Neotropical Birds of Prey: Biology and Ecology of a Forest Raptor Community

1 THE MAYA PROJECT

David F. Whitacre and William A. Burnham

In 1988, biologists of The Peregrine Fund set out to do something unprecedented—to study the ecology of the entire raptor community at a lowland forest site in the American tropics. Here we present our results, providing the first ecological portrait of virtually an entire tropical forest raptor community, results of a nine-year field study at a single locale. This effort, denoted the Maya Project, was centered on Tikal National Park in Guatemala’s remote Petén Department, site of an ancient Maya city set within an immense area of lowland tropical forest. Over the nine years from 1988 through 1996, more than 100 field workers (up to 60 at a time) scoured the forests of Tikal to document the breeding biology, behavior, diet, space and habitat use, and ecological interactions of a Neotropical raptor community. We studied 20 of the 21 forest raptors common enough to study at Tikal. These species form the core species set of lowland tropical forest raptor assemblages throughout the New World tropical mainland (the Neotropics).¹ Here we present our results for the 20 species we studied, and summarize what we learned of their biology, ecology, and behavior.

WHY STUDY A NEOTROPICAL FOREST RAPTOR COMMUNITY?

There are many utilitarian reasons to study birds of prey, and those of tropical forests in particular. Predators, including raptors, play important ecological roles, sometimes affecting abundance patterns of their prey species, and can affect the structure of entire segments of biotic communities. They exert important selection pressures that shape characteristics of their prey from anatomy and coloration to habitat use, activity patterns, and social behavior. Predators are especially significant from the standpoint of conservation efforts. Often occurring at low densities and having large home ranges, they require large habitat areas. Thus, they can be a limiting factor in conservation efforts and can play an umbrella role—protecting sufficient habitat for predators can provide adequate habitat for many less space-demanding species. For us, the simple fascination of raptors was additional justification for undertaking the work described here.

Prior to the Maya Project, only a few New World tropical forest raptors had been the subject of detailed studies of ecology or breeding biology (e.g., Beebe 1950; Rettig 1978; Delannoy and Cruz 1988; Thiollay 1991b; Alvarez-Cordero 1996). A number of studies had considered entire Neotropical raptor communities at some level. Most notably, several studies by Jean-Marc Thiollay and colleagues (e.g., Thiollay 1984, 1985, 1989b; Jullien and Thiollay 1996) considered patterns of abundance in different localities and habitats. These studies provided portraits of community composition, relative abundance patterns, and habitat affinities, and also tested methods for detecting and counting tropical forest raptors and made some initial density estimates (Thiollay 1989a). The only work approaching a detailed community-level ecological study was that of Scott Robinson (1994), who presented information on the diet, hunting behavior, and habitat use of raptors in Manu National Park in Amazonian Peru.

Why did we elect to study an entire assemblage of Neotropical forest raptors? The predator assemblages of Neotropical forests—some of the planet’s most diverse—also are among the most poorly explored. In this biological terra incognita, the field naturalist may discover on an almost daily basis information new to science. Moreover, studying many species at a single locale provided insights not available from studying the same species at disparate sites, regarding community structure and function, comparative life history patterns, and comparative population densities and spatial use. To study the ecology of the entire raptor community at a single tropical forest locale was thus an irresistible prospect.

We also believed that such a project could yield benefits for conservation. Information we gathered on basic raptor ecology, habitat needs, population dynamics, and the effects of prevalent land uses might prove useful to land and wildlife managers and provide a basis for species conservation measures. We also felt that such a project would be a fertile training ground for local people who were or might become involved in conservation and management efforts in this globally significant forest area—a belief that proved well founded.

A PLETHORA OF PREDATORS

When we consider the epitome of terrestrial vertebrate predator communities, many of us think, no doubt, of the African plains, where the megafauna we associate with earlier geological epochs lives on today. There, a large mammal assemblage unparalleled elsewhere on Earth includes a diverse suite of mammalian carnivores. However, if one considers the total community of vertebrates that eat other vertebrates, among the planet’s most diverse are those of Neotropical forests (Table 1.1). Vertebrate predators include not only mammals of the order Carnivora (cats, dogs, weasels, and the like) but also birds and reptiles, mainly snakes. Indeed four groups—the Carnivora, Falconiformes (raptors), Strigiformes (owls), and snakes (suborder Serpentes)—compose the bulk of vertebrate predators in Neotropical forests as they do in other low-latitude terrestrial communities.

For many groups of organisms, including birds, among the planet’s most species-rich forests are those of the Amazonian headwaters, a crescent-shaped area along the eastern foothills of the Andes from southern Venezuela to northern Bolivia. For a vertebrate ecologist interested in tropical forest predators, to gain an understanding of the dynamics of predator communities in the upper Amazon is among the holiest of grails. This has not yet been attempted except, for the diurnal raptors, in the paper by Robinson (1994) mentioned earlier. Meanwhile, Harry Greene (1988) has written of the vertebrate predator community of La Selva, a lowland rain forest site in Costa Rica. Site of a renowned research station, La Selva is home to at least 56 species of snakes, up to 26 species of diurnal forest raptors, 6 owls, and 14 species of mammalian Carnivora (Table 1.1). As Greene points out, several of these feed mainly on invertebrates, and some on fruits. In addition, a number of species belonging to faunal groups not listed in Table 1.1 also prey partly on vertebrates at La Selva: these include at least two species of bats, a primate, an armadillo, a peccary, two species of lizards, three of frogs, several nonraptorial birds, and some large spiders, centipedes, and katydids (Greene 1988). A full tabulation of these other groups that prey on vertebrates would probably reveal an even steeper increase in predator species richness toward the equator than that suggested in Table 1.1.

In considering the ecology of tropical forest predator communities, it can be misleading to consider only one taxonomic group, because significant interactions such as food competition may occur even among distantly related groups. For example, a number of snakes feed mainly on lizards or other snakes, as do several raptors. Hence, raptors that specialize on snakes or lizards may overlap more in diet with these reptile-eating snakes than with other raptors.

Despite our interest in the overall vertebrate predator communities of Neotropical forests, it was beyond our means to extend our studies beyond raptors to other predator groups. Such studies to date have taken place elsewhere, including a study of the diet of several cats in Amazonian Peru (Emmons 1987) and that on snake diets at La Selva, Costa Rica (Greene 1988). Ultimately, it will be desirable to study the entire vertebrate predator community at a single Neotropical forest site—indeed, at many sites.²

RAPTORS OF THE MAYA FOREST

Including vultures, migrants, winter visitors, and open country and wetland species, as many as 50 raptor species occur in Petén, Guatemala, but the list of forest-dwelling raptors is much shorter (Table 1.2). Twenty-one raptor species made up the bulk of the non-vulture forest raptor community and were common enough to study at our project site, Tikal National Park (Table 1.2). We studied the nesting biology of all but one—the Guatemalan Screech Owl (Megascops guatemalae). Beyond these species, and disregarding the vultures, only a few other raptors can be confidently assumed to nest within Tikal National Park: the Short-tailed Hawk (Buteo brachyurus), Black-and-white Hawk-eagle (Spizaetus melanoleucus), and Central American Pygmy Owl (Glaucidium griseiceps)—all of which are uncommon there. Thus, the complete non-vulture forest raptor community at Tikal consists of 21 to 24 species that occur in ecologically significant numbers, 3 or 4 of them owls (Table 1.2).

If one includes a few forest species known from within 100 km of Tikal—the Harpy Eagle (Harpia harpyja), Spectacled Owl (Pulsatrix perspicillata), and Crested Owl (Lophostrix cristata)—the community expands to 27 species, 6 of them owls. Finally, if we include the Gray Hawk (Buteo nitidus) and Ridgway’s Pygmy Owl (Glaucidium ridgwayi)—which in this region occur in human-modified, partly wooded habitats but rarely in mature forest—the forest raptor community expands to 29 species, 7 of them owls. Although the Red-throated Caracara (Daptrius americanus) is not known from Petén, it occurs in lowland forests nearby, expanding the regional lowland forest raptor list to 30 species (Table 1.2).

Of great interest, the set of raptor species we studied at Tikal is largely the same set that occurs in the hyper-diverse equatorial forests of the Amazon Basin: in essence, we studied the widely occurring set of core Neotropical lowland forest raptor species. A few additional falconiform and owl species are added closer to the equator, but the species we studied at Tikal comprise the bulk of the raptor assemblage throughout mainland Neotropical lowland forests (Table 1.2). While the ecology of our study species may vary geographically, describing their ecology at one locality is a first step toward a more global understanding (Plate 1.1).

Table 1.1 Species richness in temperate and tropical predator faunasa

Table 1.2 Raptors of the Maya Forest

THE MAYA PROJECT

The Maya Project began in 1988, and fieldwork continued yearly through 1996. Two persons were responsible for the project’s inception: Dr. William A. Burnham (Plate 1.2) and J. Peter Jenny of The Peregrine Fund. During their efforts to study the Orange-breasted Falcon (Falco deiroleucus) in Central and South America, Bill and Pete were impressed by the scant knowledge of most Neotropical raptors and the lack of proven methods for detecting, enumerating, and studying these birds in their forest home. Perceiving a need for research on these topics, they began the Maya Project, leading eventually to this book.

An initial trip was made in 1988, in which the Maya archaeological sites of Tikal (Plate 1.3), Calakmul (in southern Campeche, Mexico), and Caracol (Belize) were visited. The decision to focus at Tikal was made for several reasons, key among them being the protected status of the park and the year-round access provided by what was then the only paved, all-weather road in thousands of square miles—that linked the park with the airport at Flores, the political and commercial hub of Petén. The following year was the first one of research during a full breeding season at Tikal. Each year thereafter, through 1996, a team of researchers worked from February through July or August, studying nesting raptors at Tikal. From 1991 to 1996, we also employed a small crew of field workers during the nonbreeding season to achieve year-round coverage, especially in tracking radio-tagged birds.

A key ingredient in the project was the participation of seven graduate students from the raptor biology program of Boise State University who appear here as chapter authors. Other key participants were three students from Evergreen State College. These individuals and other participating field biologists from the United States and Mexico are listed in the acknowledgments.

A crucial ingredient was the involvement of a number of local people in field research, some of them for the entire nine-year duration of the project. Many appear here as chapter coauthors, and all are listed in the acknowledgments. Many did not have previous formal training in biology but became highly skilled biological research technicians. Several moved beyond the technician level to become full-fledged field biologists in their own right. They not only led crews in field research but also extracted and summarized data, wrote progress reports that contributed to the chapters appearing here, and in several cases presented results at international scientific meetings. The accomplishments reported here resulted in large part from the efforts of these dedicated individuals.

The Peregrine Fund assisted several local field technicians in completing high school degrees and supported four of them in a technical degree program in the conservation and management of tropical forests at the Petén branch of Guatemala’s San Carlos University. Apart from such formal education, training within the Maya Project took place via an apprenticeship model. By working shoulder to shoulder during multiple years of fieldwork, experienced graduate students and professional biologists passed on to local participants the knowledge and skills needed to conduct field research on raptors, other birds, and vegetation.

Local participants, in turn, shared with U.S. project members their extensive knowledge of woodsmanship: how to extract water from a vine when lost in the forest, the names and uses of local plants, and the like. U.S. biologists learned a great deal from the local participants about land uses prevalent in the area: slash-and-burn farming, the chicle trade, the xate (pronounced shah-tay) palm industry, and hardwood logging. In this way, the U.S. biologists gained a perspective on these economic activities from the practitioner’s point of view—a perspective essential to any true understanding of the root causes of tropical deforestation and of the sorts of changes needed to achieve lasting conservation on the tropical forest frontier. Local project members also led U.S. biologists into two other endeavors: providing environmental education for 600 schoolchildren in local villages and introducing local farmers to nitrogen-fixing cover crops that could help them improve corn yields while reducing the amount of forest cleared for farming. These were rewarding aspects of the project that made a contribution toward conservation in the area.

The large amount of field research experience received by dozens of local people as part of the Maya Project is an important legacy of the project, as many of these individuals continue to employ this experience and training. Some of them now hold important positions with Guatemalan government agencies charged with managing the floral and faunal resources of Petén, while several others work for various non-governmental organizations, conducting ecological research in the forests of the Maya Biosphere Reserve.

Our research included work with migrant and resident songbirds and allies, examining abundance patterns in a variety of natural forest types and the effects of selective logging and shifting agriculture on the forest bird community (Madrid et al. 1995; Whitacre et al. 1993, 1995a). We also compared bat assemblages in extensive forest with those of forest fragments in the farming landscape (Schulze et al. 2000b). Studies of the woody vegetation of Tikal (Schulze and Whitacre 1999) supported our studies of raptors and other birds. The overall project is further described in Burnham et al. 1994, Whitacre 1998, and Whitacre and Burnham 2001.

FIELD METHODS

During the nine years of the project, we developed a suite of methods for studying raptors in continuous tropical forest. The most challenging task was finding nests. In general, a team of two to six people focused on a given species, searching for and observing the birds until their behavior revealed a nest. For the Forest Falcons (Micrastur spp.) this entailed stealthy observation on foot through the understory beginning well before dawn. For most other species, which were often active above the canopy, we would repeatedly ascend trees in areas where activity had been noted. Observing birds flying above or perched in the canopy, we would take compass bearings and move progressively closer, often watching from several different treetops over a period of days before the birds’ behavior revealed a nest. On occasion, once a general vicinity had been determined, we were able to search through the forest on foot, scanning likely trees until a nest was found. As we searched on foot, vocalizations of adults or juveniles sometimes revealed a nest locality. In a few cases, radio tags placed on adults we had trapped led us to a nest. Finally, nests were also brought to our attention by park guards and other local people knowing of our interests.

We made heavy use of radiotelemetry. This allowed determination of home range areas and habitat use and was helpful in determining the duration of post-fledging dependency and in maintaining contact with adults from year to year, in order to find subsequent nesting efforts or to verify non-nesting. Transmitters were in the 216 kHz range, and mostly made by Holohil of Carp, Ontario. They were nearly always 3% or less of the birds’ body weight. Most transmitters were mounted as backpacks, using Teflon ribbon joined via cotton thread over the breast bone, in the hope that the thread would eventually deteriorate and break, freeing the bird of the transmitter. In several cases transmitters eventually fell off the birds, while in other cases we do not know the transmitter’s eventual fate. Some transmitters were mounted on central tail feathers, and a few juveniles were fitted temporarily with tarsal-mounted transmitters, which were replaced by tail-mount or backpack arrangements once the bird reached near-adult size. Receivers were TRX-1000, made by Wildlife Materials of Carbondale, Illinois. We used hand-held, three-element, folding directional yagi antennas from the same supplier.

We conducted most telemetry on foot, with the objective of making visual contact with the bird or pinning down its location to one or two trees. We tied locations to known geographic points and our base map by taking a GPS (global positioning system) point (usually 20 or more, averaged) in the vicinity, or making a pace and compass map to a known point. When conducting such direct pursuit telemetry, it was often necessary to climb a Maya temple or tree in order to pick up a bird’s signal. On occasion, we resorted to triangulation to estimate locations. Through experience, we learned to discern and discard spurious bearings resulting from signal bounce. At times when we could not contact birds for a triangulation fix or could not find them on foot, they may sometimes have been at the peripheries of their home ranges, especially species with large home ranges, perhaps leading to conservative estimates of home range area. At other times, failure to obtain a radio signal was due to the bird being low in the forest or behind a hill, and such instances probably had little effect on home range estimates. When visual contacts were made, a data sheet was filled out, describing the habitat, perch characteristics, and the bird’s behavior. In some cases, we made concerted efforts to observe hunting behavior of radio-tagged individuals.

Most trapping of adult raptors was conducted at nests, using the bal-chatri (Plate 1.4; Berger and Mueller 1959)—a wire cage covered with monofilament nooses that snared the raptors’ toes when they attacked a live bait animal placed within. These traps were placed on nests (sometimes temporarily removing the egg or chick for safekeeping) or on limbs or the ground nearby. For hole-nesting species, we mostly used a net, hinged via stapling to the tree, which was pulled closed over the nest hole using monofilament lines after an adult entered. Other methods occasionally used were noose carpets (a variant of the bal-chatri, placed over a nest or perch), mist nets, and Swedish Goshawk trap (Thorstrom 1996).

Nest observations were made either from the ground or from a platform in a nearby tree, often concealed via camouflage netting or foliage. Platforms were made by lashing or nailing stout limbs in place, and usually employed a small piece of plywood as a floor, allowing use of a tripod and spotting scope (Plate 1.5). Observation platforms were built at sufficient distance that the birds’ behavior was unaffected.

We conducted training sessions to give observers experience identifying prey and estimating its size. We estimated prey length by comparison with the size of the raptor’s head, beak, and feet. To estimate mass, we practiced with tracings of scores of lizards that we captured, measured, and weighed. Prey mass was generally estimated using the following categories: less than 10 g, 11–20 g, 21–50 g, 51–100 g, 101–200 g, 201–400 g, 401–600 g, 601–1000 g, and more than 1000 g, in addition to point estimates of mass. We paid careful attention to the degree of reliability of prey species identifications; where there was any doubt as to species identity, we simply listed prey as snakes, lizards, vertebrates, and so on. Most species visually identified as prey at nests were also represented among prey remains collected at nests and identified in the hand, often by comparison with museum specimens or with the aid of specialists.

We concentrated our study in the central portion of Tikal National Park, where our field camp was located. We found nests in part by chance—having noted activity of a species in a certain area, we would search there for a nest. As the study progressed, we tried to find all the nests of a given species in a specific portion of the park, such that our study nests would be those of neighboring pairs, helping us estimate nesting density and degree of territoriality. In addition, we created a 20 km² study plot in continuous, mature forest, within which we made an effort to find nests and delineate territories. On this plot, we took advantage of survey lines cut by archaeologists years earlier. This yielded a 5 × 4 km study plot with a north-south access trail every 50 m in some areas and every 100 m in others. In 1994, we devoted a two-person team to count as many occupied raptor nests as possible within this 20 km² plot. This met with limited success, as a two-person team was not sufficient for this immense task.

ANALYTICAL METHODS

Because of the difficulty of detecting birds that did not defend a territory or engage in nesting behavior, we did not attempt to estimate the total population density of any species. Rather, we estimated the number of territorial pairs per unit area. We based these estimates on the spacing among neighboring pairs, indicated by active nests. Home range size estimates based on radiotelemetry did not lead directly to density estimates except where we obtained home range data for many neighboring individuals. Telemetry, however, aided in distinguishing neighboring pairs, especially when they were not nesting, and allowed estimation of home range sizes, which helped validate pair density estimates based on inter-nest distances.

Since our study nests were not generally in predetermined study plots that had been exhaustively searched for nests, we used plotless methods to estimate densities of territorial pairs, the basic datum being the average distance between neighboring nests. To determine this, we used the minimum spanning tree method (Gower and Ross 1969; Selas 1997); each of a group of n nests is joined by a straight line to its nearest neighbor, yielding n–1 inter-nest distances and a single mean inter-nest distance. We used this mean distance to create three complementary estimates of nest density.

The first, and preferred method, we refer to as the polygon method. Using CAMRIS GIS software (Ecological Consulting, Portland, Oregon), we created a convex polygon around the largest cluster of nests believed to be nearest neighbors. We extended the polygon beyond the outermost nests a distance equal to half the mean inter-nest distance. The outer edges of these circular nest buffers were joined to form a minimum convex polygon. The area of this polygon was then determined and divided by the number of known territories within, to yield the mean exclusive area per pair—the reciprocal of pair density.

The second method we use is the Maximum Packed Nest Density (MPND) method, as it sets an upper limit on the density possible under a given mean inter-nest distance. This method assumes that nests are spaced as in a close-packed crystal lattice, with all nests at the apices of equilateral triangles. This is unlikely to be strictly true, but it is a reasonable model for cases in which nests are overdispersed, that is, spaced more regularly than expected by chance—clearly the case for most raptors at Tikal. The area per nesting pair is given by πr2 (where r is half the mean inter-nest distance), adjusted by multiplying by 1.158 to account for the interstices among the abutting circles thus generated.

The third method, referred to as the square method, is that used by Newton (1979). In this case, the mean inter-nest distance is simply squared to give the estimated space per pair. This method assumes nests are regularly spaced at the intersections of a grid of squares, but not as closely spaced as under the MPND method. All of these methods give accurate results only for densities within the group of nests yielding the mean inter-nest distance. If nests are not similarly spaced over the entire landscape, density estimates can be extrapolated to larger areas only with caution or not at all.

In most raptor species, females are larger than males, a pattern referred to as reverse sexual size dimorphism because it runs contrary to the more general pattern in birds and mammals, in which males are the larger sex. Many researchers have attempted to explain this phenomenon, with little consensus. For each species, we present values based on various measures of body size. We used the Dimorphism Index (DI) developed by Storer (1966), which, for any given measurement, is:

We multiply the above by 100, to express the DI as a percentage. This index expresses the difference between the male and female measurements as a percentage of the mean value for the two sexes. This widely used index does not bear a linear relation to the simple ratio of female-to-male measures. As females become increasingly larger than males, the DI value increases, but at a diminishing rate. Hence, this index bears a downward-concave relationship to the percentage by which females exceed males in size. From the data presented, one may also calculate the percentage by which females exceed males in size. As an indication of overall size dimorphism, we present for each species the measures used by Snyder and Wiley (1976)—the DI value³ for (cube root) body mass, and the mean of DI values for wing chord (wing length from wrist to tip), culmen (exposed beak, from front of cere to tip), and (cube root) body mass.

ABOUT THE BOOK

To provide all of our principal results in one place, we summarize here all of our published and unpublished results. Nearly half the material included is published here for the first time. In the species account chapters, our main goal is to present our results from Tikal. In addition, we conducted a thorough literature review and present enough review material so that our chapters might serve as thorough synopses of the current knowledge on these species. While attempting to give uniform coverage to each topic for each species, we also allowed chapter authors to highlight certain aspects. Thus, for some chapters, additional headings are used. Chapters vary in length because we gathered more data on some species than others.

Table 1.3 Body size data for Maya Forest raptor species and temperate North American raptor species

To facilitate comparisons with species more familiar to many readers, Table 1.3 gives some basic size data for the 29 Maya Forest raptor species and for most temperate-zone North American raptors. Using this table, the reader can select a series of touchstone species to aid in visualizing the raptors treated here. Unless otherwise noted, species names of all raptors are consistent with the Global Raptor Information Network (www.globalraptors.org). Common names of non-raptors, animals, and plants are taken from the Encyclopedia of Life (www.eol.org).

So that this book might serve as a scholarly source, we present some statistical results, including probability values—"P values." P values state the probability that a pattern existed beyond that likely due to chance. By convention, a P value of 0.05 or smaller indicates a statistically significant pattern; at this value, there is only a 1 in 20 chance that the pattern observed was due to chance.

Because much of the material presented here has not been published in scientific journals, it was important to us that this book serve as an original, scholarly source. Thus we secured a similar degree of peer review to that achieved in the normal journal publication process. We sent all chapters to one or more reviewers prior to submitting them to Cornell University Press. In turn, the Press sent each chapter to at least two academic peer reviewers. We are grateful to these reviewers, and any shortcomings of the book are the sole responsibility of the chapter authors and the editor.

Chapter 1 Notes

1. Neotropics refers to the biogeographic region extending from the Tropic of Cancer in Mexico, the Caribbean islands, south to Tierra del Fuego. Only the lowlands of this region have tropical climates.

2. The most detailed study of a tropical forest food web—that of Luquillo, Puerto Rico—is of a species-poor island ecosystem with limited predator diversity, including only three diurnal raptors and one owl (Reagan and Waide 1996); thus, it sheds limited light on trophic dynamics within the much more species-rich forests of the Neotropical mainland.

3. In calculating the DI based on body mass, it is customary to use the cube root of body mass. Because volume and body mass scale as the cube of linear dimensions, basing the body mass DI on the cube root of body mass makes it comparable to DI values based on wing or tarsus length or other linear dimensions.

2 THE MAYA FOREST

David F. Whitacre and Mark D. Schulze

To best understand the natural history of the raptors we studied, it is helpful to know something of the ecological community of which they form a part. Here we describe our study area and its present-day biota and physical environment. We also present information on the climatic and vegetational history of Neotropical lowland forests and on the evolutionary history of raptors, to help reveal how long our study species have likely occurred together. We focus on the Maya Forest or Selva Maya region—the basal, moister portions of the Yucatán Peninsula—within the context of the entire peninsula, Central America, and the American tropics as a whole.

PALEOHISTORICAL CONTEXT

It is interesting to know how long our study species and their forest home have occupied our study site at Tikal, but it is far more important to ask how long these species have coexisted, whether at Tikal or elsewhere. The key question is whether these species have occurred together over long time spans, affording opportunities for evolutionary accommodations to coexistence. Today these raptors occur together over a very large area of the humid Neotropical lowlands (see Plate 1.1). How long has this been the case? The short answer is that tropical forests have been present in the Americas since prior to the origin of raptors. Moreover, most Neotropical raptor lineages appear to have long been present in the American tropics. Thus, it seems likely that the raptors we studied have coexisted in the American tropics for several million years, and possibly throughout the entire evolutionary histories of the genera to which they belong.

Tropical forests have existed in the Neotropics throughout the past 65 million years (Behrensmeyer et al. 1992), whereas modern bird families arose by 35 million years ago (mya) and modern genera, by 23 mya (Feduccia 1996). Hence, tropical forests have been available to Neotropical raptors for as long as raptors have existed. The antiquity of living raptor species is less certain. Recently it was believed that much of the speciation producing the stunning diversity of the Neotropical region took place during the late Pleistocene, the last million years or so. However, current evidence indicates that many Neotropical species originated earlier, more than 2 mya.

The earliest falconid fossils date from 23–36 mya, and a fossil from Nebraska 23 mya is attributed to the genus Falco (Olson 1985; del Hoyo et al. 1994), though fossils of present-day Falco species are from about 2 mya (del Hoyo et al. 1994). The earliest fossils of the Accipitridae, or true hawks and eagles, also date from 36 mya (Olson 1985). By 5–10 mya, a number of accipitrid fossils have been referred to living genera (Brodkorb 1964). Although the earliest-known owls date from 55 mya, the time of appearance of modern genera of strigid (typical) owls is uncertain (Olson 1985). The earliest recognizable tytonid (Barn Owl) is from France, about 20 mya, and represents an extinct genus. Three species of Tyto (modern Barn Owls) have been named from deposits 5–15 million years old in France and Italy.

Based on this information, we conclude that the genera we studied at Tikal likely differentiated by at least 10–15 mya. Many present-day raptor species are known from Pleistocene deposits of various ages (Brodkorb 1964), and we assume that our study species had speciated by early Pleistocene, at least 2 mya. In any event, most raptor diversity at Tikal is at the generic level and thus likely represents 10–15 million years since divergence of these lineages. Most raptor genera at Tikal are endemic to the Neotropics and probably evolved there. Thus it seems likely that the raptors we studied have coexisted in the American tropics for several million years and, in many cases, throughout their entire evolutionary histories.

Systematics and Biogeography of Diurnal Raptors

Falconidae. Seven of the 10 falconid genera and two of the family’s three major clades are restricted to the Neotropics, with only Falco and the falconets Microhierax and Polihierax occurring in the Old World (Becker 1987; Kemp and Crowe 1990; Griffiths 1994a, 1994b). The Neotropics are likely the family’s center of origin and certainly the area where most higher-level diversity is concentrated, largely in forest habitats (Griffiths 1994a, 1994b).

The contribution of falconids to raptor faunas thus varies greatly in different parts of the globe and may affect evolutionary and ecological patterns seen in different tropical raptor communities. The falconid genera Micrastur, Herpetotheres, Daptrius, and Ictyber, all New World forest dwellers, present a unique set of traits, and if these birds have ecological counterparts among falconiforms in other tropical realms, those will inevitably be of accipitrid rather than falconid stock. Among other things, this means that they will be stick nesters rather than cavity nesters.

Accipitridae. Phylogenetic relationships within the Accipitridae remain uncertain. Sibley and Ahlquist (1990) review attempts to classify this large and varied group. Brown and Amadon (1968) describe a number of traditionally recognized, informal groupings within the family, and two cladistic analyses have been based largely on skeletal characters (Kemp and Crowe 1990; Holdaway 1994). Ongoing studies based on molecular genetics are revising our understanding of phylogenetic relationships among Neotropical accipitrids and demonstrate that some genera, as currently defined, contain species groups that are not monophyletic (e.g., Riesing et al. 2003; Helbig et al. 2005; Lerner and Mindell 2005; Raposo do Amaral et al. 2006), which will require redefinition of these genera.

Most generic diversity within the Accipitridae occurs in the tropics, which are populated also by genera widespread in the north temperate zones (Kemp and Crowe 1990). There is evidence of notable adaptive radiations in South America and Africa, while other areas resemble these two regions in proportion to their distance from them, and reflect the relative vagility of different taxa (Holdaway 1994). Many genera are endemic to tropical regions, and few to temperate zone regions.

Geologic and Biotic History of Central America

Central America as we know it has been recognizable for only the past few million years. Once united as portions of the supercontinent Pangea, North and South America began to rift apart 140 mya, and by 80 mya, South America had separated from Africa as well, beginning a long isolation that produced a strikingly unique South American biota (Coates 1997). While the landmass comprising most of Mexico has been above sea level for at least 140 million years, portions of Central America are mostly younger. By 65 mya, Mexico was present as a peninsula protruding from North America, and the beginnings of southern Central America came into existence as an arc of volcanic islands. By 12 mya, this volcanic chain collided with South America, and the deep ocean floor that had preserved South America’s isolation for so long lifted to form a floor only 1000 m deep (Coates 1997). Over the next few million years an archipelago formed here. By 6–8 mya, the ocean floor was only about 150 m deep, and faunal interchange between North and South America became evident, as raccoons spread southward and ground sloths northward, apparently by hopping islands. By 2.4–3.0 mya, the Central American isthmus was complete, allowing faunal interchange over a continuous landmass. The ensuing interchange of the vastly different faunas of North and South America—the Great American Interchange—ranks among the most striking biogeographic events known (Stehli and Webb 1985).

2.1 Vegetation of the Yucatán Peninsula.

Physiography and Geology of the Maya Forest Region

The Yucatán Peninsula is a gently rolling limestone platform 500 km long and 300 km wide, protruding only modestly above the sea from which it continues to emerge (Fig. 2.1). The base of the peninsula has risen from the sea more than the northern tip, commonly achieving elevations of 300–350 m in and near Tikal. Lee (1980) concludes that the Yucatán Peninsula began emerging from the sea during the Miocene (5–23 mya), from south to north such that deposits in the north are progressively younger. While the northernmost tip of the peninsula did not emerge from the sea until the last million years or so, the basal portion around Tikal has been above sea level for perhaps some 10 million years (Lee 1980).

FORESTS OF THE SELVA MAYA FROM ICE AGE TO PRESENT

Throughout the past 2 million years, global climate has fluctuated dramatically along with the wax and wane of great ice sheets in the Northern Hemisphere. No great ice coverage was experienced in Central America, except atop the tallest mountains such as Guatemala’s Cuchamatanes massif and the Talamanca range in southern Costa Rica. While a Pleistocene vegetation map for Central America is not yet possible, much has been learned from lake-bottom pollen cores, phytoliths (silica deposits laid down by some plants within their cells), and radiocarbon dating of associated organic matter. The best data are available for the past 19,000–33,000 years, but some inferences may be made back to 120,000 years, and a tentative scenario painted back to the closing of the sea portal 2.4 mya (Colinvaux 1997).

Pollen cores from two lakes a few kilometers south of Tikal suggest that forest was absent here 10,000 years ago, with a more open, arid-adapted vegetation apparently present (Leyden 1984). For a brief period, perhaps 8,000–10,000 years ago, forests here were largely of pine, oak, juniper, and temperate hardwoods, suggesting a cool interval before tropical forests similar to today’s were most recently established. Hence, tropical forests were apparently reduced or absent in some portions of the isthmus at times during the ice age. However, abundant evidence from the last glacial maximum (19,000 years ago) to the present suggests that much of Central America was more or less continuously covered with forest, though of varying composition and distribution, throughout this period (Colinvaux 1997).

Lake-bottom cores indicate that the Panamanian lowlands remained primarily tropical forest from sea to sea throughout the past 120,000 years, embracing the Wisconsin glacial period and part of the previous interglacial (Colinvaux 1997). During cooler periods, plant species today restricted to montane forest migrated downward approximately 800 m, mixing with lowland plants. A cooling of 5–6 °C in Panama, and perhaps as much as 8 °C in our study area, seems well established. Such cooling was probably the dominant glacial influence on climate and vegetation of Central America prior to the advent of the Holocene warm interval 10,000 years ago. Locally, however, low rainfall at times is believed to have been an important influence, as in our study area.

In the vicinity of Tikal, during glacial advances of the past 2 million years, tropical forest may have been replaced by savannas, with the most recent round of tropical forest expansion beginning a mere 10,000 years ago. Reclamation of the Petén by tropical forest likely represented forest expansion from moist enclaves along the foot of neighboring mountain ranges of Chiapas, Alta Verapaz, and Quiché. Thus, while the raptor community we studied may not have co-occurred most recently at the precise site of Tikal until approximately 10,000 years ago, there is reason to believe these birds have co-occurred nearby in Central America throughout the past 2 million years or more, and for much longer in South America.

The present-day forest stands of our study site are likely younger than the 10,000 years just given, because of deforestation by the Mayan during their cultural apogee. Arising in this area more than 3000 years before present (BP), the Maya civilization flourished for nearly two millennia, then collapsed mysteriously during the tenth century AD (Curtis et al. 1996). Population densities during the Maya apogee are believed to have been 200–300 individuals/km² (Rice and Rice 1990)—much higher than today—and the population of the Yucatán Peninsula may have numbered several million (Curtis et al. 1998). During the early Holocene (approximately 9000–6800 years BP), evidence indicates moist conditions and semi-deciduous lowland forest in the area. By 5780 years BP, decline of lowland forest taxa in pollen cores may indicate early human disturbance, and substantial deforestation by the Mayan by 2800 years BP appears generally accepted among archaeologists, with widespread deforestation during the Classic Period (AD 250–850; Curtis et al. 1996). In many areas a layer of Maya clay as thick as 7 m was deposited in lakes over the 2600-year period leading up to European contact; this clay is believed to reflect soil erosion resulting from widespread Maya farming (Curtis et al. 1996).

Although the degree of deforestation engendered by Maya agriculture is not well known, we consider it reasonable to assume relatively great deforestation and forest modification. We assume also that forest refugia probably remained in areas not suitable for farming, for example, on rugged karst uplands and perhaps in the extensive wetlands of northwestern Petén. Unoccupied forest buffers between competing Maya polities and forest gardens within occupied landscapes may also have served as refugia for forest species (Hayashida 2005). Multiple lines of evidence indicate forest recovery beginning 1000–1100 years ago, soon after the Maya collapse. Evidence also indicates that different population centers had different dates of prominence and collapse, suggesting that dates of deforestation and recovery may have varied across the region (ibid.). Postclassic Maya populations continued to occupy the region until European conquest in 1597, but never regained the numbers existing prior to the collapse. Pollen evidence indicates that agriculture occupied much less land during the Postclassic Period, with longer fallows in swidden plots allowing establishment of secondary forests (ibid.). Hence we assume that the forests of Tikal and much of northern Petén were but lightly influenced by people from the time of the Maya collapse in AD 900 until the past couple of centuries, when inroads were made in search of Honduras Mahogany (Swietenia macrophylla) and chicle harvested from the Chicozapote (Manilkara zapota) tree (Schwartz 1990). Even these latter activities had little effect on forest cover until 1965 or later.

Are the forests of our study area, representing 1000 years of development since presumably massive alteration by the Maya, mature, primary vegetation, or are they successional in nature? Hatshorn (1980) concluded that 1000 years since abandonment of Maya crop fields is ample time for succession to have produced forests of climax status. Based on our own vegetation studies (Schulze and Whitacre 1999), we agree. In most of Tikal’s forest, tree species that dominate the canopy are also abundant in the understory and appear to be regenerating successfully, suggesting stable species composition rather than continuing successional change. The legacy of Maya occupation and forest clearing must still be evident in regional patterns of species composition and even landscape forms—some species having achieved greater densities in response to ancient anthropogenic disturbance, and soil and topographic conditions having been modified by Maya works. However, current forest composition and species distribution patterns in Tikal likely reflect more recent disturbance processes, such as individual treefalls and larger-scale storm blowdowns, acting in ways both stochastic and predictable on a landscape of varying physical conditions.

CLIMATE OF NORTHERN PETÉN

Located at latitude 17° North, Tikal is well south of the Tropic of Cancer. Holdridge (1967) considered those portions of Central America north of 12–13° to be in the subtropical realm, corresponding to the outer tropics of geographers. While the biota is distinctly tropical, the Selva Maya experiences a seasonal climate, especially in terms of rainfall. The mean daily temperature at Tikal changes little through the year, ranging from the mid-70s to 80 °F (23–27 °C: Fig. 2.2). Average monthly maxima range from a low of 82 °F (28 °C) in December to 94 °F (nearly 35 °C) in May, with the hottest days of the year commonly falling from March–June, during the dry season and early rainy season. Temperatures drop at night, often into the 50s–60s °F (11–20 °C). The daily range of temperatures is greatest during the dry season, from February through May, with distinctly less variation during the rainy season (Fig. 2.2).

Rainfall in the Selva Maya is far more seasonal than temperature (Fig. 2.3). A dry season holds sway from roughly January through April or May. This is generally a severe dry season of reliably rainless, often windy days. It is not rare for a month or more to pass without measurable rain, though thick morning fog is frequent, sometimes depositing dew. Leaf litter becomes crunchy underfoot, and understory vegetation often takes on a wilted appearance. Onset of the rainy season is variable; on average, the rains begin gradually in April and May, building to an initial peak in June. Thereafter, rains generally decrease during July and August, with a dramatic peak in September, usually the rainiest month. Rainfall drops off but is still high in October, and declines steadily in November and December. By January or February the dry season is again underway.

Rainfall maps for the Yucatán Peninsula vary greatly; Figure 2.4 presents the one we consider most credible. Rainfall decreases markedly from south to north across the peninsula. In southern Belize, on the windward slopes of the Maya Mountains, rainfall reportedly may average 4 m annually, while in the highlands of Alta Verapaz along Petén’s southern border, up to 5 m/yr is reported. Much of southern Petén appears to receive approximately 2 m annually. Moving northward, rainfall decreases to a mean of about 1480 mm at Flores (23 years of data) and 1360 mm at Tikal (9 years of data) and thence to about 1300 mm across much of southern Campeche and Quintana Roo, and to 450 mm in the extreme northwest of the peninsula, near Mérida, Yucatán. A lens of relatively high rainfall extends northward along the Caribbean coast of Quintana Roo, producing a marked gradient of decreasing moisture from east to west across the northern portions of the peninsula (López Ornat et al. 1989; see Fig. 2.4).

2.2 Annual pattern of temperatures in Tikal National Park, Guatemala.

2.3 Mean monthly rainfall throughout the year in Tikal National Park.

Mean yearly and monthly rainfall totals are deceptive, however, as rainfall in the Maya Forest varies strongly in quantity and timing from year to year. Reliability of total rainfall is substantially greater during the latter half of the rainy season, from August to November, than during the first half, from May or June to July (Table 2.1). Rainfall is quite unpredictable during the dry season and transitional periods, and variability between years is also high. Based on nine years of data (1960–62 and 1989–94), mean annual rainfall at Tikal was 1360 mm (52 inches). However, the annual total ranged from 1019 mm (40 inches) in 1992 to 1731 mm (68 inches) in 1993, with a coefficient of variation of 18%.

Climatic Variability: A Long-Term View

A detailed climatic reconstruction for the Yucatán Peninsula for the past 3500 years showed not only pronounced rainfall variation on a multi-decadal scale, but also a pronounced dry period from 1785 to 930 years BP, with several extreme droughts occurring during this and other time spans (Curtis et al. 1996). The same study showed correlations between rainfall variations and transitional periods in Maya culture, and the researchers favored the interpretation that drought caused the collapse of Classic Maya civilization 1000 years ago (Curtis et al. 1996).

During our nine years of fieldwork at Tikal and periodic visits since, we witnessed some effects of the variable rainfall regime. During a drought in the mid-1990s, many aguadas (water holes) that are normally reliable throughout the year dried up. Since virtually all households rely on such natural or man-made rainwater receptacles, this drought resulted in occupants vacating entire villages for a time. During dry years, agricultural fire often escapes and passes through primary forest. Ground fires burned beneath the canopy in two areas of Tikal National Park in the dry season of 1993, providing a tangible example of the interaction between climate variability and forest disturbance. An extreme example was seen in 1998, when drought conditions credited to El Niño led to large forest fires in Mexico and Central America, with resultant smoke affecting air traffic as far north as Texas. Drought frequently leads to partial failure of the corn crop on which many families depend. Finally, two massive insect outbreaks we witnessed in Tikal’s forests during the 1995 rainy season may have been causally related to a preceding seven-month drought period. Larvae of the dynastine beetle (Enema endemion) removed most leaf litter from large areas of forest, and an unidentified larva, probably a lepidopteran, caused extensive defoliation of Pouteria reticulata, one of the most common trees at Tikal (D. Whitacre and M. D. Schulze, unpubl. data).

2.4 A rainfall map of the Yucatán Peninsula.

Table 2.1 Variability of monthly rainfall at Tikala

Biological Seasonality of the Selva Maya

The longest day of the year at Tikal is about 12.95 hours from official dawn to dusk, and the shortest day, about 11.08 hours. Hence the longest day is about 16.9% longer than the shortest.

Arthropods. The abundance of many insect groups at Tikal fluctuates strongly with the march of the seasons. From September 1994 to June 1997, we sampled insect abundance weekly by shining an ultraviolet light against a suspended white sheet at our field camp in the forest near the park center. We operated the light for 90 minutes at a standardized time after sunset and, using a set protocol, counted the insects of different taxonomic orders and size categories that were attracted. From a distinct population low from November through March, insect numbers increased from April to a maximum in July, about a month after the initial rainfall peak, and decreased rapidly thereafter (Fig. 2.5). Other observations confirmed a dramatic peak in abundance of some insect groups early in the rainy season. At this time, overhead lights near park facilities often attracted many insects, including large beetles and moths, and the first rains invariably provoked mating flights of winged termites and ants.

The seasonal pattern of insect abundance depicted in Figure 2.5 is typical of Central American lowland tropical forests. Virtually all studies of arthropod populations in Neotropical forests with distinct wet and dry seasons have shown that overall insect abundance or biomass is greater during at least some portion of the wet season than during the dry season (Janzen 1973b, 1973c; Wolda 1978, 1982; Gradwohl and Greenberg 1982; Smythe 1982; Tanaka and Tanaka 1982; Levings and Windsor 1985; Pearson and Derr 1986; Boinski and Fowler 1989; Poulin et al. 1992). In most studies, arthropod abundance reaches a strong annual peak soon after the onset of rains, dropping during the latter half of the rainy season to lower, often dry season levels (Gradwohl and Greenberg 1982; Smythe 1982; Tanaka and Tanaka 1982; Levings and Windsor 1985; Pearson and Derr 1986; Boinski and Fowler 1989; Poulin et al. 1992), and in some cases, mid to late rainy season populations appear to be the lowest of the year (Boinski and Fowler 1989). The multiple by which arthropod abundance or biomass during the wet season exceeds that during the dry season usually ranges between 2 and 8, but multiples as high as 2–12 (numbers) and 20–50 (biomass) were noted in a western Mexican dry forest (Lister and Aguayo 1992). A multiple of 2–4 seems most common (see summary in Lister and Aguayo 1992).

Different sampling methods at a given site often yield differences in seasonality estimates, because they selectively sample insects with divergent behavior, and because different insects occupy different microenvironments (Boinski and Fowler 1989). Insect taxa often diverge in seasonal demographic patterns within orders, families, and genera (Wolda 1978, 1982; Gradwohl and Greenberg 1982; Smythe 1982; Tanaka and Tanaka 1982; Levings and Windsor 1985; Boinski and Fowler 1989).

Seasonality of insect populations may be driven in large part by the appearance of new leaves. A flush of new leaves at the onset of the rainy season takes place at Tikal, as in other seasonal tropical forests (e.g., Wolda 1978, 1982).¹ New foliage—which is often relatively unprotected (chemically and structurally) and high in usable nutrients—often provides food for herbivorous insects (Feeny 1970; Rockwood 1974; Coley 1983; Coley and Barone 1996), and in many cases, seasonal peaks of arthropod populations seem clearly tied to episodes of new leafing (Janzen and Schoener 1968; Fogden 1972; Wolda 1978). Insect population peaks may be associated with each major peak of leafing (Boinski and Fowler 1989), often after a 2–3-week lag (Dunham 1978; Tanaka and Tanaka 1982). While the advent of rains after an intense dry season provides new vegetation as insect food, torrential rains may be partly responsible for the common decline in insect abundance during mid to late rainy season (Boinski and Fowler 1989).

Some insect groups, however, do not find the dry season inhospitable. Owing to abundant flowering of woody plants during the dry season, Janzen (1973b, 1973c) believed that solitary bees peaked in abundance at that time. Moreover, with fruit and seed production greatest during the dry season, seed predators, including certain true bugs and beetles, reached peak abundance then (ibid.). Arthropods occupying dead, hanging leaves had relatively consistent abundance throughout the year (Boinski and Fowler 1989). Among 13 major groups of leaf litter arthropods, 9 increased in abundance during the rainy season, 2 increased during the dry season, and 2 showed irregular patterns not directly linked to season (Levings and Windsor 1985).²

2.5 Seasonal patterns of insect abundance in Tikal National Park, as sampled via ultraviolet light.

For insectivorous vertebrates at Tikal, the net result of the patterns just described is a peak in prey abundance early in the rainy season, including many of the large insects that commonly serve as prey of certain raptors. This seasonal peak in arthropod abundance may affect the timing of raptor reproduction directly in the case of raptors that feed on arthropods, or indirectly by affecting the seasonal patterns of growth, reproduction, and activity of the many insectivorous vertebrates that are in turn consumed by raptors: lizards, frogs, snakes, many birds and bats, and perhaps rodents.

Birds. Skutch (1950) reviewed the nesting seasons of Central American birds. Adapted from his paper, Figure 2.6 shows that most small land birds in this region nest at the onset of the rainy season. Egg laying, peaking in April and May, slightly anticipates the initial peak of the rainy season, such that hatching and chick rearing take place during this initial rainfall peak. Our observations at Tikal agreed; most small birds here nested mainly at the outset of the rainy season. This resulted in a pulse of availability of nestling and fledgling birds in early to mid rainy season for those raptors taking such prey.

Reptiles and amphibians. In the Maya Forest, essentially all the anurans (frogs and toads) breed early in the rainy season, when surface water becomes common after months of drought (Appendix 2.1). At this time, the din of frog calls around water bodies can be deafening. Duration of egg and tadpole stages in anurans is brief, and species breeding early in the rainy season gives rise to froglets in a matter of weeks. For most snake and lizard species in our study area, hatching or birth also takes place mainly during the rainy season (Appendix 2.1). Since gestation and incubation in reptiles often require 2–3 months or longer (Zug 1993), to achieve such timing, egg laying occurs early in the rainy season or late in the preceding dry season (Appendix 2.1).³

2.6 Seasonality of bird nesting in relation to rainfall at a Central American site (adapted from Skutch 1950).

Due to seasonal reproduction and growth of young, reptile and amphibian populations typically show an annual cycle of population size, age structure, and body size. Three Anolis species studied in Panama (Sexton et al. 1971) appear representative of many members of this genus (e.g., Andrews and Rand 1982; Andrews 1983; Campbell et al. 1989). Since most reproduction occurred during the rainy season, early in this season populations were composed mainly of large adults and tiny hatchlings. Through the course of the wet season, large individuals were progressively lost from the population, and hatchlings grew in size and number; by late rainy season, fewer hatchlings appeared and many young lizards were of medium size (Plate 2.1). During the dry season there was little or no addition of new hatchlings, and young lizards continued to grow (Plate 2.2) (Sexton et al. 1971).⁴

With regard to production of new raptor prey, timing of breeding in reptiles and amphibians has different implications than that in mammals and birds. Since birds, bats, and most rodents are nearly adult size when they become independent, the breeding season produces a pulse of new raptor prey. In contrast, most neonatal reptiles and amphibians are tiny and may require months or years to reach adult size. As newborns, these species are generally too small to serve as prey for most raptors. Hence, the breeding season in amphibians and reptiles is not likely to produce an immediate pulse of raptor prey.

Activity patterns, however, probably make many reptiles and amphibians more available to raptors during the rainy season. These creatures are often so cryptic while motionless that activity must greatly enhance their detectability by raptors. Frogs and toads are clearly most active during the rainy season, and tropical lizards and snakes may also reach peak activity then (Duellman 1965; D. Whitacre, pers. observ.). In fact, it is possible that seasonal differences in activity levels of reptiles and amphibians, as well as annual rhythms of age structure and body size, may influence their availability to raptors more strongly than do seasonal changes in abundance resulting from reproductive pulses.

One study of seasonal patterns of snake activity should be especially applicable to Tikal, as it was conducted in nearby Belize. In this study, snake detections were rare during the dry season and far more frequent during the wet season,