|
|||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
|
|||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
| |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
 |
Fig. 1.
Jane Brockmann with a Florida Scrub Jay (endemic species to Florida) at the Archbold Biological Station in Lake Placid, Florida. |
 My
Background |
I
always thought of myself as predestined to follow a career
in science. I have loved animals and
biology as long as I can
remember and my parents always encouraged me in these
interests. Both my brother and I are university professors
(and both have been chairs
of our departments), he was in the Soc/Anthro Department
at St. Xaviar University in Chicago (now retired). Our parents
were
professionals; my father a chemist
who conducted food research and my mother a dietitian
and instructor in nutrition. When my parents retired
in 1976, they moved to San Miguel
Allende, Mexico where they lived for 17 years. We
enjoyed many years of traveling to Mexico to visit this wonderful
colonial town. When my
mother passed away, my father moved to Gainesville
where he died in 1996. My spouse, Tom Rider, is owner
of Goerings' Book Store in Gainesville;
we live near campus in College Park.
I spent my early years in Louisville and Indianapolis
and moved to Chicago in 1957 where I attended the
University of Chicago Lab School graduating in 1963.
I received my undergraduate degree from the Department of Biology at Tufts
University in Medford, Mass. in 1967. During my college
years I worked in the Laboratory
of Ornithology at Cornell University for the summer after my freshman year and did an undergraduate
research project on the neuroethology of cockroaches during the summer after
my junior year. I have always enjoyed bird watching and outdoor activities,
especially watching the behavior of animals. I started
graduate school in 1968 at the University
of Maryland with Jack P. Hailman on a Woodrow Wilson National Fellowship and
when Jack moved to the Department of Zoology at the University of Wisconsin,
I went with him. I received my M.S. degree in 1972 studying the territorial
behavior of a reef fish, "The function of poster-coloration in
the Beau-Gregory, Eupomacentrus leucostictus (Pisces;
Pomacentridae)." My Ph.D. (1976) was
a study of the nesting behavior of a ground-nesting wasp, "The control
of nesting behavior in the great golden digger wasp, Sphex ichneumoneus (L.)
(Hymenoptera,
Sphecidae)."
I joined the Department of Zoology, University of Florida
in 1976 and have subsequently moved up through the ranks: Assistant Professor
(1976-1981),
Associate Professor (1981-1989), Professor (1989-present) and from 1997-2001
I was Chair of the Department. In 1977-78, with a National Science
Foundation NATO Postdoctoral Fellowship, I joined the Animal Behaviour Research
Group at Oxford University, England and worked with Richard Dawkins. In 1985-86
I held a Visiting Research Biologist position in the Department of Biology,
Princeton University while on a National Science Foundation Visiting
Professorship for Women.
I have
taken two sabbaticals, one in 1984 when I returned to England to continue my
collaboration with Richard Dawkins and Alan Grafen and one in 1994 when I visited
the Department of Evolution and Ecology, University of California, Davis, CA,
to work with Marc Mangel. In 1995 I was honored to receive the Animal Behavior
Society Wm. C. Brown Animal Behavior Teaching Award followed by a University
of Florida TIP (teaching) award in 1996. In 1995 I was elected a Fellow of
the Animal Behavior Society. In 2003 I was awarded the University of Florida
Chapter
of Sigma Xi Senior Research Award and I spent a very interesting two weeks
as a Distinguished Visiting Professor at the University of Miami, Coral Gables,
Florida. In 2003-4 I lived in Arlington, VA for one year while I was the Program
Director for the Animal Behavior Program at the National Science Foundation.
 |
Fig.
2. Horseshoe crabs nesting. The female is in the middle with
an attached male behind and 4 satellite males. |
Since 1989 I have been studying the behavior, natural history and population biology of the horseshoe crab, Limulus polyphemus, at Seahorse Key, Florida (supported in part by the National Science Foundation, the Florida Foundation and the Division of Sponsored Research). In spring, female-male pairs of horseshoe crabs come to the beach at Seahorse Key to mate and lay their eggs. This island is part of the Cedar Keys National Wildlife Refuge and has proven to be a good location for studying this species. I censused permanently tagged animals for nine years as they returned to this site. No one has ever studied a marked population of this species, so it was important that I obtain basic information on the frequency of breeding, population recruitment, and patterns of life history changes as well as changes in the behavior and physical condition of individuals over the years. During several field seasons I have also gone to the Delaware Bay to study horseshoe crabs where they occur in particularly large numbers.
Horseshoe crabs nest in a narrow strip of sand in the high intertidal. My graduate student, Dustin Penn, and I examined three factors that might influence nest-site selection: predation, erosion and egg development. Batches of eggs were artificially reburied above and below the normal nesting range on the beach. We found no evidence to support the hypothesis that eggs buried in the lower beach were more likely to be consumed by predators or that they were differentially washed away by erosion and wave action. However, eggs in the lower beach were less likely to develop due to reduced oxygen and eggs reburied higher than normal were more likely to desiccate. These results mean that females are choosing nest sites that maximize egg development.
Dustin and I also discovered that males show two patterns of mating behavior: some males attach to females and other males come ashore and crowd around the nesting couples. A paternity analysis conducted in collaboration with Wayne Potts demonstrated that these satellite males fertilize a large proportion of the eggs that the female lays which means that they are engaging in sperm competition with the attached male.
Large groups of satellites form around some ("popular") couples while ignoring other ("unpopular") couples. These groups are not random: large groups occur significantly more often than expected if satellites were aggregating around couples at random. In experimental field manipulations, "popular" couples were more likely to re-attract satellites after they had been removed than "unpopular" couples. This means that differences in group size were not due to differences in the time the pair had been on the beach. Rather, differences must exist between couples in their attractiveness to satellites.
 |
Fig.
3. Three groups of horseshoe crabs. The left-hand
group is made up of one pair plus one satellite male (to
their right);
the right-hand group is only a nesting pair;
The central group contains a nesting pair and five satellite
males. You would
think that it would make more sense for some
of the males in the large group to join the couple with
no satellites, but this is not what happens. |
Measurements revealed that females, but not attached
males, of "popular" couples were larger and in better condition,
on average, than females of "unpopular" couples. Two alternative
hypotheses are that differences exist in the satellites'
ability to fertilize eggs when associating with
certain pairs and the other is that differences
exist in the quality of female's eggs or egg location.
The second hypothesis was the subject of my graduate
student Cynthia Hassler's M.S. research (1999).
She showed that there were no differences in the
success or development of eggs laid by "popular" and "unpopular" females
but her data show that the "popular" females lay more
eggs.
Nothing is known so far about how individual males
make decisions that result in the observed grouping patterns. The work
that had been done by Robert Barlow and colleagues prior to our study
showed that Limulus use visual cues.
Using cement models of nesting pairs, Rachel Schwab
(M.S. 2006) demonstrated that males use visual cues and
can detect small differences in female size, being more
attracted to larger than to smaller females. Cynthia and I conducted
an experiment that suggested chemical cues may be involved, the first
demonstration for this species. She showed that satellite males are attracted
to specific females and to chemical cues that are produced by those females.
Rachel has subsequently shown that chemicals from eggs
cause males to remain longer with a pair, but these are
not the chemicals that attract males from a distance. An
undergraduate in the lab, Lindsay Keegan is currently
following up on this problem.
Do attached and unattached males differ? A set of measurements
quickly demonstrated that attached and unattached males do not differ
in size but they do differ in condition and probably age. I conducted
experimental manipulations in the field to characterize behavioral differences
between attached and unattached males. I found that attached males that
had been detached from females were more likely to pair, they were more
likely to remain attached and if separated from a female they reattached
more quickly than unattached or satellite males. This suggests that males
that differ in mating tactics and condition and age also differ in behavior.
One major goal of the research has been to understand the adaptiveness
of these condition-dependent tactics.
Most male horseshoe crabs consistently remain attached
or unattached. The maintenance of two patterns of behavior in one population,
like attached and unattached, is an evolutionary puzzle. We know there
are costs and benefits associated with each tactic but it is unlikely
that their average success would be exactly equal. Surely, one would
have higher average fitness and the less successful would be selected
from the population. Yet, these two quite different patterns have been
maintained for generations and, furthermore, consistent differences exist
between populations in the frequency of satellites. How can such variability
be explained?
I
recently received funding from the National Science
Foundation to continue my research on horseshoe
crabs. Together with my Post-doctoral Fellow,
Sheri Johnson, we will be studying multiple mating from
the female perspective: why do some females attract
many males and some none? It is a complex
problem with many alternative hypotheses but the
external fertilization shown by this species will
allow us to manipulate the system more easily than
with other arthropods (all of which have internal
fertilization).
The study of alternative strategies in horseshoe crabs
provides new insight into the evolution of decision-making processes
in animals and how flexible, condition-dependent phenotypes evolve and
are maintained in populations. Limulus provides an excellent opportunity
for such a study because of the ease with which adults can be observed
and manipulated in the field. Their
ancient taxonomic origin and independent evolutionary history
make them particularly valuable comparative material. The vast
amount that is known about some aspects of their reproductive
physiology, immunology and development make them valuable subjects
for integrative research.
For further information on horseshoe crabs see:
Department of Natural Resources, Maryland: http://www.dnr.state.md.us/education/horseshoecrab/
University of Delaware, College of Marine Studies:
http://www.ocean.udel.edu/horseshoecrab/
New Jersey Audubon Society:
http://www.njaudubon.org/Conservation/HScrabalert.html
Marine Biology Lab, Woods Hole, MA:
http://www.mbl.edu/animals/Limulus/
Florida Marine Research Institute:
http://www.floridamarine.org/horseshoe_crab/
ERDG (an excellent general site):
http://www.horseshoecrab.org/
1983. Cohen, J. A. and H. J. Brockmann. Breeding activity and mate selection in the horseshoe crab, Limulus polyphemus. Bulletin Marine Science, 33: 274-281.
1990. Brockmann, H. J. Mating behavior of horseshoe crabs, Limulus polyphemus. Behaviour 114:206-220.
1992. Brockmann, H.J. and D. Penn. Male mating tactics in horseshoe crabs, Limulus polyphemus. Animal Behaviour 44:653-665.
1994. Brockmann, H.J., T. Coleson and W. Potts. Sperm competition in horseshoe crabs (Limulus polyphemus). Behavioral Ecology and Sociobiology 35: 153-160.
1994. Penn, D. and H. J. Brockmann. Nest-site selection in the horseshoe crab, Limulus polyphemus. Biological Bulletin 187: 373-384.
1995. Penn, D. and H. J. Brockmann. Age-biased stranding and righting in horseshoe crabs (Limulus polyphemus). Animal Behaviour 49: 1531-1539.
1996. Brockmann, H. J. Satellite male groups in horseshoe crabs. Limulus polyphemus. Ethology 102: 1-21.
2000. Brockmann, H.J., C. Nguyen and W. Potts. Paternity in horseshoe crabs when spawning in multiple male groups. Animal Behaviour 60:837-849.
2001. Hassler, C. and H.J. Brockmann. Evidence for use of chemical cues by male horseshoe crabs when locating nesting females (Limulus polyphemus). Journal of Chemical Ecology. 27: 2319-2335.
2002. Brockmann, H.J. An experimental approach to altering mating tactics in male horseshoe crabs (Limulus polyphemus). Behavioral Ecology 13:232-238.
2003. Brockmann, H.J. Nesting behavior, a shoreline phenomenon. IN The American Horseshoe Crab. Ed. By C. Shuster, R. Barlow and J. Brockmann. Harvard University Press. pp. 33-49.
2003. Brockmann, H.J. Mating behavior of male horseshoe crabs. IN The American Horseshoe Crab. Ed. By C. Shuster, R. Barlow and J. Brockmann. Harvard University Press. pp. 50-82.
2003. Shuster, C.S., R. B. Barlow and H. J. Brockmann. (eds.) The American Horseshoe Crab. Harvard University Press. 427 pp.
2005. King, T.L., M.S. Eackles, A.P. Spidle and H.J. Brockmann. Regional differentiation and sex-biased dispersal among populations of horseshoe crabs (Limulus polyphemus). Transactions of the American Fisheries Society 134:441-465.
2006. Duffy, E.E., D. Penn, M.L. Botton, H. J. Brockmannn and R.E. Loveland. Eye and clasper damage influence male mating tactics in the horseshoe crab, Limulus polyphemus. Journal of Ethology 24: 67-74.
Schwab, R., H.J. Brockmann. 2007. The role of visual and chemical cues in the mating decisions of satellite male horseshoe crabs (Limulus polyphemus). Animal Behaviour (in press).
Brockmann, H.J. and M.D. Smith. Reproductive competition and sexual selection in horseshoe crabs. IN Biology and Conservation of Horseshoe Crabs, Ed. by John Tanacredi, Mark Botton, and David Smith, Springer. (in press)
The Evolution of Alternative Tactics
Alternative mating and nesting tactics and sex ratios
are part of a much broader problem in evolution, the origin and maintenance
of discrete, alternative phenotypes. I am interested in understanding
the evolutionary factors that favor the appearance of discrete alternative
behavior patterns in populations (in one sex at one time), the factors
that influence the frequencies of the alternatives, the factors that
influence the degree of flexibility and the underlying mechanisms controlling
the alternative phenotypes.
2001. Brockmann, H.J. The evolution of alternative
strategies and tactics. Advances in the Study of Behavior. 30:1-51.
Oliveira, R., M. Taborsky and H.J.Brockmann (eds.)
Alternative Reproductive Tactics – An Integrative
Approach. Cambridge
University Press (in press) Chapters on
alternative reproductive tactics in insects; alternative
reproductive tactics and life histories.
| Past
Research |
 |
Fig.
4. A male pipe-organ mud-daubing wasp guards the entrance to a nest that a female is provisioning. He is chasing away an intruding male. |
Mating and Nesting Tactics of Wasps
Prior to studying horseshoe crabs, I spent 10 years observing the mating and nesting behavior of a solitary, mud-daubing wasp, Trypoxylon politum. This was a detailed field study on the behavior of males who guard the nests that females build, a very unusual pattern for male wasps. Females have at least five different ways of nesting, including nest-sharing and brood parasitism, and I was interested in understanding how such diversity is maintained in the population. The wasps also have a strongly male-biased sex ratio in the spring generation and a 50:50 sex ratio in the summer generation. I studied the maintenance of this unusual pattern, which seems to be related to their life-history (partial bivoltinism) and this resulted in my studying their highly variable patterns of emergence and diapause. My study of mud-daubing wasps related more broadly to issues about the evolution of alternative mating and nesting tactics, to evolutionary conflicts of interest between males and females and to the evolution of social behavior.
1984 Brockmann, H. J. The evolution of insect sociality. In J. Krebs and N. Davies (eds.) Behavioral Ecology. Sinauer Associates, Second Edition, pp. 340-361.
1987 Seger, J. and H. J. Brockmann. What is bet-hedging? In P. Harvey and L. Partridge (eds.) Oxford Surveys in Evolutionary Biology. Oxford Univ. Press, 4:182-211.
1988 Brockmann, H. J. Father of the brood (Conflict and cooperation in a mud-daubing wasp). Natural History Magazine. 97(7):32-37.
1989 Brockmann, H. J. and A. Grafen. Mate conflict and male behavior in a solitary wasp, Trypoxylon politum. Animal Behaviour 37:232-255.
1992 Brockmann, H. J. and A. Grafen. Sex ratios and life-history patterns of a solitary wasp. Behavioral Ecology and Sociobiology 30:7-27.
1992 Brockmann, H. J. Male behavior, courtship and nesting of the solitary wasp, Trypoxylon monteverdeae. Journal of the Kansas Entomological Society 65(1):66-84.
1993 Brockmann, H. J. Parasitizing conspecifics: comparison between Hymenoptera and birds. Trends in Ecology and Evolution. 8:2-3.
1997 Brockmann, H. J. Cooperative Breeding in Wasps and Vertebrates: the Role of Ecological Constraints. IN Evolution of Social Behaviour in Insects and Arachnids, J. Choe and B. Crespi (eds.). Cornell University Press. pp. 348-371.
2004 Brockmann, H.J. Variable Life-History Patterns in the Pipe-Organ Mud-daubing Wasp, Trypoxylon politum (Hymenoptera: Sphecidae). Journal of the Kansas Entomological Society (invited memorial volume for Howard E. Evans) 77: 503-527.
 |
Fig.
5. Female Diaprepes root weevil.
|
Sexual Selection in a Beetle.
The root weevil, Diaprepes abbreviatus, is a wonderful species to study; it is large, easy to observe, adapts well to the laboratory and it mates all the time. The animals are easy to observe in the field where they sometimes occur in large aggregations. Since this species is an important agricultural pest of citrus in Florida, a considerable amount is known about various aspects of its life history and ecology, but little is known about behavior. An Israeli colleague working at the USDA Lab, Ally Harrari, and I conducted a study on the diverse mating behavior of this species. We examined why females mate multiply, sperm competition and mounting by females of other females and males of other males. Laura Sirot (Ph.D. 2004) continued this research and took the study considerably further. Laura examined one of the most vexing problems in sexual selection, the role of cryptic female choice. Cryptic choice refers to choice-like processes that occur after mating that result in non-random fertilization. This process is suspected in a number of species but the evidence is slim because it is so difficult to separate the role of female choice for male traits from differences in the effectiveness of different males in fertilizing eggs (e.g. through sperm competition). She has also examined the role of copulatory courtship (courtship during their long matings such as tapping the female with the antennae) in influencing fertilization success.
1999. Harari, A. R. and H. J. Brockmann. Male beetles attracted by females mounting. Nature 401:763.
2000. Harrari, A. and H.J. Brockmann. Intrasexual mounting in the beetle Diaprepes abbreviatus. Proceeding of the Royal Society of London B 267: 2071-2079.
2003. Harari A.R., P.J. Landolt , C.W. O’Brien, H.J. Brockmann. Prolonged guarding behavior and sperm competition in the weevil Diaprepes abbreviatus (L.) (Coleoptera: Curculionidae). Behavioral Ecology 14: 89-96.
Sirot L.K., H.J. Brockmann and S.L. Lapointe. Male post-copulatory reproductive success in the beetle, Diaprepes abbreviatus. Animal Behaviour (in press).
(a)  |
(b)  |
Fig. 6. (a) Mating damselflies showing the female color polymorphism, the male mimic
or andromorph and (b) the gynomorph. |
|
Alternative Female Tactics in a Damselfly
Laura Sirot (M.S. 1999) studied a female-limited color polymorphism in a damselfly, another case of alternative strategies, but this one is controlled by a genetic polymorphism. Female damselflies of Ischnura ramburi come in two color forms, one that looks different from the males (gynomorph) and one that mimics male color and pattern (andromorph). The hypothesis to explain male mimicry was that females can avoid the very long and costly mating characteristic of this species by looking like males. In an extremely ambitious set of laboratory experiments, Laura demonstrated that mating is expensive for females, particularly for andromorphic females, when they are in a caged situation where males recognize them as females, i.e. when their mimicry is broken. Intensive field observations, experiments and censuses in a number of habitats demonstrated that andromorphic and gynomorphic females mate the same number of times but they behave differently when they encounter males. We think that the two color morphs are equally successful but use quite different approaches to maximizing success.
2001. Sirot, L. and H.J. Brockmann. Costs of sexual interactions to females in Rambur’s forktail damselfly, Ischnura ramburi (Zygoptera: Coenagrionidae). Animal Behaviour. 61:415-424.
2003. Sirot, LK and HJ Brockmann, C. Marinis and G. Muschett. Maintenance of a female-limited polymorphism in Ischnura ramburi (Zygoptera: Coenagrionidae): evaluation of hypotheses and new insights. Animal Behaviour 66: 763-775.
|
|
|
Fig. 7. Gryllus rubens has two forms. (a) Long-winged morph and (b) short-winged. The pictures are from the Singing Insects of North America http://buzz.ifas.ufl.edu/crickets.htm |
|
Sexual Selection in Crickets
On a warm summer evening, you can often hear crickets singing, which they do to attract females. But in the field cricket, Gryllus rubens, all this singing also attracts a fly that lays her larvae (her eggs hatch inside her body) on the singing male. These larvae burrow in and kill the male within seven days. In Florida where Manuel Vélez (Ph.D. 2004) conducted this study, flies are around only during the fall but the crickets sing all year. This means that spring males can sing whenever and as much as they want but autumn males are constantly faced with death whenever they sing for females. We found that fewer autumn males sing (they search for females without singing) and they are less likely to sing at dusk when flies are most active, but if an autumn male does sing, he sings more than spring males. This is because singing is more effective at attracting females in the autumn than in the spring.
Females that approach conspicuous males may also be vulnerable to predation. This means that selection for reduced male conspicuousness in the presence of these parasites may also be due to selection on females to alter their preferences in the presence of parasites. Since female crickets are also parasitized by these flies as a result of associating with calling males, we predicted that autumn females should be less attracted to male song than spring females. We tested female response to male calls in a rectangular arena in which male calling song was broadcast from a speaker. Spring females readily approached the speaker but autumn females were less likely to approach and remain in the vicinity of the speaker. These results emphasize the importance of considering how risk affects the evolution of conspicuous male behavior both directly through its effect on the male and indirectly through its effect on female responses to males.
An undergraduate in our lab, Kathleen Colby, is continuing to study seasonal variation in Gryllus rubens, focusing on male behavior and in particular on differences in accessory glands for long- and short-winged males. In females there is a known trade-off between wing length and dispersal behavior: long-winged females delay maturation of their ovaries whereas short-winged females mature immediately and are more fecund. Little is known about the differences between short- and long-winged males and whether they also differ in life-history and reproductive success.
2006. Vélez, M. and H.J. Brockmann. Seasonal variation in selection on male calling song in the field cricket, Gryllus rubens. Animal Behaviour 72:439-448.
2006. Vélez, M. and H.J. Brockmann. Seasonal variation in female phonotaxis to male calling song of the field cricket Gryllus rubens. Ethology 112:1041-1049.
My
Teaching
Undergraduate Course. Nearly all students in my undergraduate courses are planning professional careers in medical, agricultural or biological fields. My goal is to make certain that these students have the concepts and skills they need to pursue these professional goals. (I do not assume that students will be attending graduate school, but I do assume they will pursue some professional goal and so will need particular skills.)
Animal Behavior (ZOO 3513) is a wonderful course to teach because everyone finds the topic so intrinsically interesting: how and why animals behave as they do. But to understand behavior one must integrate ideas and information from a great diversity of fields (e.g. evolution, ecology, physiology, genetics, physics, etc.) indeed, all of the biological sciences, so it is a particularly valuable course for students to take late in their career. The great diversity of topics gives students the opportunity to see the larger structure of biology, to understand the major questions and to integrate the parts with the whole. They learn how to develop hypotheses, evaluate evidence, and draw conclusions. They learn the importance of variation, change and diversity; they learn to think across scales and they learn a new perspective on time. They are taught to use the library, to read original literature, to gather information from the web and other sources. I pick out the most important issues in biology and illustrate and discuss these through behavior: the nature vs. nurture problem; the extent to which genes control behavior; how variation is maintained; the evolution of sex; and the different sensory worlds of non-human animals, to name just a few. The laboratory part of the course is designed to help students understand the scientific process and to develop specific skills including description, data analysis, basic statistics, locating and evaluating literature and writing. The course is fun and interesting but it is demanding with weekly laboratory assignments and considerable reading for the lecture. It is taught every spring semester.
Graduate Courses. Graduate students from Zoology, Wildlife, Fisheries, Anthropology, Vet Med, Animal Science, Psychology, and Entomology are the most frequent participants in Ethology (ZOO 6515) and my Animal Behavior Seminar (ZOO6939) courses. Most of these students are doing (or plan to do) animal behavior research. In the early days of teaching graduate courses, I was dismayed to find that many of my students would talk about behavior without being able to provide even elementary definitions of basic concepts. For this reason I now teach two courses side-by-side: basic level and graduate level animal behavior. In this way I hope my students develop the skills and concepts they need to conduct behavioral research at the graduate level. I encourage them to question assumptions and dogma; to think deeply about what they read; to develop quantitative models; and to evaluate problems using a rigorous, hypothesis-testing approach. They learn to design experiments, analyze data, review grant proposals, critique manuscripts and write grant proposals and scientific essays and reports. The students in my graduate courses learn to integrate ideas with what is going on in other fields, to apply techniques and concepts to novel problems, and to think about behavior in new ways or from new perspectives. My courses are always interesting but a lot of work (for students and faculty alike!).
Graduate Mentoring. I take my graduate mentoring extremely seriously. Although my students study a great diversity of problems (see list) in behavior, I work closely with each in every stage of their research from thinking of the question to planning the experiments and methods, to analyzing and interpreting the data, to writing and presenting the results. I try to professionalize my students in every way possible, for example, by encouraging them to attend meetings of our national society, the Animal Behavior Society, and participating in ABS committees and workshops. During most semesters I hold a journal club each week for my students and others who wish to attend. We discuss papers, research, and issues of concern such as ethics and teaching. I always encourage my students to teach my undergraduate Animal Behavior course with me so that they will learn how to present and teach this material (and learn some of the tricks). When possible I also encourage my students to spend time in the field with me, or I go into the field with them, so that we can share approaches and methods. My students have been very successful at attracting support for their research and at finding good positions, primarily in academic settings. Graduate mentoring fosters an intense and long-term relationship with my students which I value immensely. I was enormously pleased to receive the Doctoral Dissertation/ Mentoring Award from the Graduate School in 2005.
|
|
I have always enjoyed participating in all aspects of my profession, from my national and international organizations, to the university, to my department. Every service responsibility results in my learning a great deal about how the profession functions and how to handle an entirely new set of issues and problems. It also results in a new set of friends and associates with whom my students and I enjoy interacting. I have served as an Associate Editor of Evolution (1987-90), on the Editorial Board of Animal Behaviour (1988-1991) and Editor of Ethology (1991-1997). From 2002-2005 I was one of the editors for Advances in the Study of Behavior and I am now the editor-in-chief. I have served in several capacities for the Animal Behavior Society, including Treasurer (elected 2 terms) (1982-1988), Second President-Elect (elected) 1989-1990, First President-Elect (1990-1991), President (1991-1992), and Past-President (1992-1993). I was the elected US representative to the International Ethological Conference Council (1984-1991), served as its Vice-Secretary General (1995-1999) and then as the IEC Secretary-General (1999-2003. I was elected a Councilor of the International Society of Behavioral Ecology (1998-2002) and Secretary/Treasurer of the American Institute of Biological Sciences (1998-2004). I have served on many grant review panels, evaluation panels, program reviews, and advisory committees for the National Science Foundation, the National Research Council, the Smithsonian Tropical Research Institute, the Swedish National Research Council as well as serving as an external reviewer of university departments. I have also served on peer review panels for the Atlantic States Marine Fisheries Commission and I enjoy working with the teachers and State natural resource professionals in their program Green Eggs and Sand, a Tri-State Horseshoe Crab/Shorebird Education Program. http://www.dnr.state.md.us/education/are/ges.html. In 2003-4 I was the Animal Behavior Program Director at the National Science Foundation http://www.nsf.gov/bio/ibn/. In 2000 I was honored to receive the Animal Behavior Society Distinguished Service Award.
| Graduate
Students |
 |
Fig.
8. My Students and Collaborators. From left
to right: Wayne Potts, Sharoni Shafir, Bonnie
Ploger, Rich Buchholz, Dustin Penn, Jeff
Lucas, Jane Brockmann
|
Their Research and Present Positions
Jeff Lucas (PhD 1983). Dissertation: Feeding behavior and life history strategies of antlions. NATO post-doctoral fellow, Oxford, England, 1983-1984. Assistant Professor, College of William and Mary, 1984-1986. Univ. of California at Redlands, Assistant Professor, 1986-1987. Assistant Professor, Department of Biology, Purdue University, West Lafayette, IN, 1987-1994 Associate Professor, 1994-2001. Professor 2001-present.
E-mail: jlucas@purdue.edu
Web Site: http://bilbo.bio.purdue.edu/www-ecology/faculty/lucas/index.html
Glenn Goodfriend (PhD 1983). Dissertation: Feeding behavior and the evolution of clinal variation in Jamaican land snails. Weizmann Institution of Science Post-doctoral Fellow, Rehovot, Israel, 1983-1994. Research Scientist, Smithsonian Institution, Washington, D.C. 1994-1995. Research Scientist, Geophysical Lab. Carnegie Institution of Washington, Washington, D.C. 1995-2002 (deceased).
Steven Frank (MS 1983). Thesis: Sex ratios and reproductive behavior of Florida fig wasps. Univ. of Michigan, Ph.D. 1987. Assistant Professor, Department of Biology, University of California-Irvine, 1987-1993; Associate Professor 1994-1998; Professor 1998-present.
E-mail: safrank@uci.edu
Web Site: http://stevefrank.org/
Martin Obin (MS 1983). Thesis: Territorial and marking behavior of a brood parasitic mud-daubing wasp. University of Florida, Ph.D. 1990. Assistant Professor, Gerald J. and Dorothy R. Friedman School of Nutrition Science and Policy, Tufts University Medical School, 1996-present.
E-mail: martin.obin@tufts.edu
Web Site: http://hnrcwww.hnrc.tufts.edu/scientists/people/mobin.php
Linda Fink (MS 1984, PhD 1989). Thesis: Maternal investment in the green lynx spider. Dissertation: Color polymorphism in sphingid caterpillars. Assistant Professor, Middlebury College. 1989-90. Assistant Professor, Department of Biology, Sweet Briar College, Virginia, 1990-1995; Associate Professor 1995-2001; Professor 2001-present. Chairman 2000-present.
E-mail: lfink@sbc.edu
Web site: http://www.faculty.sbc.edu/lfink/
Giselle Mora Mora (MS 1987, PhD 1991). Thesis: Mating and nesting behavior of a tropical harvestman with paternal care. Dissertation: Site-based mating system in a tropical harvestman. Assistant Professor, University of Costa Rica, San Jose, 1992-1998; Associate Professor 1998-2000. Teaching in Washington, D.C. 2000-present.
E-mail: gisellemora1@prodigy.net
Karen Masters (MS 1989). Thesis: The adaptive significance of female-biased sex ratios in the neotropical treehopper Umbonia ataliba. Princeton University, Ph.D. 2000. Conducting research for Centro Científico Tropical (CCT), Monteverde, Costa Rica on the interactions between plants and insects.
E-mail: amasters@sol.racsa.co.cr
Sharoni Shafir (MS 1991). Thesis: Honeybees foraging on artificial flowers - intransitivity of preferences and energetics. Stanford University, Ph.D. 1994. Post-doctoral Fellowship, University of Ohio, Columbus, OH. 1995-1998; Lecturer 1998-present (tenure-track), Department of Entomology, Hebrew University of Jerusalem,. Director, Triwaks Bee Research Center.
E-mail: shafir@agri.huji.ac.il
Web Site: http://departments.agri.huji.ac.il/entomology/staff_pages/shafir.html
Richard Buchholz (MS 1989; PhD 1994). Thesis: Singing behavior and ornamentation in the yellow-knobbed curassow (Crax daubentoni). Dissertation: Adaptive Functions of Fleshy Ornamentation in Wild Turkeys and Related Birds. Visiting Assistant Professor, University of Florida 1994. Assistant Professor (tenure-track) University of Northeastern Louisiana, Monroe, LA 1995-1999; Assistant Professor (tenure-track), Department of Biology, University of Mississippi 1999-2004; Associate professor 2004-present.
E-mail: byrb@olemiss.edu
Web Site: http://home.olemiss.edu/~byrb/
Bonnie Ploger (PhD 1992) Dissertation: Proximate and ultimate causes of brood reduction in brown pelicans (Pelecanus occidentalis). 1993-94, Visiting Assistant Professor, Grinnel College, Grinnel, Iowa; N.A.T.O post-doctoral fellowship, Queen's University Kingston, Ontario 1994-95. Assistant Professor, Hamline College, St. Paul, MN 1995-2001; Associate Professor 2001-present.
E-mail: bjploger@piper.hamline.edu
Web Site: http://138.192.68.68/bio/faculty/ploger/ploger.html
Dustin Penn (MS 1992) Thesis: Nesting behavior of horseshoe crabs. 1990-1997, University of Utah, Ph.D. 1997; Post-doctoral Fellowship, Univ. Utah 1997-2001. Director, Konrad Institute for Comparative Ethology, Vienna, Austria.
E-mail: d.penn@klivv.oeaw.ac.at
Web Site: http://www.oeaw.ac.at/klivv/
Laurie Eberhardt (PhD 1994). Dissertation: Sap Feeding and its Consequences for Reproductive Success and Communication in Yellow-bellied Sapsuckers. Assistant Professor (tenure-track), Valparaiso University, Valparaiso, IN 1994-2000; Associate Professor 2000-present.
E-mail: Laurie.Eberhardt@valpo.edu
Web Site: http://www.valpo.edu/biology/FACULTY.HTML
Ron Clouse (MS 1994). Thesis: Nesting Decisions of the Social Paper Wasp Myschocyttarus mexicanus. Agriculture Department Chair, PATS, Pohnpei, FM. 1994-1995. Archbold Biological Station, Res. Sci. 1996. Res. Sci., Museum of the Everglades 1996-97. Economic analyst, NYC (1998-2004). Ph.D. student, Harvard University (2004-present).
E-mail: ron@clouseart.com
Web site: http://www.clouseart.com/index.html
Jose-Luis Osorno (PhD 1996) Dissertation: Male mate desertion in the magnificent frigate-bird. Research Associate, Instituto de Ecologia, Universidad National Autonoma Mexico 1996-2004. Deceased 2004.
Cynthia Hassler (Macartney-Filgate) (MS 1999) Thesis: Satellite Male Groups in Horseshoe Crabs (Limulus polyphemus): How and why are males choosing females? Working in a computer firm in Austin, TX.
Susan Chien (MS) 1994-2000. Paternity in the Pipe-organ Mud-daubing wasp, Trypoxylon politum. Research Assistant, Brown University.
Kavita Isvaran (Ph.D. 2003) Dissertation: The evolution of lekking: insights from a species with a flexible mating system (Indian blackbuck antelope). Gardiner Post-doctoral Fellowship from Cambridge University, UK. (Post-doc with T. Clutton-Brock, Department of Zoology) 2003-2006. Assistant Professor, Centre for Ecological Sciences, Indian Institute of Science,
E-mail: kavita[at]ces.iisc.ernet.in
Web Site: http://ces.iisc.ernet.in/kavita/
Suhel Quader (Ph.D. 2003). Dissertation:
Nesting and mating decisions and their consequences in the baya weaverbird Ploceus
philippinus. Marie Curie E.U. Post-doctoral Fellowship., Cambridge, UK (Post-doc
research in Department of Zoology, Cambridge University with N. Davies) 2003-2005.
Research Scientist, British Society for the
Protection of Birds 2005-2006; Scientist, National
Centre for Biological Sciences, Bangalore, India
(2006-present).
E-mail: suhelq at ncbs.res.in
Web Site:
http://www.ncbs.res.in/~suhel/index.html
Manuel
Vélez (M.S. 1999; PhD 2004) Thesis:
Parental care strategies in the Central American cichlid Aequidens coeruleopunctatus:
tradeoffs between present and future reproduction. Dissertation: Natural and
sexual selection on call variation in the field cricket, Gryllus rubens.
Boston University School of Law (2003-2006). Law Clerk in Puerto Rico.
E-mail: velezmanuel@hotmail.com
Laura
Sirot (MS
1999, Ph. D. 2004). Thesis: Intersexual conflict and mating
avoidance in the damselfly, Ischnura ramburi. Dissertation: Sperm
competition and cryptic female choice in
a weevil, Diaprepes abbreviatus. NIH
NRSA Post-doctoral Fellowship, Department of
Molecular Biology and Genetics, Cornell University,
Ithaca, NY working with Mariana Wolfner
(2004-present).
E-mail: Ls286
at cornell.edu
Web Site:
http://www.mbg.cornell.edu/cals/mbg/research/wolfner-lab/sirot.cfm
Charles
W. Gunnels (Ph.D. 2006) 1999-2006.
Dissertation: Social and Reproductive Altruism in a
Facultatively Social Paper Wasp, Mischocyttarus
mexicanus. (Hymenoptera:
Vespidae). Teaching post-doc., University of Florida (2006-2007). Assistant
Professor, Department of Biological Sciences,
Florida Gulf Coast University, Ft. Myers, FL
(2007-present).
E-mail: cgunnels@fgcu.edu
Web Site:
http://www.zoo.ufl.edu/bgunnels/
Dimitri Blondel (M.S. 2006) Thesis: The Social Organization of the
Singing Mouse, Scotinomys xerampelinus (Rodentia,
Muridae). Ph.d. student with Steve Phelps, UF.
E-mail: dblondel@zoology.ufl.edu
Web Site: http://www.zoo.ufl.edu/phelps/people/dimitri.html
Rachel Schwab (M.S. 2006)
Thesis:
Mating group formation in the horseshoe crab (Limulus
polyphemus): are satellite males ideal-free
distributed among females of variable fecundity and
how do they assess fecundity? Naturalist, Amelia
Island Plantation (2006-2007) M.S. in teaching, UF
College of Education (2007-present)
E-mail: rlschwab13@yahoo.com
 Fig. 9. Recent graduate students.
Back row: Manuel
Vélez, Suhel Quader, Kavita Isvaran;
Front row: Laura Sirot, Jane Brockmann and Billy Gunnels |
Present Graduate Students
Matthew D. Smith (Ph.D.)
2005-present. Phenotypic variation in horseshoe
crabs.
E-mail: mdsmith@zoology.ufl.edu
Web Site:
http://www.zoo.ufl.edu/madsmith/index/Welcome.html
Hope Klug (Ph.D.) 2001-present. Filial
cannibalism in gobies and flagfish.
E-mail: hklug@zoology.ufl.edu
Web Site:
http://www.zoo.ufl.edu/user%2Dweb/grad/hklug/
Clare Rittschof (Ph.D). 2006-present.
Alternative tactics in the golden orb-weaving spider,
Nephila clavipes
E-mail: critts@zoology.ufl.edu
Daniel Sasson (Ph.D.) 2006-present. Sperm competition in horseshoe crabs E-mail: dsasson@zoology.ufl.edu
Present Post-doc
Sheri
Johnson
(Ph.D. 2007, University of Maine)
2007-present. Sheri studies reproduction
and fertilization in marine invertebrates; in
particular, the factors that result in sperm
limitation and sperm competition in marine free-spawners.
Many models predict that gametes quickly become
diluted in the water column, and as a consequence,
many eggs remain unfertilized (i.e. experience sperm
limitation). However, fertilization levels have been
shown to be very high in brooding invertebrates, which
may use feeding structures to filter dilute sperm out
of the water column, and hence experience high levels
of fertilization and sperm competition (i.e. bryozoans
and ascidians). At
UF she will be working on fertilization in horseshoe
crabs.
E-mail: slj@zoology.ufl.edu
Web Site:
http://www.marine.maine.edu/geninfo/geninfo_indiv.php?type=graduate&staff_id=126
Research Collaborators:
Their Current Research and Present Positions
Alan Grafen. Present Position: Lecturer, Animal Behavior Research Group, Oxford
0X13PS Present Research: Evolutionary theory including game theory, and formal
links between game theory and population genetics.
E-mail: alan.grafen@sjc.ox.ac.uk or alan.grafen@zoo.ox.ac.uk
Web Site: http://users.ox.ac.uk/~grafen/
or http://www.sjc.ox.ac.uk/
Ally R. Harari. Present position: Lecturer, Department of Entomology, The Volcani
Center, P.O. Box 6, 50250 Bet-Dagan, Israel.
Present research: development of control management based on phonological and
behavioral approaches in moths and beetles.
E-mail: ally@int.gov.il. or aharari@volcani.agri.gov.il
Web site: http://www.agri.gov.il/Volcani.html
Wayne Potts. Present position: Professor, Department of Biology, University
of Utah, Salt Lake City, Utah.
Present Research: the genetics of natural and sexual selection. Current research
efforts focus on the genes of the major histocompatibility complex (MHC).
E-mail: potts@biology.utah.edu.
Web Site: http://stormy.biology.utah.edu/
Jon Seger. Present position: Professor, Department of Biology, University of
Utah, Salt Lake City, Utah.
Present Research: Sex allocation, especially in Hymenoptera
E-mail: seger@bionix.biology.utah.edu
Web Site: http://www.biology.utah.edu/faculty2.php?inum=2
Tim
King. Present position: Fishery Biologist, USGS-BRD, Leetown Science center,
Aquatic Ecology Branch, Kearneysville, WV 25430
Present research: Genetic stock identification of Atlantic salmon inhabiting
North America; genetic population structure of horseshoe crabs in Delaware Bay.
E-mail: tim_king@usgs.gov
Web Site: http://www.lsc.usgs.gov/
My
Publications| 1973 | Brockmann, H. J. The function of poster-coloration in the beaugregory, Eupomacentrus leucostictus (Pisces: Pomacentridae). Zeitschrift für Tierpsychologie, 33: 13-34. |
| 1975 | Bayer, L. J. and H. J. Brockmann. Curculionidae and Chrysomelidae found in aquatic habitats in Wisconsin. Great Lakes Entomologist, 8: 219-226. |
| 1976 | Brockmann, H. J. and J. P. Hailman. Fish cleaning symbiosis: Notes on juvenile angelfishes (Pomacanthus, Chaetodontidae) and comparisons with other species. Zeitschrift f ür Tierpsychologie, 42: 129-138. |
| 1978 | Terborgh, J., J. Faaborg and H. J. Brockmann. Island colonization by lesser Antillean birds. Auk, 95: 59-72. |
| 1979 | Brockmann, H. J. and C. J. Barnard. Kleptoparasitism in birds. Animal Behaviour, 27: 487-514. |
| 1979 | Brockmann, H. J. Nest-site selection in the great golden digger wasp, Sphex ichneumoneus L. (Sphecidae). Ecological Entomology, 4: 211-224. |
| 1979 | Brockmann, H. J., A. Grafen and R. Dawkins. Evolutionarily stable nesting strategy in a digger wasp. Journal of Theoretical Biology, 77: 473-496. |
| 1979 | Brockmann, H. J. and R. Dawkins. Joint nesting in a digger wasp as an evolutionarily stable preadaptation to social life. Behaviour, 71: 203-245. |
| 1980 | Brockmann, H. J. The control of nest depth in a digger wasp (Sphex ichneumoneus L.). Animal Behaviour, 28: 426-445. |
| 1980 | Brockmann, H. J. Diversity in the nesting behavior of mud-daubers (Trypoxylon politum Say; Sphecidae). Florida Entomologist, 63: 53-64. |
| 1980 | Dawkins, R. and H. J. Brockmann. Do digger wasps commit the Concorde fallacy? Animal Behaviour, 28: 892-896. |
| 1980 | Brockmann, H. J. House sparrows kleptoparasitize digger wasps. Wilson Bulletin, 92: 394-398. |
| 1981 | Lucas, J. and H. J. Brockmann. Predatory interactions between ants and antlions. Journal Kansas Entomological Society, 54: 228-232. |
| 1983 | Brockmann, H. J. Animal behavior. In L. Johnson (ed.) Biology. W. C. Brown Co. Publ., pp. 826-859. |
| 1983 | Cohen, J. A. and H. J. Brockmann. Breeding activity and mate selection in the horseshoe crab, Limulus polyphemus. Bulletin Marine Science, 33: 274-281. |
| 1984 | Brockmann, H. J. The evolution of insect sociality. In J. Krebs and N. Davies (eds.) Behavioral Ecology. Sinauer Associates, Second Edition, pp. 340-361. |
| 1985 | Brockmann, H. J. Tool using in wasps. Psyche 92:309-329. |
| 1985 | Brockmann, H. J. Provisioning behavior of the great golden digger wasp. Journal Kansas Entomological Society, 58:631-655. |
| 1986 | Brockmann, H. J. Decision making in a variable environment: lessons from insects. In L. Drickamer (ed.) Behavior and the Dynamics of Populations. Privat Publ., Toulouse, France, pp. 95-111. |
| 1987 | Seger, J. and H. J. Brockmann. What is bet-hedging? In P. Harvey and L. Partridge (eds.) Oxford Surveys in Evolutionary Biology. Oxford Univ. Press, 4:182-211. |
| 1988 | Brockmann, H. J. Father of the brood (Conflict and cooperation in a mud-daubing wasp). Natural History Magazine. 97(7):32-37. |
| 1989 | Brockmann, H. J. and A. Grafen. Mate conflict and male behavior in a solitary wasp, Trypoxylon politum. Animal Behaviour 37:232-255. |
| 1990 | Brockmann, H. J. Mating behavior of horseshoe crabs, Limulus polyphemus. Behaviour 114:206-220. |
| 1992 | Brockmann, H. J. and A. Grafen. Sex ratios and life-history patterns of a solitary wasp. Behavioral Ecology and Sociobiology 30:7-27. |
| 1992 | Brockmann, H. J. Male behavior, courtship and nesting of the solitary wasp, Trypoxylon monteverdeae. Journal of the Kansas Entomological Society 65(1):66-84. |
| 1992 | Brockmann, H. J. and D. Penn. Male mating tactics in horseshoe crabs, Limulus polyphemus. Animal. Behaviour 44:653-665. |
| 1993 | Brockmann, H. J. Parasitizing conspecifics: comparison between Hymenoptera and birds. Trends in Ecology and Evolution 8:2-3. |
| 1994 | Brockmann, H. J., T. Coleson and W. Potts. Sperm competition in horseshoe crabs (Limulus polyphemus). Behavioral Ecology and Sociobiology 35: 153-160. |
| 1994 | Penn, D. and H. J. Brockmann. Nest-site selection in the horseshoe crab, Limulus polyphemus. Biological Bulletin 187: 373-384. |
| 1995 | Penn, D. and H. J. Brockmann. Age-biased stranding and righting in horseshoe crabs (Limulus polyphemus). Animal Behaviour. 49: 1531-1539. |
| 1996 | Brockmann, H. J. Satellite male groups in horseshoe crabs. Limulus polyphemus. Ethology 102: 1-21. |
| 1996 | Arnold, S. and H. J. Brockmann. Evolution of behavior. Approaches to studying behavioral change IN Foundations of Ethology, L. D. Houck and L. C. Drickamer (eds.). University of Chicago Press. pp. 673-682. |
| 1996 | Dyer, F. C. and H. J. Brockmann. Sensory processes, orientation and communication: Biology of the Umwelt. IN Foundations of Ethology, L. D. Houck and L. C. Drickamer (eds.). University of Chicago Press. pp. 529-538. |
| 1997 | Brockmann, H. J. Cooperative Breeding in Wasps and Vertebrates: the Role of Ecological Constraints. IN Evolution of Social Behaviour in Insects and Arachnids, J. Choe and B. Crespi (eds.). Cornell University Press. pp. 348-371. |
| 1997 | Brockmann, H. J. Xiphosurida. McGraw-Hill Yearbook of Science and Technology 1997. McGraw Hill, NY. pp. 491-493. |
| 1999 | Harari, A. R. and H. J. Brockmann. Male beetles attracted by females mounting. Nature 401:763. |
| 2000 | Harrari, A. and H.J. Brockmann. Intrasexual mounting in the beetle Diaprepes abbreviatus. Proceedings of the Royay Society of London B 267: 2071-2079. |
| 2000 | Brockmann, H.J., C. Nguyen and W. Potts. Paternity in horseshoe crabs when spawning in multiple male groups. Animal Behaviour 60:837-849. |
| 2001 | Sirot, L. and H.J. Brockmann. Costs of sexual interactions to females in Rambur’s forktail damselfly, Ischnura ramburi (Zygoptera: Coenagrionidae). Animal Behaviour. 61:415-424. |
| 2001 | Brockmann, H.J. The evolution of alternative strategies and tactics. Advances in the Study of Behavior. 30:1-51. PDF |
| 2001 | Hassler, C. and H.J. Brockmann, H.J. Evidence for use of chemical cues by male horseshoe crabs when locating nesting females (Limulus polyphemus). Journal of Chemical Ecology. 27: 2319-2335. |