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Daniel H. Janzen costaricanadventures.com Community Ecology Brittany Owens Daniel Janzen: C.V. Degree in Biology from University of Minnesota in 1961 Ph.D. from University of California, Berkeley in 1965 Teaching positions (US): -(1965-1968) University of Kansas -(1969-1972) University of Chicago -(1972-1976) University of Michigan -(1976- present) University of Pennsylvania Teaching positions (outside US): (1965-1966): Universidad de Oriente, Cumana (Venezuela) (1969): Universidad de Puerto Rico, Rio Piedras (Puerto Rico) (1973): Universidad de Los Andes (Venezuela) www.costaricanadventures.co OTS Affiliation: (1963) Student at OTS tropical biology course in Costa Rica (1965) Returned as an instructor -since has served annually as an instructor Influenced work in tropics, particularly Costa Rica www.adovsky.blogspot.com Conservation/Restoration Projects • Currently ad honorem technical advisor for Area de Conservacion Guanacaste (ACG) and Instituto Nacional de Biodiversidad (INBio) acguanacaste.ac.org Distinctions/Awards • • • • • • • • • • • • • • • 1975 Gleason Award (Amercian Botanical Society) 1984 Crafoord Prize, Coevolutionary ecology (Royal Swedish Academy of Sciences) 1987 Hijo Ilustre de Guanacaste (Governor of Guanacaste province) 1897 “Global 500” Roll of Honor (UNEP) 1989 MacArthur Fellowship 1991 Founder’s Council Award of Merit (Field Museum of Natural History) 1992 Member, National Academy of Sciences (USA) 1993 Award for Improvement of Costa Rican Quality of Life (Universidad de Costa Rica) 1994 Silver Medal Award (International Society of Chemical Ecology) 1995 Conservation Society Award 1997 Kyoto Prize, Basic Sciences Field (Inimori Foundation) 2002 Albert Einstein World Award for Science (Consejo Cultural Mundial, Mexico) 2002 Honorary Fellow of the Association for Tropical Biology, and Conservation (ATBC) 2006 National Outdoor Book Award, Design and Artistic Merit for 100 Caterpillars 2011 FBBVA Foundation Frontiers of Knowledge Award of Ecology and Conservation Biology 220 publications 55 journals ~10,000 citations • • • • • • • • • • • • • • • • • • • • • • • Publications Smith, M.A., Rodriguez, J.J., Whitfield, J.B., Deans, A.R., Janzen, D.H., Hallwachs, W., and Hebert, P.D.N. 2008. Extreme diversity of tropical parasitoid wasps exposed by iterative integration of natural history, DNA barcoding, morphology, and collections. Proceedings of the National Academy of Sciences 105:12359-12364. Burns, J.M., Janzen, D.H., Hajibabaei,M., Hallwachs,W., and Hebert, P.D.N. 2008. DNA and cryptic species of skipper butterflies in the genus Perichares in Area de Conservacion Guanacaste, Costa Rica. Proceedings of the National Academy of Sciences 105:6350-6355. Miller, J. C., Janzen, D. H. and Hallwachs, W. 2007. 100 Butterflies and moths. Harvard University Press, Cambridge, Massachusetts. 256 pp. Smith, M. A., Woodley, N. E., Janzen, D. H., Hallwachs, W., and Hebert, P. D. N. 2006. DNA barcodes reveal cryptic host-specificity within the presumed polyphagous members of a genus of parasitoid flies (Diptera: Tachinidae). Proceedings of the National Academy of Sciences 103:3657-3662. Miller, J. C., Janzen, D. H. and Hallwachs, W. 2006. 100 Caterpillars. Harvard University Press, Cambridge, Massachusetts. 264 pp. Janzen, D. H. 2005. How to conserve wild plants? Give the world the power to read them. Forward, Plant conservation: a natural history approach, eds. Krupnick, G. and Kress, J., University of Chicago Press, Chicago, 346 pp. Janzen, D. H., Hajibabaei, M., Burns, J. M., Hallwachs, W., Remigio, E. and Hebert, P. D. N. 2005. Wedding biodiversity inventory of a large and complex Lepidoptera fauna with DNA barcoding. Philosophical Transactions of the Royal Society B 360 (1462):1835-1846. Hebert, P. D. N., Penton, E. H., Burns, J. M., Janzen, D. H. and Hallwachs, W. 2004. Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences 101:14812-14817. Janzen, D. H. 2002. Tropical dry forest: Area de Conservación Guanacaste, northwestern Costa Rica. In Handbook of Ecological Restoration, Volume 2, Restoration in Practice, eds. Perrow, M. R., Davy, A. J., Cambridge University Press, Cambridge, UK, pp. 559-583. Schauff, M. E. and Janzen, D. H. 2001. Taxonomy and ecology of Costa Rican Euplectrus (Hymenoptera: Eulophidae), parasitoids of caterpillars (Lepidoptera). Journal of Hymenoptera Research 10(2):181-230. Janzen, D. H. 2001. Saving fractured oases of biodiversity (book review). Quarterly Review of Biology 76:327-330. Burns, J. M. and Janzen, D. H. 2001. Biodiversity of pyrrhopygine skipper butterflies (Hesperiidae) in the Area de Conservación Guanacaste, Costa Rica. Journal of the Lepidopterist's Society 55:15-43. Janzen, D. H. 2000. Wildlands as gardens. National Parks Magazine 74(11-12):50-51. Janzen, D. H. 2000. Costa Rica's Area de Conservación Guanacaste: a long march to survival through non-damaging biodevelopment. Biodiversity 1(2):7-20. Janzen, D. H. 1999. Gardenification of tropical conserved wildlands: Multitasking, multicropping, and multiusers. PNAS 96(11):5987-5994. Janzen, D. H. 1999. La sobrevivencia de las areas silvestres de Costa Rica por medio de su jardinificación. Ciencias Ambientales No. 16:8-18. Janzen, D. H. 1998. Gardenification of wildland nature and the human footprint. Science 279:1312-1313. Janzen, D. H. and I. D. Gauld 1997. Patterns of use of large moth caterpillars (Lepidoptera: Saturniidae and Sphingidae) by ichneumonid parasitoids (Hymenoptera) in Costa Rican dry forest. In Forests and Insects, eds. A. D. Watt, N. E. Stork and M. D. Hunter, Chapman & Hall, London, pp. 251-271. Reid, W. V., S. A. Laird, R. Gómez, A. Sittenfeld, D. H. Janzen, M. A. Gollin and G. Juma. 1993. Biodiversity Prospecting. World Resources Institute, Washington, D.C. 341 pp. Janzen, D. H. 1993. Caterpillar seasonality in a Costa Rican dry forest. In: Caterpillars. Ecological and evolutionary constraints on foraging, N. E. Stamp and T. M. Casey, eds., Chapman and Hall, New York, pp. 448-477. Janzen, D. H. 1988. Guanacaste National Park: tropical ecological and biocultural restoration. In Rehabilitating Damaged Ecosystems, vol. 11, pp. 143-192, J. Cairns, Jr., ed., CRC Press, Boca Raton, Florida. Janzen, D. H. 1986. The future of tropical ecology. Ann. Rev. Ecol. Syst. 17, 305-24. Janzen, D. H. ed. 1983. Costa Rican Natural History, University of Chicago Press, Chicago, 816 pp. www.journalogy.edu www.bio.upenn.edu Research “My research over the past 55 years has evolved from a Victorian study of natural history of tropical animal-plant interactions to an exploration of the ecology of the interface between society and tropical wildland biodiversity. The administrative structure of institutions such as wildland administrations and Ministries of the Environment, the biodiversity development of conserved wildlands to where they can pay their own direct costs, and the integration of wildlands into society, are all major and essential tools in the engineering of the tropical countryside. Costa Rica as a whole is the ecosystem, and Area de Conservacion Guanacaste, in the northwestern corner, is the place. My research is done where the organisms are, i.e., Costa Rica.” -Dan Janzen www.bio.upenn.edu • “Victorian Era Natural History Studies” Coevolution of Plants and Insects – Ant-Acacia mutualisms in the Neotropics – Spondias mombin seed dispersal by mammals – Effect of herbivory on tropical plant diversity • “Exploration of the Interface between Humans and Biodiversity” Habitat Conservation/Restoration – Area de Conservacion Guanacaste (ACG) – Instituto Nacional de Biodiversidad (INBio) www.Ibol.org What is Coevolution? • “The term coevolution is used to describe cases where two (or more) species reciprocally affect each other’s evolution” – evolution.berkeley.edu • “Coevolution is a change in the genetic composition of one species in response to a genetic change in another.” – biomed.brown.edu Coevolution • Concept of coevolutionary “races” in ecologically intimate species Mimicry http://dx.doi.org/10.1016/j.cub.2005.11.046 Predation Mutualism heliconius.cam.zoo.ak Coevolution: Ant-Acacia Mutualisms ecolibrary.org “Ant-plants” and “Plant-ants” • “Swollen Thorn Acacias” • Genus Acacia (<10% of genus in C. America) – – – – Enlarged stipular thorns which house ants Enlarged foliar nectaries Modified leaflet tips (Beltian bodies) Year-round production of leaves (even in areas with dry season) experimentation.online.co.uk Ant-plants and Plant-ants • “Obligate acacia ants” • All in genus Pseudomyrmex – Solely build colonies in swollen thorn acacias *Taxonomic Issues *Not obligate on one species of acacia, but on the life form “Fully developed interdependency” 1. Queen ant finds unoccupied sapling and chews entrance hole. 2. Lays eggs in thorn and forages for foliar nectaries. 3. Forages for Beltian bodies to feed larvae. 4. Colony grows to occupy all thorns of plant. 5. Queen becomes physogastric with workers performing daily duties. *9 months of colony growth are required to produce enough workers to police area outside of acacia and provide protection for the plant. “Fully developed interdependency” Experimentally demonstrated that the swollen-thorn acacias have evolutionarily lost their ability to withstand insect damage and competition with neighboring plants in the absence of obligate acacia-ants (Janzen 1966a) -Experimentally removed ants from acacias -Total of 50 subplots were observed for 11 months -After 7-9 months, unoccupied acacias were compared with occupied acacias in control plots -Death of unoccupied acacias usually occurs within 6-12 months due to repeated defoliation via herbivores or overshadowing by neighboring vegetation www.entangledbank.com “Fully Developed Interdependency” Mean Sucker Length Presence of Phytophagous insects Mortality Weight of Biomass (Janzen 1996) Ant-Plant Traits • “In a sense, the obligate acacia-ants are the secondary plant substances and micromorphology of the swollenthorn acacias, and they are the means through which the swollen-thorn acacias interact with a large number of the other organisms in the community… consideration of the ant x acacia interaction system leads to the hypothesis that evolutionary abilities of the entire group of organisms in the community is responsible for the structure and properties of the plants in the community, especially at the microstructure level…” (Janzen 1996) Plant-Ant Traits • “The obligate acacia-ants are completely dependent, as a colony in nature, on the swollen-thorn acacias… The obligate acaciaants have committed themselves to the fate of the swollen-thorn acacias, provided that something happens to the swollen thorn acacias so rapidly that the obligate acacia-ants cannot evolve out of the interaction system.” (Janzen 1996) Scientificamerican.com Beyond Mutualism Looking beyond the single system of ant-acacia coevolution to discuss basic properties of such systems: 1. Coevolution of mutualistic coevolution is the product of evolutionary feedback systems. -one population must supply the other with a benefit, and the receptive population must be able to favor the donating genotype of the donating population - “increasing a flow from a trickle” Beyond Mutualism 2. Within the ant-acacia system, a hypothesis for the production of a coevolutionary relationship between species is provided by examples of convergence. -There are several species of Pseudomyrmex which exhibit a range of affinity for acacias. -There are at least five independent evolutions of the acacias into the swollenthorn lifestyle. Beyond Mutualism 3. This study adds support to the idea that the chemical and microphysical properties of plants may have partly evolved as a response to insect interactions. -The services provided by the acacia-ants appear to be a substitute for many of the physical and chemical traits that other acacias use in their defense. Beyond Mutualism 4. When looking at coevolution, preadaptations on both sides are important to facilitating the coevolutionary system. -Magnification or reduction of pre-interaction properties may have led to many of the traits essential to the ant x acacia mutualism. -However, the production and consupmtion of Beltian bodies, along with the aggression of ants towards neighboring plants do not fall under this category. Beyond Mutualism 5. The existence of such coevolutionary relationship contributes to our understanding of high plant density in the tropics. -Ant-acacia coevolution appears more common in warmer, wetter areas -As one moves away from the tropics, plantinsect interspecific interactions appear to decrease in density. Coevolution Summary “The interaction system between obligate acaciaants and swollen-thorn acacias is clearly an example of mutualism between a higher plant and an animal. The present deductive discussion of the evolution of the traits of the ant and the acacia leads without reasonable doubt the conclusion that the system evolved through mutually exclusive interactive evolution, or coevolution, rather than by chance coincidence in time and space of two highly specialized yet mutually beneficial organisms.” (Janzen 1966) Habitat Conservation/Restoration Area de Conservacion Guanacaste (ACG) • 163,000 hectare national park in Costa Rica (2% of the country’s total area) • 325,000 species (2.6% of the world’s biodiversity) • Includes diverse habitats including dry forest lowlands, cloud forests, rainforest, and marine habitat www.gdfc.org ACG • 1971 established as one of the first National Parks in Costa Rica • 1999 established as UNESCO World Heritage Site • “ACG conducts research and conservation action at the leading edge of ecology, evolutionary biology, biotechnology, biodiversity development, schoolchild education, decentralized administration, and integration with local, national and international society.” www.gdfc.org ACG • Guanacaste National Park (GNP) was created with the idea that it’s long-term survival would be supported by the creation of the idea of “ownership” – “owners” being both its direct custodians and society, at large • Mission Statement of GNP included: – Restoration via use of existing fragments – Inclusion of a large enough area to facilitate intense visitation and research use – Maintenance of forested area to provide a variety of material resources and biological data to the local culture – Exposure of the local, national, and international audience to the benefits of the natural world (Janzen 2000) Orange You Glad… • • • • • 20,000 hectares of pastureland were purchased to allow restoration back into hardwood forest In 1992 an industrial- level orange plantation was established along the northern boundary of the ACG on thousands of hectares of pastureland along its northern boundary. The ACG requested an experimental 100 truckloads of orange peels from the neighboring orchard to be dumped on the reclaimed pastureland, which aided in the beginning of the forest regeneration process. ACG then negotiated a contract with the orange plantation in which organisms from the park would degrade 1000 truckloads of peel a year for 20 years in the same manner, along with providing 20 years of other environmental services such as water and biological control. In return, orange farmers would pay the ACG with 1400 hectares of Del Oro forested land neighboring the ACG forests. Controversial, but successful so far! (Janzen 2010) Gmelina Plantations • Gmelina have traditionally resulted in deforestation and erosion. • However, unweeded Gmelina plantations overshadow pastureland, killing grasses and aiding in the regeneration of forest (typically difficult to start on such lands) • ACG has partnered with local Gmelina planters, which pay the cost of the plantation, leaving it unweeded and sharing profits with the ACG. • After one 8-12 year rotation of Gmelina the planter harvests the logs and ACG conservationists herbicide the stumps, leaving the remaining unweeded understory to grow up as rainforest. • Although controversial, a grant from a conservation NGO has now put this concept into practice in the eastern ACG, providing jobs, funds for further restoration, and community involvement. (Janzen 2010) INBio Inbio.ac.cr Questions www.vivelawordpress.com • • • • • • • • • • • • • • • "Acacias." Www.entangledbank.com. N.p., n.d. Web. 15 Apr. 2013. Website "Ants Collecting Food Bodies." Www.ecolibrary.edu. N.p., n.d. Web. 15 Apr. 2013. Website "Coevolution." Www.biomed.brown.edu. Brown Biomedical Sciences, n.d. Web. 15 Apr. 2013. Website Janzen, D.H. 2000. Costa Rica’s Area de Conservacion Guanacaste: a long march to survival through non-damaging biodevelopment.Biodiversity 1(2) Janzen, D.H. 2000. "Daniel H Janzen." Wikipedia. Wikimedia Foundation, n.d. Web. 15 Apr. 2013.Website "Daniel H Janzen." Www.jouralogy.edu. N.p., n.d. Web. 15 Apr. 2013.Website "Department of Biology." Department of Biology. N.p., n.d. Web. 15 Apr. 2013.Website " Guanacaste Dry Forest Conservation." Acguanacaste.ac.cr. N.p., n.d. Web. 15 Apr. 2013.Website Janzen, D.H. 2010. Hope for tropical biodiversity through true bioliteracy. Biotropica. Janzen, D.H. 1987. How to grow a national park.Experientia 43. "Instituto Nacional De Biodiversidad." Wikipedia. Wikimedia Foundation, n.d. Web. 15 Apr. 2013.Website Janzen, Daniel H. "Coevolution of Mutualism Between Ants and Acacias in Central America." Evolution 20.3 (1966): 249-75. Web.Journal Article Janzen, Daniel H. "The Interaction of the Bull's-horn Acacia (Acacia Cornier L.) with One of Its Ant Inhabitants (Pseudomyrmex Ferruginea F.Smith) in Easter Mexico." University California Publ. Ent (n.d.): n. pag. Print.Journal Article "Moreless." Guanacaste Dry Forest Conservation Fund. N.p., n.d. Web. 15 Apr. 2013.Website "Understanding Evolution." Understanding Evolution. N.p., n.d. Web. 15 Apr. 2013.