Plant Classification Historical Perspective: Preliterate & primitive societies grouped plants in terms of uses, properties, appearance: “folk taxonomies”. Often useful classifications, but only for local area. Plant knowledge important for (agriculture & medicine) Greek & Roman Civilization Theophrastus “father of Botany”: Follower of Aristotle Recognized 4 plant groups: trees, shrubs, undershrubs, herbs Described 500 species. Best work until after the Middle ages Middle Ages Mostly copied from Theophrastus & other classical authors. Little original work Renaissance (15th & 16th centuries) New technologies: Type and printing of books. etc. herbarium (Luca Ghini, 1490-1556, Italy) Return to field observations Herbals: illustrated descriptions of medicinal plants. The 17’th Century: Age of Exploration. Botany expands beyond Europe Expeditions to the New World, Java, Africa, India etc. Many new species collected & described: 500 known in 1542 18,000 in 1682. Need for a comprehensive plant classification system felt. Carl Linnaeus (1707-1778) Swedish physician/ botanist: Devised classification system based on simple flower characters: Classes: stamen number, Orders: number of styles, etc. Published “Species plantarum” First consistent use of binomial names for species (Main contribution to modern taxonomy). Named a large number of species including many from America. Later began effort to devise a more “natural” classification. Carl Linnaeus Natural Classification Systems 18th & 19th centuries: By end of 18th Century, botanists dissatisfied with artificial systems (classified unlike plants together) Quest for classification that recognized 'natural affinities' between plants. Darwin: “Origin of Species” 1859 Theory of evolution by natural selection: Implies that plant populations are variable, change over time, & form lineages of variously related groups. Justified concept of natural groupings. Idea that classification should reflect evolution emerged. Charles Darwin Adolf Engler (1844-1936) “Die Natürlichen Pflanzenfamilien”. First classification influenced by evolutionary theory. Some herbaria & flora manuals still arranged by this obsolete system. Adolf Engler Twentieth Century Traditional Classification Systems: Takhtajan (former USSR, 1980). Cronquist (USA, New York Botanical Garden 1981). Dominant US classification of late 20th C. Flora N.C.T. uses Cronquist’s classification--See FNCT p. 1353-1356; Flora E.Tx p. 1158-1163. Armen Takhtajan Flora E.Tx. retains monocot-dicot grouping but circumscribes families largely like a 21st C. system called APG. Flowering plants classified into 2 main groups: Monocots-5 subclasses Dicots-6 subclasses Arthur Cronquist Recent Approaches to Classification: Angiosperm Phylogeny Group (APG), p. 1163. Bremer, Chase, & Stevens (1998, 2003, 2009) Tries to reclassify flowering plants into monophyletic (single-lineage) groups at the family & order-level) based on recent research. Did not attempt to formally name ranks above order. Basal families Magnoliids Monocotyledons Eudicotyledons The APG Classification (p. 1163-1165) Flowering plant groups: 3 early branches Magnoliids Monocots Eudicots Eudicots Early Magnoliids Monocots branches APG & the angiosperm phylogenetic tree Modern Trends in Plant Classification: See p. 1169-1177 (Apx. 5); 1180-1192 (Apx. 6). Increased emphasis on phylogeny. New sources of taxonomic information (DNA). Emphasis on quantitative methods. Debate between Cladistic & traditional (Phenetic) schools (Apx. 6). Increased emphasis on phylgenetic trees: they express evolutionary relationships better than classification. Current dominance of the APG system for flowering plant classification. Methods to Develop Plant Classifications (p. 1169-1172) Cladistics: Find shared derived characters (synapomorphies). Only these indicate evolutionary relationships & have taxonomic value. Rationale: groups of organisms that share derived (vs. ancestral) characters will be closely related. Goal: find branching pattern of evolution (p. 1169-1172) Traditional Approaches ('Phenetics', p. 1171): Quantify total difference between plant groups based on as many features as possible. Rational: not only phylogenetic relationships but also amount of evolutionary change important for classification (p. 1169-1172). More on Cladistics Main goal: Reconstruct branching pattern of evolution. Product: Branching diagram (phylogenetic tree) called a cladogram. Some advocate eliminating hierarchical classification altogether! (p. 1173, also the debate in Apx. 6). Bryophytes Lycophytes Ferns Gymnosperms Green algae A Cladogram Flowering plants Cladistics Terms Phylogenetic Systematics: = Cladistics Synapomorphy: Shared derived character (p. 1171). Evolved in ancestor of a group & thus present in the group but not outside it. Pleisiomorphy: Ancestral character (p. 1171). Inherited from a distant ancestor thus found both in the group and in other related groups. Clade: monophyletic group (p. 1170). Synapomorphies & Pleisiomorphies of Flowering Plants Synapomorphies: Unique to flowering plants Double fertilization. Seeds enclosed in fruits. Unique arrangement of sex parts (flower). Pleisiomorphies: Found in both flowering plants & related groups. Seeds. Lignin in cell walls. A certain chloroplast DNA mutation. Synapomorphies & Pleisiomorphies of Plants Lycophytes Ferns Gymnosperms Seeds Lignin Chloroplast DNA inversion Flowering plants Double fertilization, Fruits; flowers Critique of Cladistics (p. 1174-1175) Choice of what characters are important/derived, ancestral can be subjective. Difficulties with convergent evolution: Parallelisms (=Homeoplasies) can be confused with shared derived characters. “Evolutionary distance” not considered. Classification in form of a branching tree impractical (but good trees basis for changes to traditional classification). Loss of stability in classification. Cladistics quite successful in generating believable phylogenetic trees: Significant changes in understanding of plant evolution. Resulted in substantial revisions to classification. Classification Groups & Phylogeny (p.1170): Monophyletic group: One complete evolutionary lineageancestor & all descendants; = 'clade'. Polyphyletic group: Several lineages- members don’t share a single recent ancestor. Example: 'Liliaceae'; “dramatically polyphyletic”, p. 726. All taxonomists recognize these as inappropriate. Paraphyletic group: One ancestor but not all descendants. Examples: Reptiles, dogbanes, dicots) pp. 1171-1173). Cladistsic purists don't recognise these groups. Monophyletic Paraphyletic Polyphyletic Producing Classifications: Characters for Taxonomic Studies Pollination biology Anatomy Morphology Secondary compounds Molecular (DNA & RNA) Palynology (Pollen study) Embryology Proteins Chromosome Structure & number Molecular Systematics: the Plant Genomes Nucleus (1.1 x106 - 110 x109 kbp) Mitochondrion (200-2500 kbp) Chloroplast (135-160 kbp) Map of Corn Chloroplast Genome Critique of Molecular Taxonomy Same basic problems as morphological data (convergence, parallelism, etc.) All genes don’t evolve at same rate. Hybridization / introgression & random mutation may result in a gene’s tree not matching the species’ tree. Big advantage: large number of molecular characters available. Best trees & classifications come from studies integrating morphological, molecular & other characters. A T G C T A C G Classification of Organisms: 3 Domains Archaea: (Non-bacterial prokaryotes) Bacteria: (true bacteria) Eukarya: Traditional Kingdoms of Eukarya “Protists”: (Eukaryotic cells, often single-celled organisms. Euglenoids, Red Algae, Dinoflagelates, Diatoms, Brown algae, Green algae… “Catch-all” group: not monophyletic. Revisions in progress: 'Supergroups' of Eukarya (next slide) Fungi: (Heterotrophic, external digestion). Plants: (multicellular autotrophs). Plants in strict sense: 'embryophytes' Animals: (Heterotrophs, internal digestion). 'Supergroups' of Eukarya Alveolata Stramenopila Plants Plants & Relatives Rhizaria Early Eukaryotes Animals Opisthokonta Fungi Excavata Selected Autotrophic“Protista” Diatoms Unicellular or small-colonial. Important phytoplankton, may account for up to 25% of global photosynthesis! Critical for many aquatic ecosystems. Unique 2-part polymerized silica cell walls--Frustules. “Protista” : Brown Algae Large, conspicuous marine seaweeds (kelps). Basic body structure: thallus— multcellular, but simple. Stramenopile clade (also diatoms). Not related to plants, etc. Plstids captured via secondary endosymbiosis. “Protista”: Red Algae Structurally complex seaweeds; Mostly marine; Most abundant in tropics/ warm water. Phycobilin pigments capturegreen light in deep water. Distantly related to plants. Plastids aquired via primary endosymbiosis. Protista: Green Algae Together with higher plants, form a monophyletic group (= “Viridophytes”; G. algae minus plants are paraphyletic. Share basic cellular characters with plants. Variable habit: Unicellular, filamentous, colonial multicellular, etc. Primarily fresh water but terrestrial & marine (Ulva) examples. Plastids via primary endosymbiosis. Two phylogenetic groups (clades): Chlorophyte lineage Other green algae + “higher” plants = “streptophytes”. Green Algae Spyrogyra & Oedogonium: Filamentous green algae Green Algae Volvox: a colonial green alga Green Algae Ulvophycea: a family of marine green algae (p. 334) Green Algae Chara & relatives: “Sister group” to land plants. Multicellular with branched apical growth. Chara: Pinkston Lake, Texas Phylogeny of Viridophytes Chara & Chlorophytes relatives Other Green algae Lycophytes Ferns Seed plants Mosses & Hornworts Liverworts Seedless Vascular Plants Bryophytes Gymnosperms & Angiosperms Green Algae (Protista) •Streptophytes Embryophytes: Plants in the Strict Sense All non-green algal groups of the viridophyte clade. Monophyletic. Multicellular Terrestrial Alternation-of-generations-life cycle: 1n gametophyte alternates with 2n sporophyte. Generations unequal. Resting embryo stage at start of sporophyte generation. Multicelled meiosis organ = sporangium. Water proof wax covers surfaces: cuticle Embryophyte Plant Lifecycles Sporophyte (2N) Sporangium Mitosis Zygote (2N) Fertilization 2N 1N Meiosis ♀ Spores egg ♂ sperm Mitosis Gametophyte (1N) Cladogram of Major Embryophyte Groups Mosses, Clubmosses Group: liverworts, Hornworts (lycophytes) Ferns Pines Oaks (conifers) (angiosperms) Flowers. Seeds in fruits. Seeds. 30bp Cl-DNA inversion. Over-topping branches. Selected Derived Charactertraits Xylem & phloem. Lignin. Branched, independent sporophyte. Embryo. Sporangium. Cuticle. Multicelled sporophyte. Bryophytes: Mosses, Liverworts & Hornworts: First land plants: Ordovician (510-439) M.Y.A. had some bryophyte-like characteristics. Said to be “transitional from green algae to vascular plants”. But: diverse, successful modern plants; NOT 'throwbacks'! Worldwide in most habitats. 2nd (mosses:12,000 sp.) & 4th (liverworts: 9,000 sp.) largest plant groups. Evolutionary strategies: Stay small & remain close to water source. Survive (rather than avoid) dehydration. Produce many spores via many small sporophytes (rather than 1 large one). Gametophyte retains vegetative role. Features of Bryophytes Non-vascular plants: lack xylem & phloem. Often typical of moist areas (but dry-site species also exist)! Rely on immediate contact with moisture (but many can survive long periods of dehydration)! 3 clades Mosses liverworts hornworts Hornworts: Anthocerophyta Sporophyte Gametophyte An East Texas Hornwort! (Sabine N.F.) Sporophytes Gametophyte Liverworts: Hepatophyta A leafy liverwort Gemma cups for asexual propagation A thallose liverwort Liverworts: Hepatophyta Sporophytes Gametophyte Mosses: Bryophyta Capsule (Sporangium) Sporophyte Gametophyte Bryophytes on Ravine Wall: Sabine National Forest TX Bryophyte Life Cycle (example: moss) Sporophyte (2N) Mitosis Zygote (2N) Fertilization 2N Meiosis 1N ♀ Archegonium & egg antheridium & sperm Spores ♂ Mitosis Gametophyte (1N) Vascular Plants (Tracheophytes) All non-bryophyte embryophytes: monophyletic. Specialized water /food-conducting vascular tissues Lignin to strangthen cells in support tissues. 3 clades: Seed plants Ferns & allies Lycophytes The End! Version 15.09