Protistan Systematics Ge197 Week 1 Winter 2013

Tuesday, January 22, 13 unidentified freshwater amoeba

Tuesday, January 22, 13 modified after an original by David Patterson Marine Biological Laboratory Woods Hole, Massachusetts USA

Tuesday, January 22, 13 modified after an original from Patrick Keeling University of British Columbia Vancouver, Canada

Tuesday, January 22, 13 5

Tuesday, January 22, 13 Arcella

Tubulinids

Kreutz, M. and Foissner , W. 2006: The Sphagnum ponds of Simmelried in Germany: a biodiversity hot-spot for microscopic organisms. Protozoological Monographs Vol. 3, Shaker Verlag, Aachen, Germany

Tuesday, January 22, 13 Paraquadrula

Tubulinids

Kreutz, M. and Foissner , W. 2006: The Sphagnum ponds of Simmelried in Germany: a biodiversity hot-spot for microscopic organisms. Protozoological Monographs Vol. 3, Shaker Verlag, Aachen, Germany

Tuesday, January 22, 13 Nebela vas

Tubulinids

courtesy of Wilhelm Foissner University of Salzburg, Austria

Tuesday, January 22, 13 Stramenopiles (tripartite hairs on flagella) *

Chrysophytes

opalines

Protsville Tuesday, January 22, 13 Stramenopiles (tripartite hairs on flagella) *

Chrysophytes

opalines

Cafeteria Protsville Tuesday, January 22, 13 Stramenopiles (tripartite hairs on flagella) *

Diatoms Chrysophytes

opalines

Cafeteria Protsville Tuesday, January 22, 13 Stramenopiles (tripartite hairs on flagella) *

macrophytic Diatoms brown Chrysophytes algae

opalines

Cafeteria Protsville Tuesday, January 22, 13 modified after an original from Patrick Keeling University of British Columbia Vancouver, Canada

Tuesday, January 22, 13 Chroomonas sp.

Cryptomonads

Kugrens, P. et al. 1986: Cell form and surface patterns in Chroomonas and Cryptomonas cells (Cryptophyta) as Revealed by scanning electron microscopy J. Phyc. 22: 512-522

Tuesday, January 22, 13 Emiliania huxleyi

Haptophytes

Hallegraeff, G. 1988: , a microscopic world Brill Academic Publishers, Leiden

Tuesday, January 22, 13 Pontosphaera discopora

Haptophytes

Hallegraeff, G. 1988: Plankton, a microscopic world Brill Academic Publishers, Leiden

Tuesday, January 22, 13 Ophiaster formosus

Haptophytes

Young, J. R. et al. 2009: Coccolith function and morphogenesis: insights from appendage-bearing coccolithophores of the family Syracosphaeraceae. J. Phycol. 45: 213-226

Tuesday, January 22, 13 Michaelsarsia elegans

Haptophytes

Young, J. R. et al. 2009: Coccolith function and morphogenesis: insights from appendage-bearing coccolithophores of the family Syracosphaeraceae. J. Phycol. 45: 213-226

Tuesday, January 22, 13 Synura petersenii

Synurophytes Ochrophytes

Tuesday, January 22, 13 Mallomonas sp.

Synurophytes Ochrophytes

courtesy of Yu Iokawa Joetsu University of Education Niigata, Japan

Tuesday, January 22, 13 Coscinodiscus

Diatoms

Tuesday, January 22, 13 Bacteriastrum furcatum

Diatoms

Hallegraeff, G. 1988: Plankton, a microscopic world Brill Academic Publishers, Leiden

Tuesday, January 22, 13 Stephanopyxis palmeriana

©Tamotsu Nagumo Diatoms Nippon Dental University Tokyo, Japan Image licensed to Phycological Images

Tuesday, January 22, 13 *

dinoflagellates apicomplexans

Tetrahymena Pfisteria (red tide) ( thermophila falciparum) malaria Alveolar membrane system: sacs (alveoli) underneath outer cell membrane Tuesday, January 22, 13 spinifera

Dinoflagellates

Hallegraeff, G. 1988: Plankton, a microscopic world Brill Academic Publishers, Leiden

Tuesday, January 22, 13 Ceratocorys horrida

Dinoflagellates

courtesy of Maria Faust Smithsonian National Museum of Natural History Washington, DC, USA

Tuesday, January 22, 13 schuettii

Dinoflagellates

Hallegraeff, G. 1988: Plankton, a microscopic world Brill Academic Publishers, Leiden

Tuesday, January 22, 13 Ornithocercus thumii

Dinoflagellates

courtesy of Maria Faust Smithsonian National Museum of Natural History Washington, DC, USA

Tuesday, January 22, 13 Amphisolenia bidentata

Dinoflagellates

Hallegraeff, G. 1988: Plankton, a microscopic world Brill Academic Publishers, Leiden

Tuesday, January 22, 13 Pelagostrobilidium neptuni

Choreotrichs Spirotrichs Ciliates

Agatha et al. 2004: Pelagostrobilidium neptuni and Strombidium biarmatum: phylogenetic position inferred from morphology, ontogenesis and gene sequence data. Europ. J. Prot. 41: 65-83

Tuesday, January 22, 13 Codonella cratera

Choreotrichs (Tintinnids) Spirotrichs Ciliates

Tuesday, January 22, 13 es

Armophorids Ciliates

courtesy of Christian Bardele University of Tübingen Germany

Tuesday, January 22, 13 nasutum

Haptorians Litostomes Ciliates

Foissner et al. 1995 Taxonomische und ökologische Revision der Ciliaten des Saprobiensystems, Band 4 Bazerisches Landesamt für Wasserwirtschaft Munich, Germany

Tuesday, January 22, 13 * Rhizaria

testate amoebae

chlorarachnia cercomonads

Chlorarachnion reptans Paulinella ovalis Massisteria marina “nucleomorph”

Tuesday, January 22, 13 modified after an original from Patrick Keeling University of British Columbia Vancouver, Canada

Tuesday, January 22, 13 Acanthometra

Acantharians Radiolarians

©Peter Parks, ImageQuest Ltd. Witney, Oxon, UK

Tuesday, January 22, 13 Lychnaspis miranda

Acantharians Radiolarians

Ernst Haeckel Kunstformen der Natur

Tuesday, January 22, 13 Dictyacantha

Acantharians Radiolarians

Hallegraeff, G. 1988: Plankton, a microscopic world Brill Academic Publishers, Leiden

Tuesday, January 22, 13 Hexastylus triaxonius

Spumellarians Radiolarians

www.radiolarian.com

Tuesday, January 22, 13 Lamprocyclas maritalis

Nasellarians Radiolarians

www. radiolaria.org

Tuesday, January 22, 13 Reticulomyxa filosa

Foraminiferans

Hausmann et al. 2003: Protistology, 3rd edition ©Schweizerbart’sche Verlagsbuchhandlung

Tuesday, January 22, 13 Globigerina bulloides

Foraminiferans

Hallegraeff, G. 1988: Plankton, a microscopic world Brill Academic Publishers, Leiden

Tuesday, January 22, 13 Globigerinella sp.

Foraminiferans

©O. Roger Anderson, image licensed to micro*scope

Tuesday, January 22, 13 modified after an original from Patrick Keeling University of British Columbia Vancouver, Canada

Tuesday, January 22, 13 Giardia lamblia

Diplomonads

Tuesday, January 22, 13 Spironucleus elegans

Diplomonads

courtesy of Guy Brugerolle Blaise Pascal University of Clermont-Ferrand, France

Tuesday, January 22, 13 Trichomonas vaginalis

Trichomonads Parabasalians

Tuesday, January 22, 13 Streblomastix strix

Oxymonads

Leander, B. S. and Keeling, P. J. 2004: Symbiotic innovation in the oxymonad Streblomastix strix. J. Eukaryot. Microbiol. 51: 291-300

Tuesday, January 22, 13 Devescovina striata

Cristamonads Parabasalians

Kirby, H. 1941: Devescovenid flagellates of termites. II The genus Devescovina. Univ. Calif. Publ. Zool. 45: 1-92

Tuesday, January 22, 13 Staurojoenina assimilis

Hypermastigotes Parabasalians

Dolan, M. et al. 2004: Cysts and symbionts of Staurojoenina assimilis Kirby from Neotermes. Europ. J. Protistol. 40: 257-264

Tuesday, January 22, 13 Eucomonympha imla

Hypermastigotes Parabasalians

Carpenter, K. J. and Keeling, P. J. 2007: Morphology and phylogenetic position of Eucomonympha imla. J. Euk. Micro. 54: 325-332

Tuesday, January 22, 13 Current problems with eukaryotic systematics

• “Phylogeny by fiat” • Morphology vs. molecular systematics • Discrepancies between phylogenetic markers (rRNA vs. other genes)

• Issues with statistics (LBA) • Lack of statistical support for monophyly of major “supergroups”

Tuesday, January 22, 13 * Ways to improve phylogeny

• Increased taxonomic sampling (break up long branches) !** add more organisms

• Concatenated gene phylogenies !**add more data (meaning more genes)

• Multicharacter data sets (gene phylogenies, morphology, etc.) !** add more non-molecular data

Tuesday, January 22, 13 -enabled strategies for determining evolutionary relationships

•consensus phylogenies of single genes •consensus phylogenies of many genes •concatenated gene phylogenies

how do we assess statistical support?

do all genes have similar evolutionary

histories? (i.e., are all genes good proxies?)51

Tuesday, January 22, 13 How to read trees

Tuesday, January 22, 13 “natural” groups of organisms (or genes)

sister groups

descendants A B C

ancestor of A, B X descendant of Y

most recent common Y ancestor of A, B, C

think about your own geneology.....

Tuesday, January 22, 13 The anatomy of a tree

• Nodes • Branches • Leaves (taxa) 50 • Ingroups • Outgroups • Roots • Bootstraps • Scale bar

Tuesday, January 22, 13 Branches can rotate around nodes * Nodes are can rotate 360° - like a mobile

Tuesday, January 22, 13 Cladograms vs. phylograms

Cladograms show branching order - branch lengths are meaningless

Phylograms show branch order and branch lengths

Branch length is directly proportional to the amount of change

Tuesday, January 22, 13 57

Tuesday, January 22, 13 Rooting using an outgroup

Unrooted tree

rooted trees

Unrooted:“direction” of evolution unknown, or not inferred Rooted:“direction” of evolution known, or inferred

Tuesday, January 22, 13 Which of the four trees above depicts a different pattern?

Tuesday, January 22, 13 Which of the four trees above depicts a different pattern?

Tuesday, January 22, 13 How do we evaluate the quality of trees?

61

Tuesday, January 22, 13 How do we evaluate the quality of trees?

What parts of the tree are we confident in?

61

Tuesday, January 22, 13 Non-parametric bootstrapping

• Analyze original dataset • Generate new samples by resampling original data (n=# columns, n times) • Analyze new datasets like original • Summarize variation (“consensus”)

• Bootstrap values = variation in tree topologies (generally want >50)

Tuesday, January 22, 13 Bootstrapping

–The number of times a particular branch is formed in the tree (out of the X times the analysis is done).

–used to estimate probability and indicated on a consensus tree

–High bootstrap values don’t mean that your tree is the true tree!

Alignment and evolutionary assumptions are critical

Tuesday, January 22, 13 Monophyly, Paraphyly, and Polyphyly

“natural” groups = monophyletic

1. monophyletic: ancestor + all descendants (a.k.a. “clades”) 2. polyphyletic: multiple ancestries 3. paraphyletic: ancestor but not all descendants

check bootstrap support

What about dark groups to the right?

Tuesday, January 22, 13 Polytomy:

unresolved branching order, meaning either:

“unresolvable” using given phylogenetic methods (e.g., low bootstrap values) or multiple diverged simultaneously

Polytomies are usually represented as more than two branches coming from a single node.

Tuesday, January 22, 13 1. monophyletic: ancestor + all descendants (a.k.a. “clades”) 2. polyphyletic: multiple ancestries 3. paraphyletic: ancestor but not all descendants

check bootstrap support

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Tuesday, January 22, 13 Xenologs: Horizontal gene transfer

E. coli

Horizontal gene transfer: genes acquired not through common “vertical” ancestry

Tuesday, January 22, 13