Phylogeny of (or Tricolpates) Eudicots (or Tricolpates) “Basal eudicots”

Asterids Buxales Rosids Caryophyllales RanunculalesProteales

After Jansen et al., 2007, Proc. Natl. Acad. Sci. USA 104: 19369-19374 Phylogeny of

Asterids

Lamiids Campanulids Cornales

Cornales Ericales Campanulids: Lamiids: Aquifoliales Garryales Apiales Gentianales Dipsacales Lamiales Asterales Solanales

After APG, 2003; Judd and Olmstead, 2004, and Soltis et al., 2005 Synapomorphies for Asterids and Core Asterids

Synapomorphies for Asterids: Iridoid compounds; Unitegmic ovules; www.scielo.br/img/fbpe/jbchs/v12n2/a04fig01.gif Tenuinucellate ovules.

Synapomorphies for Core Asterids (Lamiids + Campanulids): Gamopetalous corollas; A single whorl of that alternate with the lobes; Epipetalous stamens; 2 fused carpels. ( family)

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Rhododendron macrophyllum (Pacific Rhododendron, WA State ) Ericaceae (Rhododendron family)

Textbook DVD WSJ Textbook DVD KRR & DLN Vaccinium corymbosum;

The Vaccinium contains about 450 , including blueberry, cranberry, huckleberry, etc.. Ericaceae (Rhododendron family)

Photo: Yaowu Yuan Photo: Yaowu Yuan Gautheria shallon; Salal tetragona

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Photo: Yaowu Yuan Note the more or less pendulous Menziesia ferruginea and the urn-shaped corolla. Ericaceae (Rhododendron family)

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Pieris japonica Vaccinium arboreum

Note that the anthers often have spur- or awn-like appendages. Ericaceae (Rhododendron family)

Photo: Yaowu Yuan Menziesia ferruginea Note that anther opens by 2 terminal pores –– poricidal dehiscence. Ericaceae (Rhododendron family)

Textbook DVD KRR & DLN Kalmia latifolia

Note that although corolla can be more spread and not urn-shaped, anther usually opens by terminal pores. Ericaceae (Rhododendron family)

Textbook DVD WSJ Monotropa uniflora Ericaceae includes a mycoparasitic (holoparasitic) group, the monotropoids. In fact, the entire family is strongly associated with mycorrhizal fungi. Mycotrophic: nutritionally dependent on fungi underground, which, in turn are dependent on Textbook DVD KRR other green through mycorrhizal connection. It’s a parasitic relationship, since there is no Monotropa hypopithys exchange from the Ericaceae plants; fungi obtain nutrients from nearby other roots. Ericaceae (Rhododendron family)

Ericaceae - 124 genera/4100 species, including Rhododendron, blueberry, etc..

Trees, , sometimes mycoparasitic herbs lacking chlorophyll. simple, often thick and leathery. Plants often mycotrophic. Flowers actinomorphic, but slightly zygomorphic in Rhododendron. perianths usually 5 parted; flowers more or less pendulous; corolla gamopetalous, often urn-shaped. Stamens (5-) 10; anthers poricidal with 2 terminal pores, sometimes with spur- or awn- like appendages; (stamens act like salt shaker (poricidal dehiscence often with flowers that are pendulous). Carpels 3-5, connate. Fruit a berry , or drupe. ( family)

Photo: Yaowu Yuan Phlox sp. Polemoniaceae (Phlox family)

Photo: Yaowu Yuan Photo: Yaowu Yuan

Collomia sp.

Note the three lobes; gamopetalous corolla with a narrow corolla tube. Polemoniaceae (Phlox family)

Textbook DVD KRR & DLN reptans

Note the three stigma lobes; 5 unequally inserted, epipetalous stamens; gamosepalous calyx ( are fused).

Textbook DVD KRR & DLN Phlox subulata Polemoniaceae (Phlox family)

Polemoniaceae - 18 genera/380 species, most diverse in W North America.

Herbaceous, sometimes slightly woody at base (as in some species of Phlox). Leaves simple, but compound in Polemonium. Flowers actinomorphic. perianths 5 parted; Calyx Gamosepalous (sepals fused); Corolla Gamopetalous, often forms a narrow corolla tube. 5 stamens, often unequally inserted on corolla tube. Ovary superior; Carpels 3, connate –– 3 conspicuous stigma lobes. Fruit a capsule. Solanaceae (Potato or nightshade family)

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Solanum tuberosum

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Textbook DVD WSJ Solanaceae contains many important food crops, drug plants, and popular ornamentals. Nicotiana tabacum Solanaceae (Potato or nightshade family)

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Textbook DVD WSJ Petunia hybrida

Textbook DVD KRR Physalis ixocarpa; tomatillos Solanum lycopersicon

Textbook DVD KRR Datura metel; jimsonweed Atropa belladonna; belladonna Capsicum frutescens; pepper Solanaceae (Potato or nightshade family)

Unlike other members of the Asterids, Solanaceae has alkaloids instead of iridoids as means of chemical defense. Alkaloids in various forms gives these plants various uses, including: Culinary (capsaicin = alkaloid in chilies, producing the “hot” flavor) Medicinal (atropine in belladonna) Narcotic (nicotine in tobacco) Hallucinogenic (various alkaloids in jimsonweeds) Poison (nightshade) Solanaceae (Potato or nightshade family)

Note from the previous images that Solanaceae flowers with 5 , strongly connate, forming a wheel-shaped, bell-shaped, funnel-shaped, or tubular corolla, distinctly plicate (with fold lines).

Solanum melongena (eggplant), showing the plicate corolla

It also has 5 sepals that are connate and persistent, sometimes enlarging as fruit develops.

Textbook DVD DLN Textbook DVD KRR Physalis ixocarpa; tomatillos Physalis alkekengi; Solanaceae (Potato or nightshade family)

Photo: Yaowu Yuan Photo: Yaowu Yuan Solanum spp.

The genus Solanum (more than 1000 species, 50% of the family) and a few related genera have wheel-shaped corolla, connivent anthers and anthers with terminal pores (see the next slide) –– what kind of syndrome? Solanaceae (Potato or nightshade family)

Photo: Yaowu Yuan Solanum sp.; showing the anther terminal pores Solanaceae (Potato or nightshade family)

Solanaceae - 102 genera/2510 species, including tomatoes, potatoes, peppers, eggplants, tomatillos, tobacco, belladonna, jimsonweed, petunia, etc.. Herbs, shrubs, or small trees. Leaves alternate, simple, or sometimes lobed (e.g., tomato, potato). Presence of alkaloids. Flowers actinomorphic (some zygomorphic, but not in N. America). 5 connate sepals (sometimes enlarging as fruit develops); 5 strongly connate petals (lobes plicate, folded like a fan); 5 stamens, connivent in Solanum (50% of species in this family), with wheel-shaped flowers hanging down and anthers with terminal pores –– “buzz” pollination. Stigma capitate; Carpels 2, connate, ovary superior, axile placentation. Fruit a berry or capsule. (Borage family)

Not many members of this family have been used by humans. But it’s a very prominent component of the flora of PNW.

Textbook DVD DLN Myosotis sp.; forget-me-not Boraginaceae (Borage family)

Textbook DVD KRR & DLN Photo: Yaowu Yuan Borago officinalis officinale Note the hispid hair (stiff & prickly; rough to the touch); and the as a helicoid or scorpioid cyme.

Scorpioid cyme Helicoid cyme Boraginaceae (Borage family)

Textbook DVD KRR & DLN Textbook DVD WSJ Mertensiana virginica amabile

Note the gynobasic style and the four parted ovary –– in fruit, the four schizocarps or nutlets. Gynobasic style: a style which is attached to the gynobase as well as to the carpels. Gynobase: an elongation or enlargement of the receptacle, as in the flowers of the Boraginaceae. Boraginaceae (Borage family)

Gersten and Olmstead, unpublished data. Boraginaceae (Borage family)

The Boraginaceae we teach here includes both the traditionally recognized Boraginaceae s.s. and Hydrophyllaceae.

Both Boraginaceae s.s. and formerly Hydrophyllaceae have hispid hairs and helicoid (or scorpioid) cymes as . –– synapomorphies for Boraginaceae s.l..

Difference in the gynoecium structure Ovary, 2 fused carpels: Boraginaceae s.s. with each carpel containing 2 ovules and being divided by a false septum into 2 distinct parts, forming a 4-parted ovary, each locule containing 1 ovule. Formerly Hydrophyllaceae with each carpel containing many ovules, and not being divided into 2 distinct parts. Style: Boraginaceae s.s. with a gynobasic style and capitate stigma. Formerly Hydrophyllaceae with a terminal, 2-forked style Boraginaceae (Borage family)

Boraginaceae - 134 genera/2650 species, prominent in the flora of PNW

Herbs (all of ours), shrubs, or small trees (in tropics), occasionally root parasites Leaves alternate, usually with stiff hispid hairs. Flowers actinomorphic, 5 parted. Helicoid (or scorpioid) inflorescence; Corolla gamopetalous, plicate (with fold lines); 5 stamens; Carpels 2, connate, ovary superior, in Boraginaceae s.s. 4-parted, each locule with 1 ovule, and a gynobasic style; in formerly Hydrophyllaceae each carpel with many ovule and not 4- parted, and a terminal, 2-forked style. Fruit 4 nutlets (schizocarps) in Boraginaceae s.s. or a capsule in formerly Hydrophyllaceae.