Nicaragua Field Guide

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Table of Contents

Introduction Flora Red Mangrove (Rhizophora mangle)

Cecropia Tree (Cecropia peltata) Laurel (Cordia alliodora) Carambola/Starfruit Flower (Averrhoa carambola) Teak Tree Leaf (Tectona grandis) Big Leaf Mahogany Fruit (Swietenia macrophylla) Neem (Azadirachta indica) Giant Milkweed (Calotropis procera) Piñuela (Bromelia pinguin) Turk’s Cap Mallow (Malvaviscus arboreus) Maidenhair Fern (Adiantum)

3 5 7 9 11 13 15 17 19 21 23

Fauna

Invertebrates Insects

Antlion (Myrmeleon) Azteca Ant (Azteca) Golden Carpenter Ant (Camponotus sericeiventris) Leafcutter Ant (Atta cephalotes) Bush Katydid (cf. Orophus conspersus) Arboreal Termite (Nasutitermes corniger) Montezuma’s Swallowtail (Parides montezuma)

Spiders

Golden Orb-Spider (Nephila clavipes) Wolf Spider (Lycosidae)

Vertebrates Amphibians & Reptiles

Olive Ridley Sea Turtle (Lepidochelys oliveacea) Anolis lizard (Anolis) Cane Toad (Rhinella marina) Red-Eyed Tree Frog (Agalychnis callidryas)

References

24 26 28 30 32 34 36

38 40

43 45 47 49

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Introduction

Nature diversifies deeply in Nicaragua. It was there I spent 10 days researching and documenting the wildlife and biota of the country. It was short in duration but high in biodiversity. We ventured through tangled mangroves with lime green canopies. We tented in the moist-forests where we night-walked up streams to where moths were mating and red-eye tree frogs were getting some shuteye. Insects abound, we were bitten as warned by the locals. They were a friendly bunch, from the teak-farmers where we stayed to the local families of the beach town, San Juan del Sur. I arrived Managua, from Mumbai after visiting my family on New Years. A hemisphere of my brain still asleep, I found my red 45L backpack, visa-ed through Customs, and met Professor Olson, an ecologist by training and a professor Brandeis’ Heller School of Social Policy and Management, outside the airport. We were to head to our base, San Juan del Sur, a surfer’s haven and paradise for nature aficionados. Our first overnight in the capital was at Casa Linda. Farmhouse yellow eggs, plantain, rice, and homemade cheese made for a hearty breakfast. We left for the sandy town of San Juan in a white minivan. The four of us: the Professor, Ben, Simone and I spoke over the passing trucks overflowing with sugar cane. We stopped to order snake-guards to protect our calves should snakes cross our paths. Upon arrival, we switched to a rented rusted blue stick-shift truck. The professor drove us to the dry creek-bed for the annual Christmas bird watch, pointing out curiosities along the way. We scribbled notes of sightings of bird species as we swerved through 20’ high earthen orange riverbanks dotted with holes of the national bird’s nest, the motmot. Intermittent rain nudged us to pause under canopies to inspect insects and appreciate lowland tropical tree roots. We drove to Ostional, a small beach town nearby with gigantic red mangroves. Professor Olson had been working on an informative nature walk through the mangrove with the locals. We tallied the bird count with the other ornithologist, who’d been up since dawn bird-watching. We collected the list of birds to submit for the international Christmas bird count. We slept in cabins near the mangroves situated in water, usually brackish, where the sea-tide pools and empties. They form a rich tropical ecosystem from their canopies to their plants’ roots. The trees that occupy the salt-water maintain their low salinity from activities ranging from passive filtration to active excretion of salt. For reasons unknown, their habitat stops at the subtropical regions. However, in laboratory experiments they grow just fine in fresh water. Yet, they are heavily threatened by coastal commercial and civilian beach use. The next morning, we continued to explore the mangroves and enjoyed leaping on the red mangroves’ trampoline-like limbs. We trampled carefully on cracked

grey-brown earth and kept our distance from a hive of Africanized bees, ready to sting. The hike continued to a stream where we witnessed the wide wingspan of a boat billed heron. Invasive Neem (pg. 15), which competes with the local flora, guided our walk; its seeds are spread by bats. We were back in Ostional and when we reached the bridge, the parakeets were swarming the treetops in couples as the warm tropic sun filtered through the landscape. The next morning, we left early to San Juan. The waves were mild for that time of year despite the winds. We encountered at least three blown-off roofs and toppled energy turbines. The trade winds were being squeezed across the tropics in gusts by the cold front on the southern US. The breezes delayed our journey to the jungle by a day or so. When the weather improved we shopped at the market and filled our truck. We thanked our homestay families, and began our return, driving over cracked roads broken by brooks. We tented on the Finca Guadalupe Farm located just over a dozen kilometers north of Costa Rica, a tropical paradise of immense greens and browns of a lush ecosystem. The tarp was roped and tied high between four trees. Professor Olson set up the portable sun-heated shower. I warned of an ant crossing. But my professor said, “Oh they’re just army ants, they’ll be gone once they remove their prey.” As any good entomologist, he was right; they disappeared soon after. Besides our functional, uninfested shower, we had an open air shelter, ‘furnished’ with air-mattresses, a few water sample exams and specimen collectors. The furnace and some town-bought groceries made good eats, more or less sealing leftovers from hungry arthropods. We conducted insightful late jungle walkabouts sated on full stomachs. The howler monkeys reverberated loudly through the jungle creating the feeling of being in an endless leafy green chamber. We walked through the teak plantation whose trees held giant leaves on straight, tall timber. While hiking, we spotted insects feeding on fungi grown on ant-collected herbage. We identified over 30 species of birds. It is all interconnected: systems upon systems of nature and biology, living and dead environments adapting to their inhabitants. The Nicaraguan sojourn deepened my understanding of tropical ecology and natural resources as a source of conservation of diversity and the significance of nature’s impact on life and our profound effects on the environment.

(Photo; left to right: Simone Parker ‘12, Ben Rifkin ‘12, Prof. Eric Olson, & Isaac Steinberg ‘15 - me)




Flora Jan. 10, 2015 Ostional

Red Mangrove

Rhizophora mangle Habitat Tropical Protected Seacoasts (associated with tides and brackish water)

• The red mangrove has a series of aerial roots (known as stilt roots) it uses for both stability and aeration, when submerged during high-tide. The roots can drop over 5m, giving it its other monikers, ‘the walking tree’ and the ‘jungle-gym mangrove’. • The seedlings are viviparous, meaning they develop into mini plants as seeds on the parent. Then, they float for up to a year, until the are implanted in the ground. • Mangroves provide a rich ecosystem for many aquatic and arboreal animals. At the tops are mangrove crabs (Aratus pisonii and Goniopsis pulchra) that feed off its leaves. In the aquatic pool there are coral, sponges, and numerous arthropods. Sometimes schools of fish can been seen.

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Flora Jan. 2, 2015 Finca Guadalupe

Cecropia Tree Cecropia peltata Habitat Neotropical Lowlands

• Cecropia, also known as Trumpet Trees, are Myrmecophytic trees, which mean they form a mutualistic relationship with many species of ants. These co-evolutionary relationships are neatly intertwined. Food produced by the tree (nutrient rich bodies near the petiole) is exchanged for the ants’ protection from other herbivorous insects. The tree thrives on the nutrients from the ants. Azteca ants (pg. 26) have the most specialized relationship with Cecropia trees. • To allure ants, who also eat other insects, the tree produces Müllerian and Pearl bodies. The ants retrieve and carry them back to their domiciles in stalk-like hollow nodes. These nodes are known as domatia. • Ant frass (feces), dead ants, and devoured insect carcasses leech into the host Cecropia and provide about 93% of its nitrogen.

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Flora Jan. 8, 2015 Finca Guadalupe

Laurel

Cordia alliodora Habitat Tropical America

• The Laurel, also known as salmwood or an Ecuador Laurel, is a Myrmecophytic tree and has a strong mutualism with several species of ants. Ants will form domatia, housing for the colony, in branch nodes. Scale insects will cohabitate and suckle the phloem excreting “honeydew” for the ants to eat. • The strength of the mutualistic bond appears to be tied to supply to resources. In lower-quality environments, the ants will increase the mutual net benefit at the cost of colony size.

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Flora Jan. 4, 2015 Ostional

Carambola/Starfruit Flower Averrhoa carambola Habitat Tropics and Subtropics

• The fruits are yellow and oblong with ribs (usually 4-6) and when cut horizontally form a pointed-star like shape. The fruit can be eaten raw and has a subtle sweetness. • The plant, thought to have originated in the Indo-Pacific islands, has become an important commercial fruit. Brought to the Americas by the 1800s, it was spread through the world slowly. This may have been due to the seed’s viability (limited to a few days) or the acidity of wild types. Grafting and cultivars have led to its widespread consumption.

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Flora Jan. 6, 2015 Finca Guadalupe

Teak Tree Leaf Tectona grandis Habitat Tropical Forests

• Coveted for its strong and alluring hardwood timber, teak tree plantations have spread in Central America and have contributed to deforestation and soil-erosion. The sapwood and the heartwood (the inner-most) are termite resistant. • The understory plants also suffer from lack of light because of the deciduous trees’ large leaves.

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Flora Jan. 2, 2015 Riverbed on the way to Ostional

Big Leaf Mahogany Fruit Swietenia macrophylla Habitat Lowland Tropical and Subtropical Forests

• The fruit of the Mahogany tree usually has around forty wind-dispersed seeds. They are woody, flat, and winglike. They possess an astringency that makes them offensive to some predatory rodents. • The trees grow considerably tall and wide, up to 45m in height and 2m in diameter. The timber is of great commercial use; its harvest has led to loss of habitat and has made it a vulnerable species.

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Flora Jan. 3, 2015 Ostional

Neem

Azadirachta indica Habitat Tropical and Subtropical Forests

• Neem is thought to have many ayurvedic and religious qualities in by many countries in South Asia. It has been shown in studies to have many positive health effects. • Neem is considered to be an invasive tree to Central America as it competes with local trees and seedlings. Its seeds are also dispersed by bats and birds. It grows fast and competes for resources. It is native to India and was only introduced to the Americas in the early 2000s.

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Flora Jan. 3, 2015 Ostional

Giant Milkweed Calotropis procera

Habitat Tropical and Subtropical Arid and Semi-arid Regions

• Its latex has long been used by cultures all over the world for its medicinal benefits. It has been shown to have anti-inflammatory and analgesic effects among others. • Milkweed is a host-plant for monarch butterflies and they are sometimes planted to attract them. Certain milkweeds produce anti-parasitic chemicals. Adult monarchs prefer to lay their eggs on those milkweeds, but exhibit no preference when eating milkweeds.

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Flora Jan. 3, 2015 Ostional

PiĂąuela

Bromelia pinguin Habitat Dry Soils from Central to Northern South America

• They are terrestrial bromeliads, related to pineapples. They are similar but much more tart in taste yet somewhat sweet. This tartness, due to the enzyme, papain, can tingle in your mouth and lead to bleeding if too much is eaten. Papain is used as a meat tenderizer and can be found in pineapple and papaya, too. • Bromeliads are monocots and grow from the center out spiraling. The leaves create a channeling for nutrients and water to flow to the roots. This is essential for the high turnover of nutrients in rain forests.

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Flora Jan. 8, 2015 Finca Guadalupe

Turk’s Cap Mallow Malvaviscus arboreus Habitat Tropical

• These floral relatives to the hibiscus never fully open their petals. The pollinator, usually a specialized hummingbird or bee, can best pollinate and navigate the flower’s tight morphology to nectar. • The blossom displays approach herkogamy. This is where the stigma is at the top of the central style and the pollen-holding anthers are lower. This ensures that the pollen is transferred to a specified part of its body. The pollinator is guided toward the nectar on the way.

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Flora Jan. 8, 2015 Finca Guadalupe

Maidenhair Fern Adiantum sp. Habitat Tropical Dry-Forests

• This genus of fern’s waxy cuticle helps it remain disease resistant. The wax produces a hydrophobic environment on the leaf, which is unfavorable to many phytopathogens. • The vase-life of a maidenshair fern is about 3 days. When cut it wilts quickly perhaps as another antibacterial measure. The wilting, due to the depletion of water in the leaves, is associated with vascular occlusions. The vaselife can be considerably extended with silver nitrate, up to 45 days, but without impact on the fern’s longevity.

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Insects Jan. 2, 2015 Riverbed on the way to Ostional

Antlion

Myrmeleon sp. Habitat Tropical Dry, Loose, Fine Soils

• Also known as a doodlebug because of the scribble-like lines left in the sand as the larva seeks a suitable home. 24

• The larvae construct a conical pit in finely grained soils or sand and wait until an unexpecting small arthropod encounters the pit. The antlion will flick sand at the insect, until off-kilter, it slides to the bottom of the pit where the Antlion eats its fallen prey.



Insects Jan. 8, 2015 Finca Guadalupe

Azteca Ant Azteca sp.

Habitat New World Tropics

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• These aggressive and arboreal nesting ants form colonies in the nodes of Cecropia tree branches. They are routinely seen in Cordia alliodora (pg. 7) and in the Cecropia species (pg. 5). These ants can form a mutualistic bond with their host tree. Being an aggressive ant species, they will destroy competing ant nests in the area, providing the host tree with protection from herbivory. In return,these ants enjoy a monopoly over food and shelter. The ants’ detritus that falls to the floor of the colony node when decomposed, may also provide some nutrients to the tree when decomposed. • They use the host tree’s plant matter to feed a species of mealybugs (Pseudococcidea) or scale insects (Coccidea), that share their tree-stem nodes (domatia). These Hemiptera in-turn excrete a “honeydew,” which is rich in sugars, vitamins, and amino acids, and provides food to the colony.



Insects Jan. 6, 2015 Finca Guadalupe

Golden Carpenter Ant Camponotus sericeiventris Habitat Tropical & Subtropical

• The aboreal ants establish colonies in large tree limbs and dead tree trunks. A mature colony will have hundreds or even thousands of ants. 28

• They are foraging generalist ants. They will eat everything from seeds to live prey after they are carried back to the tree nest. Few species in the genus can, in fact, form nests in the ground as well as trees.



Insects Jan. 7, 2015 Finca Guadalupe

Leafcutter Ant Atta cephalotes Habitat Tropics & Subtropics

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• These ants tend to their underground fungus garden (basidiomycete) and eat the hyphae. When starting a new colony, the queen will bring with her the fungus in her buccal pouch. The worker ants bring leaves and plant matter to their colonies on which the fungus grows. Each colony can contain up to 5 million worker ants. • Aside from the mushroom diet, the ants also chew the leaves and get nutrients from the plant juices. • The soldier ants are far larger than the workers and are specialized to use their large mandibles to protect the colony if attacked. Their large jaws have been used by indigenous peoples to suture wounds. Soldiers, as well as other ant castes, are expendable and dependent on the available resources. Given the chemical signal a caste will literally rip another caste apart.



Insects Jan. 6, 2015 Finca Guadalupe

Bush Katydid

cf. Orophus conspersus Habitat Tropical Rain-Forest

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• Its close relative, O. tessellatus, is also camouflaged as a leaf and is similar, yet smaller, in appearance. They are also usually darker and resemble dead leaves rather their green-leaf ‘in-likeness’ cousins. • The mating occurs after a musical rendezvous, where the male katydid rubs its stridulatory file across its ‘wing-scrapper‘ to a sonorous effect. An interested female katydid may reciprocate with a quitter song or remain silent.



Insects Jan. 7, 2015 Finca Guadalupe

Arboreal Termite

Nasutitermes corniger Habitat Central American Wet-Forest and Forest Perimeter

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• There are three castes: Queen/King, Workers, and Soldiers (nasutes). The workers form the nest usually on trees, using chewed wood, “carton,“ and fecal “glue.” Their guts contain protozoan symbionts that assist in the digestion of raw cellulose. • The soldiers possess glands on their heads that secrete sticky chemicals that are noxious to anteaters and may be toxic to arthropod predators.



Insects Jan. 8, 2015 Finca Guadalupe

Montezuma’s Swallowtail Parides montezuma Habitat Central American Lowlands

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• The butterflies photographed are mating. Also known as the Montezuma’s Cattleheart Butterflies, the female will lay only one egg per tree and disperse her eggs through shaded areas. Shaded leaves appear to be more suitable for larvae survival rates.



Spiders Jan. 7, 2015 Finca Guadalupe

Golden Orb-Spider Nephila clavipes

Habitat New World Forest Clearings and Secondary Growth in Lowlands and Mid-elevation

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• Also known as “banana spiders,” they are large orb weaving spiders that have a strong golden-yellow silk. The silk’s strength has been used for gun-sights and fishing lines (from Asian species). • The spider monitors the vibrations on its web, indicating entangled prey. The spider will then bite the prey and inject poison and digestive enzymes. Once immobilized, the prey will be brought back to be consumed in the webhub. Larger prey will be festooned into a silken orb before being carried off.



Spiders Jan. 7, 2015 Finca Guadalupe

Wolf Spider Lycosidae Habitat Widespread

• Pregnant females, as seen in the photograph, attach their egg sacs to their spinnerets. As the spiderlings hatch they climb onto the back of the mother spider. 40

• Their eyeshine is a prominent signature feature, its visible at night when a person’s flashlight reflects off of their tapeta in their eyes. The photic reflectivity of the eye is a function to assist in its nocturnal hunt.




Reptiles Jan. 4, 2015 La Flor Beach Natural Reserve

Olive Ridley Sea Turtle Lepidochelys oliveacea Habitat Circumtropical Seas

• Synchronized mass nesting of several thousand turtles are known as “arribada,” with individual turtles laying 100 eggs into sand cavities. • Populations are dwindling as poachers will use adults for leather trade and shells. There is local demand for its aphrodisiac qualities.

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Reptiles Jan. 7, 2015 Finca Guadalupe

Anolis lizard Anolis sp. Habitat Tropical

• The yellow dewlap under the lizard’s jaw is present on male anoles. Its evolutionary significance has not been fully understood. Weak evidence support its use for mutual specie identification. • Anoles primarily ‘sit’ on low land objects like tree trunks and wait for the unassuming arthropod to walk by. After its movements have been detected, a wild hunt for the unlucky insect will ensue.

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Amphibians Jan. 7, 2015 Finca Guadalupe

Cane Toad

Rhinella marina Habitat Tropical Lowlands

• These toads are highly susceptible to desiccation during dry season. Few risks are posed from other vertebrates because they produce Bufotenin, a highly venomous toxin affecting heart function. Its resemblance in structure to serotonin leads to hallucinogenic effects. • The toads were released in Australia to control Cane Beetles a pest to the sugar crop. The toads were ineffective and became highly invasive.

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Amphibians Jan. 7, 2015 Finca Guadalupe

Red-Eyed Tree Frog Agalychnis callidryas Habitat Wet Lowlands of Central America

• A female frog will lay on average 29-42 eggs per clutch with 3-5 clutches laid in a night. The eggs will hatch in 5 days and plop off of a leaf into the stream as tadpoles. Metamorphosis occurs around 79 days later. The frogs take most of their days ‘easy’ as they camouflage their bright colors by clenching their bodies on green leaves and are then ignored by predators. • The eggs will hatch earlier if part of the clutch is attacked. The eggs appear to use an “escape hatching response” in response to certain acoustic noises associated with frog-egg attacking animals. This is a good defense mechanism but operates at the peril of being eaten as a egg on a leaf. The premature tadpoles have to then contend with the risk of aquatic predation.

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References Archer, K. J., and A. L. J. Cole. “Cuticle, Cell Wall Ultrastructure And Disease Resistance In Maidenhair Fern.” New Phytologist 103.2 (1986): 341-48. Web. “Big-leaf Mahogany.” WWF - Big-leaf Mahogany. World Wide Fund, n.d. Web. 01 May 2015. <http://wwf.panda.org/what_we_do/endangered_species/bigleaf_mahogany/>. Benson, Kari, and Robert B. Suter. “Reflections on the Tapetum Lucidum and Eyeshine in Lycosoid Spiders.” Journal of Arachnology 41.1 (2013): 43-52. Web. Fernandes, EF, MM De Castro, BC Barbosa, and F. Prezoto. “Variation In Nesting Behavior Of The Arboreal Ant Camponotus Sericeiventris (Hymenoptera: Formicidae).” Florida Entomologist 97.3 (2014): 1237-239. Web. Forsyth, Adrian, and Kenneth Miyata. Tropical Nature. New York: Scribner, 1984. Print. Fujino, David W., Michael S. Reid, and Harry C. Kohl. “The Water Relations of Maidenhair Fronds Treated with Silver Nitrate.” Scientia Horticulturae 19.3-4 (1983): 349-55. Web. Hölldobler, Bert, and Edward O. Wilson. “Prologue.” Preface. The Leafcutter Ants: Civilization by Instinct. New York: Norton, 2011. N. pag. Print. Janzen, Daniel H. Costa Rican Natural History. Chicago: U of Chicago, 1983. Print. Kumar, Venugopalan Santhosh, and Visweswaran Navaratnam. “Neem (Azadirachta Indica): Prehistory to Contemporary Medicinal Uses to Humankind.” Asian Pacific Journal of Tropical Biomedicine 3.7 (2013): 505-14. Web. Levi, Herbert Walter, Lorna Rose Levi, and Nicholas Strekalovsky. Spiders and Their Kin. New York: St. Martin’s, 2002. Print.

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Lukmandaru, Ganis, and Koetsu Takahashi. “Variation in the Natural Termite Resistance of Teak (Tectona Grandis Linn. Fil.) Wood as a Function of Tree Age.” Annals of Forest Science 65.7 (2008): 708. Web. Nicholson, Kirsten E., Luke J. Harmon, and Jonathan B. Losos. “Evolution of Anolis Lizard Dewlap Diversity.” Ed. Tom Tregenza. PLoS ONE 2.3 (2007): E274. Web. Rausher, Mark D. “Larval Habitat Suitability and Oviposition Preference in Three Related Butterflies.” Ecology 60.3 (1979): 503. Web. Saúco, Victor Galán, H. D. Tindall, and U. G. Menini. Carambola Cultivation. Rome: Food and Agriculture Organization of the United Nations, 1993. Print. Warkentin, K. M. “Temporal Pattern Cues in Vibrational Risk Assessment by Embryos of the Red-eyed Treefrog, Agalychnis Callidryas.” Journal of Experimental Biology 209.8 (2006): 1376-384. Web. Webb, C. J., and David G. Lloyd. “The Avoidance of Interference between the Presentation of Pollen and Stigmas in Angiosperms II. Herkogamy.” New Zealand Journal of Botany 24.1 (1986): 163-78. Web. Yamamoto, Marcela, and Kleber Del-Claro. “Natural History and Foraging Behavior of the Carpenter Ant Camponotus Sericeiventris Guérin, 1838 (Formicinae, Campotonini) in the Brazilian Tropical Savanna.” Acta Ethologica 11.2 (2008): 55-65. Web. All photographs of species were taken by Isaac Steinberg ©

Special Thanks to: The Team: Professor Eric Olson, Simone Parker, and Ben Rifkin, The Friendly People of Nicaragua (including the Finca Farm and the fine people of San Juan del Sur) & My dear family and friends!




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