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Marine invertebrates
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Ernst Haeckel's 96th plate, showing some marine invertebrates. Marine invertebrates have a large variety of body plans, which are currently categorised into over 30 phyla.
Marine invertebrates are the invertebrates that live in marine habitats. Invertebrate is a blanket term that includes all animals apart from the vertebrate members of the chordate phylum. Invertebrates lack a vertebral column, and some have evolved a shell or a hard exoskeleton. As on land and in the air, marine invertebrates have a large variety of body plans, and have been categorised into over 30 phyla. They make up most of the macroscopic life in the oceans.
Contents
1 Evolution
2 Classification
3 Marine sponges
4 Marine cnidarians
5 Marine worms
6 Echinoderms
7 Marine molluscs
8 Marine arthropods
9 Other phyla
10 Minerals from sea water
11 See also
12 Notes
13 References
14 Other references
Evolution[edit]
Dickinsonia may be the earliest animal. They appear in the fossil record 571 million to 541 million years ago.
Kimberella, an early mollusc important for understanding the Cambrian explosion. Invertebrates are grouped into different phyla (body plans).
Opabinia an extinct, stem group arthropod that appeared in the Middle Cambrian[1]:124–136
See also: Avalon explosion and Cambrian explosion
The earliest animals were marine invertebrates, that is, vertebrates came later. Animals are multicellular eukaryotes,[note 1] and are distinguished from plants, algae, and fungi by lacking cell walls.[2] Marine invertebrates are animals that inhabit a marine environment apart from the vertebrate members of the chordate phylum; invertebrates lack a vertebral column. Some have evolved a shell or a hard exoskeleton.
The earliest animals may belong to the genus Dickinsonia,[3] 571 million to 541 million years ago.[4] Individual Dickinsonia typically resemble a bilaterally symmetrical ribbed oval. They kept growing until they were covered with sediment or otherwise killed,[5] and spent most of their lives with their bodies firmly anchored to the sediment.[6] Their taxonomic affinities are presently unknown, but their mode of growth is consistent with a bilaterian affinity.[7]
Apart from Dickinsonia, the earliest widely accepted animal fossils are the rather modern-looking cnidarians (the group that includes jellyfish, sea anemones and Hydra), possibly from around 580 Ma[8] The Ediacara biota, which flourished for the last 40 million years before the start of the Cambrian,[9] were the first animals more than a very few centimetres long. Like Dickinsonia, many were flat with a "quilted" appearance, and seemed so strange that there was a proposal to classify them as a separate kingdom, Vendozoa.[10] Others, however, have been interpreted as early molluscs (Kimberella[11][12]), echinoderms (Arkarua[13]), and arthropods (Spriggina,[14] Parvancorina[15]). There is still debate about the classification of these specimens, mainly because the diagnostic features which allow taxonomists to classify more recent organisms, such as similarities to living organisms, are generally absent in the Ediacarans. However, there seems little doubt that Kimberella was at least a triploblastic bilaterian animal, in other words, an animal significantly more complex than the cnidarians.[16]
The small shelly fauna are a very mixed collection of fossils found between the Late Ediacaran and Middle Cambrian periods. The earliest, Cloudina, shows signs of successful defense against predation and may indicate the start of an evolutionary arms race. Some tiny Early Cambrian shells almost certainly belonged to molluscs, while the owners of some "armor plates," Halkieria and Microdictyon, were eventually identified when more complete specimens were found in Cambrian lagerstätten that preserved soft-bodied animals.[17]
In the 1970s there was already a debate about whether the emergence of the modern phyla was "explosive" or gradual but hidden by the shortage of Precambrian animal fossils.[17] A re-analysis of fossils from the Burgess Shale lagerstätte increased interest in the issue when it revealed animals, such as Opabinia, which did not fit into any known phylum. At the time these were interpreted as evidence that the modern phyla had evolved very rapidly in the Cambrian explosion and that the Burgess Shale's "weird wonders" showed that the Early Cambrian was a uniquely experimental period of animal evolution.[1] Later discoveries of similar animals and the development of new theoretical approaches led to the conclusion that many of the "weird wonders" were evolutionary "aunts" or "cousins" of modern groups[18]—for example that Opabinia was a member of the lobopods, a group which includes the ancestors of the arthropods, and that it may have been closely related to the modern tardigrades.[19] Nevertheless, there is still much debate about whether the Cambrian explosion was really explosive and, if so, how and why it happened and why it appears unique in the history of animals.[20]
Classification[edit]
Bryozoa, from Ernst Haeckel's Kunstformen der Natur, 1904
Invertebrates are grouped into different phyla. Informally phyla can be thought of as a way of grouping organisms according to their body plan.[21][22]:33 A body plan refers to a blueprint which describes the shape or morphology of an organism, such as its symmetry, segmentation and the disposition of its appendages. The idea of body plans originated with vertebrates, which were grouped into one phylum. But the vertebrate body plan is only one of many, and invertebrates consist of many phyla or body plans. The history of the discovery of body plans can be seen as a movement from a worldview centred on vertebrates, to seeing the vertebrates as one body plan among many. Among the pioneering zoologists, Linnaeus identified two body plans outside the vertebrates; Cuvier identified three; and Haeckel had four, as well as the Protista with eight more, for a total of twelve. For comparison, the number of phyla recognised by modern zoologists has risen to 35.[22]
Historically body plans were thought of as having evolved in rapidly during the Cambrian explosion,[23] but a more nuanced understanding of animal evolution suggests a gradual development of body plans throughout the early Palaeozoic and beyond.[24] More generally a phylum can be defined in two ways: as described above, as a group of organisms with a certain degree of morphological or developmental similarity (the phenetic definition), or a group of organisms with a certain degree of evolutionary relatedness (the phylogenetic definition).[24]
As on land and in the air, invertebrates make up a great majority of all macroscopic life, as the vertebrates makes up a subphylum of one of over 30 known animal phyla, making the term almost meaningless for taxonomic purpose. Invertebrate sea life includes the following groups, some of which are phyla:
The 49th plate from Ernst Haeckel's Kunstformen der Natur, 1904, showing various sea anemones classified as Actiniae, in the Cnidaria phylum
"A variety of marine worms": plate from Das Meer by M.J. Schleiden (1804–1881)
Acoela, among the most primitive bilateral animals;
Annelida, (polychaetes and sea leeches);
Brachiopoda, marine animals that have hard "valves" (shells) on the upper and lower surfaces ;
Bryozoa, also known as moss animals or sea mats;
Chaetognatha, commonly known as arrow worms, are a phylum of predatory marine worms that are a major component of plankton;
Cephalochordata represented in the modern oceans by the lancelets (also known as Amphioxus);
Cnidaria, such as jellyfish, sea anemones, and corals;
Crustacea, including lobsters, crabs, shrimp, crayfish, barnacles, hermit crabs, mantis shrimps, and copepods;
Ctenophora, also known as comb jellies, the largest animals that swim by means of cilia;
Echinodermata, including sea stars, brittle stars, sea urchins, sand dollars, sea cucumbers, crinoids, and sea daisies;
Echiura, also known as spoon worms;
Gnathostomulids, slender to thread-like worms, with a transparent body that inhabit sand and mud beneath shallow coastal waters;
Gastrotricha, often called hairy backs, found mostly interstitially in between sediment particles;
Hemichordata, includes acorn worms, solitary worm-shaped organisms;
Kamptozoa, goblet-shaped sessile aquatic animals, with relatively long stalks and a "crown" of solid tentacles, also called Entoprocta;
Kinorhyncha, segmented, limbless animals, widespread in mud or sand at all depths, also called mud dragons;
Loricifera, very small to microscopic marine sediment-dwelling animals only discovered in 1983;
Mollusca, including shellfish, squid, octopus, whelks, Nautilus, cuttlefish, nudibranchs, scallops, sea snails, Aplacophora, Caudofoveata, Monoplacophora, Polyplacophora, and Scaphopoda;
Myzostomida, a taxonomic group of small marine worms which are parasitic on crinoids or "sea lilies";
Nemertinea, also known as "ribbon worms" or "proboscis worms";
Orthonectida, a small phylum of poorly known parasites of marine invertebrates that are among the simplest of multi-cellular organisms;
Phoronida, a phylum of marine animals that filter-feed with a lophophore (a "crown" of tentacles), and build upright tubes of chitin to support and protect their soft bodies;
Placozoa, small, flattened, multicellular animals around 1 millimetre across and the simplest in structure. They have no regular outline, although the lower surface is somewhat concave, and the upper surface is always flattened;
Porifera (sponges), multicellular organisms that have bodies full of pores and channels allowing water to circulate through them;
Priapulida, or penis worms, are a phylum of marine worms that live marine mud. They are named for their extensible spiny proboscis, which, in some species, may have a shape like that of a human penis;
Pycnogonida, also called sea spiders, are unrelated to spiders, or even to arachnids which they resemble;
Sipunculida, also called peanut worms, is a group containing 144–320 species (estimates vary) of bilaterally symmetrical, unsegmented marine worms;
Tunicata, also known as sea squirts or sea pork, are filter feeders attached to rocks or similarly suitable surfaces on the ocean floor;
Some flatworms of the classes Turbellaria and Monogenea;
Xenoturbella, a genus of bilaterian animals that contains only two marine worm-like species;
Xiphosura, includes a large number of extinct lineages and only four recent species in the family Limulidae, which include the horseshoe crabs.
Arthropods total about 1,113,000 described extant species, molluscs about 85,000 and chordates about 52,000.[25]
Marine sponges[edit]
Sponges have no nervous, digestive or circulatory system
Sponges are animals of the phylum Porifera (Modern Latin for bearing pores [26]). They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like mesohyl sandwiched between two thin layers of cells. They have unspecialized cells that can transform into other types and that often migrate between the main cell layers and the mesohyl in the process. Sponges do not have nervous, digestive or circulatory systems. Instead, most rely on maintaining a constant water flow through their bodies to obtain food and oxygen and to remove wastes.
Sponges are similar to other animals in that they are multicellular, heterotrophic, lack cell walls and produce sperm cells. Unlike other animals, they lack true tissues and organs, and have no body symmetry. The shapes of their bodies are adapted for maximal efficiency of water flow through the central cavity, where it deposits nutrients, and leaves through a hole called the osculum. Many sponges have internal skeletons of spongin and/or spicules of calcium carbonate or silicon dioxide. All sponges are sessile aquatic animals. Although there are freshwater species, the great majority are marine (salt water) species, ranging from tidal zones to depths exceeding 8,800 m (5.5 mi).
While most of the approximately 5,000–10,000 known species feed on bacteria and other food particles in the water, some host photosynthesizing micro-organisms as endosymbionts and these alliances often produce more food and oxygen than they consume. A few species of sponge that live in food-poor environments have become carnivores that prey mainly on small crustaceans.[27]
Sponge biodiversity. There are four sponge species in this photo.
Branching vase sponge
Venus' flower basket at a depth of 2572 meters
Barrel sponge
Stove-pipe sponge
Linnaeus mistakenly identified sponges as plants in the order Algae.[28] For a long time thereafter sponges were assigned to a separate subkingdom, Parazoa (meaning beside the animals).[29] They are now classified as a paraphyletic phylum from which the higher animals have evolved.[30]
Marine cnidarians[edit]
Cnidarians are the simplest animals with cells organised into tissues. Yet the starlet sea anemone contains the same genes as those that form the vertebrate head.
Cnidarians (Greek for nettle) are distinguished by the presence of stinging cells, specialized cells that they use mainly for capturing prey. Cnidarians include corals, sea anemones, jellyfish and hydrozoans. They form a phylum containing over 10,000[31] species of animals found exclusively in aquatic (mainly marine) environments. Their bodies consist of mesoglea, a non-living jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick. They have two basic body forms: swimming medusae and sessile polyps, both of which are radially symmetrical with mouths surrounded by tentacles that bear cnidocytes. Both forms have a single orifice and body cavity that are used for digestion and respiration.
Fossil cnidarians have been found in rocks formed about 580 million years ago. Fossils of cnidarians that do not build mineralized structures are rare. Scientists currently think cnidarians, ctenophores and bilaterians are more closely related to calcareous sponges than these are to other sponges, and that anthozoans are the evolutionary "aunts" or "sisters" of other cnidarians, and the most closely related to bilaterians.
Cnidarians are the simplest animals in which the cells are organised into tissues.[32] The starlet sea anemone is used as a model organism in research.[33] It is easy to care for in the laboratory and a protocol has been developed which can yield large numbers of embryos on a daily basis.[34] There is a remarkable degree of similarity in the gene sequence conservation and complexity between the sea anemone and vertebrates.[34] In particular, genes concerned in the formation of the head in vertebrates are also present in the anemone.[35][36]
Sea anemones are common in tidepools
Their tentacles sting and paralyse small fish
Close up of polyps on the surface of a coral, waving their tentacles.
As islands sink below the sea, corals growth can keep pace with the rising water, forming an atoll
Portuguese man o' war
Porpita porpita
Turritopsis dohrnii, a small, biologically immortal jellyfish transfers its cells back to childhood.[37][38]
Moon jellyfish, found in coastal waters around the world [39]
Lion's mane jellyfish, largest known jellyfish [40]
Marine worms[edit]
Further information: Marine worm and Sea worm
Arrow worms are predatory components of plankton worldwide.
Worms (Old English for serpent) typically have long cylindrical tube-like bodies and no limbs. Marine worms vary in size from microscopic to over 1 metre (3.3 ft) in length for some marine polychaete worms (bristle worms)[41] and up to 58 metres (190 ft) for the marine nemertean worm (bootlace worm).[42] Some marine worms occupy a small variety of parasitic niches, living inside the bodies of other animals, while others live more freely in the marine environment or by burrowing underground.
Different groups of marine worms are related only distantly, so they are found in several different phyla such as the Annelida (segmented worms), Chaetognatha (arrow worms), Hemichordata, and Phoronida (horseshoe worms). Many of these worms have specialized tentacles used for exchanging oxygen and carbon dioxide and also may be used for reproduction. Some marine worms are tube worms, such as the giant tube worm which lives in waters near underwater volcanoes and can withstand temperatures up to 90 degrees Celsius.
Platyhelminthes (flatworms) form another worm phylum which includes a class Cestoda of parasitic tapeworms. The marine tapeworm Polygonoporus giganticus, found in the gut of sperm whales, can grow to over 30 m (100 ft).[43][44]
Nematodes (roundworms) constitute a further worm phylum with tubular digestive systems and an opening at both ends.[45][46] Over 25,000 nematode species have been described,[47][48] of which more than half are parasitic. It has been estimated another million remain undescribed.[49] They are ubiquitous in marine, freshwater and terrestrial environments, where they often outnumber other animals in both individual and species counts. They are found in every part of the earth's lithosphere, from the top of mountains to the bottom of oceanic trenches.[50] By count they represent 90% of all animals on the ocean floor.[51] Their numerical dominance, often exceeding a million individuals per square meter and accounting for about 80% of all individual animals on earth, their diversity of life cycles, and their presence at various trophic levels point at an important role in many ecosystems.[52]
Giant tube worms cluster around hydrothermal vents
Lamellibrachia luymes, a cold seep tubeworm, lives over 250 years.
Nematodes are ubiquitous pseudocoelomates which can parasite marine plants and animals.
Bloodworms are typically found on the bottom of shallow marine waters
Bobbit worms are ambush predators that live on the seafloor
Echinoderms[edit]
Starfish larvae are bilaterally symmetric, whereas the adults have fivefold symmetry
Echinoderms (Greek for spiny skin) is a phylum which contains only marine invertebrates. The adults are recognizable by their radial symmetry (usually five-point) and include starfish, sea urchins, sand dollars, and sea cucumbers, as well as the sea lilies.[53] Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone. The phylum contains about 7000 living species,[54] making it the second-largest grouping of deuterostomes (a superphylum), after the chordates (which include the vertebrates, such as birds, fishes, mammals, and reptiles).
Echinoderms are unique among animals in having bilateral symmetry at the larval stage, but fivefold symmetry (pentamerism, a special type of radial symmetry) as adults.[55]
The echinoderms are important both biologically and geologically. Biologically, there are few other groupings so abundant in the biotic desert of the deep sea, as well as shallower oceans. Most echinoderms are able to regenerate tissue, organs, limbs, and reproduce asexually; in some cases, they can undergo complete regeneration from a single limb. Geologically, the value of echinoderms is in their ossified skeletons, which are major contributors to many limestone formations, and can provide valuable clues as to the geological environment. They were the most used species in regenerative research in the 19th and 20th centuries. Further, it is held by some scientists that the radiation of echinoderms was responsible for the Mesozoic Marine Revolution.
Echinoderm literally means "spiny skin", as this water melon sea urchin illustrates
Sea cucumbers filter feed on plankton and suspended solids
Benthopelagic sea cucumbers can lift off the seafloor and journey as much as 1,000 m (3,300 ft) up the water column
The ochre sea star was the first keystone predator to be studied. They limit mussels which can overwhelm intertidal communities.[56]
Colorful sea lilies in shallow waters
Aside from the hard-to-classify Arkarua (a Precambrian animal with echinoderm-like pentamerous radial symmetry), the first definitive members of the phylum appeared near the start of the Cambrian.
Marine molluscs[edit]
Reconstruction of an ammonite, a highly successful early cephalopod that first appeared in the Devonian (about 400 mya). They became extinct during the same extinction event that killed the land dinosaurs (about 66 mya).
Molluscs (Latin for soft) form a phylum with about 85,000 extant recognized species.[57] By species count they are the largest marine phylum, comprising about 23% of all the named marine organisms.[58] Molluscs have more varied forms than other invertebrate phylums. They are highly diverse, not just in size and in anatomical structure, but also in behaviour and in habitat. The majority of species still live in the oceans, from the seashores to the abyssal zone, but some form a significant part of the freshwater fauna and the terrestrial ecosystems.
The mollusc phylum is divided into 9 or 10 taxonomic classes, two of which are extinct. These classes include gastropods, bivalves and cephalopods, as well as other lesser-known but distinctive classes. Gastropods with protective shells are referred to as snails (sea snails), whereas gastropods without protective shells are referred to as slugs (sea slugs). Gastropods are by far the most numerous molluscs in terms of classified species, accounting for 80% of the total.[25] Bivalves include clams, oysters, cockles, mussels, scallops, and numerous other families. There are about 8,000 marine bivalves species (including brackish water and estuarine species), and about 1,200 freshwater species. Cephalopod include octopus, squid and cuttlefish. They are found in all oceans, and neurologically are the most advanced of the invertebrates.[59] About 800 living species of marine cephalopods have been identified,[60] and an estimated 11,000 extinct taxa have been described.[61] There are no fully freshwater cephalopods.[62]
Colossal squid, largest of all invertebrates [63]
The nautilus is a living fossil little changed since it evolved 500 million years ago as one of the first cephalopods.[64][65][66]
Marine gastropods are sea snails or sea slugs. This nudibranch is a sea slug.
The sea snail Syrinx aruanus has the largest shell of any living gastropod
Molluscs usually have eyes. Bordering the edge of the mantle of a scallop, a bivalve mollusc, can be over 100 simple eyes.
Common mussel, another bivalve
Generalized or hypothetical ancestral mollusc
Molluscs have such diverse shapes that many textbooks base their descriptions of molluscan anatomy on a generalized or hypothetical ancestral mollusc. This generalized mollusc is unsegmented and bilaterally symmetrical with an underside consisting of a single muscular foot.[67][68]:484–628 Beyond that it has three further key features. Firstly, it has a muscular cloak called a mantle covering its viscera and containing a significant cavity used for breathing and excretion. A shell secreted by the mantle covers the upper surface.[68] Secondly (apart from bivalves) it has a rasping tongue called a radula used for feeding. Thirdly, it has a nervous system including a complex digestive system using microscopic, muscle-powered hairs called cilia to exude mucus. The generalized mollusc has two paired nerve cords (three in bivalves). The brain, in species that have one, encircles the esophagus. Most molluscs have eyes and all have sensors detecting chemicals, vibrations, and touch. The simplest type of molluscan reproductive system relies on external fertilization, but more complex variations occur. All produce eggs, from which may emerge trochophore larvae, more complex veliger larvae, or miniature adults. The depiction is rather similar to modern monoplacophorans, and some suggest it may resemble very early molluscs.[67]:284–291[67]:298–300[69][70]
Good evidence exists for the appearance of marine gastropods, cephalopods and bivalves in the Cambrian period 541 to 485.4 million years ago. However, the evolutionary history both of molluscs' emergence from the ancestral Lophotrochozoa and of their diversification into the well-known living and fossil forms are still subjects of vigorous debate among scientists.
Marine arthropods[edit]
Head
_______________________
Thorax
_______________________
Abdomen
_______________________
Segmentation and tagmata of an arthropod[67]:518–522
Arthropods (Greek for jointed feet) have an exoskeleton (external skeleton), a segmented body, and jointed appendages (paired appendages). They form a phylum which includes insects, arachnids, myriapods, and crustaceans. Arthropods are characterized by their jointed limbs and cuticle made of chitin, often mineralised with calcium carbonate. The arthropod body plan consists of segments, each with a pair of appendages. The rigid cuticle inhibits growth, so arthropods replace it periodically by moulting. Their versatility has enabled them to become the most species-rich members of all ecological guilds in most environments.
Marine arthropods range in size from the microscopic crustacean Stygotantulus to the Japanese spider crab. Arthropods' primary internal cavity is a hemocoel, which accommodates their internal organs, and through which their haemolymph - analogue of blood - circulates; they have open circulatory systems. Like their exteriors, the internal organs of arthropods are generally built of repeated segments. Their nervous system is "ladder-like", with paired ventral nerve cords running through all segments and forming paired ganglia in each segment. Their heads are formed by fusion of varying numbers of segments, and their brains are formed by fusion of the ganglia of these segments and encircle the esophagus. The respiratory and excretory systems of arthropods vary, depending as much on their environment as on the subphylum to which they belong.
Their vision relies on various combinations of compound eyes and pigment-pit ocelli: in most species the ocelli can only detect the direction from which light is coming, and the compound eyes are the main source of information, but the main eyes of spiders are ocelli that can form images and, in a few cases, can swivel to track prey. Arthropods also have a wide range of chemical and mechanical sensors, mostly based on modifications of the many setae (bristles) that project through their cuticles. Arthropods' methods of reproduction and development are diverse; all terrestrial species use internal fertilization, but this is often by indirect transfer of the sperm via an appendage or the ground, rather than by direct injection. Marine species all lay eggs and use either internal or external fertilization. Arthropod hatchlings vary from miniature adults to grubs that lack jointed limbs and eventually undergo a total metamorphosis to produce the adult form.
Trilobites, now extinct, roamed oceans for 270 million years.[71]
Horseshoe crab, a living fossil arthropod from 450 million years ago
Crustaceans
Many crustaceans are very small, like this tiny amphipod, and make up a significant part of the ocean's zooplankton
The banded cleaner shrimp is a crustacean common in the tropics.
The Tasmanian giant crab is long-lived and slow-growing, making it vulnerable to overfishing.[72]
The Japanese spider crab has the longest leg span of any arthropod.
The evolutionary ancestry of arthropods dates back to the Cambrian period. The group is generally regarded as monophyletic, and many analyses support the placement of arthropods with cycloneuralians (or their constituent clades) in a superphylum Ecdysozoa. Overall however, the basal relationships of Metazoa are not yet well resolved. Likewise, the relationships between various arthropod groups are still actively debated.
Other phyla[edit]
Tardigrade, Lobopodia, (Onychophora)
Non-craniate (non-vertebrate) chordates: Cephalochordate, Tunicata and Haikouella. These invertebrates are close relatives of the vertebrates.
Non-craniate chordates are close relatives of vertebrates
The lancelet, a small translucent fish-like Cephalochordate, is the closest living invertebrate relative of the vertebrates.[73][74]
Fluorescent-colored sea squirts, Rhopalaea crassa. Tunicates may provide clues to vertebrate (and therefore human) ancestry.[75]
Salp chain
Gill slits in an acorn worm (left) and tunicate (right)
Minerals from sea water[edit]
There are a number of marine invertebrates that use minerals that are present in the sea in such minute quantities that they were undetectable until the advent of spectroscopy. Vanadium is concentrated by some tunicates for use in their blood cells to a level ten million times that of the surrounding seawater. Other tunicates similarly concentrate niobium and tantalum.[67]:947 Lobsters use copper in their respiratory pigment hemocyanin, despite the proportion of this metal in seawater being minute.[67]:638 Although these elements are present in vast quantities in the ocean, their extraction by man is not economic.[76]
See also[edit]
Wikimedia Commons has media related to Marine invertebrates.
Marine life
Marine biology
Marine vertebrate
List of marine aquarium invertebrate species
Wiwaxia
Notes[edit]
Jump up ^ Myxozoa were thought to be an exception, but are now thought to be heavily modified members of the Cnidaria. Jímenez-Guri, Eva; Philippe, Hervé; Okamura, Beth; Holland, Peter W. H. (July 6, 2007). "Buddenbrockia Is a Cnidarian Worm". Science. Washington, D.C.: American Association for the Advancement of Science. 317 (5834): 116–118. Bibcode:2007Sci...317..116J. doi:10.1126/science.1142024. ISSN 0036-8075. PMID 17615357. Retrieved 2008-09-03.
References[edit]
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Seven (1995 film)
From Wikipedia, the free encyclopedia
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"Seven (film)" redirects here. For the 1979 film, see Seven (1979 film).
Seven
Theatrical release poster
Directed by
David Fincher
Produced by
Arnold Kopelson
Phyllis Carlyle
Written by
Andrew Kevin Walker
Starring
Brad Pitt
Morgan Freeman
Gwyneth Paltrow
John C. McGinley
Music by
Howard Shore
Cinematography
Darius Khondji
Edited by
Richard Francis-Bruce
Distributed by
New Line Cinema
Release date
September 22, 1995 (1995-09-22)
Running time
127 minutes[1]
Country
United States
Language
English
Budget
$33 million[2]
Box office
$327.3 million[2]
Seven (sometimes stylized as SE7EN[3]) is a 1995 American neo-noir crime thriller film directed by David Fincher and starring Brad Pitt, Morgan Freeman, Gwyneth Paltrow, John C. McGinley, R. Lee Ermey, and Kevin Spacey. The film was based on a screenplay by Andrew Kevin Walker. It tells the story of David Mills (Pitt), a young detective who is partnered with the retiring William Somerset (Freeman) and soon tasked with tracking down a serial killer (Spacey) who uses the seven deadly sins as tropes in his murders.
The film's screenplay was influenced by the time Walker spent in New York City trying to make it as a writer. Principal photography took place in Los Angeles, with the last scene filmed near Lancaster, California. The film's budget was US$33 million.
Released on September 22, 1995 by New Line Cinema, Seven went on to become the seventh-highest-grossing film of the year, grossing over $327 million worldwide.[2] It was well received by critics, who praised the darkness and brutality of the film and its themes. The film was nominated for Best Film Editing at the 68th Academy Awards, but lost out to Apollo 13.
Contents
[hide]
1 Plot
2 Cast
3 Production
3.1 Pre-production
3.2 Filming
3.3 Title sequence
4 Reception
4.1 Box office
4.2 Critical response
4.3 Accolades
5 Home media
6 Novelization and comic books
7 Soundtrack
8 References
9 Further reading
10 External links
Plot[edit]
In an unnamed American city, soon-to-be-retiring detective William Somerset is partnered with short-tempered but idealistic David Mills, who recently transferred to the department, moving to the city with his wife Tracy. Mills introduces Somerset to Tracy, after which Somerset becomes her confidant. Tracy is unhappy with the city and feels it is no place to raise a child. She discloses to Somerset that she is pregnant and has yet to inform her husband. Somerset sympathizes with her, having a similar situation with his ex-girlfriend many years earlier, and advises her to tell Mills only if she plans on keeping the child.
Somerset and Mills investigate a pair of murders. The first victim is an obese man forced to eat until his stomach ruptured. The second was a wealthy defense attorney who died from both fatal bloodletting and the removal of a pound of flesh. At each crime scene, the murderer leaves behind clues for the detectives, including the word gluttony at the obese man's home and greed at the attorney's office. Somerset recognizes them as part of the seven deadly sins and realizes the murders are related. Other clues lead them to a possible perpetrator's apartment. There, they find another victim, a known drug dealer and child molester, strapped to a bed, barely alive and emaciated, with a series of pictures indicating he had been tied to the bed for an entire year. The word sloth is scrawled on the wall. The photos also indicate the killer has been planning these deaths for some time.
Somerset and Mills identify a man named John Doe, who has checked out several library books on the deadly sins. Doe flees when they go to his apartment, and Mills gives chase. Doe eventually corners Mills and holds him at gunpoint, but after a few moments, turns and escapes. At Doe's apartment, they find hundreds of handwritten journals showing Doe's apparent psychopathy, and clues leading to a fourth victim. They arrive too late to prevent the death of the victim, a prostitute killed by an unwilling man forced by Doe to wear a bladed S&M phallic device on his genitals and to rape and kill her while severely traumatizing him. They find lust written on the door. They are alerted to their next victim, an attractive young woman, presumably a model, whose face has been mutilated by Doe; she was given the option to call for help and be disfigured, or to commit suicide by taking pills. She chose suicide. The word pride is written on her wall.
Shortly after, as Somerset and Mills return to the police station, they are approached by a man covered in blood, surrendering himself. Mills recognizes him as Doe and arrests him. They discover Doe has been removing the skin on his fingers to avoid leaving behind prints; the blood on him is from a yet-to-be-identified victim. Doe, through his lawyer, advises there are two more victims and offers to take the detectives to them and confess to all the murders, but only under very specific terms, or he will otherwise plead insanity. Somerset is wary, but Mills agrees.
The two detectives, following Doe's directions, drive him to a remote desert location. Within minutes, a delivery van approaches them. Mills holds Doe at gunpoint while Somerset goes to intercept the driver, who had been instructed to bring a box to them. As Somerset recovers the box and sends the driver away, Doe begins telling Mills about how jealous he is of Mills' life and marriage to Tracy, antagonizing Mills. Somerset opens the box, and in horror, tells Mills to stay back and not listen to Doe. Doe continues to taunt Mills as Mills frantically asks what is in the box. Doe reveals that he was so jealous of Mills that he killed Tracy, her death being a result of his envy, and that her head is in the box. Doe tries to goad Mills into vengeance, to become wrath and shoot him. Somerset desperately tries to convince Mills not to shoot Doe, but then Doe reveals that Tracy was pregnant. The revelation is too much for Mills and he shoots Doe six times. Doe's death completes the seven sins. Police converge and take a devastated Mills away. The police captain reassures Somerset that Mills will be taken care of. When asked by the Captain where he will be, Somerset says he will "be around."
Cast[edit]
Brad Pitt as Detective David Mills
Morgan Freeman as Detective Lieutenant William Somerset
Gwyneth Paltrow as Tracy Mills
Kevin Spacey as John Doe
R. Lee Ermey as Police Captain
John C. McGinley as SWAT team leader California
Richard Roundtree as District Attorney Martin Talbot
Richard Schiff as Mark Swarr
Mark Boone Junior as Greasy FBI Man
Peter Crombie as Dr. O'Neill
Michael Massee as Man in Massage Parlour Booth
Leland Orser as Crazed Man in Massage Parlour
Andrew Kevin Walker as Dead Man
Production[edit]
Pre-production[edit]
The primary influence for the film's screenplay came from Andrew Kevin Walker's time spent in New York City while trying to make it as a screenwriter. "I didn't like my time in New York, but it's true that if I hadn't lived there I probably wouldn't have written Seven."[4] He envisioned actor William Hurt as Somerset and named the character after his favorite author, W. Somerset Maugham.[4]
Jeremiah S. Chechik was attached to direct at one point.[4] During pre-production, Al Pacino was considered for the Somerset role, but he decided to do City Hall. Denzel Washington and Sylvester Stallone turned down the role of Mills. Washington later regretted turning down the role.[5][6]
The ending of the screenplay, with the head in the box, was originally part of an earlier draft that New Line had rejected, instead opting for an ending that involved more traditional elements of a detective thriller film with more action-oriented elements. But when New Line sent David Fincher the screenplay to review for his interest in the project, they accidentally sent him the original screenplay with the head-in-the-box ending. At the time, Fincher had not read a script for a year and a half since the frustrating experience of making Alien 3; he said, "I thought I'd rather die of colon cancer than do another movie".[7] Fincher eventually agreed to direct Seven because he was drawn to the script, which he found to be a "connect-the-dots movie that delivers about inhumanity. It's psychologically violent. It implies so much, not about why you did but how you did it".[7] He found it more a "meditation on evil" rather than a "police procedural".
When New Line realized that they had sent Fincher the wrong draft, the President of Production, Michael De Luca, met with Fincher and noted that there was internal pressure to retain the revised version; De Luca stated that if Fincher promised to produce the movie, they would be able to stay with the head-in-a-box ending.[8] Despite this, producer Kopelson refused to allow the film to include the head-in-a-box scene.[9] Actor Pitt joined Fincher in arguing for keeping this original scene, noting that his previous film Legends of the Fall had its emotional ending cut after negative feedback from test audiences, and refusing to do Seven unless the head-in-the-box scene remained.[10]
Filming[edit]
Filming took place in Los Angeles, California.
Fincher approached making Seven like a "tiny genre movie, the kind of movie Friedkin might have made after The Exorcist." He worked with cinematographer Darius Khondji and adopted a simple approach to the camerawork, which was influenced by the television show COPS, "how the camera is in the backseat peering over people's shoulder".[7] Fincher allowed Walker on the set while filming for on-the-set rewrites.[4] According to the director, "Seven is the first time I got to carry through certain things about the camera – and about what movies are or can be".[7]
The crowded urban streets filled with noisy denizens and an oppressive rain that seems to fall without respite were integral parts of the film, as Fincher wanted to show a city that was "dirty, violent, polluted, often depressing. Visually and stylistically, that's how we wanted to portray this world. Everything needed to be as authentic and raw as possible." To this end, Fincher turned to production designer Arthur Max to create a dismal world that often eerily mirrors its inhabitants. "We created a setting that reflects the moral decay of the people in it", says Max. "Everything is falling apart, and nothing is working properly." The film's brooding, dark look was achieved through a chemical process called bleach bypass, wherein the silver in the film stock was not removed, which in turn deepened the dark, shadowy images in the film and increased its overall tonal quality.
The 'head in a box' ending continued to worry the studio after filming was completed. After the first cut of the film was shown to the studio, they attempted to mitigate the bleakness of the ending by replacing Mills' wife's head with that of a dog, or by not having Mills fire on John Doe. However, both Fincher and Pitt continued to fight for the original ending.[10] The final scenes of Mills being taken away and Somerset's quote from Ernest Hemingway were filmed by Fincher after initial filming was complete as a way to placate the studio (the original intention was for the film to suddenly end after Mills shot John Doe).[9]
Title sequence[edit]
On the film's title sequence, Fincher has said:
The sequence for Se7en did very important non-narrative things; in the original script there was a title sequence that had Morgan Freeman buying a house out in the middle of nowhere and then travelling back on a train. He was making his way back to the unnamed city from the unnamed suburban sprawl, and that's where the title was supposed to be—"insert title sequence here"—but we didn't have the money to do that. We also lacked the feeling of John Doe, the villain, who just appeared 90 minutes into the movie. It was oddly problematic, you just needed a sense of what these guys were up against. Kyle Cooper, the designer of the title sequence, came to me and said, "You know, you have these amazing books that you spent tens of thousands of dollars to make for the John Doe interior props. I'd like to see them featured." And I said, "Well, that would be neat, but that's kind of a 2D glimpse. Figure out a way for it to involve John Doe, to show that somewhere across town somebody is working on some really evil shit. I don't want it to be just flipping through pages, as beautiful as they are." So Kyle came up with a great storyboard, and then we got Angus Wall and Harris Savides—Harris to shoot it and Angus to cut it—and the rest, as they say, is internet history.[11]
Reception[edit]
Box office[edit]
Seven was released on September 22, 1995, in 2,441 theaters where it grossed US$13.9 million on its opening weekend. It went on to gross $100.1 million in North America and $227.1 million in the rest of the world for a total of $327.3 million,[12] making Seven the seventh-highest-grossing film in 1995.[13] The film also spent 4 consecutive weeks in the top spot at the U.S. box office in 1995.
Critical response[edit]
The film was well received by critics and holds an 80% positive rating at the film-review aggregator Rotten Tomatoes, based on 70 critics with an average rating of 7.7 out of 10. Its consensus reads: "A brutal, relentlessly grimy shocker with taut performances, slick gore effects, and a haunting finale."[3] The film has a rating of 65 on Metacritic based on 22 reviews.[14]
Gary Arnold, in The Washington Times, praised the cast: "The film's ace in the hole is the personal appeal generated by Mr. Freeman as the mature, cerebral cop and Mr. Pitt as the young, headstrong cop. Not that the contrast is inspired or believable in itself. What gets to you is the prowess of the co-stars as they fill out sketchy character profiles."[15] Sheila Johnston, in her review for The Independent, praised Freeman's performance: "The film belongs to Freeman and his quiet, carefully detailed portrayal of the jaded older man who learns not to give up the fight."[16] James Charisma, in a list of Spacey's greatest film performances for Paste, wrote: "Spacey’s portrayal is a perfect balancing act: John Doe is detached from the murders he commits, yet deliberate and meticulous in his execution ... Unemotional yet smug. Analytical, violent, patient, impenetrable."[17] In his review for Sight and Sound, John Wrathall wrote, "Seven has the scariest ending since George Sluizer's original The Vanishing ... and stands as the most complex and disturbing entry in the serial killer genre since Manhunter."[18] In his "Great Movies" list review, film critic Roger Ebert commented on Fincher's direction: "None of his films is darker than this one."[19]
Accolades[edit]
New Line Cinema re-released Seven in Westwood, Los Angeles, California, on Christmas Day and in New York City on December 29, 1995, in an attempt to generate Academy Award nominations for Freeman, Pitt, and Fincher, which was ultimately unsuccessful.[20]
American Film Institute lists
AFI's 100 Years...100 Thrills - Nominated
AFI's 100 Years...100 Heroes and Villains
John Doe - Nominated Villain
AFI's 10 Top 10 - Nominated Mystery Film
Year
Ceremony
Category
Recipients
Result
1995 68th Academy Awards Best Film Editing Richard Francis-Bruce Nominated
49th British Academy Film Awards Best Screenplay - Original Andrew Kevin Walker Nominated
1996 MTV Movie Awards Best Movie Seven Won
Most Desirable Male Brad Pitt Won
Best On-Screen Duo Brad Pitt and Morgan Freeman Nominated
Best Villain Kevin Spacey Won
Home media[edit]
For the DVD release, Seven was remastered and presented in the widescreen format, preserving the 2.40:1 aspect ratio of its original theatrical exhibition. Audio options include Dolby Digital EX 5.1, DTS ES Discrete 6.1, and Stereo Surround Sound.
The Seven DVD features four newly recorded, feature-length audio commentaries featuring the stars and other key contributors to the film, who talk about their experiences making Seven.
This DVD is also compatible with DVD-ROM drives. Disc One features a printable screenplay with links to the film. The Blu-ray Disc was released September 14, 2010.[21]
Novelization and comic books[edit]
In 1995, a novelization with the same title was written by Anthony Bruno based on the original film.[22]
Between September 2006 and October 2007, a series of seven comic books were published by Zenescope Entertainment with each of the seven issues dedicated to one of the seven sins. It told the story from the perspective of John Doe rather than the two homicide detectives as in the film. Each issue included contributions by a group of creators independent of each other. All seven issues were collected in trade paperback form, released on January 15, 2008, as SE7EN , edited by David Seidman and Ralph Tedesco.[23][24]
Soundtrack[edit]
The opening credit music is a spliced sample of an uncredited remix of the Nine Inch Nails song "Closer", available as "Closer (Precursor)", remixed by Coil, on the "Closer" single. The song during the end credits is David Bowie's song "The Hearts Filthy Lesson", found on his album Outside. The film's original score is by Howard Shore.
"In the Beginning" – The Statler Brothers
"Guilty" – Gravity Kills
"Trouble Man" – Marvin Gaye
"Speaking of Happiness" – Gloria Lynne – written by Buddy Scott & Jimmy Radcliffe
"Suite No. 3 in D Major, BWV 1068 Air" – written by Johann Sebastian Bach, performed by Stuttgarter Kammerorchester / Karl Münchinger
"Love Plus One" – Haircut One Hundred
"I Cover the Waterfront" – Billie Holiday
"Now's the Time" – Charlie Parker
"Straight, No Chaser" – Thelonious Monk (Taken from Monk in Tokyo)
"Portrait of John Doe" – Howard Shore
"Suite from Seven" – Howard Shore
References[edit]
Jump up ^ "Se7en (18)". British Board of Film Classification. September 27, 1995. Retrieved July 28, 2013.
^ Jump up to: a b c "Seven (1995)". Box Office Mojo. Retrieved July 17, 2014.
^ Jump up to: a b "Seven". Rotten Tomatoes.
^ Jump up to: a b c d Montesano, Anthony (February 1996). "Seven's Deadly Sins". Cinefantastique. p. 48.
Jump up ^ Doty, Meriah (September 18, 2012). "Denzel Washington regrets passing up ‘Seven’ and ‘Michael Clayton’". Yahoo! Movies. Retrieved May 13, 2017.
Jump up ^ Davis, Edward (September 26, 2012). "Denzel Washington Turned Down ‘Seven’ & ‘Michael Clayton,’ Javier Bardem Passed On ‘Minority Report’". IndieWire. Retrieved May 13, 2017.
^ Jump up to: a b c d Taubin, Amy (January 1996). "The Allure of Decay". Sight and Sound. p. 24.
Jump up ^ Salsibury, Mark (January 18, 2009). "David Fincher". The Guardian. Retrieved June 26, 2012.
^ Jump up to: a b Mottham, James (2007). The Sundance Kids: How the Mavericks Took Back Hollywood. Faber and Faber. pp. 153–155. ISBN 0865479674.
^ Jump up to: a b Smith, Grady (September 16, 2011). "How Brad Pitt fought to keep Gwyneth's head in the box in 'Se7en'". Entertainment Weekly. Retrieved June 26, 2012.
Jump up ^ Perkins, Will (August 27, 2012). "David Fincher: A Film Title Retrospective". Art of the Title. Retrieved September 8, 2012.
Jump up ^ "Seven". Box Office Mojo. Retrieved March 26, 2008.
Jump up ^ The top six grossing films of 1995 were Die Hard with a Vengeance, Toy Story, Apollo 13, GoldenEye, Pocahontas and Batman Forever.
Jump up ^ "Seven". Metacritic.
Jump up ^ Arnold, Gary (September 22, 1995). "Sinister Seven a killer of a thriller". The Washington Times.
Jump up ^ Johnston, Sheila (January 4, 1996). "Sin has seldom looked so good". The Independent.
Jump up ^ Charisma, James (August 15, 2016). "All 45 of Kevin Spacey’s Movie Performances, Ranked". Paste. Retrieved January 16, 2017.
Jump up ^ Wrathall, John (January 1996). "Seven". Sight and Sound. p. 50.
Jump up ^ Ebert, Roger (July 18, 2011). "Seven (1995)". Chicago Sun-Times.
Jump up ^ Cox, Dan (December 22, 1995). "Seven gets new dates for Oscar season". Variety.
Jump up ^ Creepy, Uncle. "First Blu-ray News: Seven". dreadcentral.com. Retrieved November 4, 2011.
Jump up ^ Bruno, Anthony (1995). Seven: a novel by Anthony Bruno based on a screenplay by Andrew Kevin Walker. New York: St. Martin's Paperbacks. p. 248. ISBN 0-312-95704-1.
Jump up ^ Comic Book Resources.com Horrific sins: SE7EN" comes to comics this September
Jump up ^ MyComicShop: Seven (2006 Se7en) comic books
Further reading[edit]
Dyer, Richard (1999). Seven. London: British Film Institute. ISBN 9780851707235.
External links[edit]
Wikiquote has quotations related to: Se7en
Seven on IMDb
Seven at the TCM Movie Database
Seven at AllMovie
Seven at Box Office Mojo
Seven at Rotten Tomatoes
Seven at Metacritic
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Alien 3 (1992)
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Seven deadly sins
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Seven (1995 film)
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