BIOL 317: Plant Identification and Classification Summer 2016 - Notes

Week 8 – Tuesday • Campanulids • typically have inferior ovary

• Dipsacales • another important family is Adoxaceae (elderberry, viburnum)

• Caprifoliaceae (honeysuckle family) • 42 genera, 890 spp. • distributed widely, but especially diverse in northern temperate zone • herbaceous or woody (shrubs or vines) • leaves - simple or compound; opposite • flowers • zygomorphic, some weakly so • sepals 5, fused • petals 5, fused § often bilabiate, with 2 upper corolla lobes and 3 lower corolla lobes (notable exception is Lonicera [honeysuckle] which has 4 upper corolla lobes and 1 lower corolla lobe) • stamens 4 or 5, epipetalous • pistil compound (2-5 carpels); ovary inferior § usually with long style and capitate stigma § usually 1 ovule/carpel; in some species, only 1 carpel matures • fruit: capsule, achene, berry, or drupe • Older circumscriptions of Caprifoliaceae, including plants with a woody habit, were paraphyletic with respect to Adoxaceae, Dipsacaceae, and Valerianaceae, families with mostly herbaceous members. In order to make monophyletic families, Sambucus and Viburnum were transferred to Adoxaceae, which all have actinomorphic flowers. Dipsacaceae and Valerianaceae were sunk into Caprifoliaceae in the classification followed here, which creates a family with woody and herbaceous members but all with zygomorphic flowers. Other phylogenetic classifications maintain Dipsacaceae and Valerianaceae, define Caprifoliaceae as a narrower clade, and erect new families Linnaeaceae, Diervillaceae, etc. • includes PNW shrubs and wildflowers (Lonicera ciliosa - orange honeysuckle, Symphoricarpos albus - snowberry, Linnaea borealis - twinflower, Valeriana sitchensis - Sitka valerian, etc.), weeds (Dipsacus fullonum - teasel, etc.), and ornamentals (Weigela sp., Abelia sp., Lonicera japonica - Japanese honeysuckle, Centranthus ruber - red valerian, Scabiosa columbaria - pincushion flower, etc.)

• Apiales • another important family is Araliaceae (English-ivy, ginseng)

• Apiaceae (carrot family) • 484 genera, 3780 spp. • distributed worldwide, but especially diverse in northern temperate zone • herbaceous • leaves - often compound, some simple; alternate; sheathing base • often aromatic - possess ethereal oils, terpenoids, and other volatile chemicals used for herbivore defense • inflorescence - umbel (pedicels originate from single point and flowers at about the same level); many with compound umbels (umbels of umbels) • flowers • actinomorphic or zygomorphic • sepals 5 • petals 5, free - secondary loss of fusion • stamens 5 • pistil compound (2 carpels); ovary inferior § 1 ovule/carpel § stylopodium - swollen style base secretes nectar • fruit: schizocarp • includes PNW wildflowers (Lomatium spp. - desert-parsley, Sanicula spp. - sanicle, Heracleum maximum - cow-parsnip, Angelica spp., etc.), weeds (Conium maculatum - poison-hemlock, etc.), food crops, herbs, and spices (Daucus carota - carrot, Pastinaca sativa - parsnip, Apium graveolens - celery, Foeniculum vulgare - fennel, Petroselinum crispum - parsley, Coriandrum sativum - cilantro, coriander, Pimpinella anisum - anise, Cuminum cyminum - cumin, Carum carvi - caraway, etc.), and ornamentals (Eryngium amethistium - sea holly, etc.)

• Asterales • often have secondary pollen presentation (also called plunger or brush pollination, or a pollen pump) • Anthers of stamens form tube through which style and stigma grow. Pollen is deposited on outside of immature stigma lobes, where it is presented to pollinators. Later, the stigma lobes mature and open to expose the inner receptive surface.

• Campanulaceae (bellflower family) • 84 genera, 2380 spp. • distributed worldwide • most herbaceous, some woody (shrubs) • leaves - simple; alternate • milky latex • flowers • actinomorphic (subfamily Campanuloideae) or zygomorphic (subfamily Lobelioideae) • sepals 5, fused • petals 5, fused § bell- or tube-shaped (subfamily Campanuloideae) § 2- or 1-lipped (subfamily Lobelioideae) • stamens 5 § secondary pollen presentation • anthers held together to form tube around pistil (subfamily Campanuloideae) • anthers fused together (syngenesious) to form tube around pistil (subfamily Lobelioideae) • pistil compound (2-5 carpels); ovary inferior • fruit: capsule or berry • includes PNW wildflowers (Campanula spp. - bellflower, Downingia spp., etc.) and ornamentals (Campanula spp., Lobelia spp., etc.)

• Asteraceae/Compositae (sunflower family) • 1620 genera, 23600 spp. - among the two largest families of angiosperms • distributed worldwide • herbaceous or woody (shrubs or trees) • leaves - usually simple, but can be highly dissected; variously arranged • inflorescence - head (capitulum) - sessile flowers densely clustered • subtended by whorl (involucre) of bracts (phyllaries) • discoid head - have only disk florets • ligulate head - have only ligulate florets • radiate head - have ray florets at margin, disk florets in center • flowers • actinomorphic (disk floret) or zygomorphic (ray or ligulate floret) • flowers sometimes individually subtended by bracts on the receptacle (chaff) • sepals none, or 2-many § if present, then highly modified into scales or bristles (pappus) • petals 5, fused § tube-shaped (disk floret) § ligule - expanded single lobe, tongue-shaped (ray or ligulate floret) • stamens 5, epipetalous § secondary pollen presentation - anthers fused together (syngenesious) to form tube around pistil • pistil compound (2 carpels); ovary inferior § 1 ovule/carpel, but only 1 carpel matures • fruit: achene • includes PNW shrubs and wildflowers (Artemisia tridentata - big sagebrush, Balsamorhiza sagitata - arrowleaf balsamroot, Arnica spp., Erigeron spp. - fleabane, etc.), weeds (Taraxacum officinale - dandelion, Centaurea spp. - knapweed, etc.), food crops (Lactuca sativa - lettuce, Cynara cardunculus - artichoke, Helianthus annuus - sunflower), and ornamentals (Chrysanthemum spp., Zinnia spp., Tagetes spp. - marigold, etc.)

Invasive Plants • Definition of "invasive species" • non-native species that spreads into wilderness or managed ecosystems, develops self-sustaining populations, and becomes disruptive to those ecosystems • Impacts of invasive plants • competition for resources, including space, light, water, nutrients, and pollinators § e.g. Rubus armeniacus (Himalayan blackberry) and Helix hedera (English-ivy) - spreading growth can smother other plants § e.g. Tamarix ramosissima (saltcedar) - deep roots can lower water table in arid regions • change ecosystem processes, including nutrient cycling, water cycling, fire regime, and soil accretion § e.g. Cytisus scoparius (Scotch broom) - nitrogen-fixer increases nitrogen content of soils and alters what other plants can compete in ecosystem § e.g. Fallopia japonica (Japanese knotweed) - more efficient translocation of nutrients from leaves to rhizomes alters availability of nutrients in riparian ecosystems § e.g. Bromus tectorum (cheatgrass) - annual grass species creates fuel which increases frequency and intensity of fires in arid ecosystems § e.g. Spartina alternifolia (saltmarsh cordgrass) - forms dense stands which can increase rate of soil build-up in wetlands, leading to loss of habitat • allelopathy - secretion or leaching of chemicals into soil which inhibit the growth of other plants § e.g. Alliaria petiolata (garlic mustard) - secretes chemicals which inhibit growth of mycorrhizal fungi required by other plants • hybridization - genetic integrity of native species may be compromised by introduction of genes from invasive species § e.g. invasive Spartina alternifolia and native Spartina foliosa • Spread of species • international travel and the movement of plant species by humans, both on purpose and accidental, vastly increased in frequency after the period of European exploration and colonization starting in the late 15th century § e.g. introduction of plant species to the Hawaiian islands • natural colonization rate: 1/100,000 years • after Polynesian settlement: 1/50,000 years • after European settlement: 1/0.25 years • how invasive species are initially introduced § woody species - 66% of invasive plants • horticulture/landscaping - 82% • agriculture - 14% • erosion control - 3% • accidental - 1% § herbaceous species - 33% of invasive plants • seed contaminant (accidental) - many • Phases of invasion • establishment phase - population is slowly spreading • expansion phase - population dramatically increases in size § causes of expansion: • major disturbance - opens opportunities for population to spread and grow in size, e.g. fire, hurricane • new genotypes - continued introduction of individuals during establishment phase or hybridization between introduced species and other species allows for novel genetic combinations to form that may allow the population to take off • perception - population may be increasing gradually, but is not noticed by people until it reaches a critical mass • saturation phase - population reaches equilibrium in ecosystem and maintains stable size • Common traits of invasive plants • capable of vegetative reproduction • fast growth and quick to reach sexual maturity • long flowering and fruiting time • early seed germination • capable of nitrogen-fixation • Strategies for control of invasive plants • institutional § prediction of potential invasiveness of species and prevention of spread before invasion § control of populations that have already invaded- very difficult • personal § remove invasive species from property § don't plant, trade, or purchase invasive species § education about invasive species § clean shoes and tires when leaving areas with invasive species that may spread by seed

Week 8 – Thursday • Caryophyllales • order sister to the Asterids • other important families include Amaranthaceae (spinach, beet), Nyctaginaceae (four-o-clock, bougainvillea), Phytolaccaceae (pokeweed), Droseraceae (sundew, Venus flytrap), and Nepenthaceae (Old World pitcher-plant)

• Caryophyllineae • long-recognized group of plants sharing unique suite of traits: • free-central or basal placentation • gave group its older name - Centrospermae • single whorl of tepals • some flowers appear to have two whorls of perianth, but only one whorl is derived from the tepals ‒ the other whorl is derived from bracts or stamens • anomalous secondary growth - due to concentric rings of vascular bundles in stems and roots • instead of single ring of vascular bundles found in most Eudicots • curved embryo • instead of straight embryo found in most angiosperms • perisperm - nutritive tissue in seed derived from nucellus (diploid) • instead of endosperm (triploid) derived from double fertilization • betalain pigments - red to yellow pigments • instead of anthocyanins found in most angiosperms (except in Caryophyllaceae, which secondarily re-gained anthocyanin production)

• Caryophyllaceae (carnation family) • 86 genera, 2200 spp. • distributed widely, but especially diverse in northern temperate zone • herbaceous • leaves – simple, opposite • nodes swollen • flowers • usually actinomorphic • sepals usually 5 - derived from tepals • petals usually 5 - derived from stamens § often bilobed § often clawed - differentiated into broader blade section at apex and narrow claw at base • stamens usually 5 or 10, some epipetalous • pistil compound (2-5 carpels); ovary superior • fruit: capsule • includes PNW wildflowers (Cerastium arvense - field chickweed, Silene spp. - campion, Minuartia spp. - sandwort, etc.), weeds (Stellaria media - chickweed, etc.), and ornamentals (Dianthus barbatus - sweet William, Dianthus caryopyllus - carnation, Gypsophila paniculata - baby's breath, Silene campion - rose campion, etc.) • Montiaceae (miners'-lettuce family) • 14 genera, 225 spp. • distributed widely, but especially diverse in western North and South America • herbaceous, often slightly succulent • leaves – simple, variously arranged; often fleshy • flowers • actinomorphic • sepals 2 - derived from bracts • petals 5 to many - derived from tepals • stamens 5 to many, some epipetalous § opposite petals • pistils compound (2-3 carpels); ovary superior • fruit: capsule • This group was once considered part of Portulacaceae. This older circumscription of Portulacaceae resulted in a paraphyletic group with respect to Cactaceae and several other families. More recent classifications split Portulacaceae into multiple smaller monophyletic families, including Montiaceae, which includes all species native to the Pacific Northwest. Portulacaceae sensu stricto includes only one genus, Portulaca, which includes the common weed P. oleracea (purslane). • includes PNW wildflowers (Lewisia rediviva - bitterroot, Claytonia lanceolata - springbeauty, Claytonia perfoliata - miners'-lettuce, Calyptridium umbellatum - pussypaws, etc.) and ornamentals (Lewisia cotyledon - Siskiyou lewisia, etc.)

• Cactaceae (cactus family) • 131 genera, 1866 spp. • distributed mainly in the New World, especially in arid regions • herbaceous or woody; some, epiphytes in tropical dry forests • primary stems succulent and photosynthetic • areoles - compressed lateral shoots producing spines • leaves – simple, alternate • on main stems - reduced and ephemeral, or none • on areoles - modified to spines § some have glochids - small, hair-like, barbed spines • flowers • actinomorphic • hypanthium • tepals many • stamens many • pistils compound (3-many carpels); ovary inferior - sunken into shoot apex • fruit: berry • includes PNW wildflowers (Pediocactus nigrispinus - snowball cactus, Opuntia fragilis - brittle prickly-pear), food crops (Opuntia ficus-indica - prickly-pear, nopal; Hylocereus undatus - dragon fruit), and ornamentals (Carnegiea gigantea - saguaro, Schlumbergera spp. - Christmas cactus, etc.)

• Polygonineae

• Polygonaceae (buckwheat family) • 43 genera, 1110 spp. • distributed widely, but especially diverse in northern temperate zone • herbaceous or woody (shrubs or trees) • leaves – simple, alternate • ocrea - fused sheathing stipule • nodes often swollen • flowers • usually small - many wind-pollinated • actinomorphic • tepals 5 or 6 • stamens 5-9 • pistil compound (2-3 carpels); ovary superior § only 1 ovule matures • fruit: achene § often angled, winged, or surrounded by enlarged tepals • includes PNW wildflowers (Bistorta bistortoides - American bistort, Eriogonum spp. - wild buckwheat, Persicaria spp. - smartweed, Polygonum spp. - knotweed, etc.), weeds (Persicaria spp., Polygonum spp., Fallopia japonica - Japanese knotweed, Rumex spp. - dock, etc.), and food crops (Fagopyrum esculentum - buckwheat, Rheum rhabarbarum - rhubarb)

• Plant rarity • Biodiversity - variety of organisms at all levels, from genes to species to ecosystems • Distribution of plant species by rarity

• most plant species are found at an intermediate abundance • although few species are very rare, these species are important in contributing to the biodiversity of ecosystems • Reasons for valuing rare species (and biodiversity in general) • aesthetic - appreciation for rare things; contribution to beauty of planet • ethical - belief that all species have the right to exist • utilitarian - potential use in medicine, agriculture, horticulture, etc.; maintain ecosystem function/services • Types of rarity • population size - small • geographic range - small § e.g. species restricted to small islands § endemism - condition of a taxon being restricted in distribution to a particular area • habitat specificity - narrow § e.g. species restricted to specific soil types or localized climates • Threats to rare species • habitat destruction • invasive species • over-exploitation • climate change • others - pollution, disease • Strategies to protect rare species • Legal protections § Federal • Endangered Species Act - 1973 - regulates land use change and possible impacts on endangered species - endangered = in danger of extinction throughout range - threatened = may become endangered in foreseeable future § WA state • Natural Heritage Program - Department of Natural Resources - monitors and protects endangered species in state • Management strategies § preserve habitat § restore habitat § ex situ preservation • maintain populations off natural habitats, e.g. in botanical gardens and seed banks