239:), 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. Additionally, anurans species have been observed to engage in multiple amplexus, which can also be referred to as a mating ball, as many toads attach themselves to a female trying to initiate amplexus. However, multiple amplexus is not common among anurans, which could indicate that the costs associated with multiple amplexus are higher than the advantages associated with it. For female anurans, the idea of multiple amplexus would probably be more advantageous, because mating with more than one male would increase fertilization chances or increase offspring genetic diversity. Multiple amplexus would typically be common in explosive breeding amphibians, when there is a larger number of adults ready to breed at a breeding site in a short period of time. When this occurs female amphibians are viewed as a very important resource for males, as there are typically more males present than females, thus leading to more chances for multiple amplexus to occur.
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females, and for males these muscles contain more oxidative fibers, which could mean that amplexus involves an increased rate of aerobic metabolism. In addition to those forelimb muscles being larger in males, male frogs also typically have keratin pads or nuptial pads, which are located on their thumbs and contribute to the success of amplexus by assisting in gripping the female for the duration of amplexus. This amplexus behaviour allows the amphibians' cloacae to be in close contact, while gametes are released. Additionally, amplexus is thought to help with the alignment of the reproductive tracts of both males and females, which furthermore contributes to successful fertilization. A female amphibian may not always be receptive to a male amphibian who is trying to initiate amplexus, as the female may not be ready to produce eggs. When the female is not ready to engage in amplexus, she will simply vibrate her body, which will then be felt by the male who is clasped to her and he will then stop the amplexus behaviour.
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male amphibians. Additionally, it is believed that the gonadotropin-releasing hormone (GnRH) has an influence on amplexus in amphibians, as it has been found to produce or initiate this behaviour in many anuran amphibians. Furthermore, both arginine vasotocin and gonadotropin-releasing hormone have been found to be involved in the sexual behaviour of male amphibians. The stress hormone corticosterone has also been identified as associated with the sexual behaviour of amplexus. A study conducted on red-spotted newts found an acute increase in corticosterone for both males and females who were engaged in the behaviour of amplexus. However, overall it was found that males have a higher level of corticosterone compared to females, as amplexus is seen as more energetically costly to males. The increase in corticosterone found in females could be attributed to amplexus preventing them from foraging for food.
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female newt. Studies have shown that male newts who have deeper tail-fins have better control of females during amplexus and are also more successful in catching the females for amplexus. Additionally, it has been found that the probability of a male newt who has a deeper tail-fin to achieve amplexus is greater than those newts who do not contain a deeper tail-fin, as male newts tend to use their tails during male-male competition. When a male newt, who is unpaired, encounters a female and male newt engaged in amplexus, the unpaired newt will try to displace the paired male newt by using wrestling tactics. A study examining the wrestling behaviour of newts found that of the observed wrestling encounters, 90% were "won" by the paired male, meaning he would retain the female newt. The study found that the invading unpaired newt rarely successfully displaces the paired male newt, engaged in amplexus.
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the female horseshoe crab has a hard shell. Additionally, males who inhabit a clean shell are more likely to enter amplexus, compared to males who contain a dirty shell, as it seems that females have a preference for clean shells on males. A male horseshoe crab's claspers may also be an important factor of consideration for the initiation of amplexus. Since claspers are used for the attachment to the female, claspers that are in good condition are more successful for the initiation of amplexus. If a male horseshoe crab has a damaged or missing clasper, then that puts the male at a disadvantage and increases the probability of being displaced by other competing male crabs.
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for the duration of amplexus. Studies have found that this reproductive behaviour of amplexus can come with different fitness costs, due to the fact that amplexus can occur for prolonged periods of time. For instance, a study found that when a male amphibian is grasping and holding onto a female amphibian, this can lead to the impairment of the female's ability to move or to feed. After conducting experiments, researchers confirmed that amplexus does decrease a female's locomotor performance (e.g., swimming, walking) as well as the feeding rates. With regard to the cost to the male engaged in amplexus, male amphibians have been found to not feed at all during amplexus.
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during amplexus. In contrast, the male's anterior claspers have been found to attach to the female's opisthosoma as well, but on the lateral edges of the opisthosoma and function to resist displacement from environmental factors. Uniquely, amplexus is most likely to occur between horseshoe crabs when
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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. During amplexus in newts, males will typically show the behaviour of tail fanning and chin rubbing which is thought to prompt the mating receptivity of the
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The competition for a female mate among males is considered intense, and it is not uncommon for a male amphibian to attack an already-amplexed pair of amphibians. When a male amphibian attacks an amplexed pair of amphibians, he is trying to force the other male to release its grasp of the female, so
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Two major hormones have been identified to be involved in amplexus. The hormone arginine vasotocin (AVT) has been identified as having an effect on the calling behaviours produced by these male amphibians when searching for a female mate, as AVT increased the amount of calling behaviour produced in
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Amplexus has been found to involve different muscles in both male and female amphibians. The forelimb muscles in both males and females have been identified as the key muscles for amplexus that most species use. These forelimb muscles that are used during amplexus are typically larger in males than
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The duration of amplexus has been found to vary across species. In some species it may last for many days, while in others it may last a few hours. Despite the variation in the duration of amplexus across species, typically all species that exhibit this behaviour have to use their forelimb muscles
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Many types of amplexus are identified in the literature. However, two types of amplexus are more common than others, known as inguinal and axillary amplexus. These two types of amplexus have been classified based on the position of the male amphibian relative to the female. When a male amphibian
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Reedy, Aaron M.; Edwards, Alex; Pendlebury, Chloe; Murdaugh, Laura; Avery, Ryan; Seidenberg, Jake; Aspbury, Andrea S.; Gabor, Caitlin R. (2014). "An acute increase in the stress hormone corticosterone is associated with mating behavior in both male and female red-spotted newts, Notophthalmus
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during sexual maturity when the male moults; these modified claspers can then help during the process of amplexus. The male's pair of posterior claspers are known for having the ability to maintain long-term amplexus which have been found to always attach to the female's
321:. Horseshoe crabs typically go ashore for amplexus in high tide, and end up on beaches where the eggs are more protected. 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
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clasps a female around her waist (inguinal region) using his forelimbs, this is considered inguinal amplexus. By contrast, when a male amphibian clasps behind the forelimbs (axillary region) of the female, this is considered axillary amplexus.
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he can then mate with her. Male amphibians are also known to show mate-guarding behaviour, which is shown after amplexus, and it is the male's attempt to prevent the female amphibian from mating with other males.
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Duffy, Erin E.; Penn, Dustin J.; Botton, Mark L.; Brockmann, H. Jane; Loveland, Robert E. (2005-08-11). "Eye and clasper damage influence male mating tactics in the horseshoe crab, Limulus polyphemus".
282:, 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
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the eggs, as they are released from the female's body. In amphibians, females may be grasped by the head, waist, or armpits, and the type of amplexus is characteristic of some taxonomic groups.
972:
Leschen, Alison S.; Grady, Sara P.; Valiela, Ivan (2006). "Fecundity and spawning of the
Atlantic horseshoe crab, Limulus polyphemus, in Pleasant Bay, Cape Cod, Massachusetts, USA".
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Mollov, Ivelin; Popgeorgiev, Georgi; Naumov, Borislav; Tzankov, Nikolay; Stoyanov, Andrei (2010). "Cases of abnormal amplexus in anurans (Amphibia: Anura) from
Bulgaria and Greece".
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Chakraborti, S.; Nag, T.C.; Das, D.; Sanyal
Chatterjee, T.; De, S.K. (2014). "Cytokeratin localization in toe pads of the anuran amphibian Philautus annandalii (Boulenger, 1906)".
446:"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"
677:
Propper, Catherine R.; Dixon, Timothy B. (1997). "Differential
Effects of Arginine Vasotocin and Gonadotropin-Releasing Hormone on Sexual Behaviors in an Anuran Amphibian".
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Verrell, Paul A. (1986). "Wrestling in the red-spotted newt (Notophthalmus viridescens): resource value and contestant asymmetry determine contest duration and outcome".
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McLister, James (2003). "The metabolic cost of amplexus in the grey tree frog (Hyla versicolor): assessing the energetics of male mating success".
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McLister, James (2003). "The metabolic cost of amplexus in the grey tree frog (Hyla versicolor): assessing the energetics of male mating success".
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Able, David J. (1999-11-04). "Scramble competition selects for greater tailfin size in male red-spotted newts (Amphibia: Salamandridae)".
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Bowcock, Haley; Brown, GregoryP.; Shine, Richard (2009-02-23). "Beastly
Bondage: The Costs of Amplexus in Cane Toads (Bufo marinus)".
76:) 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
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Sztatecsny, Marc; Preininger, Doris; Freudmann, Anita; Loretto, Matthias-Claudio; Maier, Franziska; Hödl, Walter (2012-09-23).
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Amplexus involves direct contact between male and female, distinguished from other forms of external fertilization, such as
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Unshelled? Experimental Model-Based
Neoichnology and New Evidence for A Euthycarcinoid Affinity for This Ichnospecies".
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Lod, Thierry; Lesbarr, David (2004). "Multiple paternity in Rana dalmatina , a monogamous territorial breeding anuran".
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In most anurans, the males deposit sperm onto the eggs as they are being laid, however males of the genus
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Duellman, W. E. and L. Trueb. 1986. Biology of
Amphibians. New York: McGraw-Hill Publishing Company.
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Nussbaum, Ronald A. (1980). "Phylogenetic
Implications of Amplectic Behavior in Sooglossid Frogs".
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60:(Latin "embrace") is a type of
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125:
116:
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31:
1256:Sexual acts
363:Nuptial pad
334:opisthosoma
248:possess an
172:mating ball
169:Common toad
141:common frog
1235:Categories
600:(1): 1–5.
370:References
221:tree frogs
193:Costa Rica
131:Amphibians
113:Physiology
78:fertilizes
70:amphibians
1209:129234373
1154:0289-0771
1107:0289-0003
1012:0173-9565
951:0003-3472
903:0340-5443
816:0028-1042
748:0016-6480
734:: 57–63.
649:0040-8166
606:0018-0831
564:0045-8511
472:0340-5443
225:Bufonidae
157:Bufo bufo
150:European
139:European
1115:85269977
1057:85269977
1003:1912/906
959:53183312
911:34638425
859:Archived
832:21612515
824:14740104
756:25157790
707:41855944
657:24698093
572:83622844
490:23162205
357:See also
351:Cambrian
329:claspers
245:Ascaphus
122:Hormones
58:Amplexus
1189:Bibcode
1162:1398829
982:Bibcode
796:Bibcode
699:9367717
614:3891845
481:3496481
219:), the
217:Ranidae
205:anurans
199:Anurans
1251:Mating
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544:Copeia
488:
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264:, and
1246:Frogs
1205:S2CID
1158:S2CID
1111:S2CID
1053:S2CID
955:S2CID
907:S2CID
828:S2CID
703:S2CID
610:JSTOR
568:S2CID
280:newts
274:Newts
99:Types
1150:ISSN
1103:ISSN
1008:ISSN
947:ISSN
899:ISSN
820:PMID
812:ISSN
752:PMID
744:ISSN
695:PMID
653:PMID
645:ISSN
602:ISSN
560:ISSN
548:2009
486:PMID
468:ISSN
286:and
235:and
72:and
1197:doi
1142:doi
1093:doi
1043:doi
998:hdl
990:doi
939:doi
891:doi
804:doi
736:doi
732:208
687:doi
637:doi
552:doi
520:doi
476:PMC
458:doi
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