EDITOR Ron Skylstad Leaf Litter VOLUME 2 | ISSUE 2

ASSISTANT EDITORS Ed Kowalski 1 FROM THE EDITOR Nathanial Paull 3 EDITORIAL Jason Konopinski 5 FOLIUM 9 AMPHIBIAN HEALTH & NUTRITION

LAYOUT Elizabeth Brock FEATURES 12 Herping In Anytown, USA • 15 Further Impressions from Tarapoto: EXECUTIVE DIRECTOR Frogs and Art Ron Skylstad

19 When Father Chimborazo Took PROGRAMS DIRECTOR the Frogs Away Brent L. Brock 29 Notes on the Husbandry, Captive Reproduction, and Distinct Juvenile DIRECTOR OF PROJECT SUPPORT Form of Wallace’s Flying Frog, Marcos Osorno Rhacophorus nigropalmatus

MISSION STATEMENT 41 Solace of Untouched Wilderness Tree Walkers International supports the protection, conservation, and restoration of wild amphibian populations through hands-on action both locally and internationally.

We foster personal relationships between people and nature by providing opportunities for citizens of all ages to become directly involved in global amphibian conservation.

Through this involvement, our volunteers become part of a growing and passionate advocacy for the protection and restoration of wild amphibian populations and the environmental on which they depend. COVER Rhacophorus nigropalmatus photo © Michael Ready from the editor

When one tugs at a single thing in nature, he finds it attached to the rest of the world. - John Muir -

Connection.

If we have eyes to see it, it’s everywhere: in the roles organisms play in their ecosystems, in the way our habits and decisions impact our environments (for better or worse) and the species that inhabit them, in the amphibians we choose to keep and the systems of trade and collection we inevitably support through our buying habits.

This thread of connection, of interrelatedness and influence, is woven throughout this issue of Leaf Litter.

In this issue we are introducing two regular installments. The first “Folium,” focuses on the topic of plants, including their natural history and physiology, and how we can use that information to successfully culture them in captive enclosures. The second, “Amphibian Health and Nutrition,” deals with the various health and dietary issues regarding the captive care of amphibians.

In his editorial piece, Devin Edmonds reminds amphibian keepers that their animals do not originate in a vacuum and that their worth cannot simply be reduced to a price tag. He challenges hobbyists to expand their horizons of knowledge and awareness regarding the animals they keep.

In an essay, I describe my personal experience on a field trip at the 2008 North American Amphibian Conference that was held in the Berkshires of Massachusetts. This was my introduction to both this particular region of the U.S. as well as the amphibian species that live there, and I returned home with a deeper level of insight that has changed how I understand my own native region and its amphibian species.

Tracy Hicks concludes his three-part series about a trip he took to the Tarapoto region of central Peru. In this final installment he shares his personal impressions and catalytic moments of inspiration, those “Oh, wow!” instances of understanding and connection. Through his experiences, he shows us a deeper glimpse into the region from which many captive dendrobatids originate, including the people who live there and the jungle upon which they depend.

Based on first-hand experiences with the species, John Simmons provides a detailed account of the now extinct Atelopus ignescens, looking not just at the niche it once filled in the high paramos of Ecuador, but also the cultural and social connections the people of this country have with it.

Mike Ready shares with us his observations and notes from the first-known success of captively breeding Wallace’s Flying Frog, Rhacophorus nigropalmatus. This account not only provides much insight into the captive care and reproduction of this species, but reaches back to Alfred Russel Wallace himself—over a century and a half after his original description of this frog, we discover that we still have much to learn about its habits and morphology. >>

from the editor | 1 Wrapping up this issue is an essay by high school student Emily Lisborg, who takes us along with her as she shares about a particular visit to a tropical forest on the island of St. John in the U.S. Virgin Islands and the deep sense of connection she experienced there.

It’s all connected: our desire for amphibians affects their wild populations and native habitat, affects the people who live there and how they interact with amphibians. Our relationship with amphibians can influence and shape our culture, from art to music to community festivals. Our understanding of their habits can help us more greatly appreciate them and the ecosystems in which they are found. Our knowledge of their care and reproductive habits can help us in the quest to more successfully maintain viable captive populations and put less pressure on the collection of wild ones. Our insight into their health and the mini-universes of microbes and bacteria that exist within them, as well as of the plants they naturally associate with in the wild, increases our understanding of the incredible and integral role these animals play in the biosphere.

We can tug on a single frog, but when we do, we should be prepared to find it inextricably attached to the rest of the world.

Sincerely, Ron Skylstad Editor-in-chief

2 | Leaf Litter :: Vol 2 Issue 2 Breaking the Glass Box an editorial by Devin Edmonds

Poison dart frogs seemed out of place in Joliet. As one of held less weight. When I had the opportunity to go herping the fastest growing cities in Illinois, urban sprawl had wiped outdoors, the creatures I found under logs and in ponds were clear any former wetlands in the area. On the drive into town a separate type from the kind being sold in pet stores. They I didn’t pass through any lush tropical rainforests—the tree- came without a price tag, belonging solely to the world. In like telephone poles didn’t hang with bromeliads or other my mind, these wild amphibians were unrelated to the caged epiphytic plants. Leaf litter blew across Joliet’s streets, but ones at the store. They were a different sort. was outnumbered by fast food wrappers and plastic bags, The connection I originally failed to see between captive likely tossed out the window of passing cars. Nevertheless, amphibians and their wild counterparts eventually made in a place where concrete dominated the landscape, amongst itself known. I traveled abroad, photographing frogs in the White Castle and Wal-Mart, were hundreds of the world’s wild that I kept at home in terrariums. In Peru, I saw the effect most elegantly colored and exceptionally rare amphibians, on that smuggling has on wild frog populations. Here too, in the display and for sale at a Holiday depths of the Amazon rainforest, Inn. animals were equated with money. Midwest Frog Fest is held I also travelled in the U.S. to visit annually at a hotel in Joliet, large scale reptile and amphibian and the contrast between the importers. Wild-caught frogs conference room and the tropical came in by the boxful to these life contained within strikes me warehouses of live merchandise, every year. Dozens of species and were quickly piled into stock of captive-bred exotic frogs are tanks to await shipment to pet present, their bright aposematic stores or middlemen dealers. coloration radiating from vendor These wild frogs would end up tables like ornaments on a in the hands of hobbyists like me Christmas tree. The 2008 show who, in addition to supplying the had the most diverse selection demand that supports the trade, of species yet, with tree frogs, also produce the fine captive-bred toads, and newts alongside the offspring found for sale at frog usual dominating presence of shows. I mentally followed the poison dart frogs. As always, I was trail of these animals from their struck by the beauty of the frogs, and I could see that other origin to the terrariums in my bedroom, from the wild to attendees shared my experience. I have to wonder, though: captivity. The connection became clear. when amphibians are priced as goods at a frog show in a So what’s the point of keeping frogs when we are so hotel, how easily can they be viewed as part of the natural disconnected from nature that we can only appreciate their world? The disconnect I saw here should be obvious to all beauty inside a glass box? Where does this leave us? Those who attend this event, but I’m not so sure it was. When I who are not involved in the exotic animal hobby might believe first started attending reptile and amphibian shows, it wasn’t a person should have to get mud under their nails in order to apparent to me, either. view creatures as striking as the frogs we keep in our living I grew up observing nature in cages. Weekend trips to rooms. They might feel that we must climb mountains and the pet store were routine throughout my youth. As a child walk trees over rivers, get bitten by ants and stung by bees. conditioned to view exotic animals primarily as a fascinating I don’t disagree with this view. Although we can learn about commodity, a captive amphibian’s monetary value and care a frog’s behavior in captivity -- watching them breed, feed, requirements were most important to me. Information or defend territories -- to truly know a species we must also about their natural history, ecology, and conservation status experience the natural environment in which they live.

Breaking the Glass Box | 3 Being in the field not only gives hobbyists a new appreciation for the amphibians we keep, it also liberates our minds from the concrete boundaries most of us live within daily. While searching for frogs outdoors, our senses focus in a new way, allowing us to finely tune in to the natural world. It’s in this state of mind where the sterility of the Joliet- environment is allowed to dissipate from our systems, and as a result, we can develop a new attitude about the amphibians we keep. Many have said that the terrarium serves as a way to bring the beauty of the natural world into the home. However, I see the terrarium as serving a much larger role than pure aesthetics: it invokes an interest in nature. I have watched people who were raised in urban environments travel to remote natural places as a result of their involvement in the exotic amphibian hobby. Those who take the time to do so usually come back with a different understanding of their amphibians. The mentally projected dollar signs on their tanks sometimes dissolve, being replaced by a true admiration for the tropical creatures that dwell within. Perhaps this sort of experience should be a prerequisite, viewing a species in the wild first before keeping it in captivity, reversing the role of the terrarium in the home by first creating an interest in nature. Those of us involved in this hobby have one thing in common: our interest in amphibians. We are a diverse group with varied backgrounds. As we progress in the hobby, our views tend to change as a result of communicating with other hobbyists. Unfortunately, I have too often seen this play itself out like a funnel, with attitudes of genuine interest in an animal directed toward the shallow fulfillment of collecting exotic species as if they were stamps. This effect can be reversed, but may require the temporary abandonment of the conveniences of urban life. Get outside and into the field. If you can’t afford a trip to the places your exotic frogs originate, find a nearby pond to visit each spring and watch your local Rana or Bufo species mate. Doing so may be the only way to prevent our hobby from fully becoming like the wetland- devoid, chain store-dominated, urban landscapes in which our frog shows are hosted.

4 | Leaf Litter :: Vol 2 Issue 2 folium The Allure of the Aroid by Richard Revis photos © Mike Wallitis

So, what is an Aroid? In the woodlands while others are at home are shared between this environment simplest of definitions, it is any plant in hot, humid, lowland rainforests. and that of most terrariums. One is in the family Araceae, which includes Their diversity and adaptability are the amount of light, or lack of it, that some of the more commonly known just a couple features that make many each receives. Light is at a premium in plants such as Philodendron, Monstera, members of the family equally at the rainforest and the canopy plants and Pothos (which is not really what home in the terrarium. Several other are in a constant battle to capture as most people are thinking of right now, attributes also add to their suitability much of it as they can. The result is but more on that later). and, as we discuss more details of their that very little of it reaches the forest natural history, will help you make floor. In order to survive down below, One of the most noticeable features decisions on what will work best for plants employ various strategies to that all Aroids share is the bundled your particular needs. use what little light does make it down up cluster of flowers known as the to them. One trick, actually used by inflorescence, consisting of two main For the purpose of this article many plant families, is to have leaves parts, the spathe and spadix. The we are most interested in terrarium that use light as efficiently as possible spadix is usually a cylindrical tube- compatible species, so we will focus by not only collecting it from above, shaped appendage in the center of the on those Aroids from the tropics. A but also reflecting it back up into the inflorescence that contains anywhere great many species are typically found leaf from below. The red and purple- from a few to several hundred minute growing in the forest understory. Some hued pigments found on the underside flowers. Amazingly, in some species, will spend relatively little time there of the leaves of certain species are the spadix is capable of generating heat while making their way to the top; particularly adept at accomplishing to help disperse pollinator-attracting others will never leave the forest floor. this. Leaves with this pigment on odors, or in the case of skunk cabbage, There are several characteristics that to help melt it’s way through the snow! Around the spadix is the shield-like An example of an Anthurium species utilizing pigmentation to increase light absorption. spathe. This is a modified leaf that can vary tremendously in shape, size and color depending on the species. It is the bright white “shell” on the common “Peace Lily” (no, it’s not a lily and lilies are not Aroids!) and the deep purple/brown “vase” on the Titan Arum—the giant that smells like a dead elephant. The variations in the combination of spathe and spadix can take on an alienesque appearance, the beauty of which is only exceeded by the incredible variety of foliage shape and pattern. The family of Aroids is indeed a vast one, encompassing many genera and thousands of species. It is a diverse group, being distributed throughout both the new and old world, and can be found in a wide variety of habitats. Some species thrive in cold, temperate

Folium | 5 the reverse are referred to as being Rhaphidophora have at least two then starts its marathon climb. The ‘discolor’. The result is that the plant distinctive growth habits that are so vine soon starts producing flattened can then gain more energy from the different from each other that one leaves pressed tightly against the same amount of light than one without would think they were completely trunk of the tree, appearing much like these pigments. Some examples of this different plants. It all begins with a shingles on a house. These leaves can include Syngonium erythrophyllum seed. Upon germination, the seedling be as small as a thumbnail or as wide as and Alocasia imperialis. Examining a sends out a wiry, often leafless vine a small dinner plate, and are sometimes plant’s leaf can actually tell a lot about that grows away from the light (this is splashed or veined in bright silver. what it’s requirements are, but in this called negative phototropism, and is a This shingling habit accomplishes case, if the leaves are discolor, you can fancy term that simply means it grows several things. First, by holding the be pretty well assured that it will do towards shade). So why would a plant, leaves flat against the tree they do not well in environments with lower levels in its search for light, grow toward the block any of the precious light from of light. dark? Because in this case, dark means reaching those further below. Second, Another strategy to deal with light. By growing towards a shadow, by not hanging out in mid-air the plant low light is even more amazing. the seedling is hoping to run smack dab prevents itself from becoming a ladder Many species of Monstera and into the side of a tree. When it does, it for other, faster growing vines that could otherwise out compete it. Lastly, the leaves can help trap moisture and Anthurium crystalinum nutrients for the roots along the stem. Once the vine gets close to the canopy a miraculous metamorphosis occurs: the stem of the vine starts growing much larger in order to support the massive weight of the mature foliage. The new leaves are now held away from the tree trunk and begin to resemble a more typical leaf. Thick, rope-like roots are thrown out and firmly attach to the bark of the tree to further stabilize the plant. Now, up where light is plentiful, the plant begins to unfurl huge leaves, often over three feet long and nearly as wide, to capture all of the new found energy. These canopy leaves typically have windows, or fenestrations, in the surface to allow light to penetrate down to the lower ones as well. The mature plant soon begins blooming and will set fruit to start the whole process over again. In the terrarium, this shingling habit can be used to great advantage. Even species that produce mature leaves bigger than the terrarium itself can be successfully grown in their juvenile stages. There is simply not enough space within the terrarium for them to mature and they are relatively slow enough growers and easy enough to prune that they can make excellent

6 | Leaf Litter :: Vol 2 Issue 2 additions. Another common trait shared by the two habitats is a predominance of moisture. Obviously, it rains a lot in a rainforest. All of this moisture can lead to fungal and mold infections on the leaves if they are not quickly shed. As the water drips from the canopy it collects near the forest floor. Here, there is much less air movement, so leaves stay wet longer than those higher up. The soil is frequently saturated or at least consistently moist. In order to survive here you need to be able to tolerate this moisture or you die. Similarly, many terrariums are typically moist at all times and, depending on how frequently they are sprayed, A small Central American species of Aroid. can even be downright wet. The fact that many Aroids do so well under habits makes it possible to fill a tank blue-grey, heart shaped leaves with these conditions is reflected in their with nothing but Aroids and still have darker blue veins. streamside habitats and, sometimes, a very interesting landscape. While even submerged growing conditions in it would be impossible to cover all of the wild. the terrarium worthy varieties, below Syngonium Although many species grow is a brief summary of some selected Most species in this Genus simply near the forest floor, there are many genera along with their attributes and grow too large too quickly to use in that are also found growing as challenges. all but the largest setups. The small epiphytes higher up in the canopy. distance between leaves makes them Even those that may start life out on difficult to prune to a desired size. the ground may eventually abandon Philodendron Some of the vining species, such as S. their terrestrial roots and become One of the most commonly grown rayii and S. erythrophyllum do much completely epiphytic in their mature groups, Philodendron contains many better and are easier to keep in scale stage. Others may never see the forest worthwhile species and varieties. with regular pruning. These two species floor, their seeds being distributed in Often, even large growing species can are particularly suited to low light and the droppings of birds that fall upon be successfully grown in the terrarium easily tolerate wet conditions. the mossy branches where they soon as they stunt easily and will stay small germinate. Likewise, in the terrarium, within its confines. It is not until they some types will thrive when planted on are removed and given ample space Pothos moss wrapped branches or pinned to that they will begin to enlarge. Many No, this is not what you are organic backgrounds where their thick do very well grown hydroponically, thinking. The plant sold in every roots can take hold. The sight of bright so their use in rain chambers or more nursery, department store and home white aerial roots dangling in mid-air specialized setups is of particular improvement center as “Pothos” is certainly adds a tropical look. interest. Easy to propagate from not a Pothos, but actually a species of The variety of Aroid foliage types is cuttings, they will typically produce Scindapsus. True Pothos can make virtually unmatched anywhere else in ample stock for use in additional ideal terrarium candidates but are the plant kingdom. It would be difficult enclosures. One of the most interesting difficult to find and a bit trickier to to find such striking patterns, shapes Philodendrons to grow in the terrarium propagate. The smaller species have and sizes in any other family. The is the beautiful, but rare, Philodendron nice little leaves with winged petioles variation in appearance and growth ‘Burle Marx Fantasy’. In the terrarium (the leaf stem) that gives the plant a it is a vining plant that produces small, very unique look.

Folium | 7 Scindapsus apart. With few exceptions, the main they are kept moist at all times. species used here are in their juvenile I often wonder what there is more With so many choices available it shingling stage. Monstera dubia, R. of in the US: Scindapsus (Pothos) is easy to find something that will work cryptantha and R. pachyphylla are ‘Marble Queen’ or Kudzu? Although it well in any terrarium. Aroids tend to some of the best. Rhaphidophora is not my favorite plant, what I do not be thought of as the quintessential tetrasperma is a non-shingling vine particularly like about it is what can “jungle” plant and are an easy and that looks very similar to a miniature actually make it ideal for some uses. beautiful way to add a tropical touch Monstera deliciosa (Swiss Cheese If you are looking for a plant that will to your enclosed habitat. Their ease Plant). Another great selection is M. grow rampantly, quickly fill a tank, of culture and overall hardiness siltepecana. and will grow in almost any substrate makes them the perfect choice for any (except perhaps a bed of hot coals… enthusiast. although I‘m still not 100% convinced Alocasia & Colocasia on that one), then this one is for you! Another genus known for Seriously, it can be very useful in rain absolutely stunning arrowhead- chambers or quarantine tanks where shaped leaves, it is important to make you need something that will grow selections carefully as some can grow quickly with minimal care and is so to quite an enormous size. Since the prolific it can even be thought of as growing point is actually at or just disposable. There are other members below ground, pruning is not an of the genus that have attractive leaves option to keep them under control. and do grow a bit slower, making for They either mature in the terrarium more behaved occupants. Still, keep a or are removed. Alocasia reginula pair of shears tank-side just to be safe. “Black Velvet”, A. imperialis “Elaine”, A. reversa and A. infernalis are some Anthurium excellent choices to start with. Most of them will do well in a 16-18” tall tank A magnificent genus with some of or larger. The leaf textures and patterns the most stunning leaf patterns of the can be amazing. Alocasia guttata family. Shimmering, jewel-like veins “Bullata” has deeply quilted leaves and over velvety leaves, highly textured matures to less than 18” tall as well. and quilted foliage, attractive blooms and colorful fruit—all of these can be used to describe these amazing plants. Anubias & Cryptocoryne Unfortunately, some species simply While not even similar from a grow too large for most enclosures. botanical perspective, they are used in However, there really is something very similar manners in the terrarium. beautiful for any size tank. Anthurium Both typically grow as seasonal crystallinum, A. clarinervium, A. emergents in their native habitats gracilis and A. scandens are all worth where their roots are always submerged growing. But, being semi to fully and the leaves are either submerged epiphytic in nature means that most or emergent depending on the time won’t tolerate a lot of wetness. of year. The small species of Anubias such as A. coffeefolia and A. nana are Monstera & Rhaphidophora perfect for shorelines or small pool areas. Cryptocoryne wendtii and C. I have combined these two as lucens are also good performers. The they are very similar from a terrarium scarlet red inflorescence of C. lucens is growing perspective and, unless particularly pretty. Anubias will also you have a microscope to examine grow well as full terrestrials as long as An example of the shingling growth habit certain details, can be difficult to tell utilized by many species of Aroid.

8 | Leaf Litter :: Vol 2 Issue 2 amphibian health and nutrition Choosing a Vitamin Supplement by Ed Kowalski

amphibians. However, it may be better to use a pesticide- When keeping amphibians in captivity a correct diet is free or organic certified feed as some of these feeds have one of the keys for long-term husbandry success. However, been shown to interfere with reproduction in some reptiles as the understanding of the needs of the animals increases due to trace pesticide or herbicide residues. In general, the it becomes apparent that a correct diet involves more than better the diet fed to the invertebrates, the more complete the items fed to the animals. There are other factors that the nutrition the animals receive. should be taken into consideration, such as vitamin and mineral supplementation; in fact even the light that an There are two possible ways to supplement the vitamin animal is exposed to can have an effect on the nutritional and mineral requirements of the animals: dusting and requirements of the animals. gut loading. While some people use one to the exclusion of the other there is significant benefit in overlapping the The dietary needs of amphibians have been little studied two methods. Originally gut loading referred solely to the but are believed to be similar to other animals. The reason attempt to adjust the calcium-phosphorus ratio of the feeder that the information is lacking is due to the fact that the insects. However, in today’s herpetological community nutritional needs would need to be tested by withholding the this phrase has come to mean any attempt to modify the tested nutrient and then analyzing the effects on the animal. nutritional content of the feeder invertebrates. All of the While there is an increase in the interest in specific dietary other than commercially used invertebrates (except for supplements for amphibians there is still a little guesswork earthworms maintained in a calcium-rich soil) have negative involved in the nutritional needs of the animals. At this calcium:phosphorus ratios. (The calcium to phosphorus time the majority of the commercially available nutritional ratio should be at least 2:1, improper calcium:phosphorus supplements are based on studies performed on domestic ratios are some of the more common causes of problems animals. However, even with this drawback, the majority of with calcium metabolism). The problem with trying to amphibian nutritional needs appear to be within the ranges adjust the calcium:phosphorus ratio through the use of expected for other animals (with some relatively recently an increased calcium diet is that it often does not achieve noted exceptions such as the higher vitamin A requirements a positive calcium:phosphorus ratio (it can reach positive of Wyoming Toads (Bufo baxteri)). calcium:phosphorus ratio in pinhead crickets). High calcium Nutrition is fundamental to amphibian husbandry insect diets need to be the sole diet offered to the insects for and dietary considerations need to be addressed early to at least 48 hours prior to being offered as a food item. In care for amphibians successfully. All of the commonly addition the insects need to be kept at 27 °C (80.6 °F) for the available commercially raised invertebrate species are not best result of the diet. properly balanced for use as a complete diet for amphibians If there are any other food items available, the insects and/or reptiles. These invertebrates need to have their will avoid consuming the high calcium diet negating any mineral content adjusted to insure a more complete diet. possible advantage to the high calcium diet. Additionally, All purchased feeder invertebrates should be offered a high calcium invertebrate diets will rapidly cause the death balanced diet for at least 48 hours before being offered as a of the invertebrates if water is withheld for even a few food item to insure that the invertebrate has had a chance hours. In any case high calcium insect diets begin to cause to replenish any lost proteins, fats and minerals that may increased mortality in the feeder insects after 48 hours so have been lost before purchase. Many different diets can the insects need to be fed to the animals as soon as possible be used to accomplish this, although the use of cruciferous after 48 hours to prevent the loss of the insects. In some vegetables and/or spinach should be avoided. Cruciferous amphibians increased calcium in the insects is apparently vegetables can cause problems such as goiter, and spinach not metabolized and may not be of any use to the amphibian. contains oxalates that can interfere with the metabolism of Another use of the phrase gut loading surrounds feeding the calcium. Many people use chicken mashes to feed crickets chosen insects a nutritious diet to cause the nutrients in the and other invertebrates that are to be used as food for their

Amphibian Health and Nutrition | 9 gut to be passed on to the consuming amphibian. This is also and may be over supplemented. Hypervitaminosis of A can a little more limited for increasing the nutritional content result in secondary hyperparathyroidism (which is often of the insects as the gut contents of the insects often have called metabolic bone disease) and may be evidenced by a rapid transit time. Many food items can be employed to deformation of the bones including those that are required gut load insects; however as noted above food items need to for feeding. Without proper treatment the eventual death of be offered for at least 48 hours to have any benefit. A better the animal is assured and in any case the deformities will be method for adjusting the vitamin-mineral content is by permanent. Other symptoms of hypervitaminosis A include dusting the insects with a balanced vitamin-mineral mixture. depression, anorexia, vomiting, sloughing skin and bone The major drawback to dusting the insects is that the dust lesions. Hypovitaminosis of vitamin A has been implicated either wears off or is groomed off the insects within several in “short tongue disease” in some amphibians such as spray hours of the insects being introduced into the enclosure. toads, Wyoming toads and possible Panamanian Golden If dusting the insects to provide mineral and vitamin frogs. Initial symptoms can include lethargy, anorexia, supplementation, then the maximal number of insects and weight loss (note the overlap in symptoms between should be the amount the animal can consume within half an hyper and hypovitaminosis symptoms both in A and D3). hour to maximize the nutritional value of the supplement. In severe cases hypovitaminosis A may show itself as an There are several important items that need to be closely inability to capture food. One of the possible early symptoms monitored when choosing a vitamin-mineral supplement. of improper calcium metabolization may be seizures or The first is that the supplement contains vitamin D3, as tremors in the animal. If these symptoms are noted then this is the only isomer of vitamin D that can be used by the supplementation of the insects should be reviewed as amphibians and reptiles. Many of the supplements that are soon as possible in addition to the need to consult with a manufactured for animals other than reptiles use vitamin veterinarian for support therapy. Hypovitaminosis D3 is D2, as it is much less expensive, so if the supplement does one of the problems that are often referred to as metabolic not specify D3 then a different supplement needs to be bone disease (or MBD) however it must be remembered that chosen. Additionally the ratios of vitamin A to D3 to E need this is a term that covers a variety of different conditions to be as close as possible to 100 to 10 to 1. If the ratios of that present the same symptoms. Hypervitaminosis of D3 these vitamins are significantly different from the above is also very dangerous to the animals as it can cause soft ratios then the vitamins can interfere with the uptake of tissue calcification and often shows symptoms of depression, the other vitamins. For example if the ratio of vitamin A is anorexia, and weight loss. Just as too much of some of the too high then it will compete with vitamin D3 for uptake vitamins are dangerous to the health of the animals, calcium resulting in insufficient vitamin D3 for proper calcium also has a narrow range of tolerance as a dietary supplement. metabolization. (However, in recognition of the toxicity and Obviously too little calcium can cause serious problems with problems associated with vitamin A, many supplements are amphibians but too much calcium in the diet has been linked using beta carotene, which is a much less toxic precursor to conditioned deficiencies of some trace minerals and if from which the animal can synthesize vitamin A). The the diet is high in fats, the animals will selectively target soil second concern is that many of the different vitamin-mineral dwelling arthropods that have positive calcium:phosphorus mixtures will not only oxidize over time but will also catalyze ratio (as many soils are calcium rich). an increased rate of oxidation in the mixture. Because of this Intimately linked, but often overlooked, to the concern only the freshest vitamin and mineral supplements nutritional health of amphibians are the different should be used on the feeder insects. Look for a vitamin- wavelengths of ultraviolet light. Ultraviolet light is broken mineral supplement that has either a “born on date” or down into three classifications based on wavelength. The an expiration date to ensure the freshest supplement are three types are ultraviolet A (UVA), ultraviolet B (UVB), and used. Some of these mixtures have a viable shelf life that is ultraviolet C (UVC). UVC has a wavelength of 100-280 nm measured in months due to oxidation. Exposure to light and and is used for its germicidal properties. If possible plants high temperatures will speed this process so all vitamin- and animals should never be exposed to UVC as UVC is mineral mixtures should be kept in a cool dry location and damaging to living tissue. UVA has a wavelength of 315-400 replaced on a regular basis. nm and may be important for its effects on behaviors. Birds, There is a risk of over supplementation as well as under and possibly some amphibians and reptiles may be able to supplementation of many of the important vitamins and see in the UVA spectrum. Frogs can see in the blue end of the minerals, both of which are dangerous to health of the spectrum lending credulance to the possibility that frogs can animal. Vitamins A, D3, and E are all fat-soluble vitamins possibly see in the UVA portion of the spectrum. This

10 | Leaf Litter :: Vol 2 Issue 2 ability may be important, as some of the feeder insects may Husbandry (2001 Krieger Press) is highly recommended as be more visible in this end of the spectrum due to pattern the pertinent information is beyond the scope of this article. reflecting the UVA as has been shown in birds. UVB is very In any case there should never be so many insects in the important metabolically as UVB is used to convert provi- enclosure that the number of insects stresses the animals as tamin D to vitamin D3. If there is sufficient UVB, then the this can prevent the animal from feeding. Additionally, some animal will only synthesize as much D3 as is metabolically insects (such as crickets) are known occasionally to predate required by the animal. This way the keeper does not have upon other animals if there are too many in the enclosure to worry that the use of UVB will increase the chances of and food or water for the insects is not provided. hypervitaminosis D3 as if there is sufficient vitamin D3 in The first reaction when encountering a thin or starved the diet the animal will not synthesize any excess. animal is to feed it to satiation upon a calorically dense food There is often a significant amount of discussion as to item such as waxworms in an attempt to restore the animal which food items are the most nutritious. Proper nutrition is to a better weight. Depending upon how far the starvation not only important for the long term health and reproductive has progressed this tactic could potentially cause the death success of the animals - at least one study demonstrated of the animal through refeeding syndrome. Refeeding syn- that proper nutritional health of the adult frogs may be one drome occurs when the condition of the animal is so poor of the most important items in preventing the development that blood concentrations are maintained within normal of spindly leg in offspring (personal communication ranges by lessening the levels of electrolytes within the cells. Wisnieski). Only a few of the commonly used feeder A sudden influx of food items can cause a loss of the electro- invertebrates have had any significant dietary analysis lytes in the bloodstream as the electrolytes are used to assist performed to determine the quality of the food items. the transport of nutrients into the cells. This sudden loss Of the most commonly used feeder insects the following of electrolytes after feeding can result in rapid death of the breakdowns are readily available: animal and could be an explanation for the sudden deaths

Energy (kcal/gram) Protein Fat Carbohydrate Calcium Phosphorus as fed % kcal mg/kcal Crickets (Acheta domestica) 1.0 40 54 6 0.3 2.7 Waxworm (Galleria mellonella) 2.1 27 73 0 0.1 0.9 Mealworm larvae (T. molitor) 2.1 37 60 3 0.1 1.2 Fruit Fly (D. melanogaster) 5.12* 70.1** 12.6 n/a 0.1 1.05

* This is total energy and is not corrected for non-digestible materials such as chitin.

** This includes chitin, which is only partially digestible by the majority of insect feeders (closer figure as indicated by the author of the reference is approximately 43%/kcal protein).

The amount of food required to keep an amphibian healthy of newly imported animals that are thin. To avoid these depends upon many factors including health, reproduc- complications small meals fed daily can help to improve the tive status, stress, and temperature. All of these items can condition of the animal. After a period of time of the smaller increase the metabolic rate of the animal (and in the case of meals, the quantity of the food can be increased slowly. temperature, lower it) past the expectations of the keeper. Once the amount of food approximates a normal meal then, These multiple factors leading into the amount of required depending upon the species in question, the frequency of the food have led many people to act as if feeding the required meal can be decreased to approximate the normal husband- amount of food is similar to an art form that needs to be ry applied to the species in question. learned as the keeper attains more practical experience. In many respects this is true and can be readily learned through A future article will cover not only some recent advances practical observation of the animals, however the base meta- in metabolism of vitamin A as retinol, but will also discuss bolic needs of the animals can also be determined through carotenoid metabolism. some baseline equations. If the keeper is interested in de- termining the calculated values for the animals in question then reviewing the data provided in Reptile Medicine and

Amphibian Health and Nutrition | 11 Herping in Anytown, U.S.A. by Ron Skylstad photos © Adam Butt

intense than the evergreen tones than be a swinger of birches, I noticed We met in the parking lot of I was accustomed to seeing back a few people from the group excitedly a bowling alley somewhere in rural home—maples and ferns and various crouching down in the middle of the Massachusetts. other woody plants nearly caused road. My daydream interrupted, I me to squint as they exploded in walked over to observe our first find The scene itself, especially to those chlorophyllic brilliance, each leaf of the day, a juvenile Eastern Newt passing by, probably looked a bit and frond unfolding in dramatic (Notophthalmus viridescens), called a shady: a menagerie of people gathered anticipation of early summer. My “red eft” at this stage. This specimen beside two large passenger vans (with eyes darted quickly to and fro as we would be the first of many efts we tinted windows no less), shoulders walked around the gate and proceeded would encounter that day, both on the shrugged against the damp morning down the damp dirt road, trying to road and off, their brilliant orange skin air, quiet conversation accented by identify the various unfamiliar species signaling like a beacon amongst the the occasional outburst of raucous of plants we passed. The woods dark leaf litter of the forest floor. laughter. Had someone stopped and would occasionally open up to frame asked what we were doing there, the picturesque meadows lined with hand- Before long we made our way to a explanation probably would have stacked rock walls, and I wondered shallow pond surrounded by slick sounded unbelievable: we were looking if Robert Frost himself hadn’t had a muddy banks. In decaying leaves for amphibians. hand in repairing at least one or two of around the pond we found egg masses them, placing each stone end-to-end of Ambystoma maculatum. One of my Well, not exactly at that moment. We and quietly disagreeing with the idea favorite finds at this location was an were simply getting ready to look that good fences do, in fact, make good adult Green Frog (Rana clamitans). for amphibians. The field trip we neighbors. As we continued down the For someone who grew up catching were about to embark upon was the road, so did my thoughts. Then, just small chorus frogs in rural Washington opening event for the 2008 North as I was deeply considering whether State, to see something of this stature American Amphibian Conference or not one actually could do worse and shape, the metallic sheen of its (NAAC). Having never actually visited the area of the country we were in, I One of many red efts found during our excursion. wasn’t quite sure what to expect. To be honest, I am relatively unaware of the amphibian species that reside outside of my native home of the Pacific Northwest, so I wasn’t even sure what I was supposed to be looking for. Luckily, however, it had rained for much of the night and the clouds still hung heavy in the sky. The air was warm and humidity was high. One would have to have been pretty hard pressed that day not to find some sort of frog or salamander.

We parked the vans in a gravel lot, completely surrounded by deciduous woodland. The bright shades of green before me were much more

12 | Leaf Litter :: Vol 2 Issue 2 Left and right: sub-adult Ambystoma maculatum. green skin and bronze accents…it sat brew. My guess is a particularly nasty something!” was all I could muster there before me as the quintessential combination of all three.) saying as I began to realize that before anuran, the thing one thinks of when Around mid-morning we found me was yet another type of salamander. they hear the word “frog.” We lingered ourselves perched on a rocky Like the other species we had seen that about around the pond, excited by outcropping that allowed a view of the morning, I had no idea what this one these new finds and quietly wondering valley before us. We used the time to was, exactly—I was simply happy to amongst ourselves whether the stink take a short break, posing for photo have found an amphibian on my own. in the air was from decomposing pond ops and asking the locals from our However, judging by the excitement of muck or perhaps a particularly stealthy group to identify various plants and the few people who were now gathering member of our party. provide some insight on the abundance around me, it seemed I had discovered and commonality of the amphibians something quite special. In my hands We eventually began making our way we had discovered so far. Some of was a Yellow-Spotted Salamander back to the road, which at this point us were overjoyed to find several (Ambystoma maculatum). had narrowed into a leisurely trail that clusters of Pink Lady’s Slipper orchids meandered through the woods. As we (Cypripedium acaule). In and of itself, this species wasn’t continued down the path, we happened anything rare or even all that upon an extremely quick Redback During the hike I would occasionally uncommon. I remember seeing Salamander (Plethodon cinereus). turn over a random rock or piece of pictures and paintings of them in This particular animal, however, was rotting wood, looking for any possible North American field guides when I a bit of an anomaly in that it was an sign of amphibious life that might be was little, and I would hold the page up erythristic morph, meaning it had an dwelling underneath. The handful to my parents and say, quite matter-of- unusual amount of red pigmentation. or so I had turned over up until this factly, “I want that one” (by the age of This salamander was much faster than point hadn’t yielded anything, so I was 9 I had become quite the zoo tycoon in the Ambystomids I had encountered tempted to continue walking when I the fantastic world that was my mind, back home, and getting it to hold still noticed an unusually flat rock laying spending hours each day pouring long enough for a decent photo proved next to the path. Maybe it was my through animal field guides, deciding difficult. (That night I would have a compulsive nature that just wouldn’t and redeciding which specimens I dream about trying to hold this same allow me to simply pass by, but I would place within my keep). As far as salamander still and capture a clear turned a quick about-face and, with I knew, Yellow-Spotted Salamanders photograph while simultaneously both hands, lifted over the rock. It took roamed the eastern woods in the same being asked game show questions a few seconds for my eyes to adjust and numbers bison used to roam the Great by an unknown host. I’m not sure if be able to make out the shape before Plains. Evidently, for a brief moment this rather stressful vision was due to me, taking in dark dirt and a slowly each year, they do. jet lag, low blood sugar, or too many moving form, haphazard yellow dots glasses of the local Massachusetts playing havoc with my vision. “I found This species’ existence consists

Herping in Anytown, U.S.A. | 13 primarily of staying dark and damp a small gathering of people turned around and retraced our steps along under debris and within suitable inward and focusing on yet another the trail. Who knows how many animal burrows. But each spring, find, this time a rather diminuitive other amphibians lurked among the when the nighttime temperatures Northern Two-Lined Salamander decomposed debris and freshwater warm and the new season rains begin (Eurycea bislineata). In this particular pools that surrounded us. Regardless, to fall, they make their way out of location a small stream ran beneath within a handful of hours we managed their relatively safe pockets of earth the trail and proceeded down a to discover quite an impressive and begin a long slow march toward gentle slope, spilling over in falls representation of the area’s indigenous the breeding pool. They do so in and collecting in small pools. Before amphibian population. I was amazed droves, in groups numbering into the long, another member of the group at the number of unfamiliar species of hundreds…sometimes thousands. This (smart enough to don rubber boots frogs and salamanders with which I field trip, however, wasn’t during their for the occasion) found a clutch of had been newly introduced. breeding season, hence the excitement Northern Dusky Salamander eggs of those who knew anything about (Desmognathus f. fuscus) beneath To this day, were you to hand me a map this fantastic little amphibian. Ed a rock in the stream. A female had of Massachusetts, I still wouldn’t be Kowalski, one of the group members been guarding the eggs, but upon able to tell you exactly where we were (and Lead Keeper in the Philadelphia being disturbed, she quickly dropped that day. It was one of many fields and Zoo’s Department of Herpetology) into the water and made her escape. forests, seeps and springs that fill this said that finding A. maculatum both While much of our group had already expanse of incredibly diverse country, away from its breeding pool and continued on down the trail, we providing forage and breeding for outside of its breeding season is almost continued on in our excitement to countless species of amphibians. Given impossible—in forty years of herping find other possible species. It turned this, it doesn’t really matter where we he had only found three under these out to be the right decision. In the were that day…and it doesn’t matter same circumstances. Even more, same pool we also managed to find a much where you are right now. A walk the specimen I held in my hands larval form of E. bislineata. Nearby, of discovery like this one can be taken was an immature animal, at least six others came upon another Northern anywhere, in any town. months out of the water. Even when Dusky Salamander, a larval Northern individuals of this species are found, Spring Salamander (Gyrinophilis Since returning from NAAC just a few they are usually sexually mature adults p. porphyriticus), and yet another months ago, I have stepped out my own in the breeding season (as mentioned Redback Salamander. back door to undertake similar trips. above) or recent metamorphs. I would In doing so, I have found three new like to have been able to pass this find We eventually caught up with the species of amphibians I had previously off to my incredible skill, exhaling some rest of the group, who by this time never encountered before, all within an diatribe about the particulars of the were peering at and taking pictures area I have spent most of my life. rock I had lifted and how it looked to be of blooming Northern Pitcher Plants of the same sedimentary composition (Serracenia purpurea) in a small but They’re out there, feeding and that Yellow-Spotted’s are known to sunlit bog. This was the end of the breeding. You just have to look. prefer using for shelter…but in the end line for us, so to speak, as we turned I was forced to chalk it up to dumb luck (and just a hint of overly compulsive An uncommon erythristic form of Redback salamander, Plethodon cinereus. behavior).

The trail progressively gave way to a thick canopy of leaves and trees as we continued on our way. I found myself hanging back from the main cluster of people, occasionally stopping to observe the now common efts making their way across the landscape. As I reached a bend in the trail I noticed

14 | Leaf Litter :: Vol 2 Issue 2 Further Impressions from Tarapoto: frogs and art

In the summer of 2006, Tracy Hicks spent 14 days in the area surrounding Tarapoto, Peru. In this 3-part series he chronicles the personal thoughts, emotions, and experiences that were the result of his being in one of the world’s richest Dendrobatid environments. by Tracy Hicks photos © Tim Paine unless otherwise noted

In the first of two earlier articles, I wrote about the personal catching frogs at this amazing edge. It was thrilling to hold difficulties of traveling to the Amazon basin. A half day’s a trivittatus and watch it ooze off-white toxic mucus into my drive west of Tarapoto is Yurimaguas, where the roads end hand. In my imagination, I can still feel the excitement of and the rivers converge, carrying a stream of people down to the slight numbing and tingling of my palm. Another sense- Iquitos, then on past the junctions of ever larger Amazonian memory is chasing a summersi from her breeding axil down rivers and across the vast gradual slope that shapes South to the surface of the leaf litter, then catching her another two America’s fertile pregnant belly. feet deeper where the leaf litter stopped and the soil began. And another: repeatedly bloodying my hands while grabbing Art defines culture and happens on the fertile edge of our for a summersi in the eye-level cone of spiny palm roots— understanding. exciting, even if not successful.

Accompanying me were two great friends, Tim Paine The “Oh, wow!” moments in which I discover another layer and Todd Kelley. Todd I described earlier as an expert in a work of art are one of the few things that can equal the frog wrangler, but even his wife describes him as a well- thrill of, say, coaxing a small, summersi onto my walking respected “gentle, monk-like character.” He is also one stick and passing it into Tim’s hands. of the most respected frog breeders in the United States. Tim is a scientist-artist-policeman who accumulates hard My secondary goal on this trip was to integrate art into the duty work-time in order to spend his off time studying experience, working the fertile edge and marking these and photographing the reptiles and amphibians of South moments with small acts of art, collecting mental and America. His images, which you see in these articles physical impressions of the place to shape the evolution of and major publications, are spectacular. To say I was my art. The area was almost too fertile: there was more to comfortable traveling with these guys would be a gross the jungle than I could ever do justice to in my work. understatement. Both Todd and Tim are artists at what they have chosen to do. They understood and accepted my The focus of each day was to find frogs. When we weren’t relationship to the place. in the jungle, we planned where we would be and when we would go the next day. We ate. We drank a few cervezas. The second article in this series described the “Oh, wow!” And when we weren’t in Cesar’s taxi, we slept (sleeping in an moments associated with achieving my primary goal: old taxi bouncing down a Peruvian highway is like sleeping

Above: Allobates femoralis EDITOR’S NOTE: When the first article in this series was published, what had previously been referred to as a banded form of Ranitomeya fantastica had not yet been identified as R. summersi. The change in identity has been reflected in this article but not the first two. All previous mentions of R. fantastica in this series should be understood as referring to R. summersi.

Further Impressions from Tarapoto: Frogs and Art | 15 Somewhere inside I carry the imprint of Miguel. His village was several hours up river from where the roads ended. The village had one narrow strip of a paved concrete that ran between thatch-roofed houses. Miguel’s house, like all the others, was constructed of roughly hewn mahogany. The thick, uneven planks allowed space for air, insects, and probably rats to flow in and out. His wife and children were beautiful.

After a brief introduction and explanation that we came to see and photograph frogs in the jungle, Miguel took us past fields in various states of cultivation. Each field was skirted by degraded forest—the ragged frontier where Peruvian farmers continually The aesthetics of the jungle can be like slipping on your most cut back the jungle…and it continually comfortable clothes . The jungle wraps you in an all-encompassing layer. crept back into the fields again. A photo by Tracy Hicks poignant edge: The farmers and the jungle vainly trying to impress upon one another what each needs to on a jackhammer). The rest of the time, The trails change after dark. The survive, while at the same time creating we were in the jungle. cones of light from our flashlights or a fertile area for new growth. headlamps would suddenly reveal giant While wandering through the jungle, whip spiders, Amblypygi, scuttling As soon as we hit a stretch of raw looking intently for any movement around our feet. My ultraviolet jungle, we started finding summersi… or flash of color that might be a frog, flashlight popped the eyes of spiders or so I thought. We had split up, taking peeing beside the trail was a natural out of the blackness and made the worn parallel paths only a few feet way to mark the spot. It wasn’t a matter all-too-common scorpions glow. It is apart. The area between the paths of spraying the spot with my scent—it scary to know how many scorpions proved very productive: as frogs moved was a way of reminding myself to make are around you all the time. Red, away from one group of us, they moved a mental record of the place. six-inch-long stick bugs, Phasmids, toward the other. I believe Miguel was which we occasionally glimpsed the first to catch a frog. While I was Picking things up was equally natural. during the daylight hours, were vastly looking at it, Todd grabbed another. So when I stopped beside the trail to outnumbered by other strange, red Todd and Tim reached the “Oh, wow!” pee, I took time to look for interesting bugs that were invisible during the day. quicker than I did: we were catching objects; handling them heightened the a combination of both summersi and experience of the moment, burning a I can imagine why the local people imitators. sharper mental impression of the spot. would be “creeped out” by the jungle Finding a radiating clustered seed pod, night life, but I felt right at home. From a box beside his house, Miguel I added small stones to the seed sockets periodically sold frogs to German and left it where I found it. I can still smugglers. I would have loved to see and describe that spot with great detail.

Visual association is a key to visual art. How you relate to what you see is unique to you, and what we fail to see merges quickly with what we choose not to see. On this trip we worked several jungles at night. Cesar, our guide, would drive us to the site and return to pick us up, but he, like the other local folks, wouldn’t go into the jungle at night. Left to right: Ranitomeya fantastica, R. summersi

16 | Leaf Litter :: Vol 2 Issue 2 Top to bottom: Ranitomeya imitator, Tim Paine examines and counts frogs destined to be sold to R. fantastica, R. benedicta German smugglers in a small outdoor cage beside Miguel’s home. photo by Todd Kelley bring a few of Miguel’s imitators where the recently imported Tarapoto small rubber froglet. A small species and summersi back home to study imitators originated. We had visited of Monstera layed like shingles over their obvious similarities and subtle this site repeatedly during the day, and a series of boulders. I wanted to carry differences. Instead we paid him what it is where I bloodied my hands chasing home physical impressions of this was probably six months income for an elusive summersi—a good place to specific spot. I also wanted to document being our guide. Miguel’s wife and make art. the process. I had brought a hunk of children crowded around the frog oil clay commonly used as a sculpture pen, watching as Tim gave Miguel the There was considerable science going medium. I also had my camera and money and emphasized the importance on there, too. On an earlier visit, we several small flashlights with flexible of protecting the frogs. I hope the met a Peruvian grad student who was shafts that could be clipped to most message impressed itself on Miguel. researching the frogs that breed in the anything. various cave-like structures within a I know he impressed us. Each of us boulder field. We saw entomologists I climbed down between two boulders has commented since returning on with nets working the path in. There and set up my tripod and lights. After I the beauty and pride of that simple were also dozens of numbered plastic kneaded the oil clay for a few minutes, Peruvian family. water cups were scattered around in it became soft and malleable. When I strategic spots for Rainer Schultz’s pressed it onto the Monstera, the clay One of our last forays out into the INIBICO project. retained clear impressions of a 6-inch jungle was a night hike back into square of Tarapoto jungle. a small, magical area of jungle That night, as soon as we reached just outside Tarapoto. It is my our base, I went directly to a spot I My lights drew flying insects, and the understanding that this is the area had previously marked by leaving a insects inevitably attracted bats. I

Further Impressions from Tarapoto: Frogs and Art | 17 stood there, sandwiched waist-deep in the jungle, shooting images of my hands making impressions of this magical spot while bats and bugs swarmed around my head and whip spiders crawled on my legs. The mental impression is creepy, but the physical impression, the mold, is beautiful.

Todd and Tim each stopped by to see the odd blue glow of the lights, and me, half buried. I assured them I was fine. I imagine they thought I must be insane. Time rushed by. Tim and Todd were ready to leave before I was finished, so I quickly documented two other sites within a few yards. One was the side of small tree latticed with vines where Tim had photographed an imitator. The other was the spiny palm where I had bloodied my own palms chasing the summersi I never caught.

Days later, back in Dallas, I cast plaster molds from the clay impressions. Now, two years later, I am still using them to make art that, I hope, impresses the fragility and mystery of the Amazon basin into the sense-memories of people who may never encounter a living jungle.

Top: A boulder with moss and shingling Monstera is covered with the oil clay mold material.

Center: The same spot in the jungle outside Tarapoto as above, but in this cast rubber piece is also a cast Golden toad, formerly known as Bufo periglenes, now extinct. Although the Golden toad is from an entirely different location, the foreshadowing analogy is accurate as art. www.tracyhicks.com/05309/

Bottom: The spiny palm roots before casting.

photos by Tracy Hicks

18 | Leaf Litter :: Vol 2 Issue 2 When Father Chimborazo Took the Frogs Away

text and photographs by John E. Simmons

t an altitude span of time. of 3,450 The páramos of Ecuador, meters Colombia, and Venezuela are moist (11,320 feet), bogs above the treeline that are the the thin ecological equivalent of the alpine Andean air zone in the northern hemisphere Asmelled fresh and clean but left me (Rivera Ospina, 2001). The peaty light-headed and gasping for breath as páramo soil is thick with liverworts I struggled uphill through the cushion and other bryophytes, ferns, tussock Figure 1. The jambato, Atelopus ignescens, plants and boggy grasses. It was March grasses (sigse grasses), a variety of low from Chimborazo Province, Ecuador, 14.6 km 22, 1984, and I was looking for frogs shrubby plants, and the distinctive SW of Cajabamba at 3740 m, 03 March 1984. on the Páramo de Apagua in Ecuador. cushion plants2 (figure 2). The northern All around me on the damp ground, páramos are often covered by majestic crawling in slow motion like miniature stands of frailejones (Espeletia sp.), lemurs, were thousands of little toads members of the sunflower family (Atelopus ignescens) (figure 1). Called (Asteraceae) that grow two to five the jambato (pronounced ham-BAH- meters tall over a lifespan of 100 years toe), the dull black creatures with (figure 3). Because of the altitude, the 1 fiery red bellies seemed oblivious to climate here is harsh—the average the wind, the cold, and the thin air. In temperature is about 50°F, but on my travels up and down the Andes, any given day the temperature may Figure 2. Páramo vegetation. jambatos were by far the most common go from below freezing to above 70°F. animal I had observed on the páramo. Páramos typically receive 24 to 70 or Little did I imagine that in less than more inches of rain annually. Páramo five years, the only jambatos that plants and animals must be adapted anyone would ever see again would be to withstand freezing temperatures, a few specimens floating in alcohol in intense solar radiation, and relentless museum jars. Having spent many a winds. happy day in South America looking The páramo can be spectacularly for frogs, I find it profoundly disturbing beautiful. When the sky is clear, the that the most common toad in the sun quickly warms the air and the Figure 3. Giant rosette frailejones (Espelitia), Andes could go extinct in such a short named by Alexander von Humboldt and Aimé Bonpland in 1809 for José Manuel de Ezpeleta, 1 Jambatos vary widely in coloration, especially the ventral colors (from yellow to red) which has resulted in several the viceroy of the Nuevo Granada from 1789- species being named and then synonomized over the years. Even within the species Atelopus ignescens there is a huge variation in coloration from one area to another. For some examples of Atelopus colors, see Renjifo, J.M. 1997. 1797, who supported their expedition. The Ranas y Sapos de Colombia. Editorial Colina, Instituto de Investigación de Recursos Biológicos Alexander von common name of frailejón comes from the Humboldt, Bogota, 160 pages. plant’s resemblance to monks (frailes) in the 2 A cushion plant is not a single species, but rather a dense cluster of many species of small vascular plants. páramo fog.

When Father Chimborazo Took the Frogs Away | 19 a narrow road cut, a large chunk of the earthen wall caved in with a thud in front of us. Before jambatos went extinct, they could be found in extraordinarily high densities. I found it impossible to drive on the Pan-American Highway in the northern Andes during the jambato breeding season without squashing them by the hundreds. A 1981 field study recorded as many as fifty frogs per square meter at one location (Ron, et al., 2003); a 1982 study recorded thirty individuals per square meter. Migrations of love-struck jambatos have been reported by travelers for decades. In the 1860s the Spanish Figure 4. A storm sweeping over Páramo El Angel at 3270 m. naturalist, Marcos Jiménez de la Espada, witnessed a mass migration of may prove dangerous to travelers. jambatos on the flanks of the volcano Winds laden with icy vapors blow Antisana in Ecuador. He recorded in over them with tremendous violence, his journal that “…we saw thousands of when as the natives call it, the páramo individuals…on the banks of the Mica se pone bravo (gets wild). Dense fogs on Antisana, early in 1865, that we frequently envelop man and beast; surprised during their mating period, darkness covers the earth and conceals when the males look for females, every trace of the road; snow, hail, or chasing them so blindly that, fighting to sleet comes down unmercifully; and catching them, they roll about in piles Figure 5. Six jambatos on a cushion plant in often the traveler loses his way and all mixed together.” (Cabondevilla, the Páramo de Apagua, 32 km W Pujili in Coto- wanders helplessly over endless heaths. paxi Province, Ecuador, on 22 March 1984. 1998) But this is not the worst; when worn out with fatigue and hunger, benumbed One hundred years later, James animals emerge from hiding to feed, with cold…his stomach soon becomes A. Peters (curator of amphibians and move about, and bask. In 1975, as affected as if at sea; his blood ceases to reptiles at the Smithsonian) reported part of a field study of herpetological circulate, his muscles grow stiff, and he two mass migrations of jambatos communities in the high Andes, we expires with a ghastly smile upon his that he witnessed in Ecuador (Peters, measured the temperatures of live features. Travelers thus found dead in 1973). The first was on December 31, jambatos in the páramo. We discovered these inhospitable regions, are said to 1958, when he saw thousands of the that the frogs could increase their body be emparamados (Hassaurek, 1967). little black toads “… moving across temperature from 0.5 to 4.6 °C above the Pan-American Highway north the substrate temperature within half One pleasant, sunny morning, a of Latacunga.” The other migration an hour of emergence from their hiding friend and I were looking for páramo was “in the páramos on the road places (Duellman, et al., 1986). The frogs when a storm swept over us. from Quevedo to Latacunga, on frogs need to be able to warm up fast, The jámbatos and hummingbirds January 22, 1959, at an altitude of because at any moment a darkening immediately disappeared as a cold rain 3,600 meters… the movement was… sky means rapidly falling temperatures, plummeted down. The temperature in a single direction, and definitely thick fog, and freezing precipitation plunged from 75°F to near freezing as downhill. A light drizzly rain was (figure 4). The rapid weather changes we retreated to our jeep. By the time we falling, and sufficient rain had already may catch travelers unprepared. In an reached the vehicle and got underway, fallen to make the road somewhat account published in 1867, Fredrich the rain had changed to snow and then treacherous, with standing puddles. Hassaurek cautioned that: páramos to sleet. While we were driving through In both cases, the migration seems to

20 | Leaf Litter :: Vol 2 Issue 2 be toward standing water, presumably the place of egg deposition...” Younger If you gently apply a sliver of bone and cover it with cotton, naturalists, who never had the chance being very careful not to touch it with your tongue, in a to observe jambatos in the wild short while the molar will splinter into pieces down to its prior to their decline, often find the population density data difficult to roots. I have seen this with my own eyes... believe. I once offered a photograph I took of six jambatos on a cushion quinine. When the Inca empire began bone and cover it with cotton, being very careful not to touch it with your tongue, in plant (figure 5) to a young Ecuadorian expanding in 1438 (reaching Ecuador a short while the molar will splinter into scientist who declined to use it because in 1460), Quichua was made the pieces down to its roots. I have seen this he believed that the frogs had to have lingua franca of the Andes as part with my own eyes... been posed for there to be so many of a power restructuring that also together. In fact, the little toads were involved moving ethnic groups from Atelopus ignescens was the first so common that for many years, their place of origin to new territories amphibian from Ecuador to be given a few people bothered to collect them. (MacQuarrie, 2007). By forcing formal scientific name. In 1849, Emilio Unfortunately, the last known living everyone off their ancestral lands Cornalia (1824-1882) named Atelopus individual of Atelopus ignescens was through conquest and compromise, ignescens (as Phryniscus ignescens) seen in the wild in March of 1988 and making everyone speak the same based on specimens received at the (Coloma, 2005). language, the Inca Empire solidified Museo Civico di Storia Naturale di rapidly. Nevertheless, Quichua has Milano (Cornalia, 1849) that had The problem of disappearing retained many local variations so that, been collected near Latacunga. The amphibians goes well beyond depending on the dialect, jambato may toad’s vivid, fiery red-orange belly so jambatos—43% of known amphibian be rendered as hambatu, hambato, impressed Cornalia that he derived species world-wide are either declining jambatiug, jambatyu, jambatyuj, its specific name from the Latin word or have already become extinct. What ambatyu, ambatiug, and jamp’atu. The ignescere, meaning to catch fire. The makes the disappearance of jambatos name of the city of Ambato, Ecuador generic name, Atelopus, comes from so unsettling is how well-known (140 km S of Quito) is derived from the the words ateles (imperfect) and pus the toads were and how common word jambato5. (foot), and was chosen by the French references to them are in Andean One of the earliest descriptions of herpetologists André Marie Constant cultures. The diurnal jambatos Duméril and Gabriel Bibron in 1841. were common residents in gardens jambatos was written in 1789 by Juan and parks in Andean cities—I have de Velasco (1727-1792), an Ecuadorian When a species is given a scientific collected jambatos several times within Jesuit born in Latacunga (90 km S of name, a single individual is designated the metropolis of Quito. Jambatos Quito). In his book, Fantastic Zoology as the type specimen or holotype were so prominent in popular culture (de Velasco, 1984), Velasco wrote: for the species. The identification of that in March 2002, the Ecuadorian Hambatu is the common name of a another specimen of the same species post office issued a jambato postage species of toad, which are common and can be confirmed by comparing it to stamp3. With their distinctive of diverse colors and sizes. They have a the holotype. Because the specimens coloration and stubby toes, jambatos row of poison glands at mid-body. There used by Cornalia were lost long ago, were instantly recognizable. are some very venomous ones in the a replacement type (called a neotype) mountainous deserts of the province of was designated for Atelopus ignescens The name jambato is a Quichua Ibarra that are entirely green, with red in 2000 (Coloma, et al., 2000). The 4 word . The Quichua language has eyes and as many as six fingers. They are place where Conalia’s specimens were enriched English with several words hunted because they are used as a remedy collected had become so degraded including alpaca, coca, condor, for molars with cavities that cannot be that the population of jambatos there guano, jerky, llama, puma, and removed. If you gently apply a sliver of

3 The stamp, issued in 2002, is part of the “Fauna en extinction” series and is labeled both as jambato and Atelopus ignescens.

4 Quichua is spoken by native people throughout the Andes in a grand variety of regional dialects, reflected in the various spellings of Quichua, which include Kichwa, Oquichua, Quechua, and Kechua.

5 Ironically, one Quichua-Spanish dictionary published in Ecuador illustrates the entry for jambatyuj with a line drawing of a marsupial frog (Gastrotheca), despite the precise use of the word by native Quichua speakers [Stark, L.R., and P.C. Muysken. 1977. Diccionario Español-Quichua, Quichua-Español. Publicaciones de los Museos del Banco Central del Ecuador, Quito, xvii + 366 pages].

When Father Chimborazo Took the Frogs Away | 21 Jambatos played an important role in Andean folk medicine as well (Acero Coral and Pianalto de Dalle Rive, 1985). Jambatos were used to cure espanto or susto (a form of shock often caused by the consternation of encountering a ghost), and to cure ñahui-nanay (a condition of dry, bloodshot eyes that results from being in the sun without a hat, or working in poor light). To cure ñahui-nanay, the belly of the toad was slit open and rubbed on the patient’s neck and forehead. To cure erisipela (painful, red, swollen spots on the arms, legs, or face) a live jambato was tied to the affected area of the body with a bandage, left until it Figure 6. Potato harvest on the páramo 16.5 km E of San Miguel de Salcedo at 3090 m, died, then the body was removed and Cotopaxi Province, Ecuador. hurled far away. Tadpoles were used to treat watery eyes (Coloma, 2003). was extinct. The neotype was, instead, the arrival of the rainy season. It was In the Imbabura region of Ecuador, chosen from a nearby population in the widely believed that the big jambatos it is considered a sin to kill a jambato páramo that was one of the last places carried the smaller ones around on because they were used to treat scabies, where a jambato was seen alive, near their backs when they became tired, remove warts, and (if applied to the Latacunga on the road from San Miguel but what people were really seeing forehead) cure migraines (Lombeida, de Salcedo to Lagunas de los Anteojos. was the smaller males grasping the 2005). I visited this locality in 1984, before larger females in a mating embrace There are several references to the mass extinction of jambatos (figure (once hormones were flowing, a jambatos in Ecuadorian literature. 6). I was recovering from a bad bout of male jambato—driven by amorous Because Atelopus ignescens were fever as we drove across the páramo, intentions—might cling to the back of slow moving they were thought to be but noted in my journal that “from a female for hours, days, or weeks at a dimwitted and clumsy, so Indians Salcedo on the páramo is fairly heavily time until she was ready to crawl to the sometimes use the word jambato as populated and converted to agriculture. water and deposit her string of eggs). a nickname or as an insult. Several We saw lots of llamas, in groups of Under other circumstances, the mere characters in the works of Jorge Icaza two or four, but never in herds like in presence of jambatos could be a sign of have the nickname Jambato. In Media Peru and Bolivia. Above San Miguel, bad luck. Another folk belief was that Vida Deslumbrados, Icaza describes the páramo was relatively untouched, old guinea pigs could turn into toads how “As a strong breeze opened the with a large number of lakes. It was a and then run about grunting. In the door, he was stubbornly dragged over sunny day, so Atelopus collecting was Andes, guinea pigs are an important the floor, and stuck a hand out of the good.” Just three years later, when the food source, and are traditionally kept blankets, but at once slipped like a neotype was collected on 29 November in the kitchen and fed with fresh grass jambato…” In his best known book, 1987, it was one of just 17 specimens that is collected daily, usually by the Huasipungo6, Icaza describes a group of jámbatos found, and three of them children of the household. Quichua of tired Indian workers as “…curled were already dead. children were instructed not to collect up in the corners, lulled to sleep by The indigenous people of the Andes grass from places that they saw the monotonous music of the drips, knew the jambato well, and it played jambatos because it was believed that by the incomplete orchestra of toads a significant role in folk tradition. toad urine transmitted a disease to the and crickets,” and later describes For example, the first appearance of guinea pigs that caused their stomachs “...the harmonic sound of an orchestra jambatos on the páramo heralded to swell until they died (Archetti, 1997). of toads…” The calls of several Andean

6 Published in English as The Villagers.

22 | Leaf Litter :: Vol 2 Issue 2 frogs and toads can be heard in the away.” Chimborazo is an active volcano and dry conditions in the previous 90 indigenous music of the Andes, both that dominates the landscape, and is years were recorded in the northern figuratively and literally. Figuratively, believed by many indigenous people Andes. During the last century, while frog calls are represented by the to exert a powerful influence on the the earth has warmed by an average pronounced, high-pitched repetitious environment and their lives. They of 0.5°C, the temperature increase in pulsing of many tunes. Literally, the call the mountain Taita Chimborazo the Andes of Ecuador has been closer calls may be heard in the background (Father Chimborazo). to 2.0°C. Ultraviolet radiation in the of field and festival recordings. Frog At the base of Chimborazo is the Andes has increased significantly since songs (and other natural ambient gritty, volcanic Palmira Desert (figure 1979, quite possibly beyond the limit of sounds) are included on numerous 7). In years past, where streams of tolerance for some species. 7 albums produced by native musicians . meltwater flowed down from the Luis and I made frequent stops According to James A. Peters, male glaciers on Chimborazo, the bits of at places where jambatos were once jambatos “…make a slight sound… green plant life amid the dark grey known to be common or that appeared a very low, chick-like peep, difficult ash supported small populations of to offer the right habitat for the toads. to hear even at short distances, and jambatos. In 1997, Luis and I found Jambatos deposit creamy yellow eggs in made while walking about looking that the glaciers on Chimborazo fast-running, well-oxygenated streams for a female…” (Peters, 1973) This is were in fast retreat (figures 8-9), the that are lined with vegetation to protect probably because jambatos lack the streams across the Palmira desert the tadpoles from excessive sunlight. hearing apparatus that most frogs fewer and smaller, and jambatos were Jambato tadpoles have a large suction have—they do not have an external nowhere to be found. Recent research disc behind the mouth that they use to eardrum (tympanum) or ear ossicles has revealed that glaciers in Ecuador attach themselves to the undersides of (McDiarmid, 1971). Although jambatos have lost more than 20 meters of ice stones in the swiftly flowing cold water. had limited hearing through a muscular in the past two years, which is having Wherever we found likely habitat, we connection between the subrascapula grave consequences for the natural made inquiries about jambatos among (part of the shoulder blade) and the otic environment, agriculture, and the the local Indians, including a group of capsule, the range of frequencies that numerous cities that depend on Andean native women carrying large bundles jambatos could have heard is unknown. glaciers for water (Sosa, 2007). The of sigse grass across the páramo In August 1997, I traveled with pace of glacial retreat on Chimborazo to be used for mattresses and roof the leading authority on Ecuadorian is the result of rapid climate change. In thatch, but no one could recall seeing amphibians, Luis Coloma (who wrote 1987, the year before the last jambato any jambatos. We stopped at what his doctoral dissertation on jambatos), was seen alive, the most extreme warm appeared to be a promising area just from Quito southward through the Andes to investigate persistent rumors that some isolated populations of jambatos still lingered on, despite no living individuals having been collected in ten years. After an unsuccessful morning spent looking for jambatos in Cotopaxi National Park, we stopped for a late lunch in San Miguel de Salcedo. Leaving the restaurant, Luis asked an Indian what he thought had happened to the frogs. The man replied simply, “Taita Chimborazo has taken them all

7 The distinctive call of the marsupial frog, Gastrotheca (a loud brack-bak-bak-bak), the high peeps of the robber frogs (Eleutherodactylus sp.) and glass frogs (Centrolenidae), and other anurans can be heard on the CDs Salasacamanda, Quinchuquimanda Imbabayacuna, and on most of the albums in the Ñanda Mañachi series. Figure 7. Palmira Desert, 11.3 km N Palmira, Chimborazo Province, Ecuador.

When Father Chimborazo Took the Frogs Away | 23 them in winter, but that there had not been any all summer, although there are still sapos verdes (marsupial frogs) around. Another man and a young boy riding a horse said that they used to see jambatos all the time but had not seen any for at least three years. Rural inhabitants of the Andes noticed that the jambatos were disappearing long before the scientists did and developed their own explanations for the phenomenon. Some people claim that the jambatos disappeared because a strange wind began blowing down from the mountains; others say it is a consequence of the visit of Pope John Paul II to Ecuador in 1985.

Figure 8. Glaciers on Volcán Chimborazo in March 1984. Exactly why the jambatos went extinct is a question that is difficult to answer. Several hypotheses have been proposed to explain frog extinctions, including the loss or degradation of habitat, over- exploitation of populations, climate change (resulting in warmer, dryer weather), chemical contaminants, emerging infectious diseases, or the introduction of exotic species that have out-competed the native species. As we drove past Chimborazo, Luis pointed to a small lake where a sign advertised “pesca deportiva,” or sport fishing, for introduced rainbow trout (Onchorhynus mykiss) and brown trout (Salmo trutta). However, the introductions of alien fish occurred decades before the jambatos went Figure 9. Glaciers on Volcán Chimborazo in August 1997. extinct. Likewise, 27.1% of the Andean páramo and 33.3% of the Andean beyond Palahuín, at 4,200 m elevation, “el tiempo ha cambiado y no hay” (the forests of Ecuador have been cleared, and as we were looking for frogs a weather has changed and there aren’t but this habitat destruction has been a herd of haughty llamas came by. Luis any left). As we drove up a bad dirt road gradual process, not the sort of rapid asked the Indian herder when the last over a pass at 4,500 m through Salinas change that might push a species to time was he had seen a jambato. After to the Río Pachanco, we discovered extinction so quickly. Unfortunately, a moment’s reflection he replied not that glacial retreat had reduced the there is little hard data available to for “a year or two,” which is a Quichua formerly pristine páramo river flowing support most of the hypotheses for idiom that does not mean a literal 12 down from Chimborazo to no more frog extinctions (Ron, et al., 1994). to 24 months, but rather a very long than a small trickle. When a man came Habitat loss is a serious concern time. When Luis asked the herder why by on a mule, we asked him about for biodiversity worldwide, but the the frogs were gone, he answered that jambatos. He said that he last saw pace of change it produces does not

24 | Leaf Litter :: Vol 2 Issue 2 account for the rapid extinction rates observed in most declining amphibian The chytrid fungus was found in almost all rapidly populations. Over-exploitation and the introduction of alien species may declining amphibian populations that were tested... account for a few population declines, but not the rapid extinction of entire, populations that were tested, but it took This is not to say that climate widespread species. Likewise, chemical several years before techniques were change and habitat loss are not factors contaminants have been implicated developed that could detect chytrid in amphibian extinctions (they are), in some localized areas, but not on in preserved museum specimens. but rather that climate change is not a global level. Climate change and Museum collections are an invaluable driving the spread of the pathogen infectious diseases, however, may well archive of life on earth over time. One directly. While in Cuzco, Peru, in 2004, work synergistically. reason that museums keep collecting I saw a dramatic example of how this specimens of species they already have works in nature. A colleague asked In the 1990s, several researchers is to study just such phenomena as me to stop by the Museo de Historia were quick to support the hypothesis the emergence of a new killer fungus, Natural at the Universidad Nacional that climate change, resulting in environmental contaminants, and San Antonio Abad and take a look at increased ultraviolet radiation, was the other changes that the populations some frogs that a team of high altitude primary cause of amphibian extinctions are exposed to. However, a study of biologists had discovered recently in (Blaustein, et al., 1994). While the 89 museum specimens of jambatos the central Cordillera of Peru (Seimon, UV connection looked convincing collected prior to 1987 (the year before et al., 2006). Thick glaciers have for some extinction events, it did not the last individual was seen alive) covered the Cordillera Vilcanota (near hold up as a good explanation for revealed that none were infected with Cuzco) for hundreds or thousands of others. We know that an increase the killer fungus. years, but beginning around 2000, in ultraviolet radiation and other rapid glacial retreat formed a few climatic and habitat changes may A group of frog researchers meltwater ponds at very high altitudes affect frog populations by increasing recently analyzed the patterns of where the ground had previously adult mortality, reducing reproductive the spread of the chytrid fungus by been covered by ice. The mountaineer success, and by weakening immune applying statistical analyses to data ecologists discovered that frogs had systems (which would make frogs more from museum specimens and field become established in some of these susceptible to pathogens). In 1996, studies (Lips, et al., 2008). They ponds at altitudes up to 5,348 m (more the discovery of a fungal disease in made the revealing discovery that than 17,000 feet), much higher than many dying amphibians quickly led to the fungus does not spread in direct frogs had ever been recorded before. At a new hypothesis, called the climate- response to climate change, but rather that altitude, life is extremely difficult linked epidemic hypothesis, or CLEH shows the same pattern as other for an amphibian—the team discovered (Pounds, et al., 2006). The reasoning plagues such as rabies, Lyme disease, that the frogs would freeze in the ponds was that changes in patterns of rainfall, and the Ebola virus. The researchers overnight and thaw out again each day average temperature and so forth hypothesized that chytrid fungus has with the morning sun. Unfortunately, are directly linked to the emergence been introduced at various places at the colonization of the new ponds of chytridiomycosis, a disease different times around the world, such was followed shortly by the arrival caused by a fungal pathogen named as from escaped African clawed frogs of the chytrid fungus, which caused Batrachochytrium dendrobatidis (used widely in pregnancy testing, devastating die-offs of both adults and (called Bd or chytrid for short). The research, and sold in the pet trade) and recently metamorphosed juveniles. name Batrachochytrium comes from North American bullfrogs (introduced The solar heating of the newly formed the Greek words batrachos (frog) in many places, including Ecuador, meltwater ponds that made the water and chytro (an earthen pot with the to be raised for food). From these warm enough for the frogs to live in same shape as the fungus). The name initial alien species introductions, the also created an environment in which dendrobatidis refers to the poison disease spread through the native frog the chytrid fungus could flourish. dart frogs (family Dendrobatidae) in populations in a pattern typical of which the fungus was first detected. emerging infectious diseases. A study The story of the jambato tells us The chytrid fungus was found in of the genetic signature of the chytrid a lot about how people perceive and almost all rapidly declining amphibian fungus supports the theory as well. understand their environment, both

When Father Chimborazo Took the Frogs Away | 25 Figure 10. Sapari exhibit, Quito

from folkloric and scientific points of view. For example, knowing that jambatos were more active just prior to the advent of the irregular rainy season was one of many important indicators to an indigenous agricultural society of seasonal change. A folktale such as one telling how larger frogs carry smaller ones around on their backs illustrates the importance of knowing when frogs were going to mate. The belief that toad urine causes fatal disease in guinea pigs probably came about because jambatos favor moist habitats, and wet grass is not healthy for guinea pigs, whose wild relatives graze on dry grass. As for look for is the hypothesis that is as an explanation for the jambato’s the role of jambatos in folk medicine, best supported by the available extinction shows the scientific process we know that toad skin contains a evidence, and which best explains all at work. wealth of complex chemicals, including observations. The scientific quest for toxins, hallucinogens, and substances In early 2005 I was in Quito to answers to problems is a continuous with antibacterial properties. The skin participate in a celebration of frogs and activity, which is why scientists secretions of the jambatos may well toads called Sapari. The name Sapari are reluctant to use the word fact have helped cure a number of ailments. is a clever play on words, combining (what is believed to be a fact today sapo (toad) with safari (expedition). The numerous hypotheses that may not be tomorrow). Each time I discovered that Ecuadorians are have been advanced to explain the a new bit of evidence is discovered, fascinated by and quite fond of demise of the jambatos show us how previous explanations for an observed their native frogs and toads. The science progresses. It is misleading phenomenon have to be re-evaluated. Sapari exhibit attracted more than to say that one hypothesis is wrong The shift from ultraviolet radiation to 105,000 visitors in an extended run while another is right—what scientists habitat degradation to disease vectors of 90 days at the cultural center of

26 | Leaf Litter :: Vol 2 Issue 2 the Pontificia Universidad Catolica in Archetti, E.P. 1997. Guinea-Pigs: Food, and M.W. Sears. 2008. Riding the wave: Quito (figure 10). The Sapari exhibit Symbol and Conflict of Knowledge in Ecuador. reconciling the roles of disease and climate was the brainchild of Luis Coloma, Berg Publishers, 256 pages. change in amphibian declines. PLoS Biology and included live frogs, frog art, Blaustein, A. R., P.D.Hoffman, D.G. Hokit, 6(3):441-454. frog folklore, educational programs, J.M. Kiesecker, S.C. Walls, and J.B. Hays. Lombeida, D. 2005. Los anfibios en el exhibits on frogs and frog research 1994. UV repair and resistance to solar UV-B pensamiento ecuatoriano. Ecuador: Tierra in Ecuador, as well as a tribute to in amphibian eggs: A link to population Incognita number 33:28-32. the vanished jambatos. One exhibit declines? Proceedings of the National MacQuarrie, K. 2007. The Last Days of the featured nocturnal frog species that Academy of Science. 91:1791-1795. Incas. Simon & Schuster, New York, xv + 533 had their activity cycles shifted by 12 Cabondevilla, M.A. (editor). 1998. El Gran pages. hours so that they were actively calling Viaje, Ediciones Abya-Yala, Quito, 250 pages. and mating during the day as visitors McDiarmid, R.W. 1971. Comparative were escorted through a darkened Coloma, L. 2005. El último croar del morphology and evolution of frogs room decorated like a lowland tropical jambato. Desafío 5(9):26-28; Bergman, C. of the neotropical genera Atelopus, rain forest. Children and adults could Where are the clowns? Ecuador’s harlequin Dendrophryniscus, Melanophryniscus, and use microscopes to watch developing frogs bring attention—and action—to South Oreophrynella. Bulletin of the Los Angeles marsupial frog embryos still in their American’s Vanishing Amphibians. Wildlife County Museum of Natural History, Science eggs and could see tadpoles hatching. Conservation, March/April 2007, pages 10-16. 12:1-66. The Sapari show included helpful Coloma, L. 2003. Uilli-uillis. Nuestra Peters, J.A. 1973. The frog genus Atelopus hints for making home gardens more Ciencia numero 5:3-6. in Ecuador (Anura: Bufonidae). Smithsonian frog-friendly in the hope that more Coloma, L.A., S. Lötters, and A.W. Salas. Contributions to Zoology number 145, iii + Ecuadorian species would not follow 2000. Taxonomy of the Atelopus ignescens 49 pp. the jambato into extinction. complex (Anura: Bufonidae): designation Pounds, J.A., M.R. Bustamente, L.A. Jambatos were marvelous animals, of a neotype of Atelopus ignescens Coloma, J.A.Consuegra, and M.P.L. Fogden. and the páramos seem empty without and recognition of Atelopus exiguous. 2006. Widespread amphibian extinctions them. I have worked with amphibians Herpetologica 56(3):303-324. from epidemic disease driven by global and reptiles most of my life—first in a Cornalia, E. 1849. Vertebratorum synopsis warming. Nature 39:161-167; La Marca, E., zoo, then in museums. As fascinating in museo mediolanense extantum: quæ per K.R. Lips, S.Lötters, R. Puschendorf, R. Ibáñez, and useful as captive animals and novam orbem Cajetanus Osculati collegit J.V. Rueda-Almonacid, R. Schulte, C. Marty, F. preserved specimens are for learning annis 1846-47-48 speciebus novis vel minus Castro, J. Manzanilla-Puppo, J.E. Garcia-Pérez, more about our environment, like cognitis adjectis nec non descriptionibus F. Bolaños, G. Chaves, J.A. Pounds, E. Toral, and most biologists, I prefer to see frogs atque iconibus illustrates. Typographia B.E. Young. 2005. Catastrophic population living wild and free in the habitat they Corbetta, Modoetiæ, 16 pages. declines and extinctions in neotropical evolved in. But as sad as it is to see harlequin frogs (Bufonidae: Atelopus). de Velasco, Juan. 1984. Zoologica the last remaining jambatos in a jar of Biotropica 37(2):190-201. alcohol, it is reassuring to know that we Fantástica. Historia del Reino de Quito en la Rivera Ospina, D. 2001. Páramos de have those specimens to help us figure América Meridional. Editorial El Conejo, Quito, Colombia. Banco de Occidental Credencial, out why amphibians are going extinct 144 pages. Cali, 205 pages; Acosta-Solis, M. 1984. Los worldwide. Although it is too late to Duellman, W.E. and L. Trueb. 1986. Páramos Andinos del Ecuador, Publicaciones save the jambato, what we are learning Biology of Amphibians. McGraw-Hill Book Cientificas MAS, Quito, 220 pages; Josse, C., about amphibian declines is being Company, New York, xvii + 670 pages; Péfaur, P.A. Mena, and G. Medina (editors). 2000. La applied to try to prevent the extinction J.E. and W.E. Duellman. 1980. Community Biodiversidad de los Páramos. Serie Páramo of other species. We can only hope that structure in high Andean herpetofaunas. 7, Grupo de Trabajo en Páramos del Ecuador/ we are not too late. Transactions of the Kansas Academy of Abya Yala, Quito, iii + 95 pages. Works Cited Science 83(2):45-65. Ron, S.R., W.E. Duellman, L.A. Coloma, and Hassaurek, F. 1967. Four Years Among the Acero Coral, G., M.A. Pianalto de Dalle M.R. Bustamente. 2003. Population decline of Ecuadorians. Southern Illinois University Press, Rive. 1985. Medicina Indígena Cacha- the jambato toad, Atelopus ignescens (Anura: Carbondale, xxi + 196 pages. Chimborazo. Ediciones Abya-Yala, Quito, 172 Bufonidae) in the Andes of Ecuador. Journal of pages. Lips, K.R., J. Diffendorfer, J.R. Mendelson, Herpetology 37(1):116-126.

When Father Chimborazo Took the Frogs Away | 27 Sosa, G. El Cotopaxi se deshiela. El Comercio, 17 January 2007.

Seimon, T.A., A. Seimon, P. Daszak, S.R.P. Halloy, L.M. Schloegel, C.A. Aguilar, P. Sowell, A.D. Hyatt, B.Konecky, and J.E. Simmons. 2006. Upward range extension of Andean anurans and chytridiomycosis to extreme elevations in response to tropical deglaciation. Global Change Biology 12:1-12.

28 | Leaf Litter :: Vol 2 Issue 2 on the Husbandry, Captive Reproduction, and Distinct Juvenile Form of Wallace’s Flying Frog (Rhacophorus nigropalmatus) Notes Text and photography by Michael Ready • Sandfire Dragon Ranch

One of the most curious and interesting reptiles which I met with in Borneo was a large tree-frog, which was brought me by one of the Chinese workmen. He assured me that he had seen it come down, in a slanting direction, as if it flew. On examining it, I found the toes very long and fully webbed to their very extremity, so that when expanded they offered a surface much larger than the body. The fore legs were also bordered by a membrane, and the body was capable of considerable inflation. The back and limbs were of a very deep shining green colour, the under surface and the inner toes yellow, while the webs were black, rayed with yellow. The body was about four inches long, while the webs of each hind foot, when fully expanded, covered a surface of four square inches, and the webs of all the feet together about twelve square inches. As the extremities of toes have dilated discs for adhesion, showing the creature to be a true tree-frog, it is difficult to imagine that this immense membrane of the toes can be for the purpose of swimming only, and the account of the Chinaman that it flew down from the tree becomes more credible. This is, I believe, the first instance known of a “flying frog”…. It would appear to be a new species of the genus Rhacophorus…

Alfred Russel Wallace: The Malay Archipelago, 1872

Notes on the Husbandry, Captive Reproduction, and Distinct Juvenile Form of Wallace’s Flying Frog | 29 This early report of amphibian The neck region is fairly articulated, spotting reduces subsequently to just aviation by Wallace is perhaps the allowing the frog to turn its head to a few traces. The eye color, which also first description of the behavior, and a moderate degree. Placed forward begins as a red-orange hue, gradually is a vivid account of a species that on the head, the eyes are large with fades to silver in tandem with the skin would become known as Rhacophorus horizontal pupils and a silvery white transformation. nigropalmatus. iris. This distinct juvenile form Described by George Albert As a result of our recent captive suggests use of a different microhabitat Boulenger in 1895, Rhacophorus breeding of R. nigropalmatus at during the early life stages of this nigropalmatus is commonly referred Sandfire Dragon Ranch (SDR) —a species, or perhaps a strategy to cope to as “Wallace’s flying frog.” The frog herpetoculture facility near San with different predators. It‘s also doesn’t actively fly, of course, but uses Diego, Calif.,—we discovered that the interesting that the juvenile phase of the extensive webbing of its hands juvenile form and feet, and the folds of skin along its of this species limbs to catch air, enabling a slower is quite distinct Right : Close-up of an adult fall, forward movement, and even from the adult. female Wallace’s flying frog maneuverability (Emerson & Koehl Metamorphic (Rhacophorus nigropalmatus) showing granular dorsum and 1990). A “gliding frog” is the more frogs are rusty accessory skin fringes accurate term. red to bright Below: An adult male R. orange in overall Description nigropalmatus in its typical color. Multiple A member of the family nocturnal stance, perched on an yellow and white Rhacophoridae and subfamily alocasia stem, ready to pounce on spots decorate a passing arthropod or take flight Rhacophorinae, R. nigropalmatus the dorsum, at the slightest disturbance. is an impressive and beautiful which is also amphibian. Though slender in build, R. very rugose in nigropalmatus is a relatively large frog. comparison to Males reach 90 mm, and females can the adult. It is a grow to 100 mm SVL (Inger & Stuebing fantastic looking 1997). In adults, the dorsum is emerald color phase. green and peppered with very small When it flecks of white, often with a few larger reaches a snout white spots high on the thigh or near to vent length of the heel. The dorsal skin is slightly approx 48 – 52 granular. The flanks are yellow and mm, the young fade into a white ventral region. The glider undergoes interdigital membranes, or patagia, are an ontogenetic complete, reaching almost to the tips change in color of the toes. As mentioned in Wallace’s and morphology. report, this incredible webbing is During this jet black and interspersed with lines period, the of yellow. Though the webbing is immature frog enormous, it is well concealed when the begins a gradual digits are not splayed wide. A smooth transition from flap of skin runs along the outside of rusty orange the arms and the legs, helping to better to a variable conceal the frog when at rest, as well green color, as providing greater surface area when while retaining gliding. (Emerson, Travis, & Cole 1990) the white spots. The head is short with a fairly As growth blunt snout. The rostrum of the female continues, the appears to be slightly more truncate.

30 | Leaf Litter :: Winter 2009 R. nigropalmatus is very similar in Natural History and Breeding Wallace’s flying frog (Rhacophorus color and pattern to Nyctixalus pictus, R. nigropalmatus follows the nigropalmatus) goes through a a small phytotelmic rhacophorid that breeding patterns of many other fascinating and beautiful ontogenetic change in color and skin texture. Here, shares a very similar geographic range. members of the Rhacophorinae, three individuals depict various points producing relatively large foam nests Range and Habitat in the continuum of that transition. attached to vegetation overhanging Rhacophorus nigropalmatus is Younger frogs (top) are a rusty red- temporary pools. Interestingly, the orange hue with many yellowish spots known to occur in parts of Peninsular pools have often already been used as and a fairly rugose dorsum. Half grown Thailand, Malaysia, Borneo (Brunei, wild pig, rhino, and buffalo wallows. juveniles (middle) begin to turn green, Kalimantan, Sabah and Sarawak), and but retain much of the spotting on Lee Grismer, a world-renowned Sumatra (Frost 2008). An inhabitant the dorsum, and the skin becomes herpetologist now working extensively of low elevation primary forest (sea smoother. Sub-adults (bottom) continue in Malaysia and other parts of level to 600 m), this frog spends most to transform; the overall emerald color Southeast Asia, has noted that these becomes richer and the spots and skin of its time out of sight in the upper frogs appear to prefer pools with large granulation are further reduced. canopy. The rare encounters with these amounts of buffalo urine. Additionally, dramatic looking amphibians usually Grismer gives this account of their coincide with the tropical monsoon vocalizations, which has been season, when, under heavy rainfall, undocumented otherwise: they descend from the treetops to They have an awesome breed (Malkmus et al 2002). Little to call, which sounds exactly nothing is known of its microhabitat like somebody knocking on preferences within the canopy. a hollow wooden door. It’s Most of the imported R. unmistakable. (Grismer nigropalmatus are collected in 2008, pers. comm.) Peninsular Malaysia, an area which Like many other sees a great deal of rainfall. During members of the genus, the northeastern monsoon season, during amplexus the female September through December, many produces a viscous fluid that locales receive over 300 mm of rain per she beats to a froth with the month. Temperatures are warm and rapid movement of her hind vacillate very little throughout the year, limbs. As the nest grows, averaging 31C (88F) during the day and the female oviposits her 23C (74F) at night. Relative humidity eggs within the mass. Males is also quite uniform, generally in the in amplexus may also 70% – 90% range. use their legs to help Conservation Status agitate the foam as Presently, under the International they contribute Union for Conservation of Nature sperm and (IUCN), Wallace’s flying frog is listed as seminal a species of least concern (LC) because fluid to of its fairly extensive distribution. the mass In other parts of its range, however, (Duellman populations have been assessed at & Trueb near threatened and vulnerable levels. 1986). Overall, like many other amphibian The nest is populations, there is a decreasing trend made on branches due primarily to habitat encroachment or leaves above the and deforestation (van Dijk, P.P, et al. water. The thin outer 2004; IUCN 2006). layer of the meringue- like nest hardens, while

Notes on the Husbandry, Captive Reproduction, and Distinct Juvenile Form of Wallace’s Flying Frog | 31 32 | Leaf Litter :: Winter 2009 the inside foam stays soft and liquid. treated for infections and parasites. leaves of live plants for resting spots. Over several days the eggs develop into Bacterial infections are very common We use a variety of large leaf aroids, embryos. Eventually, the nest begins to in imported rhacophorids, particularly mostly Philodendron and Epipremnum deteriorate and the foam and tadpoles those originating from South-East Asia species. Almost exclusively, our drip into the water below. (deVosjli 1996). animals choose resting sites in the Prophylactic treatment may be upper portions of the enclosures. Plants Care in captivity the best course of action for this left in plastic pots may be secured from Wild-caught R. nigropalmatus species, and perhaps all of the wild- the top of the cage to create cover in the have been imported from Malaysia caught SE Asian rhacophorids. higher regions. intermittently for a number of years. In line with Philippe de Vosjoli’s Depending on aesthetic These small, irregular shipments recommendations for establishing preferences, a variety of substrates can usually accompany larger quantities of the congener Rhacophorus be utilized. Simple unbleached paper other Malayan amphibians, reptiles, reinwardtii, we administered Panacur towel works great. Alternatively, New and invertebrates. Traditionally, the ® (Fenbendazole) and Flagl ® Zealand sphagnum moss or similar imported specimens of this species, (Metronidazole) treatments to several over a drainage layer of light expanded along with other rhacophorids from the newly imported R. nigropalmatus. In clay aggregate (LECA) is another region, have fared poorly in captivity. addition, the frogs were treated with simple method that we have used with This low survivorship is likely the antibiotic Trimethoprim sulfa. The this species. due to the poor condition of wild frogs were misted with the medication, Soil substrates are unnecessary caught animals, a result of the trauma and the water in the enclosure was for arboreal amphibians, unless one of collection, exposure to pathogens treated for five days. is attempting a naturalistic planted in crowded and unsanitary holding In our previous experience, even vivarium. If you use this type of facilities, and the stress of inadequate if outwardly healthy animals were substrate with tree frogs, it’s beneficial shipping practices. obtained, they perished from unknown to cover the surface with sphagnum The following husbandry causes in a short period of time. moss, leaf litter, pieces of bark, and parameters are methods that have Necropsies of these frogs uncovered other organic items in order to limit worked well for R. nigropalmatus in extensive bacterial infections in vital contact with the frogs. Otherwise, the our care. They are only guidelines, organs. The above protocol, however, substrate tends to coat to their skin. however, as very little is known about has greatly increased survivorship. the natural history of this species. Temperature Prophylactic treatment for Hopefully more will be learned through We have maintained this species chytridiomycosis is also a best practice expanded observation in the field and at ambient temperatures of 24C for acclimating wild-caught gliders. So in captivity to guide the development – 31C (75F - 88 F) during the day, far, PCR analysis on freshly imported of specific long-term husbandry 21C – 25C (70F – 77F) at night, adult R.nigropalmatus that we have techniques for this wonderful frog. with some gradation within the received has been negative for chytrid. vivarium. The frogs will tolerate short Acclimation A few other amphibians from the same periods of exposure to slightly higher Imported frogs are often poorly Malaysian shipments, however, were temperatures, as long as there is packed for their two-day or longer positive for the pathogen. adequate air exchange. Slightly cooler journey from Malaysia to the US. This Housing temperatures may also be acceptable seems to have improved somewhat This species, while not as nervous as long as the day period reaches in recent years. In the best cases, the or prone to jumping as many other moderate warmth. frogs are packed singly, generally in a rhacophorids, requires a sizeable The details of how R. plastic food storage container padded enclosure. Long-term housing for adult nigropalmatus spends the daylight inside with pieces of damp foam rubber animals should provide ample space. hours in the wild are unknown at this and some dried banana leaves. Due to Ideally, single animals or pairs should point. High in the canopy, they may the relatively calm demeanor of this share a vertically oriented enclosure be periodically exposed to intense species, rostral trauma and dermal 60x60x90 cm (24x24x36 in.) or larger. abrasions have been an uncommon Adequate ventilation is essential. occurrence. Along with the glass walls of the Animals are more apt to be continued on page 35 vivarium, these frogs will make use of successfully established if screened and

Notes on the Husbandry, Captive Reproduction, and Distinct Juvenile Form of Wallace’s Flying Frog | 33 Many frogs living high in the treetops make nocturnal journeys to the lower forest strata to hunt for food or to seek out mates and breeding sites. The ability to glide from tree to tree, or from the upper canopy to the undergrowth far below, may be an energetically inexpensive means of movement, and advantageous to frogs which would normally have to walk the gauntlet of arboreal predators. (Emerson & Cole, 1990) Aeronautic Amphibians Gliding behavior has evolved independently in at least two families of anurans, the hylidae and the rhacophoridae. Though the latter group is better known for its “flying frogs”, several new world hylids, including Agalychnis spurrelli, A. litodryas and Ecnomiohya miliaria are adept gliders, exhibiting both morphological and behavioral adaptations for a modified aerial descent. (Dudley, et al., 2007) By definition, “gliding” refers to a descent trajectory at less than a 45-degree angle to the ground. “Parachuting” is the term used to describe a steeper descent, falling at an angle greater than 45 degrees to the ground. (Dudley, et al., 2007) When a gliding frog leaps, instead of falling straight to the ground its modified interdigital webbing allows the frog to glide horizontally, and to maneuver through passages in the forest. For more specialized gliders, the speed of descent is slowed by their expansive patagia, arm flaps, and fringes, allowing for a softer landing with less chance of injury. These morphological qualities in conjunction with behavioral characters such as body posture and limb position, all contribute to overall flight performance for a species, which can be quantified by the distance traveled, flight speed, and maneuverability of a leap. (Emerson & Cole, 1990) A fine culmination of these traits has been noted in Wallace’s flying frog, Rhacophorus nigropalmatus. The species has been observed to glide at angles less than 25 degrees to the ground, cover extensive distances in a single glide, make 180 degree banked turns, and have a significantly reduced speed of descent. (Emerson & Cole, 1990) This fascinating aeronautic talent can be seen in the young frogs pictured here.

34 | Leaf Litter :: Winter 2009 sunlight, and may make use of that use distilled water, and avoid using any time of day and will take a wider heat and light for thermoregulation RO water as the main water source, variety of food items. In addition to and optimal UV exposure. Frogs in as these processes produce water that crickets and roaches, we have observed enclosures with an incandescent heat is too pure and may cause osmotic juvenile frogs to hand feed on tomato source seem to make use of it, and have problems with the frogs. (Wright hornworms, waxmoth larvae, and been observed to choose a sleeping spot &Whitaker 2001) freshly molted king mealworms. near the radiance. Placing the water dish on top of an Locusts and other small Lighting inverted bowl or plant pot to raise it orthopterans would be another good Ideally, a gradient of light and heat off the ground a few inches will help to food offering worth trying. should be arranged for the vivarium. keep food items from drowning in the In all cases, the length of the food This can be accomplished by placing water. To this end, adding a small piece item should not exceed the distance fluorescent tubes across the length of floating cork bark to the dish will between the frog’s eyes, which is a of the cage top, in combination with give the bugs a haul-out should they standard gauge for feeding many other a medium wattage (40w – 60w) spot fall into the water bowl. Most nights, amphibians as well. lamp above the top positioned to one the frogs will visit the water bowl and Disposition and Behavior side. The incandescent lamp should exchange their fluids. For this reason Wallace’s flying frogs are fairly provide a slightly elevated temperature, the water should be changed daily. calm animals in comparison to many ~ 32 C (90F), in the basking area. The frogs in our care, adults and other members of the rhacophoridae, Access to cover in large-leafed plants or juveniles, were exposed to a humidity which are often quite nervous and other shaded areas, at different levels range of 40 – 95%. It has been my jumpy in nature. Once acclimated to in the cage, further extends the options experience that many canopy dwelling the vivarium, these stoic frogs tend to of light and temperature exposure. frogs do not need to be kept at high choose fairly regular sleeping spots. We use fluorescent tubes with a ambient moisture levels for standard During daylight hours the frogs will color temperature between 5000K care, possibly because they already remain still on the glass walls and – 6500K, providing a natural visible possess adaptations to sun and wind large leaves in the enclosure, with spectrum. It’s very possible that these exposure in the wild. In fact, doing their arms and legs pulled tight frogs will benefit from tubes with so outside of breeding attempts may to the body to render themselves some UVB output. We are presently create unhealthy conditions. High indistinguishable amidst the foliage. experimenting with the ZooMed humidity paired with stagnant air Though decidedly nocturnal, they Reptisun ® 5.0 and 10.0 tubes in in an enclosure is a stressful, if not appear to keep their eyes slightly open combination with the Phillips 6500K lethal, combination for many arboreal during the day, just enough to keep tubes. The potential benefit of UV amphibians. Where ambient conditions tabs on their surroundings. While quite exposure is another large unknown are excessively dry, one can use a hand aware of their space, they will tolerate at this point, and is well worth spray bottle or automated misting momentary invasions into the vivarium researching for this and other canopy system a couple times a day to raise for maintenance and feeding without dwelling frogs. humidity in the enclosure. Live plants flight, sometimes turning their head also help to bolster ambient moisture Water and Humidity to get a better view of the action. At levels. Clean water is, of course, essential night, however, it’s a different story. for all amphibians. For most arboreal Diet and Feeding In the dark, they are like spring- species, including these, we provide Adult R. nigropalmatus will readily loaded acrobats. If you have to open water in a flat bottom bowl on the floor accept a variety of live invertebrate the enclosure at night, be sure to keep of the vivarium. Bottled spring water, food items, including crickets (Acheta a very close eye on the frogs and be or tap water that has been treated to domestica) and roaches (Nauphoeta ready for sudden leaps—they can be remove chlorine and chloramines using cinerea, Blaptica dubia). Wild-caught surprisingly elusive. adult gliders are quite leery of any Amquel or similar water treatment are Captive Breeding Notes movement near their cage and limit good choices for captive amphibians. In October of 2007, four weeks their feeding and most activity to late We also have good results using water after initial acclimation, two males and hours under the cover of darkness. passed through a canister of medical one female R. nigropalmatus were set Captively bred frogs, however, seem grade activated carbon to remove up for a captive breeding attempt at to be fearless: they readily feed at chlorine and other impurities. Do not SDR. The enclosure was modified to

Notes on the Husbandry, Captive Reproduction, and Distinct Juvenile Form of Wallace’s Flying Frog | 35 act as a flow-through rain chamber. Carbon-filtered tap water was run through a misting system to provide a light drizzle. A standpipe was situated in the drain to create a pool of water 6-7 cm deep over the entire bottom of the chamber. The rain period was controlled by a sprinkler timer, which was set to initiate at 4PM and shut off at 9PM After two nights of rain the frogs spawned, depositing a foam nest approximately 40 cm above the water level. A second group, set up in similar fashion, spawned one week later. In both cases the foam nest was discovered in the morning between 8AM and 9AM. At that time, the frogs were no longer in amplexus, so it was unknown as to which male (or males) contributed to the spawn, or the duration of amplexus. In these two instances, we can only deduce that it would have to have been less than 13 hours. The foam nests were gently pried off the wall of the enclosure while still soft, using a thin piece of plastic as a scraper. The nest was placed in a clear tray, which was suspended over an inch of water inside a larger bucket. A lid with light perforation was secured to the top to increase humidity. The mass was checked twice a day to monitor development (a few of the embryos could be observed against the transparent plastic). It was amazing to witness the development of these few eggs to embryos and finally tadpoles swimming through the foam. Soon, we could see more pale tadpoles moving through the froth. After five days the foam quickly liquefied and larvae dripped into the reservoir of water. The Top Left: An adult female Wallace’s flying frog (Rhacophourus nigropalmatus) next to her first clutch contained 115 tadpoles, and recently deposited foam nest. the second nest almost double at 220 Top Right: Five days after the spawn, the nest liquefies, allowing the larvae to wriggle free of tadpoles. We saw some infertile eggs the foam and drip into the water below. in the foam, but at the time of hatching Middle: Close-up of a hatchling swimming through the meringue-like foam. they had degraded enough to prevent a Bottom: A middle-stage R. nigropalmatus tadpole, showing unique color pattern on the tail count. muscle and fins. The tadpoles were transferred to

36 | Leaf Litter :: Winter 2009 three 190 L (50 gallon) breeder tanks longer received the powdered foods in turbid, stagnant, standing water— filtered by air-driven sponge filters. and were fed TetraMin-Pro flake fish conditions ideal for mosquito breeding. Submersible heaters kept the water food, Hikari sinking algae wafers, and With future spawns, we will certainly temperature between 24 – 26 C (75 – the P-Pel pellets. try some of these likely food options. 79 F). The tadpoles darkened to a The first metamorphic frog left the The tadpoles absorbed their yolks grayish-brown color, and as they grew water 50 days after hatching from the within a few days, after which time we they developed an attractive pattern on foam. The majority of the froglets were began feeding Sera Micron, spirulina the tail. Most of the time they rested out of the water within 72 days. Four powder, Hikari sinking algae wafers, on the bottom of the tank; because tadpoles were quite slow to transform, and P-Pel, an enriched sinking pellet they fed primarily from the bottom, the taking up to 120 days to finally made by Brine Shrimp Direct. Initial sinking foods were most efficient. They climb out of the hatchery. All of the losses were moderate and tadpoles remained motionless much of the time, metamorphs had no trouble climbing were observed eating dead siblings. but could be observed to gather around straight up the vertical polypropylene After two weeks the stable tadpoles the pellets and algae wafers shortly walls of the tub as soon as their front were transferred to one of our large after introducing the food to the tank. arms broke through. hatcheries. These are large capacity At night they could be seen swimming The metamorphs measured 41 – grow-out systems. Water quality is higher in the water column. 48 mm, including tail. After three to maintained by the use of fluidized In retrospect, it would have been four days the tail was fully absorbed, bed filtration, a bead filter, and UV worthwhile to try other high protein shrinking the froglets to 12 – 14 mm sterilization. food items such as live tubifex and SVL. The tadpoles were placed in one of black worms, midge and other insect The froglets were collected as they the 570 liter (150 gallon) holding tanks larvae. As mentioned previously, the left the water and transferred to glass on this system. At this point, they no tadpoles of this species are often found aquariums. Initially, groups of up to

Growing up: After a couple months underwater, these captive-bred metamorph Rhacophorus nigropalmatus are leaving the pond. Generally, they will climb out of the water without assistance, but they may also be manually removed, as soon as the front arms have emerged, and placed into a small rearing vivarium. The tail is absorbed gradually over the next three to four days, and after a week topside, they will begin hunting small invertebrates.

Notes on the Husbandry, Captive Reproduction, and Distinct Juvenile Form of Wallace’s Flying Frog | 37 Top left: A newly emerged, nearly translucent-orange Rhacophorus nigropalmatus metamorph. Top right: The juvenile form showing the ephemeral enamel-like dorsal spotting decorating a deep rusty dorsum. Left: Juvenile gliding frogs perched on a branch. At this stage their inter-digital webbing is developed enough to aid in gliding, a behavior that is exhibited at a very early stage, as well.

12 froglets were kept in 20 gallon standard aquariums, 61x30x41 cm (24x12x16”) with screen tops. These nursery tanks were outfitted with a large water bowl, sprigs of hydroponic aroid vines, and slightly damp paper towel substrate. Half of the screen top was covered with sheet plastic to contain some humidity. After three to five days the tails were absorbed and mouthparts modified. At that point the small gliders were split up further to prevent potential overcrowding issues. At seven days the frogs began feeding on one-week old crickets, which were dusted with calcium with D3 (Minerall [I]) consistently. A vitamin supplement (Herptivite and Repashy ICB) was added alternately. In some cases the food was provided in Pyrex dishes to increase food proximity. In others, the crickets were free-range, loose in the Bottom left: A transitional color stage of this species begins when the frog is about half grown tank. Both methods worked fine, but (48 – 50 mm). the dish method is more efficient and Bottom right: A ventral view of a juvenile glider, showing a bit of the developing dark patagia, helps to keep the cages cleaner. It also and enlarged adhesive toe pads. allows one to better track feeding. After gaining some size the froglets were also offered small roaches and wax moth larvae.

38 | Leaf Litter :: Winter 2009 Initial growth was slow. The young were thought to be a separate species, Novelty or Necessity?, Annual.Review of gliders seemed to grow in spurts, which would be an understandable Ecology, Evolution, and Systematics, 38:179– which usually followed graduation to assumption. 201 more substantial food items. Some Conclusion Duellman, W.and L.Trueb. 1986. Biology individuals grew more rapidly than At SDR, we hope to continue of Amphibians. Johns Hopkins University others. As a significant size difference our work in establishing the elegant Press, Baltimore developed, the smaller frogs couldn’t Wallace’s flying frog. At the time of Emerson, S.B. and M.A.R. Cole. Dec compete as well and began to lose writing, the F1 offspring are a little 1990. The Interaction of Behavioral and weight. When such size discrepancy more than half grown, and the young Morphological Change in the Evolution was evident, the frogs were relocated to males are just beginning to practice of a Novel Locomotor Type; “Flying” Frogs, enclosures with siblings of similar size. their door-knocking calls. Further Evolution, Vol 44, No. 8, 1931 – 1946 This helped considerably in getting the documentation of our observations will smaller frogs back on track. Emerson, S.B., J. Travis, and M.A.R. follow, and will include reports of any The overall time to maturity may Cole, Dec 1990. Functional Complexes and F2 offspring. be lengthy. At ten months out of the Additivity in Performance: A Test Case with Though wild collection is not water, the largest of the offspring is 65 “Flying” Frogs . Evolution, Vol 44, No. 8, 2153 thought to be a significant threat to mm. – 2157 this species, we really don’t know what A Distinct Juvenile Form effects it could have long term. It is Frost, Darrel R. 2008. Amphibian Species To our knowledge this was the first encouraging to think that a species of the World: an Online Reference. Version 5.2 captive breeding of this species. One of once regarded as difficult and variable (15 July, 2008). Electronic Database accessible the most exciting aspects of this novel to reproduce in captivity is a possibility at http://research.amnh.org/herpetology/ event was the discovery of the juvenile for sustained management. At a time amphibia/index.php. American Museum of color phase. when more than thirty percent of all Natural History, New York, USA. Just prior to leaving the water, amphibian species are threatened Grismer. L.L. 2008 La Sierra University, the tadpoles turn a rust color. At first I with extinction, anything we can do Dept of Biology, Pers. Comm. thought this was a fluke, but each one to reduce pressure on amphibians Inger, R.F. and R.B. Stuebing 1997, A Field of them climbed out a yellow, orange, should be done. Perhaps in the process Guide to the Frogs of Borneo, Natural History or rusty red metamorph. The color of developing captive husbandry and Publications, Kota Kinabalu, pp 190-191 only intensifies as they absorb their breeding protocols, we will learn tails. Over the first couple weeks, white more to complement the field study of IUCN, Conservation International, and yellow spots form on the dorsum, Rhacophorus nigropalmatus. and NatureServe. 2006. Global Amphibian and the skin becomes quite rugose. The discovery of the juvenile color Assessment. . Eventually these tubercules reduce, phase and subsequent ontogenetic Accessed on 29 Aug 2008. leaving the skin slightly granular. In color transition was thrilling, and a Malkmus, R., et al., 2002 Amphibians and the growing frogs, faint undertones of strong reminder that we still have a lot Reptiles of Mount Kinabalu (North Borneo), green could be seen when they reached to learn about the life histories of all A.R.G. Gantner Verlag K.G. pp 175, 215-216 lengths of 48 – 50 mm. At the time of amphibians. Even the ones we think McCay, M.G.. 2001. Aerodynamic writing, individuals measuring 57 – 59 we know so well have secrets to be Stability and Maneuverability of the Gliding mm SVL are almost completely green, revealed. Frog Polypedates dennysi. The Journal of but still show moderate spotting and a Bibliography and Sources Experimental Biology 204, 2817-2826 tinge of orange on the arms and legs. The largest frog mentioned above is Berry, P.Y. 1975. The Amphibian Fauna Taylor, T.H.. 1962 The Amphibian Fauna nearly solid green, with only a few of Peninsular Malaysia. Tropical Press. Kuala of Thailand, The University of Kansas Science white spots remaining. Lumpur, Malaysia Bulletin, Vol XLIII, No 8, pp 482-485 The juvenile color phase is de Vosjoli, P. 1996 How to Establish Wallace, A.R..1872 The Malay Archipelago: clearly a cryptic adaptation, but the Reinwardt’s Gliding Treefrogs (Rhacophorus The Land of the Orang-utan and the Bird of colors against a solid background reinwardtii) The Vivarium Vol 7, No 5 Paradise : a Narrative of Travel, with Studies of are absolutely stunning. It’s hard to Man and Nature. Dudley, R., et al. 2007. Gliding and the imagine that these striking young frogs Fucttional Origins of Flight: Biomechanical Macmillan and Co., London, pp 38-39 were never noted before. Perhaps they

Notes on the Husbandry, Captive Reproduction, and Distinct Juvenile Form of Wallace’s Flying Frog | 39 van Dijk, P.P., Iskandar, D. & Inger, R. 2004. Rhacophorus nigropalmatus. In: IUCN 2007. 2007 IUCN Red List of Threatened Species. . Downloaded on 25 July 2008.

Wright, K.M. and B.R. Whitaker, 2001 Amphibian Medicine and Captive Husbandry. Krieger Publishing Co. Florida, p 156

40 | Leaf Litter :: Winter 2009 Solace of Untouched Wilderness words and photos by Emily Lisborg

There was a perfect blend of green and brown between the dense flora and its remnants, which were scattered over a rich layer of grub-infested soil. The light was filtered by the trees so that just a few lonely rays reached the forest floor. In the few spots that it actually did you could see tiny lizards the size of a pinky finger basking for a moment, until they noticed they were being watched, at which point they scattered beneath the leaf litter. It was fulfilling, and being in there I didn’t think about anything but what was in front of me.

I don’t think I could ever be lonely in a forest, even if there were no people around. Some of the bolder lizards were on the trees, just hanging there. When they were scared, their throats would start to move up and down much faster and they would press themselves against the tree, trying to look bigger. They would eyeball you, and when they thought they could not last any longer they would scoot to the other side of the tree, disappointed at having to leave their precious basking spot. They would fight over the sun, at least the males would. They fought so fast it was difficult to tell if they even made physical contact; they must have, though, because the winner puffed out his crimson neck flap and started to do push-ups on the tree, marking his territory while showing off to the females.

The trees were like the ones in fairy tales, huge and imposing with gigantic roots all around their base. I was happy to be in the presence of such trees, but not a jolly skipping kind of happy or the kind of happy I feel when I get a good report card or play well in a game. It was more fulfilling than that. Everything made so much sense in there: everything was in equilibrium and everything fed off each other. A single tree had hundreds of tiny spider webs and lichen. Some of the smaller trees looked like they had

Solace of Untouched Wilderness | 41 been scraped with a knife. Their skin was peeling, only to reveal a healthier, lighter skin beneath. This pattern was all over the tree, giving the look of a collage. There were also spots that had been taken over by a primitive plant that appeared to be imbedded in the tree’s bark.

The rain brought out even the most reclusive animals, including a frog who called from a nearby plant. The plant itself clung for dear life to a branch by only a single root. Its cups held much more water than one would expect, and the crevices were deep. At its base were buds shooting out toward the light like a spiral. The bottom half of the frog looked moist and slimy from sitting in the water for so long. Deeper in the crevice was clear goo—it looked like snot and I thought it might have been skin the frog had shed, but when I looked closer I realized they were eggs: tiny frog eggs all clumped together, less the size of a small pea. They had to have been laid recently because there was no development, just perfect white circles in the center. You had to look hard for these, and you had to listen. It’s easy to miss everything, and better to walk slowly in a forest. The frog continued to call, which was especially loud and sounded much more like a bird than a frog, like somebody running their fingers over one of those cheap combs. The frog paused for about a second between each call to catch its breath, and each time his body inflated more. By the time it stopped calling it was a balloon. Had I tried to pop it, it probably would have flown in spirals before crash landing, all shriveled up and deflated.

The ground was brown with rotting and decomposing plants, and it would have been depressing but out of every single dying plant there was a new shoot of some kind: ferns sprouting like curly baby’s hair and trees emerging as small sticks with one or two meager roots. For the last half mile the trail kept getting wider and wider until it opened up to the ruins, like a small stream opening to an enormous ocean.

42 | Leaf Litter :: Vol 2 Issue 2 You could still see where the ancient walls had been, but most of it was crumpled and blanketed by nature. The back wall was still somewhat intact, but the front of the ruins was no higher than my legs. The contrast was striking: pale grey stone backing randomly scattered emerald ferns. The ferns were delicate and whenever the wind blew they danced. The forest seemed quiet, but when everyone stopped trudging there was noise everywhere. As the trail ended I looked up at the speckled sky, guarded by the trees that made me feel safe and protected, away from all the chaos.

It was just me and forest.

Solace of Untouched Wilderness | 43