Spindly Leg Syndrome, a Review by Ed Kowalski Photos © Ed Kowalski

Director’s Welcome

The Magazine of Tree Walkers International

Road to Stewardship There is a peculiar human drive to RON SKYLSTAD organize and categorize, and in the Editor conservation world it is no different. © Jason Brown When we first aligned ourselves organizationally with our goals at ED KOWALSKI TWI, we examined the individuals and organizations involved in NATHANIEL PAULL our corner of the amphibian conservation effort and looked for a Assistant Editors structure that would reflect the organization’s ethos. Eventually, we settled on “stewards” working as part of an amphibian “network.” And so, our loose grouping of captive amphibian keepers became the MARCOS OSORNO Amphibian Steward Network (ASN). Executive Director As we move forward, I ask for your help in the stewardship of two wonderful, interrelated things. First and foremost, in the continued BRENT L. BROCK care and conservation of amphibians and their habitat. In TWI, we Program Director all share a connection to our natural world through amphibians. Those who keep amphibians in captivity maintain a window into a natural habitat under immense pressures. This connection to nature MISSION STATEMENT is vital to maintaining a conservation ethos, and I am proud to be Tree Walkers International supports the part of a community through a wonderful hobby that is steeped protection, conservation, and restoration of in ethical traditions and perennially introspective in regards to its wild amphibian populations through hands-on future. action both locally and internationally. Next, I ask for your help in stewardship of the network itself. We foster personal relationships between people and nature by providing opportunities TWI and ASN are volunteer driven projects. Your involvement and for citizens of all ages to become directly generous membership dues and donations have made everything that involved in global amphibian conservation. we have done to date possible. The publication of the Dendrobates Through this involvement, our volunteers variabilis Taxon Management Guide is our first effort at managing become part of a growing and passionate animals within TWI. Taxon management will be a learn-as-we-go advocacy for the protection and restoration of wild amphibian populations and the process by fits and starts. However, I am confident that we, as a environment on which they depend. network, will see our efforts metamorphose into a viable, healthy conservation program. In the months to come, I look forward to our collective stewardship and to being a part of this process. Thank you for your ongoing support, and I’ll see you on the road ahead.

Sincerely, Marcos Osorno Executive Director

www.treewalkers.org

Vol. 1, Number 2 Winter 2007

Many who enjoy the hobby of keeping amphibians have a long cultured interest in a natural history that often has its roots sunk deeply within the sticky mud of a local wetland. My fondest childhood memories are of springtime romps to the nearest swamp to explore, inevitably leading to a cup filled with tadpoles, mud, and duckweed. My reluctance to leave the swamp led to many attempts in bringing it home with me. This took the form of contrived aquariums, terrariums, and various buried buckets, kiddie pools, sinks and even a cast iron bathtub. The small pools mostly grew algae and mosquitoes, but in the late spring after rainy evenings, a peculiar thing would happen: I would occasionally hear the distinct plop of a frog jumping into the puddle-ponds that dotted the backyard. The frogs would usually stay for a few days and then disappear. How could it be that such a small, delicate creature was able to travel the half-mile distance from the wetland to find these tiny aquatic oases? I have never forgotten the seemingly magical ability for nature to locate and utilize every potential niche, requiring only the proper invitation to do so. Since those early days in the swamp I have traffic circle in the very center of town. are 10’ x 10’, which is quite small by continued to experiment in creating After some convincing it was agreed ornamental pond standards, but a size ponds and bogs and am happy to share upon that the water feature would be that is easily manageable for a one my latest incarnation. modeled after a wetland and contain no person weekend project and requires In the summer of 2006 I set out to ornamental fish, no moving water, and nothing more than hand tools and a build a small aquatic landscape feature no mechanical filtration. little perspiration. The pond is oriented for my sister, whose home sits on a The dimensions of the pond site on the south-facing front of the house

Urban Wetland | and receives un-shaded direct sun population. For these reasons it is retain my substrate and keep it from throughout the day. This abundance important to not skimp on allocated eventually falling and collecting in the of sunshine is both a blessing and a area, and in this design it encompasses deepest portion of the pond. curse when building a pond. A body of half of the total pond area. It is planted Two substrate layers were used: water exposed to such an abundance with arrowhead, pickerel rush, and the first being organically composted of direct light will have a tendency cattail. topsoil (approximately a half-inch to grow copious amounts of algae The third zone is 5 inches deep deep) and the second consisting of unless the design and substrates of the and, upon the addition of substrate, expanded shale (3-5 inches deep). The pond instead encourage the growth of becomes an emerged bog area, allowing topsoil was used as an initial nutrient vascular plants. terrestrial bog plants to be introduced and mineral source for the plants. This pond was built with four for cover and live sphagnum moss for The expanded shale is affordable, depth zones, each of which promotes moist, but not soggy, ground cover. lightweight, does not decompose, the growth of various vascular Some of the plants included here are and provides a continuous span of emergent plants to out-compete the boneset, Joe-Pye weed, forget-me- substrate for the planting and growth algae. The deepest of these zones is nots, horsetail, and various sedges and of aquatic and emergent vegetation. I in the middle of the pond and has a rushes. am fortunate enough to have friends depth of 24 inches, which I have found The fourth and final zone is with ponds and wetlands who allowed deep enough to prevent solid freezing completely terrestrial and provides me to collect the majority of my here in Southern Pennsylvania and foraging cover as well as habitat for planting material. I dug up the plants, is important for the over wintering of prey. taking as much care as possible to not amphibians. Winter-hardy water lilies damage the roots and include an ample were planted in this zone along with a I used a flexible liner to act as my amount of natural wetland soil. I feel variety of other aquatic plants. water barrier and decided not to use that it is very important to also include an under-layment because the soil was The second zone is the marginal ecologically active “seeding mud” to not rocky, and round rocks were used kick off the ecology that will support zone and is 12 inches deep. This zone is for the landscaping. These rocks were the ecological workhorse of the pond, a technologically devoid pond. These built up on the back side of the pond intact microscopic communities work providing the majority of vegetation to retain the soil and hold up the liner. necessary for nutrient uptake and key with the plants to turn nitrogenous Each change in depth zone was backed wastes into plant matter. reproductive habitat for the amphibian by the rocks to create “planters” to

Dense growth of emergent vegetation is very important for the health of the pond when using no mechanical filtration and the pond is oriented in direct sunlight.

| Leaf Litter :: Winter 2007 CLOCKWISE FROM UPPER LEFT: The urban wetland changes throughout the growing season. It is shown here in spring, late spring, and late summer.

The seeding mud, along with the to the amount of sun. I also partially The plants began to die toward the mud attached to the plants, weighed attribute this to my organic compost end of October and after they turned in the neighborhood of 200 pounds. sub-layer (in future projects I may brown, I cut back and removed all of The plants were collected in naturally exclude this). The pond was less than the dead vegetation before it naturally occurring, non-selective “bunches” of attractive during this phase, but it fell into the water (too much decaying various species and planted with as subsided just a month later. matter in the bottom of the pond can little disturbance as possible. Some The first amphibians began to lower oxygen levels and the pH of the plants were purchased, mostly for appear following a rainy, early autumn water during periods of ice cover). I cut ornamental reasons, and included evening after the pond had acclimated the hollow stems of the cattails so that bog bean, pitcher plants, ornamental and was approximately three months they were above the water to facilitate grasses, and sedums. old. Pickerel frogs and green frogs were air exchange below the eventual ice. Immediately following the planting observed through the end of October. This past spring, as the plants woke and the addition of water, the pond These frogs had to traverse roads, from their winter slumber, the water went through an acclimation phase backyards, and sidewalks for three- remained very clear—the plants grew denoted by the occurrence of a serious quarters of a mile to make it to the in so densely that there was much less algae bloom. This was expected due pond. opportunity for algae to take hold. The

Urban Wetland | frogs also emerged after surviving their spring and hope for some reproduction is ripe for discovery: Just imagine first winter. Calls from both green frogs from the residents. a neighborhood or block in which a and pickerel frogs were heard but no Building this small puddle-pond small wetland were in each yard, or egg masses were found. Several species has proven to be a very rewarding a suburban housing development’s of dragon and damselflies have also project and has surpassed my under-utilized storm water retention been observed. expectations. I went to photograph area being converted into a wetland In the summer months, the pond for this article, and as I was sanctuary. The ecological diversity of during periods of little rainfall, it leaving that evening I found a juvenile that area would be increased many was necessary to add water due bullfrog sitting in the middle of the fold and deliver an opportunity for to the amount of transpiration by sidewalk about two blocks away, and I ecological incorporation that so many the emergent plants. A float valve wondered how many other people had of us and our children lack in our daily connected to a water line was added to had such an encounter since the pond’s lives. The conservation of amphibians maintain the desired water depth. This addition. The pond’s public placement in faraway places shrouded by rain type of device could easily be connected allows for pedestrian viewing and many forests is important, but the key to to a cistern of collected water in order people stop and look on their evening their success lies in the personal to facilitate water conservation. walks to see what is happening on that relationships we have with nature, day. New flowers are blooming and relationships that must be nurtured In the future I plan to install solar and fed on a daily basis at an intimate powered compact fluorescent lights different frogs are calling on a weekly basis. level. I encourage everyone to build in order to attract insects that will one of these ponds and feel that the help further support the amphibian The ecological potential for potential benefit for amphibians and population. I look forward to next residential and urban environments humans alike is great.

A traditional ornamental pond can function as temporary frog habitat, but the fish will consume larval amphibians and the lack of a soft, bottom substrate does not facilitate over-wintering.

| Leaf Litter :: Winter 2007 Adapting Quarantine Practices to the Amphibian Hobbyist’s Collection by Matthew Campbell photos © Matthew Campbell

uarantine is a Because our zoo hoped to work Quarantine 3: This level pertains to concept most with endangered species of native quarantine of animals for display or amphibian United States amphibians, we needed exhibition purposes with no intent to hobbyists to observe firsthand how other zoos re-release to their home range. think about were successfully addressing the It is worth noting at this point Q only when issue of quarantine of amphibians in that the average hobbyist does not receiving new an “ex situ” environment. Standards maintain animals that are likely to wild-caught have been set and adopted for be considered for reintroduction specimens. Even then, many of us have various levels of quarantine in the to the wild. As TWI works with probably been guilty of keeping those ex situ environment. The extent and hobbyists, zoos, and other groups, new wild-caught animals in proximity rigorousness of quarantine depends some individuals may eventually to our established collection. We may on the final disposition of the animals be identified who have colonies of have been lax in washing up between in the collection. Animals that are animals that may be considered for cleaning chores or perhaps we have being held with the intent of being reintroduction. For the time being even used tools with multiple animals. reintroduced to the wild at some point though, the average hobbyist can learn If you’ve always had good quarantine are subjected to the most rigorous some simple “best practices” regarding practices, I applaud you. However, quarantine. Animals which are not quarantine and general maintenance many of us know we’ve been guilty of intended to be reintroduced to the wild, of their collections that will go a long lapses in observing proper quarantine. whether they are wild-caught origin or way towards preventing introduction Some have paid the ultimate price from an existing outside collection, are of diseases to our native amphibians in having well-established animals subject to a less rigorous quarantine. as well as preventing acquisition of diseases by our captive animals. succumb to newly introduced infections There are three levels of while others have managed to dodge quarantine: What I observed at HDZ was that bullet. not only how quarantine can be set Quarantine 1: The primary concern at Last May, I was able to travel to up and maintained but how simple this level is with amphibians where attention to the design and setup of an Omaha, Nebraska’s, Henry Doorly the intent is for re-release to the wild. Zoo to view firsthand the dedicated The main goal with this level is to amphibian facility can help streamline amphibian quarantine facilities they prevent the quarantined amphibians daily care. Whether you have a bank of had recently constructed. As a full- from acquiring new diseases, which can lush display cages or are wanting to set time zookeeper, I had nominated then be introduced with them into wild up a simple breeding colony, there is populations. Also, this level strives to myself to be one of the “point men” for much you can learn from seeing a well prevent passage of disease from the designed quarantine facility. amphibian conservation projects at quarantined animals to other animals our zoo. Our zoo was looking to better in the collection or release to native Quarantine 1 standards call educate keeper staff on amphibian amphibians. for housing one species to a single husbandry and quarantine to help play Quarantine 2: This level deals with “in containment room or, at the very our part in the ongoing Association of situ” quarantine facilities and has no least, housing assemblages of animals Zoos and Aquariums (AZA) initiatives direct application to procedures that to a single room. An example of an might be adopted by the average geared towards accelerating amphibian assemblage would be restricting a hobbyist. conservation. room to only animals from a certain

Adapting Quarantine Practices to the Amphibian Hobbyist’s Collection | ABOVE: Both frogs shown were from the same breeding pair, but the frog in the top photo was raised in a vivarium without a naturally- occurring nutrient cycle and the other frog was raised in a vivarium where arthropods were supported by the soil and leaf litter.

geographic locale or country. I won’t 1. Have dedicated tools and containers animals cannot come in contact go into specifics here but instead for quarantine animals and use un- with quarantine animals and that will endeavor to point out how we as powdered latex or synthetic gloves native animals cannot in ltrate your hobbyists can adapt these procedures when handling all amphibians. quarantine area. Dispose of gloves and other waste by to our own home collections to improve 4. Service your quarantine animals only rst immersing in a bleach solution their overall health and lessen the after you have completed all other for a minimum of 24 hours before servicing tasks with your regular chance of potentially introducing putting in common garbage. collection. pathogens from our collections to our 2. Treat all waste water from What follows are some wild native amphibian communities. quarantined animals with a bleach photographic examples of the HDZ solution for a minimum of 24 hours At the very least, I would urge quarantine facility. You can use these before discharging to a sewer. hobbyists to adopt some simple photos to get an idea of how to set up 3. Ensure that quarantine animals practices to prevent disease an efficient quarantine facility of your transmission when dealing with wild- are secure and cannot escape and come in contact with other own or simply to give you ideas as to caught animals or animals of unknown collection animals or escape into how you can better organize and design origin. the surrounding native habitat. your own amphibian room. Likewise, ensure that other collection

UPPER LEFT: HDZ houses individual species in a single room. The average hobbyist need not build individual rooms; however, grouping animals in natural assemblages can help in better regulating husbandry aspects such as temperatures and photoperiod. MIDDLE: Housing your collection in a thermally stable area such as a basement or air-conditioned room is ideal. If you have difficulty keeping your temperatures stable, consider using a room air conditioner/ heater unit such as one of these. In rooms without windows or which have no central air conditioning ducts, these units can help you efficiently control your temperatures. RIGHT: Take into account the specialized needs of individual species—such as inde- pendent filtration—and incorporate these into your design. LEFT: A well-organized workspace will go a long way toward helping you streamline your maintenance and cleaning.

| Leaf Litter :: Winter 2007 If your goal is to breed, you may not always be best off with lush display food storage boxes were utilized to house both aquatic and terrestrial vivaria. Instead, utilizing simplistic setups that meet the animals’ needs species. Individual animal data and feeding records are kept on the may be the most efficient route to take. At HDZ simple polycarbonate front of the enclosures.

LEFT: Each box was plumbed with a bulkhead drain and standpipe, allowing for ease of cleaning and water changes. Setting these boxes in a utilitarian rack system allows them all to be plumbed to a common drain system, further streamlining cleaning and setup. ABOVE: Liberal use of caps, unions, and valves will mean that you can easily reconfigure your setup should you need to move it or change the layout of your tanks.

Adapting Quarantine Practices to the Amphibian Hobbyist’s Collection | UPPER LEFT: The common drain pipe empties into a receptacle where the waste water can be pumped out via a sump pump to another receptacle where it can be treated with chem- icals before being discharged to your sewer and thus to your local waste water treatment facility. Remember, during severe thunder- storms, water you empty to your sewer may get diverted to a storm drain where it is then discharged to local waterways without being treated at a wastewater treatment facility. This is one potential avenue for introducing non- native pathogens to our native amphibians. UPPER RIGHT: Wastewater from cage cleaning is pumped to these barrels where it’s treated with a bleach solution for a minimum of 24 hours before being dumped into a drain. LEFT: Use of reverse osmosis filtering of your tap water supply can remove harmful pollutants not screened out by your local water treatment plant. Also, you’re assured that pathogens not screened for are removed entirely from your water before it gets to your amphibians. Remember to reconstitute your RO water with a product like Kent RO Right before using it with your amphibians. Using a carbon pre-filter for your RO system (shown at left with blue cap) will help extend the life of its membranes. Using disinfectants where appropriate can help in preventing disease transmission. Also, prominently labeling your disinfectants can prevent costly mistakes!

| Leaf Litter :: Winter 2007 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.

The rain stopped mid-morning. an amazing variety of frogs, I sat on In the drainage ditch outside our the hotel veranda outside our room end room hylids had called in the making notes and watching the equally gentle morning rain. The day before, amazing people of Amazonia parade up our first in the jungle, wasted me. and down the main street of town. So earlier this morning I decided to The morning was pivotal for me. stay back in the Chazuta hotel while The rain was light and comforting. The Todd and Tim hiked up into the notes I started on the veranda reflected mountains. While they were out seeing both my excitement and fears. The

ABOVE: Around Chazuta Epipedobates bassleri were predominantly orange with varying amounts of red in them. Also a common leaf litter frog, their calls were frequently heard. LEFT: Looking into the mouth of the Huallaga canyon leading to Chazuta and an amazing diversity of Dendrobatids.

photos © Tim Paine unless otherwise noted

Tarapato: impressions | A view upriver of the Huallaga shortly after it merges with the Rio Mayo. Tarapoto lies in the valley at the base of the most distant range seen in the center of this photo RIGHT: Todd Kelley, left, wrangles a frog while Tim Paine photographs it. © Tracy Hicks

day before had been good, productive immediately off, climbing through was very intense so I didn’t expect to and much, much harder than I had “cultivated” fields of banana and sugar struggle too much while hiking and expected. cane, up the primeval river banks into climbing. But this first hike the entire The first full day in Peru we drove the jungle. way up and into the jungle was literally the washboard “highways” to Chazuta. I knew I was not as physically breath-taking and physically painful Almost as soon as we arrived we took capable as Tim and Todd—they are enough to scare the crap out of me. a fast boat downriver to a farm where both close to my son’s age. Over the Our group that day included our we were told we could possibly find many years since I was their age a guide, Caesar, his friend who owned some Dendrobates fantasticus. The series of heart attacks has left me with the boat, and the farmer who decided farm was “rustic.” The farmhouse itself a scarred heart muscle. Before I left to join our odd crew of frog hunters. was a platform with some planking home my cardiologist approved the A mile or so into the jungle and for walls. Several generations of a trip, but only after a new medication before the trails played out we began family were living in a space defined became available, which removed most hearing D. imitator-like calls. Todd by a few planks and sheets of hanging of the chronic angina. Without this, I and Tim immediately scrambled cloth for walls. After the strongest would not have been able to go. down into a creek bed below the trail. looking man of the group greeted us For the six months before the trip I I planted myself on the trail close to and took a dollar or two as payment for worked out diligently at the gym, three two or three calling frogs and searched our intrusion onto his land, we were to five days a week. The preparation the area without consciously moving

10 | Leaf Litter :: Winter 2007 anything but my eyes. Caesar and the giant palms were magnificent. They Let me break the story line to make local guys went a little further and sat were awe inspiring in the way I can a point here: along the trail. The frogs kept calling. imagine a magnificent cathedral would The three of us, all close friends, I tried to stay perfectly still, but soon be to a devout Catholic. I remember were in Peru to see these dendrobatid realized I was continuing to gasp for thinking how surreal it seemed to be frogs in their native habitat. Tim breath. The local guys were eating experiencing this form of religious Paine is a gifted and highly trained oranges they had gathered along the experience in a Peruvian jungle. photographer of herpetology. Todd way up from the river. Caesar brought I remember thinking how some Kelley is one of the most sensitive me one and the farmer gathered a images on Evan Twomey and Jason frog breeders you can imagine. I am “medicinal” plant for me. The plant Brown’s website (www.dendrobates. an artist whose collections document tasted medicinal, and seemed to org) now made sense in an entirely changes in life. immediately lessen the damned different way. Standing there in angina. Although we approached the the jungle for the first time, two experience of this trip from our Todd and Tim soon returned dimensions suddenly made the leap individual perspectives, we knew with a beautiful group of yellow- to three and even four. Damned that that we shared a deep respect for banded Dendrobates fantasticus. The was exhilarating! Everything was the animals in their habitat. We excitement level was high and my balancing. The sounds of the jungle collectively captured the frogs, Todd breathing problems were immediately around me were reminiscent of my wrangled and Tim photographed them. forgotten, if not completely gone. frogroom…but this was a frogroom on The frogs were then returned to their After the trail played out, we steroids. environment. Our shared goals were to became the stereotypical image of A little later and further up the document and experience the animals intrepid bushwhacking explorers with steep incline we stopped in an open with as little adverse impact on this guides clearing a trail through the thick area to rest. Much lower in the climb rich jungle environment as possible. jungle undergrowth. The occasional we had heard D. imitator calling. Now, Had the frogs not called from the clearings were where the beauty of while we rested, they began to call bags of these first D. fantasticus, they the jungle popped into focus, leaving again. This time, though, they called would all have been labeled in our my North American mouth gaping. from inside the bags of our collected D. notes as D. fantasticus. They were When seen from some distance, the fantasticus. certainly fantastic looking frogs in every way. But at least some of them had calls that are typical of the D. imitator group. So without genetic studies on this specific locality, we will never know if these frogs in the bags were a mixture of both species, all D. imitator, or even some other species. The hike back down to the farm included finding our first snake of the trip. As I was trailing the bushwhackers by a short distance, one of the local guys yelled “viperosa!” with fear in his voice. Most of the crew surged back toward me while Tim and Todd lunged forward. Tim soon came Dendrobates imitator (yellow banded): back to the macho local guys, who Dendrobates fantasticus or D. imitator? then began giggling over the small vine At first these were thought to be the snake wrapped around Tim’s hand. But former, but the staccato trill identified these frogs as D. imitator. they still felt it was poisonous and kept their distance. The terrain of the hike back down

Tarapato: impressions | 11 Dendrobates ventrimaculatus Dendrobates variabilis was found near a stream bank a found within bromeliads one short hike outside of Chazuta. to two meters above ground.

In areas, all of these dendrobatids — and more — could be found within meters of each other.

Hyloxalus (Colostethus) nexipus This monotypic (only one species in the were sometimes found in great genus) frog, Edalorhina perezi, is cryptically numbers breeding around large colored and shaped for its leaf litter world rocks and small pools found and diurnal lifestyle. at the edge of rivers and large streams.

12 | Leaf Litter :: Winter 2007 to the river took us repeatedly up and This microteiid, Cercosaura argulus, was found resting on down and across and through creek a leaf 1-1/2m above ground just meters from where the beds and back up into the thick jungle Dendrobates variabilis was found. vegetation, bushwhacking most of the way. I followed the whacked path as best I could but lost my way a time or two when the natural clearings obscured the point where the machete- wielding farmer had chopped his way back into the thick of the jungle. Caesar eventually stayed back behind the faster moving group to lead me out of the jungle. As a native Texan with lots of East Texas forest trekking, I was a little embarrassed but really appreciated Caesar’s help. Actually, I was more than a little embarrassed at my inability to keep up with the others. This was only the first day and I feared that I would inhibit my companions’ experience over the next two weeks.

I’ve debated with my ego about whether or not to include these more personal aspects of the trip. After all, my others were all resting on the porch Tim acted as our interpreter (his specific situation is uniquely personal and facing the trail and the river below. Spanish was better than Todd’s, and can be read as negative. But I believe this Todd, bless his soul, had saved me my ear for language is abysmal). We is not a totally uncommon experience. some water. The worn, chainsaw- were hungry, and despite the cost Many people have similar fears and hewn planks we sat on were more of only a few dollars, it was a truly realizations in their travels. I feel that comfortable than I can even begin to great meal of roasted chicken and the realizations on this trip and in this explain. I would have loved to stay tamales…all eaten in the front room of storyline far outweigh any possible there for the night and shared some of someone’s home. negative interpretations. the farmer’s homemade brew with him Great meals are arbitrary terms. and his generations of family—sitting We ate some very good local food Somewhere along the final few there on the porch they seemed more that would have seemed meager back hundred yards I vividly remember civilized than their tattered dirty home. But there, in the backwaters of gagging for breath and dropping to my clothes could have suggested. But after the Amazon, the food was delicious. knees. Caesar came back to check on a short rest we headed back on the The basic aspects of life there also me, and I tried to avert the dilemma river to Chazuta. lacked luxury: toilet seats are rare in by, in very broken Spanish, asking That night, Tim, Todd, and I Amazonia. Our room in Chazuta was about the orchid growing beneath me. wandered down the long main street one of the few with toilet seats we Everywhere I looked, this environment of Chazuta looking for a place to eat. encountered. was marked with plants unique to People stood around grills cooking my experience. I remember wanting So as the rain fell on this second their dinner. Some people were to collect the orchid I nearly threw- morning in Peru I sat on the hotel bartering for food. Others appeared up on…and knowing that was not a veranda reassessing the first day and to be cooking for family groups. possibility. wondering if I had made a mistake in Eventually someone smiled at us in a coming. The prospect of holding my When Caesar and I eventually way that suggested we were welcome companions back was unappealing at reached the farmhouse, the to come and eat what she was cooking. best.

Tarapato: impressions | 13 When I saw the frogs they found These bright red walking sticks were a on that morning trek up into the common sight. mountains the idea of leaving or staying slammed like a door in my face. How could I not stay and see these frogs? Yet how could I prevent holding back my friends? I figured that I could work easier areas while they took the harder trails. I could mark everyplace we went in my own way. In the hotel courtyard that afternoon I watched as Todd wrangled and Tim photographed the frogs they had collected: Dendrobates imitator, D. variabilis, D. fantasticus/imitator (from the day before), Epipedobates trivittatus, E. cainarachi, E. hahneli, and an amazing orange form of E. The veranda was on the main ducks, chicken, turkey and various wild bassleri. They had also managed to street of town. If you stood in front of birds foraging through the magenta- collect some D. ventrimaculatus- the hotel you were standing directly coated scraps. At one point a mare looking frogs that Todd, judging in the street. A short distance from and her colt were flushed out of the by their actions, deemed to be D. the other side of the street and across creek bed by a large boar that had been imitator. a narrow strip of land, the Huallaga hidden in his waller by the pink petals. After dark that evening it rained River boiled by. Going with its flow Women walked up and down the I watched a dugout canoe with two again. We had just returned to street with bundles. Some sold tamales the hotel from another dinner in young men and a dog catching fish. from bowls balanced on their heads. One man was in back and used the oar someone’s street-side home, when One older woman sold four foot long out of the dark rain a barefooted man, to maneuver in and out of the current, dried bean pods with sweet cotton-like gliding down to a spot where the water struggling under the weight of a 10- fiber inside. Kids, some in uniforms, foot-long, 6-by-8-inch beam, climbed rolled around the bank and into an most with simple packs and books, eddy where they threw their net. The the porch steps. As he passed me I hustled off down the street to school could see he only had one functioning dog barked as the occasional fish was in the morning rain and then back up caught. eye. The other was clouded white toward home in the midday shower. and distorted. I was amazed he could Between the river and where Older young people occasionally handle the load from the river to the I sat, the open land was bordered zipped up or down the street on sawmill behind the hotel. I was even with a creek that, over the years, motorcycles. On rare occasions a more surprised when I turned around had turned into a drainage ditch. As pickup truck or an old Toyota Corolla and saw an older man, who I judged to I sat writing, the local women and would splash through the puddles be the first man’s father, following with girls dumped their household trash and stop at a bar not far away, where a beam of the same size. I surmised and scraps into the ditch. While that people drank cerveza while loud salsa that they floated the beams (which I’d may sound repulsive, it was the most music played. guess weighed close to three hundred beautiful dump I’ve ever seen. The In the early afternoon, Todd pounds) into Chazuta from some banks were worn down exposing the and Tim returned down this same upriver jungle. These beams were roots of the trees. One gigantic tree street. From a long distance I could probably milled into mahogany chairs was blossoming. Magenta petals tell they were dragging and obviously now sitting in some Chazuta home. carpeted the rolling trash-covered exhausted. It was apparent that I had The tenacity of these Amazonian land and turned the scene into a far made the right decision to sleep in that people deserves our admiration. The off-Broadway play on a surreal world. morning. people we encountered all along the I watched dogs, horses, cattle, pigs, way earned our deep respect.

14 | Leaf Litter :: Winter 2007 by Danté Fenolio The Atlanta Botanical Garden’s Center for Conservation photos © Danté Fenolio

or nearly as many years within the treetops stream downward, as humans have been typically after dark, reminding exploring the tropical biologists that a community of life F forests of the planet, forms exists beyond their reach. there has been an Science has subsequently come to awareness of a special community learn that rainforests are home to of organisms that spend most or all the greatest percentage of the Earth’s of their lives high above the ground biodiversity and that the canopies and within the forest’s canopy. The of these forests are the part of the vocalizations of unseen denizens ecosystem that harbors the vast

Here are two of the tools that the CRP is famous for: the blimp that places the canopy raft and that drags the canopy sled. In addition, the one man balloon that is tethered to a long cable running up to a mile along the upper canopy is pictured. INSET: The one-man balloon is tethered to a canopy cable that spans a distance of nearly a mile. Biologists are lifted up by this canopy ascension tool, enabling them to collect samples as they move along the cable.

Canopy Amphibians | 15 BELOW AND LEFT: Trachyderma resinifictrix is a tree hole breeding species from the Amazon Basin. This species will return to water-filled tree holes that contain their tadpoles and deposit fertile clutches of eggs, which are quickly consumed. This is one of the few tree hole breeding species that has been regularly bred in captivity.

16 | Leaf Litter :: Winter 2007 majority of plants and animals living has gravitated. A number of “canopy there (Nadkarni and Lowman 1987; specialists” have evolved that no longer International Canopy Network 1999; require visits to the forest floor. This Kitching et al. 1993; Yanoviak 1999). is not to say that these species can’t In rainforest canopy communities, be encountered closer to the ground many organisms, including and on the forest floor. Rather, these amphibians, are not equally distributed amphibians no longer have the need but concentrated around resources to visit traditional egg and larval like plant-held waters (Fincke 1992; deposition sites on the forest floor. Grillitsch 1992; Yanoviak, 1999). These They don’t have to hunt prey at ground aquatic habitats draw amphibians and level nor do they need to seek mates are made when water accumulates there. Many of these species spend within tree holes, Pandanus and palm much of their time within higher levels axils, pitcher plants, and clusters of of the canopy and some appear to air plants such as bromeliads (Dunn almost exclusively reside there. The 1937; Wassersug et al. 1981; Lim 1991; trick to understanding each of these Denzer 1994; Fenolio et al., 2007). species lies in investigating how they These plant-held waters are referred to broke their attachments to the forest collectively as phytotelmata (Maguire floor. 1971). Such lofty aquatic microhabitats You might ask why a species are exactly one of the resources around would evolve in this direction? The which a special guild of amphibians answers may involve predation in

Cruziohyla craspedopus is a canopy specialist from the upper Amazon Basin. They breed in water-filled tree holes, stumps, and crotches formed in depressions between branches and the main trunk of a tree. They have fewer eggs than other species of related Agalychnis, accommodating life in a small volume of water for their tadpoles.

Canopy Amphibians | 17 The French-run Canopy Raft Program (CRP) is famous for its blimp that carries the canopy raft to a spot on the canopy where it will subsequently be tethered. The canopy raft enables scientists to study life in the canopy day and night. My experience on the raft was amazing—sleeping in sleeping bags tethered to the raft is quite interesting. Everything is fun…until it rains.

18 | Leaf Litter :: Winter 2007 traditional egg and tadpole deposition and can be crowded—private water aquatic habitats (outside of an egg) are sites. Tadpoles living in forest pools sources high within the trees would be no longer required and the tie to the and streams are subject to the an attractive alternative. Conversely, forest floor is broken. Regardless of guild of aquatic predators that will adults of some of these species may the force that drove species to begin a undoubtedly be found there. Avoiding have moved into canopy habitats lofty canopy existence, phytotelmic life these dangers might push a species where there was less competition styles have in fact been the answer for to seek new larval habitats free of for food and moist microhabitats some species. these predators. However, this is in the adult life stage. Beginning to So how has science been able to not the only proposed hypothesis for utilize arboreal aquatic sites may study these species? Because canopy an evolutionary trajectory toward have evolved after the preference for exploration techniques have arisen permanent canopy life. Competition canopy living arose. Other species relatively recently, an understanding with other amphibian species that seem to have been predisposed to a of the biology and ecology of these deposit their larvae in ground level canopy existence through an evolution amphibians has been difficult until circumstances might be a force strong around a free living larval stage. For recent times. Canopy exploration enough to drive species to look for example, Bolitoglossine salamanders techniques have been developed alternative larval sites. Terrestrial and a variety of frog lineages have through programs like the French- aquatic habitats have limited resources directly developing larvae. Larval run Canopy Raft Program, Radeau

Canopy Amphibians | 19 20 | Leaf Litter :: Winter 2007 Des Cimes (Hallé 1990). The tools include the canopy raft, a giant sled that is hung below a blimp and drug along the canopy, giant tree fort units tethered to the treetops, one-man balloons tethered to cables that run long distances along the forest canopies, traditional arborist techniques that allow scientists to ascend into the canopy, giant canopy cranes, and canopy walk systems that allow researchers to leisurely walk through the habitat and collect data. Each system has its advantages and disadvantages. Some systems tend to drive the local herpetofauna from the area via disturbance. Some don’t allow for study of extended tracts of forest. Regardless, these techniques have provided an avenue through which much insight has been gained into the biology and ecology of canopy amphibians. With each passing year, the tools improve and our ability to observe canopy amphibians grows. Even with these advancements, a few canopy species are as mysterious now as they were fifty years ago. For example, very little has been published on the biology and ecology of upper Amazon Basin species such as Nyctimantis rugiceps or Ecnomiohyla tuberculosa. At least with Nyctimantis rugiceps, we know that they breed in water-filled tree holes, generally high within the forest’s canopy. Little else is known of their ecology. Virtually nothing is known of Ecnomiohyla tuberculosa. New species remain to be discovered. For example, I was fortunate enough to participate in one of the French Canopy Raft Program’s To date, arborist techniques have served as the primary methods of canopy exploration. expeditions to NE Madagascar along Skilled climbers were brought to Madagascar by the CRP to assist groups of scientists. Nouie the Masoala Penninsula. One of the Baiben was allotted to the team I was a member of and assisted us tremendously in our new canopy species we discovered on search for new species of canopy frogs. that project is highlighted in this piece, Anodonthyla hutchisoni (Fenolio et al., 2007). We were thrilled to discover

Continued on page 24

Canopy Amphibians | 21 BELOW: Many dendrobatid frogs qualify as canopy species. Remember, canopy species aren’t defined by strictly inhabiting the upper canopy; instead, they can exploit various canopy layers and no longer have an obligatory tie to the forest floor. One of the most popular species maintained in captivity is Dendro- bates fantasticus. While this species may not be the most arboreal of the dendrobatid frogs, arguably some populations of D. fantasticus do exploit lower canopy layers. Additionally, the species does transport hatched tadpoles to phytotelmic circumstances, largely utilizing bromeliad axil pools. Often, they will also deposit clutches of eggs within bromeliads as well.

ABOVE: The “ICOS” units developed by CRP are tree fort units tethered to the upper branches of trees. They allow scientists to live in the forest canopy for days on end. Our study of vertical stratification in the herpetofauna of the Masoala Peninsula was based in these ICOS units. We performed timed surveys at various heights, searching for herpetofauna at every canopy level, while hanging onto ropes suspended from the ICOS units above us.

22 | Leaf Litter :: Winter 2007 RIGHT and BELOW: Gastrotheca cornuta is one of the seldom encountered canopy species from Mesoamerica. They spend the majority of their time high above the ground and out of the normal search radius of biologists working from the ground. The characteristic that allows this species to remain in the canopy is its reproductive biology. Females have a pouch in their dorsum that holds fertile eggs through hatching. The hatchlings develop in this pouch as well so that fully developed froglets emerge. There is no free living tadpole phase and no need to deposit eggs into an aquatic ecosystem. Tremendous strides have been made in breeding this species in captivity through the Atlanta Botanical Garden’s amphibian conservation program. The image here of a female holding eggs in her dorsal pouch was taken in their world class amphibian breeding lab.

LEFT: One of the lineages of frogs that has evolved direct development are frogs of the genus Eleutherodactylus. This is a male E. coqui from Puerto Rico guarding a developing clutch of eggs. He was found at the base of a bromeliad axil, just above the pool of water. Eleutherodactylus are often found within various canopy layers. In Puerto Rico, this species is often found in bromeliads well above the forest floor.

Canopy Amphibians | 23 a tree hole breeding microhylid frog. for permeably-skinned creatures. changes create variable and commonly The bottom line is that much work still A break from having to visit the dry conditions in the upper canopy remains before we have a good idea forest floor is only part of the story. (Fitzjarrald and Moore 1987). For how canopy amphibians make a living Adaptations are necessary to become a example, the upper forest canopy in the treetops and how diverse this denizen of the upper canopy whether tends to ventilate better and more guild actually is. they are behavioral, physiological, rapidly than lower canopy levels and Life in the canopy comes with morphological, or a mix of these. the forest floor. These upper levels its own set of difficulties, especially Drought, wind, and temperature also receive harsh sunlight exposure.

LEFT: A tree hole breeding species from isn’t to say that a species can’t be found A tree hole breeding species from Central America, Southeast Asia and Borneo is Nyctixalus at lower levels within a forest canopy. Anotheca spinosa, is a canopy specialist. The species pictus. They deposit eggs in tree holes This species can be encountered closer to deposits eggs on the walls of water-filled tree holes. Af- where tadpoles develop in the pool of the ground. The point is that traditional ter they hatch, the female returns and deposits infertile water collected at the base of the cavity. tadpole and egg deposition sites closer food eggs for the developing tadpoles. The tadpoles Breeding in water sources within a forest’s to the ground are no longer incorporated themselves are modified for phytotelmic life. Early stage canopy can enable a species to live its into the reproductive biology of this tadpoles have keritonized mouth parts for feeding on entire life high above the forest floor. This species. things like mosquito larvae and other aquatic invertebrates. With time, this keritonized mouth part disap- pears, leaving a mouth that readily accommodates egg feeding.

Bolitoglossine salamanders have eggs that directly develop, skipping an aquatic, free living larval stage. Many inhabit various canopy levels. Much of the biology and ecology of these salamanders remains a mystery. We do know that they are exceptional climbers—they are often observed climbing vertical surfaces. CLOCKWISE FROM UPPER LEFT: Bolitoglossne altamazonica, mexicana, and schizodactyla

24 | Leaf Litter :: Winter 2007 Such desiccating forces can be lethal microhabitats, amphibians may escape Morphologically, frogs might shift to amphibians that lose water as the undesirable climatic conditions toward a body form that efficiently freely through their skin as water of the canopy (Forsyth and Miyata brings the limbs into the trunk when evaporates from the surface of a pond 1987; Hutchison and Dupré 1992). the frogs are at rest, reducing surface (Jørgensen 1997). With behavioral Through physiological adaptations, area and slowing water loss. modifications, frogs might spend various species might become But there is more to the story. much of the daylight hours tucked tolerant to considerable dehydration Breeding in the canopy isn’t as simple away in water filled tree holes or wet relative to other amphibians, as has as it sounds. For species that deposit bromeliad axils. By utilizing certain been found with some African frogs. eggs and tadpoles in water sources,

Canopy Amphibians | 25 water filled tree holes and bromeliad Works Cited International Canopy Network. 1999. axils typically don’t hold the same ICAN forest canopy research planning Denzer, W. 1994. Tree hole breeding in volume of water as terrestrial pools. workshop: final report. And online at: the toad Peltophryne brevipes (Peters, 1867). Clutch size modifications might be a http://192.211.16.13/individuals/nadkarnn/ Amphibia-Reptilia 15: 224–227. necessary step in some cases to limit ESFreport.htm on 5/26/00. Dunn, E.R. 1937. The amphibian and the potential number of tadpoles living Jørgensen, C.B. 1997. 200 years of reptilian fauna of bromeliads in Costa Rica and in these restricted volumes. In fact, amphibian water economy: from Robert Panama. Copeia 1937(3): 163–167. looking at Cruziohyla craspedopus Townson to the present. Biological Review 72: and C. calcarifer, species that typically Fenolio, D.B., M.E. Walvoord, J.F. Stout, J.E. 153–237. breed in plant held waters, they have Randrianirina, and F. Andreone. 2007. A new Kitching, R.L., J.M. Bergelson, M.D. far smaller clutch sizes than their tree hole breeding Anodonthyla (Chordata: Lowmann, S. McIntyre, and G. Carruthers. closest living relatives in the genus Anura: Microhylidae: Cophylinae) from low- 1993. The biodiversity of arthropods from Agalychnis. Water volume is not altitude rainforests of the Masoala Peninsula, Australian rainforest canopies: General the only problem, food items can northeastern Madagascar. Proceedings of introduction, methods, sites, and ordinal be difficult to find in phytotelmic the Biological Society of Washington 120(1): results. Ausralian Journal of Ecology 18: circumstances. Many species have 86–98. 181–191. compensated for this by depositing Fincke, O.M. 1992. Interspecific food eggs for the developing tadpoles, Lim, K. K. P. and P.K.L. Ng. 1991. competition for tree holes: consequences whether fertile or infertile, including Nepenthiphilous larvae and breeding habits for mating systems and coexistence in Anotheca spinosa, Osteocephalus of the sticky frog Kalophrynus pleurostigma neotropical damselflies. American Naturalist oophagus, Trachyderma resinifictrix, Tschudi (Amphibia:Microhylidae). Raffles 139: 80–101. and a variety of frogs of the family Bulletin of Zoology 39(1): 209–214. Fitzjarrald, D.R. and K.E. Moore. 1987. Dendrobatidae. Another answer to Lowman, M.D. and N.M. Madkarni. 1985. Physical mechanisms of heat and mass the problem can be seen in frogs of Forest Canopies. Academic Press, San Diego. the family Microhylidae. For example, exchange between forests and atmosphere Maguire, B. Jr. 1971. Phytotelmata: biota some microhylids that breed in the in Forest Canopies, M.D. Lowman and N.M. and community structure determination forest canopy’s phytotelmata have non- Nadkarni (eds). Academic Press, San Diego, pp. in plant-held waters. Annual Review of feeding, rapidly developing tadpoles, 45–72. Systematics and Ecology 2: 439–464. eliminating competition for food and Forsyth, A. and K. Miyata. 1987. Tropical large water volume requirements. All nature: life and death in the rain forests Nadkarni, N.M. and M.D. Lowman. 1987. of these changes and modifications of Central and South America. MacMillan Canopy science: a summary of its role in can vary between canopy species Publishing Co. research and education in Forest canopies, M.D. Lowman and N.M. Nadkarni (eds). but specializations are needed to Grillitsch, B. 1992. Notes on the tadpole Academic Press, San Diego, pp. 609–613. compensate for canopy limitations. of Phrynohyas resinifictrix (Goeldi, 1907). With a little background on canopy Buccopharyngeal and external morphology Wassersug, R.J., K.J. Frogner, and R.F. amphibians, I hope I have intensified of a tree hole dwelling larva (Anura:Hylidae). Inger. 1981. Adaptations for life in tree holes your interest. This guild of permeably Herpetozoa 5(1/2): 51–66. by rhacophorid tadpoles from Thailand. Journal of Herpetology, 15(1):41–52. skinned creatures is among the least Hallé, F. 1990. A raft atop the rain forest. known of the extant amphibians. I National Geographic, 178(4):128-138. Yanoviak, S.P. 1999. Community structure hope the images you see here increase in water-filled tree holes of Panama: effects of Hutchison, V.H. and K. Dupré. 1992. your desire to contribute to the general hole height and size. Selbyana 20: 106–115. knowledge base. Time is running out Thermoregulation, in Environmental and amphibian decline may restrict physiology of the amphibians, M.E. Feder and what we learn about many of these W.W. Burggren (eds). University of Chicago species. Best of luck with your studies Press, Chicago, pp. 206–249. of treetop amphibians!

26 | Leaf Litter :: Winter 2007

Spindly Leg Syndrome, a Review by Ed Kowalski photos © Ed Kowalski

pindly leg actually compete with the tadpole for surprising that there are some common syndrome, the required nutrients. Regardless causes for SLS. or SLS, is a of the cause, the disruption prevents Tadpoles are most commonly syndrome in the proper development of the leg(s), reared in small containers of a pint Swhich an amphibian metamorphoses resulting in the condition known as or two in volume with frequent water from a larva to an adult with spindly leg. changes to prevent fouling of the water. deformation and/or non-development As there are multiple steps in In these containers the tadpoles are of one or both of the front legs. development, there can be multiple usually fed a diet of dried fish food. The This is sometimes accompanied by points where development is disrupted, problem with the diet is that the fish deformation of the hind limbs. If the resulting in the multitude of different food manufacturers will only guarantee front limb(s) develop enough to emerge “preventions” or resolutions of the a minimal analysis as opposed to an from under the skin, then the leg(s) syndrome that have been passed actual analysis for each batch of food. is/are often thin and weak-appearing, around over the years. As many Apparently one of the most common hence the name “spindly leg.” I refer individuals use similar diets, housing, causes of SLS is a deficiency of one of to spindly leg as a syndrome as it has and rearing techniques for both the the B-complex vitamins, which are multiple causes and preventions; adults and/or tadpoles, it should not be water-soluble and may rapidly leach although in general, one prevention seems anecdotally to be effective more frequently than others. To date SLS is primarily a problem with developing frogs and toads with only one recorded case in a newt. Metamorphosis is a multi-stage, highly stressful and energy-intensive period in an amphibian’s life cycle. The chemistry and genetics of limb development are beyond the scope of this article, and the details are not important here (but an excellent description of the development of the limbs can be found in Biology of Amphibians). The important steps, for our purposes, are those in which a deficiency or excess of a nutrient can block one or more of these developmental steps. In addition to nutritional problems, developmental disruption may be parasite related, D. ventrimaculatus metamorph: Note that it shows both of the classic signs of SLS. The left front as some parasites will encyst near the limb has not developed to the point where it can be visually detected while the right front limb limb bud, disrupting development has failed to complete development. Attention should be paid to the fact that SLS can affect of the limb. Other parasites may different limbs by different amounts.

28 | Leaf Litter :: Winter 2007 Nutritional Preventions Environmental Preventions • Diet of the adults • Examples of environmental cures (some are anecdotal

• Replacing or changing the adults’ supplements reports)

• Iodine addition (potentially dangerous as iodine has a very • Exposure to UVB lighting narrow safe range) • Addition of tannins via tadpole tea or a leaf (modifying

• B vitamin complex (folic acid) the pH and/or hardness)

• Vitamin E • No longer using tadpole tea or leaves

• Different fish food brands ( such as Aquarian®, Sera-Min®, • Use of reverse-osmosis (RO) water (instead of aged or Tetra-Min®) carbon filtered water)

• Live foods such as blackworms or daphnia • Use of aged tap water (instead of RO or carbon filtered water) • Algae scraped from the sides of tanks • Use of carbon filtered tap water (instead of RO or aged • Foods marketed for amphibians (like Frogbites®) tap water) • Multivitamins added to the water • Lowering the temperature at which the eggs/tadpoles • Providing non-breeding periods for the adults to sequester are reared (species dependent) nutrients • Increasing the temperature at which the eggs/tadpoles are reared

• Increasing the calcium hardness of the water

• Treating the tadpoles for parasite infections

• Rearing singly

• Communal rearing

• Communal rearing with recirculated filtered water

• Changing the water on the tadpoles daily

• Changing the water less frequently

• Never changing the tadpole water, just replace evaporated water

• Changing from polycarbonate containers to containers made from glass or a different plastic

Also the above can be combined in many different ways

out of the flake food into the water in in the 1990s, when the “prevention” of transform successfully despite being the cup (which may be why, under very choice was the addition of iodine to the fed a diet otherwise deficient in the similar conditions, a keeper who does water. B-complex vitamins. There is some not change the water as frequently may Some keepers keep the tadpoles concern with this method because there not have as high an incidence of SLS under either lengthy illumination is a genus of algae known as Prototheca compared to one who changes water (12:12 or longer day/night photoperiod) that has been implicated in causing frequently). Currently, one of the most allowing an algae growth to form in poor development of tadpoles. There is commonly used preventions for SLS is the container upon which the tadpole still some controversy about the algae’s the addition of B vitamins to the either feeds in addition to the flake foods that effects because there have been several the tadpole’s food or the water for the are the basis for most tadpole diets. studies that contained contradictory tadpoles, because deficiencies in these The algae growths may supply extra results. It is possible that Prototheca essential vitamins have been linked to B-vitamins, allowing the tadpole to is only a problem under specific SLS. This is different from the situation conditions.

Spindly Leg Syndrome, a Review | 29 Tadpoles reared in simple distilled from 0 to 100% by this change in the (and some of the possible nutrients or reverse-osmosis water (RO water) husbandry. Carbon is often used to may be required in very small trace have also been demonstrated to remove chlorine and other organic amounts). This can then manifest as develop SLS. As distilled/RO water is substances from water systems; SLS in the tadpoles at metamorphosis. very close to pure water and lacks many however, carbon is an indiscriminate If the adult diets were the same or minerals, the lack of these minerals filter and will remove many substances similar, then this could make it appear had been offered as a cause of SLS. from water. Additionally, most filter that SLS is genetically linked; but, in However, many hobbyists successfully carbons will leach phosphate, and as reality it was caused by the deficiencies rear tadpoles in distilled/RO water a result, in the small closed systems in the diet, making the results hard to to which oak leaves, alder leaves or used to rear the tadpoles, the carbon interpret. Often with today’s hobbyists, other acidifying substances have been may remove a necessary trace element the animals are fed in excess of the added. The acidification of the water and/or add just enough phosphate metabolic needs of the frog allowing with tannins should not significantly to disrupt the calcium metabolism for increased fat reserves, allowing change the mineral content of the sufficiently to cause SLS. for more frequent reproductions. This water, so the method by which this Genetics is another potential cause increased reproductive rate may exceed benefits the tadpoles in unknown. As of SLS in frogs. Some lines of frogs the rate at which the frog has time pure speculation, the increased acidity that have SLS-affected siblings also to store the required micronutrients of the water may dissolve some of the produced tadpoles that were affected for a nutritious yolk, causing SLS in minerals bound up as carbonates or with SLS. The problem with this issue developing frogs with no changes other forms in the food, increasing the is that there is strong evidence that in the husbandry routine. If this is bioavailability of the minerals over parental nutrition strongly affects the the case, then simply preventing the time. In my personal experience, the chances that SLS will develop in a frogs from breeding for a period of addition of a carbon prefilter system to clutch of tadpoles. The yolk contains a time can allow the female to replenish the tadpole water supply caused spindly huge reserve of fats, protein, vitamins the needed micronutrients. (One leg in multiple genera (Dendrobates, and minerals required for development should also consider if the vitamin- Smilesca and Mantella). In these of the tadpole. If the nutritional content mineral supplement is over 6 months instances, SLS had not appeared until of the female frog’s diet is deficient in old, because degradation of the fat- the filter was used. The incidence of any of the required nutrients, this may soluble vitamins may be causing trace spindly leg in this specific case went carry over as deficiencies in the yolk deficiencies in the adults). Exposure to improper temperatures, either too cool or too hot, (temperature ranges are species- dependent, of course) can also be a source of developmental abnormalities, including SLS. Because of this, it is important to keep the tadpoles in a temperature stable environment to prevent the risk of temperature- induced abnormalities. There are also other odd preventions of SLS, such as the aforementioned addition of iodine. The use of iodine may be problematic, as toxicity of iodine varies with the molecule used to deliver iodine, but all are toxic regardless of the source. For example, free iodine that leached from plastic containers disinfected This is the same frog as shown on page 28 shown from an angle that better illustrates the with an iodine disinfectant were underdevelopment of the limbs. shown to have resulted in the deaths

30 | Leaf Litter :: Winter 2007 of adult Dendrobates histrionicus Callaway, D.; 1995; Dendrobatid larval Krintler, K. 1988. Observations after a three hour exposure time rearing techniques in captivity survey results; concerning match legs in dendrobatids and (previous 30 minute exposures did Published by Omaha’s Henry Doorly Zoo hylids Beobachtungen zum Problem der Streichholzbeine bei Dendrobatiden und not produce symptoms or mortality). Claessen, Hugo; 1999; Developments in The concentration of iodine from the the study of spindly-leg syndrome; American Hyliden. Herpetofauna. 52(30-32) containers was in the range of 0.179- Dendrobatid Group Newsletter 41: 5-7 Malolepsy, Shawn Eric; 1993, Epidobates 0.358 PPM. tricolor in captivity; American Dendrobatid Davies, Bob; 1993, More on spindly-leg…; Group Newsletter 11: 1-4 The fact that SLS has multiple 1993; American Dendrobatid Group Newsletter preventions demonstrates that the 12: 3-4 Peaker, Malcolm; 1993, More on spindly syndrome likely has many causes. If a Diana, Stephen G.; Beasley, Val B.; Wright, leg disease; American Dendrobatid Group keeper has SLS develop in their rearing Kevin; 2001; Clinical Toxicity, In Amphibian Newsletter 12: 2-3 containers, then that person needs Medicine and Captive Husbandry; 2001, Krieger Pinzari, Alicia; 1994; Incubating to start trying a variety of different Press, Malabar Dendrobates tinctorius eggs-trial and error; husbandry and dietary techniques one American Dendrobatid Group Newsletter 14: 1-2 at a time until the corrective method for Duellman, William E.; Trueb, Linda; 1986, that specific syndrome is discovered. Biology of Amphibians; John Hopkins Press, Powell, Charles L., II; 1993; Conformation The keeper needs to be patient, because Baltimore of adult frogs causing spindly-leg in tadpoles; American Dendrobatid Group Newsletter 12: 4-5 depending upon the temperature and Fedewa, Luke A.; 2006; Fluctuating gram- species, it could take several months negative microflora in developing anurans; Schulte, R. 1980. Remarks concerning to determine if the current husbandry Journal of Herpetology; 40(1): 131-135 frog-diseases, especially in dendrobatids. or dietary change is effective in Green, D. Earle; 2001; Pathology of Bemerkungen zu Froschkrankheiten speziell eliminating SLS. In fact, small changes Amphibia; In Amphibian Medicine and Captive bei Dendrobatiden. Herpetofauna. 2(5):15-17 within or between clutches can create Husbandry; Krieger Press, 2001 Wright, Kevin; 2001; Diets for Captive or prevent SLS in a given generation Amphibians; In Amphibian Medicine and of tadpoles. Unfortunately, even if Halfpenny, Steve; 1993, Spindly leg Captive Husbandry; Krieger Press, Malabar eliminated once, SLS can reappear in syndrome; American Dendrobatid Group a collection for a completely different Newsletter 12: 1-2 Wright, Kevin; 2001; Idiopathic reason. As our understanding of the Hakvoort, J. H. M., E. J. Gouda and Syndromes; In Amphibian Medicine and Captive Husbandry; Krieger Press, Malabar syndrome broadens, hopefully we can P. Zwart. 1995. Skeletal and muscular identify husbandry principles which underdevelopment (SMUD) in young tree Wright, Kevin; Whitaker, Brent R.; 2001; will prevent SLS consistently. frogs (Dendrobatidae spp.). Proc. 5th int. symp. Nutritional Disorders; In Amphibian Medicine Pathol. Rept. Amphib and Captive Husbandry; Krieger Press, Malabar Works Cited Heselhaus, R. 1983. The problem of Bekiares, Nell A.; 1998; Green & black match legs in propagated Dendrobatids Zum poison frog tadpole feeding trials; Reptile and Problem der Streichholzvorderbeine bei Amphibian Magazine; 58: 21-26 Dendrobatiden-nachzuchten. Herpetofauna. 5(26):22-24

Spindly Leg Syndrome, a Review | 31 Carbon Dioxide in the Terrarium by Thomas Falk; Swedish Dendrobates Society This article first appeared in Tilgiftaren’ the newsletter of the Swedish Dendrobates Society. We wish to thank Lars Österdahl for translating it from Swedish.

reparing for froglet on the soil surface in a cramped where it developed normally. a short field position with its hind legs stretched Had the frogs been struck by some trip I wanted and the front part of the body raised on kind of infection, or could it be that to make sure the front legs. A little deeper in the leaf the frogs had become victims of some Pthat my four newly metamorphosed layer a dead froglet was found in about poisoning? I realised that I had not E. tricolor would have constant access the same position, hind legs stretched modified the aquarium in the same to food during my absence. It seemed and head bent upwards. The fourth way as my frog terraria. This one had a good idea to move them to an old frog was nowhere to be found and it no vent openings close to the bottom. 30 litre aquarium that contained a had probably entered the food chain of I began to suspect that the frogs had layer of old oak and beech leaves in the soil fauna. become victims of carbon dioxide varying degrees of decomposition The two still living young were poisoning. that was being used as culture of soil quickly moved to a “hospital tank” with Carbon dioxide a threat invertebrates like springtails and small wet moss. The larger of the two, the woodlice. The resident fauna was one I had found sitting on the glass, Carbon dioxide (CO2) is an supplemented with a fair number of quickly recovered and ate flies the odourless gas that normally makes vitamin dusted small fruit flies. same day. The smaller regained some up 0.04% vol. of the atmosphere. Returning from the trip I found mobility in its hind legs in the next In mammals expired air normally a froglet sitting on the glass wall, and 24 hours, but it had recurrences of contains 4% CO2 as a result of as there seemed to be a lot of fruit jerks of the hind legs, and I never saw metabolism. An increased level of flies left, I did not rush to return the it eat. It died after another two days. CO2 in the air makes it more difficult frogs to their normal terrarium. To The surviving frog was kept under for the body to get rid of this waste my dismay, when I decided to do so observation foursome weeks until it product, and the body reacts with a few days later, I found the second was returned to the original terrarium, increased breathing activity. If the

Terrarium type Drainage of heavy gas

32 | Leaf Litter :: Winter 2007 time argon gas was used instead of carbon dioxide. Argon is an inert, non-poisonous and odourless, non- combustible gas. As the oxygen inside the bottle disappeared, the frogs moved around in a much calmer way than in the CO2 experiment. After a few minutes they opened their mouths in a strange way, as though yawning. No cramps were observed, but they Cramped frog Asphyxiated frog sat shrunken. After another minute the breathing movements ceased carbon dioxide cannot be removed where they sat after feasting on the and they no longer reacted to touch. from the blood a lower pH results, fruitflies that had also been introduced. The experiment was terminated, and oxygen-saturated air was blown into which seriously affects the salt balance, When I saw that everything the chamber. After five minutes all a state called “acidosis.” Breathing looked normal I let CO2 enter the three frogs had recovered. in a CO2 level of about 5% leads to bottle through the inlet. After a few unconsciousness in a human and minutes the frogs began showing signs Interpretation higher levels can be lethal. of restlessness, jumping around in an In order to reduce the stress and The sensitivity to levels of the gas aimless manner. After another minute avoid unnecessary harm to the frogs, varies between organisms. Animals the first signs of cramps began to show, the two experiments were performed living in the soil or among fermenting and two of the small frogs fell on their more rapidly than would be the material have the highest tolerances. backs. Soon all three lay motionless case under natural conditions. The One soil living species of springtail can with their hind legs outstretched and experiments show that an increased survive in up to 35% whereas another, no visible breathing movements under level of CO2 causes unrest and cramps, living in the vegetation does not their chins. To prevent certain death whereas the low oxygen situation is tolerate more than 2%. It results from I ended the CO2 influx and ventilated characterised by a languid attitude the decomposition of leaf litter, from the bottle. No visible improvement without unrest or cramps. Therefore, it fermentation, and from respiration. occurred until after about 5 minutes must be assumed that the death of the oxygen enriched air was blown into the Carbon dioxide is over 60% three original tricolor was not a result bottle. Since no breathing was present, heavier than air. In a vivarium with of a too low oxygen level. insignificant air movement CO2 will any gaseous exchange had to take place Simple calculation shows that in accumulate in the lower parts and entirely through the skin. a closed compartment the CO2 level a gradient results with the higher The first signs of life were seen can reach lethal levels long before the concentration at the bottom. If the after about 15 minutes and after oxygen level gets too low. Therefore terrarium has no opening at a low level another 5 minutes all three frogs were oxygen depletion is no real risk in a a death trap may result. Other possible sitting upright, breathing normally. A transportation box or a terrarium. waste gases in a terrarium, like few hours after the experiment they methane or ammonia are much lighter were all feeding, and with no lasting The experiments have than air, and thus easier to vent away. symptoms. strengthened my assumption that the frogs died from CO2 that had CO2 Experiment Experiment with low Oxygen accumulated near the bottom of the In order to test the CO2 hypothesis Since the production of CO2 is tank, with a number of other factors I got three one-week-old E. tricolor often connected with the consumption playing a part: froglets from a friend. They were of oxygen it is possible that the 1. The lid lay on glass flanges that were placed in a small bottle with an inlet result of this experiment was due to meant to keep springtails etc. in. The hole under the bottom material (LECA; low oxygen rather than high carbon only ventilation was a small gauze peat moss; leaves). A fine meshed dioxide. For this reason a second type net in the cover, which made air screen covered the upper opening. The experiment was performed with the movement difficult. frogs soon found their favourite places same set up and the same frogs. This

33 | Leaf Litter :: Winter 2007 2. The lighting was a 5W compact which means when the frogs are extra for a moderate sized vivarium. If fluorescent lamp placed above sensitive to stress. As long as the the vivarium is fitted with sliding the cover glass. This gave minimal cover is lifted every day for the input doors these should al-low sufficient convection inside the tank. of fruitflies, the build-up of stagnant ventilation unless they are totally . Only a small plant was present in the CO2 rich air at the bottom is made insect proof or high in the vivarium. tank and in the lowlight it had little impossible. When the terrarium Diamond drills are best for drilling chance of absorbing a significant is left unattended for a few days, glass without cracking. It is essential amount of carbon dioxide. perhaps with extra fly bottle inside to cool the drill and the best way is to provide food in the unattended to build a Plasticine cup around the . The leaf litter was “ripe” and the period, the combined effect of the extra area to be drilled and filling this with population of soil invertebrates fermentation and the poor ventilation water. Drilling plastic is best done considerable. can be fatal. Deaths among a group of with a drill of the type that has a sharp . Small pieces of banana were froglets or a quarantine tank tend to point. Putting the drill in the freezer fermenting, thus producing CO2 (and be blamed on infections or parasites, and using it cold prevents the plastic alcohol). but the real culprit may well be an melting and sticking to it. . The tank was left almost a week unsuitable tank. An aquarium can easily be without the usual daily opening of the Avoiding Problems converted as a temporary vivarium by cover. This provided ample time for the standing it on its end and arranging a heavy carbon dioxide to accumulate at with Carbon Dioxide door and ventilation in the opening. the bottom. Thomas Falk A piece of glass is cut to the width of This article is intended as a follow-on from the aquarium and a couple of holes are How common are CO2 problems? the article that appeared in Newsletter No. 41, drilled in it and covered with mesh. March 2001 (Carbon dioxide build-up 1.) The loss of my E. tricolor froglets This is then fixed using silicone. ‘H’ is probably not a unique event in Carbon dioxide (CO2) can profiles are fitted and the upper space the hobby. One hears reports of accumulate in the bottom of a is covered with netting. Plastic should ‘unexplained’ deaths in cases where terrarium. As high levels of it not be used for the door, as this will old aquaria, plastic containers etc have are poisonous to most animals, deform. been used for temporary holding frogs, precautions must be taken to avoid CO2 can be a threat to frogs in for instance in quarantine practice problems for ground dwelling small transport boxes. The CO2 in an or directly after metamorphosis amphibians in either permanent or inadequately ventilated box can reach temporary vivaria. The CO2 must be a dangerously high level long before vented out of the vivarium and this the oxygen levels become too low. is most difficult in a tank that is only Air holes are really carbon dioxide accessible from above. It is of little use vent-holes and should be placed to increase the ventilation area at the accordingly. Most transport cases only top as this will not allow its escape, have holes in the lid that does not thus making it difficult to maintain the allow for the escape of the CO2. required high level of humidity. Holes should be made from the A ventilation hole with netting inside so that any sharp plastic edges near the bottom of the vivarium works do not damage the delicate skin of the in two ways: frogs. Do not over-do the number or 1. An inflow of air will be created by sizes of the holes. Providing the CO2 a chimney effect caused when the can escape, the low metabolic rate of lamps heat the top of the vivarium and the frogs is unlikely to exhaust the convection currents cause the CO2 to oxygen. Too much ventilation can exhaust. reduce the humidity which can harm 2. With the lamps off the CO2 will the animal. exhaust through the vent. A 10mm hole is sufficient Air movement when lid is opened

Canopy Amphibians | 34