Aesthetic Seeing in the Biology Classroom: Understanding, Selecting and Utilizing Ornithological Images to Promote Engagement and Learning in Ecology.

Learning in biology is frequently dependent upon images as a means of communication, where images are construed as any painting, drawing, photograph, map, chart, or graph from which scientific information or ideas may be extracted. Our orientation toward learning visually is to be expected, for as Aristotle pointed out, we think in images. A static image invites an intellectual response, and as the viewer sees into the image, both knowledge and personalized meaning are constructed; thus the benefits of understanding a given illustration of biological material reach beyond comprehension of the specific content (Robin, 1992).

In The Scientific Image from Cave to Computer, Robin describes how use of carefully selected images has the potential to fulfill a number of educational and cognitive functions as the viewer is engaged aesthetically. Such images may describe and explain an organism or phenomenon, demonstrate cause and effect relationships, provoke inductive reasoning about the subject matter, teach visual classification skills by example, and promote the development of explanations, forcing in turn the utilization of previously learned concepts. A further possibility is that the development of observation skills during focused work with biologically oriented images might encourage that special sensitivity to biological phenomena and discernment of fine distinctions that is a hallmark of a skilled biologist and which may mature into the aesthetic seeing espoused by Dewey (1934).

Aesthetic seeing implies that the observer goes beyond compartmentalizing and categorizing to develop a deeper knowledge of the organism or phenomenon observed, and that the fruit of this deep knowledge is the ability to "ask the right questions" (Janovy, 1996, p.20) as the observer sees beyond surface characteristics into structure, process, and complexity. In turn, aesthetic seeing has the potential for science learning to metamorphose into a transformative experience, where science concepts are naturally applied in everyday contexts and serve to enrich student lives beyond the classroom (Pugh and Girod, 2007). When this transformation has been achieved, the metaphors occupying an observer's mind become insistently scientific (Janovy, 1996; Pugh and Girod, 2007). A vacant lot, for example, brings reflections upon stages of ecological succession, a busy metropolis stimulates thinking about niches and human behavior, and experience with a garden pest raises questions and ideas for experiments. Thus a thoughtful use of images might well be one useful step toward incorporating science knowledge into student life beyond the classroom.

The majority of the image types discussed by Robin (1992) and in Edward Tufte's four volume work on effective scientific graphics ( Tufte, 1983;1990;1997;2006) are outside the scope of this article. The discussion here is limited to ornithological images as a representative sub genre of natural history illustration and the potential use of such images in the biology and environmental science classroom.

Organisms are the essential subject matter of natural history illustration, and the genre ranges from detailed technical drawings of single species of plants, birds, fish, insects, etc to complex full background paintings in which multiple species interact to produce ecological and/or behavioral narratives. As Janovy (1996) points out, natural history illustration exercises a special power over the viewer. We perceive the living world as a set of organisms: while biology and environmental science are rich in theory and abstraction, it is fascination with plants and animals that has historically exerted the greatest influence upon biological thinking, and that has fundamentally driven the science: "while we endure abstractions, we prefer to see things that are alive. Failing that, we like to see things that our minds can easily translate into living organisms. More often than not, this desire is manifested as dependence on a picture" (p. 35). A fine natural history illustration serves this need well: even complex images containing unfamiliar organisms depicted under circumstances that may be culturally, geographically, and historically far removed from the life experience of the viewer are relatively easily decoded, for the aesthetic response to an attractive or intriguing image serves a hook to engage him or her with the scientific content. Unlocking the meaning of such images confers a rare privilege upon the viewer as it bestows a glimpse of the shared intellectual passion for organisms that has animated both the scientific community and many amateur naturalists for centuries (Janovy, 1996; Robin, 1992; Wilson, 1984)

My own work with using natural history images in the science classroom has centered upon bird painting, for the long history of bird images reflects the breadth and depth of human fascination with this class of vertebrates. The artistic expression of interest in birds is evident today in the many ornithological illustrations currently found in not only in technical articles and textbooks, but in the several hundred field guides published annually, art books, gallery paintings, and an increasing number of children's books. This genre is unlikely to be replaced by photographic images of birds: although contemporary painters frequently work from large collections of photographs, the attentive selectivity of an expert observer's eye in a painting conveys a better sense of the organism and its environs than do moments frozen in time by a camera lens.

As an interest in birds has manifested across many cultures and walks of life, bird art also has a wide appeal and occupies niches within art, science, and history. The oldest known depictions of birds on cave walls in southern France and northern Spain (c. 14,000 B.C.) frequently show ravens accompanying the conclusion of a successful hunt, and bird illustrations in the psalters of the Middle Ages emphasize the symbolic and spiritual importance of these organisms. As interest in science reawakened in the Renaissance, Leonardo da Vinci executed detailed studies of bird anatomy intended ultimately to open the power of flight to humans. Five centuries later, following the establishment of ecology as a sub-discipline of biology and the public concern with the environment that followed, birds appeared as a conservation motif in the works of twentieth century landscape painters such as Robert Bateman, Lars Jonsson, and Bruno Liljefors.

The fascination of birds for man has been extensively explored by Joseph Kastner (1986) and ornithologist Edwin Streseman (1975). Among other factors, the appeal of birds stems from their availability to the casual observer, the sensory hook of bright coloration and the texture of feathers, and an almost mystical identification with the ideal of personal freedom represented by the power of flight, and evocative images of birds appear to provoke an acute identification with the natural world in many individuals (Kastner, 1986; Stresemann,1975). Long before ornithology became a methodological science in the late 18th and early 19th centuries, and birding became an extensive and commercially important leisure interest in western culture in the 20th century, there have always existed a substantial number of individuals whose lifestyle encompassed birds in some meaningful fashion. Hunting, trapping, observing, and admiring birds were activities that frequently evolved into a focused passion, producing new scientific knowledge and deeper understanding of the organisms and their environment.

Bird study provides a path to many other aspects of the biological world. Ornithologist Frank Gill (1995), wildlife ecologist Stanley Temple (1977), and population biologist Paul Ehrlich (1970; 1988) are among many authors who have pointed out that avian biology and ecology have wide implications in biology, at the level of the individual, the population, the community, and the ecosystem. Many bird illustrations explicitly or implicitly convey aspects of bird biology that demonstrate, explain, or reinforce wider concepts in ecology, and both the intrinsic appeal of the subject matter and the wide availability of bird art suggest that there is much potential for educational use of these images.

Straightforward bird portraits, such as those found in field guides, explicitly introduce the ideas of adaptation and natural selection, as they distinguish between bill, wing, and foot structures of different species. Illustrations of differences in plumage and behavior—between sexes and in different seasons—imply ideas about visual communication, evolution of displays, and sexual selection. Narrative paintings that depict several species of birds—and also include other organisms—introduce aspects of community biology, the interdependence of species, food webs and energy flow, and may portray important and relevant aspects of the local habitat or wider region where the organisms are typically encountered. The National Science Education Standards (NRC, 1996) recommend that characteristics of organisms, their life-cycles, behavior, their relationships to the environment, diversity and adaptation, interdependencies between species, and studies of populations, communities, and ecosystems be addressed at the grade school level, especially as these relate to the theory of evolution, which now occupies a central position in biology and biology teaching. Many of these topics have the potential to be introduced or demonstrated by carefully selected ornithological illustrations, and a particular advantage of so doing is that of all the vertebrates, live birds are a visible and ubiquitous presence throughout the day for most students, and thus reinforcement of these concepts and principles is close at hand.

Selection and Location of Ornithological Images

Selection of ornithological images for science education ideally requires that we find illustrations that command the attention, offer aesthetic engagement, transmit scientific information, and stimulate reasoning and questions in the viewer. Almost sixty years ago, Wilfred Blunt (1950) argued that serving the two masters of art and science simultaneously is exactly what the natural history illustrator is supposed to do: the successful aesthetic portrayal of an organism reveals its essence, focuses attention on the fine details that make it distinctive, and suggests that it is worthy of further contemplation or investigation. Achieving these goals presupposes some scientific intent on the part of the illustrator/artist: an intent that is not necessarily present in all natural history illustration and certainly not in much 'wildlife art.' However, the majority of bird painters are accomplished naturalists, and quite a number are distinguished ornithologists, rendering the task of selecting an ornithological image that possesses both scientific integrity and aesthetic sophistication relatively straightforward. Ornithological artists represent a subset of expert ecologists, for the practice of field ornithology that is necessary for the production of these images is largely rooted in ecological expertise. Some knowledge of the history of ornithology—and the concomitant growth of bird painting as a sub-field of natural history illustration—definitely facilitates the location of bird illustrations for use in science education (see Appendix 1: A Brief History of Ornithological Illustration and an Introduction to Selected Bird Artists). Names of painters whose works have potential for classroom use, and which are available as internet images is provided in Appendix 2 (Bird Artist Names for Image Searches), and a few hours of image searching will generally yield a fine collection of material.

The quantity of bird paintings available from online sources is so great that employment of some guidelines in making appropriate selections for classroom use is to be recommended. The work of Edward Tufte in the analysis and design of informational graphics provides an elegant theoretical background for the selection of scientific images that meet exacting standards of excellence for the transmission of information while providing aesthetic satisfaction and stimulation (Tufte, 1983; 1990; 1997; 2006). Within Tufte's schema of scientific information design, a scientific illustration is conceived as a 'data map' of the organism or phenomenon depicted, and his design theory for the effective display of such data rests upon two major axioms. First, the maximum possible information about the subject incorporating many different variables or dimensions of the subject matter) is to be communicated as efficiently as possible, that is , "the larger the share of the graphic's ink devoted to data, the better…Maximize the data-ink ratio, within reason" (Tufte, 1983, p.96). Second, graphics should be 'relational', explicitly showing relationships between variables or components of the illustration wherever possible. Relational images link two or more variables, communicate complex ideas with clarity, precision, and understanding, and encourage the viewer to assess possible causal relationships.

Within the area of natural history illustration, well-designed field guide plates are frequently packed with information, thus exemplifying the maximum data-ink principle, while narrative paintings offer more relational possibilities. Tufte (1983) found that only 42% of science graphics overall might be considered relational in nature, but modern ornithological illustration considerably exceeds this estimate. For example, a painting of a rainforest scene that contrasts species typically occupying the understory to those found in the canopy has a number of relational possibilities in terms of ecological, behavioral, and morphological variables, as does a painting of a mixed species group of shorebirds, showing the zones where they are typically found feeding.

Satisfying both of Tufte's axioms may present a conflict for the natural history illustrator, for the two objectives are not always easily reconciled. The influential ornithological illustrator Louis Agassiz Fuertes was forced to seek this reconciliation in the early stages of his career, while he was under the competing influences of three powerful mentors, and it is a measure of Fuertes' talent and flexibility that he managed to incorporate these apparently conflicting directives into data-rich and accurate species profiles that provide rich context without gratuitous decoration (see Pasquier & Farrand, 1991; Peck, 1982; Sutton, 1979). However, few producers of bird art rival Fuertes, and selection of images by the science educator will generally require choosing an illustration based on aesthetic features and secondarily running a 'data check' on the biological content explicitly depicted, and the potential of the image for stimulating visual reasoning and generation of new questions.

Visual Learning and Visual Literacy

A final consideration in selection of images for the classroom concerns the visual literacy of the student. A good scientific image should present a moderate cognitive challenge, and a considerable time investment is required to probe the content of, or benefit from, an information-dense graphic. Having selected a painting that is complex and information-dense, pedagogical problems frequently arise with respect to the literate use of such images. Students may not be equipped to extract the information contained, either because of a lack of visual literacy skill, or due to insufficient content knowledge. The issue of visual literacy is not trivial: one study of high school biology students indicated that the level of visual comprehension required to interpret illustrations varied by six grade levels (Blystone and Dettling, 1990).

Maximizing visual learning frequently requires a teaching approach that promotes increased interaction and engagement with the material presented. Using 'induced picture strategy' (Alesadrini and Rigney, 1981): developing and encouraging students to develop visual representations of their own knowledge (induced pictures) appears to help with understanding and effective interpretation of imposed pictures (those prepared for the student). Making graphic representations of one's own knowledge is an effective learning tool that few students utilize, and one which prepares them to extract maximum educational value from images prepared by others.

Ecology and environmental science are each pursuits that demand synthesis of a number of previously acquired concepts (McComas, 2002). A complex narrative painting may explicitly depict, or rely upon the implicit understanding of, up to 25 ecological concepts (Appendix 3: Ecological Concepts Depicted Or Referenced Within A Sample Of Bird Paintings, Table 3.2) Provoking awareness in students of their own content knowledge in the area that is required to successfully interpret the illustrations presented, by cueing with verbal or written instructions may be helpful in directing them through critical illustrations. A study of 10th grade students found that maximum visual learning occurred either when students were given no cues at all and forced to develop their own approach to the material, or when their learning was thoroughly supported through presentation of 20 questions linked to the subject of the illustration (Holliday, 1981). Partial support by presenting students with a small number of questions (Holliday used five), or accompanying the image with a prose passage did not prove effective: this is a promising area for more research and informal experimentation by teachers. A successful approach to visual literacy with older students who are fluent writers has involved the translation of a complex illustration back into words, assisting realization of the density of information contained within the image (Blystone and Dettling, 1990).

The reader may understandably consider at this point that the use of bird images in the biology classroom is unduly arduous. The educator is required to steep himself/herself in the subject matter of ornithology and the history of the discipline, develop familiarity with the associated genre of illustration, utilize exacting criteria when selecting images, and finally increase the visual literacy of the students in order to make good educational use of the image. It is thus timely to emphasize and reiterate the considerable value of utilizing images in science teaching. One major difference between expert and novice problem solvers is that experts possess the ability to form an image of a situation (Ambron, 1988), and firsthand experience with a situation in the field or laboratory is the ideal route to such visualization. Although the use of images cannot replace direct knowledge of specimens or phenomena gained in laboratory or field experiences, and may not have the same degree of potential to stimulate a lifetime interest in science, the cost and logistics of providing these experiences may prohibit their being made available to students, and learning from images has been shown to provide better understanding and retention of concepts than information delivered via lectures or reading materials. Woods (1989) cites data indicating that 90% of concepts encountered via real or simulated activities are recalled over the long term, compared to 30% of those encountered via images, and only 10% for those ideas delivered via lectures or reading materials. Thus, in the many instances when field experiences or hands-on activities cannot be delivered, using images to deliver instruction is distinctly advantageous.

Identification of Biological/Ecological 'Story Lines' in Ornithological Paintings.

One fruitful approach to the use of bird paintings in the classroom that employs both previous knowledge and critical response skills is to encourage students to construct meaning from an image containing multiple ecological variables and to frame this meaning in terms of known science concepts and in generation of topics and questions for further exploration. A good ornithological painting may not give up its treasures easily to the casual observer, and some work may be required on the part of viewer to mine the ecological concepts implied by the work, and to frame questions for further research. The careful and deliberate visual structuring of a fine narrative painting, implies that the artist has a very particular tale to tell his or her viewer, and with this in mind, I analyzed a sample of forty ornithological paintings (all of which met Tufte's criteria for excellence in scientific graphics) for the emergence of visual 'story lines' (Hunt, in prep: See Appendix 3; Table 3.1 for paintings and sources).

At least one ecological story line emerged from each of the paintings. These 'stories' are necessarily somewhat subjective, are frequently implicit, and are dependent upon some prior knowledge for elaboration. The stories that emerged from my particular perspective are listed in a brief form in Appendix 4 (Table 4.1). However, I feel it appropriate to discuss some of my own thinking about certain paintings to provide examples of such an approach to extracting themes from these images.

Even a simple canonical portrait, such as Eckelberry's White-eared Puffbird (See Appendix 3: Table 3.1 for source) tells a tale of ecomorphology and adaptation, and invites speculation about that species' classification, habitat, and distribution. Some paintings depicting multiple species, and/or those rich in behavioral interactions and habitat details contain within them the possibility of many biologically oriented stories. In such cases (e.g., Fuertes' Diurnal Birds of Prey in Flight; Pratt's Apapanes and Akepas; Weatherly's Campbell's Fairywren: See Appendix 3; Table 3.1 for sources), the three or four most obvious 'plots' have been listed. The majority of these are straightforward ornithological topics and ecological issues, however, paintings depicting recently discovered species, or rediscovery of birds previously believed to be extinct, also contain implicit stories of the nature of discovery in non-experimental branches of science (e.g., McQueen's Forest Owlet and Weatherly's Campbell's Fairywren; Audubon's Ivory-billed Woodpecker; Bewick's Great Auk. See Appendix 3: Table 3.1 for sources; Appendix 4: Table 4.1 for analysis of story line).

Surprisingly, a painting not intended as a narrative, but originally designed as a plate for a field guide to Ecuador—John O'Neill's Galapagos Finches (see Appendix 3: Table 3.1 for internet source; also published in Ecuador: The Ecotravelers' Wildlife Guide) offered the most extensive relational possibilities in this sample of paintings. Illustrating five related species of finches, and with very few habitat elements depicted, the painting explicitly depicts comparative ecomorphology, implies how biodiversity results from adaptive radiation as a result of natural selection, demonstrates niche partitioning as a result of competition, and exemplifies an approach to biology framed in terms of Darwinian principles. Implicit in the painting are notions of a common ecological community, the geography of the Galapagos islands, conservation issues resulting from global climate change and/or human impact on the environment, and the question of limiting factors to finch populations on the islands. The subtext of the development of Darwinian ideas also invites a consideration of the nature of science (Appendix 4: Table 4.1).

An unusual work in the ornithological narrative genre is Le Grande Martinet; Le Petit Martinet (Appendix 3: Table 3.1 for internet source) , by the 18th century French illustrator Francois Nicholas Martinet. Two species are portrayed (the Common Swift, Apus apus, and the Common House Martin, Delichon urbica) against a detailed architectural background. The urban environment depicted is absolutely relevant to the ecology of the bird subjects. These two primarily aerial species nest around human constructions, and as such, their population numbers are also linked to human populations, as the availability of nest sites is an important limiting factor to both species. This is one instance where the effect of urbanization upon a bird species is not deleterious to its survival: Martinet explicitly indicates how the Common Swift utilizes crevices for its nest sites, and implies by his placement of the House Martin that this species has a different preference (House Martins build a mud nest under eaves). A comparison of the morphology of the two species is visually enforced by the composition. The actual portraits explicitly describe the ecomorphological adaptations of the two species, showing the long wings and flight feathers geared to almost continuous flight, four toes pointing forward allowing gripping of vertical surfaces but not perching, and the tiny bill with a wide gape for trawling of fast-flying aerial insects. These explicit considerations of form and function invite a segue into the implicit themes of energetics and physiology. A simple 'google' search of the Common Swift, for example, will quickly reveal that the inability to perch forces parent birds to leave the nest for several days in the event they must skirt a tropical depression to which the young respond with a state of torpor, with considerably reduced respiration and body temperature. The adaptations to continuous flight also invite an exploration of long-distance migration (behavioral ecology) and the nesting sites at which the birds are depicted suggest suitable cavities for nesting as a limiting factor to populations (along with the availability of insects above 50 meters), competition for nest sites and the adverse effects on populations when urban centers undergo substantial modernization (human impact).

John Gould's Quetzal (see Appendix 3: Table 3.1 for internet source also immediately kindles consideration of limiting factors (which here is the availability of suitable trees with cavities for nesting); and also natural selection (via sexual selection and mate choice) and coevolution—a little research quickly informs the viewer that this frugivorous species is a major seed disperser of selected plants. The background also encourages speculation about the nature of the cloud forest habitat and related conservation issues. The foregoing models of some of the 'stories' emerging from relational phenomena that emerge after careful consideration of these three paintings typify an approach that might be used to mine the informational and educational potential of any image in the sample (Appendix 4: Table 4.1).

Other Uses of Bird Art in the Classroom: Beyond Conceptual Knowledge.

Paintings where concepts such as ecosystem, community, energy flow, interdependence of birds and plants, and predator-prey interactions are illustrated explicitly, are obviously suitable for incorporation into a lecture format, as examples of concepts described by the instructor. Paintings or plates where concepts are implied rather than explicitly illustrated, such as in carrying capacity, limiting factors, ecosystem fragility, and conservation issues, have great potential for use in inquiry based activities in which the student mines the concept following independent research into the species and/or habitat illustrated. This latter use of images is relatively easily implemented with access to the internet, where extensive and accurate information on birds and their habitats is widely available (more so than any other taxon due to the massive amount of data and observations reported by knowledgeable amateurs). A further use of such images lies in assessment, where the student might be asked to identify the visual representation of concepts previously taught in class (Wandersee, 2000).

The use of ornithological paintings in the classroom goes beyond illustration, reinforcement, and assessment of content knowledge, however. Ornithological images may serve as a valuable resource in conveying some of the factors of the history and nature of science. Discussing—or requiring students to research—'the story behind the painting' invokes many of the factors that have fashioned the progress of ornithology, such as the relative emphasis upon taxonomic or ecological aspects of the science at different times in history, the influence of the economic status of a nation upon the cultural demand for more ornithological knowledge, the effect of technology upon the visual representation of science, and the importance of funding for scientific exploration and the avenues through which funds have historically been obtained. Certain aspects of the sociology of science, particularly the means by which ornithological knowledge has grown through collaborations, and been disseminated among its practitioners through communities of practice is also nicely illustrated by an exploration of the lives of some of the great bird illustrators. Finally, a case study of these biographies by students, including those of contemporary illustrators, may serve to engage students with the notion of 'science as a human adventure' (AAAS, 1993), or an endeavor in which individuals with widely varied backgrounds and abilities may participate.

One essential element of scientific methodology that is frequently insufficiently addressed in many science courses is the role of observation and of reflection upon those observations. Accurate ornithological illustrations possess the potential to direct the viewer's attention to these processes. If a completed narrative painting or field plate of birds is considered as analogous to a scientific report, much understanding of these essential processes of science might be gained by subjecting the image to an analysis of the nature of the observations and thought processes that were necessary to its production, and thus emphasize to the student that engaged seeing is much more than mere 'looking.'

Understanding of the characteristics of scientific thinking is also emphasized by the American Association for the Advancement of Science (AAAS, 1993) and the National Research Council (NRC, 1996). As John Janovy (1996) wrote concerning an exhibition of George Sutton's Arctic and Mexican bird paintings that drew "shoulder to shoulder crowds "(p.34), images of organisms in landscapes serve "as a window through which (others) see the working of a scientist's mind" (p.34). Thus the disposition and characteristics necessary to produce exemplary works of ornithological illustration might be discussed with students, and should illuminate attributes of curiosity, persistence, honesty in evaluating and recording data, and the necessary skill of critical reflection upon that which is observed. Such an exercise also has potential to transfer to a wider appreciation of the ideas and data contained within other scientific images, and of the importance of visual communication skills in science.

Appendix 1: A Brief History of Ornithological Illustration and an Introduction to Selected Bird Artists

Bird painting and illustration is an enormous field, with a history spanning sixteen thousand years, from the oldest known depictions of waterfowl, cranes, eagles, owls and ravens on cave walls in southern France and northern Spain to the to tens of thousands of illustrated bird books in existence in the 21st century . The information presented here is necessarily limited to a description some of the major influences and themes that can be identified in the story of the development of bird art in the western hemisphere over the last 700 years. Potent influences in the unfolding of this tale include the maturing of scientific interest through time, how the growth of ornithological knowledge intertwined with the social, economic, and political forces shaping the activities of artists of different nations (Farber, 1997; Stresemann, 1975), changing technologies relevant to the reproduction of art, the different modes of employment and financial support available to painters of different nations and historical periods (Jackson, 1975, 1985, 1994, 2002; Pasquier & Farrand, 1991; Skipwith, 1979), and the general absence of formal training for serious ornithological painters, resulting in the informal system of mentoring that took shape in the 17th century and that continues today as a lesser known exemplar of a community of practice in a scientific discipline. Finally, bird art has always fulfilled one or more of three functions: the symbolic, the purely decorative, and the didactic or educational, and yet another subtext of the history of this field is the degree to which each of these functions has been dominant through time and across nations, and the extent to which individual painters have adopted or resisted the dominant genre.

The high points of ornithological illustration generally coincide with the vigor of scientific interest and activity at particular periods in given countries, although a dearth of scientific activity has still sometimes allowed for excellence in non-didactic bird painting. Fine examples are to be found in Chinese landscape and 'bird and flower' painting (which deviated from decorative art to form an independent genre after 800 AD), Egyptian art from 4000 BC to around 200 AD, and complex works by artists painting for the Mughal emperors in the 16th and 17th century (Miskin's The Raven addressing the assembled animals and Noah's Ark can be located on many websites, and are superb visual confections that blend accurate depictions and symbolism with a cartoonist's wit). Persian artists also had a tradition of placing birds and other animals in confections that illustrated fables or poetry, such as Habib Allah's Concourse of the Birds, one of four paintings illustrating the 13th century Persian poem The Language of the Birds, that shows the unlikely convening of a parakeet, crow, crane, grebe, heron, peacock, and stork, gathered to discuss the means of reaching paradise (see: The Conference of the Birds). Although primarily decorative or symbolic in nature, such paintings possess considerable scientific integrity in terms of both correct representation of individual birds, the natural co-occurrence of species, and a carefully selected and detailed environment as background, even though conventional scientific activity in those cultures was generally lacking.

The Origins of Scientific Ornithology and Natural History in the West

The origin of the systematic study of birds is intertwined with philosophy: the Greek philosopher Aristotle declared birds to be a suitable subject of study for the philosophic mind, a claim that was frequently reiterated as a justification for studying birds in the Renaissance, when zoology was making a tentative reappearance after the middle ages. Aristotle—a talented and astute naturalist and a careful observer—devoted volume nine of his 12 volume magnum opus Historia Animalium to birds. Much of this volume reported detailed and accurate first-hand observations, although Aristotle was unable to resist including some undocumented folklore and unsubstantiated beliefs, such as the stealing of milk from goats by nightjars, and bird hibernation. Aristotle's Historia disappeared after the disintegration of the Roman Empire, but was preserved at Thessalonica by Arab scholars, and reappeared translated from Arabic into Latin centuries later, contributing to the revival of scientific activity in the Renaissance.

During the Middle Ages of European history (5th to early 16th centuries), scientific advancement flagged in a culture dominated by the Christian church, and the notion that organisms were of any intrinsic interest likewise declined. As the idea of divine design dominated thinking, animals were perceived largely in terms of their religious and moral symbolism, and bird illustration was essentially limited to the decoration of prayer books and psalters. Although scientific activity was almost non-existent, the illustrators of these manuscripts produced some admirable bird portraits: that many of the birds depicted in these miniatures are easily identifiable testifies that their producers were gifted field observers.

In the late Middle Ages in Europe, depictions of birds for educational purposes—intended to convey visual information about the birds—first appeared, illustrations that intended to educate readers about ornithology and which largely appeared in books. The first of these was De arte venandi cum avibus by the colorful Emperor Frederick II, which attempted to document the natural history of birds based upon personal observation, free of legend and dogma, and also to refute inaccuracies in Aristotle's Historia animalium (which had by this time reappeared in a Latin translation, and was available to the intelligentsia of Europe). Frederick's illustrations were utilitarian rather than aesthetic, and show the diagnostic features of the readily identifiable birds, with some remarkable similarities to the modern field-guide. Succeeding copies of the book show the portrayals of birds actually becoming more accurate over time, indicating that ornithological awareness was progressing at a local level in Europe during the 13th and 14th centuries, and also that the artists employed by courts or the church to paint the copies were aware of this new information.

De arte venandi cum avibus was written in the mid-13th century, when distribution of new scientific information was poor to non-existent. Although twentieth century ornithologist Edwin Stresemann has called Frederick II "the first great ornithologist known to history" (Stresemann, 1975, p.9), copies of the book were so few, and communication between scientists so limited, that many of the great Renaissance writers on natural history—Belon, Gesner, and Aldrovandi—were not aware of the treatise' existence, and its potential contribution to ornithology was essentially lost.

The 16th century saw the publication of three major illustrated ornithological works in Europe, the production of which was facilitated by two innovations: the establishment of the universities in major European cities, which served as centers of intellectual activity, and the improvement in dissemination of information by relatively inexpensive books illustrated by woodcuts. The first of these works was Belon's L'histoire de la nature des oiseaux, avec leurs descriptions & Naïfs portaicts retirez du Naturel, published in Paris in 1955; the first book ever to commission ornithological drawings, and motivated by Belon's interest in comparative anatomy and taxonomy. The second was Gesner's enormously popular Qui est de avium natura (published in Zurich, also in 1955), followed by the extensively illustrated Ornithologiae hoc est de avibus historiae libri XII by Aldrovandi (Bologna, 1599–1603). All three works collectively represent the high water mark of ornithology and ornithological illustration in the Renaissance, and are notable not only for faithful renditions of the birds depicted, but for the inclusion of drawings of embryological and developmental details as an aid to understanding. Thus bird illustration was employed for the first time in the intentional development of scientific understanding, as well as in the transmission of information. It is worthy of note that all three bird books sold well (Gesner's volume in particular was a 16th century 'best-seller' and was reissued in new editions for 200 years) and that purchasers came from a wide-cross section of European society. These books were directed at a wider audience that the small scientific community: amateur naturalists were numerous in an age where some knowledge of the natural world was considered a basic component of education. Additionally, as the case today, people were curious about exotic species and it seems likely that illustrations helped make the books attractive to the less literate; Pasquier and Farrand (1991) estimate that a greater proportion of the literate population of Renaissance Europe owned bird books than is the case today.

The works of Belon, Gesner, and Aldrovandi described above dealt largely with birds native to Europe, including a smattering of descriptions of the few exotic non-European species that were known to the authors. A feature of the next century was the extensive knowledge of new species that was to inundate Europe when the age of the 'scientist as explorer' began. The 17th through the 19th centuries was a period of geographical exploration for ornithologists, facilitated by the frequent inclusion of an artist-naturalist on expeditions aimed at developing new trade routes, or simply voyages of exploration financed by countries looking to enhance prestige. For example, Georg Marcgraf explored Brazil during three separate expeditions (from 1638-43) sent out under the auspices of the Dutch East Indies Company; Mark Catesby (and later, John Abbott) studied the avifauna of the eastern United States; John William Lewin recorded the birds of Australia (then New Holland); Levaillant explored South Africa; and Napoleon sent naturalists to Egypt and Syria. Extensive expeditions left from Britain throughout the nineteenth century; in the early twentieth century the American Museum of Natural History sent Louis Agassiz Fuertes on many expeditions, and various North American universities allowed George Sutton to continue the tradition of exploration, collecting, and painting in the field in Mexico, Iceland, and the Arctic. Contemporary American bird painters continue to explore, collect, and depict the birds of Central and South America, and NOAA voyages to the Arctic and Antarctic frequently include an artist-naturalist (Hunt, in prep).

As taxonomy matured following the work of Willughby and Ray in the mid-17th century, and Linnaeus' system of classification was widely adopted after the publication of Systema Natura (1735-1758), natural historians became increasingly concerned with accurate description, detail, and evidence. French painters produced many of the best examples of 18th century bird art: Martinet, Reinhold, and Barraband, in particular (Jackson, 1994). These painters illustrated many volumes of works by Brisson, Buffon, Bonnaterre, and Levaillant; works that for the most part were motivated by scientific dissent over taxonomic issues (Farber, 1997; Ford, 1993; Stresemann, 1975).

Ornithological Illustration in 19th Century London

The greatest age of bird art—the late 19th century—essentially occurred as a cumulative result of the many specimens that were collected on expeditions, providing a massive collection of material from which bird artists could work, particularly in England (Pasquier & Farrand, 1991; Smith, 1962). A considerable number of private menageries existed in the second half of the 19th century, the park of the Zoological Society of London provided access to many birds. The collection of the British Museum included every known specimen of that time, and the increased availability of museum specimens eliminated guesswork and inaccuracies with respect to morphology and field markings. Opportunities to work from live specimens additionally increased both realism and the aesthetic interest of the work, as the character of the bird was portrayed along with its distinguishing morphological characteristics. Any discussion of nineteenth century ornithology and bird art would be incomplete without mention of John Gould (1804–1881). Gould was a self-educated British artist and naturalist adept at quick sketches capturing the character of a bird and essentials of a scene, although better known for organizing the production of illustrated bird books utilizing the many European bird artists working in London in 19th century. Originally trained as a gardener, he began his scientific career as a taxidermist for the London Zoological Society where Gould's ornithological skill was such that he rapidly earned the position of Curator of Birds for that institution, and then went on to self-publish his first book A Century of Birds from the Himalaya Mountains. This work was based upon skins in his collection, and illustrated by his wife, Elizabeth, based upon Gould's sketches and notes. The commercial success of the book spawned Gould's publishing empire: he produced ten more books by subscription, numbering ten monarchs among his subscribers. For succeeding volumes, Gould hired ornithologists and artists to travel with him as he toured the zoos and museums of Europe, and also undertook extensive fieldwork in remote regions, including tropical Asia, Africa, the New World, and Australia. Elizabeth traveled with them, illustrating birds from Gould's sketches and under his direction until she died at age 37. After her death, Gould recruited three European-trained bird artists to take her place: Henry Richter, Joseph Wolf, and William Hart. Edward Lear also contributed some work.

Gould's work was notable for its accuracy, and his publications (A Century of Birds from the Himalaya Mountains (1831); The Birds of Europe (1832–1837); A Monograph of the Ramphastidae or Family of Toucans (1833–1835); A Monograph of the Trogonidae, or Family of Trogons (1836-38); The Birds of Australia (1840–48); A Monograph of the Trochilidae or Family of Hummingbirds (1849–61); The Birds of Asia (1850–83); The Birds of Great Britain (1862–73); The Birds of New Guinea (1875–88); A Monograph of the Pittidae (1880–81)) illustrated more than 300 new species with descriptions. His early training in horticulture also gave him an eye for the attractive arrangement of plant material: illustrations in Gould books typically make an extensive use of visual hierarchy, utilizing a small area around the specimen to show details of the immediate habitat of the birds, which is then placed in a wider environment. Gould met Audubon when the latter visited London, and learned from him how to arrange a bird so that the main diagnostic features of its plumage were visible. Following Audubon's tutelage, Gould showed birds flying wherever possible, and included more chicks and nests than previous illustrators. These aesthetic improvements resulted in a corresponding increase in sales, and stimulated the growth of ornithology by magnifying public interest in birds.

Ornithological Illustration in North America

A movement in natural history that had considerable implications for ornithological illustration was the gradual move from the consideration of a 'bird as specimen' to that of 'bird as integral part of the environment', i.e., a move away from taxonomy toward a more ecologically centered ornithology. The first illustrator of the North American Avifauna, Mark Catesby (1682–1749) was one of the pioneers of this shift in emphasis. Catesby published his Natural History of the Bahamas, Florida, and the Carolinas following two collecting trips to the southeastern states (spending a total of 11 years in the field), and a further 17 years after his return to London, which were spent preparing his illustrations from field sketches and notes. Of the 220 plates in the volume, 109 depicted birds, many of which were shown in some behavioral display: calling, feeding, stretching a wing etc. Catesby's bird paintings are thus considerably more dynamic than most previous work, and also indicate a deliberate attempt to introduce some detail or cues of the animals' habitat, frequently including relevant plant material or prey items. Catesby's overall importance to ornithology and illustration is twofold: his work opened up the natural history of the American colonies, and marks the first consistent attempt to include aspects of the birds' ecology. Sixty years after Catesby's work was published, English naturalist John Abbott (1751–1841) made a permanent home in Georgia, and completed more than 5000 watercolors of natural history subjects including more than a thousand of birds, anticipating Audubon's work by a quarter of a century, and his later work included landscape elements.

The celebrated 'Father of American Ornithology', Alexander Wilson (1766–1813), published the illustrated American Ornithology in nine small volumes over six years (1808–1814). This work contains illustrations and descriptions of 262 North American species, of which 39 were entirely new species, and 23 more given the first descriptions that distinguished them from their European counterparts for which they had previously been wrongly named. Species named after Wilson include: Wilson's Storm Petrel, Wilson's Plover, Wilson's Phalarope, Wilson's Warbler, the Canada Warbler (Wilsonia Canadensis), and the Hooded Warbler (Wilsonia citrine).

Wilson, a working-class Scottish émigré who taught himself both bird identification and painting in his third decade, was a field ornithologist of a very high order. At the time of publication of American Ornithology, his illustrations were greatly admired. They are now considered 'stiff' and crude compared to Audubon's paintings, but most ornithologists consider Wilson's writing and observations to be orders of magnitude superior to those of Audubon. The two bird men met in Kentucky while Wilson was engaged in a solo promotional tour of his first volume, at a time when Audubon was managing a general store and birding and painting for pleasure. Wilson was by this time considered to be the leading authority of birds in North America, and attempted to sell the impecunious Audubon a subscription: this effort failed, but the two engaged in extensive conversation, and Audubon showed Wilson some local species. It is also clear from Audubon's own writings that exposure to Wilson's work contributed some inspiration and impetus to proceed with his own project, Birds of America (Audubon, 1986).

Audubon (1785–1851) likewise conducted extensive solitary travel throughout the southeastern states making sketches and descriptions for his Birds of America (1827–38). Like Wilson, Audubon was a colorful character and possessed a vivid and driven personality: the rise of American ornithology seems to be dominated by such rugged individuals, in contrast to the history of ornithology in Europe, where the development of the science was largely the product of social and economic forces. Audubon's Birds of America was published in five parts over nine years, and presented 435 species of birds (of the approximately 600 species known to be native to North America). The work was artistically magnificent, showing colorful and active birds in natural settings with aspects of behavior and habitat included, and was a major departure from the stiff renderings characteristic of the period. Audubon has been frequently criticized for the use of contorted and unnatural poses, however, his explicit goal was to depict plumage details that remain hidden in standard poses, and not the least of Audubon's contribution to the field was that he taught this technique to other bird painters he met in the course of his travels, including John Gould, William Swainson, and Prideaux John Selby. In spite of the 'scientific' motivation behind his use of such poses, Audubon exploited these artistically as a basis for the dramatic and sweeping compositions for which he is famous, and which mark him as the first distinctively American bird painter, able to capture both the intrinsic character of a species along with an almost visceral sense for its typical habitat.

As an scientist, Audubon's position is considerably less elevated; Streseman (1975) does not mention him as an ornithologist of importance. Five companion volumes of Ornithological Biography accompanied Birds of America, where Audubon provided detailed morphological, behavioral, and ecological descriptions of every bird that he painted. His real contribution to natural history and ornithology however, were his descriptions of many new species of birds and mammals, and his pioneer work on the use of banding as a method for tracking migration. His writing has been largely disregarded: although his descriptions are detailed and accurate, they lack sophistication and have been criticized as anthropomorphic. It is fairer to place Audubon's writings within the American tradition of nature writing, alongside the works of William Bartram, Henry David Thoreau, John Burroughs, John Muir, Ralph Waldo Emerson et al., than within the mainstream of scientific contributions of the 19th century.

The transition to 20th century bird illustration in America is marked by the works of Louis Agassiz Fuertes (1874–1827). Named by his engineer father for the great Harvard naturalist, in his undergraduate years at Cornell University, Fuertes painted some plates for field guides that at attracted the attention of leading ornithologists of late 19th century America. Three of these became powerful mentors obtaining for Fuertes both commissions and opportunities to participate in many expeditions, allowing him to explore and paint the bird life of the Yucatan, Caribbean, Columbia, Alaska, Ethiopia, and Siberia. The competing dictates of these influential mentors likely resulted in Fuertes' distinctive style, in which morphologically accurate portraits also convey the behavioral characteristics of the species while subtly suggesting a typical habitat/environment in which they are to be found. Elliott Coues—of the Smithsonian Institute and founder of the American Ornithologists Union—insisted Fuertes focus on accurate renditions of species. Frank Chapman—of the American Museum of Natural History and editor of the popular magazine Bird Lore—wanted ecological detail in Fuertes' paintings, and the classically-trained natural history artist Abbott Thayer promoted an approach to painting that utilized qualities of light to express the spirit of the subject. Fuertes responded to these varied demands by developing an ability to convey the character of a bird in very simple portrait: ornithologist and field guide editor Roger Tory Peterson has said of Fuertes:"to one who knows birds, there is more latent life in a Fuertes' bird, composed and at rest, than in a wildly animated Audubon bird" (Pasquier and Farrand, 1991).

Fuertes' illustrated a large number of books, where he made notable ornithological contributions derived from his extensive expedition work. In addition to his fine and prolific artistic output, Fuertes left another important legacy to bird painting by initiating a tradition of mentoring that has been ongoing, and continues—at least among American bird artists—today. Fuertes earned a reputation of being generous with his time and expertise, and either corresponded with or directly advised a number of younger bird artists, the most famous of which is George Miksch Sutton (the tutoring of Sutton in bird painting is described in the delightful To a Young Bird Artist (Sutton, 1979). George Sutton (1898–1982)had the distinction of being the first major bird artist to earn a doctorate in ornithology (from Cornell University, where he remained for 11 years after graduation as Curator of Birds), during and after which he participated in extensive fieldwork in Canada, the southern United States and Mexico, travels responsible for production of many exquisitely illustrated books.

Eventually becoming Professor of Zoology at the University of Oklahoma, Sutton paid his mentoring debt to Fuertes with the advising of John P. O'Neill, a renowned neotropical ornithologist of the present day and a leading bird painter. Already an accomplished birder and artist as an undergraduate, O'Neill came to Sutton's attention in the zoology program at Oklahoma, and was encouraged to participate in ornithological activities with Sutton's graduate students. When O'Neill's interest turned toward tropical birds, Sutton directed him to graduate study under George Lowery at the Louisiana State University Museum of Natural Science, which contains the largest collection of neotropical bird specimens in the world.

Nearly half a century later, O'Neill has made considerable contributions to tropical ornithology (having currently discovered 14 new species of tropical birds), and is also recognized by the art world. In addition to illustrating many tropical field guides to birds, O'Neill is an established gallery painter and has exhibited throughout the world, with paintings currently in permanent collections from Scotland to Beijing. During his long tenure at Louisiana State University, O'Neill has actively mentored many younger bird artists, including Hawaiian bird specialist Douglas H. Pratt, Daniel Lane, and children's book author Sophie Webb (Webb 2000; 2004). He continues to conduct ornithological expeditions to Peru, and to illustrate field guides, most recently Birds of Peru (Schulenberg et al., 2007), a culmination of 35 years ornithological research in that country by LSU scientists and associates from other institutions.

The painters mentioned above are a small subset of contributors to the vigorous contemporary production of ornithological art in the United States, where the scientific knowledge of the artists is at a zenith in the history of the genre. In addition, fine work continues to be produced by British artists (e.g. Martin Ridley and ornithologist Eustace Barnes), and Australian painters such as Ian Roberts, Richard Weatherley, and Lilian Marguerite Medland have made recent contributions to a three hundred year tradition of painting in the Antipodes (Olsen, 2001).

Toward the Future: Recent Trends in Ornithological Images.

In the twentieth century, a division arose between "environmental painters" such as Liljefors, Jacques, Thayer, and Clem, and those painters employed to illustrate bird guides for the scientific and lay communities (Pasquier & Farrand 1991; see also Hill, 1987; Jacques, 1983; Swanson, 1994: White, 1951). Publishers of modern American field guides have been inclined to omit backgrounds for reasons of economy, but many painters of the present day are beginning to concern themselves with depicting the ecological aspects of a bird's existence in their original paintings (Eckelberry, 1963; Hunt, in prep.), although landscape elements still rarely appear in field plates.

The increasing specialization of science affected bird illustration in the late 19th century, forcing the 'compleat naturalists' who had historically concerned themselves intermittently with bird study to narrow their focus and to metamorphose into ornithological specialists, and in bird illustration there has concomitantly been a movement back toward the depiction of "bird as specimen", particularly in field guides (Farber, 1997; Pasquier & Farrand, 1991). The emergence of scientific societies and the publication of specialized journals that resulted have both increased opportunities for bird artists while ultimately limiting their artistic expression, by emphasizing didactic elements of bird illustration at the expense of the aesthetic and symbolic aspects of bird painting (Devlin & Naismith 1974; Eckelberry, 1963; Pasquier & Farrand, 1991).

A few painters, notably Louis Agassiz Fuertes at the turn of the 20th century, have achieved a synthesis of styles. Bruno Liljefors, Francis Lee Jaques, and Robert Verity Clem are examples of 20th century artists who remained primarily painters of environments that necessarily included birds (Hill, 1987; Jaques, 1973; Pasquier & Farrand, 1991; White, 1951). In the first half of the 20th century, some painters—of which George Lodge is a fine example—adopted an either/or approach, working in the environmental tradition and also as a purely didactic museum artist (Savory, 1986). Among contemporary painters, there is a discernable correlation between the maturity of the artist and his or her inclination to move from a didactic style toward an environmental approach or at least, a synthetic compromise. Roger Tory Peterson—who set the tone of modern didactic bird painting with his introduction of the layman's field guides—became more painterly in his later years (Devlin & Naismith, 1977). Oregon painter Larry McQueen has announced his intention to do this quite overtly and John O'Neill is showing inclinations in the environmental and symbolic direction, at least in his non-academic work (Hunt, in prep.). Just as modern nature writers such as Aldo Leopold, Joseph Wood Krutch, Edward Abbey, Annie Dillard, and Gary Snyder have integrated Thoreauvian Romanticism and twentieth century biology into their work (McClintock,1994), the most recent wave of bird artists—such as James Coe, Eustace Barnes, and Sophie Webb—are similarly bringing this integration into painting (Hunt, in prep.). Such artists are incorporating exacting technical portrayals of species into a symbolic use of birds as a motif for conservation values, and in particular the intrinsic value and meaning of unmolested and unexploited landscape. This movement is in a relative infancy, but is gathering momentum in 21st century bird illustration (Hunt, in prep; Pasquier and Farrand, 1991).

Appendix 2: Bird Artist Names for Image Searches

The following list is not intended as an exhaustive list of bird painters whose works are suitable for use in the biology classroom. Many fine artists have escaped my attention. Inclusion of the names in this list simply implies that I have scrutinized individual works by these artists for scientific integrity and aesthetic merit.

Appendix 3: Ecological Concepts Depicted Or Referenced Within A Sample Of Bird Paintings

Content knowledge recommendations contained within the National Science Education Standards (NRC, 1996) and the Benchmarks for Scientific Literacy (AAAS, 1993), indicate that the teacher of ecology should address the following key topics:

  1. Environmental heterogeneity
  2. The Ecosystem
  3. Succession
  4. Population ecology
  5. The Community
  6. Life-history strategies
  7. Competition
  8. Predator-prey interactions
  9. Niche
  10. Ecological adaptation
  11. Food chains/webs
  12. Energy flow
  13. Limiting factors
  14. Carrying capacity
  15. Human impact upon the environment
  16. Ecosystem fragility
  17. Conservation of Resources
  18. Interdependencies between species
  19. Mechanisms and results of evolutionary processes
  20. Species diversity

As part of a larger study evaluating the usefulness of ornithological images, I have analyzed 40 narrative ornithological paintings and 40 field guide plates (the latter selected from four popular field guides to birds) in terms of inclusion of these 20 essential concepts plus a number of more specialized topics that may be depicted by this genre (Hunt, in prep). The narratives and field plates selected for analysis, and the number of each in which the concepts were either explicitly illustrated or implicitly referenced are tabulated below (Table 3.1, 3.2).

Appendix 4: Identification of Biological/Ecological 'Story Lines' in Ornithological Paintings.

Table 4.1. Explicit or Implied "Stories" Within a Sample Of 40 Narrative Ornithological Paintings.
Painting Biological/Ecological 'Story Line(s)' Detected
Audubon: Black-throated Guillemot, Nobbed-billed Auk, Curled-Crested Auk and Horned-billed Guillemot Adaptations of four species to the environment of the North Pacific coast; ecology and evolution of seabirds
Audubon: Ivory-billed Woodpecker Life-cycle of trees; interdependence of birds and food plants; extinction; nature of science—continual revision of beliefs in the face of new evidence
Bateman: Three Ostriches on an Egg Evolution of flightlessness and subsequent adaptations to this state implied
Bateman: The Return – Bald Eagle Implications of food shortages to the population success of these birds
Bewick: The Rook The success of this species in an agricultural environment; coevolution of human society and certain bird species
Bewick: Great Auk Adaptation of this species to northern coastal habitat; ecology and evolution of seabirds; extinction
Catesby: Passenger Pigeon Extinction; role ofhardwood forests in the story of the extinction of the Passenger Pigeon.
Catesby: Heron, eft, chigoe, cockroach… Feeding ecology of the Great Blue Heron; plasticity of diet in birds generally believed to be fish-eaters; community of organisms in freshwater habitats.
Coe: Western Rock Shorebirds Ecology of the rocky shore; reproductive ecology (sexual display and mate choice implied by seasonal plumages).
Coe: Black-bellied Plovers Ecology of the sandy beach habitat; reproductive ecology of the species (sexual display and mate choice implied by seasonal plumages).
Eckelberry: White-eared Puffbird Ecomorphology and taxonomy of species. Visual description of (then) new species; dissemination of information about new discoveries to the scientific community.
Eckelberry: Yellow-crowned Night Heron Ecology of mangrove swamp; ecomorphology and reproductive behavior (implied by paired adults) of species.
Fuertes: Diurnal Birds of Prey in Flight Role of top predators in a marsh habitat and food webs within the marsh, also implied are taxonomy of birds of prey, energetic issues relating to aerial predation, and evolution of flight
Fuertes: Rails Wetland ecology; taxonomy of Rails; reproductive ecology
Gould: Quetzal Ecology of cloud forest; reproductive ecology and behavior of the Quetzal; taxonomy of the Trogonidae.
Gould: Spotted Bowerbird Reproductive ecology and behavior; Australian birds; endemism.
Jonsson: Coastal Meadow Oystercatchers Coastal meadow ecology; reproductive ecology and behavior.
Jonsson: Atra (Desire) Marine coastal environment as habitat for birds; social behavior and flocking.
Keulemans: Pennula sandwichensis (Hawaiian Spotted Rail) Extinction; endemism; flightlessness and evolution of flight
Keulemans: Pseudonestor xanthophrys (Maui Parrotbill) Endemism; taxonomy and ecology of the Drepanidinae (Honeycreepers); conservation issues.
Lane: Boat-billed Heron Ecomorphology and taxonomy of species; mangrove ecology.
Lane: Scarlet-banded Barbets Ecomorphology and taxonomy of barbets (Family: Capitonidae); ecology of cloud forest; nature of discovery in non-experimental branches of science
Liljefors: Sea Eagle Chasing Eider Duck Predation; marine coastal environment as habitat for birds.
Liljefors: Bean Geese at Sunset Marsh ecology; social behavior and flocking; migratory behavior
Malick: High Prairie Predation; ecology of prairie; conservation issuesas an extension of the latter
Malick: Martial Eagle on Ground Hornbill Predation; African birds; ecology of desert savanna
Martinet: House Sparrows Coevolution of humans and birds; success of sparrows in urban environments
Martinet: Le Grande Martinet; Le Petit Martinet Energetics of flight; reproductive behavior and ecology of swifts and martins; effect of human activity on populations
McQueen: Anna's Hummingbird and Pacific Madrone Energetics and physiology of hummingbirds; interdependence of birds and food plants
McQueen: Forest Owlet Indian forest birds; nature of science – continual revision of beliefs in the face of new evidence; extinction and conservation issues
O'Neill: Galapagos Finches Natural selection; endemism; island biogeography, adaptive radiation
O'Neill: Black-throated Blue Warbler Reproductive ecology and behavior; life history from allocation of resources
Pratt: Brown Creeper Interdependence of plants and birds; life-cycle of trees; concept of micro-niche
Pratt: Apapanes and Akepas Endemism; island biogeography; adaptive radiation; conservation and extinction; interdependence of birds and food plants
Reinhold: Egyptian Vulture Ecomorphology and energetics of scavenging birds; competition as force driving evolution of scavenging behavior.
Reinhold: Fiscal Shrike Ecomorphology and taxonomy of species; predation and food chains; African birds
Thorburn: Gyr Falcon Energetics of large predatory birds; home range; polar ecology
Thorburn: Blue Tits on a Teasel Interdependence of plants and birds; concept of micro-niche
Weatherly: Carpenterian Grasswren Desert ecology; limiting factors; endemism
Weatherly: Campbell's Fairywren Taxonomy of Maluridae and nature of discovery in science; ecology of cloud forest; endemism