Purple pitcher plants (Sarracenia pupurea) have evolved the ability to lure prey and utilize nutrients absorbed from captured prey to fuel their metabolism.  The unique ecological role of this species provides an opportunity to study interesting interactions including, mutualism, competition and digestive symbioses.  Studies of pitcher plant ecology will provide insight into co-evolutionary theories; pollination and reproductive biology and food web interactions.  The results may also be further applied in the development of management strategies for bogs and seemingly infertile habitats.

Introduction

    The purple pitcher plant (Sarracenia pupurea), is a carnivorous plant that resides in unproductive soils and bogs; absorbing nutrients from various invertebrate prey (Heard, 1998).  The purple pitcher plant is recognized for harboring several aquatic invertebrates including mosquitos, midges, and other small species (Natase, et al. 1995).  Recently, the purple pitcher plant has been recognized for potential prey-mutualism characteristics with a mosquito (Wyeomyia smithii) (Givnish, 1989).  This recent discovery has creates new research possibilities that could potentially aid in further understanding of purple pitcher plant physiology and bog ecology.

    This study will determine if there are relationships between the frequency of prey capture, through examination of organic matter content held in the pitcher, and the frequency of pollinator encounters of the purple pitcher plant .  Purple pitcher plants tend to reside within moist bog-type habitats; therefore, studies will be performed in those ecological areas.  Although several invertebrates are of interest in this study, focus will remain on the potential pollinators that contribute to the diet of the purple pitcher plant.  Other relevant invertebrates, small vertebrates and bacteria will also be studied to pattern ecological food webs.

    Motivation for the initiation of this study includes application of co-evolutionary theories; pollination and reproductive biology of a carnivorous plant; as well as food web interactions.  The significance of these results may also be further applied in the development of management strategies for bogs and seemingly infertile habitats.
 
Background

    Purple pitcher plants (Sarracenia pupurea) have evolved the ability to lure prey and utilize nutrients absorbed from captured prey to fuel their metabolism (Chapin and Pastor, 1995).  The unique ecological role of this species provides opportunity for many interesting interactions including, mutualism, competition and digestive symbioses (Givnish, 1989).  Bacterial symbionts and prey mutualists aid the purple pitcher plant in the digestion and capture of various prey species; allowing the extraction of environmentally limiting nutrients such as nitrogen, phosphorus, sulfur and metals from digested prey (Hardwick and Giberson, 1996; Bradshaw and Creelman, 1984). The purple pitcher plant often resides in acidic swamps, bogs and wet meadows (Heard, 1998).  Pitcher plants use nutrients absorbed from prey items to exploit areas that are low in nutrient availability, discouraging competition with other non-carnivorous plants (Chapin and Pastor, 1995).

    The generalized structure of a fully developed pitcher is a cylindrical open leaf, bounded by a hood  (Pietropaolo, 1986).  The modified leaf shape collects water in a cavity where captured animals drown and are digested by autolytic digestive enzymes (Givnish, 1989).  The accumulated water is not produced by the plant itself making the pitcher dependent on the presence of rain or moisture (Bradshaw and Creelman, 1984).

    The fluid filled digestive and absorptive section harbors a number of non-prey fauna including bacteria, protozoans, rotifers and small crustaceans (Givnish, 1989).  Unlike most carnivorous species, the pitcher plant lacks digestive glands and requires non-prey inhabitants to aid in the decomposition of prey  (Hardwick and Giberson, 1996).  The prey is digested and nutrients are absorbed by the plant to increase photosynthesis and nectar production and by the non-prey inhabitants to increase digestive enzyme synthesis (Chapin and Pastor, 1995).  Nutrient uptake has been reported to have both positive and limiting effects on the plant growth of Sarracenia pupurea (Cresswell, 1993).

    Purple pitcher plant bud development creates a bend in the stalk directing the flower to open downwards (Pietropaolo, 1986).  The flower is perfect, with 5 petals, approximately 80 stamens and a style that expands into an “umbrella-shaped” structure (Pietropaolo, 1986).  Pollen falls into the concavity of the style and insects are seemingly required to move the pollen to stigmatic points (Pietropaolo, 1986).  A period of seed dormancy, often occurring during winter months, is required for optimum germination of the plant (Pietropaolo, 1986).

    The pitcher plant exhibits a prey-mutualism relationship with a mosquito (Wyeomyia smithii) that uses the pitcher fluid as a refuge for developing larvae (Lair et al., 1997).  Female mosquitos select pitcher plants by means of pitcher age, attractiveness, and size (Natase et al., 1995).  She will usually deposit the eggs into newly opened pitchers exhibiting attractive colors (Jaffe et al., 1995).  The larvae produced from the eggs will eventually make their way to the fluid below and develop into adult mosquitos (Natase et al., 1995).  The pitcher benefits from this relationship by attracting other prey items that feed on the mosquitos and their larvae (Givnish, 1989).  The pitcher may also benefit from the recruitment of mosquitos as potential pollinators (Givnish, 1989).

    This study will identify the various invertebrate interactions that occur with the purple pitcher plant.  Insect capture, frequency of pollination,  food web interactions and overall ecology will be investigated.  The cost to the plant in obtaining invertebrate prey will be examined through measurement of nectar secretion and frequency of prey capture.  The benefit to the plant will be determined through analysis of the photosynthetic rate in both high and low nutrient abundance as well as root to shoot allocation.  The analysis of these components should be able to establish if there are any mutualistic, co-evolutionary or exploitative relationships between invertebrates and the purple pitcher plant.

Literature Cited

 Bradshaw, W. E., and R. A. Creelman.  1984.  Mutualism between the carnivorous purple
      pitcher plant and its inhabitants.  The American  Midland Naturalist.  112:294-304.

Chapin, C. T. and J. Pastor.  1995.  Nutrient limitations in the northern pitcher plant
      Sarracenia pupurea.  Canadian Journal of Botany  73:728-734.

Cresswell, J. E.  1993.  The morphological correlates of Prey Capture and Resource
      parasitism in pitchers of the carnivorous plant Sarracenia pupurea.  The American
      Midland Naturalist 129:35-41.

Givnish, T.J.  1989.  Ecology and evolution of carnivorous plants.  pp. 234-290  In :  W.G
      Abrahamson (ed.)  Plant Animal Interactions.  McGraw-Hill, New  York. 480 pp.

Hardwick, M.E. and D.J. Giberson.  1996.  Aquatic insect populations in  Transplanted
      and natural populations of the purple pitcher plant, Sarracenia pupurea, on Prince
      Edward Island.  Canadian Journal of   Zoology  74(11):1956-1963.

Heard, S. B.  1998.  Capture rates of invertebrate prey by the pitcher plant Sarracenia
      pupurea L.  American Midland Naturalist  139:79-89.

Jaffe, K.; M. S. Blum; H.M. Fales; R.T. Mason; and A. Cabera.  1995.  On insect
      attractants from pitcher plants of the genus Heliamphora Sarraceniaceae).   Journal of
      Chemical Ecology  21(3):379-384.

Lair, K.P.; W.E. Bradshaw and C.M. Holzpfel.  1997.  Evolutionary divergence of  the
      genetic architecture underlying photo periodism in the pitcher plant  mosquito.
      Genetics  147:1873-1883.

Natase, A..J.; C. De la Rosa; and S.J. Newell.  1995.  Abundance of pitcher-plant
      mosquitoes, Wyeomyia smithii   (Coq.) (Diptera: Cuicidae) and midges, Meriocnemus
      knabi Coq (Diptera: Chironomidae), in relation to pitcher  characteristics of Sarracenia
      pupurea. American Midland Naturalist.  133:44-51.

Newell, S. J. and A. J. Natase.  1998.  Efficiency of insect capture by Sarracenia
      pupurea (Sarraceniaceae) the northern pitcher plant.  American Journal of  Botany 85(1): 88-91.

Pietropaolo, J. and P. Pietropaolo.  1986.  Carnivorous Plants of the World.  Timber
  Press Inc.  Portland, Oregon. 206 pages.
 
Research Design

    The anticipated null hypothesis is that the frequency of prey capture is independent of the frequency of pollination encounters.

    Insect activity will be monitored by videotaping randomly select pitcher plants.  A video camera will be mounted on a tripod and focused on the individual plants for 2-hour periods.  After the two hours, the camera will then be moved to another plant and monitored.  These videotapes will be analyzed for the identity of visitor frequency to the plants.  A CCD camera will also be used to view insect behavior over 24-hour periods; allowing the recorder to continuously document the activity surrounding the plant.  Individual invertebrates will not be marked, therefore, data will represent recorded number of visitors.  Relative ages of  the pitcher plants  will be pre-calculated to determine if there is any relationship between plant age and frequency of visitors.  Photographic equipment with macro capability will also be used for taxonomic identification by capturing high resolution snapshots of the invertebrates that frequent the purple pitcher plants.

    The gerking sampler and malaise trap will be used to identify the various invertebrates that reside in the bog habitat.  These sampling devices will be randomly placed throughout the entire area of each study site. Mesh net type traps will be also used on randomly selected plants to examine the presence of residents being harbored either in or on the pitcher plant. They will be monitored every third day for maximum quantity of insect capture without initiation of decomposition occurring.  Selected members of each collected species will be preserved in ethanol for identification in the laboratory.

    Temporal variation in plant community structure of the study areas will be determined using several, randomly placed, 50 meter, transect lines.  The transect data will represent the overall ecology and community structure within the studied habitats.  A plotless method will also be used to determine the horizontal spatial distribution and mean distance between pitcher plant individuals.  The plotless method will also determine if clustering of plants causes density changes in the frequency of prey capture or pollinator visits.

    Invertebrates perform many important functions in the soil surrounding pitcher plants.  Soil samples will be taken by digging simple core samples 10 cm adjacent to the location of randomly selected pitcher plants.  These samples will be preserved and  used to identify soil type present in the study areas. The soil will be transferred into plastic containers and further analyzed in the lab for nutrient availabilty and invertebrate abundance.

    All traps, video recording equipment, and field exercises will be conducted in various types of weather; however, the techniques used will all contain some form of protection from extreme conditions.

    Several pitcher plants containing insect larvae in the fluid will be collected from each study area.  In the laboratory these plants will be measured for actual volume of fluid contained in the pitcher, maximum possible volume, length and width of pitcher opening, and the length of the pitcher.  The total organic matter will then be dried and weighed for total organic matter content.

    A Clark style oxygen electrode, light meter and electronic balance will be used to measure the photosynthesis and respiration of the purple pitcher plant under different light intensities. A piece of tissue from a sampled plant will be allowed to respire in a recording chamber under changing the light intensity.  The consumption of oxygen will then be calculated from the data collected by the oxygen electrode recording device.

    In addition to photosynthetic productivity, an electronic balance, scalpels, aluminum foil, weighing boats, drying ovens, and large forceps will be used to measure root to shoot ratios, including total organic matter and dry organic weight.  These measurements will be used to quantify the allocation of photosynthetic productivity to the formation of new leaf tissue.  Similarly, these measurements can also quantify the “conversion of photosynthetate as a function of irradiance, water availability, amount of prey, and substrate supply of different nutrients” (Givnish, 1989).

    The composition of invertebrates will be determined through analysis of the frequency of occurrence.  Relative ages of  the pitcher plants  will be calculated to determine if there is any relationship between plant age and frequency of visitors. Temporal variation in plant community structure, horizontal spatial distribution, mean distance between pitcher plant individuals, and clustering of plants will be examined individually and related to invertebrate abundance. Finally, actual volume of fluid contained in the pitcher, maximum possible volume, length and width of pitcher opening, and the length of the pitcher will be analyzed individually and related to the abundance of organic matter in each pitcher plant.

Milestones

    The entire study will occur over the course of two years beginning in March of 1999.  Two field seasons will be used to collect data.  Initial office work will take place in March and April, including delineation of plots, transects, and study area locations. Field work will begin in May and continue through till October.

    The invertebrate sampling techniques will be used throughout the entire field season.  Video equipment, however, will be rotated between each study area for appropriate lengths of time.  Community plant structure and plotless methods will be used once every two weeks for the length of the field season to determine if there is any variation in plant community structure.  Selection and transport of soil samples and pitcher plant samples will occur at representative time intervals.

    Laboratory and data analysis will take place from November, 1999 through March 2000 and November 2000 through February, 2001.

Anticipated Significance

    The understanding of the relationship between prey capture and the reproductive biology of the purple pitcher plant is a significant step in the comprehension of food web interactions of bog-type habitats. The analysis of these components will be able to establish if there are any mutualistic, co-evolutionary or exploitative relationships between invertebrates and the purple pitcher plant.

    The conservation of rare, infertile habitats, such as bogs, is of major significance to the scientific community because of their successional role as aquatic systems and marked role in ecological systems.  Bog habitats represent the final stage of aquatic systems offering a variety of flora and fauna that are not found in any other area.  These important systems are often ignored or abused by industrial companies, because of their anoxic conditions and infertile characteristics.

Budget

    This study will require the use of a full ton truck in order to carry the equipment between the study site and the laboratory.  Field equipment used will consist of a modified gerking sampler, malaise trap, simple mesh trap, CCD camera, VHS camera, photography camera with macro capability, quadrat plots, shovels, spades and measuring tape.

    The gerking sampler and malaise trap will be used to identify the various invertebrates that reside in the bog habitat.  The simple mesh net trap will be used to examine the presence of resident invertebrates of the pitcher plant.  These techniques will be useful to determine the spatial density of fauna surrounding the plants.  The CCD and VHS cameras will be used to view the frequency of prey capture and the frequency of pollinator encounters. Video “camera’s have been used to record a diversity of insect activity including movement, flight trajectories and behavioral response” of insects (Newell and Natase, 1998). The photographic camera with macro capability will be used for taxonomic identification by capturing high resolution snapshots of the invertebrates that frequent the purple pitcher plants.  The quadrat plots and measuring tape will be used to identify community structure and richness within the study areas through random transect placement and plotless methods.  The shovels will be used to identify the type of soil and nutrient availability present in the study area by a digging a core sample.  The soil will be transferred to plastic containers and further analyzed in the lab.  Finally, the spade will be used to acquire random pitcher plant samples, surrounding vegetation, and soil that will be transported back to the lab for further analysis.

    Laboratory analysis will require the following equipment;  Clark style oxygen electrode, light meter, electronic balance, scalpels, aluminum foil, weighing boats, soil sieves, drying ovens, large forceps, and a computer.

    The Clark style oxygen electrode, light meter and electronic balance will be used to measure the photosynthesis and respiration of the purple pitcher plant under different light intensities.  The electronic balance, scalpels, aluminum foil, weighing boats, drying ovens, and large forceps will be used to measure root to shoot ratios, including total organic matter and dry organic weight.

Anticipated Environmental Impact Statement

There is to be negligible environmental impact to the study areas.  Several plants will be randomly selected, removed, and transplanted into the laboratory for further analysis, and if possible, returned to the area of origin.  As well, several small core samples of soil will be removed, however, their displacement should have little or no effect on the overall structure of the study areas.

Courses/work experience relevant to the proposal

                                    Education
____________________________________________________

Natural Resource Management - Fisheries
University of Northern British Columbia, Prince George, British Colombia
1997 -1999

Aquatic Plants BIOL 402– the course is a survey of the major aquatic plant families  with an emphasis on ecological relationships, physiology, and environmental implications of aquatic plants.  Both marine and fresh water systems are covered with an emphasis on the aquatic plants of British Columbia.

Statistics  MATH 342 – this course is a survey of statistical concepts, graphical and numerical analysis, descriptive statistics, probability, binomial analysis, confidence interval, hypothesis testing, linear regression,  and analysis of variance.

Renewable Resource Management - Diploma Lethbridge Community College, Lethbridge, Alberta 1994 -1996
 
Aquatic Biology RRM 159 – Classification, structure, life history, and life processes of aquatic invertebrates and vertebrates.  The structure of biotic communities will be considered.  Laboratory studies will emphasize bacterial algae, aquatic fungi, aquatic insects and fish identification.

Data Analysis  RRM 150 - this course introduces students to basic statistical concepts used for resource management including both graphical and numerical analysis.  Topics include descriptive statistics, elementary probability, confidence interval, hypothesis testing, and linear regression.

Plant Taxonomy  RRM154 – the course is a survey of the major plant divisions with an emphasis on ecological relationships.  The taxonomy of angiosperms and gymnosperms will be stressed in both lab and lecture.

Range Management RRM 254 – outlines western range and its relationship to domestic livestock and wildlife.  Principles of range management, plant identification, sampling techniques, analysis of data, and animal nutrition will constitute the major areas of concentration.

                                    Employment
_____________________________________________________

Senior Fisheries Technician
Ecofor Consulting  - Vanderhoof, British Columbia  June 1998 – September 1998
 

Summer Student of the Forest Management Crew
Department of Indian and Northern Affairs, Yukon Forestry Division May 1996 - August 1996
 

 Prospectus Copy

A copy of the Prospectus is attached.