Leaf Economic Spectrum

Aconitum Delphinifolium has traits of a resource acquisitive species on the leaf economic spectrum. It has relatively few leaves that are thin and deeply divided, but are positioned on the plant to capture as much sunlight as possible for photosynthesis. The leaves contain a large concentration of secondary compounds, but are loss at the end of each growing season. It is a herbaceous perennial so all above ground growth dies in the fall, and nutrients are stored in the tuber for growth the following spring.  

Herbivory and Pathogens

Aconitum is well defended against herbivory from animals through high concentrations of alkaloids, which can be fatal if ingested. These alkaloids are found in all parts of the plant except the nectar. The highest concentrations are found in the roots and pollen (Gosselin et al, 2013). The absence of the alkaloids in the nectar, and the high concentration in the pollen suggests a chemical defense against excessive pollen harvest by the pollinators. 
There are a few pathogens that are known to attack Aconitum. Crown rot, stem rot, and wilt are all caused by fungi, and they mainly cause damage to the vascular tissues. There are also some insects that attack Aconitum. The four lined plant bug and larkspur leaf miner attack the leaves of the plant, and are not deterred by the alkaloids (Pirone, 1978). 

http://www.ces.ncsu.edu/depts/ent/notes/Ornamentals_and_Turf/flowers/note09/ColLeafminer.jpg

http://bygl.osu.edu/content/fourlined-plant-bugs-2

Works Cited

Frank, Steven. "Leafminer Flies." Ornamentals and Turf. North Carolina State University, Jan. 2011. Web. 9 Apr. 2014.

Gosselin, Matthias, Denis Michez, Maryse Vanderplanck, Dorothee Roelants, Gaetan Glauser, and Pierre Rasmont. "Does Aconitum Septentrionale Chemically Protect Floral Rewards to the Advantage of Specialist Bumblebees?" Ecological Entomology 38 (2013): 400-07. Zoologie.umh. The Royal Entomological Society, 20 May 2013. Web. 9 Apr. 2014.

The Ohio State University Extension. "Fourlined Plant Bugs." Buckeye Yard & Garden Online. The Ohio State University Extension, 9 May 2013. Web. 9 Apr. 2014.

Pirone, Pascal. Diseases and Pests of Ornamental Plants. 5th ed. N.p.: John Wiley & Sons, 1978. Google Books. Web. 9 Apr. 2014.

Flowers and Pollinators

The flowers of Aconitum delphinifolium are unique in that the top sepal resembles the hood worn by monks, hence its common name monkshood. The showy flower is bisexual, and is composed of 5 violet sepals. There are 2 petals recessed under the hood in the form of nectaries. With the nectaries protected under the hood, and its stamens and pistils located at the base of the other four sepals, it is especially suited to be pollinated by bumblebees (SEINet). The bumblebee must land on the landing platform that contains the stamens and pistils, and must struggle between the sepals to reach the reward in the nectaries using its long tongue. The distance to the nectaries makes it available only to long tongued bumblebees. To reduce the chance of self-fertilization, the stamens reach maturity prior to the carpels, and can range from 25-50 per flower. There are usually 3 pistils per flower, with 10-20 ovules per pistil (SEINet). 

 http://watchingtheworldwakeup.blogspot.com/2009/08/valley-of-death-and-2-kindred-spirits.htm

(Bumblebee pollinating monkshood) 

http://cabinetofcuriosities-greenfingers.blogspot.co.uk/2012/07/bee-magnet.html

 http://watchingtheworldwakeup.blogspot.com/2009/08/valley-of-death-and-2-kindred-spirits.htm

Works Cited

Gates, Phil. "Bee Magnet." Cabinet of Curiosities. Blogspot.com, 17 July 2012. Web. 26 Mar. 2014. <http://cabinetofcuriosities-greenfingers.blogspot.co.uk/2012/07/bee-magnet.html>.

Southwest Environmental Information Network. "Aconitum." SEINet-Aconitum. Swbiodiversity.org, n.d. Web. 26 Mar. 2014. <http://swbiodiversity.org/seinet/taxa/index.php?taxon=4469&proj=Arizona>.

Watcher. "The Valley of Death, And 2 Kindred Spirits." Watching the World Wake Up. Blogspot.com, 5 Aug. 2009. Web. 26 Mar. 2014. <http://watchingtheworldwakeup.blogspot.com/2009/08/valley-of-death-and-2-kindred-spirits.html>.

Growth

Aconitum delphinifolium has biennial tubers that overwinter and break dormancy in the spring. These tubers send roots and shoots out by apical meristems. The tuberous roots consist of abnormal secondary growth where the secondary xylem appears to be “U” shaped, and parenchyma cells of secondary phloem make up the vast majority of the tuber (JianZhao, JiYaun, and Qing 2011). Rhizomes are produced from lateral buds on the tuber, which will form a daughter tuber at its extremity for the next year’s plant. Secondary xylem is produced in the stems, and a cork layer is found in at least one species of Aconitum (Dulin 2008). It is an herbaceous perennial with the parent plant only surviving one season, so it does not produce growth rings. It is unknown whether A. delphinifolium produces bulbils, but several other Aconitum species have been discovered with them. Bulbils are basically miniature bulbs, and they can be found growing in the leaf axils of some Aconitum species. They are commonly produced if the plant has failed to produce seeds, giving it three reproduction strategies (Farrell 2008). 

Works Cited

Dulin, Max. "An Investigation of Paedomorphic Secondary Xylem and Secondary Woodiness in Xanthoriza simplicissima, Coreopsis gigantea, and Mahonia bealei." Thesis Defense. Greensboro: University of North Carolina, 2008.

Farrell, Bonnie. "Histological Documentation and Ecological Implications of Bulbils in Aconitum noveborecense (Ranunculaceae), A Federally Protected, Threatened Plant Species." Thesis Defense. New Brunswick: Rutgers, October 2008.

JianZhao, Qi, Guo JiYaun, and Xu Qing. "Abnormal secondary growth and histochemical localization of alkaloids in root system of Aconitum flavum." Agricultural Science & Technology, 2011: 1313-1317.

Roots

Aconitum is a perennial with a tuberous root system. It consists of a conical tap root that is usually no more than 2 cm in diameter at its thickest point, and about 10 cm long. Each year a rhizome is produced from a lateral bud in the parent tuber, and a new tuber is formed which will have the bud for the next years plant. Often another stem tuber is produced in the late winter or early spring producing two new plants the next season (Culbreth 1927).

http://7song.com/photo?photo=240&q=Category:%20Medicinal&l=Aconitum-columbianum&g=Ranunculaceae 

Aconitum delphinifolium has a symbiotic relationship with vesicular arbuscular myccorrhizae which helps in the uptake of nutrients and water from the soil (Treu et al. 1995). The roots of Aconitum have the highest levels of the alkaloid aconitine, and are used in eastern medicine as a remedy for multiple conditions. However, the level of toxicity varies between species, and accidental poisoning is not uncommon (Chemistry 2014). 

Works Cited

7 song. "Monkshood." Northeast School of Botanical Medicine. June 13, 2006. http://7song.com/photo?photo=240&q=Category:%20Medicinal&l=Aconitum-columbianum&g=Ranunculaceae (accessed February 26, 2014).

Chemistry, American Association for Clinical. Monkshood. 2014. https://www.aacc.org/members/divisions/tdm/library/Pages/feb09-toxin.aspx (accessed February 26, 2014).

Culbreth, David. A Manual of Materia Medica and Pharmacology. Lea Brothers & Co., 1927.

Treu, R., G. Laursen, S. Stephenson, J. Landolt, and R. Densmore. "Mycorrhizae from Denali National Park and Preserve, Alaska." Mycorrhiza, 1995: 21-29.

Water

Aconitum delphinifolium is found in northern climates and prefers moist well drained soils. It is a sun loving plant, and its leaves are hypostomatic to prevent water loss (Stanescu et al, 2009). In warmer climates it performs better with some shade since it does not tolerate heat or water stress very well. It is most often found in moist meadows, and along streams which have cool soil conditions (CYSIP).

http://www.pyreneanway.com/blog/2010/08/eyne-to-nuria-and-back/?lang=en

 Since Aconitum is a perennial that grows from rootstock, it grows rapidly in the spring after the snow melts when there are ample water conditions. If water stressed the bottom leaves are often shed (Drinkwater, 2009). This makes sense because these leaves are the largest, and lose the most water through transpiration. 

References

CYSIP. "Aconitum Delphinifolium : Northern Monkshood." Flora.dempstercountry.org. Central                     Yukon Species Inventory Project, n.d. Web. 5 Feb 2014.

Cracknell, Steve. "Eyne to Núria and Back." Pyreneanway.com. Wordpress, 20 Aug. 2010. Web. 24 Feb. 2014.

Drinkwater, Robyn. "Doff Your Hat." Botanics Stories. Edinburgh Botanics, 25 Sept. 2009. Web. 24 Feb. 2014.

Stanescu, I., C. Mardari, and C. Tanase. "Notes on the morpho-anatomy of Aconitum degenii gayer." Biologie Vegetala, 2009: 55-63.

Leaves

Aconitum delphinifolium is a deciduous perennial plant that grows rapidly in the spring using the nutrition stored in its roots, this allows it to fully develop its leaves before many other plants. Its leaves are palmately divided into usually 5 segments with each segment being again divided into multiple segments giving it a needlelike appearance. 

http://www.flora.dempstercountry.org/V.B.08.Ranunculaceae/Aco.delphi/Aco.delphi.pic4.jpg

The deeply divided leaf shape is a common characteristic of many plants growing in cold and arid environments. Possible adaptation to improve hydraulic efficiency?  They are alternately arranged with the lowest leaves being the largest (25 cm), and extend from the stem by long petioles. Upper leaves tend to be smaller (5cm) and are often attached directly to the stem lacking a petiole (CYSIP). 

http://analogicalplanet.com/Pages/ContentPages/Vascular%20plants2/AconitumDelphinifolium.html

This adaptation likely allows the plant to capture the most sunlight with the fewest amounts of leaves. All of the leaves tend to be hypostomatic with the majority of their stomates located on the bottom (Stanescu et al, 2009). The alkaloid aconitine is found in the leaves which may aid as a deterrent to herbivory. 

References

CYSIP. "Aconitum Delphinifolium : Northern Monkshood." Flora.dempstercountry.org. Central                     Yukon Species Inventory Project, n.d. Web. 5 Feb 2014.

Stanescu, I., C. Mardari, and C. Tanase. "Notes on the morpho-anatomy of Aconitum degenii gayer." Biologie Vegetala, 2009: 55-63.

Aconitum delphinifolium seeds and germination

Aconitum delphinifolium flowers contain 3 carpels with approximately 10-20 ovules in each. After pollination the seeds are contained in an aggregate of 3 follicles about 1.5-2 cm in length. The number of seeds per follicle is variable, ranging from 8-12. Each seed weighs approximately 1 gram, and is stored in dry capsules that split upon maturity releasing the seeds. Since these seeds are toxic and too heavy for wind dispersal it seems that the most likely method of dispersal is by water (Molero and Puig, 1990). Seeds stored at Royal Botanic Kew Gardens for the Millennium Seed Bank Project had a 78% germination rate after being dried and froze for over 2 years.

Photo: http://www.cdhs.us/Flower%20Project/Family%20Index/Ranunculaceae%20Index/A.%20delphnifolium/A.%20delphnifolium%20Fset.htm

Aconitum seeds are dormant upon maturation and must be frozen for a period of time before being able to germinate. A study done on wild Aconitum seeds showed that those not exposed to cold temperatures failed to germinate, and those with the longest stratification periods had the highest germination rate. The period of stratification was also correlated to how quickly the seed germinated. Those with the longest duration of chilling took the shortest amount of time to germinate. Seeds that were chilled for 3 weeks took the same amount of time to germinate (21 days), whereas, seeds that were chilled for 84 days only took 11 days to germinate (Dosmann, 2002). This is a good adaptation to have to ensure that springtime has actually arrived. 

References:

Dosmann, Michael. "Stratification Improves and Is Likely Required for Germination of Aconitum Sinomonantum." Hort Technology 12.3 (2002): 423-25. Horttech. ASHS Publications. Web. 30 Jan. 2014.

Molero, Julia, and Antonieta Puig. "Seed Morphology of Iberian Species of the Genus Aconitum L." Collectanea Botanica (1990): 111-27. Diposit Digital. Web. 30 Jan. 2014.

Royal Botanic Gardens Kew. (2014) Seed Information Database (SID). Version 7.1. Available from: http://data.kew.org/sid/ (January 2014)

My Plant #1

Image: http://www.ontariowildflower.com/yukon_monkshood_keno.htm

Aconitum delphinifolium DC. is native to Alaska and is commonly referred to as larkspurleaf monkshood, or just monkshood. I became interested in monkshood because it has a rich history dating back to Greek mythology. All parts of the plant contain the alkaloid aconitine which is extremely toxic. In Greek mythology it was said that Aconitum grew on the hill of Aconitus, and it became poisonous after Cerberus, the multi-headed guard dog of Hades, fought with Hercules and its saliva fell on the plant (Kowalchik, et al). From that point on, Aconitum was a popular poison to use in hunting and warfare. The early Aleut whalers in Alaska used A. delphinifolium to prepare a poison they used on the end of their spears, reducing the effort needed to harvest the whale (Heizer). It was also used in medicine as an anesthetic, and is still used today by some as a topical pain reliever.

Monkshood can grow up to 3 feet tall, but often depends on neighboring plants for support to stay upright. Due to its toxicity it is mostly left alone by larger herbivores, however, there are some insects that depend on it as a food source. Some bumble bees use it exclusively as a food source, and in return provide pollination (Boland). 

Image: http://analogicalplanet.com/Pages/ContentPages/Vascular%20plants2/AconitumDelphinifolium.html

Rank	Scientific Name and Common Name
Kingdom	Plantae – Plants
Subkingdom	Tracheobionta – Vascular plants
Superdivision	Spermatophyta – Seed plants
Division	Magnoliophyta – Flowering plants
Class	Magnoliopsida – Dicotyledons
Subclass	Magnoliidae
Order	Ranunculales
Family	Ranunculaceae – Buttercup family
Genus	Aconitum L. – monkshood
Species	Aconitum delphiniifolium DC. – larkspurleaf monkshood

Image: http://plants.usda.gov/core/profile?symbol=ACDE2

There are over 250 species in the Aconitum Genus, with  A. delphinifolium found solely in the Northwestern part of North America. The leaves of the plant look quite similar to the common delphinium (larkspur), which is where it got its name. A. dephilnifolium was named by Augustin Pyramus de Candolle (USDA). It gets its common name monkshood due to the fact that a sepal on the flower resembles the old fashion hoods worn by monks. 

Image: http://www.flora.dempstercountry.org/0.Site.Folder/Species.Program/Species2.php?species_id=Aco.delphi

References

Boland, Todd. "Monkshood-A Deadly Beauty." Mun.ca. Memorial University, n.d. Web. 23 Jan. 2014. <http://www.mun.ca/botgarden/gardening/gardennl/Monkshood_-_A_Deadly_Beauty.pdf>.

CYSIP. "Aconitum Delphinifolium : Northern Monkshood." Flora.dempstercountry.org. Central Yukon Species Inventory Project, n.d. Web. 23 Jan. 2014. <http://www.flora.dempstercountry.org/0.Site.Folder/Species.Program/Species2.php?species_id=Aco.delphi>.

Heizer, Robert F. "Pacific Eskimo Invention in Whale Hunting." Escholarship.org. University of California, n.d. Web. 23 Jan. 2014. <http://escholarship.org/uc/item/9780c95n#page-3>.

Kowalchik, Claire, William H. Hylton, and Anna Carr. "Aconite." Rodale's Illustrated Encyclopedia of Herbs. Emmaus, PA: Rodale, 1987. 1-2. Google Books. Web. 23 Jan. 2014. <http://books.google.com/books?id=htGD3Y7WNxwC&pg=PA1&source=gbs_toc_r&cad=4#v=onepage&q&f=false>.

USDA. "Plants Profile for Aconitum Delphiniifolium (larkspurleaf Monkshood)." Plants.usda.gov. USDA, n.d. Web. 23 Jan. 2014. <http://plants.usda.gov/core/profile?symbol=ACDE2>.