Diversity and Evolution of Asterids
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Core Asterids Diversity and Evolution • will examine the second of the two well supported lineages of the core asterids of Asterids lamiids • ‘lamiid’ or Asterid I group • ‘campanulid’ or Asterid II group . viburnums, umbels, and lobelias . Reminder: pollination biology project due this Wednesday campanulids Core Asterids Early vs. Late Sympetaly euasterids II - campanulids euasterids I - lamiids • will examine the second of the two well supported lineages of the core asterids lamiids • ‘lamiid’ or Asterid I group • ‘campanulid’ or Asterid II group = NOT fused corolla tube campanulids • Asterids gained fused corolla several times • 2 separate origins of fused petals in Calendula, Asteraceae early “core” Asterids (plus several times in also in Cornaceae of Anchusa, Boraginaceae late Ericales; Cornales start fused) ”basal asterids” 1 Dipsacales “Caprifoliaceae” - what is it ? Adoxa • order within ‘campanulids or Asterid II DNA phylogeny • pre-DNA era four families group were recognized (as in Gleason/Cronquist) • 2 (or 7) families and nearly 1,100 lamiids species of herbs and shrubs largely from Caprifoliaceae North Hemisphere – paraphyletic! honeysuckle • iridoids, opposite leaves, inferior viburnum gynoecium campanulids Dipsacaceae inferior G Valerianaceae teasel valerian “Caprifoliaceae” - what is it ? “Caprifoliaceae” - what is it ? DNA phylogeny DNA phylogeny • the two families can be Adoxa viburnum elderberry differentiated by flowers: • post-DNA era up to 7 • short styled (Adoxaceae) families can be recognized • long styled (Caprifoliaceae) • will use 2 here [MI flora differs!] honeysuckle teasel valerian 2 “Caprifoliaceae” - what is it ? Adoxaceae - viburnums • the two families can be 5 genera and 200 species of opposite leaved shrubs and some herbs - mainly North Temperate differentiated by flowers: • flowers small and usually in flat-topped cymes • short styled (Adoxaceae) • 5 merous with up to 5 • long styled (Caprifoliaceae) fused carpels and short style • berry or drupaceous fruits Viburnum - viburnum Lonicera - honeysuckle Sambucus canadensis - Common elder Adoxaceae - viburnums Adoxaceae - viburnums • viburnums are simple leaved shrubs, but often lobed, or coarsely toothed Sambucus racemosa - red-berried elder • elder-berries are opposite, compounded leaved shrubs Viburnum davidii 3 Adoxaceae - viburnums Adoxaceae - viburnums • note enlarged, sterile flowers on edge of ‘head’ inflorescence for pollinator • many viburnums attraction have glands on petioles Viburnum lentago nannyberry Viburnum opulus (trilobum) high-bush cranberry Viburnum acerifolium- Viburnum rafinesquianum - another “pseudanthium” maple-leaved viburnum arrowwood or false flower Adoxaceae - viburnums *Caprifoliaceae - honeysuckles Northern hemisphere family (and tropical mountains) of 43 genera and about 900 species of shrubs, subshrubs, vines and herbs • unusual reduced woodland herb with musky scent and trifoliate, spiralled leaves Linnaea - twinflower Lonicera - honeysuckle Dipsacus - teasel Adoxa moschatellina - muskroot 4 *Caprifoliaceae - honeysuckles *Caprifoliaceae - honeysuckles ___ CA (5) CO (5) A 4-5 G (2-5) • 5 merous. long styled, and • honeysuckles include native bell-shaped or strongly (right) and introduced aggressive zygomorphic shrubs (below) or vines • inferior ovary forms berry (or reduced to achenes) Lonicera - honeysuckle Lonicera tatarica - Lonicera canadensis - tartarian honeysuckle fly honeysuckle Triosteum - horse gentian *Caprifoliaceae - honeysuckles *Caprifoliaceae - honeysuckles natives invasives Lonicera japonica Lonicera dioica - red honeysuckle Lonicera xylosteum - European fly honeysuckle Lonicera X bella - native leaf miners Lonicera reticulata - grape honeysuckle pretty honeysuckle 5 *Caprifoliaceae - honeysuckles *Caprifoliaceae - honeysuckles Linnaea borealis - twinflower • Characteristic subshrub of (circum)boreal forests where it forms large colonies. Two flowers form per inflorescence. Diervilla lonicera - bush honeysuckle Triosteum perfoliatum Horse gentian *Caprifoliaceae - honeysuckles *Caprifoliaceae - honeysuckles • the old ‘Valerianaceae’ and ‘Dipsacaceae’ • many culativated ornamentals exhibit features that will be seen in Asteraceae [“pseudanthium” or head] • congested inflorescences • bracted inflorescences Abelia Kolkwitzia Valeriana uliginosa - Valeriana edulis - Knautia marsh valerian edible valerian blue-buttons 6 *Caprifoliaceae - honeysuckles *Caprifoliaceae - honeysuckles • the old ‘Valerianaceae’ and ‘Dipsacaceae’ • the old ‘Valerianaceae’ and ‘Dipsacaceae’ exhibit features that will be seen in exhibit features that will be seen in Asteraceae [“pseudanthium” or head] Asteraceae [“pseudanthium” or head] • reduced calyx - pappus • involucral bracts • one seeded achene Knautia pappus Dipsacus fullonum - teasel Introduced and adventive Involucral bracts pappus Apiales Araliaceae - ginseng • 7 families and nearly 5,500 species of • tropical family of trees, shrubs, or herbs and shrubs - Australasia appears to herbs (temperate) be center of diversity • leaves divided or up to 3 times or more compound lamiids • iridoids, aromatic, alternate compound • distinctive aroma and chemicals leaves, umbels, corolla separate, inferior gynoecium campanulids inferior G ginseng - Araliaceae parsnip - Apiaceae Aralia hispida - bristly sarsaparilla Aralia nudicaulis - wild sarsaparilla 7 Araliaceae - ginseng Araliaceae - ginseng __ CA 5 CO 5 A 5 G (5) • umbels • flowers small, non-asterid like (no corolla tube) • 5 merous, with 5 fused carpels forming inferior ovary & berries Aralia elata Hercule’s-club, Devil’s walking stick Aralia racemosa Aralia hispida - bristly sarsaparilla : note 5 styles spikenard Araliaceae - ginseng Araliaceae - ginseng Oplopanax horridus Panax quinquefolius - American ginseng Devils club Western N Am. – Great Lakes disjunction Rich woods; leaflets are stalked. Heavily only Isle Royale in Lake Superior collected woodland species for roots. 8 Araliaceae - ginseng Araliaceae - ginseng Tetrapanax papyrifera - Hedera helix rice paper English ivy Eurasia - N. Africa; invasive in U.S. Schefflera - umbrella plants *Apiaceae - umbels *Apiaceae - umbels Large family of 430 genera and over 3700 species most • aromatic herbs with hollow stems common in north temperate regions. Economically important • dissected or compound leaves (carrot, parsnip, parsley, celery, dill, caraway). • leaves strongly sheathing sheath parsnip carrot Heracleum lanatum - cow parsnip 9 *Apiaceae - umbels *Apiaceae - umbels __ __ CA 5 CO 5 A 5 G (2) CA 5 CO 5 A 5 G (2) • flowers small in umbels, often compound • flowers small in umbels, often compound • female flowers often along edge of each umbellet • female flowers often along edge of each umbellet • 5 merous with no corolla tube common feature in Asterids with the formation of a ‘head’ • inferior gynoecium of 2 carpels separating at maturity *Apiaceae - umbels *Apiaceae - umbels __ CA 5 CO 5 A 5 G (2) • fruit dehiscent and splitting - schizocarp • 2 dry, 1-seeded mericarps held together by carpophore • fruits 5-ribbed separated by oil canals - taxonomic character for separating genera Angelica atropurpurea - great angelica • Large coarse herb of wetter areas 10 *Apiaceae - umbels *Apiaceae - umbels Cicuta bulbifera Bulblet water hemlock Cicuta maculata spotted water hemlock Although containing nasty compounds, Cicuta is not the hemlock that Socrates took (Conium). Common species of Cryptotaenia canadensis - honewort Osmorhiza longistylis - sweet cicely marshes and streams. *Apiaceae - umbels *Apiaceae - umbels Daucus carota - wild carrot, Queen Anne’s lace Note umbel with involucral bracts; larger Eryngium yuccifolium female flowers along edge of umbel; the progenitor of cultivated carrot (same Rattlesnake master species); common weed Dry prairie species with sword-shaped leaves with spiny edge. Umbels has become literally a “head” as in Asteraceae. 11 *Apiaceae - umbels *Apiaceae - umbels The mystery plant from Abilene, Texas Heracleum lanatum American cow-parsnip Eryngium leavenworthii Dipsacus fullonum – teasel (CaprifoliaceaeEryngium yuccifolium) ? Our most robust species in Wisconsin *Apiaceae - umbels *Apiaceae - umbels Pastinca sativa - wild parsnip Torilis japonica- hedge parsley Introduced and spreading along roadsides. Introduced and spreading in forests. If Dermatitic reaction from leaves is rapid collected, mount it! but only in presence of UV light which Torilin - effective blocker of testosterone causes precursor to change to phototoxin. to DHT conversion (5 alpha-reductase) [“pseudo-hermaphroditism”, no male pattern baldness, no prostate cancer] 12 Pittosporaceae Pittosporaceae “The secretory canals and some of the chemical “Rather we must suppose that the anatomical and features, notably the presence of polyacetylenes, chemical similarities between the Pittosporaceae and have led some authors to propose a close relationship Araliales illustrate the pervasive parallelism that between the Pittosporaceae and the Araliales besets efforts to establish phylogenetic relationships [Apiales]. The ovular structure would also be among the angiosperms” consistent with such a relationship.” Cronquist, 1981 Cronquist, 1981 Pittosporaceae Asterales NOT pervasive parallelisms! Recent DNA evidence • 11 families and nearly 26,000 species - Australasia appears to be center of diversity lamiids • no iridoids, latex common, inferior gynoecium, pollen presentation campanulids inferior G bellflower - chickory - Campanulaceae Asteraceae 13 Asterales *Campanulaceae -