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The rhinarium (
New Latin New Latin (also called Neo-Latin or Modern Latin) is the revival of Literary Latin used in original, scholarly, and scientific works since about 1500. Modern scholarly and technical nomenclature, such as in zoological and botanical taxonomy ...
, "belonging to the nose"; plural: rhinaria) is the furless skin surface surrounding the external openings of the nostrils in many
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s. Commonly it is referred to as the tip of the ''
snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, rostrum, or proboscis. The wet furless surface around the nostrils of the nose of many mammals is ...
'', and breeders of cats and dogs sometimes use the term ''nose leather''. Informally, it may be called a "truffle", "wet snout," or "wet nose” because its surface is moist in some
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
: for example, healthy dogs and
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s. In many species, the rhinarium has a mid-line groove (cleft)the
philtrum The philtrum ( la, philtrum from Ancient Greek ''phíltron,'' lit. "love charm"), or medial cleft, is a vertical indentation in the middle area of the upper lip, common to therian mammals, extending in humans from the nasal septum to the tuberc ...
and a wrinkled (
crenellate A battlement in defensive architecture, such as that of city walls or castles, comprises a parapet (i.e., a defensive low wall between chest-height and head-height), in which gaps or indentations, which are often rectangular, occur at interva ...
d) surface. The rhinarium is a separate sense organ: it is a touch-based chemosensory organ that connects with a well-developed
vomeronasal organ The vomeronasal organ (VNO), or Jacobson's organ, is the paired auxiliary olfactory (smell) sense organ located in the soft tissue of the nasal septum, in the nasal cavity just above the roof of the mouth (the hard palate) in various tetrapo ...
(VNO). The rhinarium is used to touch a scent-marked object containing pheromones (usually large, non-volatile molecules), and transfer these pheromone molecules down the philtrum to the VNO via the nasopalatine ducts that travel through the incisive foramen of the
hard palate The hard palate is a thin horizontal bony plate made up of two bones of the facial skeleton, located in the roof of the mouth. The bones are the palatine process of the maxilla and the horizontal plate of palatine bone. The hard palate spans t ...
. It also acts as a wind-direction detector: cold receptors in the skin of the rhinarium detect the orientation where evaporative cooling is highest, as determined by the wind direction. The study of the rhinarium's structure and associated functions has proven of considerable importance in the fields of mammalian evolution and taxonomy.Basbaum, Allan I. Kaneko, Akimichi, Shepherd, Gordon G. Westheimer, Gerald (editors). The Senses: A Comprehensive Reference. Academic press 2007. For example,
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians ( monkeys and apes, the latter includin ...
s are
phylogenetically In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
divided into those, such as lemurs, with the primitive rhinarium (
Strepsirrhini Strepsirrhini or Strepsirhini (; ) is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Co ...
) and the dry-nosed primates (
Haplorhini Haplorhini (), the haplorhines ( Greek for "simple-nosed") or the "dry-nosed" primates, is a suborder of primates containing the tarsiers and the simians (Simiiformes or anthropoids), as sister of the Strepsirrhini ("moist-nosed"). The name is so ...
, including apes and humans). In an analogous wayentirely unrelated to vertebrate morphologythe term rhinarium is sometimes applied to chemosensory structures in invertebrates. For example, microscopic sensilla in the form of flattened sense organs on the antennae of
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
s are referred to as rhinaria.


Morphology


Derivation

Morphologically, the rhinarium belongs to the
olfactory system The olfactory system, or sense of smell, is the sensory system used for smelling ( olfaction). Olfaction is one of the special senses, that have directly associated specific organs. Most mammals and reptiles have a main olfactory system and an ...
, but which part of the system it derives from is open to debate. One possibility is the main olfactory system, which captures media-borne odors; another is the "second nose," the
accessory olfactory system The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, i ...
, which samples chemicals dissolved in fluids. Arguments supporting the former position consider the rhinarium "an outward extension of the olfactory ... skin that covers the nasal passages, hichcontains nerve receptors for smell and touch." If that interpretation is correct, and the rhinarium is an extension of the olfactory epithelium lining the nasal passages, then it derives from the main system. But one opposing view traces a path from the philtrum over a notch in the upper lip, through a gap between the first
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
s and
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
e, along a "midline palatal groove" to "a canal that connects with the duct of the
vomeronasal organ The vomeronasal organ (VNO), or Jacobson's organ, is the paired auxiliary olfactory (smell) sense organ located in the soft tissue of the nasal septum, in the nasal cavity just above the roof of the mouth (the hard palate) in various tetrapo ...
," suggesting that the rhinarium belongs to the accessory system. It is unclear if moisture (
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
) there functions to trap odiferous molecules — or is the remnant of a fluid transmission system for pheromonal molecules.


Crenellations

Typically, the rhinarium is crenellated (wrinkled, crackled, or embossed), which may, in theory, increase its sensory area, but there are many exceptions and variations among different mammalian
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
, and also variations in the innervation and
sensilla A sensillum (plural ''sensilla'') is an arthropod sensory organ protruding from the cuticle of exoskeleton, or sometimes lying within or beneath it. Sensilla appear as small hairs or pegs over an individual's body. Inside each sensillum there are ...
of the rhinarium, so such generalized speculation should be treated with caution regarding this matter.


Function

Mammals with rhinaria tend to have a more acute
sense of smell The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, ...
, and the loss of the rhinarium in the haplorrhine primates is related to their decreased reliance on olfaction, being associated with other derived characteristics such as a reduced number of
turbinate In anatomy, a nasal concha (), plural conchae (), also called a nasal turbinate or turbinal, is a long, narrow, curled shelf of bone that protrudes into the breathing passage of the nose in humans and various animals. The conchae are shaped lik ...
s. The rhinarium is also very useful to animals with good sense of smell because of its role as a wind-direction detector. The cold receptors in the skin respond to the place where evaporation is the highest. Thus the detection of a particular smell is associated with the direction it comes from. The rhinarium is adapted for different purposes in different mammals, according to
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
. In aquatic mammals, the development of lobes beside the nostrils allows them to close for diving. In mammals that dig or root with their noses, the rhinarium often develops into a resilient pad, with the nostrils off to the side (or below) and capable of closing to keep out dust. Examples include the
common wombat The common wombat (''Vombatus ursinus''), also known as the coarse-haired wombat or bare-nosed wombat, is a marsupial, one of three extant species of wombats and the only one in the genus ''Vombatus''. The common wombat grows to an average of lo ...
, marsupial mole, and members of the
Chrysochloridae Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family Chrysochloridae and as such they are taxonomically distinct from the true moles, family Talpidae, and other mole-like families, all ...
. In elephants, the rhinarium has become a tactile organ. And in the walrus, a covering of stiff bristles protects it while the animal forages for shellfish. In many animals, the form and purpose of the rhinarium remains unclear. Evolutionary pressures also are not always unequivocally distinguishable, and there have been upheavals in late 20th and early 21st century taxonomy. For example, the lack of an obvious rhinarium in ''
Tarsiiformes Tarsiiformes are a group of primates that once ranged across Europe, northern Africa, Asia, and North America, but whose extant species are all found in the islands of Southeast Asia. Tarsiers (family Tarsiidae) are the only living members ...
'' has been interpreted by some scholars as the consequence of the enormous development of the eyeballs, rather than a loss of relevance of
olfaction The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, ...
,Smith, T. D., & Bhatnagar, K. P. (2004). Microsmatic primates: reconsidering how and when size matters. The Anatomical Record Part B: The New Anatomist, 279(1), 24-31. but the significance is currently debatable, because there currently is an influential body of opinion favouring inclusion of the tarsiers in the
Haplorhini Haplorhini (), the haplorhines ( Greek for "simple-nosed") or the "dry-nosed" primates, is a suborder of primates containing the tarsiers and the simians (Simiiformes or anthropoids), as sister of the Strepsirrhini ("moist-nosed"). The name is so ...
rather than in the
Strepsirrhini Strepsirrhini or Strepsirhini (; ) is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Co ...
as had been traditional.


Phylogeny


Mammalian

The rhinarium is a general mammalian feature and therefore likely to have been present in the stem mammals.


Primate

Primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians ( monkeys and apes, the latter includin ...
s are
phylogenetically In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
divided into those with a rhinarium, the
Strepsirrhini Strepsirrhini or Strepsirhini (; ) is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Co ...
(the prosimians: the lorises, and the lemurs); and those without a rhinarium, the
Haplorhini Haplorhini (), the haplorhines ( Greek for "simple-nosed") or the "dry-nosed" primates, is a suborder of primates containing the tarsiers and the simians (Simiiformes or anthropoids), as sister of the Strepsirrhini ("moist-nosed"). The name is so ...
, (the Simians:
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incomple ...
s, apes, and
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s). In place of the rhinarium, Haplorhini have a more mobile, continuous, dry upper lip.


Use of term in invertebrates

In an analogous usage unrelated to vertebrate morphology, the term rhinarium is sometimes applied to chemosensory structures in invertebrates. For example, microscopic sensilla in the flattened sense organs on the antennae of
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
s are referred to as rhinaria.Du Yongjun Yan Fushun Tang Jue. Structure and Function of Olfactory Sensilla on the Antennae of Soybean Aphids, Aphis glycines. ACTA ENTOMOLOGICA SINICA 1995, Vol. 38 Issue (1): 1-

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See also

*
Nose A nose is a protuberance in vertebrates that houses the nostrils, or nares, which receive and expel air for respiration alongside the mouth. Behind the nose are the olfactory mucosa and the sinuses. Behind the nasal cavity, air next passe ...
*
Snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, rostrum, or proboscis. The wet furless surface around the nostrils of the nose of many mammals is ...


References


Bibliography

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External links

{{wiktionary, rhinarium
Mammalian Rhinarium Group
- Lund University Mammal anatomy Dog anatomy Felidae anatomy