VENUES:

Museum of Arts & Sciences, Daytona Beach, FL
Chicago Academy of Sciences, Chicago, IL

Kansas Wetland Education Center Great Bend, KS
George A Spiva Center for the Arts; Joplin, MO
Houston Museum of Natural Science; Sugar Land, TX
Sternberg Museum of Natural History; Hays, KS
Mari Sandoz High Plains Heritage Center; Chadron, NE
Kenosha Public Museum; Kenosha, WI
Hiram Blauvelt Art Museum; Oradell, NJ
Arizona-Sonora Desert Museum; Tucson, AZ
The Wildling Art Museum; Santa Barbara, CA
The Wildlife Experience Museum;  Denver, CO

COMPLETE TOUR ITINERARY HERE
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Below are images and captions for the fifty paintings included in the exhibition. Clicking on any image will reveal a larger version of it.

HARRIS' HAWK & COMMON CHUCKWALLA (2006)

acrylic 30" x 20" Accumulations of rock make wonderful habitat for many lizard species. They provide a wide temperature gradient that makes it easy for the animals to thermoregulate, basking on a sun-exposed surface to elevate their body temperature and retreating into a cool crevice to lower it. Such crevices also afford safe fortification against many predators. Among the most rock-adapted reptiles are the five Chuckwalla species of the genus Sauromalus, found in the southwestern U.S. and northern Mexico, including a number of islands in the Sea of Cortez, where three described species are endemic. The Common Chuckwalla (S. ater) is typical of the group. With a high optimal body temperature, it spends a lot of time basking on exposed rocks, usually near a crevice which it will slide into at the hint of danger. If pressed, it will gulp air and inflate its body, making it quite impossible to pull out. Harris' Hawk (Parabuteo unicictus) is a unique raptor species of the American tropics. Normally shunning thick jungles, it haunts llanos, chaco, chaparral and scrub forest in the drier parts of that region, ranging as far north as the southern tip of Nevada. Fast and powerful, this social bird feeds on a variety of prey, from rabbits and ducks to reptiles. Incidental subjects in this painting include a Compass Barrel Cactus (Ferocactus cylindraceus ), honey ants (Myrmecosus sp.), Desert Spiny Lizard (Sceloporus magister), Turkey Vulture (Cathartes aura), and Costa's Hummingbird (Calypte costae). 


RIPARIAN RASHOMON (2009)
acrylic diptych on illustration board  15"; 15" x 20"

Two different viewpoints of the same event illustrate some of the common evasive strategies employed by frogs. The Brilliant Forest Frog (Rana warszewitschii) inhabits rain forests from Honduras to Panama. When resting upon leaf litter, its drab dorsal colors are cryptic, but bright yellow spots on its thighs flash when it leaps, and a glimpse of its brilliant underside is even more likely to startle and confuse a predator. Upon disappearing beneath the water's surface, it usually follows a wild, zig-zag course, ending up some distance from where the naïve viewer might anticipate. This painting's antagonist, the Agami Heron (Agamia agami), ranges through most of Tropical America, but does not occur in great numbers anywhere and is infrequently seen. Long of neck and short of leg, it haunts streams within heavy forests and feeds upon small fish and amphibians. Incidental subjects in this painting include a water strider (Gerris sp.), damselfly naiads (family Coenagrinionidae) and a White-necked Puffbird (Notharchus macrorhynchus).

AGARRANDO LA MAÑANA—BLACK VULTURES (1994)
India ink wash 14" x 20"

The vultures of the Americas are very different from their Old World namesakes, which are related to eagles. Of the eight New World vulture species, three can be observed in North America: the California Condor, the Turkey Vulture, and the one depicted here, the Black Vulture (Coragyps atratus). The idea for this piece was born while topping a ridge on Costa Rica’s Osa Peninsula one morning, when I spied a distant group of black vultures enjoying the dawn’s first rays in typical vulturine style. The birds were sunning in bright light, but the still thick jungle fog separating us obscured their images, simplifying their forms into boldly abstract shapes of two tones: the illuminated and the shaded. I quickly sketched some designs based on what I saw, which I planned to later organize into a painting. Black vultures are common throughout the American tropics and are plentiful and tame in many towns, so I expected to have little trouble finding cooperative models from which to render some more detailed drawings, once I had rough sketches of the basic design I wanted. As plans often are, these were soon frustrated, and as I made my way east, sunning vultures would invariably fold their wings once I removed my pack and fumbled for a pad and pencil. I was finally reduced to sleeping in the local dump outside of Antón, Panama, and positioning myself next to a favorite roost before daylight, where the final drawing for this piece was at last executed.

A KERANGAS FOREST FLOOR (2010)
acrylic on illustration board 30" x 20"

Of all of Borneo's varied ecosystems, perhaps none is more surprising than the biologically impoverished (by equatorial standards) dwarf forests that occur throughout the island, but more commonly in the west. The ecologist P. W. Richards called them “heath forests” after the similarly infertile lands of his native England, but they're better known by the Iban term kerangas, which means “land which will not support rice cultivation.” Kerangas soil is typically acidic, sandy and podzolized, or heavily leached. Essential elements enter the soil from decaying leaf litter, but most of these, magnesium, carbon, nitrogen and calcium in particular, leach away very quickly, and are only available in the top few inches. Phosphorus seems to leach away more slowly. Continual deposition of leaf litter is critical to the system, and disease, fire and logging or clearing for agriculture will convert kerangas to a barren habitat dominated by grasses and sedges known as padang (“field” in Malay). Despite the poor soil, healthy kerangas forests are dense with trees, most of them under 30 feet tall and three inches in diameter. In contrast to most equatorial forests, only a few species are represented. Dominant tree species usually belong to the mangosteen family, Clusiaceae, and to one or more of the genera Cratoxylum, Calophyllum and Ploiarium. Orchids show the greatest species diversity among kerangas plants, and terrestrial as well as epiphytic species are usually in evidence. Species of melastomes, laurels, myrtles and gingers are also commonly represented. Many kerangas plant species bear nitrogen-fixing bacterial nodules on their roots, and carnivorous plants also thrive. Borneo's kerangas forests are a center of diversity for the pitcher plant genus Nepenthes, which trap insects in leaves which are modified into water-bearing pitchers. At least one Bornean species, N. rajah, secretes a nectar that attracts tree shrews whose droppings are captured in the pitcher to nourish the plant. In perennially wet padang habitat, Bladderworts (Utricularia spp.) and Sundews (Drosera spp.) also trap small arthropods. Another famous kerangas denizen is the epiphytic ant plant (Hydnophytum spp.), which forms a symbiotic relationship with ants, providing them shelter, while receiving protection from the colony and nutrients from its wastes. This painting depicts a small patch of kerangas forest floor. Included in the leaf litter are shed leaves of the dominant tree Cratoxylum glaucum and shed needles of the podocarp (primitive conifer) Dacrydium becarii. Various mosses of the family Calymperaceae and the showy terrestrial slipper orchid Paphiopedilum javanicum grow from the soil and a single dried Nepenthes ampullaria pitcher sits on the floor while pitchers of N. stenophylla hang from epiphytic vines. Duméril's Monitor (Varanus dumerilii) occurs near rivers in various types of forest throughout the island. The hatchlings, like the one shown, are well-known for their striking coloration. It has been suggested that the colors, which begin to fade at the age of six weeks, mimic the dangerously venomous Red-headed Krait (Bungarus flaviceps), which shares its Southeast Asian range. Among Borneo's diverse and beautiful dragonflies, probably none is more conspicuous than the Red Swampdragon (Agrionoptera insignis), a member of the skimmer family, Libellulidae. Other subjects include the left-handed land snail Dyakia kintana and a Giant Forest Ant (Camponotus gigas), whose dimorphic workers forage for honeydew and other organic matter on the ground and in the canopy. At over an inch in length, the major workers of this species are among the world's biggest ants. Finally, a procession of Longipeditermes longipes termites returns to the nest with balls of lichen in tow. Both the workers and soldiers of this monotypic genus come in two sizes. Like other members of their subfamily, nasutitermes, the heads of the soldiers are distorted into nozzles, through which they can spray noxious chemicals at enemies, chiefly ants.

AJOLOTE (2011)
acrylic 14" x 9"Just where on the reptile family tree to put the worm lizards, or amphisbaenians, has long been a puzzle for taxonomists. Traditionally lumped with the lizards, these days they're more often given their own suborder alongside the snakes and lizards. What ever their systematics, their appearance and habits share more in common with earthworms than reptiles. Spending most of their lives below ground, they progress with a worm-like, peristaltic movement of their body segments. On the surface, they can move in a more typical serpentine fashion. Most amphisbaenians are found in tropical Africa and South America, but a few are found as far north as the Mediterranean, and in the Americas to Florida and northwestern Mexico. Unlike other amphisbaenians, the wormlike visage of the little-known Mexican genus Bipes is rather spoiled by the presence of a pair of stout digging forelimbs. Like the rest of their group, none of whom bear visible limbs, the 3-4 known Bipes species dig by forcing their hard little noses into the soil and moving them back and forth. The forelimbs are used to push loosened soil out of the way. They seem to subsist mostly upon termites and ants, and sometimes forage upon the surface at night. Two Bipes species occur in Michoacán and Guerrero, but the best-known of the group, B. biporus, is found in Baja California, where it is known as the Ajolote. A number of very poorly documented records from other parts of Mexico, Arizona, and as far north as Nebraska, suggest that it may be more widespread than believed. In this painting, an Ajolote forages about a rotting fencepost on termites of the genus Reticulitermes.

BLACK SKIMMER (2003)
acrylic on illustration board 22" x 30"

The peculiar skimmers are related to gulls and terns, and live near fresh or saltwater bodies in the warmer regions around the globe. The three species, one African, one Asian, and one American, are all quite similar in appearance and behavior. The lower mandibles of these birds are much longer than the upper -- an adaptation well suited to their unique method of foraging. A feeding skimmer flies just above the water, plowing the surface with its lower bill. When a fish is encountered, it is snapped up and consumed. Another peculiarity of the skimmers is the presence of a vertical pupil, something found in no other bird. This allows greater control of light entering the eye, which is useful to fowl that spend a lot of time resting on white sands reflecting the tropical sun, but which also regularly feed at night. Skimmers nest in a small scrape on a sandy beach; the female usually lays 3 to 7 eggs. The Black Skimmer (Rynchops niger) ranges along the Atlantic coast of the Americas and along rivers from the southeastern U.S. to Brazil and from southern California to Peru along the Pacific. It also nests along the Sea of Cortez.

RETICULATED PYTHON & MASKED FINFOOT (1999)
acrylic on illustration board 20” x 30”

Capable of attaining a length of thirty feet, the Reticulated Python (Python reticulatus) is by far Asia's biggest snake. Ranging through most of Southeast Asia and the Philippines, as far east as the Moluccas, the “rectic” is a versatile reptile, occurring in all manner of habitats and inclined to feed opportunistically on most any animal it encounters when hungry, from rats to deer to birds like this Masked Finfoot (Heliopais personata), an uncommon and distant relative of the coot. I agonized over whether or not to paint the female finfoot swimming beneath the surface, which is atypical behavior according to most of the literature (I've never seen this species in nature), but having read some accounts of their occasional diving, and having seen their American equivalent, the Sungrebe (Heliornis fulica), dive once, I decided to surrender to my artistic impulses. The incidental species in this piece include the butterfly Junonia almana, a robust damselfly (family Lestidae), water strider (family Gerridae), a minnow (family Cyprinidae), loach (family Cobitidae), Forest Softshell Turtle (Dogania subplana), and Water Skink (Tropidophorus berdmorei).
CALIFORNIA CONDOR (2005)
oil 72" x 108"

Giant condors were successful and important components of the North American Pleistocene fauna, but began to decline about 13,000 years ago, with the extinction of mammoths, ground sloths, and other huge mammals conventionally attributed to the Human Clovis culture. The last giant North American species, the California Condor (Gymnogyps californianus), declined to a mere 22 birds in the mid-1980s, and the bold decision was made to take the entire population into captivity. The captive propagation program has been wildly successful, increasing the population to over 400 today. More than half of these birds have been released and live, semi-wild, in California and Arizona, including a number of wild-fledged condors, the first of which hatched in 2002. The species is still far from recovered, though, and is continually dependent on human management. The mortality rate, mostly from power-line collisions and lead bullet ingestion, still exceeds wild births. Reintroduced condors show problematic behavior like extreme tameness, and lead poisoning has caused most of the Arizona birds to be re-trapped for chelation therapy.
LESSER FLAMINGOS (2005)
oil 72” x 108” Of the six species of flamingo, the Lesser Flamingo (Phoeniconaias minor) is the most abundant, with some four million individuals ranging across sub-Saharan Africa, and smaller populations on the Arabian Peninsula, India and Pakistan. At just over a yard in height, it is also the smallest species. Haunting shallow, alkaline lakes, it feeds by forcing water through lammellae inside its bill with powerful tongue movements, filtering out algae and tiny invertebrates. Throughout its range, the Lesser shares this habitat with its much larger cousin, the Greater Flamingo (P. roseus). There is little competition for food between the two, though. The lesser feeds nearer to the surface, and its bill filters out smaller organisms.

FLY RIVER TURTLE (2004)
acrylic 18” x 24”

Looking for all the world like the Mock Turtle from Alice in Wonderland, the Fly River Turtle (Carettochelys insculpta) was described to science science in 1887 from a specimen collected in New Guinea. In the 1970s a second population was discovered in the north of Australia's Northern Territory, making it the only known non-pleurodire freshwater turtle on that continent today. (The pleurodire, or side-necked, turtles of Australia, Madagascar, Africa and South America comprise a clearly-defined suborder that is distinctive in having long necks that are retracted in a horizontal “S,” laying the head sideways over one leg. The Fly River Turtle is the sole surviving species of a family that probably diverged from the ancestral softshell turtles during the late Jurassic. The group probably originated in Asia before spreading to Africa, Europe and North America during the Cenozoic era and finally dying out everywhere but New Guinea, where Carretochelys is known from fossils dating back at least 14 million years. Its stint in Australia is thought to have been a matter of tens of thousands of years, but just how it made the passage to the Northern Territory is a matter of some debate. The most physically adapted for swimming of all modern freshwater turtles, its resemblance to sea turtles is a case of convergent evolution, where similar pressures caused the same traits to evolve independently in two different groups. Living in slow-moving rivers of Southern New Guinea and Northern Australia, the Fly River Turtle is more herbivorous than most highly aquatic fresh-water turtles, supplementing its diet of aquatic plants and fallen fruit with snails and other invertebrates, carrion, and occasional fish, although the Red-striped Rainbowfishes (Melanotaenia splendida) skulking among the submerged snags have little to fear from the passing reptile. A dragonfly naiad (Aeschna sp.) clings to the same snag. Although it is protected in Australia, Indonesia and P.N.G., the Fly River Turtle has recently been smuggled out of those countries in large numbers for the pet trade. In just a couple of years this smuggling has exploded to the point that the World Wide Fund for Nature was moved to put the turtle on its “10 most wanted” endangered species list in September 2004.

PLUSH-CRESTED JAYS MOB AN ORNATE HAWK-EAGLE (2003)
acrylic on illustration board 30” x 10”
Plush-crested Jays (Cyanocorax chrysops) are common birds that exploit the edges of a variety of tropical forest types from Eastern Peru to Northen Argentina. Like most corvids, they persecute birds of prey relentlessly. In this painting a group of these jays harass the big and powerful Ornate Hawk Eagle (Spizaetus ornatus), a spectacular raptor that is rather uncommon, but found widely throughout the neotropical region. Similar eagles of the same genus are found throughout Asia and tropical Africa. Incidental creatures in this piece include a Thread-waisted Wasp (family Sphecidae), a Blue Morpho (Morpho peleides), an anole (Anolis sp.), White-necked Puffbird (Notharchus macrorhynchus), Red-legged Honeycreeper (Cyanerpes cyaneus), and a Saddleback Tamarin (Sanguinus fuscicollis).


GREAT TINAMOU (1994)
acrylic 20” x 15”
Tinamous comprise an order of chicken-like birds found in tropical America that is at least 17 million years old. Forty-seven modern tinamou species are placed in a single family. They are related to, and according to some authorities should be considered members of, the ratites, the group containing the Ostrich, Emu, Kiwis, Cassowaries and Rheas. Unlike the ratites, Tinamous have a keeled sternum and can fly, though not terribly well. It seems likely that ratites and tinamous derived from a common flying ancestor, and the rheas, ostriches and Antipodean ratites each evolved flightlessness separately, but this model is not universally accepted. Male tinamous select the nesting site, and frequently mate with multiple females, who lay their eggs in the communal nest. The eggs are incubated and the young are defended by the male. The Great Tinamou (Tinamus major) is one of the most widespread species, ranging from Guatemala to Bolivia and central Brazil. Its beautiful turquoise eggs are laid between the buttresses of a large rainforest tree like this Hymenolobium. When a male with chicks is ill at ease, he strikes an odd posture, tilting body forward with rump in the air. Once apprehension turns to fear, he engages in a distraction display, running about wildly with wings spread and head held low. The haunting, mellifluous whistle of this bird is a common evening sound in pristine neotropical lowland rainforests, and I tried to convey in this painting some of the mournful, mysterious mood it evokes in me. On the ground a Central American Jungle Runner (Ameiva festiva) basks while a yellow Eyelash Palm Pitviper (Bothriechis schlegelii) lurks in the foliage above. The huge liana draped about the tree trunks is the spectacular Monkey Ladder (Bauhinia guianensis).

GOSTOSO!--MANED WOLVES & THREE-BANDED ARMADILLO (1997)
acrylic 20" x 30"
The imperative to avoid being eaten is one of the prime drivers of evolution. Protective armor can be an effective defense, and it has evolved independently in many animal groups. Its main liability is in adding weight and reducing flexibility, limiting the creature's ability to flee from predators, thus entrenching its own value. Among modern mammals, this defense is the specialty of the South American armadillos, whose success is confirmed by the family's northward expansion over the past few million years, leaving two species in Central America, one of them ranging well into the United States. Best protected are the two species of three-banded armadillos (Tolypeutes spp.), which can roll up into perfect plated spheres. In this painting, the fortification of the species T. matacus is being tested by a pair of gangly, knock-kneed Maned Wolves (Chrysocyon brachyurus). These unusual canids of the South American plains have no close living relatives. Despite their exceptionally long legs, they are not particularly fleet of foot, but probably benefit from them by being able to see over tall grass. Normally solitary hunters, during the breeding season mated couples often forage together. The Maned Wolf's diet consists of small mammals, birds, reptiles, insects and fruit. The title of this painting, “Gostoso,” is a Brazilian soccer cheer, roughly the Portuguese equivalent of “tasty.” Incidental subjects include Pampas Grass (Cortaderia argentea), Spiny Tree Lizard (Tropidurus spinulosus), Yellow-headed Caracara (Milvago chimachima), spinetail (family Furnariidae) and Black Howler Monkey (Allouatta caraya).

ASCENSIÓN-- STRAWBERRY POISON FROG & TADPOLE (2004)
acrylic on illustration board 40” x 15”
 
To be evolutionarily successful, an individual organism must produce offspring that live long enough to produce another generation themselves. This mandates an investment on the parents' part. One option is to lay out the energy to produce great quantities of young, most of which will be eaten by predators or otherwise fail to reach adulthood. Producing fewer young requires parental care to assure their long-term survival. Reproductive approaches that produce lots of young are known as r-strategies, those that depend on parental care are called K-strategies. Among vertebrates, the frogs exhibit the greatest diversity of reproductive tactics, including extreme cases of r- and K-strategies. Most notable of the K-strategists are the poison frogs of the family Dendrobatidae, a group of beautiful and tiny diurnal amphibians found throughout the American tropics, well known for producing complex alkaloid skin secretions. The Central American Strawberry Poison Frog (Oophaga pumilio) deposits several eggs on a leaf on the forest floor, which are guarded by the male. Upon hatching, the tadpoles wriggle onto the female’s back, and are taxied up the trunk of a tree to a pre-selected bromeliad, where they are deposited into one of the water vessels formed within the axils of these arboreal epiphytes. Every few days, the female lays an unfertilized egg for each of her offspring to feed upon. Incidental creatures in this painting include an Agouti (Dasyprocta punctata), a Spectacled Antpitta (Hylopezus perspicallitus), a Racerunner (Ameiva festiva), a Lanternbug (Fulgora laternaria), a Leaf-Footed Bug (Anisosceles sp.), a leafhopper (Umbonia sp.), a Consul Butterfly (Consul fabius), and numerous ants of the species Pheidole bicornis, which are dependent upon Piper trees, like the one immediately behind the frog. 


HARMATTAN HARMONY—BLACK KITE, GRAY
PLANTAIN-EATERS & RED-BILLED HORNBILL (2002)
India ink wash and watercolor on paper 20” x 30”

Around the first of December, hot northeastern winds begin to blow fine sand from the Sahara across the savannas and deciduous scrub woods of Africa’s Sahel, that transitional zone between the Sahara Desert and the equatorial forest belt to the south. Thus begins the Harmattan, a three-month season during which the entire world appears dingy gray-brown. This annual phenomenon, along with its obverse, the late-summer southwestern monsoon, are crucial factors in forging the ecology of the region. Organisms must be able to exploit the bounty brought by the summer rains and survive the fires and desiccation wrought by the harsh Harmattan winds. In this painting, three of the Sahel's most conspicuous avian residents, a Black Kite (Milvus migrans), two Gray Plantain Eaters (Crinifer piscator), and a Red-billed Hornbill (Tockus erythrorhynchus) perch among the naked tree branches that characterize the Harmattan season.

GREEN IGUANA & LEAF-CUTTER ANTS (2011)
acrylic on illustration board 18” x 24”

One of the best-known denizens of the Neotropics, the Green Iguana (Iguana iguana) is a large and successful herbivorous lizard found in a variety of forest types from Mexico to Paraguay. Approaching the iguana in familiarity are the 50 or so species of leaf-cutter ant, which share the lizard's expansive range. The species shown is Atta cephalotes. Leaf-cutter ants are unusual, but not unique among insects in their practice of agriculture. Four different types of workers maintain the colony. The largest type, the majors, function as soldiers, defending the colony from marauders. Next in size are the mediae, which spend the day foraging for fresh leaves, which they cut into nickel-sized pieces and bring back to the colony. The sight of a mass of green leaf fragments moving slowly across the forest floor, tilting rhythmically back and forth like butterfly wings, is a common delight of New World forests. Attending the mediae are the smaller minors and minims,which protect the foraging phalanx from predators and parasites. Inside the subterranean nests, the minim workers crush the leaves, which serve as a growing medium for fungi of the family Lepiotaceae, which feed the colony. Incidental subjects include the spectacular monocot Heliconia pogonantha, a Red-capped Manakin (Pipra mentalis) and the butterfly Antirrhea pterocopha.

LANJAK DAWN—CROWNED FLYING LIZARDS & ORANG-UTAN (2009)
acrylic 20" x 30"

One of the biggest difficulties with an arboreal lifestyle is getting around. Even in the thickest forests, the canopy is rife with gaps between individual trees, caused by wind abrasion. This phenomenon, known as crown shyness, discourages transmission of tree pathogens, permits light into the understory, and facilitates tree respiration. It also seriously complicates life for arboreal animals that need to disperse and forage without descending to the dangerous forest floor. This problem has spurred many different animal groups to independently evolve patagia, or flying membranes, to enable them to glide across these gaps. Among the most accomplished forest gliders are the twenty or so lizard species known appropriately as flying lizards (Draco spp.), which are distributed throughout Southeast Asia and the Philippines. Five or six pairs of false ribs support their flying membranes, which enable them to glide for many yards with amazing dexterity. I’ve seen them launch themselves from a tree, turn around, then return to the same trunk. In addition to their primary function, these often brilliantly colored patagia are frequently employed as signals to communicate with conspecifics. Towards the end of the dry season, the males establish territories, actively guard them, and begin displaying for females by extending their long throat dewlap and one or both patagia. The Crowned Flying Lizard (D. cornutus) ranges in wooded areas on Borneo, Sumatra, western Java and the Bunguran and Sulu Archipelagos, where it forages among the treetops for the ants and termites that make up the bulk of its diet. In southern Sarawak, I found them in hilly, secondary forest. In the background of this painting, a large male Orang-utan (Pongo pygmaeus) calls from his sleeping nest. Other incidental creatures include a bark orb-web spider (Caerostris sp.), ants of the genus Bothriomyrmex, a lanternbug (Fulgora sp.), a Malaysian Bushbrown (Mycalesis fusca) and a Black and Yellow Broadbill (Eurylaimus ochromalus).


 

HARPY EAGLE & THREE-TOED SLOTH (2005)
oil triptych on linen 36" x 36"; 36" x 36"; 36" x 36"
Within the order of birds, no more powerful predator exists than the Harpy Eagle (Harpia harpyja). A large hen can weigh as much as twenty pounds. Long-tailed and surprisingly agile for such a massive bird, this uncommon eagle hunts monkeys, sloths, agoutis and other similarly-sized mammals throughout the tropical rainforests of the New World, flying in short sprints and rarely venturing above the forest canopy, except during nesting, when an eyrie high in an emergent tree cradles the single egg. The Harpy's terrible feet are unmatched in size and power: four brutal toes drive three-inch talons deep into the quarry's vital organs. Here the unfortunate victim is a Three-toed Sloth (Bradypus infuscatus), a well-known and common rainforest mammal, plucked from its favorite food source, a Cecropia tree.

GREATER ROAD-RUNNERS & CANYON TOWHEE (2011)
acrylic 15" x40"
The cuckoos are an ancient and fascinating bird group that is at least 40 million years old. This group never seemed to produce any powerful fliers, and had a tendency to evolve into running forms. Despite this, it managed to colonize Australia, New Guinea and Madagascar. Many cuckoos are brood parasites, meaning they lay their eggs in the nests of other birds, to be raised by “Foster parents,” but this is by no means universal. Cuckoos exhibit a variety of parenting strategies, including cooperative parenting in the anis and role reversal and polyandry in some of the coucals. Among the many cuckoo taxa are the Old World coucals, the Madagascan couas, the Asian malkohas, the American anis and perhaps the African turacos...maybe even the bizarre South American Hoatzin (Opisthocomus hoazin). Also included is that icon of the desert southwest, the Greater Roadrunner (Geococcyx californianus), one of two members of a genus found in Mexico and the southwestern U.S. The two species look and behave similarly, but the larger, northern one has a longer bill and prefers more open country. Roadrunners are omnivores, feeding on fruits and seeds as well as large invertebrates, reptiles and small birds and mammals, which they run down. Most of the situations I paint are hypothetical but plausible, but this piece was based upon an event I witnessed in California's Anza Borrego State Park: a roadrunner carrying a small dead bird was running in the haphazard, zig-zag evasive mode typical of its species, barely keeping half a step ahead of the mobbing of two others. Unfortunately, distinguishing whether any of the fast-moving birds were adults or juveniles was beyond my capacity, and identifying the sex of a roadrunner under any circumstances requires nearly supernatural powers. My assumption is that I watched either two juveniles chasing a parent or a mated pair chasing a young, inexperienced bird. The former scenario is probably most likely, but it was the latter that I selected to commit to illustration board. I was unable as well to identify the focal point of the fracas, but decided that one of the nondescript brown towhees would serve as good a candidate as any. While taking artistic liberties, I also moved the setting slightly east, to the Sonoran Desert, and painted the scrub towhee of that region, the Canyon Towhee (Pipilo fuscus). 

SOUTHERN CROSS—PENNANT-WINGED NIGHTJAR (1998)
India ink wash on paper 23” x 17”

A crucial ingredient to evolutionary success is the ability to find and attract a high-quality mate. This is so important that it's caused many animals, usually the males, to evolve outlandish ornamentation that is attractive indeed, but downright dangerous, too. Think of the peacock's long train that slows down his flight and impedes his ability to run through thick brush. Because really big ornaments are so risky, they almost always evolve in temporary integumentary derivatives like feathers that can be shed as soon as possible after the eggs have been fertilized. Among nature's most spectacular breeding adornments are the wings of the male Pennant-winged Nightjar (Cosmetornis vexillarius) of southern Africa. His inner primary feathers form a pair of “pennants” that can trail over two feet behind him. During his nuptial display he flies in low circles, emitting a strange katydid-like twitter. Finding an exposed perch like the termitarium in this painting, he then spreads his wings, slowly rotating them. Soon after breeding, the inner-most primary is dropped, but the rest of his moult does not continue until after the migration north of the equator for the austral winter, which often is executed in flocks. A member of the same order that includes the American nighthawks and whip-poor-wills, the Pennant-winged Nightjar feeds in the manner characteristic of the group, on insects captured in flight.

VARIOUS CENTRAL AMERICAN BUTTERFLIES (2003)
watercolor on bristol board 10” x 36”
These seven butterflies were selected and copied life-size from field studies in my Central American sketchbooks from 1992-'93. Both dorsal (upper) and ventral (under) views are shown. Lycorea cleobaea was a roadkill from Golfito, Costa Rica. Opsiphanes tamarindii was found drowned in Lake Nicaragua. Morpho peleides was found dead at La Selva Biological Station, near Puerto Viejo de Sarapiquí, Costa Rica. Caligo illioneus was roadkilled near Tocumén, Panama, a fate shared by Papilio thoas in Chiriquí Grande, Panama, and Colobura dirce near Esparza, Costa Rica. Callicore peralta was found dead on Costa Rica's Osa Peninsula.

GELADA (1998)
acrylic 26” x 10”A large and unusual monkey restricted to the mountains of central Ethiopia is the Gelada (Theropithecus gelada). Like that of its close relatives the baboons and drills, Gelada social structure is very complex and dynamic. In this species the social units seem to be held together mostly by the bonds between individuals of the same sex, especially females, who tend to associate more among themselves. Nights are spent sleeping on the faces of rocky gorges, where the animals feel secure and from where they never stray far. During the day they forage for grasses, from which they derive practically all of their nourishment -- a diet unique among primates. The adult male is easily distinguished by his long golden mane and whiskers, and a red triangle of bare skin framed by short grizzled fur on his chest. Perching elsewhere on the cliff are a lizard (Agama sp.) and a Pectinator (Pectinator spekei), an unusual rodent distantly related to the Chinchilla.

COAST HORNED LIZARD (1998)
acrylic 21" x 30"This piece was designed to look as austere as the Mojave desert that it depicts. The subject, a Coast Horned Lizard (Phrynosoma coronatum) is centered on the board within very simple arcking lines, eating honey ants (Myrmecocystus sp.), ants being the usual horned lizard fare. A total of fourteen species of horned lizards populate most of western North America, from southern British Columbia to Guatemala. They are a pretty uniform lot, small lizards (the giant among them is a Mexican brute of eight inches), flattened and covered with spines. The species depicted and one other are known to have the ability to squirt a potential predator with blood issued from the corner of their eye. This capacity was reported for over a century in popular lore and has been doubted by many, and only recently confirmed conclusively. The response seems to be an anti-coyote device, which accounts for the usual inability for non-canid creatures like us to elicit it.  

GOLDEN EAGLE & PRAIRIE FALCON (2011)
India ink wash on paper  22" x 15"

The powerful Golden Eagle (Aquila chrysaeotos) is holarctic in distribution, meaning it is found across the northern hemisphere. In North America, it is mostly a creature of the west, and is largely a predator of jackrabbits (Lepus spp.), but it is an adaptable hunter that will exploit what is available. Its large size confers upon its flight an illusion of lethargy, but in reality, it is a swift and commanding aerialist capable of taking a variety of flying prey. Mated pairs often hunt cooperatively, adding a level of advantage in such situations. I know of a pair in Utah that feed almost exclusively on Ravens (Corvus corax). Here, a Prairie Falcon (Falco mexicanus), North America's large desert falcon, evades the stoop of a marauding Golden.

JAGUAR & COLLARED PECCARIES (1994)
oil 32" x 42"The only modern big cat of the New World, the Jaguar (Panthera onca) ranges from northern Argentina to the southwestern U.S. Until the end of the Pleistocene, it occurred throughout the southeastern states, and seems to have been especially common in Florida. Over the past century, its presence north of Mexico has probably consisted of little more than occasional young males dispersing from Mexico. An old male Jaguar that had roamed southern Arizona for a number of years died in 2009. These cats are generalist predators, feeding on a wide range of quarry, but in the northern part of their range, the piglike Collared Peccary (Pecari tajacu) or Javelina is by far the most important. Here, one of these great cats prepares to charge a group of peccaries in a deciduous scrub forest. The colors and lighting of this habitat, especially during the dry season, when many of the trees have lost their leaves, are quite peculiar unto themselves and inspired me to try to characterize them with this sort of “hyper-pointilism” technique.

 

DUSKY-GILLED MUDSKIPPERS (2009)
acrylic on illustration board 6” x 12”

Life began in the sea and remained there for over 3 billion years. It's learned to thrive on land just as well, but occupying the intertidal zone, that strip between the two, continues to be a difficult trick, both ecologically and physiologically. High salinity, fluctuating tides, silty soils that are low in oxygen and nutrients, and harsh sunlight combine to make this territory a complicated and difficult living habitat. Among fish, the best adapted to this zone are the mudskippers, a subfamily of Old World tropical gobies. Capable of breathing air through their skin and mouth linings, mudskippers flop about the beach at low tide, foraging for small arthropods. At high tide, they mostly retreat to underwater burrows, where low oxygen and high ammonia levels would quickly kill most fish. This painting depicts a common Southeast Asian species, the three-inch long Dusky-gilled Mudskipper (Periophthalmus novemradiatus). Like most mudskippers, it is usually associated with mangroves, or intertidally adapted trees. Around 45 species of trees in ten genera and five families constitute the true mangroves, but plants from over a dozen other families are usually lumped into the designation as well, including a palm, a screwpine and a sedge. The many adaptations evolved by mangrove trees include numerous systems for storing gases and nutrients, and branching stilt roots and pneumatophores or “breathing tubes” rising from under the ground. Roots and stems are highly impervious to salts, and some species have evolved special glands for excreting excess salts. Many species engage in photosensitive leaf movements to limit evaporation. Mangroves form the basis for rich intertidal ecosystems called mangrove swaps or mangals. Incidental species in this painting include a fiddler crab (Uca sp.), hermit crab (Dardanus sp.), a Reef Heron (Egretta sacra), terns (Sterna spp.) and a Lesser Frigatebird (Fregata ariel).

GREAT PIED HORNBILL (2001)
acrylic 30” x 20”
The hornbills comprise a family of tropical Old World birds that are related to kingfishers but very similar in appearance and habits to the New World toucans, which are more closely related to woodpeckers. This is one of many examples in nature of convergent evolution, where two unrelated animals occupying similar niches evolve similar forms. Harder to explain is the fact that through much of their range in Asia, hornbills are known by the Malay name “tucan,” the same name given to their South American analogs by the Tupi Indians. Probably the best known hornbill is the Great Pied Hornbill (Buceros bicornis), which ranges from India through Sumatra and has been a rather common aviary bird for many years, having been bred in captivity since 1953. Like most hornbills, these are essentially forest birds, exploiting a number of different forest types up to an elevation of about 2000 meters. Usually occurring in pairs or small family parties, these birds sometimes congregate in groups of over one hundred to feed in large fruiting trees. Fruits, mostly figs, make up the bulk of their diet. During the breeding season the male bolsters the growing nestling's protein supply by delivering extra animal matter to the nest hole, where the female remains sealed until the chick is about half grown.

GREAT HELMETED HORNBILL (1998)
acrylic on illustration board 30” x 20”
The massive casques adorning the beaks of many hornbill species contain mostly air, but the Great Helmeted Hornbill (Rhinoplax vigil) of Southeast Asia has a casque of solid ivory. The function of this heavy ornament is the subject of some conjecture; it certainly makes flight more difficult, even with the counterbalancing effect of elongated central tail feathers. It has also resulted quite literally in a price being put on the birds' heads, which are coveted by Chinese artisans who for centuries have used them to create delicate carvings known as ho-ting. Some authorities claim that the males have "jousting" contests. Perhaps the extra inertia helps them peck away loose bark in search of prey. My own objective with this piece was simply to depict the effort necessary to hoist the bird's skull into the air. At the bottom of the field a startled Crested Lizard (Bronchocela cristatella) scampers away as a Little Spiderhunter (Arachnothera longirostra) forages amongst the foliage.

MALAYSIAN RHINOCEROS HORNBILL (2003)
acrylic on illustration board 18” x 30”
In the forests of the Sunda Shelf, from peninsular Malaysia through Sumatra, Java and Borneo, the braying calls and loud, huffing wingbeats of the Rhinoceros Hornbill (Buceros rhinoceros) are familiar sounds. Both sexes sport a large orange casque on the bill. This hollow ornament is larger in the male bird, and varies in shape among three distinct races: one on Java, one on Borneo, and one on Sumatra and the Malay Peninsula. In this painting a large male hops along a tree branch with a fig, possibly to present it to a mate walled up inside her nesting hole. This hopping gait is common among smaller perching birds, but I can think of no other bird as large as a hornbill that habitually moves in this fashion. Incidental creatures in this painting include a Slender Squirrel (Sundasciurus tenuis), a small flock of Oriental White-eyes (Zosterops palpebrosus), a Long-horned Beetle (Batocera sp.) and a Common Mormon Swallowtail (Papilio polytes).



PHANTOMS OF THE MOJAVE—BANDED GECKO (2010)
acrylic 20" x 15"

One of North America's most interesting waterways is the Virgin River, which flows out of southwestern Utah to form part of the Arizona-Nevada border before emptying into Lake Mead. It represents a northerly extrusion of the Mojave Desert ecological community which meets the Great Basin zone to the northwest and the Colorado Plateau to the northeast. It hosts numerous species of plants and animals that are found nowhere else, eight of them considered endangered. The river and its gorge extend the ranges of dozens of other species a hundred or more miles to the north. This painting depicts four Virgin River specialties. The Western Banded Gecko (Coleonyx variegatus) occurs in a number of desert habitats. This strictly nocturnal lizard remains well hidden until after dark. On moonlit nights its translucent body almost gleams, and it's easy to spot as it stalks its arthropod prey, its tail writhing, catlike. Here the lizard descends the woody skeleton of a dead Silver Cholla (Cylindropuntia echinocarpa) before the nocturnal, trumpet-like blossom of a Western Jimsonweed (Datura wrightii), well-known for its toxic and hallucinatory effects caused by the alkaloids atropine and scopolamine. Also visible is a Jimson Beetle (Lema daturaphila), the adults and larvae of which feed on Jimsonweed and other members of the potato family, Solanaceae.

REANIMATION—COMMON POORWILL
acrylic 30" x 20"Hibernation is one of the more effective strategies temperate animals have developed to survive winter's cold temperatures and lack of food resources. Many birds, bats and even insects opt instead for seasonal migration, exploiting distant habitats during different seasons. A few, like the Monarch Butterfly (Danaus plexippus) and the Poorwill (Phalaenoptilus nuttallii) use a combination of the two. Poorwills, relatives of nighthawks, breed in arid parts of western North America from southern Canada into northern Mexico; northern individuals seem to winter in the desert southwest. A number of bird species use daily torpor to minimize energy loss during cool nights or brief bad weather. Members of three related orders, the goatsuckers, hummingbirds and possibly the swifts, all show some abilities at metabolic adjustment, but none to the degree of the little Poorwill, which, in addition to its natural tendencies toward torpor, feeds heavily on beetles, rich in polyunsaturated fats, which remain liquid and metabolically available at low temperatures. In the laboratory, Poorwills have been observed sustaining periods of torpor for over 80 days, and in the wild as long as 25 days. A shallow shelter, open to the southern sun is selected: a patch of cactus or rock niche to which the bird develops substantial fidelity. After sundown, the torpid Poorwill's body temperature begins to fall, until the ambient temperature reaches 5.5?C, an apparent optimum hibernating level which the bird tries to maintain. Solar radiation raises the body temperature daily, presumably allowing the option to forage during warm nights. I know of no human witnesses to a Poorwill rousing from torpor in the wild, but I imagine the bird backing out of his shelter to fully bask in the final evening rays, periodically flapping his wings to elevate his body temperature. It's not known how severe a winter these birds can survive, but a sufficient winter insect population, rather than temperature, is probably the limiting factor. Instead of showing the kind of country where Poorwills are known to commonly winter, I tried in this painting to depict a habitat in the harshest extreme that I could imagine the bird toughing out. Also shown reviving are Glacier Lilies (Erythronium grandiflora), Convergent Ladybird Beetles (Hippodamia convergens), Western Boxelder Bug (Boisea rubrolineata), and snowfleas (Hypogastrura sp.), cold-adapted springtails that climb onto the snow's surface to feed on algae.


BAT-EARED FOX (2006)
acrylic on illustration board 20” x 10”

The evolutionary history of the dog family is a complex puzzle that poses many interesting questions. Zoologists have traditionally relied on physical features alone to piece together phylogenetic relationships between living animals, but the recent advent of molecular analysis has forced a lot of rethinking of evolutionary affinities. The peculiar little Bat-eared Fox (Otocyon megalotis) is one of a number of modern canids that appear to have no close relatives. Because of its primitive-looking teeth, which are very small and numerous, it was long considered to represent an ancient line. Chromosomal studies, though, suggest that its odd dentition resulted from a fairly recent mutation, and that the species arose from an ancient fox lineage that also gave rise to the similar Fennec (Fennecus zerda) of North Africa and the cat-like American gray foxes (Urocyon spp.). Bat-eared Foxes are found in African steppes and savannas from Ethiopia and Angola south. Capable diggers, they excavate complex tunnel networks where they escape from heat and enemies and raise their litters of two to six pups. They feed on insects, mostly termites, with occasional vertebrate supplements.

STARGAZING--PEREGRINE FALCON (2008)
India ink wash on paper  15" x 24"

Damage to a bird's central nervous system from injury or poisoning often manifests itself in a behavior known as stargazing. Affected birds exhibit unsteadiness and a backwards craning of the head. Terrible though this gesture appears, it does not always herald doom for its sufferer; it can be a symptom of numerous temporary or transitory maladies. Animals like the cosmopolitan Peregrine Falcon (Falco peregrinus), which feeds heavily on seabirds, are especially prone to poisoning by persistent environmental toxins, by virtue of their place in the feeding hierarchy. Rains wash poisons into the sea, where they accumulate in organisms and concentrate as they rise from one trophic level to another. For example, the mercury level in the tissues of a population of medium-sized fish can be expected to be far higher than that in the population of small fish they feed upon, and far lower than in the big fish that feed upon them. During the 1950s and '60s, many populations of Peregrine Falcons crashed due to poisoning from the persistent pesticide DDT, which stimulated production of two enzymes in the birds that broke down calcium carbonate, the compound that forms eggshells. The resulting thin-shelled eggs usually broke before hatching. The worldwide banning of DDT for agricultural use and a rigorous captive breeding and reintroduction program have restored this bird back to healthy numbers.

DISCOURAGER OF HESITANCY--KING-IN-HIS-CARRIAGE ORCHIDS (2012)
acrylic on illustration board 15” x 7½”

For plants, sex is largely a crapshoot. Early flowering plants increased the odds of transferring their male gametes to a receptive ovary from the appropriate species by producing nectar to lure insects and other mobile animals that inadvertently transported pollen from one plant to another. This made it possible to invest less energy to producing masses of pollen, but producing nectar is also energy-intensive. Certain plants living in nutrient-poor conditions circumvented this problem by deceiving their pollinators. This strategy has become especially elaborate in the orchid family, where thousands of species rely on pollinators that are lured to the flowers by color and/or scent cues, without the nectar payoff. A European orchid genus and nine Australian ones are known to use sexual deception. In these cases, a male wasp must try to copulate with the flower in order to pick up pollen and to transfer it as well. The flowers produce pheremones attractive to an insect, each orchid species attracting a different insect species. In some cases, the exact same compound is produced by orchid and wasp, and is found nowhere else in nature. The flower also bears some physical resemblance to a female insect. The little hammer orchids (Drakaea spp.) of southwestern Australia attract wasps of the family Thynnidae. The most widespread of the hammer orchids, the King-In-His-Carriage (D. glyptodon), grows in sandy heath and is pollinated by the wasp Zapilothynnus trilobatus. The flightless female wasp climbs a sedge blade or other plant when receptive, and waits for a flying male to whisk her off. Here she bears a modest resemblance to the warty dark labellum of the King-In-His-Carriage sitting atop its slender stem. When a male thynnid wasp attempts to carry off the flower's labellum, the hinged stem knocks the insect into the column, dusting the wasp or the stigma with pollinia.

A BRICK HOUSE— ENGLISH SPARROW & PAPER WASP (1992)
acrylic 27" x 23"

Commensalism is a type of ecological relationship that lies between parasitism and symbiosis, that benefits one organism but has little effect on the other. A well-known example is the relationship between remoras of the family Echeneidae and the sharks and other large marine predators that they adhere to. The remoras feed on scraps left over from the shark and receive protection from it too. The effect of the relationship on the shark seems to be small, although, as in most commensal relationships, subtle effects probably do occur. There are many kinds of bacteria that form commensal relationships with Humans—also many that are symbiotic and parasitic. A number of animals are well known Human commensals, too. Among these is the House Sparrow (Passer domesticus), an originally Eurasian bird that has followed our species in its expansion across the globe. Although it occurs today nearly everywhere that Humans do, it is rare to see a House Sparrow far from Human habitation. It was introduced to North America just over 100 years ago. The most famous importation of House Sparrows was by Eugene Schiffelin, who attempted to introduce every bird mentioned in Shakespeare's works into Central Park. Most of these introductions failed, but within a century, House Sparrows spread throughout the Western Hemisphere, radiating into numerous niches, the big, dark sparrows of the Pacific Northwest contrasting with their brightly hued eastern kin and their small, sandy-colored brethren of the desert Southwest. In this painting the nesting microhabitat of a decorative brick is shared with a common paper wasp of the genus Polistes.

PHORESY—NEOTROPICAL PSEUDOSCORPIONS & HARLEQUIN BEETLE (2014)
acrylic on panel 14” x 9”

One of the main characteristics of nearly all animals is their ability to move from one place to another. This seems obvious, but it's an important and valuable skill that is much more restricted in other kingdoms. Leaving the home turf benefits an organism by sidestepping competition with relatives, and life has evolved countless interesting ways of dispersing itself. Pseudoscorpions are tiny arachnids, a few millimeters long, that can be found in just about any habitat, nearly anywhere on Earth, although because of their size, most people have never noticed one. Dispersing very far is difficult for such minute creatures. Many small insects can fly. Some spiders, mites and caterpillars disperse by “ballooning,” floating on the breeze from a long strand of silk. Many pseudoscorpions and other small invertebrates disperse by phoresy, or commensally hitchhiking on a larger animal. The pseudoscorpion Cordylochernes scorpioides is habitually carried to better habitat on the back of a Harlequin Beetle (Acrocinus longimanis). Both animals are distributed widely through Tropical America. Like most other long-horned beetles, the larval Harlequin Beetle is a wood borer. After metamorphosis, the adult beetle emerges to the mossy surface of the dead old tree in which it spent its youth. This is the habitat of C. scorpioides, which preys on small arthropods. Upon sensing a Harlequin Beetle, a pseudoscorpion will approach it and pinch its abdomen, causing the beetle to lift its wings and allow the smaller animal to climb underneath them. A male C. scorpioides will defend his patch of beetle back against other male pseudoscorpions, and mate with females there. A large, newly-dead tree is the preferred place for Harlequin Beetles to mate and lay eggs and for C. scorpioides to live, so the ride/rider relationship is an apt match. The pseudoscorpion species Parachelifer lativittatus also regularly rides on Harlequin Beetles, further up, on the sides of the thorax. Beneath the phoretic relationship depicted, a Margay (Leopardus wiedii) crosses the forest floor. 

 PASSENGERS OF FORTUNE--CARMINE BEE-EATERS (2005)
acrylic on illustration board 40” x 15”

Commensal feeding is related to parasitism, but its effects on the host are benign. It's also related to phoresy, but instead of a means of dispersal, the hitchhiker is exploiting a good feeding platform, like the remora on the shark. On land, the riders are usually birds hawking insects disturbed by a large mammal. The striking Northern Carmine Bee-eater (Merops nubicus) of the African Sahel feeds in the typical manner of bee-eaters: searching out flying bees and wasps from a perch, winging out to capture them in midair, then returning to the perch to dispatch and eat them. This species, though, has added a couple of other methods to its feeding repertoire, including diving for small fish like a tern, and riding upon the backs of large mammals and even birds like the Ostrich (Struthio camelus), and flycatching after flying insects like the Desert Locust (Schistocerca gregaria) that emerge from the grasses as the ride passes through. The lines between symbiosis, commensalism and parasitism are blurry, and there is often overlap between them. Commensal feeders like these bee-eaters are also apt to remove the occasional parasite, like the louse fly of the family Hippoboscidae that can be seen scuttling through the ostrich's feathers. These riders are true parasites, specializing in sucking the blood of their host. Many hippoboscid fly species feed only upon a single species of bird or mammal.

CONVOY THROUGH THE CANOPY—deBRAZZA'S MONKEYS (2000)
acrylic triptych on illustration board 30” x 20”, 30”, 20”

This painting depicts a relationship that's related to the commensal feeding of the remora and shark and the bee-eater and Ostrich, but is more symbiotic, that is, both parties experience a benefit from their relationship, transitory though it may be. One of the many African monkey species known as “guenons,” de Brazza's Monkey (Cercopithecus neglectus) inhabits various types of forest, usually near rivers, from southeastern Cameroon through the southern Central African Republic and throughout most of the former Zaire. Here we see a troop moving along a massive fig tree, accompanied by Long-tailed Hornbills (Tockus albocristatus) and Oil Palm Squirrels (Protoxerus stangeri). Both of these species habitually travel with monkeys, eating insects that are disturbed by their movement, such as the giant cicada (Cicadidae) in the right panel. The sharp-eyed hornbills return the favor by making a loud racket if they spot a Crowned Eagle, warning the primates of the presence of an important predator. I've taken some artistic liberties in this piece by depicting such a large group of monkeys so close to a small settlement of Humans, the most important monkey predator of all in Central Africa. Incidental animals in this piece include a Crested Chameleon (Chameleo cristatus), Bush Viper (Atheris hispidus), Black Kite (Milvus migrans), Great Blue Touracos (Corythaeola cristata), Gray Parrots (Psittacus erithacus), Palm Swift (Cypsiurus parvus), Red-rumped Tinker Bird (Pogoniulus chrysoconus), Snowy-crowned Robin-chat (Cossypha niveicapilla), Chestnut Wattle-eyes (Platysteira castanea) and Village Weaver (Ploceus cucullatus).

BLACK AND WHITE AND RED ALL OVER—SPOTTED HYENAS AND PLAINS ZEBRA (1999)
acrylic on illustration board 20” x 17”

Although they resemble dogs, the four species of hyena are most closely related to the mongoose family. By far the largest and most powerful member of the group, the Spotted Hyena (Crocuta crocuta) ranged over most of the Old World during the Pleistocene, but today is restricted to sub-Saharan Africa. Despite this, it is still a highly successful species, outnumbering other large African predators and able to function as a scavenger, pirate, solitary hunter or pack hunter. Its incredibly powerful jaws enable it to crack large bones and exploit the marrow within, a food source inaccessible to other animals. Fetal female hyenas develop with high levels of androgens in their blood, a quirk resulting in their well-known mock male genitalia. Here a pair of hyenas pause over their meal, a Plains Zebra (Equus burchelli). Interestingly enough, it seems that the three animals we call zebras do not really constitute a discrete group, but that all modern equids evolved from a striped ancestor. Some modern forms lost their stripes, while members of two distinct lines retained theirs. This pattern probably does not aid camouflage, but rather works as a social signal to other zebras. And as for the age-old question...I say definitely white with black stripes. In the background is a small group of Bronze Mannikins (Lonchura cucullata). 

CRASH-BARRIER WALTZER—BLACK-BILLED MAGPIE (2005)
acrylic on illustration board 30” x 20”

If nature is anything, it is energy-efficient, and in any ecosystem there is a place for scavengers. In unusual situations, like the Pleistocene plains of North America and some contemporary African savannas, there are enough large animals to sustain full-time warm-blooded carrion eaters, but the majority of larger scavenging animals are generalists, feeding on a wide assortment of foods, and in that respect, western North America's Black-billed Magpie (Pica hudsonia) is typical. Long considered a race of the Old World Magpie (P. pica), the American bird is now considered a distinct species, based on DNA evidence. Thriving in a variety of situations, these handsome corvids are common through most of their range. Despite their routine habit of feeding on road-killed animals, it is surprisingly rare to find one of these intelligent birds joining those ranks. Incidental subjects in this painting include Big Sagebrush (Artemesia tridentata), Mules Ears (Wyethia amplexicaulis), a garden spider (Argiope sp.), Differential Grasshopper (Melanoplus differentialis), Green Stink Bug (Acrosternum hilare), Blue Mud Dauber Wasp (Chalybion californicum), Convergent Ladybird larva (Hippodamia convergens), looper larva (Autographa sp.), Bushtits (Psaltriparius minimus) Green-tailed Towhee (Pipilo chlorurus), and Least Chipmunk (Tamias minimus).

SPOTTED EAGLE OWL (1998)
acrylic on illustration board 30” x 20”

The eagle owls of the genus Bubo comprise a highly successful group of large predatory birds, with representatives nearly everywhere except Antarctica and the Australasian region. The group includes the arctic Snowy Owl (B. scandiacus), the huge Eurasian Eagle Owl (B. bubo) and the widespread and well-known American Great Horned Owl (B. virginianus). Through most of sub-Saharan Africa the Spotted Eagle Owl (Bubo africanus) thrives in most habitats except extreme desert and thick tropical rainforest. It is slightly smaller than its American cousin, and less likely to take large prey, usually preferring a diet of rodents, lizards and large insects. In the background a small party of Spotted Hyenas (Crocuta crocuta) scouts the savanna.

THREE MORE WORLDS— RAINBOW TROUT & OSPREY (1998)
acrylic on illustration board  30" x 20"

The Osprey Pandion haliaetus) is an unusual bird with no close relatives. It shares common ancestry with the hawks, eagles, kites and falcons, but so far it's been difficult to trace its lineage any more specifically than that. Fossils ascribed to the osprey genus go back at least 15 million years, and 30 million year-old osprey-like fossils have been found in Germany and Egypt. Its singularity has earned it its own family, Pandionidae, with but a single species (though some experts consider the Australian ospreys different enough to warrant species status). Few birds have a wider global distribution; ospreys range across every continent but Antarctica. The fact that these birds vary so little across the globe points to their wandering nature. Only a handful of animal species fit this evolutionary pattern; Homo sapiens is another one. It is extremely rare for an Osprey to eat anything that is not a fish, and this behavior has been important in its evolution. Its large, rugose feet with opposable outer toes are unique within its order, as are its nostril valves and its wings, which are structured very like those of other dive-fishing birds like pelicans and gannets: a case of convergent evolution. The Rainbow Trout (Oncorhynchus mykiss) is another creature with an expansive range, but its story is quite different. Originally native only to western North America in waters draining into the Pacific, it has proved to be an ideal species for captive propagation and transplantation as a game fish, and has been introduced throughout the continent and into waters as far away as New Zealand. This painting is in large part an overt theft of Escher's lithograph “Three Worlds,” in which the viewer gazes through leaves floating upon the water’s surface upon a koi beneath and reflections of trees above. I injected an aspect of impending doom by introducing the Osprey's reflection.

YELLOW-CROWNED NIGHT HERON (2004)
acrylic on illustration board  24" x 18"The Yellow-crowned Night Heron (Nyctanassa violacea) haunts coastal marshes from Massachusetts to Brazil, and Panamá to Perú on the Pacific side. It also occurs on the Galápagos and throughout the West Indies. Unlike its nearly cosmopolitan cousin the Black-crowned Night Heron (Nycticorax nycticorax), which hunts fish from dawn to dusk, the Yellow-crowned often stalks the crustaceans that make up the bulk of its diet at midday. In recent years this species has increased its range in the United States as far west as Illinois. Yellow-crowned Night herons also breed on Baja California as well as the eastern shore of the Sea of Cortés. The butterfly on the bank belongs to the genus Eunica.

CRASH-BARRIER WALTZER—BLACK-BILLED MAGPIE (2005)acrylic on illustration board 30” x 20”

If nature is anything, it is energy-efficient, and in any ecosystem there is a place for scavengers. In unusual situations, like the Pleistocene plains of North America and some contemporary African savannas, there are enough large animals to sustain full-time warm-blooded carrion eaters, but the vast majority of larger scavenging animals are generalists, feeding on a wide assortment of foods, and in that respect, western North America's Black-billed Magpie (Pica hudsonia) is typical. Long considered a race of the Old World Magpie (P. pica), the American bird is now considered a distinct species, based on DNA evidence. Thriving in a variety of situations, these handsome corvids are common through most of their range. Despite their routine habit of feeding on road-killed animals, it is surprisingly rare to find one of these intelligent birds joining those ranks. Incidental subjects in this painting include Big Sagebrush (Artemesia tridentata), Mules Ears (Wyethia amplexicaulis), a garden spider (Argiope sp.), Differential Grasshopper (Melanoplus differentialis), Green Stink Bug (Acrosternum hilare), Blue Mud Dauber Wasp (Chalybion californicum), Convergent Ladybird larva (Hippodamia convergens), looper larva (Autographa sp.), Bushtits (Psaltriparius minimus) Green-tailed Towhee (Pipilo chlorurus), and Least Chipmunk (Tamias minimus).

PRAIRIE SENTINEL--PRAIRIE RATTLESNAKE & AMERICAN BISON (2002)
acrylic 15” x 40”

Rattlesnakes comprise about 50 species in two unique American pitviper genera, all with tails that are tipped with a series of complex, interlocking, cornified scales, completely unlike anything else known to have been evolved by snakes—until very recently, anyway. These reptiles are not only specialized at their very tips; the musculature of the tail itself is dominated by three pairs of “shaker” muscles, two of which produce lateral, back-and-forth movements, while the third pair applies torsion, drawing the ventral edge of the rattle outward to either side. The fibers of these muscles are rich in mitochondria, sarcoplasmic reticula, capillaries and glycogen, and capable of sustaining the high respiratory levels necessary to vibrate the tail as rapidly as 100 Hz. for as long as an hour at a time. These speeds are comparable to the oscillations of sphingid moth wings. Among vertebrates, only hummingbirds can vie with the rattlesnakes in this respect. The rattling system's main function is to warn away dangerous animals like predators and large grazing animals, although in some of the small Sistrurus species, it is only audible at close range, and appears to be of little use in this area. Whatever the first proto-rattlers used their tails for, they probably enhanced an already existent behavior. Young rattlesnakes and even adult Sistrurus rattlers often engage in caudal luring, wriggling the tail to entice lizards and other potential prey to come in close. It is possible that early rattles enhanced this behavior. The tail of the recently discovered Iranian viper Pseudocerastes urarachnoides has an ornate tail lure that could be similar to the tails of early rattlesnakes. Then again, the first rattles may have been defensive. Many snakes, including some vipers, vibrate the tail defensively. When doing so against dry vegetation, the resulting sound is not unlike a rattler's. Defensive tail-shaking colubrids, like the Common Racer (Coluber constrictor), lack the specialized tail musculature, and cannot sustain the motion more than a few seconds, but the tail muscles of the Copperhead (Agkistrodon contortrix), a close cousin of the rattlesnake, have a significantly elevated respiratory capacity. The traditional view of rattler evolution posits that rattles evolved to enhance defensive tail-shaking, and since the earliest-known rattlesnake fossils were found in the American Great Plains, it's tempting to visualize the first rattler warding off vast herds of American Bison (Bison bison). Genetic mapping, though, strongly suggests that rattlesnakes first evolved in America's southeast, severely shaking this attractive theory. Today, the rattlesnakes are represented in the American Midwest by the Prairie Rattlesnake (Crotalus viridis). Incidental creatures in the painting include horseflies (Tabanus sp.) a metallic bee (Augochlora sp.), banded grasshopper (Trimerotropis sp.), skipper (Epargyreus sp.), Common Nighthawk (Chordeiles minor), Horned Larks (Eremophila alpestris), and a groundsquirrel (Spermophilus sp.).
 

STRANGE FRUIT--IVORY-BILLED WOODPECKER (2002)
acrylic 30" x 20"

At around twenty inches in length, the Ivory-billed Woodpecker (Campephilus principalis) was North America's largest woodpecker. Never a common bird, it once haunted virgin cypress swamps and bottomland forests throughout the southeastern U.S. By the end of the nineteenth century its imminent extinction was feared, and the last confirmed sighting was in the late 1950's. Sporadic events since that time point to the possibility that some of these birds may still endure: a handful of questionable photographs, numerous unconfirmed sightings, and a 2002 recording originally identified as a drumming male, but later shown to be a distant shotgun. In the spring of 2004, a brief video taken in Arkansas's White River Refuge was widely accepted at first as an Ivorybill, but later dismissed by most authorities as a Pileated Woodpecker (Dryocopus pileatus). Incidental creatures in the painting include a Zebra Butterfly (Heliconius charitonius), a Green Anole (Anolis carolinensis) and Palm Warblers (Dendroica palmarum).

SPRAWL--OUSTALET'S CHAMELEON (2007)
acrylic 18' x 24"

Ecological change is one of the drivers of evolution. Random change is detrimental to most organisms in a functioning ecosystem, but there is frequently a small minority that finds benefit in that change. The island of Madagascar, isolated since the Cretaceous, is famous for its unique flora and fauna, much of which has diminished or been extirpated as humans altered the landscape. Rampant deforestation has been devastating for most of the island's wildlife, including most of its 70+ chameleon species. Two notable chameleon species though, seem to have benefited from deforestation: the Panther Chameleon (Furcifer pardalis) and in particular, the two-foot-long Oustalet's Chameleon (F. oustaleti), a species that thrives in deforested zones and has expanded its range greatly in recent years. Lean and limber, it's an active species whose tongue can snatch small reptiles and even birds along with the large insects that make up most of its diet. In the trees it moves in typical chameleon fashion, but on the ground it can run quite quickly for a chameleon. Incidental species include Humans (Homo sapiens), Madagascan Brown Bat (Neoromicia matroka), Black Kite (Milvus migrans), Chicken (Gallus gallus), Red Fody (Foudia madagascariensis) and Lined Day Gecko (Phelsuma lineatus).

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These last six paintings are reconstructions of exinct fauna of western North America.

FAUNA OF THE BURGESS SHALE (1988-2011)
acrylic on illustration board 15" x 20"

British Columbia's Burgess Shale is one of the best-known and most important fossil beds. This half-billion-year-old formation has yielded an embarrassment of excellent fossils of soft-bodied creatures from what is frequently called the “Cambrian Explosion,” when suddenly—well, relatively suddenly—over the course of about 8 million years, the recently evolved multicellular, or metazoan organisms radiated into a multitude of different body plans, with representatives of most of the phyla that have ever turned up on the planet. For most of the preceding 3 ½ billion years, life on Earth had been unicellular. The cause of this explosion is poorly understood. Is it an illusion suggested by gaps in our knowledge of the fossil record? Was it caused by climatic or geological changes, or the evolution of Hox genes, sexual reproduction, predation or eyesight? Or was it simply a response to opportunities that could only be exploited by multicellular life? How about a combination of factors, or something else? By the beginning of the Cambrian period, 540 mya, stromatolites (1), formations of sediment trapped within layers of cyanobacteria and other microorganisms, had been common features of shallow seas for over two billion years. The appearance of grazing metazoans like trilobites and earlier creatures is thought to have been responsible for a major reduction of these structures during the Cambrian and Ordovician. True sponges (Phylum Porifera) appeared before the beginning of the Cambrian, and are represented here by Leptomitus (2) and Vauxia (3). Brachiopods (phylum Brachiopoda), which would reach an impressive peak of importance and diversity during the Ordovician, made their appearance with the Cambrian, during which they probably remained fairly uncommon. Pictured is the articulate brachiopod Billingsella (4). One of the groups that may have given rise to the arthropods is the phylum Lobopoda, in which many experts include the bizarre Hallucigenia (5), which, until recently, was normally reconstructed walking upon its spines, with a single row of tentacles running down its back. A common Cambrian lobopod was Aysheaia (6), which is thought to have fed on sponges, with which its fossilized remains are frequently associated. Also related to the arthropods was the extinct phylum Radiodonta, which included Anomalocaris (7), a huge swimming predator that reached a yard in length, as well as the five-eyed, nozzle-nosed Opabinia (8). The best known arthropods of the Cambrian were the trilobites, which appeared early on in the period, and enjoyed great success well into the Devonian. Small, primitive agnostid trilobites like Lejopyge (9) proliferated during the middle Cambrian. Though shaped like bottom-dwelling (benthic) animals, their wide oceanic distribution is suggestive of a free-swimming (pelagic) lifestyle. Perhaps they lived on the water's surface, floating on little air bubbles and filter-feeding on microorganisms. The corynexochid trilobites comprised a diverse and successful order, one of the best known of which was Olenoides (10). Another large trilobite order was the Ptychopariida, which included Asaphiscus (11), the very common Elrathia (12) and Modocia (13). Naraoia (14) was an unusual trilobite, if indeed it even belongs in that taxon. Among the arthropods whose taxonomic affinities are unclear are the benthic Habelia (15) and Burgessia (16) and the backswimming Sarotrocercus (19). The crustaceans also appeared during the Cambrian. Included in this group are the backswimming Odaraia (18), the enigmatic phyllocarid Pseudoarctolepis (20), and the crayfish-like Canadaspis (21). One of the most common arthropods of the Cambrian was the lovely little Marella (22), which may have been related to Branchiocaris (23). Members of the phylum Priapulida, very much like the burrow-dwelling Ottoia (24) still persist today. Ottoia fossils have been found containing prey like the hyolithid mollusc Hyolithes (25) (phylum Mollusca). The early segmented worm Canadia (26) (phylum Annelida) seems quite similar to some modern polychaete worms. The echinoderms comprise a large and important phylum that first arose in the early Cambrian. The primitive Gogia (27) was related to modern sea lilies. Our own phylum Chordata was represented in the Cambrian by the wormlike Pikaia (28), which was anatomically similar to modern lancelets. Some of the unusual Cambrian animals whose affinities are hard to place include the common armored Wiwaxia (29), the sessile (stationary) Dinomischus (30) and the pelagic Amiskwia (31).

APATOSAURUS (1997)acrylic 15” x 10” It's a shame to lose the best-known of all dinosaur names, but thanks to a century-old mix-up involving two specimens, one with the wrong head, being described as representing different genera, Brontosaurus was recently abandoned in favor of the precedent Apatosaurus. It was once speculated that the long necks of these sauropods allowed them to browse high in the treetops or even to breath while deep underwater, but closer study found that the neck musculature could not elevate the structure vertically. More likely, it merely allowed them to reach a considerable amount of plant food without having to move their massive bodies, as the painting illustrates. Apatosaurus lived about 150 million years ago, during the Kimmeridgian and Tithonian ages of the Jurassic period. Its fossils have been found in sites in Utah, Colorado, Oklahoma and Wyoming. This painting was originally done as an illustration for Frank DeCourten's book, “The Dinosaurs of Utah.”

BRACHIOSAURUS (1997)acrylic 15” x 10”Unlike its contemporary Apatosaurus, the robust Brachiosaurus could hold its head high, and probably specialized in browsing the leaves of tall trees that were unavailable to most other land herbivores—so much so, that it was probably restricted to well-forested areas. Its front legs were much longer than the hind legs, giving it a steeply-sloping, giraffe-like back. With high nostrils and large, blade-like teeth, its head was distinctive. Very few good fossils from this genus have turned up, all of them in western Colorado. Brachiosaurus lived during the Tithonian age of the Jurassic period—about 145 million years ago. This painting was originally done as an illustration for Frank DeCourten's book, “The Dinosaurs of Utah.”

PARASAUROLOPHUS (1998)acrylic 18” x 15” Towards the end of the Cretaceous period, there seems to have been a marked decrease in biodiversity, with a very few groups enjoying great success. Most successful of all of these were the hadrosaurs, or duck-billed dinosaurs. With its gracefully curved crest, Parasaurolophus was apparently very common through much of western North America during the Campanian age of the Cretaceous, from 76 to 73 million years ago. The function of the conspicuous crest has been the subject of much speculation. Probably it served as a social signal and possibly a resonating chamber for vocalizations as well. In the background, a pair of Avisaurus (a primitive bird) roost together. This painting was originally done as an illustration for Frank DeCourten's book, “The Dinosaurs of Utah.”

ALAMOSAURUS (1998)
acrylic 10” x 15” The sauropods (the well-known group of long-necked, terrestrial dinosaurs)

reached their peak of diversity during the Jurassic Period, but members of the group survived to the very end. One of the very last dinosaurs was Alamosaurus, which roamed what is now the southwestern U.S. during the Maastrichtian age of the Cretaceous period, up to 65 million years ago. Here an Alamosaurus herd moves along the edge of a large late Cretaceous lake.
This painting was originally done as an illustration for Frank DeCourten's book, “The Dinosaurs of Utah.”


 OSBORNODON (2006)
acrylic on illustration board 20” x 17”
 

The first carnivore-like mammals arose in North America during the last days of the dinosaurs, and radiated into a number of forms over the next 15 million years. In the early Eocene, around 50 million years ago, the first members of the modern order Carnivora appeared. Among the first of these were the early canoids, the ancestors to dogs, bears, and the weasel, raccoon and seal families. The first true canids appeared some 10 million years later. These very early dogs belonged to the now extinct subfamily Hesperocyininae, a group that included the longest surviving known dog genus, Osbornodon, which persisted from early in the Oligocene (about 34 mya) until the late Miocene (about 14 mya). It died out about 7 million years before the appearance of the first true dogs of the genus Canis. Six species of Osbornodon have been identified in various sites across western North America. They were fairly large animals, around 30 lbs., with short legs, big heads and long snouts. They probably hunted alone or in pairs. This painting depicts two Osbornodon digging after prey in the sage-covered Miocene hills of western North America.