Nuptial gift: Difference between revisions

Nuptial gifts are common in insects and other invertebrates, such as [[butterflies]], [[Ceratitis capitata|fruit flies]] and [[katydid]]s. They are less common in [[spider]]s, though the spider species ''[[Pisaura mirabilis]]'' is known for nuptial gift giving.<ref>{{cite journal |firstfirst1=Line Spinner |lastlast1=Hansen |first2=Sofia Fernández |last2=Gonzáles |first3=Søren |last3=Toft |first4=Trine |last4=Bilde |year=2008 |title=Thanatosis as an adaptive male mating strategy in the nuptial gift–giving spider ''Pisaura mirabilis'' |journal=Behavioral Ecology |volume=19 |issue=3 |pages=546–551 |doi=10.1093/beheco/arm165|doi-access=free }}</ref> However, this may in part be due to an experimental bias due to the ease of rearing it for experiments.<ref name ="prokop" /> Other taxa may commonly exhibit gift-giving behavior. Thus, more research is required to evaluate the scope of nuptial gifts in arachnid taxa. In many species of insects, birds and mammals, males acquire and donate food to females either before, during, or after copulation (termed '''mate provisioning''', courtship feeding, nuptial gift giving, or meat-for-sex). Males may relinquish body parts, produce glandular secretions, or share prey or other food to gain fitness benefits via natural and sexual selection.<ref>{{cite journal |journal= Animal Behaviour |year=2004 |volume=64 |issue=4 |pages=603–614 |title= A conceptual framework for nonkin food sharing: timing and currency of benefits|authorauthor1=Stevens, Jeffrey R. and |author2=Gilby, Ian C. |name-list-style=amp |doi=10.1016/j.anbehav.2003.04.012|s2cid=11491968 |url=http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1009&context=psychstevens }}</ref>

Courtship feeding is particularly common among many bird species.<ref>{{cite journal |title=Courtship Feeding in Birds |first=David |last=Lack |journal=The Auk |volume=57 |date=April 1940 |pages=169–178 |jstor=4078744 |issue=2 |doi=10.2307/4078744|doi-access=free }}</ref>

{{anchorMain|Great Grey Shrike}}

[[File:Pisaura mirabilis on Plantago lanceolata.jpg|thumb|''[[Pisaura mirabilis]]'' on ''[[Plantago lanceolata]]'' inflorescence]]

==== ''Pisaura mirabilis'' ====

In spiders, nuptial gifts in the form of prey are restricted to a few species from two families belonging to the superfamily [[Lycosoidea]]: [[Nursery web spider|Pisauridae]] and [[Trechaleidae]]. In both families, the male courts the female by offering a prey wrapped in silk and mating occurs while the female consumes the gift. In the species ''[[Pisaura mirabilis]]'' (Pisauridae), the gift functions as a mating effort that increases male mating success. The nuptial gift consists of a prey the male has caught and wrapped up in silk. The male offers this gift during courtship, and if the female accepts the invitation, she grabs the wrapped prey. While the female is eating, the male inserts the mating organ, and sperm is transferred.<ref>{{cite journal |journal= Behavioral Ecology |year=2001 |volume=12 |issue=6 |pages=691–697 |title= Nuptial gift in the spider Pisaura mirabilis maintained by sexual selection |author=Stålhandske, Pia |doi= 10.1093/beheco/12.6.691 |doi-access=free }}</ref>

A similar function was recently suggested for the trechaleid spider, ''Paratrechalea ornata''. In both species, males can obtain mating without a gift, but male mating success increases dramatically when a gift is offered. There is a positive correlation between male size and mating duration, and an increasing mating duration when there is a gift offered.<ref>{{Cite journal |last1=Pandulli-Alonso |first1=Irene |last2=Tomasco |first2=Ivanna H. |last3=Albo |first3=Maria J. |date=March 2022 |title=The handsome liar: Male spiders offering worthless gifts can benefit increasing mating duration |url=https://onlinelibrary.wiley.com/doi/10.1111/eth.13258 |journal=Ethology |language=en |volume=128 |issue=3 |pages=215–222 |doi=10.1111/eth.13258 |bibcode=2022Ethol.128..215P |issn=0179-1613}}</ref> In ''Pisaura mirabilis'', the male pushes up the female during mating and performs alternate pedipalp insertions into the female sperm storage organs place ventrally on the abdomen. After each insertion the male returns to a face-to-face position with the female, grabbing the gift in the chelicerae. Females often control mating duration and they often attempt to run away with the gift upon terminating the copulation.<ref name=Worthless>{{cite journal|doi=10.1186/1471-2148-11-329|title=Worthless donations: Male deception and female counter play in a nuptial gift-giving spider|year=2011|last1=Albo|first1=Maria J|last2=Winther|first2=Gudrun|last3=Tuni|first3=Cristina|last4=Toft|first4=Søren|last5=Bilde|first5=Trine|journal=BMC Evolutionary Biology|volume=11|issue=1 |pages=329|pmid=22082300|pmc=3228764 |doi-access=free |bibcode=2011BMCEE..11..329A }}</ref>

Females are less likely to succeed in stealing a wrapped – rather than an unwrapped – gift due to the silk wrapping of the gift. The silk wrapping facilitates male handling and control over the gift, as it facilitates a stronger hold of the silk covered package versus andan unwrapped insect. Male spiders have a unique opportunity for gift manipulation through the gift wrapping trait, for example by preventing female assessment of the gift content. By disguising the gift content, males may deceive females to copulate, while the female attempts to consume the gift. In ''Paratrechalea ornata'', males were observed wrapping prey carrion and occasionally inedible items such as plant seeds. In ''Pisaura mirabilis'', males have been reported to carry gifts containing empty arthropod exoskeletons or plant parts,: gifts of no nutritional value.<ref name=Worthless/> However, some studies have shown that male spiders rarely cheat in nature.<ref name ="prokop">{{cite journal|last1=Prokop|first1=P|last2=Maxwell|first2=MR|title=Gift carrying in the spider Pisaura mirabilis: nuptial gift contents in nature and effects on male running speed and fighting success.|journal=Animal Behaviour|date=2012|volume=83|issue=6|pages=1395-13991395–1399|doi=10.1016/j.anbehav.2012.03.007|doi-access=free}}</ref> Cogent reasons for this disparate behavior are being explored. It is possible there are confounding factors which do not account for ecological implications. Other research suggests that sensory bias is not the main contributing force behind gift selection in ''P. mirabilis''. Interestingly, femalesFemales did not preferentially select unwrapped gifts over wrapped gifts. <ref name ="ghislandi">{{cite journal|last1=Ghislandi|first1=PG|last2=Albo|first2=MJ|last3=Tuni|first3=C|last4=Bilde|first4=T|title=Evolution of deceit by worthless donations in a nuptial gift-giving spider.|journal=Current Zoology|date=2014|volume=60|issue=1|pages=43–51|doi=10.1093/czoolo/60.1.43|doi-51access=free}}</ref>

==== Paratrechalea''Pisaurina ornatamira'' ====

[[Tettigoniidae|Bushcricket]] males offer a spermatophylax thatcontaining containsan ampulla. The nuptial gift is also protein -rich, which the females ingest into their reproductive tract. The size of the nuptial gifts positively influences the females refractory period and the males reproduction success. The size of the gift depends on the ampulla and serves as a sperm protection. In bushcrickets ''Ephippiger ephippiger'', the females prefer older males, who have larger spermophores and better nutritional value during mating. The nutritional value is related to the female's metabolism, which stands as a benefit for females feeding on the semaphoresspermatophores. Sometimes males produce lower dosages of sperm with a lower nutritional value the fourth time they mate.<ref>{{cite journal |journal= Evolution |year=1993 |volume=47 |issue=4| pages=1203–1212 |title= Spermatophore size in bushcrickets: Comparative evidence for nuptial gifts as a sperm protection device|author=Wedell, Nina|jstor= 2409986 |doi= 10.2307/2409986 |pmid=28564284 }}</ref> It may be possible the females are using age and gift quality as a proxy for mates with good genes as their offspring are likely to have high relative fitness. The spermatophore provides protection to the ampulla by preventing it from being removed prematurely.<ref name=":52">Jay McCartney, Murray A. Potter, Alastair W. Robertson, Kim Telscher, Gerlind Lehmann, Arne Lehmann, Dagmar von-Helversen, Klaus Reinhold, Roland Achmann and Klaus-Gerhard Heller, 2008, Understanding Nuptial Gift Size in Bush-Crickets: An Analysis of the Genus Poecilimon (Tettigoniidae: Orthoptera), Journal of Orthoptera Research, Volume 17, pages 231-242</ref> It is also hypothesized to provide direct nutritional benefit to the offspring through the paternal investment hypothesis.<ref name=":52" />

=== Ornate Mothmoth ===

During mating in the [[Utetheisa ornatrix|Ornateornate Mothmoth]] (''Utetheisa ornatrix''), males provide the female with a spermatophore containing nutrients, sperm and alkaloids that serve as chemical defense from predators. Such nuptial gift accounts for up to 10% of the male's body weight and represents the total parental investment the male provides.<ref>{{cite journal|last1=Iyengar|first1=Vikram K.|last2=Eisner|first2=Thomas |title=Female choice increases offspring fitness in an Arctiid Moth (Utetheisa Ornatrix)|journal=Proceedings of the National Academy of Sciences|year=1999|volume=96|issue=26|pages=15013–15016|doi=10.1073/pnas.96.26.15013|pmid=10611329|pmc=24764|bibcode=1999PNAS...9615013I|doi-access=free}}</ref> Females receive spermatophores from several males and direct a postcopulatory selection process in which they decide what sperm will fertilize their eggs.<ref>{{cite journal|last1=Kellya|first1=Caitlin A.|last2=Norbutusb |first2=Amanda J. |last3=Lagalanteb |first3=Anthony F. |last4=Iyengara |first4=Vikram K. |title=Male courtship pheromones as indicators of genetic quality in an arctiid moth (Utetheisa ornatrix)|journal=Behavioral Ecology|year=2012|volume=23|issue=5|pages=1009-10141009–1014|doi=10.1093/beheco/ars064|doi-access=free}}</ref>

=== Six-spot''Ostrinia Burnetscapulalis'' ===

[[File:Panorpa communis with prey Diogma glabrata glabrata.jpg|thumb|left|[[Scorpionfly]] with prey]]

In the scorpionfly (''[[Panorpa cognata]]''), males offer a salivary secretion as a nuptial gift before copulation. Exchange of the salivary secretion takes place after prolonged courtship interactions. The nuptial gift is more likely to be accepted by the female if premating duration is long. Males in poor condition with a limited supply of saliva may deliberately delay initiating copulations to decrease the probability that their costly gift is rejected and, thus, wasted. Males in good condition with ample mating resources, on the other hand, may afford the risk of wasting a salivary mass and therefore take every opportunity to mate.<ref>{{cite journal |journal= Animal Behaviour |year=2009 |volume=78 |issue=2 |pages=491–497 |title= Should I stay or should I go? Condition- and status-dependent courtship decisions in the scorpionfly Panorpa cognata

|author=Engqvist, Leif|doi=10.1016/j.anbehav.2009.05.021|s2cid=51800821 |url=https://pub.uni-bielefeld.de/record/2395690 }}</ref>

Male sperm offers many nutrients to increase a female's lifetime and egg production. These nutrients include acids or sodium. Male sperm also protects females and their eggs from predators. The females would also gain a net benefit from a male's sperm. Female recipients are supposed to produce bigger offspring than those females who did not receive nuptial gifts. The eggs of female spiders who receive nuptial gifts may hatch at a higher rate than those without nuptial gifts.<ref name ="ghislandi" /> Historically, nuptial gifts were seen as nutritional substances given to females from males during mating.<ref>{{citeCite journal | last1 =Ghislandi Albo | first1 =PG M. A. J. | last2 =Albo Costa | first2 =MJ F. G. |last3 doi =Tuni|first3=C|last4=Bilde|first4=T 10.1016/j.anbehav.2010.01.018 | title =Evolution ofNuptial deceitgift-giving bybehaviour worthlessand donationsmale inmating aeffort nuptialin gift-givingthe Neotropical spider. ''Paratrechalea ornata'' (Trechaleidae) | journal=Current Zoology|date=2014 Animal Behaviour | volume =60 79 | issue =1 5 | pages =43-51}}</ref> Historically,1031 nuptial| giftsyear were= seen2010 as| nutritionals2cid substances= given39742664 to| femalesurl from= males during mating.<ref>{{Cite doihttp:/10/web.1016mnstate.2Fjedu/wisenden/Paper%208%20Spider%20nuptial%20gifts.anbehav.2010.01.018pdf}}</ref> The male benefits from a net fitness. Although the gifts are costly to find or produce, the gifts will increase attraction and copulation with other females. One of the more recently identified costs to males is reduced running speed due to gift-carrying.<ref>{{cite journal|last1=Prokop|first1=P|last2=Maxwell|first2=MR|title=Giftname carrying in the spider Pisaura mirabilis: nuptial gift contents in nature and effects on male running speed and fighting success.|journal=Animal"prokop" Behaviour|date=2012|volume=83|issue=6|pages=1395-1399|doi=10.1016/j.anbehav.2012.03.007}}</ref> This cost may be exacerbated in areas of high predation. Nuptial gifts can benefit a male by increasing his paternity share when females are promiscuous. In some insects, nuptial gifts allow the male to copulate longer and transfer more sperm to the female.<ref name=gwynne/> In fruit flies, katydids, and [[scorpion fliesscorpionflies]], nuptial gifts contain substances that reduce a female’sfemale's receptivity to additional matings. While nuptial gifts also may boost female fecundity, from a male’smale's perspective, such investment will only be beneficial if it increases the number of his own offspring.<ref name="Lewis et al"/> In bell crickets, nuptial gifts may be necessary to avoid injury or death by cannibalizing females.<ref>{{cite journal|last1=Kuriwada|first1=T|last2=Kasuya|first2=E|title=Nuptial gifts protect male bell crickets from female aggressive behavior|journal=Behavioral Ecology|date=2012|volume=23|issue=2|pages=302-306302–306|doi=10.1093/beheco/arr186|doi-access=|hdl=10.1093/beheco/arr186|hdl-access=free}}</ref> This additional benefit allows gift-giving males to surpass the fitness of other males. In this way, females are exploiting the inherent sexual dimorphism of their species.

In species where males provide females with a nuptial gift during mating, there is a particular scope for males to manipulate females to acquire matings and prolong copulation to enhance their fertilization success. Typically, females control the duration and volume of sperm transfer throughout the mating process. Research has suggested that the gifts presented by males temporarily obstruct the female's capacity to manage the copulation event.<ref>{{cite journal|last1=Sakaluk|first1=SK|title=Sensory exploitation as an evolutionary origin to nuptial food gifts in insects.|journal=Proc. R. Soc. Lond. B|date=2000|volume=267|issue=1441|pages=339-343339–343|doi=10.1098/rspb.2000.1006|pmid=10722214|pmc=1690542}}</ref> Female choice for males with nuptial gifts could lead to the evolution of male deception by the use of toketoken gifts. For instance, males can decrease the costs of mating by re-using gifts or by offering worthless gifts. Males of some dance flies may deceive females by offering inadequate or false gifts. Although males that offer inedible gifts run a higher risk of being rejected and may suffer from shorter mating periods compared to males offering edible gifts the chance of acquiring an extra mating should make deception an attractive strategy for males. Hence, males of the dance fly Rhamphomyia sulcata that use inedible token gifts to obtain mates are as successful as males offering small genuine gifts.<ref name=Worthless/>

When nuptial gifts are given it increases copulations and searches to find females to mate with. Albo and Costa conducted an experiment with ''Paratrechala ornata'' spiders to determine the function of the nuptial gift. Two groups of males were exposed to virgin females, 23 males with no nuptial gift (prey) and 21 males with a nuptial gift (prey). Mating ovulation were recorded and the researchers found that males that gave a nuptial gift had better mating success, longer copulations, and longer palpal insertions than those who did not give an nuptial gift. Longer copulations were associated with earlier egg sac construction and oviposition. The researchers' findings suggest that nuptial gift giving represents male mating effort for ''P. ornata''. Nuptial gifts would allow males to control copulation duration and to accelerate female oviposition, improving sperm supply and paternity, and minimizing possible costs of remating with polyandrous individuals.<ref name=Worthless/> Additionally, Prokop found that female mate choice is dependent on nuptial gifts rather than female reproductive status as an unmated or mated individual.<ref>{{cite journal|last1=Prokop|first1=P.|title=Insemination does not affect female mate choice in a nuptial feeding spider.|journal=Italian Journal of Zoology|date=2006|volume=73|issue=3|pages=197-201197–201|doi=10.1080/11250000600727741|doi-access=free}}</ref> This effect demonstrates sexual selection's ability to make one sex more discriminatory than the other, since females may negatively impact their output of offspring by refusing mating events with males that do not offer gifts.

=== SeeBenefits alsoto the female ===