Venomous mammal

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The Cuban solenodon (Atopogale cubana) has a venomous bite. Dixi-Solenodon cubanus.png
The Cuban solenodon (Atopogale cubana) has a venomous bite.

Venomous mammals are animals of the class Mammalia that produce venom, which they use to kill or disable prey, to defend themselves from predators or conspecifics or in agonistic encounters. Mammalian venoms form a heterogeneous group with different compositions and modes of action, from four orders of mammals: Eulipotyphla, Monotremata, Primates, and Chiroptera. To explain the rarity of venom delivery in Mammalia, Mark Dufton of the University of Strathclyde has suggested that modern mammalian predators do not need venom because they are able to kill quickly with their teeth or claws, whereas venom, no matter how sophisticated, requires time to disable prey. [1]

Contents

In spite of the rarity of venom among extant mammals, venom may be an ancestral feature among mammals, as venomous spurs akin to those of the modern platypus are found in most non-therian Mammaliaformes groups. [2]

Venom is much more common among other vertebrates; there are many more species of venomous reptiles (e.g. venomous snakes) and fish (e.g. stonefish). Some birds are poisonous to eat or touch (e.g. hooded pitohui) though no bird species is known to be venomous. [3] There are only a few species of venomous amphibians; certain salamandrid salamanders can extrude sharp venom-tipped ribs. [4] [5]

Definitions

Several definitions of venomous animals have been proposed. [6]

Bücherl states that venomous animals must possess at least one venom gland, a mechanism for excretion or extrusion of the venom, and apparatus with which to inflict wounds.

Mebs writes that venomous animals produce venom in a group of cells or gland, and have a tool, the venom apparatus, which delivers the venom by injection during a bite or sting. The venom apparatus in this definition encompasses both the gland and the injection device, which must be directly connected.

Fry et al. found that a venom is a secretion produced in a specialized gland in one animal and delivered to a target animal through the infliction of a wound. This secretion must contain molecules that disrupt normal physiological processes so as to facilitate feeding or defense by the producing animal. Additionally, the feeding secretion of hematophagous specialists (e.g. vampire bats) may be regarded as a specialized subtype of venom.

Evolutionary history and paleontology

Venomous mammals may have been more common in the past. Extratarsal spurs observed in extinct Mesozoic mammals are homologous with those seen in extant monotremes, and this feature was widespread. Whether or not these extratarsal spurs served as a venom delivery system is up for debate. While species such as Gobiconodon and Zhangeotherium possessed an extratarsal spur, they are hypothesized to be nonvenomous due to the lack of the spur being hollow or channeled, as seen in modern venomous Ornithorhyncus. Whether or not the lack of venom is due to the loss of a basal trait is difficult to determine with current fossil records. [2]

In 2005, Fox and Scott presented evidence of a venom delivery system in late Paleocene fossils found in Alberta. Bisonalveus browni was a small eutherian in the order Cimolesta. Fox and Scott argue that the anterior grooves of the upper canines in B. browni serve as venom channels. They also present a group of isolated lower mammalian teeth of unknown species that also feature these grooves. [7] These specimens were reexamined and rejected on the basis that the evidence provided may not be enough to prove the presence of venom in these species. Fox and Scott found no fossa that would be indicative of the presence of a venom gland. Also, grooved teeth are a feature of many species of nonvenomous mammals, so the presence of grooved teeth alone is not indicative of venom. There is no phylogenetic support for the presence of venom in B. browni. [8] [9]

The genus Beremendia may have featured individuals with a venom delivery system. The first lower incisor of several species exhibited a groove that may have served as a canal for venom to travel from a venom gland to the tip of the tooth. Support for a venom gland is seen in the fossa in the mandibular symphysis. Also, Beremendia is phylogenetically close to Solenodon, which has extant members that are venomous. [9] [10] Euchambersia is another example of fossil mammal with grooved teeth, and maxillary fossa to match, indicating the presence of venom glands. Euchambersia differs in that the upper canines are the teeth that are grooved, and the grooves are almost enclosed, forming a tube. [9]

More recently, nonhomologous species in the genus Nesophontes were hypothesized to be venomous. Species within Nesophontes exhibit grooved upper canines, similar to other species suspected of venom delivery. Where Nesophontes differ, however, is in the presence of grooves on the first and second upper premolars. The grooves are similar in morphology to helodermatid lizards, which feature a deep anterior groove and shallow posterior groove, as well as grooves on other teeth, which suggests that Nesophontes grooved premolars also aided in delivery of venom in addition to the upper canines. Nesophontes mandibular morphology is similar to that of existing venomous Eulipotyphlans, which further supports the hypothesis that they possessed venom. [11]

Examples

Eulipotyphla (previously known as insectivores)

With the exception of vampire bats, the Eulipotyphla are the only mammals so far observed to produce toxic saliva. These species have significantly enlarged and granular submaxillary salivary glands from which the toxic saliva is produced. [12]

The Cuban solenodon (Atopogale cubana) and Hispaniolan solenodon (Solenodon paradoxus) look similar to large shrews. They both have venomous bites; the venom is delivered from modified salivary glands via grooves in their second lower incisors. Recent study has identified the gene regulatory network responsible for the development of venom delivery systems in these small mammals. [13] Due to the overexpression of kallikreins in their saliva, solenodon bites cause vasodilation and may result in circulatory shock. [13] It was reported that death was frequent among Hispaniolan solenodons kept together in the same enclosure, with bite marks on their feet being the only observable cause. Such use in competition may be a secondary aspect of the insectivore venom. [6]

The northern short-tailed shrew is one of several venomous shrews. Blarina brevicauda.jpg
The northern short-tailed shrew is one of several venomous shrews.

The northern short-tailed shrew (Blarina brevicauda), Mediterranean water shrew (Neomys anomalus), and Eurasian water shrew (Neomys fodiens) are capable of delivering a venomous bite. Other American short-tailed shrews—the southern short-tailed shrew (Blarina carolinensis), Elliot's short-tailed shrew (Blarina hylophaga), and Everglades short-tailed shrew (Blarina peninsulae) and the Transcaucasian water shrew (Neomys teres)—possibly also have a venomous bite. Shrews cache various prey in a comatose state, including earthworms, insects, snails, and to a lesser extent, small mammals such as voles and mice. This behaviour is an adaption to winter. In this context, the shrew venom acts as a tool to sustain a living hoard, thus ensuring food supply when capturing prey is difficult. This is especially important considering the high metabolic rate of shrews. Arguments against this suggest that the venom is used as a tool to hunt larger prey. Insectivores have an enhanced dependence on vertebrate food material, which is larger and more dangerous than their power to weight ratio would allow, thus requiring an extra asset to overcome these difficulties. [12] Extant shrews do not have specialized venom delivery apparatus. Their teeth do not have channels, but a concavity on the first incisors may collect and transmit saliva from the submaxillary ducts, which open near the base of these teeth. [14]

The European mole (Talpa europaea), and possibly other species of mole, [15] have toxins in their saliva that can paralyze earthworms, allowing the moles to store them alive for later consumption. [16]

Male platypus

The calcareous spur found on the male platypus's hind limb is used to deliver venom. Platypus spur.JPG
The calcareous spur found on the male platypus's hind limb is used to deliver venom.

Both male and female platypuses ( Ornithorhyncus anatinus ) hatch with keratinised spurs on the hind limbs, although the females lose these during development. The spurs are connected to the venom-producing crural glands, forming the crural system. During the mating season these glands become highly active, producing venom to be delivered by the channeled spur. Echidnas, the other monotremes, have spurs but no functional venom glands. Although not potent enough to be lethal to humans, platypus venom is nevertheless so excruciating that victims may sometimes be temporarily incapacitated. Platypus envenomation was fairly common when the animal was still hunted for its fur. Nowadays any close contact with the animal is rare and restricted to biologists, zookeepers and anglers (who occasionally catch them in fishing lines or nets). [6]

When platypuses attack, they drive their hind legs together with considerable force so that the spurs are embedded in the flesh caught between and if venom is being produced, a few milliliters are injected by repeated jabbing. [17] The spurs have enough strength to support the weight of the platypus, which often hangs from the victim, requiring assistance for removal.

Most of the evidence now supports the proposition that the venom system is used by males on one another as a weapon when competing for females, taking part in sexual selection. During this season, males become more aggressive and are found with punctures in their bodies, especially in the tail region. Adult male platypuses largely avoid each other, outside of this mating rivalry. [6]

Platypus venom is likely retained from its distant non-monotreme ancestors, being the last living example of what was once a common characteristic among mammals. [2] Fossil records show that venom delivery systems were not sexually dimorphic in ancestral monotremes. [18] It has been hypothesized that venom spurs were once used for defense against predators. [18] Proteins derived from platypus venom are being studied for potential analgesic properties. [19]

Vampire bats

The definition of venom by Fry et al. (see Definitions) regards the feeding secretions of hematophagous (blood eating) specialists as a particular subtype of venom. In this context, the subfamily Desmodontinae represents the venomous mammals from the order Chiroptera. This group comprises the most well known venomous bat, the common vampire bat (Desmodus rotundus) and two other rare species, the hairy-legged vampire bat (Diphylla ecaudata) and the white-winged vampire bat (Diaemus youngi). These bats produce toxic saliva with anticoagulant properties and have a series of anatomical and physiological adaptations to allow nourishment based solely on blood. The majority of their prey do not perish from the attack or contact with the venom. [20]

Primates

Slow lorises (of the genera Nycticebus and Xanthonycticebus [21] ) are accepted as the only known venomous primate. [20] Slow loris venom was known in folklore in their host countries throughout southeast Asia for centuries, but dismissed by Western science until the 1990s. [20] There are nine recognised species of this small-bodied nocturnal primate. [22] They possess a dual composite venom consisting of saliva and brachial gland exudate, a malodourous fluid forming from an apocrine sweat gland on the animal's forearm. [23] Both fluids have been demonstrated as being venomous individually and creating a more potent venom when mixed. [23] Slow loris brachial gland exudate (BGE) has been shown to possess up to 142 volatile components, and possesses a variant of the cat allergen protein Fel-D1. The BGE has several ecological functions including anti-parasitic defence [24] and communication. Slow loris saliva has been shown to be cytotoxic to human skin cells in laboratory experiments without the administration of BGE. [25]

The venom is administered through morphologically distinct dentition in the form of an adapted toothcomb. [23] In the wild envenomation occurs from intraspecific competition; whereby two slow lorises fight for mates, food or territory. Slow loris inflicted wounds are a major cause of premature death in zoo and wildlife slow loris populations, [26] often resulting in festering and necrotic wounds. Slow loris envenomation in humans is rare, but can result in near fatal anaphylactic shock. [27] A suite of additional effects of the venom have been documented including mild to permanent disfigurement and mobility loss. [28] The study of slow loris venom was brought to the public attention in 2012 by the research of the primatologist Prof. K.A.I Nekaris and in her BBC documentary The Jungle Gremlins of Java. [29]

Arguably venomous mammals

Eulipotyphla (previously known as insectivores)

Hedgehogs ( Erinaceinae ) anoint their spines with a range of toxic and irritating substances. They will sometimes kill toads ( Bufo sp.), bite into the toads' poison glands and smear the toxic mixture on their spines. [30] [31]

Tenrecs, which are similar in appearance to hedgehogs but from a different line of evolutionary descent, may also have separately evolved self-anointing behaviour.

Rodentia

The African crested rat smears toxins on its flank hairs. Lophiomys imhausi.jpg
The African crested rat smears toxins on its flank hairs.

The African crested rat (Lophiomys imhausi) has a mane of long, coarse black-and-white banded hairs which extends from the top of the animal's head to just beyond the base of the tail. This mane is bordered by a broad, white-bordered strip of hairs covering an area of glandular skin on the flank. When the animal is threatened or excited, the mane erects and this flank strip parts, exposing the glandular area. The hairs in this flank area are highly specialised; at the tips they are like ordinary hair, but are otherwise spongy, fibrous, and absorbent. The rat is known to deliberately chew the roots and bark of the poison-arrow tree ( Acokanthera schimperi ), so-called because human hunters extract a toxin, ouabain, to coat arrows that can kill an elephant. After the rat has chewed the tree, it deliberately slathers the resulting mixture onto its specialised flank hairs which are adapted to rapidly absorb the poisonous mixture, acting like a lamp wick. It thereby creates a defense mechanism that can sicken or even kill predators which attempt to bite it. [31] [32] [33] [34]

Chemical defence

Skunks (Mustelidae) can eject a noxious fluid from glands near their anus. It is not only foul smelling, but can cause skin irritation and, if it gets in the eyes, temporary blindness. Some members of the mustelid family, such as the striped polecat ( Ictonyx striatus ), also have this capacity to an extent. Pangolins can also emit a noxious smelling fluid from glands near the anus. The greater long-nosed armadillo can release a disagreeable musky odor when threatened.

See also

Related Research Articles

<span class="mw-page-title-main">Lorisidae</span> Family of primates

Lorisidae is a family of strepsirrhine primates. The lorisids are all slim arboreal animals and comprise the lorises, pottos and angwantibos. Lorisids live in tropical, central Africa as well as in south and southeast Asia.

<span class="mw-page-title-main">Venom</span> Toxin secreted by an animal

Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved venom apparatus, such as fangs or a stinger, in a process called envenomation. Venom is often distinguished from poison, which is a toxin that is passively delivered by being ingested, inhaled, or absorbed through the skin, and toxungen, which is actively transferred to the external surface of another animal via a physical delivery mechanism.

<span class="mw-page-title-main">Loris</span> Subfamily of primates

Loris is the common name for the strepsirrhine mammals of the subfamily Lorinae in the family Lorisidae. Loris is one genus in this subfamily and includes the slender lorises, Nycticebus is the genus containing the slow lorises, and Xanthonycticebus is the genus name of the pygmy slow loris.

<span class="mw-page-title-main">Solenodon</span> Family of mammals

Solenodons are venomous, nocturnal, burrowing, insectivorous mammals belonging to the family Solenodontidae. The two living solenodon species are the Cuban solenodon and the Hispaniolan solenodon. Threats to both species include habitat destruction and predation by non-native cats, dogs, and mongooses, introduced by humans to the solenodons' home islands to control snakes and rodents.

<span class="mw-page-title-main">Slow loris</span> Genus of primates from Southeast Asia

Slow lorises are a group of several species of nocturnal strepsirrhine primates that make up the genus Nycticebus. Found in Southeast Asia and bordering areas, they range from Bangladesh and Northeast India in the west to the Sulu Archipelago in the Philippines in the east, and from Yunnan province in China in the north to the island of Java in the south. Although many previous classifications recognized as few as a single all-inclusive species, there are now at least eight that are considered valid: the Sunda slow loris (N. coucang), Bengal slow loris (N. bengalensis), Javan slow loris (N. javanicus), Philippine slow loris (N. menagensis), Bangka slow loris (N. bancanus), Bornean slow loris (N. borneanus), Kayan River slow loris (N. kayan) and Sumatran slow loris. A ninth species, the pygmy slow loris (X. pygmaeus), was recently moved to the new genus Xanthonycticebus. After the pygmy slow loris, the group's closest relatives are the slender lorises of southern India and Sri Lanka. Their next closest relatives are the African lorisids, the pottos, false pottos, and angwantibos. They are less closely related to the remaining lorisoids, and more distantly to the lemurs of Madagascar. Their evolutionary history is uncertain since their fossil record is patchy and molecular clock studies have given inconsistent results.

<span class="mw-page-title-main">Northern short-tailed shrew</span> Species of mammal

The northern short-tailed shrew is the largest shrew in the genus Blarina, and occurs in the northeastern region of North America. It is a semifossorial, highly active, and voracious insectivore and is present in a variety of habitats like broadleaved and pine forests among shrubs and hedges as well as grassy river banks. It is notable in that it is one of the few venomous mammals. The specific epithet, brevicauda, is a combination of the Latin brevis and cauda, meaning "short tail".

<span class="mw-page-title-main">Envenomation</span> Process of venom injection

Envenomation is the process by which venom is injected by the bite or sting of a venomous animal.

<i>Bisonalveus</i> Extinct family of mammals

Bisonalveus is an extinct genus of shrew-like mammals that were presumably ground-dwelling and fed on plants and insects. Bisonalveus fossils have been discovered in the upper Great Plains region of North America, including sites in modern-day Wyoming, North Dakota, Montana, and Alberta. The fossils have been dated to 60 million years ago, during the Tiffanian North American Stage of the Palaeocene epoch. Bisonalveus is the last known genus of the Pentacodontinae sub-family to have arisen, replacing the genus Coriphagus in the early Tiffanian. Bisonalveus itself appears to have gone extinct by the middle Tiffanian.

<span class="mw-page-title-main">Sunda slow loris</span> Species of primate found in South East Asia

The Sunda slow loris, or greater slow loris, is a strepsirrhine primate and a species of slow loris native to Indonesia, West Malaysia, southern Thailand and Singapore. It measures 27 to 38 cm from head to tail and weighs between 599 and 685 g. Like other slow lorises, it has a wet nose (rhinarium), a round head, small ears hidden in thick fur, a flat face, large eyes and a vestigial tail.

<span class="mw-page-title-main">Hispaniolan solenodon</span> Species of mammal endemic to Hispaniola

The Hispaniolan solenodon, also known as the agouta, is a small, furry, shrew-like mammal endemic to the Caribbean island of Hispaniola. Like other solenodons, it is a venomous, insect-eating animal that lives in burrows and is active at night. It is an elusive animal and was only first described in 1833; its numbers are stable in protected forests but it remains the focus of conservation efforts.

<span class="mw-page-title-main">Fel d 1</span> Secretoglobin protein

Fel d 1 is a secretoglobin protein complex that, in cats, is encoded by the CH1 and CH2 genes.

<span class="mw-page-title-main">Bengal slow loris</span> Species of primate

The Bengal slow loris or northern slow loris is a strepsirrhine primate and a species of slow loris native to the Indian subcontinent and Indochina. Its geographic range is larger than that of any other slow loris species. Considered a subspecies of the Sunda slow loris (N. coucang) until 2001, phylogenetic analysis suggests that the Bengal slow loris is most closely related to the Sunda slow loris. However, some individuals in both species have mitochondrial DNA sequences that resemble those of the other species, due to introgressive hybridization. It is the largest species of slow loris, measuring 26 to 38 cm from head to tail and weighing between 1 and 2.1 kg. Like other slow lorises, it has a wet nose (rhinarium), a round head, flat face, large eyes, small ears, a vestigial tail, and dense, woolly fur. The toxin it secretes from its brachial gland differs chemically from that of other slow loris species and may be used to communicate information about sex, age, health, and social status.

<span class="mw-page-title-main">Philippine slow loris</span> Species of primate

The Philippine slow loris is a strepsirrhine primate and a species of slow loris that is native to the north and east coastal areas of the island of Borneo, as well as the Sulu Archipelago in the Philippines. The species was first named as the Bornean slow loris in 1892, but lumped into the widespread Sunda slow loris (N. coucang) in 1952. However, it was promoted to full species status – again as the Bornean slow loris – based on molecular analysis in 2006. In 2013, two former subspecies of the Bornean slow loris were elevated to species status, and a new species—N. kayan—was recognized among the Bornean population.

<span class="mw-page-title-main">Conservation of slow lorises</span> Conservation management of the nocturnal primates in Asia

Slow lorises are nocturnal strepsirrhine primates in the genus Nycticebus that live in the rainforests of South and Southeast Asia. They are threatened by habitat loss and fragmentation from deforestation, selective logging, and slash-and-burn agriculture, as well as by collection and hunting for the wildlife trade, including the exotic pet trade, and for use in traditional medicine and as bushmeat. Because of these and other threats, all five species of slow loris are listed as either "Vulnerable" or "Endangered" by the International Union for Conservation of Nature (IUCN). Their conservation status was originally listed as "Least Concern" in 2000 because of imprecise population surveys and the frequency in which these primates were found in animal markets. Because of their rapidly declining populations and local extinctions, their status was updated and in 2007 the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) elevated them to Appendix I, which prohibits international commercial trade. Local laws also protect slow lorises from hunting and trade, but enforcement is lacking in most areas.

<i>Nycticebus kayan</i> Species of primate

The Kayan River slow loris is a strepsirrhine primate and a species of slow loris that is native to the northern and central highland region of the island of Borneo. The species was originally thought to be a part of the Bornean slow loris (N. menagensis) population until 2013, when a study of museum specimens and photographs identified distinct facial markings, which helped to differentiate it. It is distinguished by the high contrast of its black and white facial features, as well as the shape and width of the stripes of its facial markings.

<i>Nycticebus bancanus</i> Species of primate

The Bangka slow loris is a strepsirrhine primate and a species of slow loris that is native to southwestern Borneo and the island of Bangka. Originally considered a subspecies or synonym of the Bornean slow loris (N. menagensis), it was promoted to full species status in 2013 when a study of museum specimens and photographs identified distinct facial markings, which helped to differentiate it as a separate species. It is distinguished by the crimson red fur on its back, light-colored facial features, as well as the shape and width of the stripes of its facial markings.

<i>Nycticebus borneanus</i> Species of primate

Nycticebus borneanus, the Bornean slow loris, is a strepsirrhine primate and a species of slow loris that is native to central south Borneo in Indonesia. Formerly considered a subspecies or synonym of N. menagensis, it was promoted to full species status in 2013 when a study of museum specimens and photographs identified distinct facial markings, which helped to differentiate it as a separate species. It is distinguished by its dark, contrasting facial features, as well as the shape and width of the stripes of its facial markings.

<span class="mw-page-title-main">Toxungen</span>

Toxungen comprises a secretion or other body fluid of one or more biological toxins that is transferred by one animal to the external surface of another animal via a physical delivery mechanism. Toxungens can be delivered through spitting, spraying, or smearing. As one of three categories of biological toxins, toxungens can be distinguished from poisons, which are passively transferred via ingestion, inhalation, or absorption across the skin, and venoms, which are delivered through a wound generated by a bite, sting, or other such action. Toxungen use offers the evolutionary advantage of delivering toxins into the target's tissues without the need for physical contact.

Anna Nekaris is a British anthropologist and professor in primate conservation in the School of Law and Social Sciences at Oxford Brookes University. She is interested in the conservation of Asian nocturnal animals and African nocturnal primates. She was appointed an Officer of the Order of the British Empire in the 2024 New Year Honours.

References

Notes

  1. ↑ Dufton, Mark (1992). "Venomous mammals". Pharmacology and Therapeutics. 53 (2): 199–215. doi:10.1016/0163-7258(92)90009-o. PMID   1641406.
  2. 1 2 3 Jørn H. Hurum, Zhe-Xi Luo, and Zofia Kielan-Jaworowska, Were mammals originally venomous?, Acta Palaeontologica Polonica 51 (1), 2006: 1-11
  3. ↑ Gong, Enpu; Larry B Martin; David A Burnham; Amanda R Falk (12 January 2010). "The birdlike raptor Sinornithosaurus was venomous". Proc. Natl. Acad. Sci. USA. 107 (2): 766–768. Bibcode:2010PNAS..107..766G. doi: 10.1073/pnas.0912360107 . PMC   2818910 . PMID   20080749.
  4. ↑ "Venomous Amphibians (Page 1) - Reptiles (Including Dinosaurs) and Amphibians - Ask a Biologist Q&A". Askabiologist.org.uk. Retrieved 2010-07-28.
  5. ↑ Robert T. Nowak & Edmund D. Brodie Jr. (1978). "Rib penetration and associated antipredator adaptations in the salamander Pleurodeles waltl (Salamandridae)". Copeia. 1978 (3): 424–429. doi:10.2307/1443606. JSTOR   1443606.
  6. 1 2 3 4 Ligabue-Braun, R.; Verli, H.; C.R., Carlini (2012). "Venomous mammals: A review". Toxicon. 59 (7–8): 680–695. doi:10.1016/j.toxicon.2012.02.012. PMID   22410495.
  7. ↑ Fox, Richard C.; Scott, Craig S. (June 2005). "First evidence of a venom delivery apparatus in extinct mammals". Nature. 435 (7045): 1091–1093. Bibcode:2005Natur.435.1091F. doi:10.1038/nature03646. ISSN   1476-4687. PMID   15973406. S2CID   4322624.
  8. ↑ Orr, Caley M.; Delezene, Lucas K.; Scott, Jeremiah E.; Tocheri, Matthew W.; Schwartz, Gary T. (2007-06-12). "The comparative method and the inference of venom-delivery systems in fossil mammals". Journal of Vertebrate Paleontology. 27 (2): 541–546. doi:10.1671/0272-4634(2007)27[541:TCMATI]2.0.CO;2. ISSN   0272-4634. S2CID   45645935.
  9. 1 2 3 Folinsbee, Kaila E.; Müller, Johannes; Reisz, Robert R. (2007-06-12). "Canine grooves: morphology, function, and relevance to venom". Journal of Vertebrate Paleontology. 27 (2): 547–551. doi:10.1671/0272-4634(2007)27[547:CGMFAR]2.0.CO;2. ISSN   0272-4634. S2CID   54602365.
  10. ↑ Furió, Marc; Agustí, Jordi; Mouskhelishvili, Alexander; Sanisidro, Óscar; Santos-Cubedo, Andrés (2010-05-18). "The paleobiology of the extinct venomous shrew Beremendia (Soricidae, Insectivora, Mammalia) in relation to the geology and paleoenvironment of Dmanisi (Early Pleistocene, Georgia)". Journal of Vertebrate Paleontology. 30 (3): 928–942. Bibcode:2010JVPal..30..928F. doi:10.1080/02724631003762930. ISSN   0272-4634. S2CID   86133561.
  11. ↑ Turvey, Samuel T. (2010-07-14). "Evolution of non-homologous venom delivery systems in West Indian insectivores?". Journal of Vertebrate Paleontology. 30 (4): 1294–1299. Bibcode:2010JVPal..30.1294T. doi:10.1080/02724634.2010.483541. ISSN   0272-4634. S2CID   86625812.
  12. 1 2 Dufton, M.J. (1992). "Venomous mammals". Pharmacol. Ther. 53 (2): 199–215. doi:10.1016/0163-7258(92)90009-o. PMID   1641406.
  13. 1 2 Barua, Agneesh; Mikheyev, Alexander S. (2021-04-06). "An ancient, conserved gene regulatory network led to the rise of oral venom systems". Proceedings of the National Academy of Sciences. 118 (14): e2021311118. Bibcode:2021PNAS..11821311B. doi: 10.1073/pnas.2021311118 . ISSN   0027-8424. PMC   8040605 . PMID   33782124.
  14. ↑ Pournelle, G.H. (1968). "Venomous Animals and Their Venoms". In W. Bücher; E.E. Buckley; V. Deulofeu (eds.). Classification, biology, and description of the venom apparatus of insectivores of the genera Solenodon, Neomys, and Blarina. Academic Press, New York. pp. 31–42. ISBN   9780121389024.
  15. ↑ A-Z-Animals.com. "Mole (Talpidae) - Animals - A-Z Animals - Animal Facts, Information, Pictures, Videos, Resources and Links". A-Z Animals. Retrieved 2010-07-28.
  16. ↑ Mukherjee, Sarah (2008-01-25). "Science/Nature | Searching for nature's tunnellers". BBC News. Retrieved 2010-07-28.
  17. ↑ Whittington, C.M.; Belov, K. (2009). "Platypus venom genes expressed in non-venom tissues". Australian Journal of Zoology. 57 (4): 199–202. doi:10.1071/zo09046.
  18. 1 2 Whittington, Camilla M.; Belov, Katherine (April 2014). "Tracing Monotreme Venom Evolution in the Genomics Era". Toxins. 6 (4): 1260–1273. doi: 10.3390/toxins6041260 . PMC   4014732 . PMID   24699339.
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  20. 1 2 3 Nekaris, K. Anne-Isola; Moore, Richard S.; Rode, E. Johanna; Fry, Bryan G. (2013-09-27). "Mad, bad and dangerous to know: the biochemistry, ecology and evolution of slow loris venom". Journal of Venomous Animals and Toxins Including Tropical Diseases. 19 (1): 21. doi: 10.1186/1678-9199-19-21 . ISSN   1678-9199. PMC   3852360 . PMID   24074353.
  21. ↑ Nekaris, K. Anne-Isola; Nijman, Vincent (2022). "A new genus name for pygmy lorises, Xanthonycticebus gen. nov. (Mammalia, primates)". Zoosystematics and Evolution. 98 (1): 87–92. doi: 10.3897/zse.98.81942 . ISSN   1860-0743.
  22. ↑ Primates, All The World's. "Colin Groves' Taxonomy Introduction - All The World's Primates". www.alltheworldsprimates.org. Retrieved 2018-04-21.
  23. 1 2 3 Alterman, Lon (1995). "Toxins and Toothcombs: Potential Allospecific Chemical Defenses in Nycticebus and Perodicticus". Creatures of the Dark. Springer, Boston, MA. pp. 413–424. doi:10.1007/978-1-4757-2405-9_24. ISBN   9781441932501.
  24. ↑ Grow, Nanda B.; Wirdateti; Nekaris, K.A.I. (2015). "Does toxic defence in Nycticebus spp. relate to ectoparasites? The lethal effects of slow loris venom on arthropods". Toxicon. 95: 1–5. doi:10.1016/j.toxicon.2014.12.005. PMID   25528386.
  25. ↑ Gardiner, Matthew; Samuel, Priya; Nekaris, K.A.I (2017). "The Cytotoxic Effect of Slow Loris (Nycticebus) Venom on Human Cancer". Folia Primatologica. 88 (2): 10 – via google scholar.
  26. ↑ Fuller, Grace; Lukas, Kristen E.; Kuhar, Christopher; Dennis, Patricia M. (2014-03-24). "A retrospective review of mortality in lorises and pottos in North American zoos, 1980-2010". Endangered Species Research. 23 (3): 205–217. doi: 10.3354/esr00568 . ISSN   1863-5407. S2CID   37481446.
  27. ↑ Madani, George; Nekaris, K. Anne-Isola (2014-10-02). "Anaphylactic shock following the bite of a wild Kayan slow loris (Nycticebus kayan): implications for slow loris conservation". Journal of Venomous Animals and Toxins Including Tropical Diseases. 20 (1): 43. doi: 10.1186/1678-9199-20-43 . ISSN   1678-9199. PMC   4192448 . PMID   25309586.
  28. ↑ Gardiner, Matthew; Weldon, Ariana; Gibson, Nancy; Poindexter, Stephanie; Nekaris, K.A.I. "Survey of practitioners handling slow lorises (Primates: Nycticebus): an assessment of the harmful effects of slow loris bites – Journal of Venom Research". jvenomres.co.uk. Retrieved 2018-04-21.
  29. ↑ "Jungle Gremlins of Java, 2011-2012, Natural World - BBC Two". BBC. Retrieved 2018-04-21.
  30. ↑ Brodie, E.D. (1977). "Hedgehogs use toad venom in their own defence". Nature. 268 (5621): 627–628. Bibcode:1977Natur.268..627B. doi:10.1038/268627a0. S2CID   4293159.
  31. 1 2 Morelle, Rebecca (2011). "African crested rat uses poison trick to foil predators". BBC.co.uk. Retrieved November 2, 2013.
  32. ↑ Welsh, J. (2011). "Giant rat kills predators with poisonous hair". LiveScience. Retrieved August 2, 2011.
  33. ↑ "Rat makes its own poison from toxic tree". University of Oxford. 2011. Archived from the original on November 6, 2013. Retrieved November 2, 2013.
  34. ↑ Kingdon, J.; Agwanda, B.; Kinnaird, M.; O'Brien, T.; Holland, C.; Gheysens, T.; Boulet-Audet, M.; Vollrath, F. (2011). "A poisonous surprise under the coat of the African crested rat". Proc. R. Soc. B. 279 (1729): 675–680. doi:10.1098/rspb.2011.1169. PMC   3248729 . PMID   21813554.

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