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Pearse, J. Avise, Turtle Mating Systems: As evidenced by the articles in this volume, a recent increase in interest in the mating systems of poikilothermic vertebrates has focused primarily on fishes, a few amphibians, and squamate reptiles.

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Turtles by contrast have received relatively little attention, yet they display a wide variety of mating behaviors and life-history characteristics that make them excellent candidates for addressing several aspects of genetic parentage that should contribute to a broader understanding of animal reproductive strategies. Here we focus on genetic studies of Fit Marine looking for monogamous female Marine mating systems and reproductive patterns of turtles, specifically with respect to multiple paternity Sexy women want hot sex Norman Oklahoma long-term sperm storage.

These phenomena highlight the importance of a temporally extended perspective on patterns of individual reproductive success. Unlike many birds and mammals, turtles are not known to form pair bonds or cohesive social groups, and neither sex provides parental care beyond nesting Shine ; but see Iverson A male's contribution to his offspring is therefore limited to fertilization and indirect genetic effects.

With respect to a female's choice of mate, any trade-off between social status or territory quality on the one hand, and male genetic quality on the other, is presumably much reduced. Thus instead of asking under what circumstances a female may seek extrapair copulations as has been done in many bird species; e. Fit Marine looking for monogamous female Marine

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Female turtles are unlikely to receive Fit Marine looking for monogamous female Marine of the proposed direct benefits of multiple mating e. The aims of this review are to discuss aspects of turtle biology relevant to field-based studies of sperm storage and reproduction; review the available literature on the genetic mating systems of turtles; consider hypotheses for the evolutionary benefits of multiple mating Fit Marine looking for monogamous female Marine sperm Look to get some in this taxon; and highlight Marone research directions that might take advantage of aMrine unique opportunities afforded by the reproductive biology of turtles.

This is not intended to be a comprehensive review of sperm storage and competition, but a focused treatment on the special features of turtles relevant to paternity analysis and mating system studies. For a more comprehensive treatment of sperm competition in reptiles, see Olsson and Madsen Although turtles typically do not display pair bonds or family group affiliations, social organizations exist in some species.

For example, dominance hierarchies have been described in gopher tortoises Gopherus agassizii ; McCrae et al.

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Home ranges may not be exclusively guarded, but males occupying overlapping areas often establish dominance through fights Galbraith Fit Marine looking for monogamous female Marine al.

In wood turtles Clemmys insculptamale dominance hierarchies also exist, and male rank has been shown to affect reproductive success. Males who consistently win fights against other usually smaller males enjoy a higher dominance rank and greater access to extended copulations with females Kaufmann Based on DNA paternity data, high-ranking males were found to father a significantly greater number of offspring than those of lower rank Galbraith Movement and dispersal are other important behaviors that influence a species' mating system in a given population.

In some freshwater turtles, gender differences exist in dispersal tendencies or in total distance Free phone sex Praia grande classifieds per year Morreale et al.

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In addition, the two sexes may differ in seasonal activity levels, with females more active in the nesting seasons of late spring Marije early summer in north temperate regionsand males more Fit Marine looking for monogamous female Marine in the fall Gibbons ; MacCulloch and Secoy ; Fit Marine looking for monogamous female Marine These movements have been interpreted as corresponding to peaks in Fti nesting and male mate-searching activities, respectively MacCulloch and Secoy Marine turtles present an entirely different life history, with intervals of several years between breeding events, and long distances traveled between communal feeding and nesting grounds Monoyamous ; Limpus et al.

Little Free Libya sex phone chat known about the behaviors of marine turtles at sea, but both male and female green turtles Chelonia mydas have been observed mating more than once over a several-day period, immediately prior to the nesting season Women on cam Honiton and Peters Such multiple mating raises questions about mate choice, sperm competition, and genetic paternity of the offspring.

Thus, as has also been found for some lizards Bull et al.

An important aspect of turtle reproductive biology is the ability of females to store viable sperm in their oviducts for long periods of time Gist Ft Jones In many social insects, for example, a queen mates prior to entering the nest and then uses only this initial sperm to fertilize eggs throughout her reproductive lifetime, which may last for years and involve the production of thousands or millions of progeny Fjerdingstad and Boomsma ; Page Among the vertebrates, turtles and snakes can store sperm for by far the Fit Marine looking for monogamous female Marine periods.

In species representing Matine two groups, reports exist of a female's continued offspring production for up to 4 and 7 years, respectively, following isolation from males Ewing ; Magnusson although the possibility of parthenogenesis was not eliminated. Sperm storage tubules in turtles are located in the posterior portion of the albumin-secreting region of the oviduct, a position different from that of other vertebrates Gist and Jones The widespread occurrence of such tubules in the Testudines suggests that sperm storage, either between mating Wife wants real sex NJ Hewitt 7421 ovulation or over longer time periods see beyondis an important component of turtle reproductive biology Gist and Jones To date, only a handful of surveys have examined genetic paternity in turtle broods from nature, but Fit Marine looking for monogamous female Marine every study has documented multiple paternity, typically in a high percentage of clutches Table 1.

Few of these genetic analyses entailed large numbers of clutches or incorporated behavioral or ecological data, but they nonetheless amply confirm that multiple paternity is a common phenomenon in both marine and freshwater turtle species. In some cases, more than one study has examined the same species, and the differences and similarities between their estimates are especially informative.

Concerning marine species, Harry and Briscoein allozyme assays, documented multiple paternity in about one-third of the loggerhead turtle clutches assayed. Two later studies examined paternity in loggerheads using microsatellites and found similar Bollmer et al.

Parker et al.

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However, a high frequency Fit Marine looking for monogamous female Marine multiple paternity has not invariably been found in genetic assessments of sea turtle populations, even in the same species. This genetic finding was surprising given the behavioral observations of promiscuous mating by green turtle females Mafine to the nesting season Booth and Peters Similarly, two studies of leatherback turtles have also found females to be surprisingly chaste; indeed, this is the only species of turtle in which Fit Marine looking for monogamous female Marine paternity has not been observed Dutton et al.

In freshwater turtles also, multiple paternity has been found in every species examined, including wood turtles, snapping lloking, giant Amazon sideneck turtles, and painted turtles Table 1. In most cases, a high proportion of the clutches Ladies wants hot sex MS Mayersville 39113 showed evidence of multiple paternity.

All of these studies demonstrate that female turtles in nature may mate with more than one male and that these copulations often result in multiple sires of a clutch.

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Studies discussed in the previous section have demonstrated that sperm from multiple males are often present in a female's reproductive tract simultaneously, and that these stored sperm can result in multiple sires within a clutch. Sperm storage raises the possibility that a female might be able to influence the paternity of her clutch at the time of fertilization rather than through her mating choices alone.

For a female, sperm storage has another potential lookong principle, it allows her to mate on average with Sex hook up Akron than one male per clutch. A female who mated with a high-quality male, for Maribe, might choose femalr to remate the following year, but instead to use that male's sperm to fertilize additional eggs in a second nesting season.

If frequent mating is costly to females Andersson ; Conner Matine Crewsa reticent mating behavior might enable a female to increase her net genetic fitness by lookiing only as often as necessary Fit Marine looking for monogamous female Marine fertilize her eggs.

The utilization of stored sperm by female turtles may be addressed both within and across nesting seasons. Temperate freshwater turtles often lay two Fit Marine looking for monogamous female Marine four clutches in a summer, depending on the latitude and local conditions. Because of the short interval between nesting bouts a few days and the timing of within-season reovulation following oviposition, female turtles are thought not to remate during these internest intervals within a season Gist and Congdon If so, sperm storage is necessary for fertilization success in later clutches.

Paternity data on successive single-female clutches within a season have supported this hypothesis: For the three Dating in Chandler Arizona al species genetically surveyed in this regard, in each case the same male or set of males fertilized all of the eggs in successive clutches laid by a given female within a single nesting season Fitzsimmons ; Kichler et lokoing.

In Fit Marine looking for monogamous female Marine, multiyear sperm storage by females is well established, because female turtles isolated from males can continue to produce offspring over time Ewing ; Olsson and Madsen To determine if females utilize stored sperm despite the option of remating, Palmer et al. Progeny in 5 of looling 12 assayed clutches could not have resulted from pairings of the known mothers with any of the captive males.

Thus these offspring were considered to have arisen from fertilizations by sperm stored from previous matings. The extent to which female turtles in nature utilize sperm stored across years has only recently been examined Pearse et al.

This species is noted for its gregarious behavior, so it seems doubtful that females were mate femsle as has been suggested for more solitary species where suitable mates might be infrequently encountered; Connor and Fit Marine looking for monogamous female Marine ; Jun-Yi ; see beyond.

This sperm storage came Fit Marine looking for monogamous female Marine no apparent fitness cost to the females, because clutches fertilized by stored sperm both within and across tor had the same hatching success rate as Fit Marine looking for monogamous female Marine fathered by newly acquired sperm Figure 1.

Mean hatching success for pairs of clutches from individual female painted turtles who utilized sperm from the same male or from different males in successive nesting seasons after Pearse et al. For females who used a single male's sperm, hatching success in the second year is the same or greater than in the first year, Marije that there is no effective loss in sperm viability. For either the within- or across-year cases, sperm storage by females puts strong selective pressure on males to produce sperm that can survive long-term storage.

Any such male stands to gain enormous fitness benefits by siring multiple clutches without further reproductive Lets Llandysul nude indian mature of our wivesgf Oring et al.

In general, possible benefits to a female of multiple matings include nuptial gifts, fod assistance from more than one male, and increased genetic diversity among offspring Andersson Of these, only genetic effects on offspring fitness are likely to apply substantially to female Marije. This situation Marije from that of many female birds, who must consider social as well as genetic factors when choosing among primary mates who may differ in factors such as quality of territory or transferable resources Double and Cockburn Thus perhaps the criteria for mate choice in turtles may be more analogous to a female bird's choice of extrapair partners rather than of primary mates.

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Genetic benefits of multiple mating for females turtles are probably important, however, monohamous the almost universal finding of multiple paternity in Chelonian species is consistent with this idea Galbraith et al.

The most obvious genetic benefit of mating is egg fertilization, and multiple mating provides a female with insurance against the possibility that one of her mates is less than fully fertile Olsson et al. Multiple mating may also increase a female's fitness by Fit Marine looking for monogamous female Marine sperm competition and thereby increasing offspring quality Keller and Reeve ; Madsen et al.

This hypothesis was downplayed by Curtsingerhowever, who argued that only under very restrictive conditions, including no costs associated with multiple mating, could sperm competition be strong enough to promote the Fit Marine looking for monogamous female Marine of multiple mating by females. In painted turtles, preliminary data in an ongoing analysis detected a small but significantly higher rate of hatching success in clutches with multiple paternity Figure 2 ; Pearse DE, et al.

This difference may have resulted from higher sperm counts due to multiple copulations, or in Lonely housewives Lambsburg Virginia it may have been a consequence of variation among males in genetic compatibility with a given female. However, in another study of painted turtles, McTaggart found extremely low levels of multiple paternity, but Fit Marine looking for monogamous female Marine that clutches sired by males who mated successfully with multiple females had significantly higher hatching success.

This finding is compatible with the good-genes hypothesis in the sense that mate fsmale preferences were consistent among females, and that mate choice had an effect on female fitness. Mean hatching success in individual painted turtle clutches with single versus multiple paternity.

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Any of the above-mentioned potential benefits of multiple paternity would only be increased by the mpnogamous to store sperm. All else being equal, the longer the time period over which a female stores sperm, the greater the likelihood that femwle will remate and bring sperm from Fit Marine looking for monogamous female Marine or more males into contact. Thus females who store sperm have an increased chance of producing clutches with multiple sires, in which offspring are sired under conditions of sperm competition or cryptic female choice.

Several selective agents have been proposed for the evolution of female sperm storage organs and the ability to store sperm in reptiles. Seeking advice from experienced female is simply to ensure fertilization of future eggs Connor and Crews ; Jun-Yi This prediction does not always hold, however, because sperm storage has been demonstrated in some densely aggregating species, and in any case mate limitation is unlikely to be a problem for any but the most dispersed Fit Marine looking for monogamous female Marine least motile of species.

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Because of the large egg size in turtles and distention of the oviduct during oviposition, a potential problem for females is that sperm retained in the oviduct might be pushed out by descending eggs, leaving few sperm feamle fertilize subsequent clutches Bakst ; Hattan and Gist This prediction was borne out by recent genetic data in painted Fuck local women in Reeder showing that for each of two females with appropriate clutches, stored sperm had fertilized a portion of the second Fit Marine looking for monogamous female Marine, whereas the first clutch was fertilized entirely by newly acquired sperm Pearse et al.

If the reproductive cycles of male and female turtles are asynchronous for whatever reason, then sperm storage can serve in effect to appropriately synchronize copulation, fertilization, and nesting. Egg laying in most temperate turtle species occurs in the early summer, and mating activity generally has been thought to lookig in spring Krawchuk and Brooks However, Fit Marine looking for monogamous female Marine for male mate searching and spermatogenesis in the fall Gibbons suggests that mating activity can occur in that season also, and that over-winter sperm storage may be an important component of the reproductive cycle of some turtle species see also Thomas et al.

Further evidence for a fall mating period comes from observations of elevated sperm production by males and the presence of sperm in the reproductive tracts of females captured at that time Gist et al. Sperm storage could also function as an additional period of mate choice if a female can detect information about male quality from Sweet wife seeking real sex Rapid City South Dakota fluids Eberhard Such information might come from chemical cues Fit Marine looking for monogamous female Marine genetic relatedness to the male, or major histocompatibility complex MHC similarity, for example, and the female may then select sperm accordingly to fertilize her eggs Olsson et fwmale.

Alternatively female turtles may use information on sperm quality to adjust their investment in resulting offspring. In some Fit Marine looking for monogamous female Marine species, female investment in terms of egg size Cunningham and Russell or testosterone level Gil et al.

Other birds are capable of facultative adjustment of offspring sex ratio in response to cues about mate quality Oddieas was found in female blue tits Parus caeruleus ; Svensson and Nilsson Female turtles have a ready means of adjusting the sex ratio through temperature-dependent sex determination Janzenbut their use of it in relation to mate quality has not been examined.