Name: NRES 441/641 – Spring 2007 Ecology and Management of Invasive Plants First Mid-term Exam – Place your name at the top of this page in the space provided and on any other sheets of paper that you use. (Total of 50 points) I. Multiple Choice. Circle the most correct answer for each question below (one answer only). (1 pt each) 1. Which of the following elements does the Federal Executive Order 13112 NOT contain: a. provides for legal action taken by a group against a federal agency b. definition of invasive species c. details of the stakeholders involved in carrying out the order d. provisions for an Invasive Species Management Plan 2. Invasive species tend to have characteristics of: a. high phenotypic plasticity and be dioecious b. high seed production and a short juvenile period c. low seed production and long flowering periods d. seed dormancy mechanisms and be monoecious 3. In the Great Basin, juniper is an example of a _ that is also a _ a. native, invasive b. immigrant, invasive c. native, casual d. alien, transfomer 4. Callaway and Aschehoug (2000) tested the allelopathy hypothesis. Which of the following is true about the effect of diffuse knapweed (Centaurea diffusa) on either the Montana grass species or Eurasian grass species: a. activated carbon in the soil increased the biomass of the Montana grasses when grown in the presence of C. diffusa, but decreased the biomass of the Eurasian grasses b. activated carbon enhanced the uptake of phosphorus for both the Eurasian grasses and Montana grasses c. the seeds of C. diffusa attacked the seeds of the Montana grasses d. the biomass of C. diffusa decreased when grown in the presence of the Montana grasses as compared to when it was grown with Eurasian grasses 5. A difference between a noxious weed and an invasive plant is: a. there is no difference between the two; it is only a result of inconsistent scientists b. an invasive species is legally defined in the Nevada Revised Statutes – Chapter 555, while a noxious weed is not c. “noxious weed” implies that the plant has a negative economic or environmental impact, whereas the name “invasive” does not require a negative impact d. noxious weeds disperse naturally while invasive species are dispersed by humans 1 NRES 441/641 – Spring 2007 First Mid-term Exam 6. Which of the following would be evidence that a species came from a disjunct population and is therefore an alien? a. similar genetic diversity between two populations b. insects that only utilize that species c. systematic floral survey showing an absence of the species 100 years ago d. nearby pack-rat midden containing a fragment of the species 7. Changes in vegetation related to increases in atmospheric CO2 are examples of the: a. environmental change hypothesis b. disturbance and land use hypothesis c. limiting resource hypothesis d. biodiversity hypothesis 8. In a new environment, an alien plant may potentially encounter new enemies. If the new alien species is to be successful, the potential enemies would: a. feed on foliage b. be dioecious c. have a large population size d. be generalists 9. According to Alpert (2000), which of these hypotheses of invasion requires no advantage to being introduced? a. allelopathy b. vacant niche c. competition d. enemy release Note: the paper specifically mentions the competition hypothesis in this category, but I also accepted ‘vacant niche’ as a correct answer because this hypothesis also does not imply that there is an advantage simply to being introduced. 10. How would you best explain the contradictions in the biodiversity hypothesis? a. higher biodiversity implies higher levels of competition and fewer unused resources, but invaders from rich floras are more competitive and successful b. biodiversity conveys community stability, but reduces the realized niches of native plants c. areas of higher biodiversity have already undergone invasion in the past and are no longer easily invaded d. local diversity conveys resistance to invasion while between-community diversity implies higher resources and increased invasion Note – this is in the paper by Shea and Chesson. 11. Which of the following statements about alien species is true? a. aliens are currently easy to detect by remote sensing b. all alien species are dependent on disturbance by human activities c. aliens are generally considered to be colonizers d. most alien species that become naturalized also become invasive 2 NRES 441/641 – Spring 2007 First Mid-term Exam 12. The idea that a community becomes more susceptible whenever there is an increase in the amount of unutilized resources is a basic concept from: a. variable resource hypothesis b. vacant niche hypothesis c. disturbance and land use hypothesis d. environmental change hypothesis 13. Which of the statements about the micro-evolutionary change hypothesis is true? a. because of the things like genetic drift, founder effects and hybridization, microevolutionary changes that occur in aliens are always beneficial b. hybridization allows the alien species to incorporate potentially advantageous local adaptations into its genome c. this hypothesis is scientifically well-documented d. repeated introductions are necessary for alien establishment 14. Disturbances: a. create a temporary effect, and even with the introduction of invasive species, the area usually returns to its previous state b. are often caused by humans c. decrease the chance of ruderals establishing d. discourage invasions by increasing or altering available resources 15. Which of the following is NOT true of the anthropogenic hypothesis? a. there is substantial and widespread evidence for this hypothesis b. evidence in support of this hypothesis is that the most-visited natural areas have the most exotic species c. humans have intentionally introduced exotic species d. humans are vectors for introduction Note: introductions are both deliberate and accidental in this hypothesis. II. Matching. Below is a list of words and phrases. For each of the numbered statements, write on the line at the end of that statement the word or phrase from this list that best matches the statement. (1 pt each) (-)-catechin Crowding index Introgressants Realized niche Allelopathy Allopolyploid Saltation Fundamental niche Genetic bottleneck Hybridization Naturalized Phylogenetic distance C3 Ruderal Transformer Weed C4 1. Subset of invasive plants that change ecosystems. Transformers 2. One reason why plants may be more invasive in some communities than in others (Strauss et al 2006). Phylogenetic distance (Strauss et al is the paper about relatedness and invasion ability) 3. Plants having this photosynthetic pathway are predicted by the global change hypothesis to benefit from an increase in atmospheric CO2. C3 3 NRES 441/641 – Spring 2007 First Mid-term Exam 4. A decrease in reproductive barriers is a consequence of this process. Hybridization 5. A life history strategy in which the goal is to grow quickly and have a high fecundity, particularly under stress. Ruderal 6. The theoretical limits of existence for a species along n resource axes. Fundamental niche 7. An individual formed by two different species that can still back cross with 1 or more parent. Introgressant 8. An alien species that can reproduce consistently and can create self-replacing populations without human-intervention. Naturalized 9. The actual limits of existence for a species along n resource axes. Realized niche 10. When one plant releases chemicals that are toxic to another. Allelopathy III. Short Answer: Data Interpretation. This section involves a series of graphs and questions related to those graphs. On a blank piece of paper, answer each of the questions related to the graphs in 1 or 2 sentences. Be sure to clearly number each question (for example, “Short answer 1a”), make sure your name is at the top of each page, and number each page in consecutive order. (10 pts). At the top of the next page of this exam are two figures. Fig A shows growth of grass tillers of five U.S. native grasses grown in competition with Centaurea maculosa, both with and without activated carbon in the soil. Fig B shows growth of C. maculosa with and without activated carbon, and with and without native competitors. “Naïve” plants are from un-invaded native populations, “experienced” plants are from populations growing with C. maculosa in the wild. Answer the following questions about these graphs. (a) What is the effect of activated charcoal on the growth of native grasses grown with C. maculosa? What does this imply about the effect of C. maculosa on native prairie grasses? (b) What do you notice about the effect of C. maculosa on ‘naïve’ plants VS ‘experienced’ plants, in general? (c) What do you notice about the growth of C. maculosa when grown with naïve VS experienced native competitors? (d) What do the results you noted in (b) and (c) tell you about the long-term response of vegetation to invasion by C. maculosa? (e) The authors of this paper suggest that ‘maternal effects’ could be a confounding factor in this study. How would you design an experiment to verify that maternal effects do not come into play? 4 NRES 441/641 – Spring 2007 First Mid-term Exam Fig B Fig A (From Callaway et al 2005. Journal of Ecology 93:576-583). 5 NRES 441/641 – Spring 2007 First Mid-term Exam Data interpretation a) Figure A shows the percent change in tiller growth (compared to no competition) for native grasses grown with Centauria maculosa with and without activated charcoal in the soil – the format is the same as for the assigned paper Callaway and Ascheoug 2000. In the absence of activated charcoal, C. maculosa has a strong negative impact on growth of most of the native grasses. Activated charcoal reduces this negative effect, implying that the negative impact is due to an allelochemical released by C. maculosa (since activated charcoal absorbs chemicals from the soil). b) This question is based on Figure A again. Both naïve and experienced plants are negatively impacted by C. maculosa and activated charcoal reduces the negative effect on both naïve and experienced plants, but in general the negative impact of C. maculosa is greater on naïve plants than it is on experienced ones. c) Figure B shows biomass of C. maculosa grown alone (-N) and in competition with a native grass (+N), either with or without activated carbon, and using competitors from either experienced or naïve native populations. When grown with experienced plants, C. maculosa biomass is decreased regardless of whether activated carbon is present in the soil or not; when grown with naïve plants, C. maculosa only experiences a decrease in biomass when activated carbon is present. This implies that experienced plants have a competitive effect on C. maculosa even when the allelochemical is present, but naïve plants only have a competitive effect when the allelochemical is reduced by activated carbon. d) Taken together, the evidence in a) through c) implies that there is adaptation for resistance to the allelochemical released by C. maculosa. Experienced populations (which have been growing with C. maculosa) have undergone natural selection where more allelochemical-resistant plants have produced more offspring, resulting in a reduced effect of C. maculosa on the population. This may mean that there will be evolution of native communities to become more resistant to C. maculosa through time, and that those plants surviving the initial invasion may be able to maintain or even increase their population, thus reducing the invasive ability of C. maculosa. e) Maternal effects are caused when differences in the environment experienced by the parent plant result in non-genetically based differences in seed (weight, protein content, etc) which in turn cause differences in the growth of the resulting seedlings. To overcome this effect, grow seed from naïve and experienced populations for one generation in a common environment (common garden) taking care to avoid cross-pollenation, harvest seed from those plants, grow the second generation seeds, and conduct the experiment on the resulting plants. 6 NRES 441/641 – Spring 2007 First Mid-term Exam Short Essay (CHOOSE ONE). On blank pieces of paper, answer one question in about 10-15 sentences. Be sure to clearly label the question at the top of the page (for example, “Short Essay #1”), make sure your name is at the top of each page, and number each page in consecutive order. If you require more paper to answer the questions, get additional sheets from the instructor. (15 pts) 1. Throughout this class, we have discussed the difficulties of concisely defining invasive species. In the Richardson et al. (2000) paper, the authors first breakdown the barriers that a species must take if it is to become invasive. These stages are introduction, naturalization and invasion. Using those stages, they then ascribe different statuses to alien species such as alien plant, casual alien plant, naturalized plant, invasive plants, and transformers depending on which barriers they have surmounted. First explain the introduction, naturalization and invasion stages. Next, detail the types of barriers that an alien must overcome to move to the next step. Then describe three of the five statuses listed above in detail. Points breakdown for essay 1: 6 points for explaining the three stages of invasion (introduction = moving plant to new location and having it survive; naturalization = plant population reproduces and becomes self-sustaining; invasion = plant population expands and disperses far from initial introduction, moving into new areas). 3 points for mentioning 3 of the 4 barriers that must be overcome in relation to each stage of invasion (introduction = geographic barrier; naturalization = environmental and reproductive barriers; invasion = dispersal barrier). 6 points for describing three of the five statuses (alien = plant introduced by humans; casual alien = can survive and maybe reproduce but only present because of repeated introductions; naturalized = established, can reproduce and maintain population, has become part of the flora; invasive = population is expanding into new areas, including undisturbed habitats, and may begin to be a pest; transformer = plant is abundant or different enough to cause dramatic changes in the ecosystem) 2. Many of the hypotheses explaining what makes species invasive discussed in class are not mutually exclusive of one another. From the list, pick two hypotheses that you believe are not independent of one another and explain how they are not mutually exclusive. To better illustrate, use either data presented in class or create a scenario that supports your argument. Points breakdown for essay 2: 2 points for naming two hypotheses. 4 points for describing the two hypotheses. 6 points for explaining how the two are related. 3 points for examples/scenario. The hypotheses are: vacant niche, superior competitor, allelopathy, environmental change, variable resources, biodiversity, microevolution, enemy release, disturbance, and anthropogenic. This question was very open-ended, so almost anything was acceptable as long as you were able to explain the hypotheses, explain the link you saw between them, and give examples. 7