Sorry DC and Baltimore, no periodical cicadas for you this year, unless you take a road trip: Brood XIII (13) and Brood XIX (19), Magicicada spp

 

Holes about the size of dimes beneath trees appearing in March and April foretell the emergence of periodical cicadas.

 

This week we interrupt our six-legged adventures in Borneo and return to North America, to learn about all the hubbub surrounding the emergence of periodical cicadas. Last week Bug of the Week was bombarded with fearful and hopeful questions about the appearance of periodical cicadas here in the DMV. Cicadaphiles are eagerly anticipating an emergence, but irate entomophobes have called me out like this, “… but back in 2021 you told us they only come out every 17 years! Now, I have to get out of town again.” People have also heard that there is a “double brood” occurring this year. Does twice as much anguish or joy loom ahead for us in the DMV? Not so much, here is what will happen.

2024 is the year that the ginormous Great Southern Brood, Brood 19, will emerge from extreme Southern Maryland in a broad swath that ranges south to Mississippi, west to Texas, and north to the central U.S. This is arguably the largest brood of periodical cicadas in the U.S., right up there with Brood 10 which we enjoyed in our region in 2021. Brood 19, The Great Southern Brood, is fascinating and is comprised of four species of periodical cicadas which emerge every thirteen years. Also this year, the Northern Illinois Brood, Brood 13, will appear in a handful of midwestern states. Brood 13 is comprised of three species that emerge every 17 years. In a few locations in Illinois, Brood 19 and Brood 13 are expected to emerge simultaneously with seven species in total, producing one heck of a cicadapalooza and a fascinating biological event. The last time this happened with Broods 13 and 19 was 13 X 17 = 221 years ago, way back when the Louisiana Purchase brought a whole bunch of new territory to our growing nation.

Map of current known distributions of periodical cicada broods in the Mid-Atlantic. Source: https://ift.tt/uwLVpD4

What does the co-emergence of two broods in the same location mean? Will there be double the number of cicadas causing joy and terror? Will egg-laying damage in orchards, tree nurseries, and landscapes be dreadful and elevated? Will birds and small insectivorous mammals feast and enjoy greater survival and reproduction and will regular food items like caterpillars get a real break from beaks and toothy jaws? And will cicadas from Broods 13 and 19 mate and hybridize, resulting in who knows what? A fantastic ecological and evolutionary experiment brought to us by periodical cicadas and Mother Nature awaits us, if, indeed, there is significant overlap of the two broods. But let’s circle back to Brood 19 and cicadas in the DMV. Back in 2011, the last time Brood 19 visited our region way down in St. Mary’s County, Bug of the Week went on a quest to commune with periodical cicadas. The following is an excerpt from that episode posted in the spring of 2011 when the emergence of Brood 19 was underway.

Billions of cicada nymphs from Broods 19 and 13 will emerge in more than 15 states in the US this spring.

“As a cicada lover, I was more than a little jealous of our neighbors in southern and central states that were able to witness this amazing event. Fortunately, a couple of weeks ago, I began hearing reports of cicadas emerging in St. Mary’s County here in Maryland. These reports were confirmed last week when intriguing images of periodical cicadas arrived in my email. Upon closer examination of the distribution map of Brood XIX, one remarkable location was an outlier to the other populations of the Great Southern Brood, a tiny speck in St. Mary’s County. One major concern and oft discussed aspect of periodical cicada lore is their declining range. During the history of human occupation of North America, several localized broods of cicadas have disappeared, gone extinct. In his classic treatise on the biology and ecology of periodical cicadas, C. L. Marlatt mentioned the disappearance and shrinking distributions of several cicada broods in the United States. Experts suggest that fragmentation and elimination of cicada habitat due to farming and urbanization may be linked to vanishing cicadas in some locations. So, to witness this brood of magnificent creatures that may someday wink out of existence in Maryland, I loaded up my car with camera gear and set off in search of Maryland’s Brood XIX - the St. Mary’s survivors. After a disappointing search that consumed most of one day and yielded a single sighting, I was richly rewarded two days later with the otherworldly songs of thousands of cicadas in treetops festooned with egg-laying females and courting males near the small hamlet of Dameron in southern St. Mary’s County. 

The life of a cicada is mysterious and precarious. When their development is nearly complete in spring, they construct an escape tunnel to the surface of the earth. Soil temperatures in the middle sixties seem to be a signal that the world above ground is warm enough to support flight and reproduction. Many nymphs emerge at night and make a mad dash for vertical structures such as trees and shrubs, however, lampposts, street signs, and slowly moving people seem to work just as well. After climbing up and away from the soil, they attach to a firm object to begin the process of molting. Their outer skin or exoskeleton splits along a predetermined line on their back and the beautiful adult cicada wiggles free from the shell. The freshly molted adult is almost pure white except for bright red eyes and patches of black behind the head. Before its skin hardens, the cicada must expand its wings or it will be unable to fly and seek a mate. After wings and legs have hardened, cicadas scurry or fly to the treetops. Emergence from the earth and the final molt are perilous times for cicadas. Many cicadas survive interment underground for thirteen years only to perish attempting to molt or while trying to reach the safety of the trees.

During winter, periodical cicadas are still deep underground where they feed on plant roots. In late winter and early spring, their presence will be marked by exit holes beneath trees. Fully developed nymphs will take a peek at the world above ground before emerging when soil temperatures are in the mid-sixties. They will ascend vertical structures like trees and shed their exoskeletons to become adults. To the treetops they climb to chorus and mate during May and June. After mating, females deposit eggs in small branches. Eggs hatch into the next generation of cicadas destined to emerge with their brood-mates in 13 or 17 years.

Male cicadas produce their otherworldly sounds by vibrating a tymbal organ on their abdomen.

Male cicadas have evolved a unique structure called a tymbal. These paired organs are located on the sides of their bodies just beneath the wings. The tymbal is vibrated much like a drumhead to produce sound. Males produce a variety of calls for different purposes. If threatened by a predator such as a bird or a squirrel, a loud squawking noise is made in an attempt to startle the predator and make an escape. The principal function of the tymbal is to produce calls that assist in finding a mate and winning her affection. One type of call attracts both males and females to a common assembly place such as a large tree. When guys and gals get eye to eye, the male will use three distinct and different courtship songs to try and convince the gal that he should be the father of her nymphs. If the lady likes his advances, she will signal her approval by flicking her wings with an audible click.

After mating, the female cicada will move to tender young branches to lay eggs. Using a saber-like structure on her abdomen called an ovipositor, the female gouges groves into the woody tissue and lays 20 to 30 eggs in an egg nest. This process is repeated on one or more plants. Females lay as many as 600 eggs. After incubating for more than a month, eggs hatch and tiny nymphs a few millimeters long dive to the earth beneath the tree. In a matter of minutes, they burrow into the soil, find roots, and insert a small straw-like proboscis into the roots. Cicada nymphs hunker down underground sipping sap and slowly growing larger. Despite what you might have heard, cicadas are not blind. Their red eyes see fine. They do not bite, although, if very thirsty, one may probe you a bit with its tiny beak in search of moisture.

Female cicadas use a sharp, tubelike appendage called an ovipositor to slice small branches into which they deposit a cluster of eggs.

Cicadas have survived in North America for millions of years. Early records of cicadas date back to colonial times. A report from the April 3, 1751 edition of the Maryland Gazette noted that “We are informed from many Places, that the Caterpillars appear already in vast numbers, and in some Places the Locusts have been found in great plenty, just under the surface of the Earth, almost at their full growth: May God avert our impending Calamities.”  Well, we all know that locusts are not really found “just under the surface of the earth,” but in April cicadas ready to emerge would be found in droves. If you do the math, it is most likely that the locusts in this report were really the fully developed nymphs of Brood XIX cicadas ready to make their appearance. How marvelous! So, let’s hope that 2011 is another successful season for St. Mary’s survivors and, that in 260 years after another 20 broods, these wonders of nature will still be with us.” Fingers crossed that a return to St. Mary’s County this spring will find Brood 19 alive and well having survived another 13 years.

Acknowledgements

Bug of the week thanks our friends at the Weather Channel for providing the inspiration for this episode. Paula Shrewsbury helped hunt and wrangle cicadas and provided images. The marvelous references “The periodical cicada” by C. L. Marlatt,  “Reproductive character displacement and speciation in periodical cicadas, with description of a new species, 13-year Magicicada neotredecim” by D. C. Marshall and J. R. Cooley, and “Advances in the Evolution and Ecology of 13- and 17-Year Periodical Cicadas” by  Chris Simon, John R. Cooley, Richard Karban, and Teiji Sota were used to prepare this episode. To learn more about these really cool insects, please visit the following websites:

https://www.cicadamania.com/

https://cicadas.uconn.edu/

Rainforest raiders: Fierce Leptogenys ants

 

On the rainforest floor, a column of Leptogenys workers like this one transport food to the colony and help relocate the colony from one hunting zone to another.

 

This week we return to the rainforests of Borneo in the spectacular Tabin Wildlife Reserve where we met millipedes, stingless bees, and lanternflies in previous episodes. Here on the rainforest floor several hundred species of ants can be found, but one of the most ferocious members of the ant clan are the predatory ants in the genus Leptogenys. Although Leptogenys share many characteristics with their army ant cousins which we met in the rainforests of Costa Rica, they are members of a different subfamily of ants known as the Ponerinae. They contrast to the true army ants that are members of the subfamily Dorylinae. Despite their taxonomic distinction, they share many attributes in common with true army ants, not the least of which is their ability to conduct massive swarm raids to plunder an impressive variety of prey as food for their colony. Their smorgasbord includes a wide variety of prey - small invertebrates like spiders, centipedes, millipedes, earth worms, flatworms, and many insects, and some vertebrates including frogs and snakes.

Life and death in Tabin Wildlife Reserve includes fierce Leptogenys ants that plunder several kinds of prey on the rainforest floor. A slow-motion video clip of the frenetic column reveals several workers with unidentified insects clamped tightly in their jaws. Are these other ants, termites, or some tasty insect morsal? The lens fails to reveal. Moments later along the same column, the band of sisters transport a tattered moth back to the nest to fill the hungry mouths of waiting ant larvae and the queen mother.

Cooperation is the name of the game when sister Leptogenys capture prey like this moth and move it back to the nest.

Leptogenys colonies can be massive with more than 50,000 workers attending the colony and its queen. Swarm raids to capture food for the colony may consist of tens of thousands of workers. The raid begins at dusk when workers leave the nest to forage during the night. The raiding column begins as a single line of workers which then bifurcates repeatedly to form a massive fan-shaped front of terror that may span as much as 300 square meters of rainforest floor. When a worker encounters potential prey, a massive attack ensues as scores of workers rapidly respond to subdue the prey. The bites and wicked stings of hundreds of nestmates immobilize the victim. Using sharp, powerful jaws, workers dismember the prey and transport the pieces of meat back to the nest to feed the queen and developing brood. Less savory body parts such as wings and legs may be discarded.

As prey are expunged from one part of the rainforest, the colony is regularly relocated and remains in the same location for only a few days. The emigration from an old site to a new one is an orderly process where helpless pupae and larvae are the first to be carried to the new nest site by workers. Young workers bring up the rear. The queen is also part of the entourage, while a picket line of wary workers guards the route as the colony moves to its new location. Famed entomologist and sociobiologist E. O. Wilson once remarked “ants have the most complicated social organization on earth next to humans.” We could learn something about cooperation by watching Leptogenys at work.  Stay tuned for more ant stories from Borneo at Bug of the Week.

Acknowledgements

Several great articles including “A South East Asian Ponerine of the Genus Leptogenys (Hym., Form.) with Army Ant Life Habits” by U. Maschwitz, S. Steghaus-Kovac, R. Gaube and H. Hänel, and “Topology of the foraging trails of Leptogenys processionalis – Why are they branched?” by K.N. Ganeshaiah and T. Veena provided insight into Leptogenys, as did “The Ants” by Bert Hölldoppler and Edward O. Wilson.  We thank Oliver at Tabin Wildlife Reserve for braving rainforest trails to help us find insects and the keen eyes of Dan, Kristie, Bob, Pat, and Paula for spotting insects. Special thanks to Chien C. Lee at Wild Borneo Photography for help in identifying the star of this episode.

Encounters with Bornean kelulut - stingless bees, Meliponini

 

Pollen baskets loaded? Check. One last sip of nectar? Check. Passion flower pollinated? Check. Time to return to the hive to feed the sisters. The passion flower says thanks.

 

Cracks in masonry walls often provide a suitable nest site for stingless bees. The horn-shaped entrance made of propolis (wax and other substances) narrows the entry way to the hive, making entry from intruders more difficult.

In recent weeks we visited the magnificent millipedes and elegant sap-sucking lanternflies in the Bornean rainforests. This week let’s meet stingless bees, pollinators helping to sustain biodiversity in the plant world. Here in the US our notion of bees is usually western honeybee-centric, visions of rectangular white boxes packed with racks of honeycomb, busy workers gathering pollen and nectar, and a queen producing legions of brood all guarded by fearless female warriors capable of a delivering a fierce sting. Although they are closely related to western honeybees, tropical stingless bees, which in Borneo and other parts of Malaysia are known as kelulut, are quite a bit different. In the natural world, they typically nest in tree hollows and earthen crevices, but with invasion of their realm by modern man cracks and crevices in wooden, cinder block, and masonry walls have become popular nesting sites. Often these hollows have rather large openings and to limit access to the colony and facilitate defense, voids are narrowed to trumpet-shaped entrances constructed with a sticky substance called propolis, a mixture of wax and other materials. This defensible portico may be help stingless bees repel ants and other creatures that would love to raid the colony and plunder honey, pollen, and baby bees inside.  

Honey produced by western honeybees, Apis mellifera, is an important commercial enterprise in many parts of the world. Honey produced by solitary bees has been described as sour or bitter sweet. It is valued as much for its medicinal properties that sources claim include anti-bacterial, anti-carcinogen and anti-oxidant properties, as it is for its flavor. The domestication of stingless bees is known as meliponiculture. The value of locally produced kelulut honey is so great that in 2019 the Malaysian government launched a National Kelulut Honey Industry Plan to assist beekeepers to supply stingless bee honey to a world market, hopefully at a hefty profit. 

An excited tourist reported swarms of unidentified insects outside his cabin at Tabin Wildlife Reserve. At first glance, these appeared to be tiny flies, but on closer inspection the frenetic horde proved to be a species of small stingless bees that built their colony in the wall void of the cabin. In a more natural setting on an ancient tree in the Sun Bear Conservation Centre in Sepilok, Malaysia, a trumpet shaped cone of propolis marked the entryway to a colony of mellow stingless bees. Nearby, stingless bees provided the vital service of pollination to a passiflora blossom while gathering nectar and pollen for their colony.

Almost home, a stingless bee with pollen baskets fully loaded, is on final approach to an ancient tree where her nestmates guard the horn-shaped entryway the hive.

Although stingless bees lack the ability to sting, they are by no means defenseless. In past encounters with solitary bees in the tropical rainforests of Belize, they proved otherwise. In a previous episode we reported on a large dark Belizean bee that employed a highly effective defense. The nest entrance of this species was guarded by several workers carefully watching the movements of people nearby. When one person ventured a little too close to the colony, bees mounted a surprising attack. The assault consisted of dozens of workers flying into faces and hair of nearby humans. Bees paid special attention to eyes, noses, and ears where, despite the absence of stings, their annoying bites were unpleasant and persistent, forcing the intruders to vacate the premises near the hive. 

Recently, scientists have discovered that stingless bee soldiers are a unique caste, some 30% heavier than their nest-mates. Their job is to guard the colony against marauding enemies, including species of robber bees that commonly raid stingless bee colonies. Their coup de main involves grabbing an invader by antenna or wing and refusing to let go, thereby thwarting an attack. In his book, The Insect Societies, E.O. Wilson describes accounts of stingless bees attacking human intruders. Some species eject an irritating liquid that causes a burning sensation to skin. This trick has earned them the local name of cagafogos or “fire defecators” in Brazil. So potent is this defense that it may dissuade very aggressive attackers like army ants from entering nests. In Bornean rainforests and nearby open areas stingless bees were common visitors to many kinds of flowering plants, gathering nectar and pollen. Although past encounters with stingless bees in Central America were spicy, we were able to observe and enjoy mellow Bornean stingless bees without drama. Lucky us. 

Acknowledgements

References used in this episode include the following: “A morphologically specialized soldier caste improves colony defense in a neotropical eusocial bee” by Christoph Grütera, Cristiano Menezesb, Vera L. Imperatriz-Fonsecab, and Francis L. W. Ratnieksa, and the fascinating "The Insect Societies" by E.O. Wilson. Once again, we thank our rainforest guides, Leo on the Kinabatangan River, Larry at Mulu National Park, and Oliver at Tabin Wildlife Reserve for helping us find insects, and adventurers Dan, Kristie, Bob, Pat, and Paula for spotting insects and providing ideas about creatures featured in these episodes. 

To learn more about stingless bee farming in Borneo, please visit this website: https://www.mysabah.com/wordpress/stingless-bee-kelulut-farming-sabah/

Lanternflies: vandals in vineyards, beauties in Borneo

 

Does the bulb at the tip of Pyrops intricatus’ headgear really produce light?

 

Over the past several years, we learned much about the mischief caused by the spotted lanternfly, Lycorma delicatula, as it spreads, continuing its depredations of vineyards, and creating problems such as fouling plants with honeydew, attracting stinging insects, and generally freaking-out humans with its vast numbers and bumbling behaviors. To see another side of lanternflies, we return to Borneo, a biodiversity hotspot where last week we met a fascinating collection of millipedes and centipedes.  

Lanternflies belong to a group of insects known as the Hemiptera. Unlike caterpillars or beetles that munch leaves with jaws, Hemiptera use soda-straw-like sucking mouthparts to remove sweet phloem sap from plants. Immature feeding stages of Hemiptera are called nymphs, rather than larvae. Within the taxonomic order Hemiptera, lanternflies belong to a family called Fulgoridae. How do fulgorids come by the name lanternfly? One account has it that famed German naturalist and illustrator Maria Sibylla Merian coined the moniker “lanternfly” after encountering these strange insects with bizarre ornamentation on their heads while on an expedition to Surinam in the early 1700s. She mistakenly believed these adornments could produce light at night, perhaps for the purpose of mating. If you look at the images herein, it certainly looks like those horns could light up, right? Alas, they don’t emit light, but the name lanternfly has stuck to this day.

A nighttime walk in the rainforest provides an exciting chance to search for lanternflies. A pair of ghost-like Pyrops sultanus rest on the trunk of a tree. In the morning light their magnificent head decoration is revealed. One look at the bulb on the tip of Pyrops intricatus’ headgear explains why they were believed to be luminescent. A bright blue horn decorates the head of Pyrops whiteheadi and for obvious reasons some call Zanna nobilis the crocodile lanternfly. A pretty Penthicodes lanternfly rests on reed near the riverbank. Nothing goes to waste in the rainforest. Once their time in the rainforest is through, ants quickly recycle the remains of lanternflies.

As we learned with spotted lanternfly, these sap-feeders excrete vast quantities of the carbohydrate rich waste product known as honeydew. We also learned that several stinging insects, hornets, yellow jackets, paper wasps, and even honeybees are attracted to trees infested with lanternflies to gather honeydew, a food for themselves and their brood. Do Bornean lanternflies produce honeydew attractive to other creatures? You bet they do. Some of the species depicted in this episode in the genus Pyrops participate in an ecological association known as trophobiosis, a deal between different organisms where one produces food for another. The species providing food is called a trophobiont. Observations by naturalists and researchers revealed that lanternflies are attended by geckos and cockroaches. Assuming a post near the derriere of lanternflies, they collect sweet honeydew as it Is produced. Clever beasts. Scientists today still remark that little is known about the biology and ecology of lanternflies. Hopefully, enough unique rainforest habitats can be saved to prevent the loss of these gorgeous creatures and their compatriots in Bornean rainforests and other biodiversity hotspots around the world.

Acknowledgements

We thank our dauntless guides, Leo on the Kinabatangan River, Larry at Mulu National Park and Oliver at Tabin Wildlife Reserve for braving the rivers and rainforest trails to help us find insects. The keen eyes of Dan, Kristie, Bob, Pat, and Paula helped spot lanternflies featured in this episode. The interesting articles “Review of the effusus group of the Lanternfly genus Pyrops Spinola, 1839, with one new species and notes on trophobiosis (Hemiptera: Fulgoromorpha: Fulgoridae) by Jérôme Constant, and “The Beauty of Life - Lantern Flies” by Aub Podlich contributed greatly to this article, as did images provided by Chien C. Lee at Wild Borneo Photography.

Recyclers of the rainforest: Bornean myriapods, millipedes (Diplopoda), and their nemesis, giant centipedes (Chilipoda)

 

Rainforest romance ensues when randy red millipedes (Trachelomegalus modestior) of Borneo get face-to-face.

 

Clusters of strange “noodle” millipedes (Pseudodesmus sp.) frequent decaying logs in the Bornean rainforest.

With terrestrial insect life in the wild mostly battened down for winter in much of North America, it’s time to get out of town to warmer places. We head to the tropics to meet some of the most fascinating arthropods on the planet. First stop, Borneo, the third largest island on earth where we visit Gunung Mulu National Park and Tabin Wildlife Reserve. At some 130 million years of age, the Bornean rainforest is one of the oldest and most diverse places on the planet. Populated by some 10,000 species of plants that provide food and homes for an estimated 3,000 species of arthropods, Mulu and Tabin are ideal locations to discover insects and their allies.

Guess why these rainforest recyclers in the genus Stenoniodes are called tractor millipedes.

Year-round warmth and stupendous rainfall make tropical rainforests one of the most productive ecosystems on earth and this phenomenal bounty of plant material provides untold opportunities for many of Mother Nature’s most important recyclers. Premier amongst the arthropod recyclers are millipedes, a coterie some 400 million years old. These detritivores shred and consume organic matter including mosses, algae, fallen trees, and decaying vegetation that carpet the forest floor.

Millipedes belong to the subphylum of the arthropods called Myriapoda, those with “many feet.” Do they really have a thousand feet? Nah, they don’t really have feet, but they do have legs and the record number of legs for a millipede is somewhere north of 700. However, most millipedes have fewer than 400 legs. As millipedes molt and grow, body segments with two pairs of legs are added.

When under attack by a predator, giant pill millipedes (Sphaerotheriida) go armadillo and curl into a defensive ball to foil attacks by would-be predators.

Millipedes live many years and can produce hundreds of offspring during their lifetime. Millipedes do not bite or sting, but several species such as those we met in a previous episode secrete noxious chemicals from glands lining the margins of their body. Noxious chemicals are not the sole defense of these rainforest recyclers. Some like the giant pill millipedes can curl into a ball, tucking their head and vulnerable underbelly beneath a sphere of hard armored plates impenetrable to many rainforest predators.

But danger prowls the rainforest floor in another member of the myriapod clan, Pacific giant centipedes. These fierce predators bear but one pair of legs on each body segment. Two highly modified legs just beneath the head called toxicognaths are poison claws which deliver a highly toxic, lethal bite to hapless prey, including many arthropods and some small vertebrates. In addition to millipedes, spiders, scorpions, mice and lizards may be on menu.

Highly modified legs called toxicognaths deliver lethal venom to victims of predacious centipedes.

Centipedes locate prey by sensing vibrations in the surrounding substrate. Once detected, a vicious assault ensues. First, a potential meal is snared by legs near the head of the centipede and then embraced with several sets of legs while the toxicognaths inject venom into the prey. While the venom takes effect, the perfidious embrace prevents the prey’s escape. Once subdued, the prey is chopped into small pieces by two pairs of jaws and the tasty meal enters the centipede’s maw. Pacific giant centipedes now occupy several continents and dozens of islands around the world.

Should you have the good fortune to encounter one of these carnivorous creatures, use caution. The poison-claw-pinch of the centipede is extremely painful and is potentially lethal, so if you encounter one, please avoid the urge to cuddle it. Next week, we return to the rainforest to meet more six-legged wonders.

A walk in the Bornean rainforest reveals strange serenades and a rich assortment of millipedes, key recyclers of organic matter. Along a shady boardwalk a pair of red millipedes (Trachelomegalus) get romantic and glam for the camera. Nearby, “noodle” millipedes (Pseudodesmus) cluster on a decaying log. Romance is also afoot on the forest floor where tractor millipedes (Stenoniodes) get it on. When harassed by a predator or bug geek, giant pill millipedes (Sphaerotheriida) go armadillo and curl into a defensive ball. But danger awaits these recyclers and other small animals in the form of giant predacious centipedes (Scolopendra). The underslung venomous claw of centipedes delivers a lethal puncture to prey or fiercely painful and potentially life-threatening wound to humans.   

Acknowledgements

Several offerings by iNaturalist and Wikipedia were used to prepare this episode as were “An Introduction to Insects” by D. J. Borer, D. M. DeLong, and C. A. Triplehorn, and “Secret Weapons” by T. Eisner, M. Eisner, and M. Siegler.  We thank our intrepid guides, Larry at Mulu National Park and Oliver at Tabin Wildlife Reserve for braving the rainforest after dark and Dan, Kristie, Bob, Pat, and Paula for spotting, wrangling, and documenting creatures featured in this episode.