Field Research Station at Fort Missoula

Through the Office of the President at The University of Montana, land and an existing building (appraised at $2,048,000) was offered to the Department of Geology and Division of Biological Sciences to commence development of a Field Research Station. This multi-disciplinary research facility, located 12 minutes from our main campus, consists of a 9,000 sq. ft building and 100 ha of surrounding land. It will provide an ideal bridge between our current University facilities and the largest expanse of wilderness surrounding any university in the lower 48 states, and allow controlled, semi-natural, field and laboratory experiments for the students, research associates, and faculty in the Geological and Biological Sciences. The building, however, is old but structurally sound, and without internal structure. The University of Montana has thus also provided us with $700,000 for renovations to the building (which are due to be completed by December 1998). This outlay of capital is truly unique within Montana, a state where capital expenditures for research are very rare and limited.

The task of turning a former horse stable into a functional research facility was assigned to the architectural firm of MacArthur, Means & Wells, of Missoula MT. That this firm would design an avian research facility could be considered providence. More on this later.

 Floor plan (courtesy MacArthur, Means & Wells)

The Division of Biological Sciences at FRSFM

The University of Montana has been aggressively building an active group of ecological and evolutionary biologists. Our research programs have been limited by the physical infrastructure and computational resources available to us on campus. The Field Research Station will enable us to house animals under semi-natural conditions and to investigate a variety of research topics in controlled environments that have not been previously available at The University of Montana. The research activities of the major users are described below in more detail. It should be noted that the faculty oversee two extensive avian regional and national databases; the BBIRD (Breeding Biology Research and Monitoring Database) program is directed by Tom Martin and includes data on more than 30,000 nests of more than 150 bird species and more than 100 habitat variables and nesting success information. The US Forest Service Northern Region Landbird Monitoring Program is directed by Richard Hutto, and includes count and habitat data from more than 10,000 points distributed throughout the northern region. These two databases are used by these faculty, and by University of Montana postdoctoral and research associates and graduate students. They will also be accessible by scientists throughout the country via the Internet for additional analyses. The Field Research Station's computer network also will be integrated with the network of the main campus at The University of Montana allowing access to the data sets stored at the research facility from campus and vice versa.

The University of Montana is in an unique position to provide a broad inter-disciplinary approach to investigation of these topics because it arguably holds the largest inter-disciplinary group of faculty working on birds in the world. In total, the faculty includes 9 individuals with active research programs on birds who mentor ca. 30 graduate students. Four of these faculty form a key central role (Dial, Greene, Hutto, Martin) in the development and management of the facility proposed here and represent a diverse array of disciplines (habitat selection, foraging behavior, social behavior, parental care behavior, flight, morphology, physiological ecology, life history evolution, species coexistence). In addition, the faculty has recently added other individuals with vigorous research programs and we are currently undergoing a search for a new faculty member that will add additional vigor to our group. All of these individuals have nationally and internationally-recognized research programs.

Below are brief outlines of research problems that the four principal investigators (listed alphabetically) plan to attack at the new Field Research Station. 

Kenneth P. Dial - Flight Laboratory and Director of the Field Research Station at Fort Missoula

Most animals fly. There are over 1 million flying insects and more than 10,000 of the 13,000 species of endothermal vertebrates (bird and mammals) fly. While we marvel at the locomotor abilities of birds, our understanding of both internal and external mechanisms responsible for these organisms taking to the air is just beginning to unfold. With the advent of new biomechanical and electrophysiological recording devices we are now able to peer inside the animal as it performs various locomotor behaviors. Not only has our research shed new light on many aspects of bird flight, but the results are of interest to others studying animal locomotion (Taylor, 1996) and also aerospace engineers in the design and flight-control of current and future aircraft (Dial, 1994 and Dial, 1995)

All aspects of research in our lab are centered around the quantification of movement and behavior of flying vertebrates. As such, we generate extraordinary volumes of digital data from high speed video cameras as well as the concurrent recording of a variety of biomechanical and neurophysiological processes. We are anxious to utilize the requested proposed computer network as our need for computer storage space, computational studies, and modeling of flight continues to escalate. In our current Flight Lab and in our new field facility we study aspects of both steady-state flight (i.e., straight and level flight over a range of speeds) and nonsteady-state flight (i.e., maneuvering, takeoff and landing) to better understand the biomechanical, physiological, and aerodynamic parameters associated with the biology of flying animals. Over the past 10 years, researchers working in the University of Montana Flight Laboratory have focused on the neuromuscular control (Dial et al., 91, Dial, 92, 93, Boggs and Dial 93), biomechanics (Biewener et al., 92, Dial and Biewener, 95, Biewener and Dial, 96, Warrick et al. 98, Warrick and Dial 98), scaling (Tobalske and Dial, 96,97), respiratory biomechanics during flight (Boggs et al., 97a,b), ecology (Tobalske et al., 97, Tobalske, 96, Warrick 98) and evolution (Gatesy and Dial, 96a,96b) of avian flight. We have used various forms of conventional video and high-speed 16mm film to record the kinematics of avian (Tobalske and Dial, 96) and chiropteran locomotion (Boggs and Dial, unpubl). This includes the only records of high-speed movie x-ray (cineradiography) of the skeletal apparatus (Jenkins et al. 88), muscle activity patterns during different modes of flight, measurements of skeletal stress and muscle strain in various species of birds (starlings, magpies, pigeons, parakeets, quail, pheasants, turkeys, and ducks) during flight.

(See website: http://biology.dbs.umt.edu/flightlab/flightlab.htm)

Field measurements of mechanical power output: We have developed the technology to measure the mechanical power output of birds during flight. This is a direct method to determine the cost of flight at various speeds and during different behaviors. Our goal is to extend these techniques using telemetric signal processing in order that the study subjects may fly freely within an aviary. To date there are no reliable data available on the cost of flight and flight styles in any free-flying vertebrates. In order to understand the selective pressures on animal design it is imperative to understand the relative cost associated with each behavior. The requested digital storage and analysis system will be absolutely necessary to store the massive data bases built from digital video, digital biomechanical recordings and modeling.

Scaling energetics and biodiversity: We are currently designing projects to better understand why the vast majority of flying birds are small (25-75 g). Given that the cost of locomotion per unit mass decreases with whole body mass, a clear dilemma is why selective forces result in the dominance of smaller-sized species. We are now able to predict why flying birds don't exceed the 10-12 kg range (using muscle power output recordings within a family), but no one has been able to clearly address (quantitatively) how and why most birds are small. Using established and new biomechanical techniques created in our lab, we plan to study maneuvering flight, cost of flight, and predator-prey strategies in order to enhance our understanding of the design, physiology and evolution of the avian body.

Aerial flight displays and body size: Courtship flight displays in small birds appears to be common and ubiquitous. I will collaborate with several of my colleagues to document the frequency, context, and diversity of aerial flight displays based on the energetics of flying and body sizes. In particular, with Dr. Greene, we will integrate courtship and agonistic behaviors during flight displays with concurrent vocal displays by several species of passeriform birds. This will require high-speed filming, vocal recordings, and a clear understanding of the reproductive effort and success of the individuals under study. Here be believe we can provide new insight of multiple display criteria in the differential reproductive success of birds. The above projects are going to require substantial digital storage for video, biomechanical, and neurophysiological data generated from experiments. Also, public access to the video of birds in the field and in the flight arenas will be available through the server.

Public education: Over the past two years, I have been on a quest to discuss basic yet important biological themes to the general public. To this end, I am the host and scientific consultant for a nature program ("All Bird TV") which airs on the Discovery Communication's Animal Planet to an international audience. The current world-wide subscriber volume is over 40 million people. We have successfully highlighted the work of six researchers (Drs. Greene, Hutto, Marks and graduate students: Perkins, Osborne, and Tewksbury) within the University of Montana's Avian Biology Program in our 26 episode series on bird biology (see website http://www.discovery.com/). In addition, my research has been highlighted on four internationally televised programs: Discovery Magazine, Scientific American Frontiers, Nova, and Chronkite/Ward.

Erick Greene - Evolution of communication and signaling

My research has focused on understanding the adaptive significance of behavior and signaling in animals (Greene and Meagher 1998, Greene et al. 1987, Greene 1989), the evolution of behavioral and morphological plasticity (Greene 1999, 1996, 1989), as well as understanding ecological, physiological, and behavioral factors that influence the distribution and abundance of animals (Weathers and Greene 1998). During the past five years, I have been focusing on vocal communication, plumage signaling, and conservation biology of Lazuli Buntings, a neotropical migrant songbird (Greene et al. 1996, Muehter, Greene and Ratcliffe 1997, Greene in press, Greene, Jolivette and Redmond in press). This species is a fascinating model for studying the evolution of social signals because the males have delayed acquisition of two important social signals: (1) delayed plumage maturation, in which young males take several years to attain the bright plumage of older adult males; and (2) delayed song learning, in which yearling males develop a song only after they arrive on the breeding grounds.

My intensive studies of individually marked, free-living buntings have shown considerable individual variation in both of these important social signals. For example, in habitats in which competition among males for limited territories is intense, we have found that (1) yearling males are likely to copy the song of one older neighbor (Greene et al. 1996), and (2) yearling males with the dullest plumage are more likely to attain a good territory and attract a social mate than brighter yearling males. However, in habitats in which territories are not limited, and competition among males for territories is negligible, we have found that yearling males tend to develop unique songs that are unlike adult neighbors, and they tend to be brighter. Model presentation experiments have shown that in high male-male competition environments, the dull plumage of yearling males elicits lower aggression from older territorial males (Muehter et al. 1997). Thus the dull plumage of young birds may be adaptive in that it allows them to settle near older males without incurring the cost of fighting with them (Greene et al., in review, Science).

Although these detailed field studies have revealed previously undocumented variability in song learning behavior and habitat-related plumage patterns, we have now reached the limits of which questions we can address in the field. In particular, in studying the adaptive significance of signals, such as plumage signals and vocal behavior, which are clearly important in social interactions, we need to be able to experimentally manipulate social conditions (e. g. flock size and sex ratios, interaction levels by varying food amount, quality, and spatial dispersion). We are excited since we now have the opportunity to conduct controlled experiments in aviaries at our newly emerging Field Research Station.

Evolution of phenotypic plasticity, polymorphism, and signal polymorphism. In addition to my research on signal evolution in birds, I have been investigating the ecological and evolutionary causes and consequences of phenotypic plasticity (Greene 1999). I am currently working on a group of fascinating caterpillars in the inchworm genus Nemoria. I discovered that individuals in one species, Nemoria arizonaria, can develop into two very different morphs (an oak flower mimic or an oak twig mimic), depending upon what they eat as larvae (Greene 1989, 1996, 1999). The genus is very large, with over 70 species in North America. It is an fascinating model system, since different species have made evolutionary shifts onto different larval hosts, and their larval morphology and developmental programs have changed in interesting ways. I am planning to examine how developmental flexibility and constraints have influenced radiation and host shifting in this group. This research requires that I be able to quantify the morphology of caterpillar integument (e.g. color, patterns on the skin, shape, size, and location of protuberances on the skin) from digital images taken from a microscope. This morphological analysis will be similar to those performed by Douglas Emlen (described below), and it will generate extremely large data sets that will require a powerful computer for image capture, storage, and analysis.

In addition to significantly enhancing my own research projects, the computer network we are requesting would also allow us to more easily collaborate. I am excited at the prospect of being able to collaborate on a wide variety of projects with my colleagues, such as visual and vocal displays during courtship (with Ken Dial and Tom Martin), function of the many non-song vocalizations (eg. chips and buzzes) of flocking birds (with Dick Hutto) and in response to predators (with Tom Martin), and correlations of song characteristics (loudness and frequency characteristics) and parasitism risk, by species of song birds parasitized by Brown-headed Cowbirds (with Tom Martin). The equipment we are requesting would provide us with the ability to do these.

Richard L. Hutto - Foraging and Community Ecology

My research career has revolved around foraging ecology, and the extent to which differences in the mode of food acquisition allows coexistence in both mammal (Hutto 1978) and bird communities (Hutto 1981, 1985a, 1988, 1990), and the extent to which food abundance can explain patterns of habitat use in birds (Hutto 1980, 1985b). Most recently, I have developed a strong interest in the way documented effects of land use practices on bird communities (e.g., Tobalske et al. 1990, Hutto 1993, 1995, 1997, Villaseñor and Hutto 1995) might be the result of increased costs associated with food acquisition. As emphasized by Martin (1986), mechanistic studies must focus on the activities of individuals if we are to gain a better understanding of the potential evolutionary force of competition. I am moving into the use of high-speed video to allow us, for the first time, to evaluate whether changes in parameters such as vegetation architecture, competitive mileau, or predation risk can act through changes in foraging efficiency of individual birds to make an otherwise suitable habitat unsuitable.

Competition for food by arboreal insectivores: whether breeding bird communities are structured (which species are present and where they occur within a habitat) by competition for food is a still-unresolved subject. Because food is so abundant during the breeding season, most biologists believe that the foraging activity of individuals of one species cannot reduce food to an extent where it might affect the foraging success of another species. Nonetheless, the economics of foraging suggests that competition for food is not only possible, but an important force in structuring bird communities (Hutto 1985a). Yet, past studies have been overly simplified in their quantification of foraging differences among coexisting species. By using multiple-camera video technology to record the 3-dimensional position and feeding rates of free-ranging birds that use a series of trees in the riparian bottom land adjacent to field station, we will determine whether feeding locations and rates of a given species are affected by the number and duration of visits by other species that use the same trees and we can quantify subtle but important differences in foraging.

Coexistence mechanisms: One of the most interesting aspects of the foraging behavior of the vast majority of nonbreeding insectivorous forest birds worldwide is the fact that they forage in mixed-species flocks (Hutto 1987, 1994). The diversity of species that participate in these social foraging groups begs the question of how they can coexist at a time of food scarcity while often foraging in the same individual trees. Subtle differences in the manner in which the different species achieve the same overall foraging rate (Hutto 1988) might be the primary mechanism they use to reduce overlap in resource use. Again, we plan to use high-speed video to document the kinds of foraging differences that exist among flock mates, how those differences are associated with differences in morphology, and whether such foraging differences promote coexistence; (b) a second focus would involve the set of small Empidonax flycatchers that coexist in Montana in summer. This group is interesting because some are syntopic (e.g., Hammond's and Dusky flycatcher) but the seem to do basically the same thing. High-speed video is expected to reveal subtle but distinct differences in foraging technique, flight style, and strike distances that cannot be revealed easily with the naked eye. This research will soon be possible in the aviaries at the Field Research Station.

Effects of timber harvesting: In a continuing collaborative effort with Plum Creek Timber Company, the Salish and Kootenai Tribes, U.S. Forest Service, Bureau of Land Management, and Montana Fish, Wildlife and Parks (see description of program at http://www.umt.edu/biology/dbs/landbird.htm), we will couple field observations of bird foraging behavior in harvested and unharvested forests with controlled laboratory observations at the field station to better understand how changes in forest architecture might affect the foraging efficiency of some common forest bird species. As part of these collaborative efforts, I oversee the management of The US Forest Service Northern Region Landbird Monitoring Program. This massive regional data base (the largest in the country) of bird-habitat relationships includes count and habitat data from more than 10,000 points distributed throughout the northern region. In addition, scientists from around the world would be allowed access to this database through the Internet Server.

Public education: I am playing an active role in digesting and packaging scientific findings from research conducted at the Field Research Station and elsewhere for students and the public at large. For example, I am incorporating ornithological information into a web page for students (and eventually the general public) through a pilot project with The University of Montana's Information Technology Resource Center. In addition, I serve as the host for the Connecticut Public Television series "BirdWatch," which reaches 85% of the nation through PBS television stations. I plan to use the Field Research Station's computing network to provide sate-of-the art access to scientific information about science in general, and birds in particular.

Tom Martin- Life History Evolution and Habitat Selection

Life History: Recently, I have re-emphasized a hypothesis by Skutch (1949) and pointed out that one mechanism that may influence life history evolution is constraints of nest predation risk on parental activity at the nest site (Martin 1996a). My graduate students and I (Martin and Ghalambor, in press; Ghalambor and Martin, in review; Conway and Martin, in review) have shown that variation in incubation activity among species is better explained by variation in nest predation than by the long-accepted hypothesis of microclimate. Similar effects during the nestling feeding period could help explain evolution of clutch size differences. Recent work by Julliard et al. (1997) suggest that birds exhibit phenotypic plasticity in clutch size in response to nest predation. However, the mechanism underlying such responses are unclear. One likely possibility is that birds monitor presence (i.e., encounter rates) of predators and decrease their activity and clutch sizes with increasing encounters. Yet, if birds decrease activity during the nestling stage, then they should increase the amount of food that they bring per trip (increased food loading) (Martin 1996). Tests of such mechanisms are critical for understanding new evidence showing the critical importance of nest predation and nest sites on life history traits (Martin 1993a, 1993b, 1995, Martin and Clobert 1996, Julliard et al. 1997). Video surveillance of nesting birds that differ in their risk of predation can allow tests of whether nest visitation rates are decreased and food loading is increased. Video images can be uploaded into the computer and food loads can be measured using digitized image analysis with measured bill size being used as a calibrator. However, such image analyses require extensive data storage space and high speed computing capabilities to allow rapid processing of large numbers of images.

Risk of nest predation may also constrain acoustical properties of nestlings; examination of acoustics of nestling begging calls among 24 coexisting bird species shows that amplitude (loudness) and frequency (affecting attenuation) are strongly correlated with risk of nest predation (Briskie, Martin, and Martin, in prep.). This acoustical work is continuing in new projects in Latin America where nest predation is thought to strongly shape life history traits, but rigorous tests are missing. 

Foraging and Male Guarding: If predators constrain the time that parents can get on and off the nest, it can constrain energy acquisition and influence foraging behavior while off the nest; if off bouts are constrained, then females should exhibit more rapid and intensive foraging while off the nest. Field data suggest that females forage much faster during incubation off-bouts than prior to incubation (Dobbs and Martin, in press), but consequences of predation on such behavior is unknown. Females of several species appear to signal males to guard the females during incubation off-bouts (Barber et al., In press; Martin, pers. obs.). Acoustical analysis of call notes of females when leaving the nest in conjunction with Dr. Greene can aid understanding of these pair interactions. Future paternity does not explain the behavior (Barber et al., in press), but a reasonable alternative is that guarding males may allow females to be more efficient at foraging by reducing their need to be vigilant. Video monitoring of birds that are exposed versus not exposed to nest predators in the presence and absence of males can assay variation in their incubation on and off bouts and the subsequent effects on foraging behavior, speed, and efficiency, as well as the consequences for female physiological condition as measured by changes in body mass. Such video monitoring of female foraging in the context of nest stage and variation in predators, mates, and food can strongly enhance understanding of the role of food and foraging in life histories and habitat selection. 

Habitat Selection and Species Coexistence: Work on habitat selection and species coexistence of birds historically has focused on the importance of food and competition. Work in the Martin lab has shown that nest predation may play a strong role in both habitat selection and species coexistence (Martin 1993c, 1996b, 1998). Most recent work, however, is suggesting that species interactions may include a complex interplay between time budgets, food and predation. Within and among species interactions may influence time budgets (males may need to spend more time in territorial defense both intra-and interspecifically with increased abundance of conspecifics or ecologically-similar heterospecifics) that then influence ability of males to feed females during incubation and to feed nestlings (Martin and Martin, in review). However, feeding rates can also be constrained by nest predation (Martin and Ghalambor, in press; Ghalambor and Martin, in review). Songs of ecologically similar species may converge to increase spacing and acoustical analysis of songs in the presence and absence of conspecifics and heterospecifics can be used to examine such possibilities. Moreover, play-back experiments can be used to directly test response and interactions of ecologically similar species to each other. In addition, detailed video analysis of male and female behavior in the presence and absence of ecologically-similar heterospecifics, nest predators and variation in food supply can provide significant new insight into the mechanisms underlying potential costs of habitat selection and species coexistence. 

BBIRD (Breeding Biology Research and Monitoring Database) Program: The BBIRD program uses standardized sampling protocols to monitor nest fate and associated habitat of birds throughout North America. The program includes collaborative involvement of scientists from Universities, Smithsonian, and federal agencies at 42 sites in 27 States. The database currently includes data on fate and habitat of more than 30,000 nests of more than 150 bird species, including more than 100 variables per nest and the database is growing rapidly. This database allows examination of landscape level processes with true replication for the first time ever; allows examination of variation in life history traits across the geographic range of species; allows identification of habitat features that are critical to habitat selection and successful reproduction of species; and allows comparisons of nesting success of species under differing land management conditions. We anticipate that the University of Montana Field Station at Fort Missoula (Phase I renovations are scheduled to be completed in November 1998) will soon be a vigorous research facility that will be well used by many faculty and students. In addition to the four co-PIs described above, we include brief descriptions of the research programs of two additional faculty who would make use of the facility.

Douglas J. Emlen - Development and evolution

My research utilizes behavioral, genetic and ecological experimental methods to explore the evolutionary significance of exaggerated or unusual morphological traits in insects (Emlen 1994, 1996, 1997a, b, Moczek and Emlen in press). Beetles of the genus Onthophagus (Coleoptera: Scarabaeidae) exhibit extraordinary variation in male morphology. Males in most species have Ahorns,@ and the horns vary interspecifically in size and shape, as well as in the location on the animal from which they extend (e.g. some species have horns on the head, others have horns on the thorax). Male investment in horns can be extreme, and this raises two questions: (1) does production of a horn cost a male in any detectable way (e.g. by necessitating reductions in the sizes of other traits like wings or eyes; Nijhout and Emlen 1998), and (2) can such costs (defined here as allocation tradeoffs) bias, or otherwise influence, the directions of evolution in this genus? 

Addressing these questions requires large controlled-breeding experiments, with huge numbers of families, and offspring within families. All individuals must be measured for a variety of morphological traits, including the areas of the eyes, wings, forelegs, hindlegs, antennae and mouthparts. To accomplish this, I employ a dissecting microscope with a digital camera and image analysis software (funded by NSF MONTS Program award #291832). I also plan to conduct many of my future breeding experiments (NSF IBN-98-07932) at the proposed field station.

Delbert Kilgore - Respiratory physiology

In my lab, we are studying the sources and potential fitness value of phenotypic plasticity in the respiratory control systems of birds and mammals. Specifically, we are interested in (1) the mechanisms underlying the attenuated ventilatory response of mammals and birds living at high altitude or in underground burrows to changes in their respiratory environments (Kilgore et. al. 1985, Colby et al. 1987, Williams et al. 1995, Walsh et. al. 1996, Frappell et al. in press), (2) the effect of respiratory environment during development on plasticity in the respiratory control systems of these organisms (Williams and Kilgore 1992), and (3) the fitness value of the observed plasticity.

Jeffrey Marks - Ecology and reproductive behavior