocarpus curvipes var. elegans 2 Thysanocarpus curvipes var. elegans 1 Thysanocarpus curvipes var. curvipes Thysanocarpus curvipes 2 Thysanocarpus curvipes var. elegans 4 83 Thysanocarpus curvipes var. elegans 3 75 Thysanocarpus curvipes var. elegans 5

  11. Thysanocarpus in Schizopetaleae... • Apart from Thysanocarpus, Schizopetaleae includes exclusively plants with long, narrow, bicarpellate, many-seeded fruits. • Most members of the tribe are similar to Thysanocarpus in being glabrous or with only sparse, simple trichomes. The tribe also includes a number of taxa with similar ecology.

  12. Streptanthella longirostris Pseudoturritis turrita Draba Draba muralis Draba hystrix Aubrieta deltoidea 76 70 Draba funiculosa Arabis alpina 62 Arabis bryoides 98 Arabis procurrens 23 Arabis 79 53 Arabis hirsuta Phylogeny of Arabideae from ITSdata. Consensus of 59694 most parsimonious trees. WinClada settings: hold= 100001 mult= 10000 hold/= 10 Support estimated from 100 bootstrap replicates with 100 mults each. Arabis soyeri 47 Arabis scabra 57 Arabis flagellosa 94 Arabis nuttallii 55 Arabis blepharophylla 24 § Erophila 100 Draba spathulata Draba verna 37 Draba species Draba reptans Draba 99 § Tomostima Draba cuneifolia 47 Draba araboides Draba platycarpa 79 Draba jaegeri 30 Draba ramosissima Draba hitchcockii 65 Draba pterosperma Draba nemorosa 9 92 Heterodraba unilateralis 31 Athysanus pusillus Draba rigida 70 Draba bruniifolia 67 Draba lasiocarpa 53 Draba cryophila 82 Draba bellardii Draba aizoides 55 Draba cuatrecasana Draba elata 4 Draba yunnanensis 88 Draba surculosa 9 Draba species 11 Draba koeiei 98 Draba olgae Draba radicans 32 Draba aubrietioides Draba setosa 68 1 Draba cachemirica 68 Draba oreades Draba asprella Draba Draba juniperina Draba crassifolia 29 Draba densifolia 23 Draba pulvinata 45 18 Draba pycnophylla Draba brachystylis 8 Draba species 5 Draba sobolifera 63 Draba subalpina 32 Draba species 6 Draba barclayana 41 Draba standleyi Draba boyacana Draba confertifolia 3 Draba stylosa 60 Draba violacea Draba depressa Draba pickeringii Draba farsetioides 20 Draba litamo Draba hallii

  13. Streptanthella longirostris Arabis alpina Draba hystrix Arabis Draba 98 100 Arabis blepharophylla 47 Arabis soyeri 39 Aubrieta deltoidea Pseudoturritis turrita 93 Draba muralis 99 Heterodraba unilateralis 42 Athysanus pusillus Draba cuneifolia §s Abdra & Tomostima Draba reptans 34 Draba platycarpa Phylogeny of Arabideae from trnL data. Consensus of 37225 most parsimonious trees. WinClada settings: hold= 100001 mult= 10000 hold/= 10 --final swapping after mult completion was terminated early Support estimated from 100 bootstrap replicates with 100 mults each. Draba araboides 61 Draba brachycarpa 25 Draba species Draba stylaris 83 Draba carinthiaca Draba ladina Draba farsetioides 52 89 Draba fladnizensis 99 Draba aizoides 75 Draba cuatrecasana Draba pennell hazenii Draba laurentiana 51 Draba hirta 59 24 Draba arseniewii Draba lanceolata Draba lactea 65 Draba alpina 80 Draba winterbottomii Draba altaica Draba radicans 59 47 Draba macbeathiana Draba alberti 85 Draba nemorosa Draba hispida 79 Draba lasiocarpa 21 100 Draba lutescens 58 Draba huetii 3 Draba jucunda 73 Draba gracillima 93 Draba ellipsoidea Draba eriopoda Draba hyperborea 72 Draba aubrietioides 39 Draba stenocarpa Draba trinervis 58 53 93 Draba olgae Draba 0 Draba odudiana § Erophila Draba longisiliqua 30 90 Draba verna 11 Draba hederifolia Draba mollissima 87 Draba heterocoma 94 Draba cuspidata 63 Draba natolica 25 Draba scardica 41 Draba hispanica 8 Draba cretica 12 Draba oreadum 24 68 Draba parnassica Draba lacaitae 21 Draba species 91 Draba streptocarpa Draba rositae Draba incana 51 Draba spectabilis 14 97 Draba jaegeri 70 Draba incerta Draba magellanica 76 Draba norvegica Draba kotschyi 14 Draba breweri Draba aurea 6 Draba gilliesii Draba tucumanensis 43 Draba paysonii Draba hitchcockii Draba glabella Draba athoa Draba pusilla Draba aureola 40 Draba species 99 Draba juniperina 23 18 Draba micropetala Draba stenopetala 48 Draba wurdackii 62 24 Draba alyssoides Draba lapaziana 38 Draba obovata Draba pauciflora 13 58 Draba subalpina Draba kassii 47 Draba melanopus Draba bhutanica

  14. Phylogeny of Arabideae from combined ITS and trnL data. Streptanthella longirostris Draba muralis Draba Arabis alpina Draba hystrix 73 Aubrieta deltoidea 78 Pseudoturritis turrita Arabis bryoides 100 Arabis procurrens 59 47 Arabis soyeri Arabis hirsuta 41 Arabis Consensus of 100001 most parsimonious trees. WinClada settings: hold= 100001 mult= 10000 hold/= 10 Support estimated from 100 bootstrap replicates with 100 mults each. Arabis scabra 57 Arabis flagellosa 94 67 Arabis blepharophylla Arabis nuttallii Heterodraba unilateralis 100 Athysanus pusillus Draba hederifolia 73 § Erophila Draba verna 89 Draba spathulata 157 Draba species Draba globosa 39 41 Draba kassii Draba burkei 54 Draba incana 50 Draba jaegeri 88 Draba incerta Draba magellanica Draba bellardii 71 Draba cryophila 48 39 67 Draba aizoides Draba cuatrecasana Draba Draba lutescens Draba huetii 12 Draba lasiocarpa Draba rigida Draba bruniifolia 39 Draba brachycarpa 100 Draba reptans Draba platycarpa 68 §s Abdra & Tomostima 50 Draba cuneifolia Draba araboides Draba mollissima 77 Draba heterocoma 83 Draba cuspidata 58 Draba natolica 18 Draba scardica 32 Draba hispanica 8 Draba cretica 16 Draba oreadum 20 56 Draba parnassica Draba lacaitae

  15. Athysanus and Draba • Although some Draba have shortened, nearly orbicular fruits, the single-seeded, unicarpellate fruits of Athysanus are unique in Arabideae. • In other gross features, Athysanus is quite dissimilar from most Draba; most Draba are alpine to subalpine plants, typically perennial and yellow-flowered.

  16. Athysanus and Draba • The only New World Draba that are white-flowered and annual are in the Abdra/Tomostima clade, which also includes the New World’s only arid and semiarid Draba. • Old World white-flowered annual Draba include § Erophila. • Our analyses place either § Erophila or §s Abdra & Tomostima basal or sister to the remaining Draba, but resolution is insufficient to allow analysis of evolution of these characters.

  17. Conclusions • Fruit similarities between Athysanus and Thysanocarpus represent convergent evolution. Trichomes better predict relationships in this case. • Athysanus is sister to Heterodraba, and the two are either nested within or sister to Draba, depending on the data set used. • Thysanocarpus is weakly supported as a member of Schizopetaleae, but does not show a close relationship with any other genera within the tribe.

  18. The Future: Beyond Phylogeny • In addition to Athysanus and Thysanocarpus, Lepidium, Idahoa, Lunaria, Alyssum, Thlaspi, and Noccaea are other prominent genera with reduced fruits, all apparently independently derived, • So now we have a pattern that seems to demand explanation! • Some possibilities: • reduced per-fruit investment allows quicker reproduction and better fine-tuning of fruit production to resource availability • fewer seeds/fruit may mean larger seeds, which may provide rapid seedling establishment needed in temporally- limited habitats • fewer seeds/fruit may have dispersal advantages by lowering the density of fruits • few-seeded fruits with large photosynthetic area may be able to “pay for themselves”

  19. Cited works: • Al-Shehbaz IA, Beilstein MA, Kellogg EA (2006) Systematics and phylogeny of the Brassicaceae (Cruciferae): an overview. Plant Systematics and Evolution 259, 89-120. • Bailey CD, Koch MA, Mayer M, Mummenhoff K, O’Kane SL, Warwick SI, Windham MD, Al-Shehbaz IA (2006) A Global nrDNA ITS Phylogeny of the Brassicaceae. Molecular Biology and Evolution, in review. • Bailey CD, Price RA, Doyle JJ (2002) Systematics of the Halimolobine Brassicaceae: Evidence from Three Loci and Morphology. Systematic Botany27, 318-332. • Beilstein MA, Al-Shehbaz IA, Kellogg EA (2006) Brassicaceae phylogeny and trichome evolution. American Journal of Botany93, 607-619. • de Candolle AP (1821) Cruciferae. Systema Naturale2, 139-700. • Goloboff PA (2000) NONA (NO NAME) ver. 2. Published by the author. • Koch M, Mummenhoff K (2001) Thlaspi s.str. (Brassicaceae) versus Thlaspi s.l.: morphological and anatomical characters in the light ot ITS nrDNA sequence data. Plant Systematics and Evolution227, 209-225. • Nixon KC (1999-2002) WinClada ver. 1.0000. Published by the author. • Schulz OE (1936) Cruciferae. In: Die natürlichen Planzenfamilien eds. Engler A, Harms H). Wilhelm Engelmann, Leipzig, Germany. • Simmons MP, Ochoterena H, Carr TG (2001) Incorporation, Relative Homoplasy, and Effect of Gap Characters in Sequence-Based Phylogenetic Analyses. Systematic Biology50, 454-462. • Thompson JD, Higgins DG, Gibson TJ (1994) CLUSTAL W: improving the sensitivity of progressive sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research 22, 4673-4680. • Young ND, Healy J (2003) GapCoder automates the use of indel characters in phylogenetic analysis. BMC Bioinformatics 4, 6. • Acknowledgements: • NMSU Dept of Biology and NSF EF-0542228 (CDB)

  20. Questions?

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