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Sponge

Sponges (also known as sea sponges), the members of the phylum Porifera (meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like mesohyl sandwiched between two thin layers of cells.

Sponges have unspecialized cells that can transform into other types and that often migrate between the main cell layers and the mesohyl in the process. Sponges do not have complex nervous, digestive or circulatory systems like humans. Instead, most rely on maintaining a constant water flow through their bodies to obtain food and oxygen and to remove wastes. Believed to be some of the most basal animals alive today, sponges were possibly the first to branch off the evolutionary tree from the last common ancestor of all animals, which would make them the sister group of all other animals.

The branch of zoology that studies sponges is known as spongiology.

Etymology
The term sponge derives from the Ancient Greek word.

Overview

Sponges are similar to other animals in that they are multicellular, heterotrophic, lack cell walls and produce sperm cells. Unlike other animals, they lack true tissues and organs. Some of them are radially symmetrical, but most are asymmetrical. The shapes of their bodies are adapted for maximal efficiency of water flow through the central cavity, where the water deposits nutrients and then leaves through a hole called the osculum. 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. A great majority are marine (salt-water) species, ranging in habitat from tidal zones to depths exceeding 8800 m, though there are freshwater species. All adult sponges are sessile, meaning that they attach to an underwater surface and remain fixed in place (i.e., do not travel). While in their larval stage of life, they are motile.

Many sponges have internal skeletons of spicules (skeletal-like fragments of calcium carbonate or silicon dioxide), and/or spongin (a modified type of collagen protein). 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. 90% of all known sponge species that have the widest range of habitats including all freshwater ones are demosponges that use spongin; many species have silica spicules, whereas some species have calcium carbonate exoskeletons. Calcareous sponges 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.

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.

Most sponges reproduce sexually, but they can also reproduce asexually. Sexually reproducing species release sperm cells into the water to fertilize ova released or retained by its mate or "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. Some 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.

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.

Distinguishing features
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.

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.

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 do not have a nervous system similar to that of vertebrates but may have one that is quite different. 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.

Cell types
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. 18 distinct cell types have been identified. 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.

Other types of cells live and move within the mesohyl:

 * Lophocytes are amoeba-like cells that move slowly through the mesohyl and secrete collagen fibres.
 * Collencytes are another type of collagen-producing cell.
 * Rhabdiferous cells secrete polysaccharides that also form part of the mesohyl.
 * Oocytes and spermatocytes are reproductive cells.
 * Sclerocytes secrete the mineralized spicules ("little spines") that form the skeletons of many sponges and in some species provide some defense against predators.
 * In addition to or instead of sclerocytes, demosponges have spongocytes that secrete a form of collagen that polymerizes into spongin, a thick fibrous material that stiffens the mesohyl.
 * Myocytes ("muscle cells") conduct signals and cause parts of the animal to contract.
 * "Grey cells" act as sponges' equivalent of an immune system.
 * Archaeocytes (or amoebocytes) are amoeba-like cells that are totipotent, in other words, each is capable of transformation into any other type of cell. They also have important roles in feeding and in clearing debris that block the ostia.

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

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.

Water flow and body structures
Most sponges work rather like chimneys: they take in water at the bottom and eject it from the osculum 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.

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.

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 1 mm in diameter.

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.

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 1 m 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.

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 10 cm tall and 1 cm 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.

[[File:Porifera calcifying 01.png|thumb|Sponge with calcium carbonate skeleton.

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Skeleton
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.

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.

Movement
Although adult sponges are fundamentally sessile animals, some marine and freshwater species can move across the sea bed at speeds of 1 - 4 mm 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.

Respiration, feeding and excretion

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.

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.

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.

Carnivorous sponges

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 8840 m, 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 17 - 23 m, alongside the more usual filter-feeding sponges. The cave-dwelling predators capture crustaceans under 1 mm 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.

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.

Endosymbionts
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.

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

"Immune" system
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.

Asexual

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.

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.

Sexual
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.

A few species release fertilized eggs into the water, but most retain the eggs until they hatch. By retaining the eggs, the parents can transfer symbiotic microorganisms directly to their offspring through vertical transmission, while the species who release their eggs into the water has to acquire symbionts horizontally (a combination of both is probably most common, where larvae with vertically transmitted symbionts also acquire others horizontally). There are four types of larvae, but all are lecithotrophic (non-feeding) balls of cells with an outer layer of cells whose flagella 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.

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.

Meiosis
The cytological progression of porifera oogenesis and spermatogenesis (gametogenesis) is very similar to that of other metazoa. Most of the genes from the classic set of meiotic genes, including genes for DNA recombination and double-strand break repair, that are conserved in eukaryotes are expressed in the sponges (e.g. Geodia hentscheli and Geodia phlegraei). Since porifera are considered to be the earliest divergent animals, these findings indicate that the basic toolkit of meiosis including capabilities for recombination and DNA repair were present early in eukaryote evolution.

Life cycle
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 0.2 mm per year and, if that rate is constant, specimens 1 m 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.

Coordination of activities
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.

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.

Habitats

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.

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.

The different classes of sponge live in different ranges of habitat:

 * {|class="wikitable"

! Class !! Water type !! Depth !! Type of surface ! Calcarea ! Glass sponges ! Demosponges
 * Marine ||less than 100 m ||Hard
 * Marine ||Deep ||Soft or firm sediment
 * Marine, brackish; and about 150 freshwater species ||Inter-tidal to abyssal; a carnivorous demosponge has been found at 8840 m ||Any
 * }

As primary producers
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.

Defenses

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.

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.

Glass sponges produce no toxic chemicals, and live in very deep water where predators are rare.

Predation
Spongeflies, also known as spongillaflies (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.

Bioerosion
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 1 m per year from reefs, creating visible notches just below low-tide level.

Diseases
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.

Collaboration with other organisms
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.

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.

Sponge loop
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.

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.

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.

Sponge holobiont
Besides a one to one symbiotic relationship, it is possible for a host to become symbiotic with a microbial consortium, resulting in a diverse sponge microbiome. 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.

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.

Taxonomy
Carl 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 subkingdom Parazoa ("beside the animals") separated 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.

The phylum Porifera is further divided into classes mainly according to the composition of their skeletons:
 * Hexactinellida (glass sponges) have silicate spicules, the largest of which have six rays and may be individual or fused. The main components of their bodies are syncytia in which large numbers of cell share a single external membrane.
 * Calcarea have skeletons made of calcite, a form of calcium carbonate, which may form separate spicules or large masses. All the cells have a single nucleus and membrane.
 * Most Demospongiae have silicate spicules or spongin fibers or both within their soft tissues. However, a few also have massive external skeletons made of aragonite, another form of calcium carbonate. All the cells have a single nucleus and membrane.
 * Archeocyatha are known only as fossils from the Cambrian period.

In the 1970s, sponges with massive calcium carbonate skeletons were assigned to a separate class, Sclerospongiae, otherwise known as "coralline sponges". However, in the 1980s it was found that these were all members of either the Calcarea or the Demospongiae.

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.

Classes
Sponges were traditionally distributed in three classes: calcareous sponges (Calcarea), glass sponges (Hexactinellida) and demosponges (Demospongiae). However, studies have now shown that the Homoscleromorpha, a group thought to belong to the Demospongiae, has a genetic relationship well separated from other sponge classes. Therefore, they have recently been recognized as the fourth class of sponges.

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:
 * {|class="wikitable"

! Class !! Type of cells !! Spicules !! Spongin fibers !! Massive exoskeleton !! Body form ! Hexactinellida May be individual or fused ||Never ||Never ||Leuconoid ! Demospongiae Made of aragonite if present. ||Leuconoid ! Calcarea May be individual or large masses ||Never ||Common. Made of calcite if present.||Asconoid, syconoid, leuconoid or solenoid ! Homoscleromorpha
 * Mostly syncytia in all species||Silica
 * Single nucleus, single external membrane ||Silica ||In many species ||In some species.
 * Single nucleus, single external membrane||Calcite
 * Single nucleus, single external membrane||Silica ||In many species ||Never ||Sylleibid or leuconoid
 * }

Fossil record

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.

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. The earliest sponge symbionts are known from the early Silurian.

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. 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.

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. 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.

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.

Relationships to other animal groups

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.

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.

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: 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.
 * Homoscleromorpha are most closely related to Eumetazoa;
 * calcareous sponges are the next closest;
 * the other demosponges are evolutionary "aunts" of these groups; and
 * the chancelloriids, bag-like animals whose fossils are found in Cambrian rocks, may be sponges.

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.

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. '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.

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.

In March 2021, scientists from Dublin found additional evidence that sponges are the sister group to all other animals, while in May 2023, Schultz et al. found patterns of irreversible change in genome synteny that provide strong evidence that ctenophores are the sister group to all other animals instead.

Notable spongiologists

 * Stephen Hillenburg

 * Céline Allewaert
 * Patricia Bergquist
 * James Scott Bowerbank
 * Maurice Burton
 * Henry John Carter
 * Max Walker de Laubenfels
 * Arthur Dendy
 * Édouard Placide Duchassaing de Fontbressin
 * Randolph Kirkpatrick
 * Robert J. Lendlmayer von Lendenfeld
 * Edward Alfred Minchin
 * Giovanni Domenico Nardo
 * Eduard Oscar Schmidt
 * Émile Topsent

By dolphins
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.

Skeleton
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, overfishing brought both the animals and the industry close to extinction.

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.

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).

Antibiotic compounds
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.

Other biologically active compounds

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: Sponge


text:
Talk:Sponge

Semi-protected edit request on 13 May 2016
I'm a bit new to editing Wikipedia, but I'm pretty sure this is how you make a request. Correct me if I'm wrong. Regardless I think Porifera incertae sedis should be added to the list of classes. — Preceding unsigned comment added by CltrAltDelicious (talk • contribs) 01:35, 14 May 2016 (UTC)

Semi-protected edit request on 26 December 2016
porifera has radial symmetry 117.192.205.128 (talk) 09:59, 26 December 2016 (UTC)
 * Red information icon with gradient background.svg Not done: please provide reliable sources that support the change you want to be made. DRAGON BOOSTER   ★  10:30, 26 December 2016 (UTC)

add photo of skeletal structure?
hi...new to wikipedia here. I'd like to add a photo showing skeletal structure of a live sponge. There is lots of discussion of spicules and skeletal structure in the article without illustration.

is the photo I would like to add. I'm too new to edit this page.

Jessica Rosenkrantz (talk) 13:24, 28 March 2017 (UTC)

External links modified
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 * Added archive https://web.archive.org/web/20081006123601/http://www.bio.georgiasouthern.edu/Bio-home/Gleason/Ruzicka%26Gleasonfulltext.pdf to http://www.bio.georgiasouthern.edu/Bio-home/Gleason/Ruzicka%26Gleasonfulltext.pdf
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Overview section contains ambuguities related to classification
The Overview section needs improvement regarding the taxonomic relationships among different kinds of sponges. For example, the section refers to Demosponges, sponges with silica, sponges with calcium carbonate skeletons, and Calcareous sponges. It is unclear whether these are four mutually exclusive taxa, whether some of these taxa overlap partially, or whether some of these taxa are completely subsumed under supraordinate taxa. Someone who understands the classification (assuming the classification is not controversial) and also can write clearly should edit the section with eye toward using clear and simple language to indicate the interrelationships among the sponge species groupings noted above. ~PB — Preceding unsigned comment added by Pb1618 (talk • contribs) 05:18, 4 August 2019 (UTC)

Temporal range
The infobox says Ediacaran-recent but the evolution section (and other articles like Cryogenian and Animal) state that sponges were likely around earlier during or after the Marinoan glaciation. Should the infobox be changed, at least with "possible Cryogenian record"? – filelakeshoe (t / c) 🐱 14:21, 29 August 2019 (UTC)

Too technical
Please, experts, tell me what the first sentence is supposed to convey to a scientifically literate nonspecialist? I can tell you: virtually nothing. The introduction to a WP article is supposed to be meaningful to an educated reader. This is hopeless. Will someone please put in some plain English? And thanks. Zaslav (talk) 20:00, 24 March 2020 (UTC)

Is Porifera as sister group to all other Metazoa debate really definitively resolved? (April 2020)
This article seems to cite at least twice the 2017 article which claims that Porifera are the sister group to all Metazoa without any qualifications, as if it were incontrovertibly established fact. (In particular in the first paragraph, rather than say something more neutral, like "some sources suggest that...".) However another 2017 paper https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5664179/ claims that Ctenophores are the sister group to all Metazoa, and this other paper is not mentioned in the article at all, which seems to give an incomplete picture of the discussion of the topic among professional biologists and thus seems to possibly be a disservice to the readers and users of Wikipedia. Neither study can seriously be considered definitive, since neither study uses whole-genome data, and thus do not use all available biological information. (Nor does either study have sufficiently many outgroups, and focusing only on genes transcribed into amino acids completely neglects information which could be contained in pseudogenes about lost characteristics.) Both studies make questionable arguments and use questionable methods, and therefore the questions regarding the phylogenetic relationships among Metazoa can not be considered to be definitely resolved as the introduction of this article seems to purport. — Preceding unsigned comment added by 157.131.142.229 (talk) 20:34, 28 April 2020 (UTC)

 * The section on "relationships to other animal groups" needs to be re-written. Right at the end of this section the article references 2023 research 2023 research that convincingly places ctenophores as the sister group to all other animals including sponges. This research contradicts most of what precedes in the article including the diagrams (though one of those simply omits ctenophores), so it is odd to leave it hanging in this way. The research does not conclude the question of sponge monophyly but is not inconsistent with it and does support parahoxozoan monophyly. Anropa (talk) 08:56, 6 April 2024 (UTC)

"List of sponges" listed at Redirects for discussion
A discussion is taking place to address the redirect List of sponges. The discussion will occur at Redirects for discussion/Log/2020 August 22 until a consensus is reached, and readers of this page are welcome to contribute to the discussion. Steel1943 (talk) 17:16, 22 August 2020 (UTC)

a link in the in the introductory section is incorrect
The first paragraph makes mention of the common ancestor of all animals, which should link to the page on the urmetazoan. Instead it links to the last universal common ancestor. — Preceding unsigned comment added by 2600:1700:CA31:1040:C34:75DA:9EEC:EEE2 (talk) 00:01, 26 October 2020 (UTC)

Sponges do not appear to be sessile after all
At least not exclusively. Apparently it was known that they were facultatively mobile for a while, but this seems to be the first report of mobile sponges in the wild.

https://www.cell.com/current-biology/fulltext/S0960-9822(21)00353-5?_returnURL=https%3A%2F%2Flinkinghub.elsevier.com%2Fretrieve%2Fpii%2FS0960982221003535%3Fshowall%3Dtrue

I'd add this myself but I've mostly just done simple grammar and clarity edits in the past and am not sure of the standards for what to cite and how to cite it, especially for a scientific article. — Preceding unsigned comment added by Gazeboist (talk • contribs) 06:16, 29 April 2021 (UTC)

Sponge loop section; missing original reference in text
Dear editors,

I read the Sponge Wikipedia with great interest as sponge biologist. However, I find it strange that under the 'sponge loop' paragraph, the mechanism is attributed to Rix et al.2018 (ref 71), Rix et al. 2017 (ref 72 and 75? is actually same ref) and Rix et al. 2016 (ref 74). However, the original sponge loop hypothesis is published in Science in 2013 (de Goeij et al. 2013, ref 73), and in fact first proposed in my PhD thesis in 2009. I think it is important to credit the original publication more than wikipedia does now. I am also part of the Rix and colleagues publications and those are great works, but not the original contribution to the sponge loop theory and mechanism. Please let me know how I can help to edit this, since I am not familiar with editing Wikipedia pages. — Preceding unsigned comment added by JM de Goeij (talk • contribs) 13:27, 25 May 2021 (UTC)

Science
Sponges, the members of the phylum Porifera (/pəˈrɪfərə/; meaning 'pore bearer'), are a basal animal clade as a sister of the Diploblasts.[3][4][5][6][7] They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like mesohyl sandwiched between two thin layers of cells. The branch of zoology that studies sponges is known as spongiology.[8] 122.106.252.176 (talk) 09:31, 30 November 2021 (UTC)

Monophyletic or paraphyletic?
Is this phylum monophyletic or paraphyletic? The article dosen't seem to give a clear answer. 2001:1970:4885:CC00:0:0:0:767B (talk) 21:44, 11 April 2023 (UTC)

 * I am not sure as phylogenetics showing it as paraphyletic have existed for a long time and even as recently as a week ago. at the same time there are many that just consider them monophyletic instead. for the time it is probably best to stick with the monophyletic model while showcasing the paraphyletic alternatives until a better consensus is established. Zacharyman1mil (talk) 23:02, 30 April 2023 (UTC)

Parazoan, Myriazoan, or neither?
On Taxonomy/Porifera, poriferans have been placed in either Parazoa or Myriazoa. gives sources for both placements. I think the taxobox should give the most specific placement, so I'm restoring Animalia for now. jlwoodwa (talk) 01:03, 8 August 2023 (UTC)

Role of Bacterial Endosymbionts
I just wanted to add something I recently read wherein cryptic (AKA hidden) encrusted sponges were found to absorb high quantities of dissolve organic matter. This is much smaller organic "food" than was previously thought and the authors indicate that the role of endosymbiont bacteria (they do not specify which) play heavily into this. "We conclude ... 3 sponge-microbe associations are ... 'dissolved organic matter (DOM)-feeders'" Sponges are very important for carbon cycling and dissolved carbon re-uptake in reef ecosystems. https://doi.org/10.3354/meps07403] Spacefatty (talk) 17:53, 23 October 2023 (UTC)

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/

title: Talk:Sponge


text:
Wikipedia talk:Blocked IPs

Something to keep in mind is that a huge number of IPs that get blocked are dynamic IPs for home broadband connections. A knowledgeable user will kill their DHCP lease and get a new one for a different IP, thus avoiding the block; less savvy users who get assigned these IPs are the ones who suffer, especially with long-term blocks. If you then ban a subnet in order to get one wily bug you are most likely nuking the entire neighborhood along with it. Perusal of the blocklist reveals many people blocked for trifling reasons, not defacement.

Brion VIBBER, Lee Daniel Crocker, Tim Shell, Larry_Sanger, Koyaanis Qatsi appear to be the de facto editorial/censor board. Should there be links here to the "reasons why"? Not necessarily for disputes, but to establish an accountability trail.

That will matter later, I'm sure.

Well, I've blocked IPs too, so go ahead and call me a "censor," but it's not something I did for political reasons so the term would be inappropriate. Anyway, I can't speak for anyone else, but I blocked people who were consistently adding patent nonsense. One of the IPs also adds his share of relevant info, as Axel Boldt (& LDC) pointed out, so I took him off the list. Originally I thought the ban was just for a short time, like 30 minutes or something, but it turns out the blocks aren't lifted automatically, and I just found out how to lift them manually. So that's that. There's been discussion of the ways that IP blocking should and should not be used on wikipedia-l, if you're interested. That's also archived at nupedia, if you'd care to take a look at what's been said already. Koyaanis Qatsi

 * See examples of discussion in VANDALISM IN PROGRESS and Talk:VANDALISM IN PROGRESS; after I've overhauled the software a bit, there will be a comment field in the IP-blocking tool for a quick explanatory note there as well. Note also that anyone with the ability to block IPs also has the ability to unblock IPs -- and everyone can see who did which. (Larry, by the way, is no longer involved in Wikipedia.) Brion VIBBER, Tuesday, April 2, 2002

---

Thanks, that is very helpful, especially for newcomers learning the ropes. Without Larry it will be interesting to see how "the ability to block IPs" and "the ability to unblock IPs" will be managed. Presumably some political process for deciding this must evolve... "electing sherrifs" or whatever... although probably not soon, as we are still figuring out editorial stances on some very fundamental stuff.

I notice Blocked IPs doesn't log all changes in the history, only the manual edits. Is that a bug or a feature? (I think it's a bug.) Koyaanis Qatsi
 * All the automatic logs are like this (see also log:Uploads and log:Page Deletions), but the pages in the log: namespace aren't manually editable. Keeping histories of automatic changes there is of dubious value -- that's exactly what the logs themselves contain -- but the blocked IP log seems to be a different situation, and perhaps should be handled differently. Magnus, you wrote it, what do you think? Brion VIBBER

I just blocked 212.138.47.12 after this person deleted content from at least three Islam related articles (Qur'an, Dialect, & Muslim language). It would be nice if I could have stated this as part of the blocking process and for this article to show up in Recent Changes with my explanation automatically in an edit comment. This is an issue of policy that I will take up with the wikipedia mailing list. BTW it also would be nice to be able to more easily track what these vandals have done so that the other damage this person has probably done can be fixed -- this is another issue that will have to be taken to the list. --maveric149, Saturday, April 20, 2002

 * Try http://www.wikipedia.com/wiki/special:contributions&theuser=212.138.47.12 (except the minor edits won't be shown). AxelBoldt
 * Cool! Now it would be nice to have that link generated automatically whenever an IP is added to the list. --maveric149

 * For the record, Dialect wasn't exactly Islam-related (though 212.138.47.15's edits to Muslim language spoke vaguely of Arabic dialects; it's likely that this is the same person with either a dynamic IP or using two computers on the same network)... In Dialect, he/she removed two things: a contextless quote in Italian (which I found reasonable to remove, though it should have been noted in the talk page or at least an edit summary), and a cross-reference to idiolect (for which there is no article yet). I restored the xref to idiolect, since that's something that should be noted. However, I agree that their changes to Qur'an and Muslim language were unreasonable -- large amounts of information deleted.

 * (Oh, and it's not important that minor edits won't be shown, since anonymous users can't make minor edits!) Brion VIBBER

- I placed 210.131.75.142 back on the blocked list due to the fact that within 10 minutes of me removeing this IP from the list this person started doing the same thing again -- creating numerous blank pages are messing with the formatting of articles. Last night I warned this person again and this morning I see that my warning was unheaded, so I blocked the IP. --maveric149

I am concerned that we are blocking IPs for valid reasons and then not going back after an appropriate period of time to see if particular IP blocks can be lifted. Some of the infractions listed don't seem to warrent more than a week long IP block (others seem to have only warrented a couple of days and yet others arguably seem to only warrent warnings of an IP block). For example, 195.92.67.66 was blocked for placing an ad on a wikipedia page and for inputing an obsenity within an article. I personally would have warned the person a couple of times especially since this a person using this same IP made a great contribution to the Insurance article (which makes me suspect that the IP had changed hands, is the IP of a public computer, or a shared IP on a network -- possibly blocking a valued contributor). Another example that worries me is the case of 144.134.241.96 whose first and only "contribution" was to create Maurice Wilkins which only had obcenities for content. Who knows, this person may have just been experimenting with wiki wiki not knowing what this project was about. Unfortunetely we probably will never know because this person's IP was blocked the next day. Does limited damage like this warrent an IP block? How about first trying to give a friendly warning, then a stern warning, and then a final warning before blocking an IP? It would be nice if the wikiware gods provided us with a warning feature to automate the process. Any thoughts? --maveric149

Because it is so easy to become a member whit a username and no ip is shown, how to you deal whit that? giskart 09:44 Oct 20, 2002 (UTC)

 * I think Brion is working on a way to allow Admins to block users by IP but without exposing the IP at least to non-Admins. But this really hasn't been a problem yet. --mav

 * I can't honestly say I've been working on it, but it is on my to-do list. As mav says, it's not really something that's come up; I don't think we've ever had a persistent vandal who bothered to log in. Trolls and jerks who refuse to work NPOV, occasionally, but not outright vandals. --Brion 20:08 Oct 20, 2002 (UTC)

I notice the first few entries in the logfile are dated December 31, 1969. Tualha 14:10, 19 Dec 2003 (UTC)

title: Wikipedia talk:Blocked IPs


text:
Pannonia

Pannonia was a province of the Roman Empire bounded on the north and east by the Danube, coterminous westward with Noricum and upper Italy, and southward with Dalmatia and upper Moesia. Pannonia was located in the territory that is now western Hungary, western Slovakia, eastern Austria, northern Croatia, north-western Serbia, northern Slovenia, and northern Bosnia and Herzegovina.

Background
In the Early Iron Age, Transdanubia was inhabited by the Pannonians or Pannonii, a collection of Illyrian tribes. The Celts invaded in the Late Iron Age and Gallo-Roman historian Pompeius Trogus writes that the Celts met with heavy resistance from the locals and were not able to overrun the southern part of Transdanubia. Some tribes advanced as far as Delphi, with the Scordisci settling in Syrmia (279 BC) upon being forced to withdraw. The arrival of the Celts in Transdanubia disrupted the flow of amber from the Baltic Sea region, through the Amber Road, to the Illyrians. They founded many villages. Those that held prominent economic significance developed into oppida. Independent tribes minted their own coins with the faces of their leaders. These were at first modelled on Macedonian and, later, Roman currency.

Upon the Scordisci's withdrawal and settlement, they and the Dardani (in Dardania) both became strong powers opposing each other. The Dardani consistently raided Macedon and developed close ties to Rome. Philip V, who was a vehement enemy of the Dardani, allied with the Scordisci and in 179 BC persuaded the Bastarnae (at the Danube Delta) to break into Italy and subdue them on the way. Despite Philip's defeat at the hands of the Romans in 197 BC and the failure of the Bastarnae, in this time the Dardani's power crumbled under the pressure from the Macedonians and Scordisci. Finally, Perseus annihilated them, giving way to hundred years of Scordisci hegemony in the Balkans. During this time, the tribe started raiding the new province of Macedonia, and —Strabo says— expanded as far as Paeonia, Illyria and Thrace.

Aquileia's foundation in 181 BC was the first step towards the Roman takeover of Pannonia. The town functioned as the starting station of the Amber Road and the starting point of attacks in that direction. The Scordisci, in alliance with the Dalmatae were in armed conflict with the Romans as early as 156 BC and 119 BC. In both wars, the Romans failed to take Siscia (now Sisak, Croatia), which laid in a key position. After these setbacks, Rome instead turned towards Noricum which had both iron and silver mines.

As part of a new Celtic migration wave at the end of the 2nd century BC, the Boii left Northern Italy and established themselves as an important power at the Danube. According to the Posidonius's record of the Cimbri migration (preserved by Strabo), they were first repulsed by the Boii, then by the Scordisci, and then by the Taurisci towards the Helvetii. This describes the balance of power in the region. In the early 1st century BC, the Dacians emerged as a new dominant power. While their hold on the area between the Danube and the Tisza river was loose, they had considerable influence in the territories beyond. In 88 BC, Scipio Asiaticus (consul 83 BC) defeated the Scordisci so badly that they retreated to the eastern part of Syrmia. Taking advantage of this situation, the Dacian king Burebista vanquished them sometime between 65 and 50 BC, and subsequently the Boii and the Taurisci too. Thanks to the ebb of these entities, several local tribes regained their independence and influence. In context of Mithridates VI Eupator's unfulfilled plan to invade Italy from the north (64 BC), the territory he was to cross is noted to have belonged to the Pannonians. Immediately after Burebista's death (c. 44 BC), Dacia's kingdom dissolved too, leaving no entity in the region that Rome would make allowances for.

Roman conquest
The Pannonians were driven into conflict due to their support of the Dalmatae in their strife against Rome, but weren't long-term and known enemies. The tribes north of the Drava River didn't participate in nor this, nor the subsequent fights. In 35 BC, Octavian led a campaign against the Iapydes and the Pannonians, in which he captured Siscia in a month-long siege and occupied a large part of the Sava River valley. This was in accordance with Caesar's plan of creating a base for an invasion of Dacia, not realized due to his assassination. However, Octavian only used the hoax of the "Dacian threat" as a pretense to gain control over a large amount of land in the Second Triumvirate.

In 15 BC, the future-emperor Tiberius defeated the Scordisci, forcing them to become allies. This was in response to Pannonian and Scordisci incursions the previous year. The following events were part of the Roman Empire's efforts to reach the Danube and are sometimes known thematically as Bellum Pannonicum.

In 14 BC, the Pannonians rose up. Vipsanius Agrippa was sent to the region after another rebellion in 13 BC. After his death the following year, the campaign was taken over by Tiberius, who celebrated his triumph in 11 BC. The province of Illyricum was established between the Sava and the Adriatic Sea. In 10 BC, Tiberius returned to quell a new uprising of the Pannonians and Dalmatae. After winning in 9 BC, he sold the youth of the Breuci and Amantini as slaves in Italy and held an ovation. His operations between 12 and 9 BC included constant expeditions into territories north of the Drava and almost certainly brought the whole Transdanubia under Roman control even though there's no direct evidence to that.

"Through Tiberius Nero, then my stepson and legate, I brought under Roman authority Pannonian peoples which no Roman army had approached before I became princeps and advanced the boundaries of lIIyricum to the bank of the Danube."Pannonia was invaded by the Dacians in 10 BC. The Romans launched campaigns through the Danube in order to secure it as the imperial border and defend the threatened new land. Lucius Domitius Ahenobarbus's (consul 16 BC) operation in 1 AD extended as far as the Elbe. In 10 AD, Cornelius Lentulus Augur was able to debar not just the Dacians, but also the Sarmatians "from access to the Danube", says Florus. Locally more important was the offensive of Marcus Vinicius against the tribes east of the Danube Bend, showing an intent of "monopolizing" the Northern Transdanubian region politically. The last decade of the century saw the Marcomanni under their king Maroboduus—settling north of Pannonia. Augustus planned a two-sided attack on them, with one army approaching their territory from the Rhine and another one under Tiberius crossing the Danube at Carnuntum.

Before witnessing any result, Tiberius had to rush back in 6 AD and face a new uprising. The unfolding Bellum Batonianum lasted for three years. The Breuci (under Bato the Breucian) and Daesitiates (under Bato the Daesitiate and Pinnes) took the leading role, while the tribes north of the Drava stayed out again. The insurgents attempted to invade Italy and Macedonia, but due to their lack of success they united to besiege Sirmium (now Sremska Mitrovica, Serbia). There, Caecina Severus defeated the insurgents, who retreated into the Fruška gora Mountains. He annihilated them the following year when they tried to intercept him on his way to join Tiberius at Siscia. Tiberius competently initiated a scorched-earth policy which was unsatisfactory for Augustus, who sent more generals, including Germanicus and Plautius Silvanus (consul 2 BC) to the war theatre. A capitulation was forced out in 8 AD, and Bato the Breucian delivered Pinnes to the Romans, becoming a vassal king of his tribe. However, the revolt flared up once again as the Daesitiates captured and executed Bato the Breucian, and persuaded his people to continue the resistance. Silvanus reconquered them and ousted Bato the Daesitiate into the Dinaric Alps, where he laid down arms in 9 AD.

Consolidation and establishment of administration
Illyricum was divided into Dalmatia (initially called Illyricum Superius) and Pannonia (initially Illyricum Inferius) in 8 or 9 AD.

According to Suetonius, with the Bellum Batonianum, Tiberius finally defeated all peoples between the Danube and the Adriatic Sea. No Illyrian resistance is known after this, not due to the natives' compliance with the new status quo, but due to their extreme exhaustion. The eligible Pannonian youth were conscripted and commanded to other provinces. The communities taking part in the uprising were afterward relocated and organized into civitates under military supervision.

The military occupation of Pannonia may have been carried out in gradual steps. The Romans felt it necessary to resettle certain tribes to the territory of the peoples north of the Drava, which, for them, had no economic, but strategic significance. Augustus formed a kind of alliance where the Romans would act as supervisors, and it was not until his death (14 AD) that legions would be moved over from South Pannonia.

The second emperor Tiberius ((r. 17 – 37 AD)) founded multiple coloniae in the province and developed its road network. However, due to these land's unsuitability for cultivation, it was a hard task to persuade veterans to comply with settling there, and he had to silence a mutiny right when assuming power. He sent his son Drusus Julius Caesar to create tranquility and depose Maroboduus, who needed Roman support for his war against Arminius. This ultimately caused the rise of Vannius (20 AD), who ruled over an extended realm.

It was Claudius ((r. 41 – 54 AD)) who finished Pannonia's occupation and began to construct of the local limes. Systematic integration into the Empire accompanied by the establishment of settled Roman life progressed subsequently. In 50 AD, Vannius was overthrown by Vangio and Sido, who enjoyed the emperor's support. By this date, the nomadic Sarmatian population of the Iazyges had taken possession of the Danube–Tisza Interfluve, helping the Romans by being a buffer state against the dangerous Dacians.

At first, the primary goal of the Roman administration was the conclusion of the barbarian conflicts outside the province. In Nero's time ((r. 54 – 68 AD)) as many as 100,000 barbarians were moved from Pannonia to Moesia by Plautius Silvanus Aelianus, and 50,000 may have been settled in Pannonia by Tampius Flavianus. During his important governorship, money began to circulate in the Barbaricum and the line of the limes was stabilized.

Under the Flavians
The Year of the Four Emperors (69 AD) passed with peace in Pannonia. Flavianus declared for Vespasian and led his legions to Italy against Vitellius. Vespasian ((r. 69 – 79 AD)) invested greatly in the construction of the limes. Discarding the Augustan strategy where the legions' role was with maintaining order in their provinces, the Flavian emperors continually moved them to the border. This way they were prevented from interfering in domestic policy, while the conquests were already pacified. Systematic circulation of money in the region situated north of the Drava shows that by this time Roman civilization had firmly taken root there.

Domitian's ((r. 81 – 96)) emperorship saw expensive wars with the barbarians, as a result of which the military emphasis shifted to the Danube frontier. At the end of 85 or the beginning of 86, the reemerging Dacians under Decebalus raided Moesia, killing its governor and eradicating a legion. After a brief stay, Domitian left Cornelius Fuscus to deal with the situation. After clearing the province of raiders, Fuscus undertook a disastrous campaign and lost his life (86). Finally, in 88, Tettius Julianus defeated Decebalus and the sides agreed to make peace. Vangio and Sido were most likely dead by now, the Marcomanni and Quadi denied vassal duties. When the emperor's punitive expedition (partially sent through Dacian territory) was repelled in 89, he—despite the damages suffered—settled for mild terms with Decebalus, instead committing his forces elsewhere. In the same year, he held his triumphs over the Dacians and Chatti, but not over the disloyal Danubian Germans. When the Romans started supporting the Lugii against them, they made a pact with the Iazyges. This produced another war, almost completely unknown except for another catastrophe and destruction of a legion at the hands of the nomads. In 92 or 93, he finished the war, but held only an ovation, indicating he probably had further plans in Pannonia.

Under the Antonines
We hear of war with the Danubian Germans again under Nerva ((r. 96 – 98)). Between 103 and 107, Trajan ((r. 98 – 117)) executed the division of the province into Pannonia Inferior and Pannonia Superior. This allowed the Empire to better combat the radically different Germanic and Sarmatian tribes. While Superior had most urbanized areas and a shorter frontier with three legions, Inferior contained one municipium and one legion, virtually being a border zone. Under his reign, the placement of garrison and the main lines of commerce became permanent.

The creation of Roman Dacia had a great effect on Pannonia. In Trajan's Dacian Wars, the Iazyges allied with the Romans, seeking to retain Oltenia where they were expelled by Decebalus. A brief confrontation in 107 was resolved Hadrian, then-governor of Pannonia Inferior and it may have been agreed that the nomads would instead take possession of the region between the Tisza and the Apuseni Mountains, not incorporated into the new province. However, taking advantage of Trajan's death and the preoccupation of the Empire with the Parthian war, they joined forces with the relative Roxolani and attacked again in 117, to which Dacia's governor, Julius Quadratus Bassus fell victim. Hadrian ((r. 117 – 138)) traveled to the spot and invested Marcius Turbo as governor of both Dacia and Pannonia Inferior to defeat the barbarians. The Roxolani were pacified first. Turbo's authorization was over in 119 as Iazyx peace envoys appeared in Rome. The postal connection between the two provinces through the Danube–Tisza Interfluve—which aggravated relations with the Sarmatians—was completed.

War with the Quadi broke out again in the last years of Hadrian's reign, which his adopted son and joint governor of the Pannonian provinces, Aelius Caesar successfully handled until he died in 138. Command of Pannonia Superior was taken over by Haterius Nepos, who ended the war with a Roman victory, becoming the last person to be awarded with ornamenta triumphalia. Under Antoninus Pius's ((r. 138 – 161)) quiet reign, some coins were issued propagating not the ending of a new campaign but the reestablishment of foederatus relationship by the investiture of a new Quadi king. Discharges and detachments of troops happened.

Findings of hoards of coins likely buried during the rule of Marcus Aurelius ((r. 161 – 180)) evidence turmoil due to barbarian attacks. Large-scale population movements in Northern and Eastern Europe related to the Goths highly endangered Rome's clients, who wanted the Empire to give its lands to settlement and extend its protection over the tribes. Rome was unwilling to grant these requests. The Romans may not have been aware of the dangerous situation at the start of the Parthian war of Lucius Verus because they sent a whole legion and many vexillationes away from Pannonia. It is thanks to the diplomatic efforts made by regional governors that tensions were eased until the dispatched forces could get back. When the threat became fully clear, Marcus even raised new legions. The first attack came in the winter of 166-167, from the Lombards and Ubii, between Brigetio and Arrabona. It was quickly repulsed by two auxiliary units. Cassius Dio tells of a legation of 11 tribes led by the Marcomanni subsequently petitioning the governor of Pannonia Superior, Iallius Bassus to concede. This may have been the last attempt at making peace, as next, a barbarian coalition formed to fight Rome.

In 168, Marcus and Verus returned to Aquileia and set up their base there. The Marcomanni and Quadi broke through the border and the Alps' crosses, besieging the city and burning the small town of Opitergium. The peak of the Antonine Plague in the peninsula was at this time, causing Verus's death. The next years' heavy fighting resulted in the death of governor of Moesia Superior and Dacia Claudius Fronto and praetorian prefect Macrinius Vindex. Claudius Pompeianus and future-emperor Pertinax returned part of the spoils taken by the enemy and led the offensive starting from 172. Against severe losses, the Romans forced first the Quadi, then the Marcomanni to surrender (172-173), while the military emphasis shifted to the Iazyges. Despite the winter incursion of the Iazyges was crushed (173-174), the Quadi overthrew their Roman-installed king and started to support the nomads. While the two nations tried to negotiate, Marcus eventually defeated both of them in separate campaigns.

The second phase of the war started in 177. The attacking barbarians were kept in check, with Marcus and his son, the newly acclaimed Commodus ((r. 177 – 192)) coming to Pannonia. A decisive campaign by Tarrutenius Paternus in 179 convinced the Iazyges to make peace. In the same year, the land of the Danubian Germans was occupied by a force Cassius Dio claims to be 40,000 men—the number of soldiers stationed in Pannonia Inferior and Pannonia Superior combined. Control over tribes was taken over by prefects. Valerius Maximianus, born in Pannonia, was an important general here. Any possible plans with the creation of two new provinces—Marcomannia and Sarmatia—were aborted after the death of Marcus in 180. Commodus returned to the old border and client system, to which new residents were seemingly willing to join. As the barbarians pillaged during the war, taking cattle and captives away en masse, the destruction and loss of life in Pannonia was huge.

Commodus vigorously started to strengthen the limes with new fortifications. Minor raids on the province continued to occur, prompting a third campaign over the Danube at about. This campaign was smaller, and its leader, Tigidius Perennis, achieved a victory. Another victorious expedition was conducted in 188.

Under the Severans
During the Year of the Five Emperors (193), no attack was made on Pannonia. According to Herodian, Septimius Severus ((r. 193 – 211)) calmed the barbarian tribes via negotiations before marching off his troops to Italy and gaining the throne. In the coming years, the arrival of foreign groups led to new conflicts, but these were centered on Dacia and Pannonia only experienced collateral effects. The Severans' rule was supported by the Pannonian military and other provinces of the collective "Illyricum" region, which became politically important. In 202, a thorough visit to Pannonia by the imperial house was organized. Partly during this tour and throughout Severus' reign, the province benefited from many constructions. The road network was fully repaired, civilian and military buildings were inaugurated, military camps were improved and cities were protected with walls thus increasing their rank.

Administration
Pannonia Superior was under the consular legate, who had formerly administered the single province, and had three legions under his control. Pannonia Inferior was at first under a praetorian legate with a single legion as the garrison; after Marcus Aurelius, it was under a consular legate, but still with only one legion. The frontier on the Danube was protected by the establishment of the two colonies Aelia Mursia and Aelia Aquincum by Hadrian.

Under Diocletian and his successors, a fourfold division of the country was made:
 * Pannonia Prima in the northwest, with its capital in Savaria, it included Pannonia Superior and the major part of Central Pannonia between the Raba and Drava,
 * Pannonia Valeria in the northeast, with its capital in Sopianae, it comprised the remainder of Central Pannonia between the Raba, Drava and Danube,
 * Pannonia Savia in the southwest, with its capital in Siscia,
 * Pannonia Secunda in the southeast, with its capital in Sirmium

Diocletian also moved parts of today's Slovenia out of Pannonia and incorporated them in Noricum. In 324 AD, Constantine I enlarged the borders of Roman Pannonia to the east, annexing the plains of what is now eastern Hungary, northern Serbia and western Romania up to the limes that he created: the Devil's Dykes.

In the 4th-5th century, one of the dioceses of the Roman Empire was known as the Diocese of Pannonia. It had its capital in Sirmium and included all four provinces that were formed from historical Pannonia, as well as the provinces of Dalmatia, Noricum Mediterraneum and Noricum Ripense.

Loss
In the 4th century, the Romans (especially under Valentinian I) fortified the villas and relocated barbarians to the border regions. In 358 they won a great victory over the Sarmatians, but raids didn't stop. In 401 the Visigoths fled to the province from the Huns, and the border guarding peoples fled to Italia from them, but were beaten by Uldin in exchange for the transferring of Eastern Pannonia. In 433 Rome completely handed over the territory to Attila for the subjugation of the Burgundians attacking Gaul.

After Roman rule

During the Migration Period in the 5th century, some parts of Pannonia were ceded to the Huns in 433 by Flavius Aetius, the magister militum of the Western Roman Empire. After the collapse of the Hunnic empire in 454, large numbers of Ostrogoths were settled by Emperor Marcian in the province as foederati. The Eastern Roman Empire controlled southern parts of Pannonia in the 6th century, during the reign of Justinian I. The Byzantine province of Pannonia with its capital at Sirmium was temporarily restored, but it included only a small southeastern part of historical Pannonia.

Afterwards, it was again invaded by the Avars in the 560s, and the Slavs, who first may settled c. 480s but became independent only from the 7th century. In 790s, it was invaded by the Franks, who used the name "Pannonia" to designate the newly formed frontier province, the March of Pannonia. The term Pannonia was also used for Slavic polity like Lower Pannonia that was vassal to the Frankish Empire.

Cities and auxiliary forts
The native settlements consisted of pagi (cantons) containing a number of vici (villages), the majority of the large towns being of Roman origin. The cities and towns in Pannonia were:

Now in Austria:
 * Carnuntum (Petronell, Bad Deutsch-Altenburg)
 * Vindobona (Vienna)

Now in Bosnia and Herzegovina:
 * Saldae (Brčko)
 * Serbinum or Servitium (Gradiška)
 * Castrum and Canabea (Doboj)

Now in Croatia:
 * Ad Novas (Zmajevac)
 * Andautonia (Ščitarjevo)
 * Aqua Viva (Petrijanec)
 * Aquae Balisae (Daruvar)
 * Certissa (Đakovo)
 * Cibalae (Vinkovci)
 * Cornacum (Sotin)
 * Cuccium (Ilok)
 * Halicanum (Sveti Martin na Muri)
 * Iovia or Iovia Botivo (Ludbreg)
 * Marsonia (Slavonski Brod)
 * Mursa (Osijek)
 * Siscia (Sisak)
 * Teutoburgium (Dalj)

Now in Hungary:
 * Ad Flexum (Mosonmagyaróvár)
 * Ad Mures (Ács)
 * Ad Statuas (Vaspuszta)
 * Ad Statuas (Várdomb)
 * Alisca (Szekszárd)
 * Alta Ripa (Tolna)
 * Aquincum (Óbuda, Budapest)
 * Arrabona (Győr)
 * Brigetio (Szőny)
 * Caesariana (Baláca)
 * Campona (Nagytétény)
 * Cirpi (Dunabogdány)
 * Contra-Aquincum (Budapest)
 * Contra Constantiam (Dunakeszi)
 * Gorsium-Herculia (Tác)
 * Intercisa (Dunaújváros)
 * Iovia (Szakcs)
 * Lugio (Dunaszekcső)
 * Lussonium (Dunakömlőd)
 * Matrica (Százhalombatta)
 * Morgentianae (Tüskevár (?))
 * Mursella (Mórichida)
 * Quadrata (Lébény)
 * Sala (Zalalövő)
 * Savaria (Szombathely)
 * Scarbantia (Sopron)
 * Solva (Esztergom)
 * Sopianae (Pécs)
 * Ulcisia Castra (Szentendre)
 * Valcum (Fenékpuszta)

Now in Serbia:
 * Acumincum (Stari Slankamen)
 * Ad Herculae (Čortanovci)
 * Bassianae (Donji Petrovci)
 * Bononia (Banoštor)
 * Burgenae (Novi Banovci)
 * Cusum (Petrovaradin)
 * Graio (Sremska Rača)
 * Onagrinum (Begeč)
 * Rittium (Surduk)
 * Sirmium (Sremska Mitrovica)
 * Taurunum (Zemun)

Now in Slovakia:
 * Gerulata (Rusovce)

Now in Slovenia:
 * Celeia (Celje)
 * Neviodunum (Drnovo)
 * Poetovio (Ptuj)

Economy
The country was fairly productive, especially after the great forests had been cleared by Probus and Galerius. Before that time, timber had been one of its most important exports. Its chief agricultural products were oats and barley, from which the inhabitants brewed a kind of beer named sabaea. Vines and olive trees were little cultivated. Pannonia was also famous for its breed of hunting dogs. Although no mention is made of its mineral wealth by the ancients, it is probable that it contained iron and silver mines.

Slavery
Slavery held a less important role in Pannonia's economy than in earlier established provinces. Rich civilians had domestic slaves do the housework while soldiers who had been awarded with land had their slaves cultivate it. Slaves worked in workshops primarily in western cities for rich industrialist. In Aquincum, they were freed in a short time.

Religion
Pannonia had sanctuaries for Jupiter, Juno and Minerva, official deities of empire, and also for old Celtic deities. In Aquincum there was one for the mother goddess. The imperial cult was also present. In addition, Judaism and eastern mystery cults also appeared, the latter centered around Mithra, Isis, Anubis and Serapis.

Christianity began to spread inside the province in the 2nd century. Its popularity didn't decrease even during the big persecutions in the late 3rd century. In the 4th century, basilicas and funeral chapels were built. We know of the Church of Saint Quirinus in Savaria and numerous early Christian memorials from Aquincum, Sopianae, Fenékpuszta, and Arian Christian ones from Csopak.

Legacy
The ancient name Pannonia is retained in the modern term Pannonian plain.

title: Pannonia