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Selective Mate Choice by Females of Harpobittacus Australis (Mecoptera: Bittacidae)

Selective Mate Choice by Females of Harpobittacus Australis (Mecoptera: Bittacidae) BY FEMALES OF SELECTIVE MATE CHOICE HARPOBITTA CUS A USTRALIS (MECOPTERA: BITTACIDAE)* BY JOHN ALCOCK Department of Zoology State AZ 85281 Arizona University, Tempe, The use of in courtship is a rare nuptial gifts relatively phe- nomenon among insects but it is well developed in the Bittacidae where it has been studied in detail by Thornhill (1976, 1977, 1979). of In an investigation Hylobittacus apicalis Thornhill (1976) demon- strated that females exercise choice in the selection of mates, prefer- those that offer gifts; males that gave their mates large ring superior nutritious prey were permitted to copulate longer and transfer more sperm than males that offered small or unpalatable food presents. that the of This paper outlines limited evidence duration feeding and females of Australian copulation by mecopteran, Harpobittacus australis Klug, is also linked to the quality of the nuptial gifts pro- vided by the male. METHODS The was studied in National Park in species Warrumbungle northern New South Wales between 18-23 December 1978, a period of dry, warm weather when insects were active. Males and females appeared in the late morning and afternoon and perched on flowers, and on a hillside a small brook. I for grasses shrubs by searched males that were a and one carrying prey present, upon finding remained with it until the prey was discarded or until the insect moved and was lost to sight. As soon as a male was detected, I estimated the length of the prey in mm, recorded the manner in which it was carried and made notes on social interactions that any occurred. RESULTS observations of the sexual behavior of H. in My australis are general with those of Bornemissza Males carry agreement (1966). nuptial gifts impaled on the beak or gripped with the hind tarsi *Manuscript received by the editor November 2, 1979. 213 214 Psyche [June-September while from their front they hang a perch with legs. Prey carried by the had a mean estimated of 8.8 mm hindlegs length (N 9)and included three bulky moths and one large lepidopterous larva. Prey pierced by the beak averaged 3.8 mm in length (N 11) and con- sisted entirely of delicate mirids and various soft-winged beetles. with from Males prey release a sex pheromone glands in the an attractant that draws both males and females posterior abdomen, to them. In 19.5 hr of observation, I recorded 10 male-male encoun- ters and 31 male-female interactions. Male visitors uniformly to steal from the attempted the prey calling male and succeeded on two occasions. Prey males is common bittacids piracy by among Some females may also visit calling males solely to (Thornhill 1979). secure a food gift. One male-female interaction ended when the female removed the prey from the male and flew away. In another female for 3 case, the probed the male’s prey about min after coming to his the male without success to effect a perch; attempted copula- tion and eventually a tussle for control of the ensued prey leading the pair to tumble off the perch and onto the ground. The female left; the male returned to his perch and probed the prey briefly before discarding it, apparently because the female had drained it of its contents. In more interactions between males and females, the arriv- typical ing female alighted causing the male to transfer the prey to his of mouthparts. In my observations, unlike those Bornemissza (1966) the female with her beak usually probed nuptial gift briefly (N on only three occasions a female refrained from "testing" the 26); prey and all three involved unusually large nuptial gifts whose potential value might have ben assessed visually. As the female the the male to with her. probed gift, attempted couple In fifteen the female refused to to cases, permit mating occur, withdrawing from the male after the initial probe of his present. In cases in which copulation occurred the male usually succeeded in pulling the prey from the female with his hindlegs shortly after insertion of his genitalia. The female, which had been facing the male in this initial phase of the interaction, would then drop down so that the bodies of the copulating pair formed an "L". aver- (The age time from the start of an encounter to assumption of the "L" position was 3.1 min.). While the female hung below the male she employed her hindlegs to pull vigorously at the male’s hindlegs which were the holding prey 1979] Alcock 215 Harpobittacus australis away from the female’s mouthparts. Males appeared reluctant to extend their legs and so permit the female to feed. The time required for the female to draw the to her nuptial gift averaged 0.95 min with a maximum of 4.6 min When the male’s were (N 11). legs fully extended the female could reach the prey with her she mouthparts; fed upon it for as little as min to as much as 17.25 min. Imme- diately upon cessation of feeding the female disengaged from her and flew off. partner Selective Mate Choice by Females It seems likely that the probing of the prey by the female at the start of an encounter between the sexes permits the female to assess the value of the male’s nutritional/caloric present. Females offered small prey or insects that had been consumed an earlier by partner of the male frequently rejected the male outright, before leaving copulation could begin. If we consider the first observed reaction of a female to a prey offered by a male, copulation occurred only once in ten cases in which the a 3-4 prey was tiny mm mirid or beetle. In contrast, if the were estimated to be 6 mm or prey larger, mating occurred in 8 of 10 cases 7.3, P (X < .05). Five of twenty prey were offered to more than one females (2-4) in sequence. All five prey were 6 mm or larger. In one case, the prey was discarded the male after the of by immediately second two lengthy copulations. In the remaining four cases, the male dropped his gift only after it had been probed and rejected by a female, presumably because she found it an empty husk whose contents had been removed earlier females. by Thus the quality of the must be of a certain minimum nuptial gift standard if the female is to permit the male to with her at copulate all. In addition, Bornemissza felt that the size of the prey was (1966) related to the duration of copulation in H. australis. Table shows that there is indeed a correlation between the estimated size of a prey and the duration of the of (1) feeding phase copulation and (2) the overall length of the copulation. Large food items evidently contain within them sufficient materials to keep the female occupied for some time, during which period the male presumably transfers to her. sperm Still more for the that the of support hypothesis food value the gift determines how long a female will feed on the and therefore prey how long she will copulate comes from comparisons of the response 216 Psyche [June-September of several females offered the same In the three prey. cases in which a male presented the same food item to three different females, copulation lengths were (a) 10.2, 7.8, and 0 min, 15.7, 17.2, and 0 (b) and min, (c) 12.3, 3.8 and 0 min and the feeding phase lasted (a) 10.2, 4.8, and 0 min, (b) 15.3, 14.8, and 0 min, and (c) 12.3, 3.8 and 0 min. Table I. The relation between the estimated length of the prey offered a female as a nuptial gift and the time that she fed upon the prey and the duration of copulation. Size of Mean time of Mean time of gift N feeding Range copulation Range 3-4 mm 10 0.2 min 0- 1.8 1.1 min 0- 8.0 6-8 mm 6 3.6 0-17.3 6.2 0-19.3 10-14 mm 4 12.3 10.2-15.3 13.7 10.2-16.8 Correlation between size of and time, .01 prey feeding 0.70, p < Correlation between size of and copulation time, 0.62, .01 prey p < Data collected from the first observed interaction between a male carrying a nuptial present and female. DISCUSSION The species of bittacids studied to date exhibit reasonably similar sexual behavior the of including use pheromones, nuptial gift giv- ing, female sampling of the before present copulation, and copula- tions of variable length but with means of about 10-20 min (Thornhill, 1977). An unusual feature of the mating behavior of Harpobittacus is the removal of the prey from the female by the male for a short onset period after the of copulation. In some other bittacids females feed on the continuously nuptial gift. The apparent struggle between copulating partners of H. australis for possession of the prey demonstrates the male and female interests are not identical. Perhaps the risk of prey thievery by females favors males that are cautious about their relinquishing valuable presents; per- haps they refuse to let mates feed until potential sperm have begun to be accepted by their partners. Sperm transfer in Hylobittacus apicalis is proportional to the duration of to about 20 copulation (up min) and it would not be surprising if this were also true for H. australis, whose females feed 217 1979] Alcock Harpobittacus austral& in copulo for from 1-17 min. Certainly females discriminate among males on the basis of volume of the to edible nuptial gift, refusing at all with males very small or that couple offering presents prey have been drained by previous mates. This mecopteran, therefore, probably provides another example of a species whose females exer- cise mate choice, accepting sperm preferentially from individuals a investment the form of a food that that make large parental in gift or female survival. may promote egg development ACKNOWLEDGEMENTS the This study was conducted while author was a visiting lecturer at Monash I thank the members of the Depart- University. Zoology ment at Monash for their help, my sons Joey and Nicky for their assistance with field work, and Randy Thornhill for reading the manuscript. REFERENCES F. BORNEMISSZA, G. on the and behaviour of two species of 1966. Observations hunting mating flies Mecoptera). Aust. J. Zool. 14: 371-382. scorpion (Bittacidae: THORNHILL, R. 1976. Sexual selection and nuptial feeding behavior in Bittacus apicalis (Insecta: Mecoptera). Amer. Nat. 110: 529-548. 1977. The and sexual behavior of flies comparative predatory hanging (Mecop- tera: Occ. Papers Mus. Zool., Univ. Mich. 677: 1-43. Bittacidae). 1979. Adaptive female-mimicking behavior in a scorpionfly. Science 205: 412-414. International Journal of Peptides Advances in International Journal of BioMed Stem Cells Virolog y Research International International Genomics Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 Journal of Nucleic Acids International Journal of Zoology Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 Submit your manuscripts at http://www.hindawi.com The Scientific Journal of Signal Transduction World Journal Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 International Journal of Advances in Genetics Anatomy Biochemistry Research International Research International Microbiology Research International Bioinformatics Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 Enzyme Journal of International Journal of Molecular Biology Archaea Research Evolutionary Biology International Marine Biology Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Psyche: A Journal of Entomology Hindawi Publishing Corporation

Selective Mate Choice by Females of Harpobittacus Australis (Mecoptera: Bittacidae)

Psyche: A Journal of Entomology , Volume 86 – Jan 1, 1979

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Hindawi Publishing Corporation
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This article is in the public domain. This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.
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Abstract

BY FEMALES OF SELECTIVE MATE CHOICE HARPOBITTA CUS A USTRALIS (MECOPTERA: BITTACIDAE)* BY JOHN ALCOCK Department of Zoology State AZ 85281 Arizona University, Tempe, The use of in courtship is a rare nuptial gifts relatively phe- nomenon among insects but it is well developed in the Bittacidae where it has been studied in detail by Thornhill (1976, 1977, 1979). of In an investigation Hylobittacus apicalis Thornhill (1976) demon- strated that females exercise choice in the selection of mates, prefer- those that offer gifts; males that gave their mates large ring superior nutritious prey were permitted to copulate longer and transfer more sperm than males that offered small or unpalatable food presents. that the of This paper outlines limited evidence duration feeding and females of Australian copulation by mecopteran, Harpobittacus australis Klug, is also linked to the quality of the nuptial gifts pro- vided by the male. METHODS The was studied in National Park in species Warrumbungle northern New South Wales between 18-23 December 1978, a period of dry, warm weather when insects were active. Males and females appeared in the late morning and afternoon and perched on flowers, and on a hillside a small brook. I for grasses shrubs by searched males that were a and one carrying prey present, upon finding remained with it until the prey was discarded or until the insect moved and was lost to sight. As soon as a male was detected, I estimated the length of the prey in mm, recorded the manner in which it was carried and made notes on social interactions that any occurred. RESULTS observations of the sexual behavior of H. in My australis are general with those of Bornemissza Males carry agreement (1966). nuptial gifts impaled on the beak or gripped with the hind tarsi *Manuscript received by the editor November 2, 1979. 213 214 Psyche [June-September while from their front they hang a perch with legs. Prey carried by the had a mean estimated of 8.8 mm hindlegs length (N 9)and included three bulky moths and one large lepidopterous larva. Prey pierced by the beak averaged 3.8 mm in length (N 11) and con- sisted entirely of delicate mirids and various soft-winged beetles. with from Males prey release a sex pheromone glands in the an attractant that draws both males and females posterior abdomen, to them. In 19.5 hr of observation, I recorded 10 male-male encoun- ters and 31 male-female interactions. Male visitors uniformly to steal from the attempted the prey calling male and succeeded on two occasions. Prey males is common bittacids piracy by among Some females may also visit calling males solely to (Thornhill 1979). secure a food gift. One male-female interaction ended when the female removed the prey from the male and flew away. In another female for 3 case, the probed the male’s prey about min after coming to his the male without success to effect a perch; attempted copula- tion and eventually a tussle for control of the ensued prey leading the pair to tumble off the perch and onto the ground. The female left; the male returned to his perch and probed the prey briefly before discarding it, apparently because the female had drained it of its contents. In more interactions between males and females, the arriv- typical ing female alighted causing the male to transfer the prey to his of mouthparts. In my observations, unlike those Bornemissza (1966) the female with her beak usually probed nuptial gift briefly (N on only three occasions a female refrained from "testing" the 26); prey and all three involved unusually large nuptial gifts whose potential value might have ben assessed visually. As the female the the male to with her. probed gift, attempted couple In fifteen the female refused to to cases, permit mating occur, withdrawing from the male after the initial probe of his present. In cases in which copulation occurred the male usually succeeded in pulling the prey from the female with his hindlegs shortly after insertion of his genitalia. The female, which had been facing the male in this initial phase of the interaction, would then drop down so that the bodies of the copulating pair formed an "L". aver- (The age time from the start of an encounter to assumption of the "L" position was 3.1 min.). While the female hung below the male she employed her hindlegs to pull vigorously at the male’s hindlegs which were the holding prey 1979] Alcock 215 Harpobittacus australis away from the female’s mouthparts. Males appeared reluctant to extend their legs and so permit the female to feed. The time required for the female to draw the to her nuptial gift averaged 0.95 min with a maximum of 4.6 min When the male’s were (N 11). legs fully extended the female could reach the prey with her she mouthparts; fed upon it for as little as min to as much as 17.25 min. Imme- diately upon cessation of feeding the female disengaged from her and flew off. partner Selective Mate Choice by Females It seems likely that the probing of the prey by the female at the start of an encounter between the sexes permits the female to assess the value of the male’s nutritional/caloric present. Females offered small prey or insects that had been consumed an earlier by partner of the male frequently rejected the male outright, before leaving copulation could begin. If we consider the first observed reaction of a female to a prey offered by a male, copulation occurred only once in ten cases in which the a 3-4 prey was tiny mm mirid or beetle. In contrast, if the were estimated to be 6 mm or prey larger, mating occurred in 8 of 10 cases 7.3, P (X < .05). Five of twenty prey were offered to more than one females (2-4) in sequence. All five prey were 6 mm or larger. In one case, the prey was discarded the male after the of by immediately second two lengthy copulations. In the remaining four cases, the male dropped his gift only after it had been probed and rejected by a female, presumably because she found it an empty husk whose contents had been removed earlier females. by Thus the quality of the must be of a certain minimum nuptial gift standard if the female is to permit the male to with her at copulate all. In addition, Bornemissza felt that the size of the prey was (1966) related to the duration of copulation in H. australis. Table shows that there is indeed a correlation between the estimated size of a prey and the duration of the of (1) feeding phase copulation and (2) the overall length of the copulation. Large food items evidently contain within them sufficient materials to keep the female occupied for some time, during which period the male presumably transfers to her. sperm Still more for the that the of support hypothesis food value the gift determines how long a female will feed on the and therefore prey how long she will copulate comes from comparisons of the response 216 Psyche [June-September of several females offered the same In the three prey. cases in which a male presented the same food item to three different females, copulation lengths were (a) 10.2, 7.8, and 0 min, 15.7, 17.2, and 0 (b) and min, (c) 12.3, 3.8 and 0 min and the feeding phase lasted (a) 10.2, 4.8, and 0 min, (b) 15.3, 14.8, and 0 min, and (c) 12.3, 3.8 and 0 min. Table I. The relation between the estimated length of the prey offered a female as a nuptial gift and the time that she fed upon the prey and the duration of copulation. Size of Mean time of Mean time of gift N feeding Range copulation Range 3-4 mm 10 0.2 min 0- 1.8 1.1 min 0- 8.0 6-8 mm 6 3.6 0-17.3 6.2 0-19.3 10-14 mm 4 12.3 10.2-15.3 13.7 10.2-16.8 Correlation between size of and time, .01 prey feeding 0.70, p < Correlation between size of and copulation time, 0.62, .01 prey p < Data collected from the first observed interaction between a male carrying a nuptial present and female. DISCUSSION The species of bittacids studied to date exhibit reasonably similar sexual behavior the of including use pheromones, nuptial gift giv- ing, female sampling of the before present copulation, and copula- tions of variable length but with means of about 10-20 min (Thornhill, 1977). An unusual feature of the mating behavior of Harpobittacus is the removal of the prey from the female by the male for a short onset period after the of copulation. In some other bittacids females feed on the continuously nuptial gift. The apparent struggle between copulating partners of H. australis for possession of the prey demonstrates the male and female interests are not identical. Perhaps the risk of prey thievery by females favors males that are cautious about their relinquishing valuable presents; per- haps they refuse to let mates feed until potential sperm have begun to be accepted by their partners. Sperm transfer in Hylobittacus apicalis is proportional to the duration of to about 20 copulation (up min) and it would not be surprising if this were also true for H. australis, whose females feed 217 1979] Alcock Harpobittacus austral& in copulo for from 1-17 min. Certainly females discriminate among males on the basis of volume of the to edible nuptial gift, refusing at all with males very small or that couple offering presents prey have been drained by previous mates. This mecopteran, therefore, probably provides another example of a species whose females exer- cise mate choice, accepting sperm preferentially from individuals a investment the form of a food that that make large parental in gift or female survival. may promote egg development ACKNOWLEDGEMENTS the This study was conducted while author was a visiting lecturer at Monash I thank the members of the Depart- University. Zoology ment at Monash for their help, my sons Joey and Nicky for their assistance with field work, and Randy Thornhill for reading the manuscript. REFERENCES F. BORNEMISSZA, G. on the and behaviour of two species of 1966. Observations hunting mating flies Mecoptera). Aust. J. Zool. 14: 371-382. scorpion (Bittacidae: THORNHILL, R. 1976. Sexual selection and nuptial feeding behavior in Bittacus apicalis (Insecta: Mecoptera). Amer. Nat. 110: 529-548. 1977. The and sexual behavior of flies comparative predatory hanging (Mecop- tera: Occ. Papers Mus. Zool., Univ. Mich. 677: 1-43. Bittacidae). 1979. Adaptive female-mimicking behavior in a scorpionfly. Science 205: 412-414. International Journal of Peptides Advances in International Journal of BioMed Stem Cells Virolog y Research International International Genomics Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 Journal of Nucleic Acids International Journal of Zoology Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 Submit your manuscripts at http://www.hindawi.com The Scientific Journal of Signal Transduction World Journal Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 International Journal of Advances in Genetics Anatomy Biochemistry Research International Research International Microbiology Research International Bioinformatics Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 Enzyme Journal of International Journal of Molecular Biology Archaea Research Evolutionary Biology International Marine Biology Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation Hindawi Publishing Corporation http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014 http://www.hindawi.com Volume 2014

Journal

Psyche: A Journal of EntomologyHindawi Publishing Corporation

Published: Jan 1, 1979

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