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>

{{anchorMain|Great Grey Shrike}}

The male [[great grey shrike]], a raptor-like passerine bird, gives prey ([[rodents]], [[birds]], [[lizards]], or large [[insects]]) to females immediately before copulation. Shrikes are well known for impaling prey on thorns and sharp sprigs. Great grey shrike females select a mate according to the size of prey impaled, with larders thus serving as an [[extended phenotype]] of a male. If the amount of food stored by the males can drive female [[mate choice]], food provided by males before copulation may also influence the female's decision to copulate. This applies both to the male's own partner and to other females.<ref>{{cite journal |journal= Animal Behaviour |year=2005 |volume=69 |issue=3 |pages=529–533 |title= Do males of the great grey shrike, ''Lanius excubitor'', trade food for extrapair copulations?|authorauthor1=Tryjanowski, Piotr and |author2=Hromada, Martin |name-list-style=amp |doi=10.1016/j.anbehav.2004.06.009|s2cid=54298849 }}</ref>

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

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 an 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=1395–1399|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-access=free}}</ref>

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

In decorated crickets, ''[[Gryllodes sigillatus]]'' (Orthoptera: Gryllidae), the nuptial food gift is a [[spermatophylax,]] (a large, gelatinous, sperm-free mass) that surrounds a smaller, sperm-containing ampulla. Together, the spermatophylax and the ampulla constitute the male’smale's [[spermatophore]], which is transferred to the female during copulation and remains attached outside her body at the base of her ovipositor. After mating, the female detaches the spermatophylax from the ampulla and feeds on it while the ampulla remains attached and is emptied of sperm.<ref>{{cite journal |journal= Animal Behaviour |year=2012 |volume=83 |issue=2 |pages=369–375 |title= Glycine in nuptial food gifts of decorated crickets decreases female sexual receptivity when ingested, but not when injected|authorauthor1=Gordon, Darcy G.; |author2=Gershman, Susan N. and |author3=Sakaluk, Scott K. |name-list-style=amp |doi=10.1016/j.anbehav.2011.11.005|s2cid=20094851 }}</ref> Once the female has consumed or discarded the spermatophylax, she removes the sperm ampulla, terminating sperm transfer. Female cooperation is required for successful spermatophore transfer, and thus, males cannot impose copulations on females.<ref>{{cite journal |journal= Phytochemistry

|year=2007 |volume=74 |issue=4 |pages=1065–1072 |title= Sequential mate choice in decorated crickets: females use a fixed internal threshold in pre- and postcopulatory choice|doi=10.1016/j.anbehav.2007.01.017 |last1= Ivy |first1= Tracie M. |last2= Sakaluk |first2= Scott K.|s2cid=28796676 }}</ref>

[[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" />

During mating in the [[Utetheisa ornatrix|ornate moth]] (''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=1009–1014|doi=10.1093/beheco/ars064|doi-access=free}}</ref>

[[File:Zygaena filipendula 240503.jpg|thumb|[[Six-spot burnet]]]]

Nuptial gifts are widespread in insects such as the [[six-spot burnet]] (''Zygaena filipendulae''), and comprise food items or glandular products offered as paternal investment in offspring and/or to promote mating. Female Zygaena may use this gift for her own defense and to protect her eggs.<ref>{{cite journal |journal= Animal Behaviour |year=2011 |volume=72 |issue=13 |pages=1585–1592 |title= Cyanogenic glucosides in the biological warfare between plants and insects: The Burnet moth-Birdsfoot trefoil model system|authorauthor1=Zagrobelny, Mika; and |author2=Birger, Lindberg Møller |name-list-style=amp | doi=10.1016/j.phytochem.2011.02.023|pmid=21429539 |bibcode=2011PChem..72.1585Z }}</ref>

[[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>{{Cite journal | last1 = Albo | first1 = M. A. J. | last2 = Costa | first2 = F. G. | doi = 10.1016/j.anbehav.2010.01.018 | title = Nuptial gift-giving behaviour and male mating effort in the Neotropical spider ''Paratrechalea ornata'' (Trechaleidae) | journal = Animal Behaviour | volume = 79 | issue = 5 | pages = 1031 | year = 2010 | pmids2cid = | pmc =39742664 | url = http://web.mnstate.edu/wisenden/Paper%208%20Spider%20nuptial%20gifts.pdf}}</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 name ="prokop" /> 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=302–306|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.

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