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55-661: The Ver is a 28 km (17 mi) long chalk stream in Hertfordshire , England . It is a tributary of the River Colne . The source is in the grounds of Lynch Lodge, Kensworth Lynch on the west side of the A5 trunk road and stays on the west side for some half mile or so. It then crosses through a culvert into Markyate Cell, afterwards crosses under the A5 in culvert and runs through Markyate . The river exits above ground at

110-435: A vertebral column (commonly known as a spine or backbone ), which evolved from the notochord . It is a paraphyletic grouping including all animals excluding the chordate subphylum Vertebrata , i.e. vertebrates . Well-known phyla of invertebrates include arthropods , mollusks , annelids , echinoderms , flatworms , cnidarians , and sponges . The majority of animal species are invertebrates; one estimate puts

165-528: A small proportion of the Metazoa that to speak of the kingdom Animalia in terms of "Vertebrata" and "Invertebrata" has limited practicality. In the more formal taxonomy of Animalia other attributes that logically should precede the presence or absence of the vertebral column in constructing a cladogram , for example, the presence of a notochord . That would at least circumscribe the Chordata. However, even

220-494: A source of information for forensic investigators. Two of the most commonly studied model organisms nowadays are invertebrates: the fruit fly Drosophila melanogaster and the nematode Caenorhabditis elegans . They have long been the most intensively studied model organisms , and were among the first life-forms to be genetically sequenced. This was facilitated by the severely reduced state of their genomes , but many genes , introns , and linkages have been lost. Analysis of

275-431: A wide range of invertebrate species, including annelids, molluscs, nematodes and arthropods. One type of invertebrate respiratory system is the open respiratory system composed of spiracles , tracheae, and tracheoles that terrestrial arthropods have to transport metabolic gases to and from tissues. The distribution of spiracles can vary greatly among the many orders of insects, but in general each segment of

330-552: Is also included within invertebrates: the Arthropoda, including insects, spiders , crabs , and their kin. All these organisms have a body divided into repeating segments, typically with paired appendages. In addition, they possess a hardened exoskeleton that is periodically shed during growth. Two smaller phyla, the Onychophora and Tardigrada , are close relatives of the arthropods and share some traits with them, excluding

385-580: Is believed to be the oldest extant bridge in Hertfordshire. It is Grade II listed . According to a contemporary account of the Second Battle of St Albans in 1461, another bridge existed on this site previously (recorded in 1505 as Pons de la Maltemyll - Malt Mill Bridge). It is thought that the Romans had built a bridge here by the 3rd century AD. The ford alongside the current bridge, which

440-500: Is easily seen in snails and sea snails , which have helical shells. Slugs appear externally symmetrical, but their pneumostome (breathing hole) is located on the right side. Other gastropods develop external asymmetry, such as Glaucus atlanticus that develops asymmetrical cerata as they mature. The origin of gastropod asymmetry is a subject of scientific debate. Other examples of asymmetry are found in fiddler crabs and hermit crabs . They often have one claw much larger than

495-485: Is fairly stable and rarely deviates from 10 °C (50 °F). On cold winter mornings, water vapour from the relatively warm stream condenses in the cold air above to form fog. Chalk is slightly soluble in rainwater because rain is naturally slightly acidic . The products of chalk weathering are dissolved in rainwater and are transported in stream flow. Chalk streams transport little suspended material (unlike most rivers), but are considered " mineral -rich" due to

550-770: Is known to have existed for 2,000 years and is traditionally believed to be Alban 's crossing point on his way to his execution, was substantially restored in 2001 by local residents' associations. Ye Olde Fighting Cocks public house in St Albans, which claims to be the oldest pub in England, was moved in the early 1600s to the side of the Ver next to the Abbey Mill at the bottom of the Abbey Orchard, and remains in this location today. The river south of Redbourn has been

605-474: Is not always precise among non-biologists since it does not accurately describe a taxon in the same way that Arthropoda , Vertebrata or Manidae do. Each of these terms describes a valid taxon, phylum , subphylum or family . "Invertebrata" is a term of convenience, not a taxon; it has very little circumscriptional significance except within the Chordata . The Vertebrata as a subphylum comprises such

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660-636: Is particularly salient in eusocial species but applies to other invertebrates as well. Insects recognize information transmitted by other insects. The term invertebrates covers several phyla. One of these are the sponges ( Porifera ). They were long thought to have diverged from other animals early. They lack the complex organization found in most other phyla. Their cells are differentiated, but in most cases not organized into distinct tissues. Sponges typically feed by drawing in water through pores. Some speculate that sponges are not so primitive, but may instead be secondarily simplified. The Ctenophora and

715-610: Is rated good or fail. Water quality of the River Ver in 2019: Chalk stream The beds of the rivers are generally composed of clean, compacted gravel and flints , which are good spawning areas for Salmonidae fish species. Since they are fed primarily by aquifers , the flow rate, mineral content and temperature range of chalk streams exhibit less seasonal variation than other rivers. They are mildly alkaline and contain high levels of nitrate , phosphate , potassium and silicate . In addition to algae and diatoms ,

770-737: The Acanthocephala , or spiny-headed worms, the Gnathostomulida , Micrognathozoa , and the Cycliophora . Also included are two of the most successful animal phyla, the Mollusca and Annelida. The former, which is the second-largest animal phylum by number of described species, includes animals such as snails , clams , and squids , and the latter comprises the segmented worms, such as earthworms and leeches . These two groups have long been considered close relatives because of

825-737: The Cnidaria , which includes sea anemones , corals , and jellyfish , are radially symmetric and have digestive chambers with a single opening, which serves as both the mouth and the anus. Both have distinct tissues, but they are not organized into organs . There are only two main germ layers, the ectoderm and endoderm , with only scattered cells between them. As such, they are sometimes called diploblastic . The Echinodermata are radially symmetric and exclusively marine, including starfish (Asteroidea), sea urchins , (Echinoidea), brittle stars (Ophiuroidea), sea cucumbers (Holothuroidea) and feather stars (Crinoidea). The largest animal phylum

880-536: The Nemertea , or ribbon worms, and the Sipuncula . Another phylum is Platyhelminthes , the flatworms. These were originally considered primitive, but it now appears they developed from more complex ancestors. Flatworms are acoelomates , lacking a body cavity, as are their closest relatives, the microscopic Gastrotricha . The Rotifera , or rotifers, are common in aqueous environments. Invertebrates also include

935-595: The Phanerozoic . Fossils of invertebrates are commonly used in stratigraphy. Carl Linnaeus divided these animals into only two groups, the Insecta and the now-obsolete Vermes ( worms ). Jean-Baptiste Lamarck , who was appointed to the position of "Curator of Insecta and Vermes" at the Muséum National d'Histoire Naturelle in 1793, both coined the term "invertebrate" to describe such animals and divided

990-666: The Protozoa , Porifera , Coelenterata , Platyhelminthes , Nematoda , Annelida , Echinodermata , Mollusca and Arthropoda . Arthropoda include insects , crustaceans and arachnids . By far the largest number of described invertebrate species are insects. The following table lists the number of described extant species for major invertebrate groups as estimated in the IUCN Red List of Threatened Species , 2014.3. The IUCN estimates that 66,178 extant vertebrate species have been described, which means that over 95% of

1045-591: The River Thames , River Lea and River Colne ): Chalk streams of the Yorkshire Wolds : Chalk streams of Lincolnshire : There are several chalk streams in the Lincolnshire Wolds including Chalk streams of Kent: Chalk streams of Norfolk: Chalk streams of Suffolk: Invertebrate Invertebrates is an umbrella term describing animals that neither develop nor retain

1100-457: The Tonian . Trace fossils such as tracks and burrows found in the late Neoproterozoic Era indicate the presence of triploblastic worms, roughly as large (about 5 mm wide) and complex as earthworms . Around 453 MYA, animals began diversifying, and many of the important groups of invertebrates diverged from one another. Fossils of invertebrates are found in various types of sediment from

1155-545: The gill chamber of their fish hosts ). Neurons differ in invertebrates from mammalian cells. Invertebrates cells fire in response to similar stimuli as mammals, such as tissue trauma, high temperature, or changes in pH. The first invertebrate in which a neuron cell was identified was the medicinal leech , Hirudo medicinalis . Learning and memory using nociceptors in the sea hare, Aplysia has been described. Mollusk neurons are able to detect increasing pressures and tissue trauma. Neurons have been identified in

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1210-414: The 1930s following the archaeological excavations of Verulamium by Sir Mortimer Wheeler and his wife Tessa . During the 1960s and 1970s it suffered serious problems as a result of water extraction upstream. Although these abated temporarily after the closure of one of the pumping stations, as of 2005 the upstream part of the river dries up completely during the summer, and the rest of the river may suffer

1265-718: The 1968 edition of Invertebrate Zoology , it is noted that "division of the Animal Kingdom into vertebrates and invertebrates is artificial and reflects human bias in favor of man's own relatives." The book also points out that the group lumps a vast number of species together, so that no one characteristic describes all invertebrates. In addition, some species included are only remotely related to one another, with some more related to vertebrates than other invertebrates (see Paraphyly ). For many centuries, invertebrates were neglected by biologists, in favor of big vertebrates and "useful" or charismatic species . Invertebrate biology

1320-595: The 20th and 21st centuries, much of that management has been aimed at producing the best conditions for fly fishing , and most specifically, dry fly fishing . The chalk streams hold a good number of wild salmonid fish species such as the brown trout ( Salmo trutta ), and grayling ( Thymallus thymallus ). In addition to these there are also considerable numbers of stocked brown trout and stocked rainbow trout ( Oncorhynchus mykiss ). The rich invertebrate life and characteristic transparent shallow water make chalk rivers and streams particularly suited to fly fishing. Many of

1375-447: The UK has caused the chalk streams to dry up. This has caused ecological damage and stagnant sewage that flows through the rivers and increasing phosphate levels. Although chalk streams are generally watercourses originating from chalk hills, including winterbournes , streams, and rivers, the term chalk stream is used even for larger rivers, which would normally be considered too large for

1430-538: The air sacs in their abdomen, are able to control the flow of air through their body. In some aquatic insects, the tracheae exchange gas through the body wall directly, in the form of a gill , or function essentially as normal, via a plastron . Despite being internal, the tracheae of arthropods are shed during moulting ( ecdysis ). Only vertebrate animals have ears, though many invertebrates detect sound using other kinds of sense organs. In insects, tympanal organs are used to hear distant sounds. They are located either on

1485-460: The body can have only one pair of spiracles, each of which connects to an atrium and has a relatively large tracheal tube behind it. The tracheae are invaginations of the cuticular exoskeleton that branch ( anastomose ) throughout the body with diameters from only a few micrometres up to 0.8 mm. The smallest tubes, tracheoles, penetrate cells and serve as sites of diffusion for water , oxygen , and carbon dioxide . Gas may be conducted through

1540-941: The call of her host, a male cricket. Depending on where the song of the cricket is coming from, the fly's hearing organs will reverberate at slightly different frequencies. This difference may be as little as 50 billionths of a second, but it is enough to allow the fly to home in directly on a singing male cricket and parasitise it. Like vertebrates, most invertebrates reproduce at least partly through sexual reproduction . They produce specialized reproductive cells that undergo meiosis to produce smaller, motile spermatozoa or larger, non-motile ova . These fuse to form zygotes , which develop into new individuals. Others are capable of asexual reproduction, or sometimes, both methods of reproduction. Extensive research with model invertebrate species such as Drosophila melanogaster and Caenorhabditis elegans has contributed much to our understanding of meiosis and reproduction. However, beyond

1595-615: The chalk stream springs are also used as sites for watercress production, due to the constant temperature and clean, alkaline , mineral-rich spring water. The Mid Hants Watercress Railway in Hampshire is so named on account of its use for transporting watercress to London from local chalk streams. A number of the chalk aquifers and associated groundwater sources related to chalk streams and rivers are used for water abstraction by local and national water utility companies. The over-abstracting of chalk streams by private water companies in

1650-826: The common presence of trochophore larvae, but the annelids were considered closer to the arthropods because they are both segmented. Now, this is generally considered convergent evolution , owing to many morphological and genetic differences between the two phyla. Among lesser phyla of invertebrates are the Hemichordata , or acorn worms, and the Chaetognatha, or arrow worms. Other phyla include Acoelomorpha , Brachiopoda , Bryozoa , Entoprocta , Phoronida , and Xenoturbellida . Invertebrates can be classified into several main categories, some of which are taxonomically obsolescent or debatable, but still used as terms of convenience. Each however appears in its own article at

1705-561: The conclusion that in vertebrates are a group that deviates from the normal, vertebrates. This has been said to be because researchers in the past, such as Lamarck, viewed vertebrates as a "standard": in Lamarck's theory of evolution, he believed that characteristics acquired through the evolutionary process involved not only survival, but also progression toward a "higher form", to which humans and vertebrates were closer than invertebrates were. Although goal-directed evolution has been abandoned,

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1760-566: The described animal species in the world are invertebrates. The trait that is common to all invertebrates is the absence of a vertebral column (backbone): this creates a distinction between invertebrates and vertebrates. The distinction is one of convenience only; it is not based on any clear biologically homologous trait, any more than the common trait of having wings functionally unites insects, bats, and birds, or than not having wings unites tortoises , snails and sponges . Being animals, invertebrates are heterotrophs, and require sustenance in

1815-574: The dissolved calcium and carbonate ions . The surface water of chalk streams is commonly described as "gin clear". The channel bed consists of angular flint gravel derived from the natural flint deposits found embedded within the chalk geology that contains relatively low amounts of clay and silt deposits. The unique characteristics of chalk stream ecology are due to stable temperature and flow regimes combined with highly transparent water and lack of sand grade sediment particles. The chalk streams have been intensively managed for many generations. In

1870-466: The distinction of invertebrates and vertebrates persists to this day, even though the grouping has been noted to be "hardly natural or even very sharp." Another reason cited for this continued distinction is that Lamarck created a precedent through his classifications which is now difficult to escape from. It is also possible that some humans believe that, they themselves being vertebrates, the group deserves more attention than invertebrates. In any event, in

1925-457: The few model systems, the modes of reproduction found in invertebrates show incredible diversity. In one extreme example, it is estimated that 10% of orbatid mite species have persisted without sexual reproduction and have reproduced asexually for more than 400 million years. Social behavior is widespread in invertebrates, including cockroaches, termites, aphids, thrips , ants, bees, Passalidae , Acari , spiders, and more. Social interaction

1980-534: The figure at 97%. Many invertebrate taxa have a greater number and diversity of species than the entire subphylum of Vertebrata. Invertebrates vary widely in size, from 10  μm (0.0004 in) myxozoans to the 9–10 m (30–33 ft) colossal squid . Some so-called invertebrates, such as the Tunicata and Cephalochordata , are actually sister chordate subphyla to Vertebrata, being more closely related to vertebrates than to other invertebrates. This makes

2035-470: The following links. The earliest animal fossils appear to be those of invertebrates. 665-million-year-old fossils in the Trezona Formation at Trezona Bore, West Central Flinders, South Australia have been interpreted as being early sponges. Some paleontologists suggest that animals appeared much earlier, possibly as early as 1 billion years ago though they probably became multicellular in

2090-555: The form of the consumption of other organisms. With a few exceptions, such as the Porifera , invertebrates generally have bodies composed of differentiated tissues. There is also typically a digestive chamber with one or two openings to the exterior. The body plans of most multicellular organisms exhibit some form of symmetry , whether radial, bilateral, or spherical. A minority, however, exhibit no symmetry. One example of asymmetric invertebrates includes all gastropod species. This

2145-401: The ground, where the chalk layer acts as an aquifer . The groundwater flows through the chalk bedrock, re-emerging lower down the slope in springs . The chalk acts as a temporary reservoir by regulating the amount of water supplied to the springs. This is why many chalk streams in the UK have stable flow regimes that vary only slightly over time. The temperature of the emerging surface water

2200-495: The hardened exoskeleton. The Nematoda , or roundworms, are perhaps the second largest animal phylum, and are also invertebrates. Roundworms are typically microscopic, and occur in nearly every environment where there is water. A number are important parasites. Smaller phyla related to them are the Kinorhyncha , Priapulida , and Loricifera . These groups have a reduced coelom, called a pseudocoelom. Other invertebrates include

2255-474: The head or elsewhere, depending on the insect family . The tympanal organs of some insects are extremely sensitive, offering acute hearing beyond that of most other animals. The female cricket fly Ormia ochracea has tympanal organs on each side of her abdomen. They are connected by a thin bridge of exoskeleton and they function like a tiny pair of eardrums, but, because they are linked, they provide acute directional information. The fly uses her "ears" to detect

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2310-529: The notochord would be a less fundamental criterion than aspects of embryological development and symmetry or perhaps bauplan . Despite this, the concept of invertebrates as a taxon of animals has persisted for over a century among the laity , and within the zoological community and in its literature it remains in use as a term of convenience for animals that are not members of the Vertebrata. The following text reflects earlier scientific understanding of

2365-563: The original two groups into ten, by splitting Arachnida and Crustacea from the Linnean Insecta, and Mollusca, Annelida, Cirripedia , Radiata , Coelenterata and Infusoria from the Linnean Vermes. They are now classified into over 30 phyla , from simple organisms such as sea sponges and flatworms to complex animals such as arthropods and molluscs. Invertebrates are animals without a vertebral column. This has led to

2420-422: The other. If a male fiddler loses its large claw, it will grow another on the opposite side after moulting . Sessile animals such as sponges are asymmetrical alongside coral colonies (with the exception of the individual polyps that exhibit radial symmetry); Alpheidae claws that lack pincers; and some copepods , polyopisthocotyleans , and monogeneans which parasitize by attachment or residency within

2475-441: The respiratory system by means of active ventilation or passive diffusion. Unlike vertebrates, insects do not generally carry oxygen in their haemolymph . A tracheal tube may contain ridge-like circumferential rings of taenidia in various geometries such as loops or helices . In the head , thorax , or abdomen , tracheae may also be connected to air sacs. Many insects, such as grasshoppers and bees , which actively pump

2530-426: The river systems in England. Each is given an overall ecological status, which may be one of five levels: high, good, moderate, poor and bad. There are several components that are used to determine this, including biological status, which looks at the quantity and varieties of invertebrates , angiosperms and fish. Chemical status, which compares the concentrations of various chemicals against known safe concentrations,

2585-616: The river's length (most notably in the unspoilt stretch between Redbourn and St Albans): the entire Ver valley was a national centre for the watercress growing industry. The poplar and willow plantations around Pre Mill and the former Pre Hotel on the A5183 (just west of St Albans on the edge of the Gorhambury Estate), which are a significant feature of the landscape, indicate a more modern industry: that of cricket bat manufacturing. The Environment Agency measure water quality of

2640-676: The same fate within a few years; compared to the current situation with the "great flow of water" that was reported to exist in 1885, with a depth of 12 ft (3.7 m) at Dolittle Mill on the Redbourn Road . In 2004 a proposal for remedial work was being developed for the St Albans lakes. The Romans built the city of Verulamium alongside the Ver at a time when it was navigable, expressing their preference for building in valleys rather than on hills. The bridge in St Michael's Street, adjacent to Kingsbury Mill, dates from 1765 and

2695-472: The site of several watermills down the centuries, mainly for grinding corn but also put to such diverse uses as paper making, fulling cloth, silk spinning and diamond lapping. Eleven mills are known to have existed, of which a number can still be seen today, either as mills (working or otherwise), or converted (for example into private homes or parts of public houses). Particularly worthy of note are: Disused watercress beds can be seen at various points along

2750-539: The southern end of Markyate , and on through Flamstead , Redbourn , St Albans and Park Street , finally joining the River Colne at Bricket Wood . The Ver is a chalk stream, which is partly a seasonal winterbourne north of Redbourn. However, many of its natural features have been compromised as a result of being canalised during the construction of the artificial lakes at Verulamium Park in St Albans in

2805-422: The streams provide a suitable habitat for macrophytes (including water crowfoot ) and oxygen levels are generally supportive of coarse fish populations. Of the 210 rivers classified as chalk streams globally, 160 are in England. A list of chalk streams in England gives a total of 224. Chalk is a highly porous and permeable rock, and rain falling onto chalk topography percolates directly into

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2860-757: The term stream . The Somme in northern France is a chalk stream on a larger scale. Winterbournes are known by different names depending on region: Chalk streams of the Southern England Chalk Formation in Berkshire , Hampshire , Wiltshire , Dorset and the Isle of Wight : Chalk streams of the Southern England Chalk Formation in the Chiltern Hills , Hertfordshire and Surrey (tributaries of

2915-511: The term "invertebrates" rather polyphyletic , so the term has little meaning in taxonomy . The word "invertebrate" comes from the Latin word vertebra , which means a joint in general, and sometimes specifically a joint from the spinal column of a vertebrate. The jointed aspect of vertebra is derived from the concept of turning, expressed in the root verto or vorto , to turn. The prefix in- means "not" or "without". The term invertebrates

2970-411: The term and of those animals which have constituted it. According to this understanding, invertebrates do not possess a skeleton of bone, either internal or external. They include hugely varied body plans . Many have fluid-filled, hydrostatic skeletons, like jellyfish or worms. Others have hard exoskeletons , outer shells like those of insects and crustaceans . The most familiar invertebrates include

3025-410: Was not a major field of study until the work of Linnaeus and Lamarck in the 18th century. During the 20th century, invertebrate zoology became one of the major fields of natural sciences, with prominent discoveries in the fields of medicine, genetics, palaeontology, and ecology. The study of invertebrates has also benefited law enforcement, as arthropods, and especially insects, were discovered to be

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