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GRASSES AS INVASIVE SPECIES Clay Antieau, MS, Phc Botanist, Horticulturist, Environmental Educator

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GRASSES AS INVASIVE SPECIES Clay Antieau, MS, Phc Botanist, Horticulturist, Environmental Educator GRASSES AS INVASIVE SPECIES Clay Antieau, MS, PhC Botanist, Horticulturist, Environmental Educator WFCA Eighth Western Native Plant Conference November 2019 THE GRASS FAMILY (Poaceae) • Genera : 700 – 800 • Species: 7,500 – 11,000 (4th largest) • Comparable in size to Aves (Birds); ~ twice as large as Mammalia; half the size of Orchid or Aster families • First appeared in pollen record 55 – 70 mya (Paleocene) • Only angiosperm family found natively on all seven continents (Deschampsia Antarctica) • Includes cereal crops (forage, food, sugar, beer...); many ecological dominants • All major civilizations developed around cultivated grasses (Asia to Middle East to New World) Among the World’s Worst Invasives: Grasses • Imperata cylindrica (cogon grass) • Phragmites australis (common reed) • Phalaris arundinacea (reed canarygrass) • Sorghum halepense (Johnson grass) • Spartina species and hybrids (cordgrasses) • Cortaderia species and hybrids (pampas grass) • Arundo donax (giant reed) • Microstegium vimineum (Japanese stilt grass) • Bromus tectorum (cheat) • Taeniatherum caput-medusae (medusahead) • Cynodon dactylon (Bermuda grass) • Neyraudia reynaudiana (silk reed) Important North American Invasive Grasses (adapted from www.fs.fed.us/database/feis) Aegilops cylindricus (goat grass) Eragrostis curvula (weeping lovegrass) Aegilops triuncialis (barbed goat grass) Eragrostis lehmanniana (Lehmann love Agropyron desertorum (desert wheat grass) grass) Elytrigia repens (quack grass) Agropyron cristatum (crested wheat Festuca arundinacea (tall fescue) grass) Imperata cylindrica (cogon grass) Ammophila arenaria (European Microstegium vimineum (Japanese stilt beachgrass) grass) Arrhenatherum elatius (tall oatgrass) Oplismenus undulatifolius (wavyleaf Arundo donax (giant reed) basket grass) Brachypodium sylvaticum (false Pennisetum setaceum (fountain grass) brome) Phalaris arundinacea (reed Bromus hordeaceus (soft chess) canarygrass) Bromus inermis (smooth brome) Phalaris aquatica (Harding grass) Bromus japonicus (Japanese brome) Poa pratensis (Kentucky bluegrass) Bromus tectorum (cheat) Ripidium ravennae (ravenna grass) Cortaderia jubatua(pampas grass) Sorghum halepense (Johnson grass) Cynodon dactylon (Bermuda grass) Spartina alterniflora (smooth cordgrass) Dactylis glomerata (orchard grass) Taeniatherum caput-medusae Echinochloa crus-galli (barnyard grass) (medusahead) Cogon grass (Imperata cylindrica) rhizomes: > 60% of total biomass (40 tons/ac); up to 4 feet deep Cogon grass (Imperata cylindrica) • Infests more than 500 million acres worldwide; every continent except Antarctica; expanding…. • USA introductions: accidental (crate packing) [1911, Mobile (Grand Bay), AL] and intentional (forage) (1921 MS; 1930 FL) Cogon grass (Imperata cylindrica) Ornamental cultivar ‘Red Baron’ can revert back to the wild form and cause infestations. Japanese stilt grass (Microstegium vimineum) Chris Evans, Illinois Wildlife Action Plan, Bugwood.org Giant reed (Arundo donax), Sabino Canyon, AZ photo by Mark Hengesbaugh VENTENATA (Ventenata dubia) •Annual grass, similar to cheat (Bromus tectorum) in habit, habitat, ecology •Formerly in Spokane, Yakima, Klickitat counties; now in Kittitas, Skamania, King, Cowlitz counties photos by Matt Lavin Slender false brome (Brachypodium sylvaticum) Oregon, California, Ontario Slender false brome (Brachypodium sylvaticum) in Oregon as of January 23, 2007 Estimates suggest infestation of at least 10,000 acres California Valley and Foothill California Valley and Foothill Grassland Ecosystem resembled PNW perennial bunchgrass Grassland Ecosystem…GONE! ecosystems. THEN: Dominated by Stipa pulchra; S. cernua, Danthonia californica, Festuca californica, Sitanion jubatum, Elymus glaucus, Melica californica, etc. NOW: 1)introduction of Mediterranean annual grasses and forbs (Bromus, Hordeum, Avena, Vulpia, Briza, Brachypodium…..) 2)overgrazing 3)drought 4)severe wildfire --human-mediated disturbance climax vegetation (“disclimax”)…. permanent, incapable of returning to previous climax vegetation of perennial bunchgrass otherwise expected for regional climate and soils = novel ecosystem Bromus, Lolium, Vulpia California Brachypodium California INVASIVE GRASSES COMPROMISE ECOSYSTEM HEALTH AND INTEGRITY Altered Key Ecosystem Processes and Services include: • nutrient cycling and carbon cycling [Aegilops triuncialis (Drenovsky and Batten 2007)] • sediment erosion and deposition rates (spartina; giant reed) • disturbance intensities and frequencies [cheat (reduces fire cycle from 60–110 yrs to 3–5 yrs); medusahead; cogongrass (max temp 856º F)] • evapotranspiration, water cycling, hydroperiods (reed canarygrass; giant reed; cogongrass) • soil chemistry; soil biological processes (velvet grass) • primary productivity (ryegrass) • genetic integrity (bromes; wheatgrasses) • resilience to disturbance (incl. biol. invasions) (bromes; cogongrass) • habitat availability for native plants/animals/other organisms (reed canarygrass; quackgrass; cogongrass; Japanese stiltgrass; tall fescue…) • biodiversity and associated food web interactions/characteristics Apical Meristems Intercalary Meristems WHY CAN GRASSES BE SO INVASIVE?? Simple modular growth form giant reed (Arundo donax) (phytomer concept: repeated building blocks containing fixed regions of cell division and origins of differentiation; basic unit of growth): node, internode, sheath, blade, and axillary bud; intercalary meristems at bases of blade, sheath and internode. Diversity derives from myriad modified versions of this basic, simple body plan. annual bluegrass (Poa annua) WHY CAN GRASSES BE SO INVASIVE?? Wind pollinated or apomictic; highly reduced (inexpensive) floral and fruit structures; efficient wind, water, and animal (human) seed dispersal; extended seed dormancy: extraordinary capacity for sexual reproduction Phleum pratensis: 1,152,000 seeds/lb vs Trifolium repens: 768,000 seeds/lb Bromus tectorum: 400 lbs/ac vs 200-300 lbs/ac for cool season grasses field corn 3,000 seeds/lb hairy vetch 16,000 seeds/lb tall fescue Bermuda grass seeds (hulled) 227,000 seeds/lb 2,071,000 seeds/lb WHY CAN GRASSES BE SO INVASIVE?? Simple modular growth form (phytomer concept: basic unit of growth): node, internode, sheath, blade, and axillary bud; intercalary meristems at bases of blade, sheath and internode. Diversity from myriad modified versions of this basic body plan. Wind pollinated or apomictic; highly reduced (inexpensive) floral and fruit structures; efficient wind, water, and animal (human) seed dispersal; extended seed dormancy (many, not all). Generally do not produce secondary toxic compounds (glycosides, alkaloids, cyanogenics, etc) and instead avoid/tolerate grazing through morphological and anatomical mechanisms (but see allelochemicals and endophytic fungi, below). = extraordinary capacity for vegetative growth (tillers, stolons, rhizomes, bulbils, corms, fibrous roots) and sexual reproduction…. PHOTOSYNTHETIC APPARATUS C3: mesophyll is C4: coordinated anatomical and physiological loosely packed; bundle features concentrate CO2 around the C3 sheath cells lack photosynthetic machinery through use of a solar- chloroplast. powered biochemical cycle; mesophyll is tightly packed and radiately arranged; bundle sheath cells with chloroplasts. PEP shuttles CO2 to bundle sheath cells. 3% of vascular plant species (C4) account for ~25% of terrestrial photosynthesis. GRASSES ARE PRODUCTIVE Mollisols: Soils formed under prairie or grassland vegetation; grasses add abundant raw organic matter to the soil, mostly by in situ root death. More than 25% of the USA land area is covered by Mollisols, fertile soils now used predominantly for agriculture. Zedler. U. WI Arboretum Leaflet 15. Aboveground productivity, C:N ratios, and root:shoot ratios are higher, on average, in native A Palouse-region soil profile, grasses > non-native grasses > ID native forbs. Abdala, Caldas, Haridasan, Eiten (1998). (Brazil) WHY CAN GRASSES BE SO INVASIVE?? Pre-adapted genotypes (~11,000 species World-wide!) Rapid evolutionary capacity (post-immigration evolution and microevolution) [Novack and Mack (2005): founder effects but high intra-population diversity from multiple introductions and novel genotypes; Longland and Ashleu 2007: evidence for local adaptation]; rapid speciation; niche specialization Chromosomal Diversification: reduction, polyploidy, and hybridization are rampant (80% of all grasses are estimated to be polyploid); notable phenotypic plasticity within taxa (drought, cold, grazing, fire, soil fertility, soil moisture, climate change…) Adaptation to drought and open (sunny) spaces and disturbance Allelochemicals [Hogal and Sanford (2006): Bromus tectorum root extracts reduced growth of Hilaria jamesia by 60%; Lolium and Pinus ponderosa] Endophytic fungi: asymptomatic fungi that occur intra- and inter-cellularly and produce secondary metabolites (alkaloids, indole–diterpenoids…) Escape from biotic constraints FERAL CEREAL RYE (Secale cereale) Beginning in early 1960’s Compared to cultivated varieties (~60 generatons): ShareAlike 4.0 International (CC) • shattering spikelets • delayed flowering • more tillers • shorter (smaller leaves; thinner stems) • smaller, more seeds Burger, J.C. and N.C. Ellstrand. 2014. Rapid evolutionary divergence of an invasive weed from its crop ancestor and evidence for local diversification. J. Systematics and Evolution 52 (6): 750–764. MANAGEMENT OPTIONS Cultural: prevention Physical: mowing, burning, grazing, hand-pulling mature stands, digging Biological:
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