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Egg case (Chondrichthyes)

From Wikipedia, the free encyclopedia

Egg case of a skate
Embryo active inside egg case.

An egg case or egg capsule, often colloquially called a mermaid's purse, is the casing that surrounds the eggs of oviparous chondrichthyans. Living chondricthyans that produce egg cases include some sharks, skates and chimaeras. Egg cases typically contain one embryo, except for big skate and mottled skate egg cases, which contain up to 7 embryos.[1] Oviparity is completely absent in the superorder Squalomorphii.[2][3] Egg cases are also thought to have been produced by some extinct chondricthyan groups, such as hybodonts and xenacanths.

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Transcription

Next time someone asks you who you think you are, just give them the facts. You're a mammalian amniotic tetrapodal sarcopterygiian osteichthyan gnathostomal vertebrate cranial chordate. Yeah, it's a mouthful. And in order to understand what it means, you're going to have to understand the most complex group of animals on earth, and what it takes to get from this to this. The phylum Chordata accounts for all 52,000 species of vertebrates on earth and several thousand species of invertebrates. Together they range from tiny, brainless filter feeders all the way up to Scarlett Johansson. Now, you know by now that when we talk about classifying animals, we're really talking about their shared ancestry, each new branch on this tree marking an important new evolutionary milestone. And just like with tissue layers and segmentation in simpler animals, there are traits we can look for to track the evolution of chordates. By the time all of those traits appear in one organism, we'll have arrived at the most complex class within the most complex phlyum: the mammals. But first let's start with the fundamentals. We've talked before about synapomorphic traits: traits that set a group of animals apart from its ancestors and from other groups that came from the same ancestors. Chordates share four synapomorphies that make us who we are. Each of them is present at some point in every chordate's life cycle. How about a volunteer to demonstrate these traits? Ah, I see that the lancelets are raising their...mouthparts. The lancelets, also known as cephalochordata, literally "head-cords," are one of the three sub-phyla of chordates. And unlike almost all other chordates, these tiny, brainless, invertebrate filter feeders retain all four of these characteristics for their entire lives. You probably already know where most of these traits are going to appear, since the phylum is named after it: the spinal cord, or at least something that resembles a spinal chord. First there's the notochord, a structure made of cartilage that runs between an animal's digestive tube and its nerve cord. In most vertebrates, a skeleton develops around the notochord and allows the muscles to attach. In humans, the notochord is reduced to the disks of cartilage that we have between our vertebrae. Second, we have the nerve cord itself, called the dorsal hollow nerve cord a tube made of nerve fibers that develops into the central nervous system. This is what makes chordates different from other animal phyla, which have solid, ventral nerve cords, meaning they run along the front or stomach side. Third, all chordates have pharyngeal slits. In invertebrates like our lancelet here, they function as filters for feeding. In fish and other aquatic animals, they're gill slits, and in land-dwelling vertebrates like us they disappear before we're born, but that tissue develops into areas around our jaws, ears, and other structures in the head and neck. And finally, we can't forget our fourth synapomorphy, the post-anal tail, which is exactly what it sounds like. It helps propel aquatic animals through the water, it makes our dog look happy when she wags it, and in humans it shrinks during embryonic development into what is known as the coccyx, or tailbone. It's right here. And trust me, when it comes to tail placement, post-anal is the way to go. These four traits all began to appear during the Cambrian explosion more than 500 million years ago, and today they're shared by members of all three chordate subphyla even if the animals in those subphyla look pretty much nothing like each other For instance, our new friends here in cephalochordata are the oldest living subphylum, but you can't forget the other invertebrate group of chordates, urochordata, literally "tail-cords." There are over 2,000 species here, including sea squirts. If you're confused about why this ended up in a phylum with us, it's because they have tadpole-like larvae with all four chordate characteristics. The adults, which actually have a highly developed internal structure, with a heart and other organs, retain the pharyngeal slits but all other chordate features disappear or reform into other structures. The third and last and most complex subphylum, is the Vertebrata, and has the most species in it because its members have a hard backbone, which has allowed for an explosion in diversity, from tiny minnows to the great blue whale. You can see how fantastic this diversity really is when you break down Vertebrata into its many, many classes from slimy, sea-snakey things to us warm and fuzzy mammals. And as these classes become more complex, you can identify the traits they each developed that gave them an evolutionary edge over the ones that came before. For example, how's this for an awesome trait: a brain! Vertebrates with a head that contains sensory organs and a brain are called craniates. They also always have a heart with at least two chambers. Since this is science, you're going to have to know that there's going to be an exception for every rule that you're going to have to remember. and the exception in this case is the Myxini, or hagfish the only vertebrate class that has no vertebrae but is classified with us because it has a skull. This snake-like creature swims by using segmented muscles to exert force against its notochord. Whatever, hagfish. Closely-related to it is the class petromyzontida, otherwise known as lampreys, the oldest living lineage of vertebrates. Now these have a backbone, made of cartilage, and maybe even more important, a more complex nervous system. With the advent of the backbone we see vertebrates getting larger, developing more complex skeletons, and becoming more effective at catching food and avoiding predators. But do you notice anything missing? Lampreys and other early vertebrates are agnathans, literally "no-jaws." And if you want to be able to chew food, it really helps to have a jaw and teeth. Most scientists think that the jaw evolved from structures that supported the first two pharyngeal slits near the mouth. And teeth? Well, the current theory is that they evolved from sharp scales on the face! Gnathostomes, or "jaw-mouths", arrived on the scene 470 million years ago, and one of the oldest and most successful groups of gnathostomes that have survived to the present day are the class chondrichthyes, the "cartilage fish". You know them as sharks, skates and rays, and as their name says, their skeleton is made up mostly of cartilage, but they show the beginnings of a calcified skeleton. Chondrichthyans haven't changed much over the past 300 million years or so, and their success stems from the paired fins that allow for efficient swimming, and those jaws for biting off delicious hunks of flesh. If we're going to eventually get to mammals, we need bones, and we find those with the evolution of fish. Meet Osteichthyes, which technically means "the bony fish" unlike cartilaginous fish, members of this group have a mineralized endoskeleton. Now, Osteichthyes is sometimes considered a superclass, because it includes a whole slew of diverse classes that descended from it. There's actually some controversy among taxonomists about what to call it, but the main thing to know is that the majority of all vertebrates fall under Osteichthyes, and that includes you. It's broken up into two main groups, which themselves include a bunch of classes. The first is Actinopterygii, or ray-finned fishes, and with 27,000 species, pretty much every fish you've ever heard of is in this group. Ray-finned fishes evolved in fresh water, spread out into the oceans, and some eventually came back to fresh water. In the second group, things start to get really strange and interesting. These are the lobe-finned fishes, or sarcopterygii, a name derived from bones surrounded by muscle found in their pectoral and pelvic fins. And that sounds like something that could be used for walking! Lobe-fins include the coelacanths, which consist of one living species lungfish, which gulp air into their lungs; and tetrapods, which have adapted to land with four limbs. So this is weird, right? Even though land animals clearly are not fish, since tetrapods evolved from bony fish, they are filed under this group. These taxonomists, man. I want to party with them sometime. But first: Imagine that you're a fisherman off the coast of South Africa in the western Indian Ocean about 75 years ago. Just put that in your brain and hold on to it. And you've just pulled up a fish that no one has ever seen. Not only that, you've caught a fish that was thought to have become extinct 75 million years ago. This is exactly what happened in 1938, when Captain Hendrick Goosen hauled up a coelacanth, and it has mystified scientists ever since. A second population has since been identified near Indonesia in 1999, but the deep-sea creatures remain extremely rare. The coelacanth fascinates scientists because of its paired lobe fins. They extend from the body like legs and move in an alternating pattern. In other words, they move more like a horse than like a fish. And in fact those paired fins are supported by the very same bones that we have in our arms and legs. The coelacanth also has a hinged joint in the skull so it can widen its mouth to eat large prey, as well as thick scales that don't exist on any living fish. It's not good eating, but why would you want to eat what's essentially a living fossil? Alright, now we're talking about tetrapods, which of course means "four feet," and getting those four feet onto land was really awesome for those early creatures because that meant that they could escape the increasingly brutal and predatory world of the ocean. Tetrapods gradually replaced their fins with limbs, and developed entirely new body parts that were never seen before, like necks, with the help of additional vertebra, that separated the body from the head. The first tetrapods are today found in the class Amphibia, which were the first creatures to develop a three-chambered heart. There are more than 6,000 known species of amphibians like frogs and salamanders, most of which begin life as tadpoles in water, and then later develop legs, lungs and a digestive system, and often migrate to land for adulthood. But amphibians lay eggs that don't have shells, so they dehydrate quickly, so they have to be laid in water. So this leads us to our next evolutionary milestone for chordates: the amniotic egg. Amniotes are tetrapods that have eggs adapted for life on land a group that includes reptiles, birds and mammals. The amniotic egg was crucial for the success of land-dwellers, allowing embryos to develop in their own "private pond" of the amniotic sac, often surrounded by a hard shell in the case of reptiles and birds. The class Reptilia represents the earliest amniotes. Like amphibians, they have a 3-chambered heart, but they're totally terrestrial. And here's where we find our dinosaurs, snakes, turtles and lizards. You often hear reptiles described as "cold-blooded." This does not mean that their blood is cold. They're actually ectothermic, which means that they absorb external heat as their main source of body heat. Hence the lizard that likes to lay in the sun all day. The oldest group of reptiles, the archosaurs, mostly disappeared when most of the dinosaurs died out 65 million years ago. But two lineages of archosaurs survived. One includes the modern reptiles crocodiles and alligators, and the other is a type of dinosaur that we now call birds, the class Aves. There are big, obvious differences between these two surviving archosaurs: one is designed for eating and fighting big animals, while the other is designed for flying around and being graceful and stuff. The not-so-obvious but equally important difference is that birds are endotherms, which means that they can crank up their metabolism to regulate their body temperature. Making all that heat requires a big furnace, which is provided thanks to the evolution of a four-chambered heart. There's only one other group of animals that developed this trait, independently of birds, by the way, and it allowed them to spread through the planet. I'm talking, of course, about the class Mammalia, otherwise known as amniotes that have hair, three special ear bones and mammary glands. And most mammals have evolved to dispense with the hard egg shell altogether the embryo avoiding predation and other environmental dangers by developing inside the mother's body. In this class of chordates you'll find me, dame Judi Dench, your dog, your cat, Shamu the orca, African elephants, the South American pudu, and 5,300 other known species of mammals. It all began with a simple ancestor more than 500 million years in this crazy chordate phylum, but we've finally made it! And now you know exactly who you are. Thank you for watching this episode of Crash Course Biology, We hope that it was helpful and that you feel like a smarter person There's review stuff next to me that you can click on to go to those parts of the video that you maybe want to watch a little bit more of; to reinforce it in your brain. Thank to everybody who helped put this together. And if you have any questions for us you can get in touch with us on Facebook or Twitter. Or, of course, down in the comments below. Goodbye.

Description

Egg cases are made of collagen protein strands,[4] and are often described as feeling rough and leathery.[5] Some egg cases have a fibrous material covering the outside of the egg case, thought to aid in attachment to substrate.[1][6] Egg cases without a fibrous outer layer can be striated, bumpy, or smooth and glossy.[1][6] With the exception of bullhead shark eggs, egg cases are typically rectangular in shape with projections, called horns, at each corner.[1][6] Depending on the species, egg cases may have one or more tendrils.[5]

Development

Shortly after internal fertilization, the fertilized ovum enters the partially formed egg case located in the oviduct.[6] After the ovum enters, the rest of the egg case forms around it.[6] Shortly after the egg case finishes developing, it is deposited outside the body; common locations include kelp forests and rocky seafloors. Egg cases are typically produced in pairs, each with one fertilized embryo inside, with the exception of a few species that produce egg cases with more than one viable embryo.[1][6]

Gestation can take anywhere from a few months to over a year. After a period of development, typically a week or two, small slits open on each side of the egg case to aid water flow.[6] The embryo fans its tail constantly to promote exchange with surrounding water.[6]

Sharks

Egg case of a Port Jackson shark

Oviparity in sharks can be categorized as single or retained.[2] With single oviparity, the egg cases are extruded soon after fertilization.[2] With retained oviparity, eggs are kept within the oviduct for a period of time before depositing outside of the body as an unhatched egg case.[2] It is thought that viviparity is the ancestral condition for sharks, and that it evolved through the elongation of retention time of retained oviparity.[3]

Oviparous sharks are known to regularly produce unfertilized eggs when kept in captivity without males.[7]

Bullhead sharks

Bullhead shark egg cases are shaped like an auger, with two spiral flanges. This allows the egg cases to become wedged in the crevices of rocky sea floors, where the eggs are protected from predators; however, some bullhead sharks deposit their eggs on sponges or seaweed.[8][5] Hatchlings are considered large for sharks, reaching over 14 cm in length by the time they leave the egg case.[5] Bullhead shark eggs typically hatch after 7 to 12 months, depending on the species.[5] The female Japanese bullhead shark has been known to deposit their eggs in one location along with other females, called a "nest".[5] The egg case of the Mexican hornshark features a tendril and more rigid flanges, suggesting that egg case design of this species is evolving towards anchoring with tendrils and away from wedging into crevices.[5] As a member of the order Heterodontiformes, the whitespotted bullhead sharks is thought to be oviparous, but egg cases have never been observed.[5]

Egg case of a brownbanded bamboo shark

Carpet sharks

The bamboo sharks (Hemiscylliidae) and the zebra shark (Stegostomatidae) lay eggs on the bottom, while the other carpet sharks give live birth. The egg cases are oval and covered with adhesive fibers that serve to secure them to the bottom.

Ground sharks

Egg cases of a catshark

Some catsharks (Scyliorhinidae) and the finback catsharks in the genus Proscyllium are the only members of their order that lay eggs. The egg cases of catsharks are purse-shaped with long tendrils at the corners that serve to anchor them to structures on the sea floor.

The size of egg cases vary; those of the small-spotted catshark or lesser spotted dogfish, Scyliorhinus canicula, are around 5 centimetres (2 in) long, while those of the greater spotted dogfish, S. stellaris, are around 10 centimetres (4 in). That excludes the four long tendrils found in each corner, which assist in anchorage. Egg cases from rays vary in that they have points rather than tendrils. The colours and shapes of egg cases also vary greatly from species to species.

Skates

Egg case of a big skate

The skates (Rajidae, Arhynchobatidae, Anacanthobatidae) are the only rays that are oviparous.[9] Females lay egg cases onto the sea floor after fertilization occurs in utero. While in utero, a protected case forms around the embryo which is called the egg case.[1][10] Studies have been done where egg cases were removed from gravid females to ensure proper identification in regard to skate species.[1] Egg cases have distinguishable characteristic traits that are unique to that species, thus making it a great tool for identifying a skate. The two most distinguishable features on the egg case are the keel and the absence or presence of a fibrous covering. A keel runs laterally along both sides of the outer edge of the egg case; it is a flexible structure. Keels will also run the length of the horns on some skate species. Some egg cases have broad keels (greater than 10% of the maximum egg case width) while others have narrow keels (less than 10% of the maximum egg case width).[1] Many egg cases are covered with a layer of fiber; some will have a fine layer while others have a thick layer.

Big skate

Big skate egg cases are larger than most other skate egg cases; typically ranging from 210 to 280 mm in length and 110 to 180 mm in width.[1][11] Big skates egg cases are approximately 15% of the overall length of the female skate.[1] The egg case is very smooth and lacks external fibrous material.[1] This egg case can be easily identified from all others in that it is the only one to have a steep ridge; giving the case a convex shape.[1] The keel on the egg case is considered very broad; representing 30–33% of the width of the egg case.[1]

Big skates are one of only two skates known to have multiple embryos inside an egg case; up to 7 embryos have been found inside a single case. But most big skate egg cases contain 3–4 embryos.[12]

Longnose skate

The longnose skate, Raja rhina, is considered a larger skate species; reaching a maximum size range of 145 cm total length.[12] Although their egg cases are smaller than that of the big skate, their cases are also considered large; ranging 93–102 mm in length.[1] Egg cases contain a single embryo. Longnose skate egg cases found in the field are brown in color. The external side is covered with a fibrous material, which is thicker on the top side and thinner on the bottom side of the case. The case is smooth underneath the fibrous material.[1]

Chimaeras

Egg case of a Cape elephantfish

All known chimaeras produce egg cases.[13] The egg cases of chimaeras are spindle- or bottle-shaped with fins on the sides. They are laid on the bottom of the sea floor. Chimaeras (subclass Holocephali), some sharks, and skates are among the 43% of known Chondrichthian species to exhibit oviparity.[14] However, there are some key morphological differences that are specific to chimaeras. The holocephalan egg capsule, or egg case, has a bulbous center flanked laterally by flattened collagen tissue. The flattened collagen tissue joins on the anterior end of the egg capsule to form a tail.[14] Sharp projections located on the anterior and posterior end of the egg case serve to better secure the egg case in between rocks, as well as protection against potential predators.[15]

Extinct chondrichthyans

Illustration of an Early Jurassic estuarine ecosystem, with Palaeoxyris egg cases attached via tendril to Neocalamites stems, with hybodonts in the background

The egg case genera Palaeoxyris and Fayolia, which are thought to have been produced by hybodonts and xenacanths respectively, two groups of extinct shark-like cartilaginous fish more closely related to modern sharks and rays than to chimaeras, resemble those of bulldog sharks in having a spiral collarettes running around them. Both Palaeoxyris and Fayolia taper towards their ends (with the tapering being more pronounced in Palaeoxyris), with one end having a tendril. Unlike modern sharks, these eggs are typically found in freshwater environments.[16]

Threats

Predation on egg cases is thought to be a major source of mortality for developing oviparous sharks, skates and chimaeras.[17] In general, predation is the leading cause of mortality for marine fish eggs, due to their abundance and high nutritional value.[17] Parental care ends when the egg case is released from the body, so the embryo relies on its tough, leathery exterior as its only source of protection.[17] Some gastropods are known to feed on egg cases by boring into the exterior.[17] Sharks are also common predators of egg cases.[6]

References

  1. ^ a b c d e f g h i j k l m n o Ebert, David A., Davis, Chante D. (2007). "Descriptions of skate egg cases (Chondrichthyes: Rajiformes: Rajoidei) from the eastern North Pacific". Zootaxa 1393: 1-18.
  2. ^ a b c d Carrier, J.C; Musick, J.A.; Heithaus, M.R. (2012). Biology of Sharks: Second Edition. Taylor & Francis Group. p. 296.
  3. ^ a b Carrier, J.C; Musick, J.A.; Heithaus, M.R. (2004). Biology of Sharks: First Edition. Taylor & Francis Group. p. 270.
  4. ^ Evans, David H. (June 1981). "The egg case of the oviparous elasmobranch, Raja Erinacea, does osmoregulate" (PDF). Journal of Experimental Biology. 92. doi:10.1242/jeb.92.1.337.
  5. ^ a b c d e f g h Compagno, Leonard (2002). "Sharks of the World". FAO Species Catalogue for Fishery Purposes. 2: 31–50.
  6. ^ a b c d e f g h i Treloar, M.A.; Laurenson, L.J.B.; Stevens, J.D. (2006). "Description of Rajid egg cases from southeastern Australian waters". Zootaxa. 1231: 53. doi:10.11646/zootaxa.1231.1.3.
  7. ^ "Egg Identification". Aquarium of the Pacific. 2008.
  8. ^ Buch, Robert. "Heterodontus francisci". Florida Museum.
  9. ^ "Most Commonly Asked Questions". Florida Museum of Natural History.
  10. ^ "Raja binoculata (Big Skate, Big Skate)". Zipcodezoo.com. Archived from the original on 8 September 2008.
  11. ^ "Raja binoculata (Big Skate, Big Skate)". Zipcodezoo.com. Archived from the original on 8 September 2008.
  12. ^ a b Ebert, D.A., Smith, W.D., and Cailliet, G.M. (2008). "Reproductive biology of two commercially exploited skates, Raja binoculata and R. rhina, in the western Gulf of Alaska". Fisheries Research, 94:48-57. doi:10.1016/j.fishres.2008.06.016
  13. ^ García-Salinas, Pablo; Gallego, Victor; Asturiano, Juan F. (23 July 2021). "Reproductive Anatomy of Chondrichthyans: Notes on Specimen Handling and Sperm Extraction. II. Sharks and Chimaeras". Animals. 11 (8): 2191. doi:10.3390/ani11082191. ISSN 2076-2615. PMC 8388383. PMID 34438648.
  14. ^ a b Fischer, Jan, Martin Licht, Jürgen Kriwet, Jörg W. Schneider, Michael Buchwitz, and Peter Bartsch. "Egg capsule morphology provides new information about the interrelationships of chondrichthyan fishes." Journal of Systematic Palaeontology 12.3 (2013): 389-99.
  15. ^ Klimley, A. Peter (2013). The Biology of Sharks and Rays. Chicago, Illinois: The University of Chicago Press. pp. 286–288. ISBN 978-0226442495.
  16. ^ Fischer, Jan; Licht, Martin; Kriwet, Jürgen; Schneider, Jörg W.; Buchwitz, Michael; Bartsch, Peter (3 April 2014). "Egg capsule morphology provides new information about the interrelationships of chondrichthyan fishes". Journal of Systematic Palaeontology. 12 (3): 389–399. Bibcode:2014JSPal..12..389F. doi:10.1080/14772019.2012.762061. ISSN 1477-2019. S2CID 84827548.
  17. ^ a b c d Lucifora, L.O.; Garcia, V.B. (2004). "Gastropod predation on egg cases of skates (Chondrichthyes, Rajidae)". Marine Biology. 145: 917–922. doi:10.1007/s00227-004-1377-8. S2CID 53508039.

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This page was last edited on 12 February 2024, at 06:29
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