Amplexus

Amplexus (Latin "embrace") is a type of mating behavior exhibited by some externally fertilizing species (chiefly amphibians and horseshoe crabs) in which a male grasps a female with his front legs as part of the mating process, and at the same time or with some time delay, he fertilizes the eggs as they are released from the female's body.[1] In amphibians, females may be grasped by the head, waist, or armpits, and the type of amplexus is characteristic of some taxonomic groups. Amplexus involves direct contact between male and female, distinguished from other forms of external fertilization such as broadcast spawning, where sperm and eggs are freely shed into water without direct contact by individuals. In order for amplexus to be initiated, male frogs must first find a mate by attracting one through calls, typically in the evening.[2] Once a male has successfully attracted a mate the process of amplexus begins while the unsuccessful males are forced to continue their search for a mate through further calls.[2] The competition for a female to mate with among males is considered intense as it is not uncommon for a male amphibian to attack an already-amplexed pair of amphibians.[3] When a male amphibian attacks an already-amplexed pair of amphibians he is trying to force the other male to release its grasp of the female so he can then mate with her.[4] Male amphibians are also known to show mate-guarding behaviour, which is shown after amplexus and it is the males attempt to prevent the female amphibian from mating with other males.[4] The duration of amplexus has been found to vary across different species as in some species it has been found to last for many days but in other species, it has been found to last a few hours.[5] Despite the variation in the duration of amplexus across different species, typically all the species that exhibit this behaviour have to use their forelimb muscles for the whole duration of amplexus.[5]Additionally, amplexus is thought to help with the alignment of the reproductive tracts of both males and females which furthermore contributes to the achievement of successful fertilization.[1]
Amphibians


Anurans
Amplexus chiefly occurs aquatically, but some more terrestrial anurans (frogs and toads) like the disc-tongued frogs (Discoglossidae) perform amplexus on land. In crown-group anurans like the true frogs (Ranidae), the tree frogs (Hylidae), and the true toads (Bufonidae), amplexus is axillary (in the armpits). Other anurans (the Archaeobatrachia, Sooglossidae and Myobatrachidae), show the ancestral state which is inguinal or lumbar amplexus (abdominal, in front of the hindlegs). Some species show cephalic amplexus where the head of the female is held while others show complete lack of amplexus.[6]
In most anurans, the males deposit sperm onto the eggs as they are being laid, however males of the genus Ascaphus possess an intromittent organ, unique among anurans, for internal fertilization. Internal fertilization does occur in a few other genera, including Nectophrynoides, Mertensophryne, and Eleutherodactylus.[6][7]
Newts
In the case of newts the process of amplexus is often observed soon after the newts become seasonally active. In the western USA, for example, this time is typically soon after the onset of the winter rainy season, when intermittent streams and vernal pools become available as breeding habitat. The rough-skinned newt is a specific widespread example of a newt in the western USA that can be observed in quiet stream pools and shallow ponds engaging in amplexus.[8]
Horseshoe crabs

Amplexus occurs in all four species of horseshoe crab. The first pair of walking legs is used to tightly clasp the female in all species, and the second pair is also employed in all but Limulus polyphemus.[9]
Other animals
Fossil evidence suggests that a certain euthycarcinoid (an extinct arthropod) from the Cambrian may also have mated by amplexus.[10]
See also
References
- ^ a b Mollov, Ivelin; Popgeorgiev, Georgi; Naumov, Borislav; Tzankov, Nikolay; Stoyanov, Andrei (2010). "Cases of abnormal amplexus in anurans (Amphibia: Anura) from Bulgaria and Greece". Biharean Biologist.
- ^ a b McLister, James (2003). "The metabolic cost of amplexus in the grey tree frog (Hyla versicolor): assessing the energetics of male mating success". NRC Canada. doi:10.1139/Z03-013.
- ^ Sztatecsny, Marc; Preininger, Doris; Freudmann, Anita; Loretto, Matthias-Claudio; Maier, Franziska; Hödl, Walter (2012-09-23). "Don't get the blues: conspicuous nuptial colouration of male moor frogs (Rana arvalis) supports visual mate recognition during scramble competition in large breeding aggregations". Behavioral Ecology and Sociobiology. 66 (12): 1587–1593. doi:10.1007/s00265-012-1412-6. ISSN 0340-5443.
- ^ a b Sztatecsny, Marc; Preininger, Doris; Freudmann, Anita; Loretto, Matthias-Claudio; Maier, Franziska; Hödl, Walter (2012-09-23). "Don't get the blues: conspicuous nuptial colouration of male moor frogs (Rana arvalis) supports visual mate recognition during scramble competition in large breeding aggregations". Behavioral Ecology and Sociobiology. 66 (12): 1587–1593. doi:10.1007/s00265-012-1412-6. ISSN 0340-5443.
- ^ a b McLister, James (2003). "The metabolic cost of amplexus in the grey tree frog (Hyla versicolor): assessing the energetics of male mating success". NRC Canada. doi:10.1139/Z03-013.
- ^ a b Duellman, W. E. and L. Trueb. 1986. Biology of Amphibians. New York: McGraw-Hill Publishing Company.
- ^ Linzey, D. 2001. Vertebrate Biology, McGraw Hill Publishers, New York.
- ^ C. Michael Hogan (2008) Rough-skinned Newt (Taricha granulosa) Archived 2009-05-27 at the Wayback Machine, Globaltwitcher, ed. N. Stromberg
- ^ Botton, Mark L.; Shuster, Carl N.; Sekiguchi, Koichi; Sugita, Hiroaki (February 1996). "Amplexus and Mating Behavior in the Japanese Horseshoe Crab, Tachypleus tridentatus". Zoological Science. 13 (1): 151–159. doi:10.2108/zsj.13.151.
- ^ Collette, Joseph H.; Gass, Kenneth C.; Hagadorn, James W. (May 2012). "Protichnites eremita Unshelled? Experimental Model-Based Neoichnology and New Evidence for A Euthycarcinoid Affinity for This Ichnospecies". Journal of Paleontology. 86 (3): 442–454. doi:10.1666/11-056.1.