BambooBamboo EvolutionEvolution andand Classification:Classification: RecentRecent AdvancesAdvances Lynn G. Clark Iowa State University U.S.A. MajorMajor TopicsTopics •• BamboosBamboos areare grasses!grasses! •• BambooBamboo evolutionevolution andand classificationclassification—— updatesupdates •• CurrentCurrent directionsdirections BamboosBamboos areare grasses!grasses! theclyde, Flickr L Attigala What makes a grass a grass? Flagellariaceae Joinvilleaceae Ecdeiocoleaceae Poaceae (11,000+ species) Micro-canals in the tectum trnT inversion Grass-type embryo in in the LSC region of lateral position and the chloroplast caryopsis fruit type Crepet & Feldman 1991 Ancestral grasses Origin of the grasses 70-80 mya in forests of the S hemisphere Silica bodies (phytoliths) Prasad et al. 2005 Anomochlooideae Pharoideae Puelioideae Bambusoideae Origin of the (bamboos) grass-type spikelet Pooideae and lodicules (bluegrasses, wheat) (petals) Ehrhartoideae = Spikelet (rices and allies) clade Aristidoideae (wiregrasses) Panicoideae (maize, panic grasses) Chloridoideae (love grasses) Danthonioideae (pampas grasses) Micrairoideae Arundinoideae (reeds) Anomochlooideae Pharoideae Puelioideae Bambusoideae (bamboos) Loss of a stigma (3 to 2); Pooideae (bluegrasses, wheat) spikelets with multiple Ehrhartoideae florets (rices and allies) Aristidoideae (wiregrasses) Panicoideae (maize, panic grasses) Chloridoideae (love grasses) Danthonioideae (pampas grasses) Micrairoideae Arundinoideae (reeds) Ancestral or early-diverging grasses (Anomochlooideae, Pharoideae, Puelioideae) Clayton 1969 EJ Judziewicz Anomochlooideae Pharoideae Puelioideae BEP Clade Bambusoideae (bamboos) Pooideae (bluegrasses, wheat) Ehrhartoideae (rices and allies) Aristidoideae Stamens (wiregrasses) reduced to 3; Panicoideae + 55 mya (maize, panic grasses) Chloridoideae (love grasses) PACMAD Clade Danthonioideae (pampas grasses) Oldest grass macrofossil Micrairoideae ca. 55 mya Crepet & Feldman 1991 Arundinoideae (reeds) Ancestral grasses (30) BEP Clade Initial (5000 species) Bambusoideae diversification (1400) 55-35 mya Pooideae (Eocene) (3300) in forests and Ehrhartoideae then open (120) habitats Aristidoideae* (350) Initial diversification Panicoideae* (3300) 35-28 mya (Oligocene- Chloridoideae* Miocene) (1400) in open habitats Danthonioideae (250) PACMAD Clade (5500 species); Micrairoideae* (155) C4 photosynthesis* Arundinoideae (40) Ancestral grasses (30) BEP Clade ca. 30 mya (Bouchenak-Khelladi et al. 2010) Initial (5000 species) Bambusoideae diversification (1400) 55-35 mya Pooideae (Eocene) (3300) in forests and Ehrhartoideae then open (120) habitats Aristidoideae* (350) Initial diversification Panicoideae* (3300) 35-28 mya (Oligocene- Chloridoideae* Miocene) (1400) in open habitats Danthonioideae (250) PACMAD Clade (5500 species); Micrairoideae* (155) C4 photosynthesis* Arundinoideae (40) Pharoideae Anomochlooideae Bambusoideae fusoid cell March & Clark 2011 Olyra ciliatifolia in the forest understory Ancestral grasses (30) ca. 30 mya (Bouchenak-Khelladi et al. 2010) BEP Clade Bambusoideae Brachyelytrum, (bamboos) temperate Pooideae forests (wheat, bluegrasses) Ehrhartoideae (rices and allies) Aristidoideae (350) Panicoideae (3300) Chloridoideae (1400) PACMAD Clade Danthonioideae (250) Streptogyna, Micrairoideae tropical (155) forests Arundinoideae (40) BambooBamboo evolutionevolution andand classificationclassification ——updatesupdates Bamboos (subf. Bambusoideae) (>1,400 species) arm cells X Londoño The only major grass lineage to diversify in forests Bambuseae (tropical woody bamboos, 784) Olyreae (herbaceous bamboos, 122) Arundinarieae (temperate woody bamboos, 533) Three principal lineages of Bambusoideae Olyreae—herbaceous bamboos Patricia de Oliveira 21 genera, ca. 122 described species, primarily Latin American crenate (olyroid) Olyreae, lower leaf surface silica body veins stomate cruciform silica body Olyreae have unisexual spikelets and unusual adaptations in some Eremitis Pariana underground spikelets Olyra standleyi gregarious monocarpy insect pollination? Raddiella vanessae, smallest “hidden” bamboo Diandrolyra inflorescences Olyreae distribution Latitudinal diversity H. Iltis rest of tribe Olyra latifolia Buergersiochloa (1 species) Arundinarieae (temperate woody bamboos) G Reiners Temperate woody bamboos: 28 genera, 533 species De-Zhu Li, Chris Stapleton Major defining characters: •Leptomorph, monopodial rhizomes (but pachymorph in some) •Tetraploidy (2n=48) •Basipetal vegetative branch development •Molecular evidence Temperate woody bamboos E North America Afro-Indian montane E Asia 3 species ~20 species ~510 species Diversity within Arundinarieae J Triplett J Triplett Chimonocalamus Pleioblastus Phyllostachys Indosasa J Triplett L Attigala Sarocalamus Arundinaria A. densifolia Fargesia Major lineages of the temperate clade Summary phylogenetic tree based on up to 12 cpDNA regions VI. Arundinaria clade Zeng et al., 2010 Triplett & Clark, 2010 Attigala et al., unpubl. IV. Shibataea clade V. Phyllostachys clade High support III. Chimonocalamus Moderate support VII. Thamnocalamus Low support or single species I. T. tessellatus VIII. Indocalamus wilsonii IX. Gaoligonshania X. Indocalamus sinicus II. African alpine bamboos XI. Sri Lankan alpine bamboos Lakshmi Attigala Hibanobambusa Data from J. Triplett Sasa veitchii x Phyllostachys nigra AFLPs—from both parents Chloroplast phylogeny (5 markers) Maternally inherited AFLPs Arundinaria AFLP analysis 4 primer pairs (data from J. Triplett) (River cane) Jimmy Triplett with A. gigantea (Switch cane) (Hill cane) Natural hybridization also occurs! Bambuseae (tropical woody bamboos) Tropical woody bamboos: 66 genera, 784 species Ximena Londoño, Major defining characters: Emmet Judziewicz •Pachymorph rhizomes •Tetraploidy or hexaploidy •Acropetal or bidirectional vegetative branch development •Molecular evidence Bambuseae distribution Diversity hotspots Woody bamboo diversity is strongly associated with mountains Major lineages of the tropical woody bamboos Paleotropical woody bamboos Neotropical Arthrostylidiinae + Guaduinae clade ? Neotropical Chusqueinae clade Major lineages of the tropical woody bamboos Paleotropical woody bamboos Neotropical Arthrostylidiinae + Guaduinae clade ? Neotropical Chusqueinae clade Evolutionary relationships within Chusqueinae Euchusquea clade ▪Multiple, dimorphic * buds E 138 spp ▪Connate lemma tips N1 ▪Spikelet structure 11 spp ▪Papillate * subsidiary cells N2 10 spp Amanda Fisher Key innovation in Chusqueinae? single buds but no multiple, dimorphic buds in the branch development Euchusquea clade (138 spp) in “Neurolepis” (21 spp) J Triplett single bud in most bamboos giving rise to complex branching patterns Toni Grieb Branching patterns in Chusquea (the Euchusquea clade) The Euchusquea clade shows high species diversity yet low genetic variation Low variation (>5K nucleotides, 260 PICs) Recent origin of Euchusquea clade? Data from A. Fisher BEAST software MolecularMolecular datingdating analysisanalysis ofof ChusqueaChusquea Bayesian framework chloroplast sequence data C. 1. Secondary calibration from a scandens group molecular dating analysis of Poaceae 14 – 24.8 mya (Kelchner & Fisher unpublished) 2. Neotropical woody bamboo fossil 3.6 – 11.8 mya (Brea & Zucol 2007) Data from A. Fisher MolecularMolecular datingdating analysisanalysis ofof ChusqueaChusquea C. scandens group Estimated divergence of the Euchusquea clade in the late Miocene to Pliocene (3.9-9.16 mya) Estimated divergence of Chusquea In the Miocene (10.8-21 mya) Data from A. Fisher Major lineages of the tropical woody bamboos Paleotropical woody bamboos Neotropical Arthrostylidiinae + Guaduinae clade ? Neotropical Chusqueinae clade Summary phylogeny of the V Actinocladum Arthrostylidiinae (molecular) Merostachys IV Filgueirasia Alvimia Atractantha III Aulonemia Colanthelia Leaf anatomy/ Arthrostylidium/ morphology Rhipidocladum II Elytrostachys Arthro/Rhip R. geminatum I Glaziophyton Guaduinae What did the ancestor of Arthrostylidiinae look like and where in South America did the subtribe originate? Glaziophyton Tom Soderstrom X Londoño Other Arthrostylidiinae Leaf anatomy and micromorphology in Arthrostylidiinae Ana Paula Santos-Gonçalves Describing bamboo diversity 1999: 3 species of Otatea (Guaduinae) (Judziewicz et al. 1999) + Ecological niche modeling Population genetic analysis 2011: 8 species of Otatea (Guaduinae) (Ruiz & Sosa 2010; Ruiz et al. 2011; Ruiz, in press) Major lineages of the tropical woody bamboos Paleotropical woody bamboos hexaploidy Neotropical Arthrostylidiinae + Guaduinae clade ? tetraploidy Neotropical Chusqueinae clade Paleotropical woody bamboos Racemobambosinae 4 subtribes, 47 genera, 407 species T Rodd Hickeliinae S Dransfield Jatmi Dransfield KM Wong NE India Pics Bambusinae Melocanninae Relationships within paleotropical woody bamboos Melocanninae Bambusinae & other PWB? Analyses with broader sampling are ongoing (by Chokthaweepanich et al.)…but natural hybridization is documented within Bambusinae (Goh et al., in press). SummarySummary ofof mostmost significantsignificant recentrecent advancesadvances inin bamboobamboo evolutionevolution SummarySummary ofof mostmost significantsignificant recentrecent advancesadvances inin bamboobamboo evolutionevolution • Estimated origin of the bamboos ca. 30 mya; 3 major lineages (herbaceous, temperate woody and tropical woody) SummarySummary ofof mostmost significantsignificant recentrecent advancesadvances inin bamboobamboo evolutionevolution