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Home , Acaena, Acaena magellanica, Acaena microphylla, Adenostoma, Aeonium, Agrimonia gryposepala

Diversity and Evolution of . . . , currants, . . . • continue survey through the eudicots or tricolpates • vast majority of eudicots are Rosids (polypetalous) and core (sympetalous) eudicots

rosid asterid Eudicots • unlike Asterids, Rosids (in orange) now represent a diverse set of families * • before examining the large Rosid group, look at a small but important of flowering - Saxifragales

Paeonia

Sedum *Saxifragales • small group of 16 families and about 2500 sister to Rosids • ancient lineage from 120 mya and underwent rapid radiation

Paeonia

Sedum *Saxifragales

• part of this ancient radiation may involve this small of holo-parasites - Cynomoriaceae *Saxifragales • they generally can be identified by their two or more separate or semi-fused carpels, but otherwise quite variable

Paeonia

Sedum Paeoniaceae 1 / 33 species • like many of these families, Paeonia exhibits an Arcto- distribution Paeoniaceae 1 genus / 33 species • small with primitive features of perianth and • hypogynous with 5-8 separate carpels developing into follicles Cercidiphyllaceae 1 genus / 2 species • small (kadsura-) restricted to eastern and Japan . . . • . . . but in and from Tertiary Cercidiphyllaceae 1 genus / 2 species • unisexual, wind-pollinated but do produce follicles 27 genera and 80 species - witch • family of trees and shrubs in subtropical and temperate areas but only 1 species in Wisconsin - witch found in rich

Hamamelis virginiana - witch hazel Hamamelidaceae CA 4-5 CO 4-5 A 4-5 G (2)

• 4-5 merous and insect pollinated in the fall (images from Sept)

• are ribbon-like

Hamamelis virginiana - witch hazel Hamamelidaceae CA 4-5 CO 4-5 A 4-5 G (2)

• ovary is generally inferior or half-inferior with the tops somewhat separated

• woody, dehiscent at top

Hamamelis virginiana - witch hazel Previous year’s fruit 1 genus and 27 species - sweet gums • small family of trees - sweet gum is familiar in North America; Arcto- Tertiary distribution

• clusters of small follicles

Liquidambar styraciflua * 30 genera and 700 species • family of herbs, Northern Hemisphere in distribution • common - scapose

• prefer wet woods, swampy conditions, or - cool wort drippy cliffs as in the driftless region of SW WI

Micranthes - saxifrage *Saxifragaceae CA 5 CO 5 A 5or10 G (2)

• 5 merous • Superior pistil is made of 2 carpels, separated, at least from the middle up; perigynous often present

2 styles

Micranthes pensylvanica - swamp saxifrage *Saxifragaceae

Note cup-like hypanthium

Mitella - Bishop’s-cap

Heuchera richardsonii prairie alumroot *Saxifragaceae cordifolia Foamflower Endangered boreal sp.

Chrysosplenium - golden saxifrage Grossulariaceae 1 genus and 150 species - temperate regions • characterized by lobed leaves, raceme , and fleshy (currants and gooseberries)

Ribes americanum - American black currant Grossulariaceae CA 5 CO 5 A 5 G (2)

• flowers 5 merous with large and petals smaller

• inferior of 2 fused carpels

well developed hypanthium Grossulariaceae • Currants identified by long racemes of many flowers

Ribes americanum - American black currant swamp currant Grossulariaceae • Gooseberries identified by paired flowers; stems often spiny

Ribes cynosbati - prickly gooseberry, dogberry gooseberry * 34 genera and 1370 species - temperate or warm temperate regions of the world • succulent herbs or small shrubs - jade plants

• CAM (crassulacean acid metabolism) type of photosynthesis

• Wisconsin species are introduced, although yellow sedum is spreading - Gold- in sandy soils stonecrop,Yellow sedum *Crassulaceae CA 5 CO 5 A 10 G 5

• 5 merous with stamens 2X number of sepals (3,4, or 6 merous species occur)

• carpels separate and produce follicles when mature

• nectary scales usually evident at base of each carpel Sedum acre - Gold-moss stonecrop,Yellow sedum *Crassulaceae • major radiation of genera in Mediterranean climates (e.g., )

Aeonium *Crassulaceae

Echeveria Rosids Rosids are one of two large groups of dicots; the other group are the Asterids

Rosids:

separate petals

Asterids:

fused petals * • two major groups within Rosids - we will start with Fabids N2 fixing • include all N fixing plants 2 • the order Rosales: not well defined morphologically (roses, , marijuana, nettles, figs)

1. N2 fixing via actinomycetes (Frankia) 2. loss of corolla in order; petals in = stamens! 3. serrated leaves (glandular +/-) *Rosaceae 100 genera and almost 3000 species distributed worldwide but most common in the north temperate regions - commercial fruits • Comprise herbs, shrubs, or trees and with alternate simple or pinnately or palmately compound leaves

Stipules well developed in compound leaves *Rosaceae CA 5 CO 5 A ∞ G [variable!] • 5 merous, with numerous stamens • gynoecium is variable and used to define subfamilies *Rosaceae CA 5 CO 5 A ∞ G [variable!] • hypanthium present in all species

Bracts on calyx (epicalyx) often present *Rosaceae The gynoecium is variable – four basic types

1. group

2. group

3. Cherry group

4. group

Gynoecium variability encompasses size of receptacle, position of ovary, size of hypanthium, and the resulting fruit types: *Rosaceae

follicles drupelets

achenes aggregate of achenes

drupes pomes *Rosaceae – spiraea group opulifolius - ninebark CA 5 CO 5 A ∞ G 2-8 apocarpic, superior pistils short hypanthium - perigynous follicle fruits *Rosaceae – spiraea group

Spiraea alba - meadow-sweet - hardhack *Rosaceae – rose group CA 5 CO 5 A ∞ G ∞

Herbs with compound leaves

Plants with stolons (running stems above ground) or running

Flowers apocarpic with many carpels

Hypanthium well-developed or receptacle elongated - perigynous

One-seeded achenes *Rosaceae – rose group CA 5 CO 5 A ∞ G ∞

Achenes often modified into aggregate of achenes (from one ) as in the or fleshy drupelets as in raspberry, dewberry

Rubus idaeus - American raspberry sp. - strawberry *Rosaceae – rose group

Fragaria virginiana - wild strawberry triflorum - prairie smoke *Rosaceae – rose group

Fragaria virginiana - wild strawberry gryposepala - common agrimony, harvest lice

2 achenes, but hypanthium disperses as a unit with “velcro”-like barbs from top of hypanthium *Rosaceae – rose group

Potentilla argentea silverweed

Potentilla simplex Common cinquefoil *Rosaceae – rose group

Potentilla breweri complex Western cinquefoils *Rosaceae – rose group

Rubus parviflorus thimbleberry *Rosaceae – rose group

Rubus parviflorus Rubus hispidus thimbleberry

Rubus allegheniensis *Rosaceae – rose group Beach rose

Rosa multiflora Multiflora rose Invasive weed Swamp rose *Rosaceae – cherry group CA 5 CO 5 A ∞ G 1 Shrubs and trees with simple leaves, often with glands along (cherries, plums, , )

Prunus serotina - black cherry *Rosaceae – cherry group CA 5 CO 5 A ∞ G 1 Gynoecium superior with one carpel = monocarpic - perigynous Fruit a drupe = fleshy, with one bony *Rosaceae – cherry group

Prunus serotina wild black cherry

Prunus virginiana choke cherry *Rosaceae – cherry group

Prunus pumila - sand cherry *Rosaceae – cherry group

Prunus americana Wild plum *Rosaceae – apple group — CA 5 CO 5 A ∞ G (3-5)

Shrubs or trees with showy 5 merous flowers

Gynoecium inferior of 3 to 5 fused carpels

Malus pumila - apple *Rosaceae – apple group — CA 5 CO 5 A ∞ G (3-5)

Hypanthium thickens in fruit to form pome fruit Calyx (& CO + A) inserted at top of ovary = epigynous flower

ovary

Malus pumila - apple *Rosaceae – apple group melanocarpa black chokeberry

Pyrus communis (introduced) *Rosaceae – apple group laevis Serviceberry, Juneberry *Rosaceae – apple group

Crataegus crus-galli - cockspur hawthorn

Crataegus mollis - downy hawthorn Xiang et al. . doi:10.1093/molbev/msw242 *RosaceaeMBE

Malus baccata * Malus domestica * delavayi trilobatus * Pyrus bretschneideri DNA tree * alnifolia * Sorbus * * Cydonia oblonga sinensis • spiraea group is polyphyletic japonica * villosa * * Sorbus keissleri amphidoxa * japonica * indica Amelanchier alnifolira * denticulata * - not first diverging group * * germanica californica * oblonga stipulata * Gillenieae * alabamensis * * * ramosissima Kerrieae scandens * uniflora * Prinsepia utilis * cerasiformis Exochordeae racemosa sorbifolia * * • rose core group is * millefolium Sorbarieae * fasiculatum * * * Prunus dulcis * * Prunus persica Prunus yedoensis Amygdaleae * Maddenia hypoleuca monophyletic, but others are * * Maddenia wilsonii * Prunus spinulosa * Pygeum topengii floribundus Lyonothamneae laevigata * * Sibiraea tomentosa * caespitosum scattered around * * * Spiraeeae dioicus * discolor Stephanandra chinensis * Stephanandra tanakae * affinis * * Neillia sinensis Neillieae glabra * australis * * * pinnatus * repens Agrimonieae officinalis • cherry group and apple group * * spinosum * abyssinica * sericea * Spenceria ramalana californica * * purpurascens Potentilla aurea form a monophyletic clade * *

Duchesnea indica * * * Potentilla freyniana bifurca * * adpressa * * Sibbaldia parviflora alpina Potentilleae * * Alchemilla fissa Fragaria ananassa * * palustre * arguta * * altaica * * Chamaerhodos erecta glabra * mongolica * * * * Rosa davurica Roseae * * Geum triforum * Acomastylis elata Colurieae * henryi * * Taihangia rupestris geoides Xiang et al. 2017 paradoxa * * * * Rubeae palmata * * Ulmarieae betuloides * * australis Dryad- mexicana Dryadeae * * Purshia stansburyana octopetala Outgroup oideae

FIG. 1. A summary of Rosaceae phylogeny and Rosaceae fruit morphologies. On the left is a summary tree with results from five coalescence analyses of 882, 571, 444, 256, and 113 gene sets, respectively, and a concatenation analysis using the 113-gene supermatrix. Topologies consistent in all six trees are drawn in black lines. Grey lines show uncertain relationships, with some trees support the topology. For further information see figure 3 and supplementary figure S14, Supplementary Material online. Asterisks (*) indicate 100% supports in all six trees. Diamonds indicate more than 90% supports in at least five trees and more than 85% supports in all six trees. Squares indicate more than 80% supports in at least three trees

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L-CretaceousPal Oli 110 100 90 80 70 60 50 40 30 Age (Mya) MBE Crataegus Fragaria fruit evolution? What this does tell about us Xiang et et Xiang • • once twice or • achenes follicles pomes al. al. and are ancestral 2017 evolved many times drupes evolved