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103-462: Limulus is a genus of horseshoe crab , with one extant species, the Atlantic horseshoe crab ( Limulus polyphemus ). One fossil species is currently assigned to the genus though several other species have been named, which have since been assigned to other genera. Currently valid species include: Doubtful species include: Tentative species: This prehistoric arthropod –related article

206-466: A whole-genome duplication (WGD) event. This was followed by at least two, possibly three, WGDs in a common ancestor of the living horseshoe crabs. This gives them unusually large genomes for invertebrates (the genomes of C. rotundicauda and T. tridentatus being approximately 1.72 Gb each). Evidence for the duplication events includes similarity in structure between chromosomes ( synteny ), and clustering of homeobox genes . Over time, many of

309-555: A Berriasian–Barremian warm-dry phase, an Aptian–Santonian warm-wet phase, and a Campanian–Maastrichtian cool-dry phase. As in the Cenozoic, the 400,000 year eccentricity cycle was the dominant orbital cycle governing carbon flux between different reservoirs and influencing global climate. The location of the Intertropical Convergence Zone (ITCZ) was roughly the same as in the present. The cooling trend of

412-469: A cavity known as the food groove that begins near the pusher legs and extends to the animal's mouth . The end of the groove is closed off by the animal's chilaria . To break up any food , each pair of coxa moves in the opposite direction parallel to the ones in front of and behind it. This motion happens while feeding and walking, pushing food towards the mouth. Horseshoe crabs catch soft prey with claws on their second to fifth legs and place them in

515-615: A geologic signature associated with the mass extinction that lies between the Mesozoic and Cenozoic Eras . The Cretaceous as a separate period was first defined by Belgian geologist Jean d'Omalius d'Halloy in 1822 as the Terrain Crétacé , using strata in the Paris Basin and named for the extensive beds of chalk ( calcium carbonate deposited by the shells of marine invertebrates , principally coccoliths ), found in

618-463: A horseshoe crab is divided into three main parts: the cephalothorax , abdomen , and telson . The largest of these, the cephalothorax, houses most of the animal's eyes , limbs, and internal organs . It is also where the animal gets its name, as its shape somewhat resembles that of a horseshoe . Horseshoe crabs have changed little in appearance since they first evolved in the Triassic , earning them

721-439: A long, tubular heart located in the middle of their body. Like the hearts of vertebrates , the hearts of these animals have two separate states: a state of contraction known as systole , and a state of relaxation known as diastole . At the beginning of systole, blood leaves the heart through a large artery known as the aorta and numerous arteries parallel to the heart. Next, the arteries dump blood into large cavities of

824-517: A million times more sensitive to light at night than during the day. At the front of the animal along the cardiac ridge are a pair of eyes known as median ocelli . Their retina is even less organized than those of the compound eyes having between 5 and 11 photoreceptors paired with one or two secondary visual cells called arhabdomeric cells. Arhabdomeric cells are equivalent to eccentric cells as they function identically. The median ocelli are unique due to having two distinct visual pigments . While

927-674: A straight shell, flourished in the seas along with reef-building rudist clams. Inoceramids were also particularly notable among Cretaceous bivalves, and they have been used to identify major biotic turnovers such as at the Turonian-Coniacian boundary. Predatory gastropods with drilling habits were widespread. Globotruncanid foraminifera and echinoderms such as sea urchins and starfish (sea stars) thrived. Ostracods were abundant in Cretaceous marine settings; ostracod species characterised by high male sexual investment had

1030-563: A system of closed-off veins and arteries , gasses are transported through a cavity called the hemocoel. The hemocoel contains hemolymph , a fluid that fills all parts of the cavity and serves as the animal's blood. Rather than using iron -based hemoglobin , horseshoe crabs transport oxygen with a copper -based protein called hemocyanin , giving its blood a bright blue color. The blood also contains two types of cells: amebocytes that are utilized in clotting, and cyanocytes that create hemocyanin . Horseshoe crabs pump blood with

1133-564: A ~0.6 °C increase in temperature. The latter warming interval, occurring at the very end of the Cretaceous, was triggered by the activity of the Deccan Traps. The LKEPCI lasted into the Late Palaeocene , when it gave way to another supergreenhouse interval. The production of large quantities of magma, variously attributed to mantle plumes or to extensional tectonics , further pushed sea levels up, so that large areas of

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1236-419: Is biramous or divided into two separate branches. The branch closest to the front bears a flat end that looks like a leaf. This end is called the flabellum. The branch towards the back is far longer and looks similar to a walking leg. However, rather than ending in just a claw, the back branch has four leaf-like ends that are arranged like a petal . The final segment of the cephalothorax was originally part of

1339-410: Is J-shaped, lined with a cuticle , and can be divided into three main sections: the foregut, midgut, and hindgut. The foregut is contained in the animal's cephalothorax and comprises the esophagus , crop, and gizzard . The esophagus moves food from the mouth to the crop where it is stored before entering the gizzard. The gizzard is a muscular, toothed organ that serves to pulverize the food from

1442-926: Is a cladogram showing the internal relationships of Limulidae (modern horseshoe crabs) based on morphology. It contains both extant and extinct members. Carcinoscorpius rotundicauda Tachypleus gigas † Tachypleus decheni † Tachypleus syriacus Tachypleus tridentatus † Heterolimulus gadeai † Volanalimulus madagascarensis Limulus polyphemus † Limulus coffini † Crenatolimulus paluxyensis † Crenatolimulus darwini † Keuperlimulus vicensis † Casterolimulus kletti † Victalimulus mcqueeni † Allolimulus woodwardi † Mesolimulus crespelli † Mesolimulus walchi † Mesolimulus tafraoutensis † Mesolimulus sibiricus † Tarracolimulus rieki † Yunnanolimulus henkeli † Yunnanolimulus luopingensis The common ancestor of arachnids and xiphosurans (the group that includes horseshoe crabs) underwent

1545-403: Is a stub . You can help Misplaced Pages by expanding it . Horseshoe crab See text Horseshoe crabs are arthropods of the family Limulidae and are the only surviving xiphosurans . Despite their name, they are not true crabs or even crustaceans . Rather, they are chelicerates . This makes them more closely related to arachnids like spiders , ticks , and scorpions . The body of

1648-619: Is found in England, northern France, the low countries , northern Germany , Denmark and in the subsurface of the southern part of the North Sea . Chalk is not easily consolidated and the Chalk Group still consists of loose sediments in many places. The group also has other limestones and arenites . Among the fossils it contains are sea urchins , belemnites , ammonites and sea reptiles such as Mosasaurus . In southern Europe,

1751-434: Is likely due to a combination of two things: Like all arthropods , horseshoe crabs have segmented bodies with jointed limbs, which are covered by a protective cuticle made of chitin . They have heads composed of several segments, which eventually fuse as an embryo. Horseshoe crabs are chelicerates, meaning their bodies are composed of two main parts ( tagma ): the cephalothorax and the opisthosoma . The first tagma,

1854-473: Is mainly due to coastal habitat destruction and overharvesting. To ensure their continuous existence, many areas have enacted regulations on harvesting and established captive breeding programs. The fossil record of xiphosurans extends back to the Late Ordovician , or around 445 million years ago. For modern horseshoe crabs, their earliest appearance was approximately 250 million years ago during

1957-469: The Arachnida . This assumption was challenged when a genetics-based phylogeny found horseshoe crabs to be the sister group to the ricinuleids , thereby making them an arachnid . In response, a more recent paper has again placed horseshoe crabs as separate from the arachnids. This new study utilized both new and more complete sequencing data while also sampling a larger number of taxa . Below

2060-482: The Early Triassic . Because they have seen little morphological change since then, extant (surviving) forms have been described as " living fossils ". Horseshoe crabs resemble crustaceans but belong to a separate subphylum of the arthropods , Chelicerata . Horseshoe crabs are closely related to the extinct eurypterids (sea scorpions), which include some of the largest arthropods to have ever existed, and

2163-713: The Mancos Shale of western North America. These shales are an important source rock for oil and gas , for example in the subsurface of the North Sea. In northwestern Europe, chalk deposits from the Upper Cretaceous are characteristic for the Chalk Group , which forms the white cliffs of Dover on the south coast of England and similar cliffs on the French Normandian coast. The group

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2266-711: The North American Cordillera , as the Nevadan orogeny was followed by the Sevier and Laramide orogenies . Gondwana had begun to break up during the Jurassic Period, but its fragmentation accelerated during the Cretaceous and was largely complete by the end of the period. South America , Antarctica , and Australia rifted away from Africa (though India and Madagascar remained attached to each other until around 80 million years ago); thus,

2369-553: The Selli Event . Early Aptian tropical sea surface temperatures (SSTs) were 27–32 °C, based on TEX 86 measurements from the equatorial Pacific. During the Aptian, Milankovitch cycles governed the occurrence of anoxic events by modulating the intensity of the hydrological cycle and terrestrial runoff. The early Aptian was also notable for its millennial scale hyperarid events in the mid-latitudes of Asia. The BAWI itself

2472-728: The Turonian Age, based on isotopic evidence. However, this has subsequently been suggested to be the result of inconsistent isotopic proxies, with evidence of polar rainforests during this time interval at 82° S. Rafting by ice of stones into marine environments occurred during much of the Cretaceous, but evidence of deposition directly from glaciers is limited to the Early Cretaceous of the Eromanga Basin in southern Australia . Flowering plants (angiosperms) make up around 90% of living plant species today. Prior to

2575-467: The chelicerae , which give chelicerates their name. In horseshoe crabs, these look like tiny pincers in front of the mouth. Behind the chelicerae are the pedipalps , which are primarily used as legs. In the final molt of males, the ends of the pedipalps are modified into specialized, grasping claws used in mating. Following the pedipalps are three pairs of walking legs and a set of pusher legs for moving through soft sediment. Each of these pusher legs

2678-483: The tuatara ) disappeared from North America and Europe after the Early Cretaceous , and were absent from North Africa and northern South America by the early Late Cretaceous . The cause of the decline of Rhynchocephalia remains unclear, but has often been suggested to be due to competition with advanced lizards and mammals. They appear to have remained diverse in high-latitude southern South America during

2781-999: The Albian regularly expanded northward in tandem with expansions of subtropical high pressure belts. The Cedar Mountain Formation's Soap Wash flora indicates a mean annual temperature of between 19 and 26 °C in Utah at the Albian-Cenomanian boundary. Tropical SSTs during the Cenomanian-Turonian Thermal Maximum were at least 30 °C, though one study estimated them as high as between 33 and 42 °C. An intermediate estimate of ~33-34 °C has also been given. Meanwhile, deep ocean temperatures were as much as 15 to 20 °C (27 to 36 °F) warmer than today's; one study estimated that deep ocean temperatures were between 12 and 20 °C during

2884-486: The Cenomanian between the Albian and Turonian. The Cretaceous is divided into Early and Late Cretaceous epochs , or Lower and Upper Cretaceous series . In older literature, the Cretaceous is sometimes divided into three series: Neocomian (lower/early), Gallic (middle) and Senonian (upper/late). A subdivision into 12 stages , all originating from European stratigraphy, is now used worldwide. In many parts of

2987-571: The Cenozoic Era   —   the ichthyosaurs , last remaining temnospondyls ( Koolasuchus ), and nonmammalian cynodonts ( Tritylodontidae )   —   were already extinct millions of years before the event occurred. Coccolithophorids and molluscs , including ammonites , rudists , freshwater snails , and mussels , as well as organisms whose food chain included these shell builders, became extinct or suffered heavy losses. For example, ammonites are thought to have been

3090-645: The Coniacian through the Maastrichtian. During the Cretaceous, the late- Paleozoic -to-early-Mesozoic supercontinent of Pangaea completed its tectonic breakup into the present-day continents , although their positions were substantially different at the time. As the Atlantic Ocean widened, the convergent-margin mountain building ( orogenies ) that had begun during the Jurassic continued in

3193-626: The Cretaceous is sharply defined, being placed at an iridium -rich layer found worldwide that is believed to be associated with the Chicxulub impact crater , with its boundaries circumscribing parts of the Yucatán Peninsula and extending into the Gulf of Mexico . This layer has been dated at 66.043 Mya. At the end of the Cretaceous, the impact of a large body with the Earth may have been

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3296-570: The Cretaceous is usually a marine system consisting of competent limestone beds or incompetent marls . Because the Alpine mountain chains did not yet exist in the Cretaceous, these deposits formed on the southern edge of the European continental shelf , at the margin of the Tethys Ocean . During the Cretaceous, the present North American continent was isolated from the other continents. In

3399-537: The Cretaceous seas. Stagnation of deep sea currents in middle Cretaceous times caused anoxic conditions in the sea water leaving the deposited organic matter undecomposed. Half of the world's petroleum reserves were laid down at this time in the anoxic conditions of what would become the Persian Gulf and the Gulf of Mexico. In many places around the world, dark anoxic shales were formed during this interval, such as

3502-672: The Jurassic, the North Atlantic already opened, leaving a proto-ocean between Europe and North America. From north to south across the continent, the Western Interior Seaway started forming. This inland sea separated the elevated areas of Laramidia in the west and Appalachia in the east. Three dinosaur clades found in Laramidia (troodontids, therizinosaurids and oviraptorosaurs) are absent from Appalachia from

3605-576: The LKEPCI. During this period of relatively cool temperatures, the ITCZ became narrower, while the strength of both summer and winter monsoons in East Asia was directly correlated to atmospheric CO 2 concentrations. Laramidia likewise had a seasonal, monsoonal climate. The Maastrichtian was a time of chaotic, highly variable climate. Two upticks in global temperatures are known to have occurred during

3708-532: The Late Cretaceous, where lizards remained rare, with their remains outnumbering terrestrial lizards 200:1. Choristoderes , a group of freshwater aquatic reptiles that first appeared during the preceding Jurassic, underwent a major evolutionary radiation in Asia during the Early Cretaceous, which represents the high point of choristoderan diversity, including long necked forms such as Hyphalosaurus and

3811-555: The Late Cretaceous-Early Palaeogene Cool Interval (LKEPCI). Tropical SSTs declined from around 35 °C in the early Campanian to around 28 °C in the Maastrichtian. Deep ocean temperatures declined to 9 to 12 °C, though the shallow temperature gradient between tropical and polar seas remained. Regional conditions in the Western Interior Seaway changed little between the MKH and

3914-628: The MKH. Mean annual temperatures at the poles during the MKH exceeded 14 °C. Such hot temperatures during the MKH resulted in a very gentle temperature gradient from the equator to the poles; the latitudinal temperature gradient during the Cenomanian-Turonian Thermal Maximum was 0.54 °C per ° latitude for the Southern Hemisphere and 0.49 °C per ° latitude for the Northern Hemisphere, in contrast to present day values of 1.07 and 0.69 °C per ° latitude for

4017-526: The MKH. The poles were so warm that ectothermic reptiles were able to inhabit them. Beginning in the Santonian, near the end of the MKH, the global climate began to cool, with this cooling trend continuing across the Campanian. This period of cooling, driven by falling levels of atmospheric carbon dioxide, caused the end of the MKH and the transition into a cooler climatic interval, known formally as

4120-404: The Maastrichtian, bucking the trend of overall cooler temperatures during the LKEPCI. Between 70 and 69 Ma and 66–65 Ma, isotopic ratios indicate elevated atmospheric CO 2 pressures with levels of 1000–1400 ppmV and mean annual temperatures in west Texas between 21 and 23 °C (70 and 73 °F). Atmospheric CO 2 and temperature relations indicate a doubling of pCO 2 was accompanied by

4223-546: The Mesozoic) ended with the Cretaceous–Paleogene extinction event , a large mass extinction in which many groups, including non-avian dinosaurs, pterosaurs , and large marine reptiles , died out, widely thought to have been caused by the impact of a large asteroid that formed the Chicxulub crater in the Gulf of Mexico. The end of the Cretaceous is defined by the abrupt Cretaceous–Paleogene boundary (K–Pg boundary),

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4326-541: The South Atlantic and Indian Oceans were newly formed. Such active rifting lifted great undersea mountain chains along the welts, raising eustatic sea levels worldwide. To the north of Africa the Tethys Sea continued to narrow. During most of the Late Cretaceous, North America would be divided in two by the Western Interior Seaway , a large interior sea, separating Laramidia to the west and Appalachia to

4429-570: The Southern and Northern hemispheres, respectively. This meant weaker global winds, which drive the ocean currents, and resulted in less upwelling and more stagnant oceans than today. This is evidenced by widespread black shale deposition and frequent anoxic events . Tropical SSTs during the late Albian most likely averaged around 30 °C. Despite this high SST, seawater was not hypersaline at this time, as this would have required significantly higher temperatures still. On land, arid zones in

4532-576: The Tethys to the Arctic Ocean and enabling biotic exchange between the two oceans. At the peak of the Cretaceous transgression , one-third of Earth's present land area was submerged. The Cretaceous is justly famous for its chalk ; indeed, more chalk formed in the Cretaceous than in any other period in the Phanerozoic . Mid-ocean ridge activity—or rather, the circulation of seawater through

4635-441: The abdomen but fused in the embryo. On it are two flap-like appendages known as chilaria . If severed from the body, lost legs or the telson may slowly regenerate, and cracks in the body shell can heal. The opisthosoma or abdomen of a horseshoe crab is composed of several fused segments. Similar to a trilobite , the abdomen is made up of three lobes: a medial lobe in the middle, and a pleural lobe on either side. Attached to

4738-425: The ancestors of modern-day birds also diversified. They inhabited every continent, and were even found in cold polar latitudes. Pterosaurs were common in the early and middle Cretaceous, but as the Cretaceous proceeded they declined for poorly understood reasons (once thought to be due to competition with early birds , but now it is understood avian adaptive radiation is not consistent with pterosaur decline ). By

4841-531: The animal when walking around or swimming. The lateral eyes can be found directly behind the compound eyes and become functional just before a horseshoe crab larvae hatch. The telson's photoreceptors are unique as they're spaced throughout the structure rather than located in a fixed spot. Together with UV -seeing median ocelli, these photoreceptors have been found to influence the animal's circadian rhythm . Like all arthropods , horseshoe crabs have an open circulatory system . This means that instead of using

4944-489: The cephalothorax or prosoma , is a fusion of the head and thorax . This tagma is also covered by a large, semicircular, carapace that acts like a shield around the animal's body. It is shaped like the hoof of a horse, giving this animal its common name. In addition to the two main tagmata, the horseshoe crab also possesses a long tail-like section known as the telson . In total, horseshoe crabs have 6 pairs of appendages on their cephalothorax . The first of these are

5047-412: The compound eyes comprise around 1,000 individual units known as ommatidia . Each ommatidium is made up of a ring of retinal and pigment cells that surround something known as the eccentric cell. This secondary visual cell gets its name from the way it behaves. The eccentric cell is coupled with the dendrites of normal retinal cells so that when a normal cell depolarizes in the presence of light ,

5150-408: The continental crust were covered with shallow seas. The Tethys Sea connecting the tropical oceans east to west also helped to warm the global climate. Warm-adapted plant fossils are known from localities as far north as Alaska and Greenland , while dinosaur fossils have been found within 15 degrees of the Cretaceous south pole . It was suggested that there was Antarctic marine glaciation in

5253-508: The crop and regurgitate any indigestible particles. The foregut terminates in the pyloric valve and sphincter , a muscular door of sorts that separates it from the midgut. Cretaceous The Cretaceous was a period with a relatively warm climate , resulting in high eustatic sea levels that created numerous shallow inland seas . These oceans and seas were populated with now- extinct marine reptiles , ammonites , and rudists , while dinosaurs continued to dominate on land. The world

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5356-456: The duplicated genes have changed through processes of neofunctionalization or subfunctionalization , meaning their functions are different from what they originally were. Several hypotheses have been given as possible reasons why a size difference exists between male and female horseshoe crabs This phenomenon is known as sexual size dimorphism and results in the females having a larger average size than males. The existence of this trend

5459-463: The earliest crown group birds. Acanthomorph fish, the most diverse group of modern vertebrates, appeared in aquatic habitats around the middle of the Cretaceous. During the Early Cretaceous, flowering plants appeared and began to rapidly diversify, becoming the dominant group of plants across the Earth by the end of the Cretaceous, coincident with the decline and extinction of previously widespread gymnosperm groups. The Cretaceous (along with

5562-544: The early and mid-Cretaceous (becoming extinct during the late Cretaceous Cenomanian-Turonian anoxic event ), plesiosaurs throughout the entire period, and mosasaurs appearing in the Late Cretaceous. Sea turtles in the form of Cheloniidae and Panchelonioidea lived during the period and survived the extinction event. Panchelonioidea is today represented by a single species; the leatherback sea turtle . The Hesperornithiformes were flightless, marine diving birds that swam like grebes . Baculites , an ammonite genus with

5665-462: The east coasts of North America and South Asia . Horseshoe crabs are often caught for their blood , which contains Limulus amebocyte lysate , a chemical used to detect bacterial endotoxins . Additionally, the animals are used as fishing bait in the United States and eaten as a delicacy in some parts of Asia . In recent years, horseshoe crabs have experienced a population decline. This

5768-437: The east, then receded late in the period, leaving thick marine deposits sandwiched between coal beds. Bivalve palaeobiogeography also indicates that Africa was split in half by a shallow sea during the Coniacian and Santonian, connecting the Tethys with the South Atlantic by way of the central Sahara and Central Africa, which were then underwater. Yet another shallow seaway ran between what is now Norway and Greenland, connecting

5871-427: The eccentric cell does too. A horseshoe crab's compound eyes are less complex and organized than those of most other arthropods . Ommatidia are arranged messily in what's been deemed an "imperfect hexagonal array" and have a highly variable number of photoreceptors (between 4 and 20) in their retina . Although each ommatidium typically has one eccentric cell, there are sometimes two, and occasionally more. All

5974-449: The end of a horseshoe crab's abdomen is a long, tail-like spine known as a telson . It is highly mobile and serves a variety of functions. Horseshoe crabs have a variety of eyes that provide them with useful visual information. The most obvious of these are two large compound eyes found on top of the carapace . This feature is unusual, as all other living chelicerates have lost them in their evolution . In adult horseshoe crabs,

6077-747: The end of the AACS, which ended around 111 Ma with the Paquier/Urbino Thermal Maximum, giving way to the Mid-Cretaceous Hothouse (MKH), which lasted from the early Albian until the early Campanian. Faster rates of seafloor spreading and entry of carbon dioxide into the atmosphere are believed to have initiated this period of extreme warmth, along with high flood basalt activity. The MKH was punctuated by multiple thermal maxima of extreme warmth. The Leenhardt Thermal Event (LTE) occurred around 110 Ma, followed shortly by

6180-501: The end of the Cretaceous. The high sea level and warm climate of the Cretaceous meant large areas of the continents were covered by warm, shallow seas, providing habitat for many marine organisms. The Cretaceous was named for the extensive chalk deposits of this age in Europe, but in many parts of the world, the deposits from the Cretaceous are of marine limestone , a rock type that is formed under warm, shallow marine conditions. Due to

6283-463: The end of the period only three highly specialized families remained; Pteranodontidae , Nyctosauridae , and Azhdarchidae . The Liaoning lagerstätte ( Yixian Formation ) in China is an important site, full of preserved remains of numerous types of small dinosaurs, birds and mammals, that provides a glimpse of life in the Early Cretaceous. The coelurosaur dinosaurs found there represent types of

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6386-407: The enlarged ridges—enriched the oceans in calcium ; this made the oceans more saturated, as well as increased the bioavailability of the element for calcareous nanoplankton . These widespread carbonates and other sedimentary deposits make the Cretaceous rock record especially fine. Famous formations from North America include the rich marine fossils of Kansas 's Smoky Hill Chalk Member and

6489-528: The extinction fed on insects , larvae , worms , and snails, which in turn fed on dead plant and animal matter. Scientists theorise that these organisms survived the collapse of plant-based food chains because they fed on detritus . In stream communities , few groups of animals became extinct. Stream communities rely less on food from living plants and more on detritus that washes in from land. This particular ecological niche buffered them from extinction. Similar, but more complex patterns have been found in

6592-449: The eye's photoreceptors, both rods and cones , have a single visual pigment with a peak absorption of around 525 nanometers. This differs from insects or decapod crustaceans , as their photoreceptors are sensitive to different spectrums of light. Horseshoe crabs have relatively poor vision, and to compensate for that, have the largest rods and cones of any known animal, about 100 times the size of humans'. Furthermore, their eyes are

6695-431: The first functions similarly to the pigment in the compound eyes, the second has a peak absorption of around 360 nanometers, allowing the animal to see ultraviolet light . Other, more rudimentary eyes in horseshoe crabs include the endoparietal ocelli, the two lateral ocelli, two ventral ocelli, and a cluster of photoreceptors on the abdomen and telson. The endoparietal, lateral, and ventral ocelli are very similar to

6798-558: The first records of the gharial-like Neochoristodera , which appear to have evolved in the regional absence of aquatic neosuchian crocodyliformes. During the Late Cretaceous the neochoristodere Champsosaurus was widely distributed across western North America. Due to the extreme climatic warmth in the Arctic, choristoderans were able to colonise it too during the Late Cretaceous. In the seas, rays , modern sharks and teleosts became common. Marine reptiles included ichthyosaurs in

6901-416: The food groove to be ground up. For harder prey, Horseshoe crabs use a pair of stout, cuspid gnathobases (informally known as "nutcrackers") on the back of their sixth legs. After the food is sufficiently torn up, it is moved by the chelicerae into the mouth for further digestion . Horseshoe crabs are some of the only living chelicerates with guts that can process solid food. Its digestive system

7004-533: The four extant species is estimated to have lived about 135 million years ago in the Cretaceous . Limulidae is the only extant family of the order Xiphosura , and contains all four living species of horseshoe crabs: After Bicknell et al. 2021 and Lamsdell et al. 2020 The horseshoe crab's position within Chelicerata is complicated. However, most morphological analyses have placed them outside

7107-503: The genus Berriasella , but its use as a stratigraphic indicator has been questioned, as its first appearance does not correlate with that of C. alpina . The boundary is officially considered by the International Commission on Stratigraphy to be approximately 145 million years ago, but other estimates have been proposed based on U-Pb geochronology, ranging as young as 140 million years ago. The upper boundary of

7210-625: The group Maniraptora , which includes modern birds and their closest non-avian relatives, such as dromaeosaurs , oviraptorosaurs , therizinosaurs , troodontids along with other avialans . Fossils of these dinosaurs from the Liaoning lagerstätte are notable for the presence of hair-like feathers . Insects diversified during the Cretaceous, and the oldest known ants , termites and some lepidopterans , akin to butterflies and moths , appeared. Aphids , grasshoppers and gall wasps appeared. Rhynchocephalians (which today only includes

7313-453: The hemocoel surrounding the animal's tissues . Larger cavities lead to smaller cavities, allowing the hemocoel to oxygenate all the animal's tissues. During diastole, blood flows from the hemocoel to a cavity known as the pericardial sinus. From there, blood re-enters the heart and the cycle begins again. Horseshoe crabs breathe through modified swimming appendages beneath their abdomen known as book gills . While they appear smooth on

7416-412: The high sea level, there was extensive space for such sedimentation . Because of the relatively young age and great thickness of the system, Cretaceous rocks are evident in many areas worldwide. Chalk is a rock type characteristic for (but not restricted to) the Cretaceous. It consists of coccoliths , microscopically small calcite skeletons of coccolithophores , a type of algae that prospered in

7519-551: The highest rates of extinction and turnover. Thylacocephala , a class of crustaceans, went extinct in the Late Cretaceous. The first radiation of the diatoms (generally siliceous shelled, rather than calcareous ) in the oceans occurred during the Cretaceous; freshwater diatoms did not appear until the Miocene . Calcareous nannoplankton were important components of the marine microbiota and important as biostratigraphic markers and recorders of environmental change. The Cretaceous

7622-404: The lamellae are routinely aerated by rhythmic movement of the book gills. These movements create a current that enters through two gaps between the cephalothorax and abdomen and exits on either side of the telson . Horseshoe crabs first break up their food using bristles known as gnathobases located at the coxa or base of their walking limbs . Gnathobases on the right and left legs form

7725-714: The last epoch of the Jurassic, the Tithonian, continued into the Berriasian, the first age of the Cretaceous. The North Atlantic seaway opened and enabled the flow of cool water from the Boreal Ocean into the Tethys. There is evidence that snowfalls were common in the higher latitudes during this age, and the tropics became wetter than during the Triassic and Jurassic. Glaciation was restricted to high- latitude mountains, though seasonal snow may have existed farther from

7828-575: The late Valanginian (~ 134 million years ago) found in Israel and Italy, initially at low abundance. Molecular clock estimates conflict with fossil estimates, suggesting the diversification of crown-group angiosperms during the Late Triassic or the Jurassic, but such estimates are difficult to reconcile with the heavily sampled pollen record and the distinctive tricolpate to tricolporoidate (triple grooved) pollen of eudicot angiosperms. Among

7931-465: The late Cretaceous, and all else that depended on them suffered, as well. Herbivorous animals, which depended on plants and plankton as their food, died out as their food sources became scarce; consequently, the top predators , such as Tyrannosaurus rex , also perished. Yet only three major groups of tetrapods disappeared completely; the nonavian dinosaurs , the plesiosaurs and the pterosaurs . The other Cretaceous groups that did not survive into

8034-574: The latest Albian. Approximately 94 Ma, the Cenomanian-Turonian Thermal Maximum occurred, with this hyperthermal being the most extreme hothouse interval of the Cretaceous and being associated with a sea level highstand. Temperatures cooled down slightly over the next few million years, but then another thermal maximum, the Coniacian Thermal Maximum, happened, with this thermal event being dated to around 87 Ma. Atmospheric CO 2 levels may have varied by thousands of ppm throughout

8137-711: The l’Arboudeyesse Thermal Event (ATE) a million years later. Following these two hyperthermals was the Amadeus Thermal Maximum around 106 Ma, during the middle Albian. Then, around a million years after that, occurred the Petite Verol Thermal Event (PVTE). Afterwards, around 102.5 Ma, the Event 6 Thermal Event (EV6) took place; this event was itself followed by the Breistroffer Thermal Maximum around 101 Ma, during

8240-421: The median ocelli, except like the compound eyes, they only see in visual light with a peak absorbance of around 525 nanometers. The endoparietal eye further differs due to being a fusion of two separate ocelli . This eye is found not far behind the median eyes and sits directly on the cardiac ridge. The two ventral ocelli are located on the underside of the cephalothorax near the mouth and likely help to orient

8343-517: The middle Cretaceous, becoming the dominant group of land plants by the end of the period, coincident with the decline of previously dominant groups such as conifers. The oldest known fossils of grasses are from the Albian , with the family having diversified into modern groups by the end of the Cretaceous. The oldest large angiosperm trees are known from the Turonian (c. 90 Mya) of New Jersey, with

8446-470: The most promising candidates for fixing the Jurassic–Cretaceous boundary. In particular, the first appearance Calpionella alpina , coinciding with the base of the eponymous Alpina subzone, has been proposed as the definition of the base of the Cretaceous. The working definition for the boundary has often been placed as the first appearance of the ammonite Strambergella jacobi , formerly placed in

8549-824: The oceans. Extinction was more severe among animals living in the water column than among animals living on or in the seafloor. Animals in the water column are almost entirely dependent on primary production from living phytoplankton, while animals living on or in the ocean floor feed on detritus or can switch to detritus feeding. The largest air-breathing survivors of the event, crocodilians and champsosaurs , were semiaquatic and had access to detritus. Modern crocodilians can live as scavengers and can survive for months without food and go into hibernation when conditions are unfavorable, and their young are small, grow slowly, and feed largely on invertebrates and dead organisms or fragments of organisms for their first few years. These characteristics have been linked to crocodilian survival at

8652-601: The oldest records of Angiosperm macrofossils are Montsechia from the Barremian aged Las Hoyas beds of Spain and Archaefructus from the Barremian-Aptian boundary Yixian Formation in China. Tricolpate pollen distinctive of eudicots first appears in the Late Barremian, while the earliest remains of monocots are known from the Aptian. Flowering plants underwent a rapid radiation beginning during

8755-400: The outside are flat and broad, while the ones on the inside are more narrow. Closest to the front is a plate-like structure made of two fused appendages. This is the genital operculum and is where horseshoe crabs keep their reproductive organs . Following the operculum are five pairs of book gills . While mainly used for breathing, horseshoe crabs can also use their book gills to swim. At

8858-435: The outside, the insides of these book gills are lined with several thin "pages" called lamellae. Each lamella is hollow and contains an extension of hemocoel , allowing gasses to diffuse between a Horseshoe crab's blood and external environment. There are roughly 80–200 lamellae are present in each gill, with all ten of them giving the animal with a total breathing surface area of about two square meters. When underwater,

8961-461: The perimeter of each pleural lobe is a flat, serrated structure known as the flange. The flange on either side is connected by the telson embayment, which itself is attached to the medial lobe. Along the line where these lobes meet are six sets of indentations known as apodeme . Each of these serves as a muscle attachment point for the animal's twelve movable spines. On the underside of the abdomen are several biramous limbs. The branches closest to

9064-523: The poles. After the end of the first age, however, temperatures began to increase again, with a number of thermal excursions, such as the middle Valanginian Weissert Thermal Excursion (WTX), which was caused by the Paraná-Etendeka Large Igneous Province's activity. It was followed by the middle Hauterivian Faraoni Thermal Excursion (FTX) and the early Barremian Hauptblatterton Thermal Event (HTE). The HTE marked

9167-412: The principal food of mosasaurs , a group of giant marine lizards related to snakes that became extinct at the boundary. Omnivores , insectivores , and carrion -eaters survived the extinction event, perhaps because of the increased availability of their food sources. At the end of the Cretaceous, there seem to have been no purely herbivorous or carnivorous mammals . Mammals and birds that survived

9270-547: The punctuation mark at the end of a progressive decline in biodiversity during the Maastrichtian age. The result was the extinction of three-quarters of Earth's plant and animal species. The impact created the sharp break known as the K–Pg boundary (formerly known as the K–T boundary). Earth's biodiversity required substantial time to recover from this event, despite the probable existence of an abundance of vacant ecological niches . Despite

9373-744: The rise of angiosperms, during the Jurassic and the Early Cretaceous, the higher flora was dominated by gymnosperm groups, including cycads , conifers , ginkgophytes , gnetophytes and close relatives, as well as the extinct Bennettitales . Other groups of plants included pteridosperms or "seed ferns", a collective term that refers to disparate groups of extinct seed plants with fern-like foliage, including groups such as Corystospermaceae and Caytoniales . The exact origins of angiosperms are uncertain, although molecular evidence suggests that they are not closely related to any living group of gymnosperms. The earliest widely accepted evidence of flowering plants are monosulcate (single-grooved) pollen grains from

9476-451: The severity of the K-Pg extinction event, there were significant variations in the rate of extinction between and within different clades . Species that depended on photosynthesis declined or became extinct as atmospheric particles blocked solar energy . As is the case today, photosynthesizing organisms, such as phytoplankton and land plants , formed the primary part of the food chain in

9579-422: The strong regionality of most biostratigraphic markers, and the lack of any chemostratigraphic events, such as isotope excursions (large sudden changes in ratios of isotopes ) that could be used to define or correlate a boundary. Calpionellids , an enigmatic group of planktonic protists with urn-shaped calcitic tests briefly abundant during the latest Jurassic to earliest Cretaceous, have been suggested as

9682-633: The terrestrial fauna of the late Cretaceous Hell Creek Formation . Other important Cretaceous exposures occur in Europe (e.g., the Weald ) and China (the Yixian Formation ). In the area that is now India, massive lava beds called the Deccan Traps were erupted in the very late Cretaceous and early Paleocene. Palynological evidence indicates the Cretaceous climate had three broad phases:

9785-407: The title of " living fossil ". Only four species of horseshoe crab are extant today. Most are marine , though the mangrove horseshoe crab is often found in brackish water. Additionally, certain extinct species transitioned to living in freshwater . Horseshoe crabs primarily live at the water's bottom but they can swim if needed. In the modern day, their distribution is limited, only found along

9888-483: The trunk having a preserved diameter of 1.8 metres (5.9 ft) and an estimated height of 50 metres (160 ft). During the Cretaceous, ferns in the order Polypodiales , which make up 80% of living fern species, would also begin to diversify. On land, mammals were generally small sized, but a very relevant component of the fauna , with cimolodont multituberculates outnumbering dinosaurs in some sites. Neither true marsupials nor placentals existed until

9991-406: The two may be sister groups . The difficult-to-classify chasmataspidids are also thought to be closely related to horseshoe crabs. The radiation of horseshoe crabs resulted in 22 known species, of which only 4 remain. The Atlantic species is sister to the three Asian species, the latter of which are likely the result of two divergences relatively close in time. The last common ancestor of

10094-789: The ultimate end of the Tithonian-early Barremian Cool Interval (TEBCI). During this interval, precession was the dominant orbital driver of environmental changes in the Vocontian Basin. For much of the TEBCI, northern Gondwana experienced a monsoonal climate. A shallow thermocline existed in the mid-latitude Tethys. The TEBCI was followed by the Barremian-Aptian Warm Interval (BAWI). This hot climatic interval coincides with Manihiki and Ontong Java Plateau volcanism and with

10197-643: The upper Cretaceous of Western Europe . The name Cretaceous was derived from the Latin creta , meaning chalk . The twofold division of the Cretaceous was implemented by Conybeare and Phillips in 1822. Alcide d'Orbigny in 1840 divided the French Cretaceous into five étages (stages): the Neocomian , Aptian, Albian, Turonian, and Senonian, later adding the Urgonian between Neocomian and Aptian and

10300-628: The very end, but a variety of non-marsupial metatherians and non-placental eutherians had already begun to diversify greatly, ranging as carnivores ( Deltatheroida ), aquatic foragers ( Stagodontidae ) and herbivores ( Schowalteria , Zhelestidae ). Various "archaic" groups like eutriconodonts were common in the Early Cretaceous, but by the Late Cretaceous northern mammalian faunas were dominated by multituberculates and therians , with dryolestoids dominating South America . The apex predators were archosaurian reptiles , especially dinosaurs , which were at their most diverse stage. Avians such as

10403-407: The world, alternative local subdivisions are still in use. From youngest to oldest, the subdivisions of the Cretaceous period are: The lower boundary of the Cretaceous is currently undefined, and the Jurassic–Cretaceous boundary is currently the only system boundary to lack a defined Global Boundary Stratotype Section and Point (GSSP). Placing a GSSP for this boundary has been difficult because of

10506-654: Was followed by the Aptian-Albian Cold Snap (AACS) that began about 118 Ma. A short, relatively minor ice age may have occurred during this so-called "cold snap", as evidenced by glacial dropstones in the western parts of the Tethys Ocean and the expansion of calcareous nannofossils that dwelt in cold water into lower latitudes. The AACS is associated with an arid period in the Iberian Peninsula . Temperatures increased drastically after

10609-454: Was largely ice-free, although there is some evidence of brief periods of glaciation during the cooler first half, and forests extended to the poles. Many of the dominant taxonomic groups present in modern times can be ultimately traced back to origins in the Cretaceous. During this time, new groups of mammals and birds appeared, including the earliest relatives of placentals & marsupials ( Eutheria and Metatheria respectively), and

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