Development and Identity of Sexually Dimorphic Reproductive Signals

and Responses by African Elephants

A National Science Foundation Sponsored Research Project

Name, Institution and Addresses of PI's

Bruce A. Schulte, Ph.D., Department of Biology, Georgia Southern University, Statesboro, Georgia (NSF-DBI-02-17062)

Thomas Goodwin, Ph.D., Department of Chemistry, Hendrix College, Conway, Arkansas (NSF-DBI-02-16862)

L.E.L. (Bets) Rasmussen, Ph.D., Department of Biochemistry & Molecular Biology, Oregon Graduate Institute School of Science and Engineering, Oregon Health & Sciences University, Beaverton, Oregon (NSF-DBI-02-17068)

Student Involvement (see graduate student page for all graduate students in the Schulte lab)

GEORGIA SOUTHERN GRADUATE STUDENTS INVOLVED IN THE PROJECT:

Helen Loizi, MSc 2004. Helen conducted biological assays with male African elephants using compounds from follicular phase urine at the Riddle's Elephant Sanctuary in Arkansas. Helen also conducted field studies at Addo Elephant National Park examining the development of chemotactile behaviors in male and female elephants. Helen completed her thesis in 2004 (available as pdf through the Henderson Library, Thesis link)

Katie Bagley, MSc 2004. Katie was supported on the grant from May 2003 - May 2004. Ms. Bagley conducted biological assays with captive elephants at Riddle's Elephant Sanctuary with the assistance of three undergraduate students and at Jacksonville Zoo, Knoxville Zoo, Lion Country Safari, Miami MetroZoo and North Carolina Zoo with the aid of Lauren Stanley. Ms. Bagley studied elephants at AENP from September to December 2003 and was assisted again by undergraduate student Lauren Stanley. This was the first time either of them had traveled outside of the USA. Both students worked closely with the park personnel. Ms. Bagley completed her MSc degree in December 2004 and was employed as a High School science teacher in Georgia. She is employed at Zoo Atlanta. A paper in Animal Behaviour was published in 2006 emanating from her thesis research. Thesis link.

Christen Merte, MSc 2006. Ms. Merte examined the development of behaviors in female African elephants across time using a longitudinal technique in addition to a cross sectional sampling method. She also is compared investigative behavior of male and female elephants of each age class as they entered a water hole. Ms. Merte completed her fieldwork at Addo Elephant National Par in July 2005 and presented a talk at a zoological Congress in Grahamstown, South Africa. Ms. Merte graduated in the spring 2006. Thesis link.

Dhaval Vyas, MSc 2006. Mr. Vyas completed a semester of coursework in the spring 2004 and journeyed to Tanzania in July. He has been conducting research on elephant behavior and bird/mammal biodiversity at a waterhole on the Ndarakwai Ranch, owned and operated by Mr. Peter Jones. This study is permitted through TAWIRI/COSTECH. Mr. Vyas will conclude his study in June 2005 after assisting in the training of new graduate student Erek Napora. Mr. Vyas graduated in the summer 2006. Thesis link.

Jordana Meyer, MSc 2006. Ms. Meyer entered Georgia Southern in the fall semester 2004. She has completed one year of coursework, received fiscal awards from the Academic Excellence and Graduate Student funds at Georgia Southern. She conducted behavioral studies at AENP from May-October 2005, examining the developmental relationship between play and chemical communication. She also performed a study with captive female African elephants in North America where she studied how behavior changes with hormonal status. Ms. Meyer graduated in December 2006 and now collaborates with our team on a related project (examining fecal hormone levels and behavior in African elephants) in AENP. Thesis link.

Erek Napora, MSc 2007. Mr. Napora entered Georgia Southern in the spring semester 2005. He conducted a study on developmental aspects of chemosensory behavior in African elephants and on elephant damage to woody vegetation on Ndarakwai Ranch, Tanzania from May 2005 to July 2006. Erek presented some of his research findings at the 2006 Animal Behavior Society meeting in Snowbird, Utah. Mr. Napora graduated in the spring 2007 and is now living in northeastern Tennessee. Contact. His MS thesis is available online.

High school teacher Juli Sergi, Stacie Castelda, Erek Napora and Bruce Schulte at the Ndarakwai waterhole

Stacie Castelda, Master of Science candidate, 2008. Stacie is from Iowa and graduated from Luther College. She spent 2006-7 in Tanzania observing the responses of African elephants to putative pheromone candidates. In addition, she continued to track elephant damage to the woody vegetation on Ndarakwai Ranch as well as monitoring elephant damage to local farm fields. Stacie plans to graduate in spring 2008.

Russell Blogg, Master of Science candidate, 2008. Russ graduated from Illinois Wesleyan University and was a scholarship swimmer. He is examining the chemosensory development of African elephants in the 10-20 year age class at Addo Elephant National Park in South Africa. He also has collected data on known males before, during and after bouts of musth. Russ plans to present some of his findings at the IEF meeting in Orlando in 2007. He should graduate in spring 2008 and shortly thereafter will be married!

Rhea Esposito, Master of Science candidate, 2008. Rhea is from Boulder, Colorado and graduated from Oberlin College. She has published on Antarctic diatoms. Rhea is examining chemosensory and social behavior at the family level with elephants in AENP. She also will use her computer skills to examine temporal patterns in elephant behavior over the 4+ years of data that we have acquired at AENP. Rhea plans to graduate in spring 2008.

Nabil Nasseri, Master of Science candidate, 2009. Nabil graduated from Texas A&M - Corpus Christi (TAMU-CC) with a B.S. in Biology. He joined our program in 2006. Nabil is studying the responses of African elephants to potential deterrent compounds in comparison to natural elephant chemo-signals. In addition, he is determining the impact of elephant damage on the herpetofauna at Ndarakwai Ranch. Nabil plans to graduate in the spring term 2009.

Rebekah Hoffmeier, Master of Science candidate, 2010. Bekah graduated from Michigan State University with a B.S. in Zoology, Animal Behavior. Bekah has studied olive baboons in Kenya and chacma baboons in Botswana. She also has worked with Belding's ground squirrels and hyenas. Bekah's interests are in conservation behavior to understand better and to reduce human-wildlife conflict. She will be working at our Tanzanian study site on human-elephant conflict issues.

GEORGIA SOUTHERN UNDERGRADUATES INVOLVED IN THE PROJECT:

Amy Gray,Undergraduate, Georgia Southern. BS candidate, class 2004. Amy assisted Helen in South Africa. Amy presented a poster at the annual Society for Integrative and Comparative Biology (SICB) conference. Amy continued her career in graduate school at Georgia Southern University where she completed her MSc under Dr. Ray Chandler.

Matthew Groover, graduate spring 2006 Georgia Southern University,. Matthew assisted Erek in Tanzania and conducted a senior research project during his stay. He presented his research as a 4890 undergraduate research talk and as a poster at the 2006 GSU Research Day.

Lauren Stanley, Undergraduate, Georgia Southern. BS candidate, class 2003. Lauren assisted Katie with the captive and wild elephant studies. She is now living in the greater Atlanta area after graduating from Georgia Southern.

Kimberly Wollett, graduated spring 2006 Georgia Southern University. Kimberly will assisted Jordana in South Africa and conducted a senior research project during her stay. She presented her research as a 4890 undergraduate research talk and as a poster at the 2006 GSU Research Day.

UNDERGRADUATE STUDENTS AT HENDRIX COLLEGE INVOLVED IN PROJECT:

Patrick Brown, Undergraduate, Hendrix College, Arkansas. In March, 2005, Patrick presented a research poster at the Spring National American Chemical Society Meeting in San Diego, CA. 'Use of SPME, Reverse Phase SPE, and GC-MS in a Search for African Elephant Urinary Pheromones'.

Adam Carver, Dustin Freyaldenhoven, Leighton Satterfield (2003-4), Presented posters at National American Chemical Society Meeting in Anaheim, two of these gentlemen are in medical school and one is in chemical engineering.

Joi Chen, Stephen Jackson (2006-7), Presented posters at NACS in Chicago; co-authors on chapter in Chemical Signals in Vertebrates 11; graduate 2008, one headed to PhD program in chemistry, one to medical school.

Whitney Dill, Ben Davis, Adam Hicks (2002-3): Presented posters at the National American Chemical Society (NACS) Meeting in New Orleans; Attending medical school. Co-authors on one journal paper.

Mindy Eggert In March, 2005, Mindy presented a research poster at the Spring National American Chemical Society Meeting in San Diego, CA. 'Comparison of Different SPME Fibers for the GC-MS Analysis of Trace Organics in a Biological Matrix (Urine)' [This was basically an examination of the use of our new macros for mass spectral data analysis from African elephant urine.]

Maria Evola (2004-5), Presented poster at National Conference for Undergraduate Research in VA; co-author on chapter in Chemical Signals in Vertebrates 11, attending veterinary school.

Mimi Heineman (2004) Assisted with bioassays and chemical analyses; she is now in veterinary school.

Sam House (2005-6), Presented poster at NACS Atlanta, CA; co-author on one publication & chapter in Chemical Signals in Vertebrates 11; enrolled in Master's in Forensic Science.

Grant Morshedi (2003-4), Presented poster at NACS Anaheim; CA. Co-author on 1 journal paper as well as chapter in Chemical Signals in Vertebrates 10 & 11; attending medical school.

Daniel Mwanza (2002) Presented poster at NACS in San Diego, CA; co-author on one journal paper & chapter in Chemical Signals in Vertebrates 10.

Margaret Weddell (2005-7), Co-author on posters at NACS Atlanta and Chicago, co-author one one paper & chapter in Chemical Signals in Vertebrates 11, graduated in 2007, enrolled in PhD program in chemistry.

STUDENTS FROM OTHER UNIVERSITIES INVOLVED IN PROJECT:

Maureen Correll, B.S. degree from the College of William and Mary, Spring 2003. Maureen was a biology major, studied abroad at James Cook University in Australia, and conducted research on mate choice behavior and sexual selection in zebra finches with Dr. John Swaddle at W&M. Maureen is assisting Helen Loizi in the field study during the summer 2003.

Michelle Malament, undergraduate at Miami University of Ohio. Michelle is a double major in biology and environmental science. During the summer 2003, Michelle is assisting Katie Bagley to conduct biological assays.

Summary of Study

Polygamous animals often display extreme sexual dimorphism, to the point where males and females may live in very different social structures. In mammals, females typically raise the offspring often with little or no male assistance. African elephants (Loxodonta africana) illustrate this pattern very well. Intersexual interaction is especially prevalent during the breeding season when individuals use a variety of communicative behaviors to compete for and select mates as well as maintain social bonds. Chemical signals and related tactile behaviors are primary means of evaluating reproductive condition and attracting mates. Our study is examining the development of chemical signal production and sex-specific responses related to reproductive and social signals. In addition, we are searching for specific pheromones, that is, chemical signals that mediate inter- and intra-sexual behavior in African elephants.
We are studying female-to-male signals, female-to-female signals and then comparing the two developmental pathways. Answers are sought by: (i) Collecting urine from captive and wild female African elephants; (ii) Performing behavioral bioassays of urine samples for the responses of both male and female conspecifics; (iii) Chemically analyzing bioactive urine fractions to identify organic compounds; (iv) Bioassaying selected urinary components with captive and then wild African elephants; (vi) Observing wild African elephants for reproductive chemotactile communication.
The proposed research will identify proximate chemical causes of reproductive behavior and elucidate sexually dimorphic developmental patterns of communication. This study will lay the groundwork for future investigation on male based signals and of ultimate questions on this topic. The findings may assist in the conservation of endangered elephants in practice and other species in principle.

Conservation Issues

Elephants, along with much of the Earth's biodiversity, are either in immediate peril (e.g., Asian species) or have a problematic outlook of continued human intrusion and disturbance (Barnes 1999). Elephants tend to promote diversity (Western 1989), but the trade-off of more elephants may be greater conflict with humans (Barnes et al. 1995). Yet, Hoare (1999) proposed that human-elephant conflict was less density dependent and more individual animal specific. His 'male behaviour hypothesis' suggests that males are the likely cause of most crop raiding because of the male social structure and in our view, developmental pattern. Crop raiding by females may be related to matriarchal behavior, which is sometimes reflective of aggressive male attributes (Hoare 1999, 2001). In practice, killing problem animals may have little impact on human-elephant conflict (Hoare 2001) and hence, alternative solutions should be sought. In recent years the importance of understanding behavior for conservation has received increasing attention (e.g., see ABS Conservation Committee website (Dr. Schulte serves on this committee).
Conservation relies upon sound scientific data and our study will provide such information on the developmental patterns of reproductive communication in elephants. Because chemical signals are so important to elephants and most mammals, deciphering their identity and message will move the field of mammalian behavior forward and should facilitate conservation efforts. For such direct application to be possible, we must know the signals, the context and the variation in response by age, sex and status. By working with captive and wild African elephants, we can achieve our stated objectives and set the stage for further investigation on signal function and potential applications for conservation.

Study Sites

A major portion of our study requires the collection of urine from captive elephants and the bioassaying of urine and identified compounds at facilities with captive elephants. We are working with numerous facilities, including Riddle's Elephant Sanctuary, Baltimore Zoo, Cameron Park Zoo, Indianapolis Zoo, Jacksonville Zoo, Knoxville Zoo, Lion Country Safari, Louisville Zoo, Miami MetroZoo, Nashville Zoo, North Carolina Zoo, Sedgwick County Zoo, Seneca Park Zoo, Six Flags Marine World, Wildlife Safari Park. For our field studies, we are collaborating with researchers and management personnel from several locations in Africa. Currently, we are located at Addo Elephant National Park in South Africa operated by South African National Parks (SANParks). We have been kindly assisted by Professor Graham Kerley and his students including Dr. Anna Whitehouse and Katie Gough at the Nelson Mandela Metropolitan University (NMMU) in Port Elizabeth and the Terrestrial Ecology Research Unit (TERU). The field aspect of the project expanded to a second permanent field site in 2004. One of our graduate students, Dhaval Vyas, had worked in Tanzania as an undergraduate at Bucknell University. Mr. Vyas put Dr. Schulte in touch with Peter Jones, the owner of Ndarakwai Ranch. Arrangements were made for Mr. Vyas to conduct research at Ndarakwai contingent upon receiving TAWIRI approval. We have been working under TAWIRI and COSTECH approval for the past four years at Ndarakwai Ranch. Graduate students Erek Napora and Stacie Castelda each lived on site for a year; Nabil Nasseri is currently in residence conducting his research.

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