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Core 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 . . . , umbels, and lobelias . . .

Reminder: 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 in Calendula, Asteraceae early “core” Asterids (plus several times in also in Cornaceae of Anchusa, Boraginaceae late Ericales; Cornales start fused) ”basal asterids”

1 ” - 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 largely from Caprifoliaceae North Hemisphere – paraphyletic! • iridoids, opposite , inferior campanulids

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 :

• post-DNA era up to 7 • short styled () 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 • or drupaceous

Viburnum - viburnum Lonicera - honeysuckle 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’ for pollinator • many viburnums attraction have glands on petioles

Viburnum lentago nannyberry

Viburnum opulus (trilobum) high-bush cranberry Viburnum acerifolium- Viburnum rafinesquianum - another “pseudanthium” -leaved viburnum arrowwood or false

Adoxaceae - viburnums *Caprifoliaceae - 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 - 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 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 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 • 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 • 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 , 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 - 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

*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)

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 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 from Abilene, Heracleum lanatum American cow-parsnip Eryngium leavenworthii Dipsacus fullonum – teasel (CaprifoliaceaeEryngium yuccifolium) ? Our most robust species in

*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 []. 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, presentation

campanulids

inferior G

bellflower - chickory - Campanulaceae Asteraceae

13 Asterales *Campanulaceae - bellflowers A family mostly of herbs, but some secondarily woody, widely distributed in the temperate regions and in the montane tropics. Contains 65 genera and over 2200 species, with half belonging to Campanula and Lobelia. lamiids • alternate leaves • milky latex Secondary pollen presentation in Campanula • 5 merous with inferior ovary campanulids

inferior G

bellflower - chickory - Campanulaceae Asteraceae

*Campanulaceae - bellflowers *Campanulaceae - bellflowers The family is divided into two distinct subfamilies - Subfamily Campanuloideae Campanuloideae and Lobelioideae - distinguished by ___ floral symmetry, staminal fusion, and carpel number CA (5) CO (5) A 5 G (3-5) Campanula and relatives have actinomorphic flowers, not fused, and 3-5 fused carpels. Note the 3 styles of Campanula to the left.

Campanula - bellflower Lobelia - lobelia Subfamily Campanuloideae Subfamily Lobelioideae

This is a 5 carpellate species

14 *Campanulaceae - bellflowers *Campanulaceae - bellflowers

Campanula americana - tall bellflower Campanula rotundifolia Campanula rapunculoides Bluebell - circumboreal European bellflower

*Campanulaceae - bellflowers *Campanulaceae - bellflowers Subfamily Lobelioideae Lobelia and relatives have zygomorphic ___ flowers, stamens fused into a tube in CA (5) COZ (5) A (5) G (2) which the pollen is shed, and 2 fused carpels. Style pushes pollen out through the tube. Style

Staminal tube

Triodanis perfoliata - Venus looking glass

15 *Campanulaceae - bellflowers *Campanulaceae - bellflowers

male phase “beard

Lobelia cardinalis Lobelia cardinalis Cardinal flower Cardinal flower female phase

*Campanulaceae - bellflowers *Campanulaceae - bellflowers

Long considered a cure for syphilis – but probably not true

Lobelia siphilitica L. - Great blue lobelia Lobelia siphilitica L. - Great blue lobelia

16 *Campanulaceae - bellflowers *Campanulaceae - bellflowers

Indian tobacco • asthma • muscle disorders • food poisoning • nicotine replacement

• piperidine alkaloids • lobeline primarily

Lobelia inflata Centropogon - Andes Lobelia - Hawaii Indian tobacco Nicocure (vomit-wort) Lobelia - Mt. Kenya

Menyanthaceae - bog buckbean Menyanthaceae - bog buckbean Aquatic, or semi-aquatic family. Flowers 5 merous with fringed petals. Aquatic, or semi-aquatic family. Flowers 5 merous with fringed petals. Gynoecium of 2 fused superior carpels. – long placed in Gentianaceae! Gynoecium of 2 fused superior carpels.

The DNR forced the Botany Dept in Menyanthes trifoliata 2010 to dig up the Botany Garden pond Bog buckbean that had Nymphoides Nymphoides - convergence!

17 Stylidiaceae - trigger flowers Goodeniaceae

Australasian and South American Australasian – and beaches!

Stylidium Donatia Scaevola - note opened corolla tube

Calyceraceae • closest family to Asteraceae - the composites; note biogeography

South American only

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