Faunivorous
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A carnivore , or meat-eater (
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
, ''caro'', genitive ''carnis'', meaning
meat Meat is animal Tissue (biology), tissue, often muscle, that is eaten as food. Humans have hunted and farmed other animals for meat since prehistory. The Neolithic Revolution allowed the domestication of vertebrates, including chickens, sheep, ...
or "flesh" and ''vorare'' meaning "to devour"), is an
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
or
plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
whose
nutrition Nutrition is the biochemistry, biochemical and physiology, physiological process by which an organism uses food and water to support its life. The intake of these substances provides organisms with nutrients (divided into Macronutrient, macro- ...
and
energy Energy () is the physical quantity, quantitative physical property, property that is transferred to a physical body, body or to a physical system, recognizable in the performance of Work (thermodynamics), work and in the form of heat and l ...
requirements are met by
consumption Consumption may refer to: * Eating *Resource consumption *Tuberculosis, an infectious disease, historically known as consumption * Consumer (food chain), receipt of energy by consuming other organisms * Consumption (economics), the purchasing of n ...
of animal tissues (mainly
muscle Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...
,
fat In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food. The term often refers specif ...
and other
soft tissue Soft tissue connective tissue, connects and surrounds or supports internal organs and bones, and includes muscle, tendons, ligaments, Adipose tissue, fat, fibrous tissue, Lymphatic vessel, lymph and blood vessels, fasciae, and synovial membranes. ...
s) as
food Food is any substance consumed by an organism for Nutrient, nutritional support. Food is usually of plant, animal, or Fungus, fungal origin and contains essential nutrients such as carbohydrates, fats, protein (nutrient), proteins, vitamins, ...
, whether through
predation Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
or
scavenging Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...
.


Nomenclature


Mammal order

The technical term for mammals in the
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
Carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...
is ''carnivoran'', and they are so-named because most member species in the group have a carnivorous diet, but the similarity of the name of the order and the name of the diet causes confusion. Many but not all carnivorans are meat eaters; a few, such as the large and small cats (
Felidae Felidae ( ) is the Family (biology), family of mammals in the Order (biology), order Carnivora colloquially referred to as cats. A member of this family is also called a felid ( ). The 41 extant taxon, extant Felidae species exhibit the gre ...
) are ''obligate'' carnivores (see below). Other classes of carnivore are highly variable. The ursids (bears), for example: while the Arctic
polar bear The polar bear (''Ursus maritimus'') is a large bear native to the Arctic and nearby areas. It is closely related to the brown bear, and the two species can Hybrid (biology), interbreed. The polar bear is the largest extant species of bear ...
eats meat almost exclusively (more than 90% of its diet is meat), almost all other bear species are
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
, and one species, the
giant panda The giant panda (''Ailuropoda melanoleuca''), also known as the panda bear or simply panda, is a bear species endemic to China. It is characterised by its white animal coat, coat with black patches around the eyes, ears, legs and shoulders. ...
, is nearly exclusively
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
. Dietary carnivory is not a distinguishing trait of the
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
. Many mammals with highly carnivorous diets are ''not'' members of the order Carnivora.
Cetacean Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
s, for example, all eat other animals, but are paradoxically members of the almost exclusively plant-eating hooved mammals.


Carnivorous diet

Animals that depend solely on animal flesh for their nutrient requirements in nature are called ''
hypercarnivore A hypercarnivore is an animal that has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant exampl ...
s'' or ''obligate carnivores'', whilst those that also consume non-animal food are called ''
mesocarnivore A mesocarnivore is an animal whose diet consists of 30–70% meat with the balance consisting of non-vertebrate foods which may include insects, fungi, fruits, other plant material and any food that is available to them. Mesocarnivores are from ...
s'', or ''facultative carnivores'', or ''
omnivores An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...
'' (there are no clear distinctions). A carnivore at the top of the
food chain A food chain is a linear network of links in a food web, often starting with an autotroph (such as grass or algae), also called a producer, and typically ending at an apex predator (such as grizzly bears or killer whales), detritivore (such as ...
(adults not preyed upon by other animals) is termed an
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
, regardless of whether it is an ''obligate'' or ''facultative'' carnivore. In captivity or domestic settings, obligate carnivores like cats and crocodiles can, in principle, get all their required nutrients from processed food made from plant and synthetic sources. Outside the animal kingdom, there are several genera containing
carnivorous plant Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods, and occasionally small mammals and birds. They have adapted to grow in waterlo ...
s (predominantly insectivores), several phyla containing
carnivorous fungi A carnivorous fungus or predaceous fungus is a fungus that derives some or most of its nutrients from trapping and eating microscopic or other minute animals. More than 200 species have been described, belonging to the phyla Ascomycota, Mucoromyc ...
(preying mostly on microscopic
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s, such as
nematode The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
s,
amoebae An amoeba (; less commonly spelled ameba or amœba; : amoebas (less commonly, amebas) or amoebae (amebae) ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and r ...
, and
springtail Springtails (class Collembola) form the largest of the three lineages of modern Hexapoda, hexapods that are no longer considered insects. Although the three lineages are sometimes grouped together in a class called Entognatha because they have in ...
s) and carnivorous protists.


Subcategories of carnivory

Carnivores are sometimes characterized by their type of prey. For example, animals that eat mainly
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s and similar
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s are called ''
insectivore file:Common brown robberfly with prey.jpg, A Asilidae, robber fly eating a hoverfly An insectivore is a carnivore, carnivorous animal or plant which eats insects. An alternative term is entomophage, which can also refer to the Entomophagy ...
s'', while those that eat mainly soft-bodied
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s are called ''
vermivore Vermivore (from Latin vermi, meaning "worm" and vorare, "to devour") is a Zoology, zoological term for animals that primarily eat worms (including annelids, nematodes, and other worm-like animals). Animals with such a diet are known to be vermivor ...
s''. Those that eat mainly
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
are called ''
piscivore A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...
s''. Carnivores may alternatively be classified according to the percentage of meat in their diet. The diet of a
hypercarnivore A hypercarnivore is an animal that has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant exampl ...
consists of more than 70% meat, that of a
mesocarnivore A mesocarnivore is an animal whose diet consists of 30–70% meat with the balance consisting of non-vertebrate foods which may include insects, fungi, fruits, other plant material and any food that is available to them. Mesocarnivores are from ...
30–70%, and that of a
hypocarnivore A hypocarnivore is an animal that consumes less than 30% meat for its diet, the majority of which consists of fungi, fruits, and other plant material. Examples of living hypocarnivores are the grizzly bear (''Ursus horribilis''), black bear (''U ...
less than 30%, with the balance consisting of non-animal foods, such as
fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
, other plant material, or
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
.
Omnivore An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...
s also consume both animal and non-animal food, and apart from their more general definition, there is no clearly defined ratio of plant vs. animal material that distinguishes a ''facultative carnivore'' from an ''omnivore''.


Obligate carnivores

Obligate or "true" carnivores are those whose diet requires nutrients found only in animal flesh in the wild. While obligate carnivores might be able to
ingest Ingestion is the consumption of a substance by an organism. In animals, it normally is accomplished by taking in a substance through the mouth into the gastrointestinal tract, such as through eating or drinking. In single-celled organisms, inges ...
small amounts of plant matter, they lack the necessary
physiology Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
required to fully digest it. Some obligate carnivorous mammals will ingest vegetation as an
emetic Vomiting (also known as emesis, puking and throwing up) is the forceful expulsion of the contents of one's stomach through the mouth and sometimes the nose. Vomiting can be the result of ailments like food poisoning, gastroenteritis, preg ...
, a food that upsets their stomachs, to self-induce vomiting. Obligate carnivores are diverse. The amphibian
axolotl The axolotl (; from ) (''Ambystoma mexicanum'') is a neoteny, paedomorphic salamander, one that Sexual maturity, matures without undergoing metamorphosis into the terrestrial adult form; adults remain Aquatic animal, fully aquatic with obvio ...
consumes mainly worms and larvae in its environment, but if necessary will consume algae. All wild
felids Felidae ( ) is the family of mammals in the order Carnivora colloquially referred to as cats. A member of this family is also called a felid ( ). The 41 extant Felidae species exhibit the greatest diversity in fur patterns of all terrestria ...
, including feral
domestic cats The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the d ...
, require a diet of primarily animal flesh and organs. Specifically, cats have high protein requirements and their metabolisms appear unable to synthesize essential nutrients such as
retinol Retinol, also called vitamin A1, is a fat-soluble vitamin in the vitamin A family that is found in food and used as a dietary supplement. Retinol or other forms of vitamin A are needed for vision, cellular development, maintenance of skin and ...
,
arginine Arginine is the amino acid with the formula (H2N)(HN)CN(H)(CH2)3CH(NH2)CO2H. The molecule features a guanidinium, guanidino group appended to a standard amino acid framework. At physiological pH, the carboxylic acid is deprotonated (−CO2−) a ...
,
taurine Taurine (), or 2-aminoethanesulfonic acid, is a naturally occurring amino sulfonic acid that is widely distributed in animal tissues. It is a major constituent of bile and can be found in the large intestine. It is named after Latin (cogna ...
, and
arachidonic acid Arachidonic acid (AA, sometimes ARA) is a polyunsaturated omega−6 fatty acid 20:4(ω−6), or 20:4(5,8,11,14). It is a precursor in the formation of leukotrienes, prostaglandins, and thromboxanes. Together with omega−3 fatty acids an ...
; thus, in nature, they must consume flesh to supply these nutrients.


Characteristics of carnivores

Characteristics commonly associated with carnivores include strength, speed, and keen senses for hunting, as well as teeth and claws for capturing and tearing prey. However, some carnivores do not hunt and are
scavenger Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
s, lacking the physical characteristics to bring down prey; in addition, most hunting carnivores will scavenge when the opportunity arises. Carnivores have comparatively short digestive systems, as they are not required to break down the tough cellulose found in plants. Many hunting animals have evolved eyes facing forward, enabling depth perception. This is almost universal among mammalian predators, while most reptile and amphibian predators have eyes facing sideways.


Prehistory of carnivory

''
Predation Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
'' (the eating of one living
organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...
by another for
nutrition Nutrition is the biochemistry, biochemical and physiology, physiological process by which an organism uses food and water to support its life. The intake of these substances provides organisms with nutrients (divided into Macronutrient, macro- ...
) predates the rise of commonly recognized carnivores by hundreds of millions (perhaps billions) of years. It began with
single-celled organism A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and e ...
s that phagocytozed and digested other cells, and later evolved into
multicellular organism A multicellular organism is an organism that consists of more than one cell (biology), cell, unlike unicellular organisms. All species of animals, Embryophyte, land plants and most fungi are multicellular, as are many algae, whereas a few organism ...
s with specialized cells that were dedicated to breaking down other organisms. Incomplete digestion of the prey organisms, some of which survived inside the predators in a form of
endosymbiosis An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
, might have led to
symbiogenesis Symbiogenesis (endosymbiotic theory, or serial endosymbiotic theory) is the leading evolutionary theory of the origin of eukaryotic cells from prokaryotic organisms. The theory holds that mitochondria, plastids such as chloroplasts, and possibl ...
that gave rise to
eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
s and eukaryotic
autotroph An autotroph is an organism that can convert Abiotic component, abiotic sources of energy into energy stored in organic compounds, which can be used by Heterotroph, other organisms. Autotrophs produce complex organic compounds (such as carbohy ...
s such as
green Green is the color between cyan and yellow on the visible spectrum. It is evoked by light which has a dominant wavelength of roughly 495570 nm. In subtractive color systems, used in painting and color printing, it is created by a com ...
and
red algae Red algae, or Rhodophyta (, ; ), make up one of the oldest groups of eukaryotic algae. The Rhodophyta comprises one of the largest Phylum, phyla of algae, containing over 7,000 recognized species within over 900 Genus, genera amidst ongoing taxon ...
.


Proterozoic origin

The earliest predators were
microorganism A microorganism, or microbe, is an organism of microscopic scale, microscopic size, which may exist in its unicellular organism, single-celled form or as a Colony (biology)#Microbial colonies, colony of cells. The possible existence of unseen ...
s, which engulfed and "swallowed" other smaller
cell Cell most often refers to: * Cell (biology), the functional basic unit of life * Cellphone, a phone connected to a cellular network * Clandestine cell, a penetration-resistant form of a secret or outlawed organization * Electrochemical cell, a de ...
s (i.e.
phagocytosis Phagocytosis () is the process by which a cell (biology), cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs ph ...
) and digested them internally. Because the earliest fossil record is poor, these first predators could date back anywhere between 1 and over 2.7  bya (billion years ago). The rise of
eukaryotic The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
cells at around 2.7 bya, the rise of multicellular organisms at about 2 bya, and the rise of
motile Motility is the ability of an organism to move independently using metabolic energy. This biological concept encompasses movement at various levels, from whole organisms to cells and subcellular components. Motility is observed in animals, mi ...
predators (around 600 Mya – 2 bya, probably around 1 bya) have all been attributed to early predatory behavior, and many very early remains show evidence of boreholes or other markings attributed to small predator species. The sudden disappearance of the
precambrian The Precambrian ( ; or pre-Cambrian, sometimes abbreviated pC, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of t ...
Ediacaran biota The Ediacaran (; formerly Vendian) biota is a taxonomic period classification that consists of all life forms that were present on Earth during the Ediacaran Period (). These were enigmatic tubular and frond-shaped, mostly sessile, organis ...
at the
end-Ediacaran extinction The end-Ediacaran extinction is a mass extinction believed to have occurred near the end of the Ediacaran period, the final period of the Proterozoic eon. Evidence suggesting that such a mass extinction occurred includes a massive reduction in dive ...
, who were mostly bottom-dwelling
filter feeder Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a s ...
s and grazers, has been hypothetized to be partly caused by increased predation by newer animals with hardened skeleton and mouthparts.


Paleozoic

The degradation of
seafloor The seabed (also known as the seafloor, sea floor, ocean floor, and ocean bottom) is the bottom of the ocean. All floors of the ocean are known as seabeds. The structure of the seabed of the global ocean is governed by plate tectonics. Most of ...
microbial mat A microbial mat is a multi-layered sheet or biofilm of microbial colonies, composed of mainly bacteria and/or archaea. Microbial mats grow at interfaces between different types of material, mostly on submerged or moist surfaces, but a few surviv ...
s due to the
Cambrian substrate revolution The "Cambrian substrate revolution" or "Agronomic revolution", evidenced in trace fossils, is a sudden diversification of animal burrowing during the early Cambrian period. Before this "widening of the behavioural repertoire", bottom-dwelling ...
led to increased active predation among animals, likely triggering various
evolutionary arms race In evolutionary biology, an evolutionary arms race is an ongoing struggle between competing sets of co-evolving genes, phenotypic and behavioral traits that develop escalating adaptations and counter-adaptations against each other, resembling the ...
s that contributed to the rapid diversification during the Cambrian explosion.
Radiodont Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and were used for a variety ...
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s, which produced the first
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
s such as ''
Anomalocaris ''Anomalocaris'' (from Ancient Greek , meaning "unlike", and , meaning "shrimp", with the intended meaning "unlike other shrimp") is an extinct genus of radiodont, an order of early-diverging stem-group marine arthropods. It is best known fro ...
'', quickly became the dominant carnivores of the Cambrian sea. After their decline due to the Cambrian-Ordovician extinction event, the niches of large carnivores were taken over by
nautiloid Nautiloids are a group of cephalopods (Mollusca) which originated in the Late Cambrian and are represented today by the living ''Nautilus'' and ''Allonautilus''. Fossil nautiloids are diverse and species rich, with over 2,500 recorded species. Th ...
cephalopod A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
s such as ''
Cameroceras ''Cameroceras'', from Ancient Greek καμάρα (''kamára''), meaning "chamber", and κέρας (''kéras''), meaning "horn", is an extinct genus of endocerid cephalopod which lived in equatorial oceans during the entire Ordovician period. Lik ...
'' and later
eurypterid Eurypterids, often informally called sea scorpions, are a group of extinct marine arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period, 467.3 Myr, mil ...
s such as ''
Jaekelopterus ''Jaekelopterus'' is a genus of predatory eurypterid Eurypterids, often informally called sea scorpions, are a group of extinct marine arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the ...
'' during the
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
and
Silurian The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...
periods. The first
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
carnivores appeared after the
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
of
jawed fish Gnathostomata (; from Ancient Greek: (') 'jaw' + (') 'mouth') are jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all extant taxon, extant vertebrates, including all living bony fishes (bot ...
, especially
armor Armour (Commonwealth English) or armor (American English; see American and British English spelling differences#-our, -or, spelling differences) is a covering used to protect an object, individual, or vehicle from physical injury or damage, e ...
ed
placoderm Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos'''Plate (animal anatomy), plate' and δέρμα 'derma'''skin') are vertebrate animals of the class (biology), class Placodermi, an extinct group of prehistoric fish known from Pal ...
s such as the massive ''
Dunkleosteus ''Dunkleosteus'' is an extinct genus of large arthrodira, arthrodire ("jointed-neck") fish that existed during the Late Devonian period, about 382–358 million years ago. It was a pelagic fish inhabiting open waters, and one of the first verteb ...
''. The dominance of placoderms in the
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
ocean forced other fish to venture into other niches, and one clade of
bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
, the
lobe-finned fish Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
, became the dominant carnivores of
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include non-salty mi ...
wetland A wetland is a distinct semi-aquatic ecosystem whose groundcovers are flooded or saturated in water, either permanently, for years or decades, or only seasonally. Flooding results in oxygen-poor ( anoxic) processes taking place, especially ...
s formed by early
land plant The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophyt ...
s. Some of these fish became better adapted for breathing air and eventually giving rise to
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
s. These early tetrapods were large
semi-aquatic In biology, being semi-aquatic refers to various macroorganisms that live regularly in both aquatic and terrestrial environments. When referring to animals, the term describes those that actively spend part of their daily time in water (in ...
piscivores A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...
and
riparian A riparian zone or riparian area is the interface between land and a river or stream. In some regions, the terms riparian woodland, riparian forest, riparian buffer zone, riparian corridor, and riparian strip are used to characterize a ripar ...
ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture their prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey u ...
s that hunt
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...
arthropods (mainly
arachnid Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...
s and
myriopod Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. Although molecular evidence and similar fossils suggests a diversific ...
s), and one group in particular, the
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
s, became terrestrial apex predators that hunt other tetrapods. The dominance of temnospondyls around the wetland habitats throughout the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
forced other amphibians to evolve into
amniote Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
s that had adaptations that allowed them to live farther away from water bodies. These amniotes began to evolve both carnivory, which was a natural transition from insectivory requiring minimal adaptation; and
herbivory A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
, which took advantage of the abundance of
coal forest Coal forests were the vast swathes of freshwater swamp and riparian forests that covered much of the lands on Earth's tropical regions during the late Carboniferous ( Pennsylvanian) and Permian periods.Cleal, C. J. & Thomas, B. A. (2005). "Pala ...
foliage A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, f ...
but in contrast required a complex set of adaptations that was necessary for digesting on the
cellulose Cellulose is an organic compound with the chemical formula, formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of glycosidic bond, β(1→4) linked glucose, D-glucose units. Cellulose is an important s ...
- and
lignin Lignin is a class of complex organic polymers that form key structural materials in the support tissues of most plants. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidit ...
-rich plant materials. After the
Carboniferous rainforest collapse The Carboniferous rainforest collapse (CRC) was a minor extinction event that occurred around 305 million years ago in the Carboniferous period. The event occurred at the end of the Moscovian and continued into the early Kasimovian stages of th ...
, both
synapsid Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
and
sauropsid Sauropsida ( Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern reptiles and birds (which, as theropod dino ...
amniotes quickly gained dominance as the top terrestrial animals during the subsequent
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
period. Some scientists assert that sphenacodontoid synapsids such as ''
Dimetrodon ''Dimetrodon'' ( or ; ) is an extinct genus of sphenacodontid synapsid that lived during the Cisuralian (Early Permian) Epoch (geology), epoch of the Permian period, around 295–272 million years ago. With most species measuring long and ...
'' "were the first terrestrial vertebrate to develop the curved, serrated teeth that enable a predator to eat prey much larger than itself".


Mesozoic

In the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
, some
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s such as ''
Tyrannosaurus rex ''Tyrannosaurus'' () is a genus of large theropoda, theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It live ...
'' are thought probably to have been obligate carnivores. Though the theropods were the larger carnivores, several carnivorous mammal groups were already present. Most notable are the gobiconodontids, the
triconodontid Triconodontidae is an extinct family of small, carnivorous mammals belonging to the order Eutriconodonta, endemic to what would become Asia, Europe, North America and probably also Africa and South America during the Jurassic through Cretaceous p ...
''
Jugulator ''Jugulator'' is the thirteenth studio album by English heavy metal band Judas Priest. It was released in Japan on 16 October 1997 and the rest of the world on 28 October 1997. It was their first studio album since ''Painkiller'' in 1990 an ...
'', the deltatheroidans and ''
Cimolestes ''Cimolestes'' (from Ancient Greek , 'chalk robber') is a genus of early eutherians with a full complement of teeth adapted for eating insects and other small animals. Paleontologists have disagreed on its relationship to other mammals, in part b ...
''. Many of these, such as ''
Repenomamus ''Repenomamus'' (Latin: "reptile" (reptilis), "mammal" (mammalis)) is a genus of opossum- to badger-sized Gobiconodontidae, gobiconodontid mammal containing two species, ''Repenomamus robustus'' and ''Repenomamus giganticus''. Both species are kn ...
'', ''Jugulator'' and ''Cimolestes'', were among the largest mammals in their faunal assemblages, capable of attacking dinosaurs.


Cenozoic

In the early-to-mid-Cenozoic, the dominant predator forms were mammals:
hyaenodonts Hyaenodontidae ("hyena teeth") is a family of placental mammals in the extinct superfamily Hyaenodontoidea. Hyaenodontids arose during the early Eocene and persisted well into the early Miocene. Fossils of this group have been found in Asia, Nor ...
, oxyaenids,
entelodont Entelodontidae is an extinct family of pig-like artiodactyls (even-toed ungulates) which inhabited the Northern Hemisphere (Asia, Europe, and North America) from the late Eocene to the early Miocene epochs, about 38-19 million years ago. Their ...
s,
ptolemaiida Ptolemaiida is a taxon of wolf-sized Afrotheria, afrothere mammals that lived in northern and eastern Africa during the Paleogene. The oldest fossils are from the latest Eocene strata of the Jebel Qatrani Formation, near the Fayum oasis in Egypt. ...
ns, arctocyonids and
mesonychia Mesonychia ("middle claws") is an extinct taxon of small to large-sized carnivorous ungulates related to artiodactyls. Mesonychians first appeared in the early Paleocene, went into a sharp decline at the end of the Eocene, and died out entirely w ...
ns, representing a great diversity of
eutheria Eutheria (from Greek , 'good, right' and , 'beast'; ), also called Pan-Placentalia, is the clade consisting of Placentalia, placental mammals and all therian mammals that are more closely related to placentals than to marsupials. Eutherians ...
n carnivores in the northern continents and
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
. In
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
,
sparassodonts Sparassodonta (from Greek to tear, rend; and , gen. , ' tooth) is an extinct order of carnivorous metatherian mammals native to South America, related to modern marsupials. They were once considered to be true marsupials, but are now though ...
were dominant, while
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
saw the presence of several
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
predators, such as the
dasyuromorphs Dasyuromorphia (, meaning "hairy tail" in Greek) is an order comprising most of the Australian carnivorous marsupials, including quolls, dunnarts, the numbat, the Tasmanian devil, and the extinct thylacine. In Australia, the exceptions inc ...
and thylacoleonids. From the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
to the present, the dominant carnivorous mammals have been carnivoramorphs. Most carnivorous mammals, from
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the gray wolf. Also called the domestic dog, it was selectively bred from a population of wolves during the Late Pleistocene by hunter-gatherers. ...
s to ''
deltatheridium ''Deltatheridium'' (meaning ''triangle beast'' or ''delta beast'') is an extinct species of metatherian. It lived in what is now Mongolia during the Upper Cretaceous, ''circa'' 80 million years ago. A study in 2022 strongly suggested that ''De ...
s'', share several dental adaptations, such as
carnassial Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...
teeth, long
canine Canine may refer to: Zoology and anatomy * Animals of the family Canidae, more specifically the subfamily Caninae, which includes dogs, wolves, foxes, jackals and coyotes ** ''Canis'', a genus that includes dogs, wolves, coyotes, and jackals ** Do ...
s and even similar tooth replacement patterns. Most aberrant are thylacoleonids, with a diprodontan dentition completely unlike that of any other mammal; and
eutriconodonts Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia (including pre-contact India), Africa, Europe, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack ''et al.'' in 1973 ...
like gobiconodontids and ''Jugulator'', with a three-cusp anatomy which nevertheless functioned similarly to carnassials.


See also

*
Mesocarnivore A mesocarnivore is an animal whose diet consists of 30–70% meat with the balance consisting of non-vertebrate foods which may include insects, fungi, fruits, other plant material and any food that is available to them. Mesocarnivores are from ...


References


Further reading

* {{Authority control Biological interactions Animals by eating behaviors Ethology *