{"id":352,"date":"2013-09-24T08:26:00","date_gmt":"2013-09-24T12:26:00","guid":{"rendered":"https:\/\/sites.bu.edu\/warkentinlab\/?page_id=352"},"modified":"2019-09-18T14:10:36","modified_gmt":"2019-09-18T18:10:36","slug":"red-eyed-treefrogs","status":"publish","type":"page","link":"https:\/\/sites.bu.edu\/warkentinlab\/research\/red-eyed-treefrogs\/","title":{"rendered":"Red-eyed treefrogs"},"content":{"rendered":"<h2>Life cycle \u2013 natural history<\/h2>\n<a href=\"\/warkentinlab\/files\/2013\/09\/RETFlifecycle-550.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/09\/RETFlifecycle-550.jpg\" alt=\"Red-eyed treefrog life cycle\" title=\"Red-eyed treefrog life cycle\" class=\"size-full wp-image-355\" width=\"550\" height=\"401\" \/><\/a>\n<p>Red-eyed treefrogs live in low to mid-elevation rainforests from the Yucatan to Colombia. In the rainy season, males come to ponds and establish territories in plants near the water. They defend their territories and compete with other males by calling, shaking the plants, and fighting. When it rains, females carrying eggs come to the ponds to choose a mate. The pairs lay eggs on plants hanging over the water and tadpoles fall into the water upon hatching. Eggs in the trees can be eaten by wasps, snakes, or katydids, or killed by pathogenic fungus. They can also drown, if they fall into the water, or die from dehydration if the weather is too dry. Embryos can hatch early to escape from attacks by egg predators and pathogens, or in response to abiotic threats, but they typically hatch later if undisturbed. Thus tadpoles enter the water at different ages, sizes and stages of development. Tadpoles are eaten by many aquatic predators, including dragonfly nymphs, giant water bugs, freshwater shrimp, and fishes. Tadpoles that were induced to hatch early are more likely to be killed by aquatic predators and less likely to survive to metamorphosis. After a month or more in the water, the tadpoles metamorphose into froglets. Froglet size varies a lot, depending on conditions in the pond. So does their shape when they leave the water; metamorphs emerge with longer tails in response to water bugs. On land, froglets face a new set of predators, including semi-terrestrial spiders. Metamorphs on land remain relatively inactive near the pond while they absorb their tails, then climb up into the trees and disappear. We know very little about their lives as juveniles.<\/p>\n<p><a href=\"\/warkentinlab\/files\/2013\/09\/CicloDeVida-LifeCycle.pdf\">Ciclo de vida de la rana arbor\u00edcola de ojos rojos\/Life cycle of the red-eyed treefrog<\/a> [pdf with image and text in English and Spanish \u2013 for printable poster-size file for educational use contact Warkentin]<\/p>\n<h2>Embryonic development<\/h2>\n<a href=\"\/warkentinlab\/files\/2013\/09\/AcOviposition-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/09\/AcOviposition-200.jpg\" alt=\"Oviposition\/fertilization\" title=\"AcOviposition-200\" class=\"size-full wp-image-529\" width=\"200\" height=\"278\" \/><\/a>\n<p>Red-eyed treefrogs lay eggs at night. Most eggs are laid between about 10 pm and 2 am, although this varies with the weather. The female provides the embryos with a large amount of water, in the egg jelly. She usually breaks the eggs she is carrying into several egg masses or clutches, taking time to absorb more water in between laying each set of eggs. Egg color varies among females, and is affected by their diet, but\u00a0 it is consistent within the set of eggs laid by a female on the same night. This can be used to distinguish egg masses that could be siblings (same color) from those that are clearly not siblings (different colors). The photos below are from several different clutches.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/09\/IMG_1108-2005-EarlyCleavage-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/09\/IMG_1108-2005-EarlyCleavage-200.jpg\" alt=\"Early cleavage (0 d, 7:45 am)\" title=\"EarlyCleavage-200\" class=\"size-full wp-image-532 \" width=\"200\" height=\"201\" srcset=\"https:\/\/sites.bu.edu\/warkentinlab\/files\/2013\/09\/IMG_1108-2005-EarlyCleavage-200.jpg 200w, https:\/\/sites.bu.edu\/warkentinlab\/files\/2013\/09\/IMG_1108-2005-EarlyCleavage-200-150x150.jpg 150w\" sizes=\"(max-width: 200px) 100vw, 200px\" \/><\/a>\n<p>By morning, early cleavage is well underway and individual cells are visible at the top of each egg. The large mass of yolk at the bottom is not yet divided. Individual eggs are surrounded by relatively thick layers of jelly, and embedded at different levels in the overall clutch structure.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/YolkPlug-1d730am-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/YolkPlug-1d730am-200.jpg\" alt=\"Yolk plug stage\" title=\"YolkPlug-1d,730am-200\" class=\"size-full wp-image-546\" width=\"200\" height=\"176\" \/><\/a>\n<p>During the evening and night, embryos progress through later cleavage stages and early gastrulation stages. By morning, gastrulation is almost complete, with just a small yolk plug remaining. This is often hard to see, as it usually points toward the interior of the clutch, but is visible in the bottom right egg. Note the &#8220;two-toned&#8221; look, with the animal pole (up) and vegetal pole (down), which is distinct from the more uniform color of newly laid eggs.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/NeuralFolds1d1230pm-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/NeuralFolds1d1230pm-200.jpg\" alt=\"Neural folds (1 d, 12:30 pm)\" title=\"NeuralFolds(1d,1230pm)-200\" class=\"size-full wp-image-550\" width=\"200\" height=\"189\" \/><\/a>\n<p>Here, neural folds have formed. The embryos now float freely in the perivitelline fluid within their capsule with the heavier yolk down and developing embryo up, however the clutch is positioned.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/NeuralTube1d233pm-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/NeuralTube1d233pm-200.jpg\" alt=\"Neural tube stage (1 d, 2:30 pm)\" title=\"NeuralTube,1d,233pm-200\" class=\"size-full wp-image-549\" width=\"200\" height=\"197\" \/><\/a>\n<p>The neural folds have closed to form the neural tube and the developing head is more apparent, with gill plates forming. At this stage, embryos keep their proto-head toward the air-exposed surface, where there is more oxygen, and put their developing tail toward the back. If displaced, they will use ciliary rotation to return to this position.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/IMG_0606-Tailbud1d915pmTurq200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/IMG_0606-Tailbud1d915pmTurq200.jpg\" alt=\"Tailbud stage (1 d, 9:15 pm)\" title=\"IMG_0606-Tailbud(1d,915pmTurq)200\" class=\"size-full wp-image-552\" width=\"200\" height=\"203\" \/><\/a>\n<p>By evening, embryos have reached the tailbud stage. The perivitelline space inside the egg is larger, and the jelly layer around each egg is thinner.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/IMG_0663-Tailbud1d1005pmYellow200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/IMG_0663-Tailbud1d1005pmYellow200.jpg\" alt=\"Tailbud stage (1 d, 10:05 pm)\" title=\"IMG_0663-Tailbud(1d,1005pmYellow)200\" class=\"size-full wp-image-554\" width=\"200\" height=\"158\" \/><\/a>\n<p>Tailbud stage embryos from yellow eggs \u2013 the color variation extends to embryo tissue, not just yolk reserves.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/MuscularResponse2d145pm-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/MuscularResponse2d145pm-200.jpg\" alt=\"Muscular response (2 d, 1:45 pm)\" title=\"MuscularResponse(2d,145pm)-200\" class=\"size-full wp-image-557\" width=\"200\" height=\"157\" \/><\/a>\n<p>At age 2 days, embryos become more active. Their tails are curled to the side and they will twitch or squirm if disturbed (e.g. sprayed with water). Their hearts begin to beat and their gills are forming but initially their blood is not red. The clutch structure has changed, so individual eggs, with little surrounding jelly are at the surface, attached to an underlying jelly core.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/GillCirculation3eggs-2d630pm-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/GillCirculation3eggs-2d630pm-200.jpg\" alt=\"Gill circulation (2 d, 6:30 pm)\" title=\"GillCirculation(3eggs-2d,630pm)-200\" class=\"size-full wp-image-556\" width=\"200\" height=\"238\" \/><\/a>\n<p>Later in the day, the gills are longer with distinct (but short) branches and pale blood cells can be seen circulating through them. The eyes are becoming distinct but have little pigment.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/greyeyeearlyCT3d820am-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/greyeyeearlyCT3d820am-200.jpg\" alt=\"Grey eyes (cornea clear, 3 d 8:20 am)\" title=\"greyeye(earlyCT)3d,820am-200\" class=\"size-full wp-image-561 \" width=\"200\" height=\"183\" \/><\/a>\n<p>In the morning at age 3 days, the eyes appear grey, with melanophores on the iris in an incomplete circle. The lens of the eye is still cloudy. Gill branches are elongate and blood is pale red. There are scattered melanophores on the tail fin and part-way down the belly, but no pigmentation over the heart. The yolk is bulbous, with prominent superficial blood vessels.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/CorneaTransparent3d7pm-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/CorneaTransparent3d7pm-200.jpg\" alt=\"Dark eyes (3 d, 6:57 pm)\" title=\"CorneaTransparent(3d,7pm)-200\" class=\"size-full wp-image-563\" width=\"200\" height=\"197\" \/><\/a>\n<p>By evening at 3 days old, there is tail fin circulation. More obvious though, is the much darker pigmentation of the eyes and\u00a0 brighter red blood. There are scattered\u00a0 melanophores across the heart. Gill branches are continuing to elongate.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/KW-1165-blackeyes34d827am-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/KW-1165-blackeyes34d827am-200.jpg\" alt=\"Operculum formed (4 d, 8:27 am)\" title=\"KW-1165-blackeyes3(4d,827am)-200\" class=\"size-full wp-image-566\" width=\"200\" height=\"173\" \/><\/a>\n<p>By the morning at age 4 days, embryos have scattered shiny iridophores on the iris and over the heart. Overall, they are more darkly pigmented than at 3 d, and their yolk is less bulbous. Blood vessels are still prominent in the yolk sac, but show more branching.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/GreenEggs4d1010am-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/GreenEggs4d1010am-200.jpg\" alt=\"Hatching-competent (4 d, 10:10 am)\" title=\"GreenEggs4d(1010am)-200\" class=\"size-full wp-image-565\" width=\"200\" height=\"165\" \/><\/a>\n<p>At 4 days, embryos begin responding to predator attack by hatching, although escape rates are quite variable from clutch to clutch. All embryos hatch (eventually) if flooded. Individuals may wait an hour or more, but none drown.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/blackeyes14d824am-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/blackeyes14d824am-200.jpg\" alt=\"Hypoxia slows development (4 d, 8:24 am)\" title=\"blackeyes1(4d,824am)-200\" class=\"size-full wp-image-567 \" width=\"200\" height=\"156\" \/><\/a>\n<p>Embryos are packed closely together in egg clutches and oxygen only enters through the air-exposed surface. Embryos that are at least 15% exposed do fine, but those mostly buried under their siblings (upper left) develop more slowly.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/IMG_0954-gills5d11am-v2cleanB-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/IMG_0954-gills5d11am-v2cleanB-200.jpg\" alt=\"Waiting to hatch (5 d, 11 am)\" title=\"IMG_0954-gills5d,11am-v2cleanB-200\" class=\"size-full wp-image-570\" width=\"200\" height=\"185\" \/><\/a>\n<p>At 5 days old, dense iridophores cover the iris and heart, and melanophores are abundant on the sides and belly. The yolk is more streamlined, but undivided, and blood vessels are inconspicuous. Beaks begin to keratinize. Embryos hatch readily if attacked by wasps or snakes, with ~80% escape rates, but rarely hatch if undisturbed.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/GutCoils6d1am.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/GutCoils6d1am.jpg\" alt=\"Gut coils (6 d, 1 am)\" title=\"GutCoils(6d,1am)\" class=\"size-full wp-image-572\" width=\"200\" height=\"203\" \/><\/a>\n<p>At age 6 d the yolk is divided into gut coils. The gut is packed full of yolk, so animals have substantial energy reserves and they cannot feed immediately upon hatching.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/KeratinizedMouthparts6d1am-200.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/KeratinizedMouthparts6d1am-200.jpg\" alt=\"Keratinized mouthparts (6 d,1 am)\" title=\"KeratinizedMouthparts(6d,1am)-200\" class=\"size-full wp-image-573\" width=\"200\" height=\"183\" \/><\/a>\n<p>Mouthparts are more strongly keratinized and overall pigmentation is darker (melanophores) and shinier (iridophores). At this point embryos are highly sensitive to disturbance and more difficult to handle without inducing hatching although, if undisturbed, many will not hatch until the evening they are 7 (almost 8) days old.<\/p>\n<div style=\"clear: both;\"><\/div>\n<a href=\"\/warkentinlab\/files\/2013\/10\/KW-080619-6187-4d6dHatchlings-2001.jpg\"><img loading=\"lazy\" src=\"\/warkentinlab\/files\/2013\/10\/KW-080619-6187-4d6dHatchlings-2001.jpg\" alt=\"New hatchlings (4 d and 6 d)\" title=\"4 and 6 d old Agalychnis callidryas hatchings, Gamboa, Panama\" class=\"size-full wp-image-575\" width=\"200\" height=\"236\" \/><\/a>\n<p>Tadpoles that hatched near the onset of hatching competence (4 d, left) enter the water smaller and less developed than those hatching later. Most undisturbed embryos hatch at 6 d (right) or 7 d.<br style=\"clear: both;\" \/><a target=\"_blank\" href=\"\/warkentinlab\/files\/2013\/09\/AgalychnisDevelopmentalStages.pdf\" rel=\"noopener\">Photographic guide<\/a> to development of <em>A. callidryas<\/em>, with stages according to <a target=\"_blank\" href=\"http:\/\/www.jstor.org\/discover\/10.2307\/3890061\" rel=\"noopener\">Gosner 1960<\/a> and Pyburn 1963 (Texas J Sci 15:155-170). This was compiled from images of multiple clutches in Gamboa, Panama, and shows some of the variation in development at a given age.<\/p>\n<div style=\"clear: both;\"><\/div>\n","protected":false},"excerpt":{"rendered":"<p>Life cycle \u2013 natural history Red-eyed treefrogs live in low to mid-elevation rainforests from the Yucatan to Colombia. In the rainy season, males come to ponds and establish territories in plants near the water. They defend their territories and compete with other males by calling, shaking the plants, and fighting. When it rains, females carrying [&hellip;]<\/p>\n","protected":false},"author":7562,"featured_media":0,"parent":55,"menu_order":3,"comment_status":"closed","ping_status":"closed","template":"","meta":[],"_links":{"self":[{"href":"https:\/\/sites.bu.edu\/warkentinlab\/wp-json\/wp\/v2\/pages\/352"}],"collection":[{"href":"https:\/\/sites.bu.edu\/warkentinlab\/wp-json\/wp\/v2\/pages"}],"about":[{"href":"https:\/\/sites.bu.edu\/warkentinlab\/wp-json\/wp\/v2\/types\/page"}],"author":[{"embeddable":true,"href":"https:\/\/sites.bu.edu\/warkentinlab\/wp-json\/wp\/v2\/users\/7562"}],"replies":[{"embeddable":true,"href":"https:\/\/sites.bu.edu\/warkentinlab\/wp-json\/wp\/v2\/comments?post=352"}],"version-history":[{"count":29,"href":"https:\/\/sites.bu.edu\/warkentinlab\/wp-json\/wp\/v2\/pages\/352\/revisions"}],"predecessor-version":[{"id":645,"href":"https:\/\/sites.bu.edu\/warkentinlab\/wp-json\/wp\/v2\/pages\/352\/revisions\/645"}],"up":[{"embeddable":true,"href":"https:\/\/sites.bu.edu\/warkentinlab\/wp-json\/wp\/v2\/pages\/55"}],"wp:attachment":[{"href":"https:\/\/sites.bu.edu\/warkentinlab\/wp-json\/wp\/v2\/media?parent=352"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}