LEAF LITTER ARTHROPODS OF MESOAMERICA (LLAMA)

2009 LLAMA expeditions in Guatemala.
2008 LLAMA expeditions in Chiapas, Mexico.
Preliminary results for Chiapas baiting samples.
Ants of Chiapas.
Results of 2007 Reconnaissance trip to Chiapas and Guatemala.

LEAF LITTER ARTHROPODS OF MESOAMERICA (LLAMA)

Terrestrial arthropods make up a large fraction of global biodiversity but are among the least known of multicellular organisms. In the American tropics, Costa Rica has been a focus for inventory effort on a wide variety of arthropods (INBio, Project ALAS), but we remain ignorant of biodiversity patterns at larger spatial scales. In particular we know very little of species richness, degrees of endemism, and patterns of faunal turnover across the rest of Mesoamerica, a region scientists have long recognized as one of the most biogeographically interesting and imperiled landscapes on the planet. LLAMA is a program of arthropod sampling that will build on Costa Rican knowledge and extend it from Nicaragua to southern Mexico, in both lowland and highland areas. The sampling from highland areas is particularly important since these are known to be conservation priorities because of high endemism levels and vulnerability to global warming.

LLAMA will focus on two signature taxa - ants (Formicidae) and weevils (Curculionidae) - and one subcomponent of tropical forest ecosystems - the leaf litter layer of the forest floor. Ants and weevils are diverse, are important in ecosystem-level processes, represent different trophic guilds, have a well-developed taxonomic infrastructure on which to build, and have a large user community among ecologists and evolutionary biologists. Forest floor leaf litter is an easily sampled zone of concentrated biodiversity. Over 60% of total ant diversity and numerous poorly-known but diverse radiations of weevils live in this thin layer at the interface of soil and vegetation. It is an important zone for carbon movement, where carbon is either respired or stored in the soil, and litter arthropods play an important role in litter decomposition and dynamics.

Following reconnaissance and initial taxonomic work in year 1, LLAMA will sample four regions over four years: southern Mexico, Guatemala, Honduras, and Nicaragua. Each year the project will sample three mountains or mountain ranges, with low, medium, and high elevation sites on each mountain. At each site two plots will be sampled, one in mature and one in secondary forest. Each plot will be sampled with a combination of quantitative methods that allow diversity analysis and non-quantitative methods that maximize species capture. Each year there will be a 6-week field season, followed by 2-weeks of sample processing, workshops, and lectures at a Latin American host institution. The field crew will consist of graduate and undergraduate students from both PI and host institutions. All ants and weevils will be sorted and non-focal taxa will be distributed to collaborating taxonomists. At the end of the project, prepared specimens will be distributed to host institutions and other research collections.

All specimen data will be made electronically available. Each year all species and morphospecies of Formicidae and Curculionidae will be reported on-line, complete with Automontage images. For ants, special emphasis will be placed on the systematics of the hyperdiverse taxa Pheidole and Dacetini, and a thorough revision and molecular phylogenetic analysis of Mesoamerican Stenamma. For weevils the focus will be on members of the Lymantini, including descriptions of a number of known new generic lineages, and a thorough revision and phylogenetic analysis of the cryptorhynchine genus Tylodinus.

Broader impacts of the project are in education, international collaboration and development, and biodiversity conservation. Each year an international team of students will be immersed in a field experience with taxonomists, encouraging their development through training, inspiration, guidance, and social networks. Two PhD candidates will receive support and training for their research programs. PI's Longino and Anderson, in conjunction with collaborators from Latin American host institutions, will plan and deliver lectures and workshops each year during the sample processing time, fostering international scientific ties that will greatly benefit global biodiversity science. The scientific research in reserves and remaining forest patches will raise their status as sites that can attract economic activity, and the results on species turnover and endemism will inform conservation decisions and highlight the biotic importance of Mesoamerican cloud forests.

Mexico: El Colegio de la Frontera Sur, ECOSUR, San Cristobal de las Casas, Chiapas (http://www.ecosur.mx/). ECOSUR is a postgraduate institution of multidiscplinary research and education in the southern border regions of Mexico. There are three speciality areas of study, one of which is biodiversity conservation.

Expedition Year: 2008.

Collaborator: Dr. Jorge Leonel León Cortés, Lepidoptera biodiversity and systematics; conservation biology of fragmented habitats.

E-mail: jleon@sclc.ecosur.mx

Guatemala: Universidad del Valle, Guatemala City (http://www.uvg.edu.gt/), and Universidad de San Carlos de Guatemala (http://www.usac.edu.gt/). UVG and USAC are the main institutions of advanced learning in Guatemala. The UVG Department of Biology has a focus on insect biodiversity and systematics. USAC has an active biodiversity research group and plays a key role in managing natural areas in the country.

Expedition Year: 2009.

Collaborator: Dr. Jack Schuster, Biodiversity, systematics and evolution of Passalidae (Coleoptera).

E-mail: jschuste@uvg.edu.gt

Honduras: Regional Biodiversity Institute, Pan-American School of Agriculture (Centro Zamorano de Biodiversidad), Zamorano (http://www.zamorano.edu/). The Pan-American School of Agriculture is an internationally recognized and accredited institution dedicated to the responsible management of natural resources. It is the home of the Regional Biodiversity Institute, a recent initiative to integrate biodiversity knowledge and capacity building across Central America.

Expedition Year: 2010.

Collaborator: Dr. Jorge Ivan Restrepo, Administrator at Centro Zamorano de Biodiversidad.

E-mail: restrepo@alumni.ksg.harvard.edu

Nicaragua: Museo Entomologico, Leon (http://www.bio-nica.info/topic/index.html).

Expedition Year: 2011.

Collaborator: Dr. Jean-Michel Maes, Biodiversity and systematics of insects, Curator of insect collection.

E-mail: jmmaes@ibw.com.ni

Bulk samples of non-focal taxa will be distributed to five taxonomic collaborators. They will maintain a distribution network for material. If you have questions about access to this material, please contact the collaborator directly.

Arachnida, Myriapoda	Gonzalo Giribet Museum of Comparative Zoology Harvard University, 16 Divinity Avenue Cambridge, MA 02138, USA ggiribet@oeb.harvard.edu
Hemiptera	Harry Brailovsky Departamento de Zoología, Instituto de Biología, U.N.A.M. Circuito Exterior s/n, Ciudad Universitaria, Copilco, Coyoacán A.P. 70-233 México, Distrito Federal. C.P. 04510 coreidae@ibiologia.unam.mx
Coleoptera	Zachary Falin Division of Entomology, KUNHM/BRC Snow Entomological Collection 1501 Crestline Dr. Suite 140 University of Kansas Lawrence, Kansas 66049 ksem@ku.edu
Diptera	Steve Marshall Department of Environmental Biology University of Guelph Guelph, ON, Canada, N1G 2W1 smarshal@uoguelph.ca
Hymenoptera	Michael Sharkey Department of Entomology S-225 Ag. Sci. North University of Kentucky Lexington, KY 40546-0091, USA msharkey@uky.edu

Winkler samples are an efficient method of sampling leaf litter ants. The method involves sifting bulk samples of leaf litter and rotten wood by agitating them vigorously in a bag above a coarse mesh screen. Litter arthropods are concentrated in the finer "siftate" that passes through the screen. Arthropods are then extracted from the siftate by a passive extraction method, in which the siftate is placed in thin mesh sacks and then suspended and enclosed within an outer cloth "Winkler bag." The Winkler bag tapers to a cup of ethanol. After loading the sample the Winkler bag is closed at the top and suspended in a sheltered location. Arthopods fall from the litter and accumulate in the ethanol, and the sample is taken off after 3 days.

Methodological variations involve how the bulk litter is selected. A "MaxiWinkler" sample involves a subjective selection of litter to sift, and is not based on a particular area or volume of bulk litter. The person taking the sample moves through the habitat selecting pockets of moist litter. These accumulations of litter and rotten wood occur between buttresses, along the sides of rotten logs, around dead stumps, and in soil depressions. When approximately 6l of siftate is obtained, the material is placed in a cloth bag and returned to the lab or field camp to be suspended in a Winkler bag.

A "miniWinkler" sample involves sifting the litter from a one square meter quadrat. The litter and dead wood in the quadrat are first aggressively minced with a machete. Then the litter is sifted until all the litter in the quadrat is sifted or 2l of siftate are obtained, whichever comes first. When there is more than enough litter to produce 2l of siftate, the different kinds of litter in the quadrat (e.g. leaves on soil versus litter from a rotten log) are subsampled so that all are represented (a somewhat subjective process that attempts to include the diversity of litter types found in the quadrat). Quadrats are placed according to a protocol of transects, grids, stratification, randomization, or other procedures. Thus the method is much less subjective and less dependent on the habits of the sifter. However, miniWinkler samples are less efficient than maxiWinklers in terms of species capture per unit of effort.

At each field site two transects of miniWinkler samples will be taken. Each transect will be 245m long and contain 50 quadrats at 5m spacing. Starting point and direction of transect will be chosen subjectively, attempting to keep the transect within a habitat patch. If the patch is too small, or an impassable area is encountered, the transect may change direction or be broken into more than one segment.

Each year there will be an opportunity for four U.S. undergraduates to be part of the field sampling crew.

The work: Students will be part of a field crew composed of 4 U.S. undergraduates, 4 students (or equivalent) from host institutions in the countries where we will be working, and two graduate students. Undergraduates will be under the direct supervision of the graduate students. Anderson and Longino will be present during the first and last two weeks of the field season. Travel will be from mid-April to mid-June. The first 45 days will be spent in field, working 4 days per site, with travel days in between, to cover a total of 9 sites. On arrival at a site, field crews will spend the first day taking a series of samples of sifted leaf litter from meter-square plots and hanging them up for extraction. The following three days will be spent applying various other insect sampling techniques in the study plots.

This work will be physically demanding. You will need to carry bulky sampling supplies into rugged terrain. Some sites may be swelteringly hot, others cold and rainy. You will need to move off trails into dense forested habitats. Long hikes may be required to reach sampling sites. There will be mud and mosquitos. If you are applying, this should sound like fun to you.

Housing is likely to be a combination of cheap hotels in small towns, rustic field stations, and possibly camping.

Following the field work at the 9 field sites, the field crew will move to the host institution for a 10-day period of workshops and sample sorting. There will be workshops and lectures on the biology and taxonomy of weevils and ants, and on quantitative biodiversity inventory methods. During this time all samples will be processed and sorted into a standard set of fractions. Training will be provided in how to sort the samples. Applicants must be comfortable with using dissecting microscopes, have sufficient fine motor skills to separate tiny arthropods into separate vials, and have the patience to sit and sort samples for about 10 days.

The reward: the grant will cover round trip airfare to the host country, all food and lodging costs during the field and lab work, and a stipend of $800. Evergreen students will get 16 units of upper division science credit for Spring Quarter. You will get to work in some of the most interesting and diverse habitats on the planet. You will work with professional entomologists and conservation biologists. You will develop a network of friends and colleagues among your fellow students, the graduate students, senior researchers, and all the other people you will meet in the host country. You will learn some Spanish. You will learn a lot about tropical arthropods.

Evergreen students: Evergreen students will be given strong preference. Only if there are not enough suitable candidates from Evergreen will non-Evergreen students be considered. Students will have an independent contract with Longino for Spring quarter, receiving 16 units of upper division science credit.

The 2010 field season: The 2010 field work will be carried out in Honduras and will be approximately the months of May and June. Similar opportunities will be available in 2011 (Nicaragua).

The application: We are looking for students with a sincere interest in the study of tropical arthropods. If you are not interested in arthropods you will find the work dull and boring.

In your application, please include the following:
1. Name and contact information.
2. Educational background.
3. An essay that discusses why you are interested in the position, how the experience could help your career goals, and what relevant qualifications and experience you have. (Two page limit)
4. Spanish language ability
5. Field experience (backpacking, camping, hiking, etc.)
6. Names and contact information for 2 people willing to be references for you. References should be college teachers (include what courses you took with them and when), employers, or supervisors.

Due date: 16 November 2009. Submit applications electronically to John Longino, longinoj@evergreen.edu. Decisions will be made by early December 2009.

Branstetter, M. G. 2009. The ant genus Stenamma Westwood (Hymenoptera: Formicidae) redefined, with a description of a new genus Propodilobus. Zootaxa 2221:41-57.

Longino, J. T. 2009. Additions to the taxonomy of New World Pheidole (Hymenoptera: Formicidae). Zootaxa 2181:1-90.

Quinet, Y., R. Hamidi, M. X. Ruiz-Gonzalez, J.-C. de Biseau, and J. T. Longino. 2009. Crematogaster pygmaea (Hymenoptera: Formicidae: Myrmicinae), a highly polygynous and polydomous Crematogaster from northeastern Brazil. Zootaxa 2075:45-54.

Kumar, A., J. T. Longino, R. K. Colwell, and S. O'Donnell. 2009. Elevational patterns of diversity and abundance of eusocial paper wasps (Vespidae) in Costa Rica. Biotropica 41:338-346.

Longino, J. T., D. J. Cox. 2009. Pheidole bilimeki reconsidered (Hymenoptera: Formicidae). Zootaxa 1985:34–42.

Astrin, J. J., and P. E. Stüben. 2008. Phylogeny in cryptic weevils: molecules, morphology and new genera of western Palaearctic Cryptorhynchinae (Coleoptera : Curculionidae). Invertebrate Systematics 22:503–522.

Colwell, R. K., G. Brehm, C. L. Cardelus, A. C. Gilman, and J. T. Longino. 2008. Global warming, elevational range shifts, and lowland biotic attrition in the wet tropics. Science 322:258-261.

Adams, R. M. M., and J. T. Longino. 2007. Nesting biology of the arboreal fungus-growing ant Cyphomyrmex cornutus and behavioral interactions with the social-parasitic ant Megalomyrmex mondabora. Insectes Sociaux 54:136-143.

Longino, J. T. 2007. A taxonomic review of the genus Azteca (Hymenoptera: Formicidae) in Costa Rica and a global revision of the aurita group. Zootaxa 1491:1-63.

Longino, J. T., and F. Fernández C. 2007. A taxonomic review of the genus Wasmannia. Pages 271-289 in R. R. Snelling, B. L. Fisher, and P. S. Ward, editors. Advances In Ant Systematics (Hymenoptera: Formicidae): Homage To E. O. Wilson – 50 Years Of Contributions. Memoirs of the American Entomological Institute, 80.

Dunn, R. R., N. J. Sanders, M. C. Fitzpatrick, E. Laurent, J.-P. Lessard, D. Agosti, A. N. Andersen, C. Bruhl, X. Cerda, A. M. Ellison, B. L. Fisher, H. Gibb, N. J. Gotelli, A. Gove, B. Guenard, M. Janda, M. Kaspari, J. T. Longino, J. Majer, T. P. McGlynn, S. B. Menke, C. L. Parr, S. M. Philpott, M. Pfeiffer, J. Retana, A. V. Suarez, and H. L. Vasconcelos. 2007. Global ant (Hymenoptera: Formicidae) biodiversity and biogeography – a new database and its possibilities. Myrmecological News 10:77-83.

Longino's database of over 87,000 species occurrences in over 2200 species has been integrated with AntWeb at the California Academy of Sciences, providing fully georeferenced specimen data available for browsing on Google Earth.

Longino continues to develop the Pheidole working group. Web-based resources include a complete taxonomic catalogue in the form of a down-loadable parsed Excel file, an illustrated glossary of morphology, and a Lucid key to the nearly 700 New World Species.

Longino's Ants of Costa Rica website continues to grow and improve, the latest addition being a treatment of the genus Azteca.

This work is supported by National Science Foundation grant DEB-0640015, Biodiversity Surveys and Inventories Program. Any publications that rely on the LLAMA project for specimens or other forms of support should include a statement to that effect, with the grant number, in the Acknowledgments. Please send the full bibliographic reference of any such publications, preferably in a form that can be imported into EndNote, to John T. Longino longinoj@evergreen.edu.

NSF Acknowledgement and Disclaimer

This web site is based upon work supported by the National Science Foundation grant DEB-0640015.

Any opinions, findings and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation (NSF).