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Mate Sampling Strategy in a Field Cricket Animal Behaviour 81 (2011) 519e527 Contents lists available at ScienceDirect Animal Behaviour journal homepage: www.elsevier.com/locate/anbehav Articles Mate sampling strategy in a field cricket: evidence for a fixed threshold strategy with last chance option Oliver M. Beckers*, William E. Wagner, Jr School of Biological Sciences, University of Nebraska-Lincoln article info The strategy females use to sample potential mates can influence mate choice and thus sexual selection. fi Article history: We examined the mate sampling strategy of the cricket Gryllus lineaticeps. In our rst set of experiments, Received 13 July 2010 we simultaneously presented three different chirp rates to females. The set consisted of three trials, each Initial acceptance 12 August 2010 covering a different range of chirp rates. Independent of chirp rate range, female G. lineaticeps preferred Final acceptance 15 November 2010 rates that were above 3.0 chirps/s to rates that were below 3.0 chirps/s. Females did not discriminate Available online 28 December 2010 among chirp rates that were below this threshold and did not discriminate among chirp rates that were MS. number: A10-00483 above this threshold, suggesting that they express a fixed threshold sampling strategy. In our second experiment, females were presented sequentially with a fast chirp rate and then a slow chirp rate. When Keywords: the interval between presentations was 20 min, females showed significantly weaker responses to the acoustic communication slow rate than to the fast rate. However, when the interval between presentations was 24 h, female female preference responses to the slow and fast rate did not significantly differ. The latter result suggests that females fixed threshold strategy Gryllus lineaticeps lower their threshold of acceptance when they have not recently experienced highly attractive song last chance option types. This lower acceptance threshold is probably adaptive, as it would allow females to avoid paying mate choice high search costs, and to reproduce, only when low-quality males are available. Our results are consistent search strategy with a rarely considered sampling strategy (fixed threshold with last chance option strategy) and highlights the importance of the timing of social experience for mate sampling. Ó 2010 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved. How females choose a mate is a central issue for understanding Numerous sampling strategies have been proposed (e.g. Janetos the causes and consequences of sexual selection. Most research on 1980; Wittenberger 1983; Real 1990; Dombrovsky & Perrin 1994; female mate choice has focused on female mating preferences, and Luttbeg 1996; Wiegmann et al. 1996), most of which involve we know that females often prefer to mate with males with comparing the traits of males that are sampled or comparing male particular types of traits (e.g. Basolo 1990; Wilkinson & Reillo 1994; traits to an internal standard (Janetos 1980; Moore & Moore 1988). Gerhardt et al. 2000). Mate choice, however, is the result of inter- For all strategies, females are assumed to receive direct or indirect actions between environmental conditions (e.g. predation risk; benefits from mating with males with preferred traits (preferred Godin & Briggs 1995), mating preferences and sampling strategies males are thus assumed to be of higher quality). Four of the more (Wagner 1998). Much less is known about how females sample, commonly proposed strategies are (1) the best-of-N strategy, (2) the assess and compare potential mates (i.e. female sampling or search sequential comparison strategy, (3) the fixed threshold strategy and strategies) than about female mating preferences. Understanding (4) the variable threshold strategy. In the best-of-N strategy, a female female sampling strategies is important because different strategies samples a number (N) of males and then returns to the male with the can lead to different mate choices. For example, the number of highest quality (Janetos 1980). In the sequential comparison strategy, males sampled may differ depending upon the sampling strategy a female continues searching as long as each new male encountered a female uses, and females that sample more males are likely to is of higher quality than the previous male. Once a female encounters show a greater bias in their mate choices than females sampling a new male that is of lower quality than the previous male sampled, fewer males (Janetos 1980; Wagner 1998). The sampling strategies she chooses the previously encountered male (Wittenberger 1983). that females use can thus affect both how sexual selection acts on Finally, in the fixed threshold strategy (Janetos 1980) and the variable male traits and the benefits of mate choice itself. (or adjustable) threshold strategy (Janetos 1980; Reid & Stamps 1997), a female continues searching until she encounters a male whose quality exceeds an internal threshold value. The difference between these two strategies is how the threshold value is * Correspondence: O. M. Beckers, University of Nebraska, School of Biological fi Sciences, Lincoln, NE 68588, U.S.A. determined. In the xed threshold strategy, females compare males E-mail address: [email protected] (O.M. Beckers). against a fixed internal standard of acceptance (Janetos 1980). 0003-3472/$38.00 Ó 2010 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved. doi:10.1016/j.anbehav.2010.11.022 520 O.M. Beckers, W.E. Wagner Jr / Animal Behaviour 81 (2011) 519e527 This means that the female uses the same threshold acceptance In a second set of experiments, we tested whether female G. lin- criteria regardless of the mating opportunities that are available to eaticeps use a last chance option when males are not readily available. her (Reid & Stamps 1997). In the variable threshold strategy, the Previous results have shown that females show weaker responses to standard changes in response to the information gathered during a slow chirp rate if they have experienced a fast chirp rate within the sampling, such as in response to information about the costs of previous 20 min (Wagner et al. 2001). We tested whether this effect searching and the distribution of male quality. We consider the one- of acoustic experience weakens over time, which would be consistent step decision strategy (Janetos 1980), sequential search strategy (Real with a last chance option as proposed by Janetos (1980). 1990) and optimal stopping rule (Dombrovsky & Perrin 1994)as variations of the variable threshold strategy. GENERAL METHODS Mate sampling strategies have to balance the benefits obtained by finding a preferred mate with the costs associated with increased Animals sampling effort (Janetos 1980; Real 1990). These costs include the time and energy used for sampling, the risk of predation during We used females from a laboratory population of G. lineaticeps sampling and opportunity costs, such as reduced time available for maintained at the University of Nebraska-Lincoln. We collected adult foraging, the loss of previously sampled mates due to their death, females from Academy, California, U.S.A. (Wagner & Basolo 2007a)to emigration, or selection by other females (Daly 1978; Parker 1983; establish the laboratory population. Most field-collected females had Real 1990). In addition, the ability of a female to remember the mated before capture in the field and laid fertile eggs in the labora- characteristics and locations of previously sampled males may tory. Individuals hatching from those eggs constituted the first constrain the sampling strategy that a female uses (Janetos 1980). In laboratory generation. We actively managed the matings of subse- fact, all of the above-described strategies, with the exception of the quent laboratory generations by pairing males and females from fixed threshold strategy, require some degree of memory, a trait long unrelated families to reduce inbreeding and maintain genetic diver- assumed to be limited to vertebrates (but see Dukas 2008). While the sity. We tested females of the second and older laboratory genera- fixed threshold strategy does not require that females remember tions for our experiments and tested no more than three females previously sampled males, it has a major drawback: a female that from a given family in a treatment group. The number of full-sibling uses this strategy may risk not mating if the threshold is set too high, families represented in each treatment ranged from 16 to 28. or, alternatively, risk accepting a low-quality male if the threshold is Juvenile crickets were reared in family containers until the set too low (Jennions & Petrie 1997). However, it has been suggested penultimate instar, at which time we transferred them to individual that there could be a time-dependent decrease in the threshold value, containers. Once separated into individual containers, the nymphs which would improve the success of this strategy by granting a ‘last were acoustically isolated from singing adult males. We checked chance option’ for the female if no high-quality males are available the individual containers daily and recorded the date of the moult (Janetos 1980; Real 1990; Jennions & Petrie 1997). To our knowledge, to adulthood. Females and males used in our experiments were the ‘last chance option’ has not been demonstrated in any mating tested between 7 and 12 days after adult moult. system. Family and individual containers had a paper towel substrate We
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