title: Sponge

text: Although most of the approximately 5,000–10,000 known species of sponges feed on bacteria and other microscopic food in the water, some host photosynthesizing microorganisms as endosymbionts, and these alliances often produce more food and oxygen than they consume. A few species of sponges that live in food-poor environments have evolved as carnivores that prey mainly on small crustaceans.


title: Sponge

text: Most species use sexual reproduction, releasing sperm cells into the water to fertilize ova that in some species are released and in others are retained by the "mother". The fertilized eggs develop into larvae, which swim off in search of places to settle. Sponges are known for regenerating from fragments that are broken off, although this only works if the fragments include the right types of cells. A few species reproduce by budding. When environmental conditions become less hospitable to the sponges, for example as temperatures drop, many freshwater species and a few marine ones produce gemmules, "survival pods" of unspecialized cells that remain dormant until conditions improve; they then either form completely new sponges or recolonize the skeletons of their parents.


title: Sponge

text: In most sponges, an internal gelatinous matrix called mesohyl functions as an endoskeleton, and it is the only skeleton in soft sponges that encrust such hard surfaces as rocks. More commonly, the mesohyl is stiffened by mineral spicules, by spongin fibers, or both. Demosponges use spongin; many species have silica spicules, whereas some species have calcium carbonate exoskeletons. Demosponges constitute about 90% of all known sponge species, including all freshwater ones, and they have the widest range of habitats. Calcareous sponges, which have calcium carbonate spicules and, in some species, calcium carbonate exoskeletons, are restricted to relatively shallow marine waters where production of calcium carbonate is easiest. The fragile glass sponges, with "scaffolding" of silica spicules, are restricted to polar regions and the ocean depths where predators are rare. Fossils of all of these types have been found in rocks dated from . In addition Archaeocyathids, whose fossils are common in rocks from , are now regarded as a type of sponge.


title: Sponge

text: The single-celled choanoflagellates resemble the choanocyte cells of sponges which are used to drive their water flow systems and capture most of their food. This along with phylogenetic studies of ribosomal molecules have been used as morphological evidence to suggest sponges are the sister group to the rest of animals. Some studies have shown that sponges do not form a monophyletic group, in other words do not include "all and only" the descendants of a common ancestor. Recent phylogenetic analyses suggested that comb jellies rather than sponges are the sister group to the rest of animals. However reanalysis of the data showed that the computer algorithms used for analysis were misled by the presence of specific ctenophore genes that were markedly different from those of other species, leaving sponges as either the sister group to all other animals, or an ancestral paraphyletic grade.


title: Sponge

text: The few species of demosponge that have entirely soft fibrous skeletons with no hard elements have been used by humans over thousands of years for several purposes, including as padding and as cleaning tools. By the 1950s, though, these had been overfished so heavily that the industry almost collapsed, and most sponge-like materials are now synthetic. Sponges and their microscopic endosymbionts are now being researched as possible sources of medicines for treating a wide range of diseases. Dolphins have been observed using sponges as tools while foraging.


title: Sponge

text: Sponges constitute the phylum Porifera, and have been defined as sessile metazoans (multicelled immobile animals) that have water intake and outlet openings connected by chambers lined with choanocytes, cells with whip-like flagella. However, a few carnivorous sponges have lost these water flow systems and the choanocytes. All known living sponges can remold their bodies, as most types of their cells can move within their bodies and a few can change from one type to another.


title: Sponge

text: Even if a few sponges are able to produce mucus – which acts as a microbial barrier in all other animals – no sponge with the ability to secrete a functional mucus layer has been recorded. Without such a mucus layer their living tissue is covered by a layer of microbial symbionts, which can contribute up to 40–50% of the sponge wet mass. This inability to prevent microbes from penetrating their porous tissue could be a major reason why they have never evolved a more complex anatomy.


title: Sponge

text: Like cnidarians (jellyfish, etc.) and ctenophores (comb jellies), and unlike all other known metazoans, sponges' bodies consist of a non-living jelly-like mass (mesohyl) sandwiched between two main layers of cells. Cnidarians and ctenophores have simple nervous systems, and their cell layers are bound by internal connections and by being mounted on a basement membrane (thin fibrous mat, also known as "basal lamina"). Sponges have no nervous systems, their middle jelly-like layers have large and varied populations of cells, and some types of cells in their outer layers may move into the middle layer and change their functions.


title: Sponge

text: A sponge's body is hollow and is held in shape by the mesohyl, a jelly-like substance made mainly of collagen and reinforced by a dense network of fibers also made of collagen. The inner surface is covered with choanocytes, cells with cylindrical or conical collars surrounding one flagellum per choanocyte. The wave-like motion of the whip-like flagella drives water through the sponge's body. All sponges have ostia, channels leading to the interior through the mesohyl, and in most sponges these are controlled by tube-like porocytes that form closable inlet valves. Pinacocytes, plate-like cells, form a single-layered external skin over all other parts of the mesohyl that are not covered by choanocytes, and the pinacocytes also digest food particles that are too large to enter the ostia, while those at the base of the animal are responsible for anchoring it.


title: Sponge

text: Many larval sponges possess neuron-less eyes that are based on cryptochromes. They mediate phototaxic behavior.


title: Sponge

text: Glass sponges present a distinctive variation on this basic plan. Their spicules, which are made of silica, form a scaffolding-like framework between whose rods the living tissue is suspended like a cobweb that contains most of the cell types. This tissue is a syncytium that in some ways behaves like many cells that share a single external membrane, and in others like a single cell with multiple nuclei. The mesohyl is absent or minimal. The syncytium's cytoplasm, the soupy fluid that fills the interiors of cells, is organized into "rivers" that transport nuclei, organelles ("organs" within cells) and other substances. Instead of choanocytes, they have further syncytia, known as choanosyncytia, which form bell-shaped chambers where water enters via perforations. The insides of these chambers are lined with "collar bodies", each consisting of a collar and flagellum but without a nucleus of its own. The motion of the flagella sucks water through passages in the "cobweb" and expels it via the open ends of the bell-shaped chambers.


title: Sponge

text: Some types of cells have a single nucleus and membrane each, but are connected to other single-nucleus cells and to the main syncytium by "bridges" made of cytoplasm. The sclerocytes that build spicules have multiple nuclei, and in glass sponge larvae they are connected to other tissues by cytoplasm bridges; such connections between sclerocytes have not so far been found in adults, but this may simply reflect the difficulty of investigating such small-scale features. The bridges are controlled by "plugged junctions" that apparently permit some substances to pass while blocking others.


title: Sponge

text: Most sponges work rather like chimneys: they take in water at the bottom and eject it from the osculum ("little mouth") at the top. Since ambient currents are faster at the top, the suction effect that they produce by Bernoulli's principle does some of the work for free. Sponges can control the water flow by various combinations of wholly or partially closing the osculum and ostia (the intake pores) and varying the beat of the flagella, and may shut it down if there is a lot of sand or silt in the water.


title: Sponge

text: Although the layers of pinacocytes and choanocytes resemble the epithelia of more complex animals, they are not bound tightly by cell-to-cell connections or a basal lamina (thin fibrous sheet underneath). The flexibility of these layers and re-modeling of the mesohyl by lophocytes allow the animals to adjust their shapes throughout their lives to take maximum advantage of local water currents.


title: Sponge

text: The simplest body structure in sponges is a tube or vase shape known as "asconoid", but this severely limits the size of the animal. The body structure is characterized by a stalk-like spongocoel surrounded by a single layer of choanocytes. If it is simply scaled up, the ratio of its volume to surface area increases, because surface increases as the square of length or width while volume increases proportionally to the cube. The amount of tissue that needs food and oxygen is determined by the volume, but the pumping capacity that supplies food and oxygen depends on the area covered by choanocytes. Asconoid sponges seldom exceed in diameter.


title: Sponge

text: Some sponges overcome this limitation by adopting the "syconoid" structure, in which the body wall is pleated. The inner pockets of the pleats are lined with choanocytes, which connect to the outer pockets of the pleats by ostia. This increase in the number of choanocytes and hence in pumping capacity enables syconoid sponges to grow up to a few centimeters in diameter.


title: Sponge

text: The "leuconoid" pattern boosts pumping capacity further by filling the interior almost completely with mesohyl that contains a network of chambers lined with choanocytes and connected to each other and to the water intakes and outlet by tubes. Leuconid sponges grow to over in diameter, and the fact that growth in any direction increases the number of choanocyte chambers enables them to take a wider range of forms, for example "encrusting" sponges whose shapes follow those of the surfaces to which they attach. All freshwater and most shallow-water marine sponges have leuconid bodies. The networks of water passages in glass sponges are similar to the leuconid structure.


title: Sponge

text: In all three types of structure the cross-section area of the choanocyte-lined regions is much greater than that of the intake and outlet channels. This makes the flow slower near the choanocytes and thus makes it easier for them to trap food particles. For example, in "Leuconia", a small leuconoid sponge about tall and in diameter, water enters each of more than 80,000 intake canals at 6 cm per "minute". However, because "Leuconia" has more than 2 million flagellated chambers whose combined diameter is much greater than that of the canals, water flow through chambers slows to 3.6 cm per "hour", making it easy for choanocytes to capture food. All the water is expelled through a single osculum at about 8.5 cm per "second", fast enough to carry waste products some distance away.


title: Sponge

text: In zoology a skeleton is any fairly rigid structure of an animal, irrespective of whether it has joints and irrespective of whether it is biomineralized. The mesohyl functions as an endoskeleton in most sponges, and is the only skeleton in soft sponges that encrust hard surfaces such as rocks. More commonly the mesohyl is stiffened by mineral spicules, by spongin fibers or both. Spicules, which are present in most but not all species, may be made of silica or calcium carbonate, and vary in shape from simple rods to three-dimensional "stars" with up to six rays. Spicules are produced by sclerocyte cells, and may be separate, connected by joints, or fused.


title: Sponge

text: Some sponges also secrete exoskeletons that lie completely outside their organic components. For example, sclerosponges ("hard sponges") have massive calcium carbonate exoskeletons over which the organic matter forms a thin layer with choanocyte chambers in pits in the mineral. These exoskeletons are secreted by the pinacocytes that form the animals' skins.


title: Sponge

text: Although adult sponges are fundamentally sessile animals, some marine and freshwater species can move across the sea bed at speeds of per day, as a result of amoeba-like movements of pinacocytes and other cells. A few species can contract their whole bodies, and many can close their oscula and ostia. Juveniles drift or swim freely, while adults are stationary.


title: Sponge

text: Sponges do not have distinct circulatory, respiratory, digestive, and excretory systems – instead the water flow system supports all these functions. They filter food particles out of the water flowing through them. Particles larger than 50 micrometers cannot enter the ostia and pinacocytes consume them by phagocytosis (engulfing and intracellular digestion). Particles from 0.5 μm to 50 μm are trapped in the ostia, which taper from the outer to inner ends. These particles are consumed by pinacocytes or by archaeocytes which partially extrude themselves through the walls of the ostia. Bacteria-sized particles, below 0.5 micrometers, pass through the ostia and are caught and consumed by choanocytes. Since the smallest particles are by far the most common, choanocytes typically capture 80% of a sponge's food supply. Archaeocytes transport food packaged in vesicles from cells that directly digest food to those that do not. At least one species of sponge has internal fibers that function as tracks for use by nutrient-carrying archaeocytes, and these tracks also move inert objects.


title: Sponge

text: It used to be claimed that glass sponges could live on nutrients dissolved in sea water and were very averse to silt. However, a study in 2007 found no evidence of this and concluded that they extract bacteria and other micro-organisms from water very efficiently (about 79%) and process suspended sediment grains to extract such prey. Collar bodies digest food and distribute it wrapped in vesicles that are transported by dynein "motor" molecules along bundles of microtubules that run throughout the syncytium.


title: Sponge

text: Sponges' cells absorb oxygen by diffusion from water into cells as water flows through body, into which carbon dioxide and other soluble waste products such as ammonia also diffuse. Archeocytes remove mineral particles that threaten to block the ostia, transport them through the mesohyl and generally dump them into the outgoing water current, although some species incorporate them into their skeletons.


title: Sponge

text: In waters where the supply of food particles is very poor, some species prey on crustaceans and other small animals. So far only 137 species have been discovered. Most belong to the family Cladorhizidae, but a few members of the Guitarridae and Esperiopsidae are also carnivores. In most cases little is known about how they actually capture prey, although some species are thought to use either sticky threads or hooked spicules. Most carnivorous sponges live in deep waters, up to , and the development of deep-ocean exploration techniques is expected to lead to the discovery of several more. However, one species has been found in Mediterranean caves at depths of , alongside the more usual filter feeding sponges. The cave-dwelling predators capture crustaceans under long by entangling them with fine threads, digest them by enveloping them with further threads over the course of a few days, and then return to their normal shape; there is no evidence that they use venom.


title: Sponge

text: Most known carnivorous sponges have completely lost the water flow system and choanocytes. However, the genus "Chondrocladia" uses a highly modified water flow system to inflate balloon-like structures that are used for capturing prey.


title: Sponge

text: Freshwater sponges often host green algae as endosymbionts within archaeocytes and other cells, and benefit from nutrients produced by the algae. Many marine species host other photosynthesizing organisms, most commonly cyanobacteria but in some cases dinoflagellates. Symbiotic cyanobacteria may form a third of the total mass of living tissue in some sponges, and some sponges gain 48% to 80% of their energy supply from these micro-organisms. In 2008 a University of Stuttgart team reported that spicules made of silica conduct light into the mesohyl, where the photosynthesizing endosymbionts live. Sponges that host photosynthesizing organisms are most common in waters with relatively poor supplies of food particles, and often have leafy shapes that maximize the amount of sunlight they collect.


title: Sponge

text: A recently discovered carnivorous sponge that lives near hydrothermal vents hosts methane-eating bacteria, and digests some of them.


title: Sponge

text: Sponges do not have the complex immune systems of most other animals. However, they reject grafts from other species but accept them from other members of their own species. In a few marine species, gray cells play the leading role in rejection of foreign material. When invaded, they produce a chemical that stops movement of other cells in the affected area, thus preventing the intruder from using the sponge's internal transport systems. If the intrusion persists, the grey cells concentrate in the area and release toxins that kill all cells in the area. The "immune" system can stay in this activated state for up to three weeks.


title: Sponge

text: Sponges have three asexual methods of reproduction: after fragmentation, by budding, and by producing gemmules. Fragments of sponges may be detached by currents or waves. They use the mobility of their pinacocytes and choanocytes and reshaping of the mesohyl to re-attach themselves to a suitable surface and then rebuild themselves as small but functional sponges over the course of several days. The same capabilities enable sponges that have been squeezed through a fine cloth to regenerate. A sponge fragment can only regenerate if it contains both collencytes to produce mesohyl and archeocytes to produce all the other cell types. A very few species reproduce by budding.


title: Sponge

text: Gemmules are "survival pods" which a few marine sponges and many freshwater species produce by the thousands when dying and which some, mainly freshwater species, regularly produce in autumn. Spongocytes make gemmules by wrapping shells of spongin, often reinforced with spicules, round clusters of archeocytes that are full of nutrients. Freshwater gemmules may also include photosynthesizing symbionts. The gemmules then become dormant, and in this state can survive cold, drying out, lack of oxygen and extreme variations in salinity. Freshwater gemmules often do not revive until the temperature drops, stays cold for a few months and then reaches a near-"normal" level. When a gemmule germinates, the archeocytes round the outside of the cluster transform into pinacocytes, a membrane over a pore in the shell bursts, the cluster of cells slowly emerges, and most of the remaining archeocytes transform into other cell types needed to make a functioning sponge. Gemmules from the same species but different individuals can join forces to form one sponge. Some gemmules are retained within the parent sponge, and in spring it can be difficult to tell whether an old sponge has revived or been "recolonized" by its own gemmules.


title: Sponge

text: Most sponges are hermaphrodites (function as both sexes simultaneously), although sponges have no gonads (reproductive organs). Sperm are produced by choanocytes or entire choanocyte chambers that sink into the mesohyl and form spermatic cysts while eggs are formed by transformation of archeocytes, or of choanocytes in some species. Each egg generally acquires a yolk by consuming "nurse cells". During spawning, sperm burst out of their cysts and are expelled via the osculum. If they contact another sponge of the same species, the water flow carries them to choanocytes that engulf them but, instead of digesting them, metamorphose to an ameboid form and carry the sperm through the mesohyl to eggs, which in most cases engulf the carrier and its cargo.


title: Sponge

text: A few species release fertilized eggs into the water, but most retain the eggs until they hatch. There are four types of larvae, but all are balls of cells with an outer layer of cells whose flagellae or cilia enable the larvae to move. After swimming for a few days the larvae sink and crawl until they find a place to settle. Most of the cells transform into archeocytes and then into the types appropriate for their locations in a miniature adult sponge.


title: Sponge

text: Glass sponge embryos start by dividing into separate cells, but once 32 cells have formed they rapidly transform into larvae that externally are ovoid with a band of cilia round the middle that they use for movement, but internally have the typical glass sponge structure of spicules with a cobweb-like main syncitium draped around and between them and choanosyncytia with multiple collar bodies in the center. The larvae then leave their parents' bodies.


title: Sponge

text: Sponges in temperate regions live for at most a few years, but some tropical species and perhaps some deep-ocean ones may live for 200 years or more. Some calcified demosponges grow by only per year and, if that rate is constant, specimens wide must be about 5,000 years old. Some sponges start sexual reproduction when only a few weeks old, while others wait until they are several years old.


title: Sponge

text: Adult sponges lack neurons or any other kind of nervous tissue. However, most species have the ability to perform movements that are coordinated all over their bodies, mainly contractions of the pinacocytes, squeezing the water channels and thus expelling excess sediment and other substances that may cause blockages. Some species can contract the osculum independently of the rest of the body. Sponges may also contract in order to reduce the area that is vulnerable to attack by predators. In cases where two sponges are fused, for example if there is a large but still unseparated bud, these contraction waves slowly become coordinated in both of the "Siamese twins". The coordinating mechanism is unknown, but may involve chemicals similar to neurotransmitters. However, glass sponges rapidly transmit electrical impulses through all parts of the syncytium, and use this to halt the motion of their flagella if the incoming water contains toxins or excessive sediment. Myocytes are thought to be responsible for closing the osculum and for transmitting signals between different parts of the body.


title: Sponge

text: Sponges contain genes very similar to those that contain the "recipe" for the post-synaptic density, an important signal-receiving structure in the neurons of all other animals. However, in sponges these genes are only activated in "flask cells" that appear only in larvae and may provide some sensory capability while the larvae are swimming. This raises questions about whether flask cells represent the predecessors of true neurons or are evidence that sponges' ancestors had true neurons but lost them as they adapted to a sessile lifestyle.


title: Sponge

text: Sponges are worldwide in their distribution, living in a wide range of ocean habitats, from the polar regions to the tropics. Most live in quiet, clear waters, because sediment stirred up by waves or currents would block their pores, making it difficult for them to feed and breathe. The greatest numbers of sponges are usually found on firm surfaces such as rocks, but some sponges can attach themselves to soft sediment by means of a root-like base.


title: Sponge

text: Sponges are more abundant but less diverse in temperate waters than in tropical waters, possibly because organisms that prey on sponges are more abundant in tropical waters. Glass sponges are the most common in polar waters and in the depths of temperate and tropical seas, as their very porous construction enables them to extract food from these resource-poor waters with the minimum of effort. Demosponges and calcareous sponges are abundant and diverse in shallower non-polar waters.


title: Sponge

text: Sponges with photosynthesizing endosymbionts produce up to three times more oxygen than they consume, as well as more organic matter than they consume. Such contributions to their habitats' resources are significant along Australia's Great Barrier Reef but relatively minor in the Caribbean.


title: Sponge

text: Many sponges shed spicules, forming a dense carpet several meters deep that keeps away echinoderms which would otherwise prey on the sponges. They also produce toxins that prevent other sessile organisms such as bryozoans or sea squirts from growing on or near them, making sponges very effective competitors for living space. One of many examples includes ageliferin.


title: Sponge

text: A few species, the Caribbean fire sponge "Tedania ignis", cause a severe rash in humans who handle them. Turtles and some fish feed mainly on sponges. It is often said that sponges produce chemical defenses against such predators. However, experiments have been unable to establish a relationship between the toxicity of chemicals produced by sponges and how they taste to fish, which would diminish the usefulness of chemical defenses as deterrents. Predation by fish may even help to spread sponges by detaching fragments. However, some studies have shown fish showing a preference for non chemically defended sponges, and another study found that high levels of coral predation did predict the presence of chemically defended species.


title: Sponge

text: Sponge flies, also known as spongilla-flies (Neuroptera, Sisyridae), are specialist predators of freshwater sponges. The female lays her eggs on vegetation overhanging water. The larvae hatch and drop into the water where they seek out sponges to feed on. They use their elongated mouthparts to pierce the sponge and suck the fluids within. The larvae of some species cling to the surface of the sponge while others take refuge in the sponge's internal cavities. The fully grown larvae leave the water and spin a cocoon in which to pupate.


title: Sponge

text: The Caribbean chicken-liver sponge "Chondrilla nucula" secretes toxins that kill coral polyps, allowing the sponges to grow over the coral skeletons. Others, especially in the family Clionaidae, use corrosive substances secreted by their archeocytes to tunnel into rocks, corals and the shells of dead mollusks. Sponges may remove up to per year from reefs, creating visible notches just below low-tide level.


title: Sponge

text: Caribbean sponges of the genus "Aplysina" suffer from Aplysina red band syndrome. This causes "Aplysina" to develop one or more rust-colored bands, sometimes with adjacent bands of necrotic tissue. These lesions may completely encircle branches of the sponge. The disease appears to be contagious and impacts approximately 10 percent of "A. cauliformis" on Bahamian reefs. The rust-colored bands are caused by a cyanobacterium, but it is unknown whether this organism actually causes the disease.


title: Sponge

text: In addition to hosting photosynthesizing endosymbionts, sponges are noted for their wide range of collaborations with other organisms. The relatively large encrusting sponge "Lissodendoryx colombiensis" is most common on rocky surfaces, but has extended its range into seagrass meadows by letting itself be surrounded or overgrown by seagrass sponges, which are distasteful to the local starfish and therefore protect "Lissodendoryx" against them; in return the seagrass sponges get higher positions away from the sea-floor sediment.


title: Sponge

text: Shrimps of the genus "Synalpheus" form colonies in sponges, and each shrimp species inhabits a different sponge species, making "Synalpheus" one of the most diverse crustacean genera. Specifically, Synalpheus regalis utilizes the sponge not only as a food source, but also as a defense against other shrimp and predators. As many as 16,000 individuals inhabit a single loggerhead sponge, feeding off the larger particles that collect on the sponge as it filters the ocean to feed itself. Other crustaceans such as hermit crabs commonly have a specific species of sponge, "Pseudospongosorites", grow on them as both the sponge and crab occupy gastropod shells until the crab and sponge outgrow the shell, eventually resulting in the crab using the sponge's body as protection instead of the shell until the crab finds a suitable replacement shell.


title: Sponge

text: Most sponges are detritivores which filter organic debris particles and microscopic life forms from ocean water. In particular, sponges occupy an important role as detritivores in coral reef food webs by recycling detritus to higher trophic levels.


title: Sponge

text: The hypothesis has been made that coral reef sponges facilitate the transfer of coral-derived organic matter to their associated detritivores via the production of sponge detritus, as shown in the diagram. Several sponge species are able to convert coral-derived DOM into sponge detritus, and transfer organic matter produced by corals further up the reef food web. Corals release organic matter as both dissolved and particulate mucus, as well as cellular material such as expelled "Symbiodinium".


title: Sponge

text: Organic matter could be transferred from corals to sponges by all these pathways, but DOM likely makes up the largest fraction, as the majority (56 to 80%) of coral mucus dissolves in the water column, and coral loss of fixed carbon due to expulsion of "Symbiodinium" is typically negligible (0.01%) compared with mucus release (up to ~40%). Coral-derived organic matter could also be indirectly transferred to sponges via bacteria, which can also consume coral mucus.


title: Sponge

text: Besides a one to one symbiotic relationship, it is possible for a host to become symbiotic with a microbial consortium. Sponges are able to host a wide range of microbial communities that can also be very specific. The microbial communities that form a symbiotic relationship with the sponge can amount to as much as 35% of the biomass of its host. The term for this specific symbiotic relationship, where a microbial consortia pairs with a host is called a holobiotic relationship. The sponge as well as the microbial community associated with it will produce a large range of secondary metabolites that help protect it against predators through mechanisms such as chemical defense.


title: Sponge

text: Some of these relationships include endosymbionts within bacteriocyte cells, and cyanobacteria or microalgae found below the pinacoderm cell layer where they are able to receive the highest amount of light, used for phototrophy. They can host over 50 different microbial phyla and candidate phyla, including Alphaprotoebacteria, Actinomycetota, Chloroflexota, Nitrospirota, "Cyanobacteria", the taxa Gamma-, the candidate phylum Poribacteria, and Thaumarchaea.


title: Sponge

text: Linnaeus, who classified most kinds of sessile animals as belonging to the order Zoophyta in the class Vermes, mistakenly identified the genus "Spongia" as plants in the order Algae. For a long time thereafter sponges were assigned to a separate subkingdom, Parazoa ("beside the animals"), separate from the Eumetazoa which formed the rest of the kingdom Animalia. They have been regarded as a paraphyletic phylum, from which the higher animals have evolved. Other research indicates Porifera is monophyletic.


title: Sponge

text: The phylum Porifera is further divided into classes mainly according to the composition of their skeletons:


title: Sponge

text: In the 1970s, sponges with massive calcium carbonate skeletons were assigned to a separate class, Sclerospongiae, otherwise known as "coralline sponges".


title: Sponge

text: However, in the 1980s it was found that these were all members of either the Calcarea or the Demospongiae.


title: Sponge

text: So far scientific publications have identified about 9,000 poriferan species, of which: about 400 are glass sponges; about 500 are calcareous species; and the rest are demosponges. However, some types of habitat, vertical rock and cave walls and galleries in rock and coral boulders, have been investigated very little, even in shallow seas.


title: Sponge

text: Sponges were traditionally distributed in three classes: calcareous sponges (Calcarea), glass sponges (Hexactinellida) and demosponges (Demospongiae). However, studies have shown that the Homoscleromorpha, a group thought to belong to the Demospongiae, is actually phylogenetically well separated. Therefore, they have recently been recognized as the fourth class of sponges.


title: Sponge

text: Sponges are divided into classes mainly according to the composition of their skeletons: These are arranged in evolutionary order as shown below in ascending order of their evolution from top to bottom:


title: Sponge

text: Although molecular clocks and biomarkers suggest sponges existed well before the Cambrian explosion of life, silica spicules like those of demosponges are absent from the fossil record until the Cambrian. An unsubstantiated 2002 report exists of spicules in rocks dated around . Well-preserved fossil sponges from about in the Ediacaran period have been found in the Doushantuo Formation. These fossils, which include: spicules; pinacocytes; porocytes; archeocytes; sclerocytes; and the internal cavity, have been classified as demosponges. Fossils of glass sponges have been found from around in rocks in Australia, China, and Mongolia. Early Cambrian sponges from Mexico belonging to the genus "Kiwetinokia" show evidence of fusion of several smaller spicules to form a single large spicule. Calcium carbonate spicules of calcareous sponges have been found in Early Cambrian rocks from about in Australia. Other probable demosponges have been found in the Early Cambrian Chengjiang fauna, from . Fossils found in the Canadian Northwest Territories dating to may be sponges; if this finding is confirmed, it suggests the first animals appeared before the Neoproterozoic oxygenation event.


title: Sponge

text: Freshwater sponges appear to be much younger, as the earliest known fossils date from the Mid-Eocene period about . Although about 90% of modern sponges are demosponges, fossilized remains of this type are less common than those of other types because their skeletons are composed of relatively soft spongin that does not fossilize well.


title: Sponge

text: A chemical tracer is 24-isopropylcholestane, which is a stable derivative of 24-isopropylcholesterol, which is said to be produced by demosponges but not by eumetazoans ("true animals", i.e. cnidarians and bilaterians). Since choanoflagellates are thought to be animals' closest single-celled relatives, a team of scientists examined the biochemistry and genes of one choanoflagellate species. They concluded that this species could not produce 24-isopropylcholesterol but that investigation of a wider range of choanoflagellates would be necessary in order to prove that the fossil 24-isopropylcholestane could only have been produced by demosponges.


title: Sponge

text: Although a previous publication reported traces of the chemical 24-isopropylcholestane in ancient rocks dating to , recent research using a much more accurately dated rock series has revealed that these biomarkers only appear before the end of the Marinoan glaciation approximately , and that "Biomarker analysis has yet to reveal any convincing evidence for ancient sponges pre-dating the first globally extensive Neoproterozoic glacial episode (the Sturtian, ~ in Oman)". While it has been argued that this 'sponge biomarker' could have originated from marine algae, recent research suggests that the algae's ability to produce this biomarker evolved only in the Carboniferous; as such, the biomarker remains strongly supportive of the presence of demosponges in the Cryogenian.


title: Sponge

text: Archaeocyathids, which some classify as a type of coralline sponge, are very common fossils in rocks from the Early Cambrian about , but apparently died out by the end of the Cambrian .


title: Sponge

text: It has been suggested that they were produced by: sponges; cnidarians; algae; foraminiferans; a completely separate phylum of animals, Archaeocyatha; or even a completely separate kingdom of life, labeled Archaeata or Inferibionta. Since the 1990s archaeocyathids have been regarded as a distinctive group of sponges.


title: Sponge

text: It is difficult to fit chancelloriids into classifications of sponges or more complex animals. An analysis in 1996 concluded that they were closely related to sponges on the grounds that the detailed structure of chancellorid sclerites ("armor plates") is similar to that of fibers of spongin, a collagen protein, in modern keratose (horny) demosponges such as "Darwinella". However, another analysis in 2002 concluded that chancelloriids are not sponges and may be intermediate between sponges and more complex animals, among other reasons because their skins were thicker and more tightly connected than those of sponges. In 2008 a detailed analysis of chancelloriids' sclerites concluded that they were very similar to those of halkieriids, mobile bilaterian animals that looked like slugs in chain mail and whose fossils are found in rocks from the very Early Cambrian to the Mid Cambrian. If this is correct, it would create a dilemma, as it is extremely unlikely that totally unrelated organisms could have developed such similar sclerites independently, but the huge difference in the structures of their bodies makes it hard to see how they could be closely related.


title: Sponge

text: In the 1990s sponges were widely regarded as a monophyletic group, all of them having descended from a common ancestor that was itself a sponge, and as the "sister-group" to all other metazoans (multi-celled animals), which themselves form a monophyletic group. On the other hand, some 1990s analyses also revived the idea that animals' nearest evolutionary relatives are choanoflagellates, single-celled organisms very similar to sponges' choanocytes – which would imply that most Metazoa evolved from very sponge-like ancestors and therefore that sponges may not be monophyletic, as the same sponge-like ancestors may have given rise both to modern sponges and to non-sponge members of Metazoa.


title: Sponge

text: Analyses since 2001 have concluded that Eumetazoa (more complex than sponges) are more closely related to particular groups of sponges than to other sponge groups. Such conclusions imply that sponges are not monophyletic, because the last common ancestor of all sponges would also be a direct ancestor of the Eumetazoa, which are not sponges. A study in 2001 based on comparisons of ribosome DNA concluded that the most fundamental division within sponges was between glass sponges and the rest, and that Eumetazoa are more closely related to calcareous sponges (those with calcium carbonate spicules) than to other types of sponge. In 2007 one analysis based on comparisons of RNA and another based mainly on comparison of spicules concluded that demosponges and glass sponges are more closely related to each other than either is to the calcareous sponges, which in turn are more closely related to Eumetazoa.


title: Sponge

text: Other anatomical and biochemical evidence links the Eumetazoa with Homoscleromorpha, a sub-group of demosponges. A comparison in 2007 of nuclear DNA, excluding glass sponges and comb jellies, concluded that:


title: Sponge

text: The sperm of Homoscleromorpha share features with the sperm of Eumetazoa, that sperm of other sponges lack. In both Homoscleromorpha and Eumetazoa layers of cells are bound together by attachment to a carpet-like basal membrane composed mainly of "typ IV" collagen, a form of collagen not found in other sponges – although the spongin fibers that reinforce the mesohyl of all demosponges is similar to "type IV" collagen.


title: Sponge

text: The analyses described above concluded that sponges are closest to the ancestors of all Metazoa, of all multi-celled animals including both sponges and more complex groups. However, another comparison in 2008 of 150 genes in each of 21 genera, ranging from fungi to humans but including only two species of sponge, suggested that comb jellies (ctenophora) are the most basal lineage of the Metazoa included in the sample. If this is correct, either modern comb jellies developed their complex structures independently of other Metazoa, or sponges' ancestors were more complex and all known sponges are drastically simplified forms. The study recommended further analyses using a wider range of sponges and other simple Metazoa such as Placozoa.


title: Sponge

text: The results of such an analysis, published in 2009, suggest that a return to the previous view may be warranted. 'Family trees' constructed using a combination of all available data – morphological, developmental and molecular – concluded that the sponges are in fact a monophyletic group, and with the cnidarians form the sister group to the bilaterians.


title: Sponge

text: A very large and internally consistent alignment of 1,719 proteins at the metazoan scale, published in 2017, showed that (i) sponges – represented by Homoscleromorpha, Calcarea, Hexactinellida, and Demospongiae – are monophyletic, (ii) sponges are sister-group to all other multicellular animals, (iii) ctenophores emerge as the second-earliest branching animal lineage, and (iv) placozoans emerge as the third animal lineage, followed by cnidarians sister-group to bilaterians.


title: Sponge

text: In March 2021, scientists from Dublin found additional evidence that sponges are the sister group to all other animals.


title: Sponge

text: A report in 1997 described use of sponges as a tool by bottlenose dolphins in Shark Bay in Western Australia. A dolphin will attach a marine sponge to its rostrum, which is presumably then used to protect it when searching for food in the sandy sea bottom. The behavior, known as "sponging", has only been observed in this bay, and is almost exclusively shown by females. A study in 2005 concluded that mothers teach the behavior to their daughters, and that all the sponge-users are closely related, suggesting that it is a fairly recent innovation.


title: Sponge

text: The calcium carbonate or silica spicules of most sponge genera make them too rough for most uses, but two genera, "Hippospongia" and "Spongia", have soft, entirely fibrous skeletons. Early Europeans used soft sponges for many purposes, including padding for helmets, portable drinking utensils and municipal water filters. Until the invention of synthetic sponges, they were used as cleaning tools, applicators for paints and ceramic glazes and discreet contraceptives. However, by the mid-20th century, over-fishing brought both the animals and the industry close to extinction.


title: Sponge

text: Many objects with sponge-like textures are now made of substances not derived from poriferans. Synthetic sponges include personal and household cleaning tools, breast implants, and contraceptive sponges. Typical materials used are cellulose foam, polyurethane foam, and less frequently, silicone foam.


title: Sponge

text: The luffa "sponge", also spelled "loofah", which is commonly sold for use in the kitchen or the shower, is not derived from an animal but mainly from the fibrous "skeleton" of the sponge gourd ("Luffa aegyptiaca", Cucurbitaceae).


title: Sponge

text: Sponges have medicinal potential due to the presence in sponges themselves or their microbial symbionts of chemicals that may be used to control viruses, bacteria, tumors and fungi.


title: Sponge

text: Lacking any protective shell or means of escape, sponges have evolved to synthesize a variety of unusual compounds. One such class is the oxidized fatty acid derivatives called oxylipins. Members of this family have been found to have anti-cancer, anti-bacterial and anti-fungal properties. One example isolated from the Okinawan "plakortis" sponges, plakoridine A, has shown potential as a cytotoxin to murine lymphoma cells.


title: Sumerian language

text: Sumerian (Cuneiform: "native tongue") is the language of ancient Sumer. It is one of the oldest attested languages, dating back to at least 3000 BC. It is accepted to be a local language isolate and to have been spoken in ancient Mesopotamia, in the area that is modern-day Iraq.


title: Sumerian language

text: Akkadian, a Semitic language, gradually replaced Sumerian as a spoken language in the area around 2000 BC (the exact date is debated), but Sumerian continued to be used as a sacred, ceremonial, literary and scientific language in Akkadian-speaking Mesopotamian states such as Assyria and Babylonia until the 1st century AD. Thereafter it seems to have fallen into obscurity until the 19th century, when Assyriologists began deciphering the cuneiform inscriptions and excavated tablets that had been left by its speakers.


title: Sumerian language

text: Archaic Sumerian is the earliest stage of inscriptions with linguistic content, beginning with the Jemdet Nasr (Uruk III) period from about the 31st to 30th centuries BC. It succeeds the proto-literate period, which spans roughly the 35th to 30th centuries.


title: Sumerian language

text: Some versions of the chronology may omit the Late Sumerian phase and regard all texts written after 2000 BC as Post-Sumerian. The term "Post-Sumerian" is meant to refer to the time when the language was already extinct and preserved by Babylonians and Assyrians only as a liturgical and classical language for religious, artistic and scholarly purposes. The extinction has traditionally been dated approximately to the end of the Third Dynasty of Ur, the last predominantly Sumerian state in Mesopotamia, about 2000 BC. However, that date is very approximate, as many scholars have contended that Sumerian was already dead or dying as early as around 2100 BC, by the beginning of the Ur III period, and others believe that Sumerian persisted, as a spoken language, in a small part of Southern Mesopotamia (Nippur and its surroundings) until as late as 1700 BC. Whatever the status of spoken Sumerian between 2000 and 1700 BC, it is from then that a particularly large quantity of literary texts and bilingual Sumerian-Akkadian lexical lists survive, especially from the scribal school of Nippur. They and the particularly intensive official and literary use of the language in Akkadian-speaking states during the same time call for a distinction between the Late Sumerian and the Post-Sumerian periods. Sumerian school documents from the Sealand Dynasty were found at Tell Khaiber, some of which contain year names from the reign of a king with the Sumerian throne name Aya-dara-galama.


title: Sumerian language

text: The standard variety of Sumerian was "Emegir" ( ). A notable variety or sociolect was "Emesal" ( eme-sal),


title: Sumerian language

text: possibly to be interpreted as "fine tongue" or "high-pitched voice" . Other terms for dialects or registers were "eme-galam" "high tongue", "eme-si-sa" "straight tongue", "eme-te-na" "oblique[?] tongue", etc.


title: Sumerian language

text: "Emesal" is used exclusively by female characters in some literary texts (that may be compared to the female languages or language varieties that exist or have existed in some cultures, such as among the Chukchis and the Garifuna). In addition, it is dominant in certain genres of cult songs such as the hymns sung by Gala priests. The special features of "Emesal" are mostly phonological (for example, "m" is often used instead of "g̃" [i.e. ], as in "me" instead of standard "g̃e" for "I"), but words different from the standard language are also used ("ga-ša-an" rather than standard "nin", "lady").


title: Sumerian language

text: Sumerian is a language isolate. Ever since decipherment, it has been the subject of much effort to relate it to a wide variety of languages. Because it has a peculiar prestige as one of the most ancient written languages, proposals for linguistic affinity sometimes have a nationalistic background. Such proposals enjoy virtually no support among linguists because of their unverifiability. Sumerian was at one time widely held to be an Indo-European language, but that view later came to be almost universally rejected.


title: Sumerian language

text: It has also been suggested that the Sumerian language descended from a late prehistoric creole language (Høyrup 1992). However, no conclusive evidence, only some typological features, can be found to support Høyrup's view.


title: Sumerian language

text: A more widespread hypothesis posits a Proto-Euphratean language that preceded Sumerian in Southern Mesopotamia and exerted an areal influence on it, especially in the form of polysyllabic words that appear "un-Sumerian"—making them suspect of being loanwords—and are not traceable to any other known language. There is little speculation as to the affinities of this substratum language, or these languages, and it is thus best treated as unclassified. Researchers such as Gonzalo Rubio disagree with the assumption of a single substratum language and argue that several languages are involved. A related proposal by Gordon Whittaker is that the language of the proto-literary texts from the Late Uruk period ( 3350–3100 BC) is really an early Indo-European language which he terms "Euphratic".


title: Sumerian language

text: The Sumerian language is one of the earliest known written languages. The "proto-literate" period of Sumerian writing spans c. 3300 to 3000 BC. In this period, records are purely logographic, with no phonological content. The oldest document of the proto-literate period is the Kish tablet. Falkenstein (1936) lists 939 signs used in the proto-literate period (late Uruk, 34th to 31st centuries).


title: Sumerian language

text: Records with unambiguously linguistic content, identifiably Sumerian, are those found at Jemdet Nasr, dating to the 31st or 30th century BC. From about 2600 BC, the logographic symbols were generalized using a wedge-shaped stylus to impress the shapes into wet clay. This "cuneiform" ("wedge-shaped") mode of writing co-existed with the pre-cuneiform archaic mode. Deimel (1922) lists 870 signs used in the Early Dynastic IIIa period (26th century). In the same period the large set of logographic signs had been simplified into a logosyllabic script comprising several hundred signs. Rosengarten (1967) lists 468 signs used in Sumerian (pre-Sargonian) Lagash. The pre-Sargonian period of the 26th to 24th centuries BC is the "Classical Sumerian" stage of the language.


title: Sumerian language

text: The cuneiform script was adapted to Akkadian writing beginning in the mid-third millennium. Over the long period of bi-lingual overlap of active Sumerian and Akkadian usage the two languages influenced each other, as reflected in numerous loanwords and even word order changes.


title: Sumerian language

text: Depending on the context, a cuneiform sign can be read either as one of several possible logograms, each of which corresponds to a word in the Sumerian spoken language, as a phonetic syllable (V, VC, CV, or CVC), or as a determinative (a marker of semantic category, such as occupation or place). (See the article Transliterating cuneiform languages.) Some Sumerian logograms were written with multiple cuneiform signs. These logograms are called diri-spellings, after the logogram 'diri' which is written with the signs SI and A. The text transliteration of a tablet will show just the logogram, such as the word 'diri', not the separate component signs.


title: Sumerian language

text: Not all epigraphists are equally reliable, and before publication of an important treatment of a text, scholars will often arrange to collate the published transcription against the actual tablet, to see if any signs, especially broken or damaged signs, should be represented differently.


title: Sumerian language

text: The key to reading logosyllabic cuneiform came from the Behistun inscription, a trilingual cuneiform inscription written in Old Persian, Elamite and Akkadian. (In a similar manner, the key to understanding Egyptian hieroglyphs was the bilingual (Greek and Egyptian with the Egyptian text in two scripts) Rosetta stone and Jean-François Champollion's transcription in 1822.)


title: Sumerian language

text: In 1838 Henry Rawlinson, building on the 1802 work of Georg Friedrich Grotefend, was able to decipher the Old Persian section of the Behistun inscriptions, using his knowledge of modern Persian. When he recovered the rest of the text in 1843, he and others were gradually able to translate the Elamite and Akkadian sections of it, starting with the 37 signs he had deciphered for the Old Persian. Meanwhile, many more cuneiform texts were coming to light from archaeological excavations, mostly in the Semitic Akkadian language, which were duly deciphered.


title: Sumerian language

text: By 1850, however, Edward Hincks came to suspect a non-Semitic origin for cuneiform. Semitic languages are structured according to consonantal forms, whereas cuneiform, when functioning phonetically, was a syllabary, binding consonants to particular vowels. Furthermore, no Semitic words could be found to explain the syllabic values given to particular signs. Julius Oppert suggested that a non-Semitic language had preceded Akkadian in Mesopotamia, and that speakers of this language had developed the cuneiform script.


title: Sumerian language

text: In 1855 Rawlinson announced the discovery of non-Semitic inscriptions at the southern Babylonian sites of Nippur, Larsa, and Uruk.


title: Sumerian language

text: In 1856, Hincks argued that the untranslated language was agglutinative in character. The language was called "Scythic" by some, and, confusingly, "Akkadian" by others. In 1869, Oppert proposed the name "Sumerian", based on the known title "King of Sumer and Akkad", reasoning that if Akkad signified the Semitic portion of the kingdom, Sumer might describe the non-Semitic annex.


title: Sumerian language

text: Credit for being first to scientifically treat a bilingual Sumerian-Akkadian text belongs to Paul Haupt, who published "Die sumerischen Familiengesetze" (The Sumerian family laws) in 1879.


title: Sumerian language

text: Ernest de Sarzec began excavating the Sumerian site of Tello (ancient Girsu, capital of the state of Lagash) in 1877, and published the first part of "Découvertes en Chaldée" with transcriptions of Sumerian tablets in 1884. The University of Pennsylvania began excavating Sumerian Nippur in 1888.


title: Sumerian language

text: The bewildering number and variety of phonetic values that signs could have in Sumerian led to a detour in understanding the language – a Paris-based orientalist, Joseph Halévy, argued from 1874 onward that Sumerian was not a natural language, but rather a secret code (a cryptolect), and for over a decade the leading Assyriologists battled over this issue. For a dozen years, starting in 1885, Friedrich Delitzsch accepted Halévy's arguments, not renouncing Halévy until 1897.


title: Sumerian language

text: François Thureau-Dangin working at the Louvre in Paris also made significant contributions to deciphering Sumerian with publications from 1898 to 1938, such as his 1905 publication of "Les inscriptions de Sumer et d'Akkad". Charles Fossey at the Collège de France in Paris was another prolific and reliable scholar. His pioneering "Contribution au Dictionnaire sumérien–assyrien", Paris 1905–1907, turns out to provide the foundation for P. Anton Deimel's 1934 "Sumerisch-Akkadisches Glossar" (vol. III of Deimel's 4-volume "Sumerisches Lexikon").


title: Sumerian language

text: In 1908, Stephen Herbert Langdon summarized the rapid expansion in knowledge of Sumerian and Akkadian vocabulary in the pages of "Babyloniaca", a journal edited by Charles Virolleaud, in an article "Sumerian-Assyrian Vocabularies", which reviewed a valuable new book on rare logograms by Bruno Meissner. Subsequent scholars have found Langdon's work, including his tablet transcriptions, to be not entirely reliable.


title: Sumerian language

text: In 1944, the Sumerologist Samuel Noah Kramer provided a detailed and readable summary of the decipherment of Sumerian in his "Sumerian Mythology".


title: Sumerian language

text: Friedrich Delitzsch published a learned Sumerian dictionary and grammar in the form of his "Sumerisches Glossar" and "Grundzüge der sumerischen Grammatik", both appearing in 1914. Delitzsch's student, Arno Poebel, published a grammar with the same title, "Grundzüge der sumerischen Grammatik", in 1923, and for 50 years it would be the standard for students studying Sumerian. Poebel's grammar was finally superseded in 1984 on the publication of "The Sumerian Language: An Introduction to its History and Grammatical Structure", by Marie-Louise Thomsen. While much of Thomsen's understanding of Sumerian grammar would later be rejected by most or all Sumerologists, Thomsen's grammar (often with express mention of the critiques put forward by Pascal Attinger in his 1993 "Eléments de linguistique sumérienne: La construction de du/e/di 'dire") is the starting point of most recent academic discussions of Sumerian grammar.


title: Sumerian language

text: More recent monograph-length grammars of Sumerian include Dietz-Otto Edzard's 2003 "Sumerian Grammar" and Bram Jagersma's 2010 "A Descriptive Grammar of Sumerian" (currently digital, but soon to be printed in revised form by Oxford University Press). Piotr Michalowski's essay (entitled, simply, "Sumerian") in the 2004 "The Cambridge Encyclopedia of the World's Ancient Languages" has also been recognized as a good modern grammatical sketch.


title: Sumerian language

text: There is relatively little consensus, even among reasonable Sumerologists, in comparison to the state of most modern or classical languages. Verbal morphology, in particular, is hotly disputed. In addition to the general grammars, there are many monographs and articles about particular areas of Sumerian grammar, without which a survey of the field could not be considered complete.


title: Sumerian language

text: The primary institutional lexical effort in Sumerian is the Pennsylvania Sumerian Dictionary project, begun in 1974. In 2004, the PSD was released on the Web as the ePSD. The project is currently supervised by Steve Tinney. It has not been updated online since 2006, but Tinney and colleagues are working on a new edition of the ePSD, a working draft of which is available online.


title: Sumerian language

text: Assumed phonological or morphological forms will be between slashes //, with plain text used for the standard Assyriological transcription of Sumerian. Most of the following examples are unattested.


title: Sumerian language

text: Modern knowledge of Sumerian phonology is flawed and incomplete because of the lack of speakers, the transmission through the filter of Akkadian phonology and the difficulties posed by the cuneiform script. As I. M. Diakonoff observes, "when we try to find out the morphophonological structure of the Sumerian language, we must constantly bear in mind that we are not dealing with a language directly but are reconstructing it from a very imperfect mnemonic writing system which had not been basically aimed at the rendering of morphophonemics".


title: Sumerian language

text: The existence of various other consonants has been hypothesized based on graphic alternations and loans, though none have found wide acceptance. For example, Diakonoff lists evidence for two l-sounds, two r-sounds, two h-sounds, and two g-sounds (excluding the velar nasal), and assumes a phonemic difference between consonants that are dropped word-finally (such as the "g" in zag > "za") and consonants that remain (such as the "g" in "lag"). Other "hidden" consonant phonemes that have been suggested include semivowels such as and , and a glottal fricative or a glottal stop that could explain the absence of vowel contraction in some words—though objections have been raised against that as well. A recent descriptive grammar by Bram Jagersma includes , , and as unwritten consonants, with the glottal stop even serving as the first-person pronominal prefix.


title: Sumerian language

text: Very often, a word-final consonant was not expressed in writing—and was possibly omitted in pronunciation—so it surfaced only when followed by a vowel: for example the /k/ of the genitive case ending "-ak" does not appear in "e lugal-la" "the king's house", but it becomes obvious in "e lugal-la-kam" "(it) is the king's house" (compare liaison in French).


title: Sumerian language

text: The vowels that are clearly distinguished by the cuneiform script are , , , and . Various researchers have posited the existence of more vowel phonemes such as and even and , which would have been concealed by the transmission through Akkadian, as that language does not distinguish them. That would explain the seeming existence of numerous homophones in transliterated Sumerian, as well as some details of the phenomena mentioned in the next paragraph. These hypotheses are not yet generally accepted.


title: Sumerian language

text: There is some evidence for vowel harmony according to vowel height or advanced tongue root in the prefix i/e- in inscriptions from pre-Sargonic Lagash, and perhaps even more than one vowel harmony rule. There also appear to be many cases of partial or complete assimilation of the vowel of certain prefixes and suffixes to one in the adjacent syllable reflected in writing in some of the later periods, and there is a noticeable, albeit not absolute, tendency for disyllabic stems to have the same vowel in both syllables. These patterns, too, are interpreted as evidence for a richer vowel inventory by some researchers. What appears to be vowel contraction in hiatus (*/aa/, */ia/, */ua/ > a, */ae/ > a, */ue/ > u, etc.) is also very common.


title: Sumerian language

text: Syllables could have any of the following structures: V, CV, VC, CVC. More complex syllables, if Sumerian had them, are not expressed as such by the cuneiform script.


title: Sumerian language

text: Ever since its decipherment, research of Sumerian has been made difficult not only by the lack of any native speakers, but also by the relative sparseness of linguistic data, the apparent lack of a closely related language, and the features of the writing system. Typologically, as mentioned above, Sumerian is classified as an agglutinative, split ergative, and subject-object-verb language. It behaves as a nominative–accusative language in the 1st and 2nd persons of the incomplete tense-aspect, but as ergative–absolutive in most other forms of the indicative mood. Sumerian nouns are organized in two grammatical genders based on animacy: animate and inanimate. Animate nouns include humans, gods, and in some instances the word for "statue". Case is indicated by suffixes on the noun. Noun phrases are right branching with adjectives and modifiers following nouns.


title: Sumerian language

text: Sumerian verbs have a tense-aspect complex, contrasting complete and incomplete actions/states. The two have different conjugations and many have different roots. Verbs also mark mood, voice, polarity, iterativity, and intensity; and agree with subjects and objects in number, person, animacy, and case. Sumerian moods are: indicative, imperative, cohortative, precative/affirmative, prospective aspect/cohortative mood, affirmative/negative-volitive, unrealised-volitive?, negative?, affirmative?, polarative, and are marked by a verbal prefix. The prefixes appear to conflate mood, aspect, and polarity; and their meanings are also affected by the tense-aspect complex. Sumerian voices are: active, and middle or passive. Verbs are marked for three persons: 1st, 2nd, 3rd; in two numbers: singular and plural. Finite verbs have three classes of prefixes: modal prefixes, conjugational prefixes, and pronominal/dimensional prefixes. Modal prefixes confer the above moods on the verb. Conjugational prefixes are thought to confer perhaps venitive/andative, being/action, focus, valency, or voice distinctions on the verb. Pronominal/dimensional prefixes correspond to noun phrases and their cases. Non-finite verbs include participles and relative clause verbs, both formed through nominalisation. Finite verbs take prefixes and suffixes, non-finite verbs only take suffixes. Verbal roots are mostly monosyllabic, though verbal root duplication and suppletion can also occur to indicate plurality. Root duplication can also indicate iterativity or intensity of the verb.


title: Sumerian language

text: The Sumerian noun is typically a one or two-syllable root ("igi" "eye", "e" "house, household", "nin" "lady"), although there are also some roots with three syllables like "šakanka" "market". There are two grammatical genders, usually called human and non-human (the first includes gods and the word for "statue" in some instances, but not plants or animals, the latter also includes collective plural nouns), whose assignment is semantically predictable.


title: Sumerian language

text: The adjectives and other modifiers follow the noun ("lugal maḫ" "great king"). The noun itself is not inflected; rather, grammatical markers attach to the noun phrase as a whole, in a certain order. Typically, that order would be:


title: Sumerian language

text: An example may be /dig̃ir gal-gal-g̃u-ne-ra/ ("god great (reduplicated)-my-plural-dative" = "for all my great gods"). The possessive, plural and case markers are traditionally referred to as "suffixes", but have recently also been described as enclitics or postpositions.


title: Sumerian language

text: The plural markers are /-(e)ne/ (optional) for nouns of the human gender. Non-human nouns are not marked by a plural suffix. However, plurality can also be expressed with the adjective "ḫi-a" "various", with the plural of the copula /-meš/, by reduplication of the noun ("kur-kur" "all foreign lands") or of the following adjective ("a gal-gal" "all the great waters") (reduplication is believed to signify totality) or by the plurality of only the verb form. Plural reference in the verb form occurs only for human nouns.


title: Sumerian language

text: More endings are recognised by some researchers; e.g. Bram Jagersma notes a separate adverbiative case in /-eš/ and a second locative used mostly with infinite verb forms.


title: Sumerian language

text: Additional spatial or temporal meanings can be expressed by genitive phrases like "at the head of" = "above", "at the face of" = "in front of", "at the outer side of" = "because of" etc.: "bar udu ḫad-ak-a" = "outer.side sheep white-genitive-locative" = "in the outer side of a white sheep" = "because of a white sheep".


title: Sumerian language

text: The embedded structure of the noun phrase can be further illustrated with the phrase "sipad udu siki-ak-ak-ene" ("the shepherds of woolly sheep"), where the first genitive morpheme ("-a(k)") subordinates "siki" "wool" to "udu" "sheep", and the second subordinates "udu siki-a(k)" "sheep of wool" (or "woolly sheep") to "sipad" "shepherd".


title: Sumerian language

text: Sumerian has a combination decimal and sexagesimal system (for example, 600 is 'ten sixties'), so that the Sumerian lexical numeral system is sexagesimal with 10 as a subbase. Numerals and composite numbers are as follows:


title: Sumerian language

text: The Sumerian finite verb distinguishes a number of moods and agrees (more or less consistently) with the subject and the object in person, number and gender. The verb chain may also incorporate pronominal references to the verb's other modifiers, which has also traditionally been described as "agreement", although, in fact, such a reference "and" the presence of an actual modifier in the clause need not co-occur: not only e-še ib-ši-du-un" "I'm going to the house", but also e-še i-du-un" "I'm going to the house" and simply "ib-ši-du-un" "I'm going to it" are possible.


title: Sumerian language

text: The Sumerian verb also makes a binary distinction according to a category that some regard as tense (past vs present-future), others as aspect (perfective vs imperfective), and that will be designated as TA (tense/aspect) in the following. The two members of the opposition entail different conjugation patterns and, at least for many verbs, different stems; they are theory-neutrally referred to with the Akkadian grammatical terms for the two respective forms – "ḫamṭu" (quick) and "marû" (slow, fat). Finally, opinions differ on whether the verb has a passive or a middle voice and how it is expressed.


title: Sumerian language

text: The verbal root is almost always a monosyllable and, together with various affixes, forms a so-called verbal chain which is described as a sequence of about 15 slots, though the precise models differ. The finite verb has both prefixes and suffixes, while the non-finite verb may only have suffixes. Broadly, the prefixes have been divided in three groups that occur in the following order: "modal prefixes", ""conjugation prefixes"", and "pronominal and dimensional" prefixes. The suffixes are a future or imperfective marker /-ed-/, pronominal suffixes, and an /-a/ ending that nominalizes the whole verb chain. The overall structure can be summarized as follows:


title: Sumerian language

text: Note also that more than one pairing of a pronominal prefix and a dimensional prefix may occur within the verb chain.


title: Sumerian language

text: The meaning, structure, identity and even the number of "conjugation prefixes" have always been a subject of disagreements. The term "conjugation prefix" simply alludes to the fact that a finite verb in the indicative mood must always contain one of them. Some of their most frequent expressions in writing are "mu-, i-" (ED Lagaš variant: "e-"), "ba-", "bi-" (ED Lagaš: "bi- or be"), "im-", "im-ma-" (ED Lagaš "e-ma-"), "im-mi-" (ED Lagaš "i-mi" or "e-me-"), "mi-" (always followed by pronominal-dimensional "-ni-") and "al-", and to a lesser extent "a-", "am-", "am-ma-", and "am-mi-"; virtually all analyses attempt to describe many of the above as combinations or allomorphs of each other.


title: Sumerian language

text: The starting point of most analyses are the obvious facts that the 1st person dative always requires "mu-", and that the verb in a "passive" clause without an overt agent tends to have "ba-". Proposed explanations usually revolve around the subtleties of spatial grammar, information structure (focus), verb valency, and, most recently, voice. "Mu-", "im-" and "am-" have been described as ventive morphemes, while "ba-" and "bi-" are sometimes analyzed as actually belonging to the pronominal-dimensional group (inanimate pronominal /-b-/ + dative /-a-/ or directive /-i-/). "Im-ma-", "im-mi-", "am-ma-" and "am-mi-" are then considered by some as a combination of the ventive and /ba-/, /bi-/ or otherwise a variety of the ventive. The element "i-" has been argued to be a mere prothetic vowel, "al-" a stative prefix, "ba-" a middle voice prefix, etcetera.


title: Sumerian language

text: The dimensional prefixes of the verb chain basically correspond to, and often repeat, the case markers of the noun phrase. Like the latter, they are attached to a "head" – a pronominal prefix. The other place where a pronominal prefix can be placed is immediately before the stem, where it can have a different allomorph and expresses the absolutive or the ergative participant (the transitive subject, the intransitive subject or the direct object), depending on the TA and other factors, as explained below. However, this neat system is obscured by the tendency to drop or merge many of the prefixes in writing and possibly in pronunciation as well.


title: Sumerian language

text: The generally recognized dimensional prefixes are shown in the table below; if several occur within the same verb complex, they are placed in the order they are listed in.


title: Sumerian language

text: The morphemes /-n-/ and /-b-/ are clearly the prefixes for the 3rd person singular animate and inanimate respectively; the 2nd person singular appears as "-e-" in most contexts, but as /-r-/ before the dative (-ra-), leading some to assume a phonetic /-ir-/ or /-jr-/. The 1st person may appear as "-e-", too, but is more commonly not expressed at all (the same may frequently apply to 3rd and 2nd persons); it is, however, cued by the choice of "mu-" as conjugation prefix (/mu-/ + /-a-/ → "ma-"). The 1st, 2nd and 3rd plural infixes are "-me-", "-re"?"-" and "-ne-" in the dative and perhaps in other contexts as well, though not in the pre-stem position (see below).


title: Sumerian language

text: An additional exception from the system is the prefix "-ni-" which corresponds to a noun phrase in the locative – in which case it doesn't seem to be preceded by a pronominal prefix – and, according to Gábor Zólyomi and others, to an animate one in the directive – in the latter case it is analyzed as pronominal /-n-/ + directive /-i-/. Zólyomi and others also believe that special meanings can be expressed by combinations of non-identical noun case and verb prefix. Also according to some researchers /-ni-/ and /bi-/ acquire the forms /-n-/ and /-b-/ (coinciding with the "absolutive–ergative" pronominal prefixes) before the stem if there isn't already an absolutive–ergative pronominal prefix in pre-stem position: "mu-un-kur" = /mu-ni-kur/ "he went in there" (as opposed to "mu-ni-kur" = "mu-ni-in-kur" = /mu-ni-n-kur/ "he brought in – caused [something or someone] to go in – there".


title: Sumerian language

text: The general principle for pronominal agreement in conjugation is that in "ḫamṭu" TA, the transitive subject is expressed by the prefix, and the direct object by the suffix, and in the "marû" TA it is the other way round; as for the intransitive subject, it is expressed, in both TAs, by the suffixes and is thus treated like the object in "ḫamṭu" and like the subject in "marû" (except that its third person is expressed, not only in "ḫamṭu" but also in "marû", by the suffixes used for the "object" in the "ḫamṭu" TA). A major exception from this generalization are the plural forms – in them, not only the prefix (as in the singular), but also the suffix expresses the transitive subject.


title: Sumerian language

text: Additionally, the prefixes of the plural are identical to those of the singular – /-?-/ or /-e-/, /-e-/, /-n-/, /-b-/ – as opposed to the "-me-", "-re-?", "-ne-" that are presumed for non-pre-stem position – and some scholars believe that the prefixes of the 1st and second person are /-en-/ rather than /-e-/ when they stand for the object. Before the pronominal suffixes, a suffix /-e(d)-/ with a future or related modal meaning can be inserted, accounting for occurrences of "-e" in the third-person singular "marû" of intransitive forms; because of its meaning, it can also be said to signal "marû" in these forms.


title: Sumerian language

text: Examples for TA and pronominal agreement: ("ḫamṭu" is rendered with past tense, "marû" with present): /i-gub-en/ ("I stood" or "I stand"), /i-n-gub-en/ ("he placed me" or "I place him"); /i-sug-enden/ ("we stood/stand"); /i-n-dim-enden/ ("he created us" or "we create him"); /mu-e?-dim-enden/ ("we created [someone or something]"); i-gub-be = /i-gub-ed/ ("he will/must stand"); ib-gub-be = /i-b-gub-e/ ("he places it"); /i-b-dim-ene/ ("they create it"), /i-n-dim-eš/ ("they created [someone or something]" or "he created them"), /i-sug-eš/ ("they stood" or "they stand").


title: Sumerian language

text: Confusingly, the subject and object prefixes (/-n-/, /-b-/, /-e-/) are not commonly spelled out in early texts, although the "full" spellings do become more usual during the Third Dynasty of Ur (in the Neo-Sumerian period) and especially during the Late Sumerian period. Thus, in earlier texts, one finds "mu-ak" and "i-ak" ("e-ak" in early dynastic Lagash) instead of "mu-un-ak" and "in-ak" for /mu-n-ak/ and /i-n-ak/ "he/she made", and also "mu-ak" instead of "mu-e-ak" "you made". Similarly, pre-Ur III texts also spell the first- and second-person suffix /-en/ as "-e", making it coincide with the third person in the "marû" form.


title: Sumerian language

text: The verbal stem itself can also express grammatical distinctions. The plurality of the absolutive participant can be expressed by complete reduplication of the stem or by a suppletive stem. Reduplication can also express "plurality of the action itself", intensity or iterativity.


title: Sumerian language

text: The nominalizing suffix /-a/ converts non-finite and finite verbs into participles and relative clauses: "šum-ma" "given", "mu-na-an-šum-ma" "which he gave to him", "who gave (something) to him", etc.. Adding /-a/ after the future/modal suffix /-ed/ produces a form with a meaning similar to the Latin gerundive: "šum-mu-da" = "which will/should be given". On the other hand, adding a (locative-terminative?) /-e/ after the /-ed/ yields a form with a meaning similar to the Latin "ad" + gerund (acc.) construction: "šum-mu-de" = "(in order) to give".


title: Sumerian language

text: The copula verb /me/ "to be" is mostly used as an enclitic: "-men", "-men", "-am", "-menden", "-menzen", "-(a)meš".


title: Sumerian language

text: The imperative mood construction is produced with a singular "ḫamṭu" stem, but using the "marû" agreement pattern, by turning all prefixes into suffixes: "mu-na-an-sum" "he gave (something) to him", "mu-na-e-sum-mu-un-ze-en" "you (plur.) gave (something) to him" – "sum-mu-na-ab" "give it to him!", "sum-mu-na-ab-ze-en" "give (plur.) it to him!" Compare the French "vous le lui donnez", but "donnez-le-lui!"


title: Sumerian language

text: The basic word order is subject–object–verb; verb finality is only violated in rare instances, in poetry. The moving of a constituent towards the beginning of the phrase may be a way to highlight it, as may the addition of the copula to it. The so-called anticipatory genitive ("e-a lugal-bi" "the owner of the house/temple", lit. "of the house, its owner") is common and may signal the possessor's topicality. There are various ways to express subordination, some of which have already been hinted at; they include the nominalization of a verb, which can then be followed by case morphemes and possessive pronouns ("kur-ra-ni" "when he entered") and included in "prepositional" constructions ("eg̃er a-ma-ru ba-ur-ra-ta" "back – flood – conjugation prefix – sweep over – nominalizing suffix – [genitive suffix?] – ablative suffix" = "from the back of the Flood's sweeping-over" = "after the Flood had swept over"). Subordinating conjunctions such as "ud-da" "when, if", "tukum-bi" "if" are also used, though the coordinating conjunction "u" "and", a Semitic adoption, is rarely used. A specific problem of Sumerian syntax is posed by the numerous so-called compound verbs, which usually involve a noun immediately before the verb, forming a lexical or idiomatic unit (e.g. "šu...ti", lit. "hand-approach" = "receive"; "igi...du", lit. "eye-open" = "see"). Some of them are claimed to have a special agreement pattern that they share with causative constructions: their logical object, like the causee, receives, in the verb, the directive infix, but in the noun, the dative suffix if animate and the directive if inanimate.


title: Sumerian language

text: This text was inscribed on a small clay cone c. 2400 BC. It recounts the beginning of a war between the city-states of Lagaš and Umma during the Early Dynastic III period, one of the earliest border conflicts recorded. (RIME 1.09.05.01)


title: Jerry Lawler

text: Jerry O'Neil Lawler (born November 29, 1949), better known as Jerry "The King" Lawler, is an American color commentator and professional wrestler. He is currently signed to WWE, although he has not performed as a full-time commentator since April 2020.


title: Jerry Lawler

text: Prior to joining WWE in 1992 (then known as the World Wrestling Federation), he wrestled in numerous territories, winning many championships, including multiple World Heavyweight Championships, throughout his career. Lawler is a one-time AWA World Heavyweight Champion and a three-time WCWA World Heavyweight Champion: he unified the titles by defeating Kerry Von Erich at Superclash III, forming the USWA Unified World Heavyweight Championship, a championship which he held 28 times. He is also known for his feud with comedian Andy Kaufman and he portrayed himself in the 1999 film "Man on the Moon".


title: Jerry Lawler

text: Lawler has held more recognized championships than any professional wrestler in history though he has never won any championships in WWE having wrestled sporadically whilst primarily providing color commentary, since joining the company. In 2007, Lawler was inducted into the WWE Hall of Fame.


title: Jerry Lawler

text: Lawler was born November 29, 1949, in Memphis, Tennessee, where he graduated from Treadwell High School. When he was 19, his father, Jerome Lawler, died.


title: Jerry Lawler

text: While working in Memphis, Tennessee, as a disc jockey, Lawler's artistic ability attracted the attention of local wrestling promoter Aubrey Griffith.


title: Jerry Lawler

text: The two made an agreement in which Lawler would give Griffith free publicity in exchange for free wrestling training. Lawler debuted as a wrestler in 1970, and won his first championship in September 1971 by winning a battle royal. He soon won the NWA Southern Tag Team Championship under the managerial service of Sam Bass with partner Jim White. In 1974, Lawler began feuding with Jackie Fargo, who had been his trainer and mentor. This led to a match for the NWA Southern Heavyweight Championship. On July 24, 1974, Lawler won the belt and the title of "King of Wrestling."


title: Jerry Lawler

text: During 1975, Lawler teamed with a variety of partners such as Mr Wrestling II, Don Greene, and Bob Orton, Jr. He won the NWA Macon Tag Team Championship twice during this period. While Lawler began his career as a heel, he became a face after splitting from Bass at the end of 1974.


title: Jerry Lawler

text: In 1977, promoter Jerry Jarrett broke away from Nick Gulas' NWA Mid-America promotion and formed his own promotion, the Continental Wrestling Association. Lawler—Gulas' biggest star—opted to join Jarrett. The CWA quickly outperformed NWA Mid-America, which ultimately folded in 1981. Lawler was both a co-owner of the CWA and its top star.


title: Jerry Lawler

text: On November 12, 1979, while working in the CWA, Lawler defeated Superstar Billy Graham to become the CWA World Champion. In 1980, coming off the back end of a feud with The Fabulous Freebirds, his career was put on hold due to a broken leg suffered in a game of touch football, but he returned to the ring after several months.


title: Jerry Lawler

text: In 1982, Lawler began a notorious feud with comedian Andy Kaufman. At the time, Kaufman wrestled women as part of his skits and had declared himself the Intergender Heavyweight Champion. On April 5, Lawler, who had taken exception to the skits, wrestled Kaufman in Memphis. During the course of the match, Lawler delivered two piledrivers to Kaufman, the second after the bell rang, sending him to the hospital and nearly breaking his neck. On July 29, Lawler slapped Kaufman in the face on an episode of "Late Night with David Letterman". Kaufman responded by shouting profanities and throwing his coffee at Lawler.


title: Jerry Lawler

text: Years later, Lawler appeared as himself in the Kaufman biopic "Man on the Moon"; the film revealed that Lawler's feud with Kaufman had been staged. Lawler later claimed that not only was his entire feud with Kaufman staged, but also the two were actually very good friends.


title: Jerry Lawler

text: In 1989, the CWA merged with World Class Championship Wrestling to form the United States Wrestling Association.


title: Jerry Lawler

text: On March 7, 1983, Lawler won the AWA International Championship by defeating Austin Idol. On May 30, 1983, Bill Dundee defeated Jerry Lawler for the AWA Southern Heavyweight Championship. The feud quickly escalated and on June 6, 1983, the two met in a Loser Leaves Town Match for the title, in which Lawler won. Lawler defeated Ken Patera on July 25 to begin his second reign as the International Champion. Lawler became the NWA Mid America Champion on April 12, 1984, when he defeated Randy Savage for the title. He later returned to the United States, where he defeated Bill Dundee on July 29, 1986, to begin a new reign as the AWA International Champion. Lawler feuded with Tommy Rich, Austin Idol, and Paul E. Dangerously throughout early 1987. The animosity began after controversy over an AWA World Championship title shot involving Nick Bockwinkel. During the feud, the trio defeated Lawler in a steel cage match and cut his hair, which caused a riot in the Mid-South Coliseum.


title: Jerry Lawler

text: Lawler won the AWA World Heavyweight Championship from Curt Hennig on May 9, 1988. During his reign, Lawler feuded with World Class Championship Wrestling's Champion Kerry Von Erich. He defeated Von Erich on December 15, 1988, at Superclash III to unify the two titles. Soon after, Lawler's issues with Verne Gagne led to his departure from the AWA, most notably Lawler claiming that he was never paid for the match at Superclash.


title: Jerry Lawler

text: Lawler continued feuding with Kerry Von Erich in WCCW. He lost to Von Erich in a steel cage match on November 25, 1988. He would defeat Von Erich by disqualification to retain the WCCW heavyweight title. He wrestled Mil Mascaras to a draw on July 28.


title: Jerry Lawler

text: In 1981, Lawler wrestled for Championship Wrestling From Florida feuding with Dory Funk Jr. and Terry Funk. In 1985, Lawler traveled to Hawaii, where he won the NWA Polynesian Pacific title on January 25, 1986, defeating Lars Anderson. He dropped the title to Tui Selinga on March 26 in Honolulu, Hawaii.


title: Jerry Lawler

text: In March 1989, he wrestled for New Japan Pro-Wrestling. He lost to IWGP Heavyweight Champion Tatsumi Fujinami on March 16, 1989.


title: Jerry Lawler

text: In 1989, Lawler made his debut in the Memphis brand United States Wrestling Association, where he won the USWA Unified World Heavyweight Championship 28 times from 1989 until its doors closed in 1997. While champion, he feuded with The Soultaker, Jimmy Valiant, Kamala, Eddie Gilbert, and Jeff Jarrett.


title: Jerry Lawler

text: In 1992, Lawler teamed with Jeff Jarrett in a feud against The Moondogs. The feud between Jarrett/Lawler and The Moondogs was voted the 1992 PWI Feud of the Year by "Pro Wrestling Illustrated". He won the USWA World Tag Team Championship with Jarrett four times and twice with Bill Dundee.


title: Jerry Lawler

text: Lawler dropped the USWA Unified World Heavyweight for the last time to Dutch Mantel on August 8, 1997 in an All Body Hair match. The promotion closed in November 1997.


title: Jerry Lawler

text: Lawler began his WWF career in December 1992 as an announcer on "Superstars". USWA began a working relationship with the WWF. He made his in-ring debut at the 1993 Royal Rumble when he participated in the namesake match, which was won by Yokozuna. From 1993 to 1995, he feuded with Bret Hart and the rest of the Hart family. The feud began at King of the Ring when Lawler interrupted Hart's victory ceremony and attacked Bret.


title: Jerry Lawler

text: Lawler claimed that he was the only true king in the World Wrestling Federation (WWF), and the two were scheduled to wrestle at SummerSlam to settle the dispute. At the event, however, Lawler came to the ring on crutches and claimed that he could not wrestle because of injuries suffered in a car accident. Hart faced Lawler's "court jester", Doink the Clown instead, and beat him by submission. Lawler then attacked Hart, revealing that he was not injured. Hart defeated Lawler by submission but refused to release the Sharpshooter. As a result, the referee reversed the decision and awarded the title of "Undisputed King of the World Wrestling Federation" to Lawler. The two would continue to work throughout the fall on the house show circuit, including in steel cages.


title: Jerry Lawler

text: Simultaneously, in a form of cross-promotion, Lawler engaged in a bitter feud with Vince McMahon (who at the time was never advertised as the actual owner of the World Wrestling Federation) back in the USWA. There, Lawler played the babyface to his hometown Memphis audience, whereas McMahon (who had always played face in the WWF) was being portrayed as a smug heel intent on dethroning Lawler as the king of professional wrestling. As part of the cross-promotion, McMahon, Bret and Owen Hart, Giant González, Tatanka, and "Macho Man" Randy Savage would begin appearing on USWA television to further the feud. While the program continued in the USWA, the feud between Lawler and McMahon would not be acknowledged on WWF television.


title: Jerry Lawler

text: The Hart family (Bret, Owen, Bruce, and Keith) was scheduled to face a team captained by Lawler in an elimination match at Survivor Series. However, Shawn Michaels had to take Lawler's place because Lawler was facing legal troubles. He was indicted for raping and sodomizing a 15-year-old girl, though charges were dropped when the alleged victim recanted her story.


title: Jerry Lawler

text: As a result, the feud between Lawler and Vince McMahon back in the USWA was also abruptly discontinued. Lawler did not face Bret Hart at another pay-per-view until the first In Your House, when he beat Hart after Hakushi and his manager Shinja interfered. This set up a "Kiss My Foot" match at King of the Ring 1995, which Bret won. As a result, Lawler was forced to kiss Bret's feet. The feud took one final turn when Lawler introduced his "dentist" Isaac Yankem, D.D.S.. After Hart defeated Yankem by disqualification, however, the feud quickly disappeared.


title: Jerry Lawler

text: Following the end of his legal troubles which kept him out of Survivor Series 1993, Lawler eventually returned to the WWF at WrestleMania X, which was also his first appearance as a commentator on a WWF pay-per-view. During the main event of the night, "Rowdy" Roddy Piper served as special guest referee for the second WWF World Heavyweight Championship match. During this Lawler began making disparaging remarks about him. Lawler would continue to berate Piper on later episodes of "Monday Night Raw", including bringing a skinny kid into the ring dressed as Piper and forcing him to kiss his feet. This ultimately led to a match between the two at King of the Ring 1994 which Lawler lost.


title: Jerry Lawler

text: In the fall of 1994, Lawler initiated a feud with Doink the Clown. Lawler popped the balloons carried by Doink's midget sidekick, Dink. After Doink and Dink retaliated, Lawler introduced a midget sidekick of his own, who he named Queasy. In the following weeks, Doink added two more sidekicks, Wink and Pink, while Lawler introduced Sleazy and Cheesy. This led to an elimination match at Survivor Series 1994, which Lawler's team won. After the match, however, Lawler's team turned on him, joining with Doink's team to attack Lawler.


title: Jerry Lawler

text: In late 1994 and early 1995, Lawler wrestled briefly in Smoky Mountain Wrestling (SMW) while still continuing to commentate sporadically for the WWF. During his absences, Shawn Michaels filled in for him as color commentator on "Monday Night Raw". He defeated Tony Anthony for the promotion's top title in January 1995. Lawler was the last SMW Heavyweight Champions defeating Brad Armstrong on Boxing Day 1995. The promotion closed its doors on December 30.


title: Jerry Lawler

text: By 1996, Lawler wrestled occasionally on "WWF Superstars" where he would take on jobbers while holding a microphone in the ring to, essentially, "do the commentary on (his) very own matches" while occasionally serving as the official cornerman for Isaac Yankem D.D.S. After a brief feud with The Ultimate Warrior, Lawler began feuding with Jake Roberts after making fun of Roberts' real life drug and alcohol problems. The two met in a match at SummerSlam 1996, which Lawler won. After the match, Lawler poured Jim Beam whiskey down Roberts' throat.


title: Jerry Lawler

text: In early 1997, Lawler was involved in a working relationship between the WWF and Extreme Championship Wrestling (ECW). In June, Lawler entered the King of the Ring tournament for the first time and advanced to the semi-final round where he was defeated by Mankind. By the fall, the WWF introduced a new "light-heavyweight division" to compete with World Championship Wrestling's cruiserweight division. Lawler's son, Brian Christopher, was one of the major superstars in the division, although the WWF played up an angle where both Lawler and Christopher would deny their family relationship, even though the two would aide each other in matches and so on. USWA folded in November of that year.


title: Jerry Lawler

text: By 1998, Lawler rarely wrestled in the WWF and focused on commentary. Despite their feud in the USWA in 1993, by 1998, Vince McMahon had turned heel in the WWF for the first time and left the announce position, to which Lawler began praising McMahon's name on commentary as part of his own heel persona, much to the chagrin of Jim Ross. It was McMahon's departure from the commentary team which led to the strong on-screen chemistry between Lawler and Ross in subsequent years. This played a key role in a change of Lawler's character; although he still supported the heels, he showed a sense of right and wrong, and would condemn actions of heels when they went too far.


title: Jerry Lawler

text: Lawler during this period would continue commentating and rarely wrestled for WWF. He wrestled between 1999 and 2001 mainly in house shows. On June 22; 2000, he made an appearance on "SmackDown!" teaming with The Kat to defeat Dean Malenko and Terri Runnels. This would be the first time in two years he wrestled on WWF television. A week later, he defeated Malenko on "Raw."


title: Jerry Lawler

text: He would turn face by 2000 (while wrestling). This began when Lawler surprisingly attacked Tazz when the latter started bullying Jim Ross. At SummerSlam, The King wrestled Tazz in defense of Ross. Lawler won the match. A feud with Tazz begun. At Unforgiven, he lost to Tazz in a Leather Strap match. Also, Lawler feuded with Tazz's partner Raven until January 2001.


title: Jerry Lawler

text: With the creation of the XFL in 2001, Lawler was given the job as an announcer for the new football league. Lawler claims that he never wanted to announce for the XFL (he would admit on-air that he knew and cared almost nothing about the sport), but that he agreed to it after McMahon and Kevin Dunn asked him.


title: Jerry Lawler

text: In February 2001, Lawler's then-wife Stacy "The Kat" Carter was involved in a storyline where Right to Censor, a group of wrestlers purportedly wanting to rein in the vulgarity of the "Attitude Era," during which she demanded equal time for the "right for nudity". RTC's leader Steven Richards offered a match with Lawler at No Way Out. If Lawler won, Kat got naked and if Richards won, Kat became RTC property. Richards won the match at No Way Out. The next night, Lawler teamed with the APA (Bradshaw and Faarooq) as they defeated Right to Censor members Bull Buchanan, The Goodfather and Val Venis on "Raw." This would be Lawler's last WWF match.


title: Jerry Lawler

text: Lawler has stated several theories as to why he was allowed to leave. His first involves the ascent of Carter's career alongside the downfall of Chyna's. In his contention, Chyna was jealous of his wife's push inside the company, in part due to the Right to Censor storyline, and in part because of his wife's offer to pose for "Playboy" magazine. Until that time, Chyna had been the second major wrestling star from the World Wrestling Federation to have done a piece in "Playboy" (after Sable); during Chyna's debut on the adult magazine, she had suddenly broken her friendship with Carter.


title: Jerry Lawler

text: In interviews, he has also stated that there may have been an alternate reason, namely, that the company wanted to fire him. He also criticized McMahon for the cavalier attitude he had given him on the day he quit. In his recollection, Ross was fired by McMahon with exactly the same demeanor while struck with a bout of Bell's palsy in 1994, a time when Ross' "usefulness" had run out. Lawler felt that Carter's release was an attempt to remove him as well, stating that the company was well-aware that he would walk out alongside his wife. Lawler's replacement with Paul Heyman launched a theory in which Heyman was promoted at Lawler's expense. Internet rumors circulated that the company was on better terms with Heyman than Lawler, and used remarks Lawler had made in criticism of ECW to launch a theory whereby Heyman wanted Lawler out. Lawler has stated repeatedly that he has no resentment towards Heyman, accusing the media and internet theorists of spreading false information.


title: Jerry Lawler

text: In 1997, Lawler became heavily involved in the working relationship between the WWF and Extreme Championship Wrestling (ECW). In interviews and commentary, he referred to the promotion as "Extremely Crappy Wrestling". His frequent insults toward ECW eventually led to the promotion "invading" "Monday Night Raw" in February 1997. He wrestled in two matches. The first on July 19 at Heat Wave 1997 in a steel cage match with Rob Van Dam and Sabu in a no contest against Rick Rude, The Sandman, and Tommy Dreamer. He wrestled against Dreamer at ECW's Hardcore Heaven pay-per-view in August, which Dreamer won.


title: Jerry Lawler

text: During his absence from the WWF, Lawler made appearances on the independent circuit in both Australia and the States, as well as joining the fledgling Xcitement Wrestling Federation (XWF) promotion alongside WCW veteran Tony Schiavone as a color commentator. On October 26, 2001, he appeared on WWA The Inception as a commentator alongside Jeremy Borash, as well as wrestling in a battle royal. He also made appearances with the International Wrestling Cartel and with Maryland Championship Wrestling, where he held the promotion's world title after defeating The Bruiser on November 2. He would return to WWF after a nine-month absence.