Spined pygmy shark
Encyclopedia
The spined pygmy shark (Squaliolus laticaudus) is a species
of dogfish shark
in the family
Dalatiidae
found widely in all oceans. With a maximum known length of 28 cm (11 in), it is one of the smallest living sharks. This shark has a slender, cigar-shaped body with a sizable conical snout, a long but low second dorsal fin
, and an almost symmetrical caudal fin. It and its sister species S. aliae
are the only sharks with a spine on the first dorsal fin and not the second. Spined pygmy sharks are dark brown to black, with numerous bioluminescent
organs called photophore
s on their ventral surface. The shark is believed to use these photophores to match ambient light conditions and thus disguise its silhouette from predators.
Usually inhabiting nutrient-rich waters over upper continental and insular slopes, the spined pygmy shark feeds on small bony fishes and squid
. Like its prey it is a diel vertical migrator
, spending the day at close to 500 m (1,640.4 ft) deep and moving towards a depth of 200 m (656.2 ft) at night. Reproduction is presumably aplacental viviparous, with female giving birth to litters of 4 pups. This diminutive shark has no economic value. The International Union for Conservation of Nature (IUCN) has assessed this species as of Least Concern
, as it faces little threat from commercial fisheries and has a wide distribution.
Steamer
Albatross
. It was described by American ichthyologists Hugh McCormick Smith
and Lewis Radcliffe
in a 1912 paper for the scientific journal
Proceedings of the United States National Museum, based on two specimens collected in Batangas Bay
, south of Luzon
in the Philippines
. One of these, a 15 cm (5.9 in) long adult male, was designated the type specimen.
Smith and Radcliffe coined the new genus
Squaliolus for this shark, and gave it the specific epithet laticaudus, from the Latin
latus meaning "broad" or "wide", and cauda meaning "tail". The spined pygmy shark may also be referred to as the dwarf shark or the bigeye dwarf shark. Based on similarities in their claspers (male intromittent organ
s), the closest relative of the spined pygmy shark and the related S. aliae is thought to be the pygmy shark
(Euprotomicrus bispinatus).
, it occurs off Bermuda
, the United States
, Suriname
, southern Brazil
, and northern Argentina
in the west, and off northern France
, Madeira
, Cape Verde
, and the Azores
in the east. In the Indian Ocean
, this species has only been recorded off Somalia
. In the Pacific Ocean
, it is found off southern Japan
, Taiwan
, and the Philippines
. The spined pygmy shark is found at depths of 200–500 m (656.2–1,640.4 ft) and seldom approaches the surface, unlike the related pygmy shark and cookiecutter shark
(Isistius brasiliensis). This shark prefers areas of high biological productivity over upper continental and insular slopes. It may also be found over outer shelves
, but avoids central ocean basins. The range of this species does not overlap that of the pygmy shark, which has a similar ecology, and is also largely separate from that of the cookiecutter shark.
s are minute and uniform in size.
The two species of Squaliolus are the only sharks with a spine on the first dorsal fin
but not the second. The spine is sexually dimorphic, being typically exposed in males and enclosed by skin in females. The first dorsal fin is tiny and originates over the trailing margin of the pectoral fins. The second dorsal fin is low, with a base twice as long as that of the first, and originates over the anterior half of the pelvic fin bases. The pectoral fins are short and triangular, with the rear margin slightly curved. The pelvic fins are long and low, and there is no anal fin. The caudal peduncle is slender and laterally expanded into weak keels. The caudal fin is broad and paddle-like, with the upper and lower lobes of similar size and shape, and a deep notch in the trailing margin of the upper lobe.
The dermal denticles are flat and blocky, not elevated on stalks or bearing marginal teeth. The coloration is dark brown to black, with light fin margins. The underside is densely carpeted by light-emitting photophore
s, which extend to the tip of the snout and around the eyes and nostrils, and thin to almost non-existent on the back. This species has on average only 60 vertebrae, the fewest of any shark.
Idiacanthus, the lanternfish
Diaphus
, and the bristlemouth Gonostoma) and squid
(including members of the genera Chiroteuthis
and Histioteuthis
). Catch records suggest that the spined pygmy shark follows its prey on their diel vertical migration
s, spending the day close to a depth of 500 m (1,640.4 ft) and ascending towards a depth of 200 m (656.2 ft) at night. The ventral photophores of the spined pygmy shark have been theorized to function in counter-illumination
, a form of camouflage in which the shark disguises its silhouette from would-be predators by matching the ambient light welling down from above. There is no evidence that this shark swallows its shed teeth like the pygmy and cookiecutter sharks.
The spined pygmy shark is aplacental viviparous like the rest of its family, with the developing embryo
s being sustained by a yolk sac
until birth. Adult females have two functional ovaries that may each contain up to 12 mature eggs. However, the actual litter size is much smaller; a pregnant female caught off southern Brazil in 1999 contained four near-term pups. The young are born at 9–10 cm (3.5–3.9 in) long. Males mature sexually at a length of 15 cm (5.9 in), and females at a length of 17–20 m (669.3–787.4 in). The spined pygmy shark was widely considered to be the smallest living shark species until the discovery of the dwarf lanternshark
(Etmopterus perryi), though the pygmy ribbontail catshark
(Eridacnis radcliffei) is also known to mature at a size comparable to these two species. Whether one of these sharks is definitively smaller than the others cannot yet be stated with certainty, because of the difficulties involved in assessing reproductive maturity in sharks.
of trawl fisheries, but are generally too small to be captured. In light of its wide distribution and the absence of substantial threats from human activity, the International Union for Conservation of Nature (IUCN) has assessed this species as of Least Concern
.
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
of dogfish shark
Squaliformes
Squaliformes is an order of sharks that includes about 97 species in seven families.Members of the order have two dorsal fins, which usually possess spines, no anal fin or nictitating membrane, and five gill slits. In most other respects, however, they are quite variable in form and size...
in the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
Dalatiidae
Dalatiidae
Dalatiidae is a family of sharks in the order Squaliformes, commonly known as kitefin sharks . Members of this family are small, under long, and are found worldwide. They have cigar-shaped bodies with narrow heads and rounded snouts...
found widely in all oceans. With a maximum known length of 28 cm (11 in), it is one of the smallest living sharks. This shark has a slender, cigar-shaped body with a sizable conical snout, a long but low second dorsal fin
Dorsal fin
A dorsal fin is a fin located on the backs of various unrelated marine and freshwater vertebrates, including most fishes, marine mammals , and the ichthyosaurs...
, and an almost symmetrical caudal fin. It and its sister species S. aliae
Smalleye pygmy shark
The smalleye pygmy shark is a little-known species of dogfish shark in the family Dalatiidae, found in water deep near Japan, the Philippines, and Australia. It migrates vertically daily, spending the day in deep water and the night in shallower water. One of the smallest shark species, the...
are the only sharks with a spine on the first dorsal fin and not the second. Spined pygmy sharks are dark brown to black, with numerous bioluminescent
Bioluminescence
Bioluminescence is the production and emission of light by a living organism. Its name is a hybrid word, originating from the Greek bios for "living" and the Latin lumen "light". Bioluminescence is a naturally occurring form of chemiluminescence where energy is released by a chemical reaction in...
organs called photophore
Photophore
A photophore is a light-emitting organ which appears as luminous spots on various marine animals, including fish and cephalopods. The organ can be simple, or as complex as the human eye; equipped with lenses, shutters, color filters and reflectors...
s on their ventral surface. The shark is believed to use these photophores to match ambient light conditions and thus disguise its silhouette from predators.
Usually inhabiting nutrient-rich waters over upper continental and insular slopes, the spined pygmy shark feeds on small bony fishes and squid
Squid
Squid are cephalopods of the order Teuthida, which comprises around 300 species. Like all other cephalopods, squid have a distinct head, bilateral symmetry, a mantle, and arms. Squid, like cuttlefish, have eight arms arranged in pairs and two, usually longer, tentacles...
. Like its prey it is a diel vertical migrator
Diel vertical migration
Diel vertical migration, also known as diurnal vertical migration, is a pattern of movement that some organisms living in the ocean and in lakes undertake each day. Usually organisms move up to the epipelagic zone at night and return to the mesopelagic zone of the oceans or to the hypolimnion zone...
, spending the day at close to 500 m (1,640.4 ft) deep and moving towards a depth of 200 m (656.2 ft) at night. Reproduction is presumably aplacental viviparous, with female giving birth to litters of 4 pups. This diminutive shark has no economic value. The International Union for Conservation of Nature (IUCN) has assessed this species as of Least Concern
Least Concern
Least Concern is an IUCN category assigned to extant taxon or lower taxa which have been evaluated but do not qualify for any other category. As such they do not qualify as threatened, Near Threatened, or Conservation Dependent...
, as it faces little threat from commercial fisheries and has a wide distribution.
Taxonomy and phylogeny
The spined pygmy shark was one of many new species discovered during the course of the 1907–1910 Philippine Expedition of the U.S. Fish CommissionUnited States Fish Commission
The United States Commission of Fish and Fisheries was established on February 9, 1871 , as an independent commission with a mandate to investigate the causes for the decrease of commercial fish and aquatic animals in U.S...
Steamer
Steamboat
A steamboat or steamship, sometimes called a steamer, is a ship in which the primary method of propulsion is steam power, typically driving propellers or paddlewheels...
Albatross
USS Albatross (1882)
The second USS Albatross, often seen as USFC Albatross in scientific literature citations, was an iron-hulled, twin-screw steamer in the United States Navy and reputedly the first vessel ever built especially for marine research....
. It was described by American ichthyologists Hugh McCormick Smith
Hugh McCormick Smith
Hugh McCormick Smith was an American ichthyologist and administrator in the Bureau of Fisheries.-Biography:...
and Lewis Radcliffe
Lewis Radcliffe
Lewis Radcliffe was a naturalist, malacologist, and ichthyologist. He was Deputy Commissioner of the United States Bureau of Fisheries until 1932 and was the assistant naturalist under Hugh McCormick Smith for the 1907-1910 Philippines Expedition. During his life, he described numerous new species...
in a 1912 paper for the scientific journal
Scientific journal
In academic publishing, a scientific journal is a periodical publication intended to further the progress of science, usually by reporting new research. There are thousands of scientific journals in publication, and many more have been published at various points in the past...
Proceedings of the United States National Museum, based on two specimens collected in Batangas Bay
Batangas Bay
Batangas Bay is a semi-enclosed body of water located in the Philippines. The municipalities of Mabini, Bauan, San Pascual, Tingloy on Maribacan Island and Batangas City are located on the coast of the bay. The water surface area of the bay is about 220 km² , and the coastline has an...
, south of Luzon
Luzon
Luzon is the largest island in the Philippines. It is located in the northernmost region of the archipelago, and is also the name for one of the three primary island groups in the country centered on the Island of Luzon...
in the Philippines
Philippines
The Philippines , officially known as the Republic of the Philippines , is a country in Southeast Asia in the western Pacific Ocean. To its north across the Luzon Strait lies Taiwan. West across the South China Sea sits Vietnam...
. One of these, a 15 cm (5.9 in) long adult male, was designated the type specimen.
Smith and Radcliffe coined the new genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
Squaliolus for this shark, and gave it the specific epithet laticaudus, from the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
latus meaning "broad" or "wide", and cauda meaning "tail". The spined pygmy shark may also be referred to as the dwarf shark or the bigeye dwarf shark. Based on similarities in their claspers (male intromittent organ
Intromittent organ
An intromittent organ is a general term for an external organ of a male organism that is specialized to deliver sperm during copulation. Intromittent organs are found most often in terrestrial species, as most aquatic species fertilize their eggs externally, although there are...
s), the closest relative of the spined pygmy shark and the related S. aliae is thought to be the pygmy shark
Pygmy shark
The pygmy shark, Euprotomicrus bispinatus, the second-smallest of all the shark species after the dwarf lanternshark, is a sleeper shark of the Dalatiidae family, the only member of the genus Euprotomicrus...
(Euprotomicrus bispinatus).
Distribution and habitat
The spined pygmy shark has a wide distribution around the world. In the Atlantic OceanAtlantic Ocean
The Atlantic Ocean is the second-largest of the world's oceanic divisions. With a total area of about , it covers approximately 20% of the Earth's surface and about 26% of its water surface area...
, it occurs off Bermuda
Bermuda
Bermuda is a British overseas territory in the North Atlantic Ocean. Located off the east coast of the United States, its nearest landmass is Cape Hatteras, North Carolina, about to the west-northwest. It is about south of Halifax, Nova Scotia, Canada, and northeast of Miami, Florida...
, the United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...
, Suriname
Suriname
Suriname , officially the Republic of Suriname , is a country in northern South America. It borders French Guiana to the east, Guyana to the west, Brazil to the south, and on the north by the Atlantic Ocean. Suriname was a former colony of the British and of the Dutch, and was previously known as...
, southern Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...
, and northern Argentina
Argentina
Argentina , officially the Argentine Republic , is the second largest country in South America by land area, after Brazil. It is constituted as a federation of 23 provinces and an autonomous city, Buenos Aires...
in the west, and off northern France
France
The French Republic , The French Republic , The French Republic , (commonly known as France , is a unitary semi-presidential republic in Western Europe with several overseas territories and islands located on other continents and in the Indian, Pacific, and Atlantic oceans. Metropolitan France...
, Madeira
Madeira
Madeira is a Portuguese archipelago that lies between and , just under 400 km north of Tenerife, Canary Islands, in the north Atlantic Ocean and an outermost region of the European Union...
, Cape Verde
Cape Verde
The Republic of Cape Verde is an island country, spanning an archipelago of 10 islands located in the central Atlantic Ocean, 570 kilometres off the coast of Western Africa...
, and the Azores
Azores
The Archipelago of the Azores is composed of nine volcanic islands situated in the middle of the North Atlantic Ocean, and is located about west from Lisbon and about east from the east coast of North America. The islands, and their economic exclusion zone, form the Autonomous Region of the...
in the east. In the Indian Ocean
Indian Ocean
The Indian Ocean is the third largest of the world's oceanic divisions, covering approximately 20% of the water on the Earth's surface. It is bounded on the north by the Indian Subcontinent and Arabian Peninsula ; on the west by eastern Africa; on the east by Indochina, the Sunda Islands, and...
, this species has only been recorded off Somalia
Somalia
Somalia , officially the Somali Republic and formerly known as the Somali Democratic Republic under Socialist rule, is a country located in the Horn of Africa. Since the outbreak of the Somali Civil War in 1991 there has been no central government control over most of the country's territory...
. In the Pacific Ocean
Pacific Ocean
The Pacific Ocean is the largest of the Earth's oceanic divisions. It extends from the Arctic in the north to the Southern Ocean in the south, bounded by Asia and Australia in the west, and the Americas in the east.At 165.2 million square kilometres in area, this largest division of the World...
, it is found off southern Japan
Japan
Japan is an island nation in East Asia. Located in the Pacific Ocean, it lies to the east of the Sea of Japan, China, North Korea, South Korea and Russia, stretching from the Sea of Okhotsk in the north to the East China Sea and Taiwan in the south...
, Taiwan
Taiwan
Taiwan , also known, especially in the past, as Formosa , is the largest island of the same-named island group of East Asia in the western Pacific Ocean and located off the southeastern coast of mainland China. The island forms over 99% of the current territory of the Republic of China following...
, and the Philippines
Philippines
The Philippines , officially known as the Republic of the Philippines , is a country in Southeast Asia in the western Pacific Ocean. To its north across the Luzon Strait lies Taiwan. West across the South China Sea sits Vietnam...
. The spined pygmy shark is found at depths of 200–500 m (656.2–1,640.4 ft) and seldom approaches the surface, unlike the related pygmy shark and cookiecutter shark
Cookiecutter shark
The cookiecutter shark , also called the cigar shark, is a species of small dogfish shark in the family Dalatiidae. This shark occurs in warm, oceanic waters worldwide, particularly near islands, and has been recorded from as deep as . It migrates vertically up to every day, approaching the...
(Isistius brasiliensis). This shark prefers areas of high biological productivity over upper continental and insular slopes. It may also be found over outer shelves
Continental shelf
The continental shelf is the extended perimeter of each continent and associated coastal plain. Much of the shelf was exposed during glacial periods, but is now submerged under relatively shallow seas and gulfs, and was similarly submerged during other interglacial periods. The continental margin,...
, but avoids central ocean basins. The range of this species does not overlap that of the pygmy shark, which has a similar ecology, and is also largely separate from that of the cookiecutter shark.
Description
One of the world's smallest sharks, the spined pygmy shark attains a maximum recorded length of 22 cm (8.7 in) for males and 28 cm (11 in) for females. This species has an elongated, spindle-shaped body with a long, bulbous, moderately pointed snout. The eyes are large, with the upper rim of the orbit almost straight. Each nostril is preceded by a short flap of skin. The mouth has thin, smooth lips and contains 22–31 tooth rows in the upper jaw and 16–21 tooth rows in the lower jaw. The upper teeth are narrow and smooth-edged with single upright cusps. The bases of the lower teeth are broad and interlocked to form a continuous cutting surface, with each tooth bearing a single upright, smooth-edged, knife-like cusp. The openings of the five pairs of gill slitGill slit
Gill slits are individual openings to gills, i.e., multiple gill arches, which lack a single outer cover. Such gills are characteristic of Cartilaginous fish such as sharks, rays, sawfish, and guitarfish. Most of these have five pairs, but a few species have 6 or 7 pairs...
s are minute and uniform in size.
The two species of Squaliolus are the only sharks with a spine on the first dorsal fin
Dorsal fin
A dorsal fin is a fin located on the backs of various unrelated marine and freshwater vertebrates, including most fishes, marine mammals , and the ichthyosaurs...
but not the second. The spine is sexually dimorphic, being typically exposed in males and enclosed by skin in females. The first dorsal fin is tiny and originates over the trailing margin of the pectoral fins. The second dorsal fin is low, with a base twice as long as that of the first, and originates over the anterior half of the pelvic fin bases. The pectoral fins are short and triangular, with the rear margin slightly curved. The pelvic fins are long and low, and there is no anal fin. The caudal peduncle is slender and laterally expanded into weak keels. The caudal fin is broad and paddle-like, with the upper and lower lobes of similar size and shape, and a deep notch in the trailing margin of the upper lobe.
The dermal denticles are flat and blocky, not elevated on stalks or bearing marginal teeth. The coloration is dark brown to black, with light fin margins. The underside is densely carpeted by light-emitting photophore
Photophore
A photophore is a light-emitting organ which appears as luminous spots on various marine animals, including fish and cephalopods. The organ can be simple, or as complex as the human eye; equipped with lenses, shutters, color filters and reflectors...
s, which extend to the tip of the snout and around the eyes and nostrils, and thin to almost non-existent on the back. This species has on average only 60 vertebrae, the fewest of any shark.
Biology and ecology
The diet of the spined pygmy shark consists mainly of bony fishes (including the dragonfishStomiidae
Stomiidae is a family of deep-sea ray-finned fish, including the barbeled dragonfishes, stareaters and loosejaws.Stomiids are generally elongated fish with black or near-black bodies, but they are highly variable in form, and are sometimes grouped into multiple different families as a result. The...
Idiacanthus, the lanternfish
Lanternfish
Cooper Lanternfishes are small mesopelagic fish of the large family Myctophidae. One of two families in the order Myctophiformes, the Myctophidae are represented by 246 species in 33 genera, and are found in oceans worldwide. They are aptly named after their conspicuous use of bioluminescence...
Diaphus
Diaphus
Diaphus is a genus of lanternfish in the family Myctophidae.- Species :* Gilbert's large lantern fish, Diaphus adenomus Gilbert, 1905* Diaphus aliciae Fowler, 1934* Andersen's lantern fish, Diaphus anderseni Tåning, 1932...
, and the bristlemouth Gonostoma) and squid
Squid
Squid are cephalopods of the order Teuthida, which comprises around 300 species. Like all other cephalopods, squid have a distinct head, bilateral symmetry, a mantle, and arms. Squid, like cuttlefish, have eight arms arranged in pairs and two, usually longer, tentacles...
(including members of the genera Chiroteuthis
Chiroteuthis
Chiroteuthis is a genus of chiroteuthid squid, comprising two subgenera. The hectocotylus is absent from all members of the genus; instead, a penis extending from the mantle opening is utilised. The genus is characterised by enlarged, lidded photophores present at the end of the tentacular club...
and Histioteuthis
Histioteuthis
Histioteuthis is a genus of squid and the only member of the Histioteuthidae family. It goes by the English name cock-eyed squid, because in all species the right eye is normal-sized, round, blue and sunken; whereas the left eye is at least twice the diameter of the right eye, tubular,...
). Catch records suggest that the spined pygmy shark follows its prey on their diel vertical migration
Diel vertical migration
Diel vertical migration, also known as diurnal vertical migration, is a pattern of movement that some organisms living in the ocean and in lakes undertake each day. Usually organisms move up to the epipelagic zone at night and return to the mesopelagic zone of the oceans or to the hypolimnion zone...
s, spending the day close to a depth of 500 m (1,640.4 ft) and ascending towards a depth of 200 m (656.2 ft) at night. The ventral photophores of the spined pygmy shark have been theorized to function in counter-illumination
Countershading
Countershading, or Thayer's Law, is a form of camouflage. Countershading, in which an animal’s pigmentation is darker dorsally, is often thought to have an adaptive effect of reducing conspicuous shadows cast on the ventral region of an animal’s body...
, a form of camouflage in which the shark disguises its silhouette from would-be predators by matching the ambient light welling down from above. There is no evidence that this shark swallows its shed teeth like the pygmy and cookiecutter sharks.
The spined pygmy shark is aplacental viviparous like the rest of its family, with the developing embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
s being sustained by a yolk sac
Yolk sac
The yolk sac is a membranous sac attached to an embryo, providing early nourishment in the form of yolk in bony fishes, sharks, reptiles, birds, and primitive mammals...
until birth. Adult females have two functional ovaries that may each contain up to 12 mature eggs. However, the actual litter size is much smaller; a pregnant female caught off southern Brazil in 1999 contained four near-term pups. The young are born at 9–10 cm (3.5–3.9 in) long. Males mature sexually at a length of 15 cm (5.9 in), and females at a length of 17–20 m (669.3–787.4 in). The spined pygmy shark was widely considered to be the smallest living shark species until the discovery of the dwarf lanternshark
Dwarf lanternshark
The dwarf lanternshark is a little-known species of dogfish shark in the family Etmopteridae and possibly the smallest shark in the world, reaching a maximum known length of . It is known to be present only on the upper continental slopes off Colombia and Venezuela, at a depth of...
(Etmopterus perryi), though the pygmy ribbontail catshark
Pygmy ribbontail catshark
The pygmy ribbontail catshark is a species of finback catshark, family Proscylliidae, distributed patchily in the western Indo-Pacific from Tanzania to the Philippines. It occurs around the edges of continental and insular shelves at a depth of , typically on or near mud bottoms...
(Eridacnis radcliffei) is also known to mature at a size comparable to these two species. Whether one of these sharks is definitively smaller than the others cannot yet be stated with certainty, because of the difficulties involved in assessing reproductive maturity in sharks.
Human interactions
Spined pygmy sharks have no commercial value; they sometimes appear in the bycatchBycatch
The term “bycatch” is usually used for fish caught unintentionally in a fishery while intending to catch other fish. It may however also indicate untargeted catch in other forms of animal harvesting or collecting...
of trawl fisheries, but are generally too small to be captured. In light of its wide distribution and the absence of substantial threats from human activity, the International Union for Conservation of Nature (IUCN) has assessed this species as of Least Concern
Least Concern
Least Concern is an IUCN category assigned to extant taxon or lower taxa which have been evaluated but do not qualify for any other category. As such they do not qualify as threatened, Near Threatened, or Conservation Dependent...
.