Terminal Investment Hypothesis Research Articles

Age, but not an immune challenge, triggers terminal investment in the Pacific field cricket, Teleogryllus oceanicus.

The terminal investment hypothesis proposes that, when individuals are faced with a threat to survival, they will increase investment in current reproduction. The level of the threat necessary to elicit terminal investment (the dynamic terminal investment threshold) may vary based on other factors that also influence future reproduction. Here, we tested whether there is an interactive effect of age and an immune challenge on the dynamic terminal investment threshold in the Pacific field cricket, Teleogryllus oceanicus. We measured the courtship call, mating attractiveness, ejaculate size, and offspring production of T. oceanicus males. We found only limited support for the dynamic terminal investment threshold: there was no consistent evidence of a positive interaction between male age and immune challenge intensity. However, we found evidence for age-related terminal investment: older males produced a larger spermatophore than younger males. Older males also had a slower calling rate compared to younger males, suggesting a potential trade-off between these two pre- and post-copulatory traits. As some, but not all, reproductive traits responded plastically to cues for terminal investment, our research highlights the importance of considering a broad range of pre-and post-copulatory traits when exploring the potential for terminal investment to occur.

Impact of food availability and predator cues on meadow vole response to social vs. non-social odorants

Abstract The risk of predation and food deprivation may alter the degree to which animals associate with conspecifics. We examined if food deprivation, the risk of predation, or simultaneous exposure to both altered meadow voles’ preference for odour cues in a way that adheres to the terminal investment, safety in numbers, or avoidance hypotheses. Satiated or food-deprived meadow voles were given the choice to investigate either opposite-sex conspecific bedding, same-sex conspecific bedding, clean bedding, or self-bedding when exposed to mink urine or olive oil. Mink urine and food deprivation did not impact the amount of time meadow voles spent with each type of bedding, but meadow voles did begin investigating more quickly when experiencing either or both stressors. However, food deprivation and mink urine did not have an additive impact on any measured variable. Further research is needed to determine if the terminal investment hypothesis is the hypothesis that best describes the mating behaviour of meadow voles facing one or multiple stressors.

Reproductive effort and terminal investment in a multispecies assemblage of Amazon electric fish

AbstractThe terminal investment hypothesis (TIH) predicts that individuals with favorable prospects for future reproduction (i.e., high residual reproductive value, RRV) should moderate current reproductive investment in favor of growth, survival, and future reproduction, whereas those with low RRV should “terminally invest” by diverting somatic resources towards current reproduction at the expense of future reproduction. However, support for the TIH in wild animal populations is fragmentary, and the ecological contexts of terminal investment remain poorly known. We report a remarkable case of simultaneous terminal investment involving five sympatric species of the electric knifefish genus Brachyhypopomus, from Amazonian floodplain and terra firme stream habitats. We found that terminal investment is synchronized by seasonal breeding, in response to circannual environmental variation in mortality risk. Four species exhibit a uniseasonal iteroparous (annual) life history with complete post‐reproductive mortality after a single breeding season. One species (Brachyhypopomus beebei) exhibits a 2‐year multiseasonal iteroparous life history with breeding in two seasons and post‐reproductive mortality after the second. In mature females and (most) males of the annual species, as well as in both mature female and male second‐year (but not first‐year) B. beebei, we documented an increase in two metrics of reproductive effort (size‐adjusted gonad mass and electric signal amplitude) and a concomitant reduction in somatic condition (size‐adjusted somatic mass), all in response to proximity to the end of the common breeding season, when RRV approximates zero. In mature first‐year B. beebei, we documented neither an increase in reproductive effort nor a decline in somatic condition, implying an alternative strategy of reproductive restraint. Our findings support Kirkwood's disposable soma theory, which posits that death by reproductive exhaustion can be delayed if terminal investment is replaced by reproductive restraint, allowing individuals to survive and breed in a subsequent season. Deferral of the terminal investment response in annual species, and the origin of a gonadal regression‐regeneration sequence, may open pathways for rapid evolutionary transitions to multiseasonal iteroparity. Excepting the age (year‐group) dependency of terminal investment in B. beebei, we were unable to identify intrinsic cues or extrinsic environmental cues for the terminal investment response in Brachyhypopomus.

Age-specific fecundity under pathogenic threat in an insect: Terminal investment versus reproductive restraint.

The terminal investment hypothesis predicts that as an organism's prospects for survival decrease, through age or when exposed to a pathogenic infection, it will invest more in reproduction, which should trade-off against somatic maintenance (including immunity) and therefore future survival. Attempts to test this hypothesis have produced mixed results, which, in addition, mainly rely on the assessment of changes in reproductive effort and often overlooking its impact on somatic defences and survival. Alternatively, animals may restrain current reproduction to sustain somatic protection, increasing the chance of surviving for additional reproductive opportunities. We tested both of these hypotheses in females of the yellow mealworm beetle, Tenebrio molitor, an iteroparous insect with reproductive tactics similar to that of long-lived organisms. To achieve this, we mimicked pathogenic bacterial infections early or late in the life of breeding females by injecting them with a suspension of inactivated Bacillus cereus, a known natural pathogen of T. molitor, and measured female age-specific fecundity, survival, body mass and immunity. Inconsistent with a terminal investment, females given either an early or late-life immune challenge did not exhibit reduced survival or enhance their reproductive output. Female fecundity declined with age and was reduced by the early but not the late immune challenge. Both early and late-life fecundity correlated positively with life expectancy. Finally, young and old females exhibited similar antibacterial immune responses, suggesting that they both restrained reproduction to sustain immunity. Our results clearly demonstrate that age-specific reproduction of T. molitor females under pathogenic threat is inconsistent with a terminal investment. In contrast, our results instead suggest that females used a reproductive restraint strategy to sustain immunity and therefore subsequent reproductive opportunities. However, as infections were mimicked only, the fitness benefit of this reproductive restraint could not be shown.

Late-life reproduction in an insect: Terminal investment, reproductive restraint or senescence.

The terminal investment, reproductive restraint or senescence theories may explain individual late-life patterns of reproduction. The terminal investment hypothesis predicts that individuals increase reproductive allocation late in life as prospects for future survival decrease. The other two hypotheses predict reduced reproduction late in life, but for different reasons. Under the Reproductive Restraint hypothesis, individuals restrain their reproductive effort to sustain future survival and gain more time for reproducing, whereas under the Senescence process, reproduction is constrained because of somatic deterioration. While these hypotheses imply that reproduction is costly, they should have contrasted implications in terms of survival after late reproduction and somatic maintenance. Testing these hypotheses requires proper consideration of the effects of age-dependent reproductive effort on post-reproduction survival and age-related somatic functions. We experimentally tested these three hypotheses in females of the mealworm beetle, Tenebrio molitor, an iteroparous and income breeder insect. We manipulated their age-specific allocation into reproduction and observed the effects of this manipulation on their late-life fecundity, post-reproduction survival and immunocompetence as a measurement of somatic protection. We found that females exhibit age-related decline in fecundity and that this reproductive senescence is accelerated by a cost of early reproduction. The cost of reproduction had no significant effect on female longevity and their ability to survive a bacterial infection, despite that some immune cells were depleted by reproduction. We found that female post-infection survival deteriorated with age, which could be partly explained by a decline in some immune parameters. Importantly, females did not increase their reproductive effort late in life at the expense of their late-life post-reproduction survival. Late-life reproduction in T. molitor females is senescing and not consistent with a terminal investment strategy. Rather, our results suggest that females allocate resources according to a priority scheme favouring longevity at the expense of reproduction, which is in line with the reproductive restraint hypothesis. Such a priority scheme also shows that a relatively short-lived insect can evolve life-history strategies hitherto known only in long-lived animals. This puts in perspective the role of longevity in the evolution of life-history strategies.

Macronutrient intake and simulated infection threat independently affect life history traits of male decorated crickets.

Nutritional geometry has advanced our understanding of how macronutrients (e.g., proteins and carbohydrates) influence the expression of life history traits and their corresponding trade‐offs. For example, recent work has revealed that reproduction and immune function in male decorated crickets are optimized at very different protein:carbohydrate (P:C) dietary ratios. However, it is unclear how an individual's macronutrient intake interacts with its perceived infection status to determine investment in reproduction or other key life history traits. Here, we employed a fully factorial design in which calling effort and immune function were quantified for male crickets fed either diets previously demonstrated to maximize calling effort (P:C = 1:8) or immune function (P:C = 5:1), and then administered a treatment from a spectrum of increasing infection cue intensity using heat‐killed bacteria. Both diet and a simulated infection threat independently influenced the survival, immunity, and reproductive effort of males. If they called, males increased calling effort at the low infection cue dose, consistent with the terminal investment hypothesis, but interpretation of responses at the higher threat levels was hampered by the differential mortality of males across infection cue and diet treatments. A high protein, low carbohydrate diet severely reduced the health, survival, and overall fitness of male crickets. There was, however, no evidence of an interaction between diet and infection cue dose on calling effort, suggesting that the threshold for terminal investment was not contingent on diet as investigated here.

An immune challenge of female great tits decreases offspring survival and has sex-specific effects on offspring body size

Investment in immunity is expected to decrease (costly immunity) or enhance (terminal investment) reproductive performance. Here, we tested the effects of activation of the immune system in female great tits (Parus major) on (1) their reproductive effort and (2) the survival and body condition of their offspring, controlling for chick sex. We injected females tending 3-day-old chicks with sheep red blood cells (SRBC) or saline (control) and recorded their provisioning rates 6 days later, during the expected peak of antibody production. We measured tarsus length and body mass in 11-day-old chicks and monitored changes in brood size. We found that female provisioning rates were unaffected by the SRBC challenge. An analysis without an outlier, however, showed a significant challenge-by-hatch date interaction. This interaction indicated that female provisioning rates decreased with hatch dates in the SRBC but not in the control nests, suggesting a stronger effect in later breeders. Chick body mass was not affected by female immunisation nor by its interaction with chick sex. However, we found a significant challenge-by-sex interaction on offspring tarsus. In SRBC nests, the difference in tarsus length between male and female chicks was lower than in controls, suggesting sex-dependent effects of the challenge on offspring structural growth. Finally, chick mortality was greater in SRBC nests compared with controls, but chick survival probability was not affected by sex. Overall, our results support the costly immunity but not the terminal investment hypothesis in the great tit.

Modification of reproductive schedule in response to pathogen exposure in a wild insect: Support for the terminal investment hypothesis.

Trade-offs in the time and energy allocated to different functions, such as reproductive activities, can be driven by alterations in condition which reduce resources, often in response to extrinsic factors such as pathogens or parasites. When individuals are challenged by a pathogen, they may either reduce reproduction as a cost of increasing defence mechanisms or, alternatively, modify reproductive activities so as to increase fecundity thereby minimizing the fitness costs of earlier death, a behaviour consistent with the terminal investment hypothesis (TIH). The TIH predicts that individuals with decreased likelihood of future reproduction will maximize current reproductive effort, which may include shifts in reproductive timing. We examined how wild, adult female click beetles (Agriotes obscurus) responded after exposure to the fungal pathogen Metarhizium brunneum. Field-collected beetles exposed to a high concentration of M.brunneum died earlier and in greater numbers than those exposed to a low concentration. Using a multivariate approach, we examined the impact of pathogen challenge on lifespan and a suite of reproductive traits. Stepdown regression analysis showed that only female lifespan differed among the fungal treatments. Fungal-induced reductions in lifespan drove changes in the reproductive schedule, characterized by a decrease in preoviposition period. Moving the start of egg laying forward allowed the females to offset the costs of a shortened lifespan. These changes suggest that there is a threshold for terminal investment, which is dependent on strength of the survival threat. From an applied perspective, our findings imply that exposing adult click beetles to M.brunneum to reduce their population density might not succeed and is an approach that needs further investigation.

All or nothing: Switch to high current reproductive investment under risk of starvation in male kelp crab.

One of the key features in reproduction of polygynous species is seminal recovery after mating. However, it is poorly known how environmental factors affect the recuperation period of seminal material. This study aims to test plasticity in recovery of seminal reserves in response to distinct environmental conditions of the kelp crab Taliepus dentatus. Male crabs were maintained after depletion of seminal reserves in one of eight different treatments in a factorial design of temperature (12 and 16°C), food availability (with alimentation and food deprivation), and time period (15 and 30 days), simulating different environmental situations in the laboratory to which the crab might be exposed to along its distribution. Temperature and food availability modulated the seminal recovery period in T. dentatus. Complete replenishment was reached within 30 days in all treatments (i.e., 12 and 16°C each with alimentation and food deprivation), but the highest recovery index was found in crabs without food provision (16°C). In this condition, the recovery index was twice as high compared with males maintained at a similar temperature but with feeding. Prolonged starvation at 16°C may be extremely stressful conditions for male crabs, during which risk to die probably triggered a concentration of the reproductive effort in favor of immediate reproduction. This suggests that plasticity of energy allocation toward reproduction may be expressed during extremely suboptimal conditions, which might be a similar strategy as proposed by the terminal investment hypothesis. The generally relatively fast seminal recovery regardless of the temperature may explain the kelp crab's continuous mating throughout the year.

Terminal investment strategies following infection are dependent on diet.

When future reproductive potential is threatened, for example following infection, the terminal investment hypothesis predicts thatindividuals will respondby investing preferentially in current reproduction. Terminal investment involves reallocating resources to current reproductive effort, so it is likely to be influenced by the quantity and quality of resources acquired through diet. Dietary protein specifically has been shown to impact both immunity and reproduction in a range of organisms, but its impact on terminal investment is unclear. We challenged females from ten naturally derived fruit fly (Drosophila melanogaster) genotypes with the bacterial pathogen Pseudomonas aeruginosa. We then placed these on either a standard or isocaloric high-protein diet, and measured multiple components of reproductive investment. As oogenesis requires protein, and flies increase egg production with protein intake, we hypothesized that terminal investment would be easier to observe if protein was not already limiting. Oral exposure to the pathogen triggered an increase in reproductive investment. However, whereas flies feeding on a high-protein diet increased the number of eggs laid when exposed to P. aeruginosa, those fed the standard diet did not increase the number of eggs laid but increased egg-to-adult viability following infection. This suggests that the specific routes through which flies terminally invest are influenced by the protein content of the maternal diet. We discuss the importance of considering diet and natural routes of infection when measuring nonimmunological defences.

Flirting with danger: predation risk interacts with male condition to influence sexual display

AbstractSexual signaling coevolves with the sensory systems of intended receivers; however, predators may be unintended receivers of sexual signals. Conspicuous aerial displays in some species may place males at high risk of predation from eavesdropping predators. There are three different hypotheses to explain how signaling males can deal with increased predation risk: (1) males invest in survival by decreasing signal conspicuousness; (2) males invest in reproduction by increasing signal conspicuousness; and (3) male response is condition-dependent according to his residual reproductive value. Here, we used blue-black grassquits (Volatinia jacarina) to test these hypotheses, asking whether males modify leap displays under different levels of predation risk. Grassquit males develop an iridescent nuptial plumage and spend considerable time emitting a multimodal signal: while leaping from a perch, males clap their wings above their heads and emit a high-pitched short song. We exposed males to predator and nonpredator playbacks while video recording their displays. We found interactions between predation risk and 2 male condition variables (ectoparasite infestation and proportion of nuptial plumage coverage) that influenced display behavior. Less parasitized males and those with higher proportion of nuptial plumage showed no change in display behavior, while more parasitized males and those with lower proportion of nuptial plumage increased the vigor of displays under predation risk. In other words, males with low residual reproductive value increased reproductive effort when there was a high risk of extrinsic death. Our study provides some empirical support for the terminal investment hypothesis.

Inbreeding alters context-dependent reproductive effort and immunity in male crickets.

Infection can cause hosts to drastically alter their investment in key life-history traits of reproduction and defence. Infected individuals are expected to increase investment in defence (e.g., by increasing immune function) and, due to trade-offs, investment in other traits (e.g., current reproduction) should decrease. However, the terminal investment hypothesis postulates that decreased lifespan due to infection and the associated reduction in the expectation for future offspring will favour increased investment towards current reproduction. Variation in intrinsic condition will likely influence shifts in reproductive investment post-infection, but this is often not considered in such assessments. For example, the extent of inbreeding can significantly impact an individual's lifetime fitness and may influence its reproductive behaviour following a threat of infection. Here, we investigated the effects of inbreeding status on an individual's reproductive investment upon infection, including the propensity to terminally invest. Male crickets (Gryllodes sigillatus) from four genetically distinct inbred lines and one outbred line were subjected to a treatment from an increasing spectrum of simulated infection cue intensities, using heat-killed bacteria. We then measured reproductive effort (calling effort), survival and immune function (antibacterial activity, circulating haemocytes and haemocyte microaggregations). Inbred and outbred males diverged in how they responded to a low-dose infection cue: relative to unmanipulated males, outbred males decreased calling effort, whereas inbred males increased calling effort. Moreover, we found that inbred males exhibited higher antibacterial activity and numbers of circulating haemocytes compared with outbred males. These results suggest that an individual's inbreeding status may have consequences for context-dependent shifts in reproductive strategies, such as those triggered by infection.

Age-related reproduction of female Mongolian racerunners (Eremias argus; Lacertidae): Evidence of reproductive senescence.

The reproductive maturation hypothesis, the terminal investment hypothesis, and the senescence hypothesis are the most extensively evaluated hypotheses proposed to explain age-related patterns of reproduction in iteroparous organisms. Here, we evaluated these hypotheses for the Mongolian racerunner (Eremias argus), a short-lived lacertid lizard, by comparing reproductive traits between females that completed reproductive cycles under the same laboratory conditions in two consecutive years (2008 and 2009). Reproductive females gained linear size (snout-vent length) not only as they got 1 year older but also during the breeding season. Larger females generally laid eggs earlier and invested more in reproduction than did smaller ones. Females switched from laying smaller eggs in the first clutch to larger eggs in the subsequent clutches but kept clutch size and postpartum body mass constant between successive clutches in a breeding season and between years. Females that laid more clutches or eggs in 2008 did not lay fewer clutches or eggs in 2009. Of the traits examined, only clutch frequency, annual fecundity, and annual reproductive output were susceptible to ageing. Specifically, the clutch frequency was reduced by 1.1 clutches, annual fecundity by 3.1 eggs and annual reproductive output by 1.0 g in 2009 compared with 2008. Our results suggest that the reproductive maturation hypothesis better explains patterns of reproduction in young or prime-aged females of E. argus, whereas the senescence hypothesis better explains reproductive patterns in old females. The terminal investment hypothesis does not apply to any trait examined because no trait value was maximized in old females.

Declining fruit production before death in a widely distributed tree species, Sorbus aucuparia L.

Trees are commonly thought to increase their seed production before death. We tested this terminal investment hypothesis using long-term data on rowan trees ( Sorbus aucuparia ) and found no support. Rather, seed production declined significantly before death, which points to the potential detrimental effects of reproductive senescence on regeneration in stands of old trees. Aging poses a fundamental challenge for long-lived organisms. As mortality changes with with age due to actuarial senescence, reproductive senescence may also lead to declines in fertility. However, life history theory predicts that reproductive investment should increase before mortality to maximize lifetime reproductive success, a phenomenon termed terminal investment. To date, it is unclear whether long-lived, indeterminantly growing trees experience reproductive senescence or display terminal investment. We investigated fruit production of rowan (Sorbus aucuparia L.), widely distributed trees that live up to 150 years, as they approached death. In our study population in Poland’s Carpathian Mountains, 79 trees that died produced up to 20% fewer fruits in the years before their demise compared to 199 surviving trees of the same population. The pattern of reproductive investment in S. aucuparia is suggestive of age-independent reproductive senescence rather than terminal investment. These findings highlight that the understanding of the generality of life history strategies across diverse taxa of perennial plants is still in its infancy.

Queenless honey bees build infrastructure for direct reproduction until their new queen proves her worth.

The terminal investment hypothesis predicts that individuals will alter their reproductive investments based on reproductive prospects. This hypothesis, however, has never been tested at the colony-level, where reproductive prospects for thousands of individuals can change instantly with the death of a single individual: the queen. A honey bee queen's death also changes the reproductive mechanism; if the queen is not replaced, then workers reproduce directly, by producing males in reproductive comb-drone comb-before the colony dies. To test how workers respond to reproductive uncertainty, I made colonies queenless and measured comb building throughout the stages of rearing a replacement queen. Queenless workers built only drone comb throughout queen rearing, even when the colony possessed a virgin or mated queen. Only when the new queen started laying fertilized eggs did workers stop building drone comb. Despite queenless colonies being more likely to eventually rear a replacement queen (88%), than to fail (12%), workers still built drone comb for direct reproduction, even when the colony was provided with ample drone comb. Therefore, workers "pessimistically" invest in drone comb to prepare for direct reproduction, even while rearing a replacement queen to prevent that outcome. When faced with reproductive uncertainty, honey bees may "hope" for the best, but they prepare for the worst.

Age-dependent variation in the terminal investment threshold in male crickets.

The terminal investment hypothesis proposes that decreased expectation of future reproduction (e.g., arising from a threat to survival) should precipitate increased investment in current reproduction. The level at which a cue of decreased survival is sufficient to trigger terminal investment (i.e., the terminal investment threshold) may vary according to other factors that influence expectation for future reproduction. We test whether the terminal investment threshold varies with age in male crickets, using heat-killed bacteria to simulate an immune-inducing infection. We measured calling effort (a behavior essential for mating) and hemolymph antimicrobial activity in young and old males across a gradient of increasing infection cue intensity. There was a significant interaction between the infection cue and age in their effect on calling effort, confirming the existence of a dynamic terminal investment threshold: young males reduced effort at all infection levels, whereas old males increased effort at the highest levels relative to naïve individuals. A lack of a corresponding decrease in antibacterial activity suggests that altered reproductive effort is not traded against investment in this component of immunity. Collectively, these results support the existence of a dynamic terminal investment threshold, perhaps accounting for some of the conflicting evidence in support of terminal investment.

A dynamic threshold model for terminal investment.

Although reproductive strategies can be influenced by a variety of intrinsic and extrinsic factors, life history theory provides a rigorous framework for explaining variation in reproductive effort. The terminal investment hypothesis proposes that a decreased expectation of future reproduction (as might arise from a mortality threat) should precipitate increased investment in current reproduction. Terminal investment has been widely studied, and a variety of intrinsic and extrinsic cues that elicit such a response have been identified across an array of taxa. Although terminal investment is often treated as a static strategy, the level at which a cue of decreased future reproduction is sufficient to trigger increased current reproductive effort (i.e., the terminal investment threshold) may depend on context, including the internal state of the organism or its current external environment, independent of the cue that triggers a shift in reproductive investment. Here, we review empirical studies that address the terminal investment hypothesis, exploring both the intrinsic and extrinsic factors that mediate its expression. Based on these studies, we propose a novel framework within which to view the strategy of terminal investment, incorporating factors that influence an individual's residual reproductive value beyond a terminal investment trigger - the dynamic terminal investment threshold.

Ant queens increase their reproductive efforts after pathogen infection.

Infections with potentially lethal pathogens may negatively affect an individual's lifespan and decrease its reproductive value. The terminal investment hypothesis predicts that individuals faced with a reduced survival should invest more into reproduction instead of maintenance and growth. Several studies suggest that individuals are indeed able to estimate their body condition and to increase their reproductive effort with approaching death, while other studies gave ambiguous results. We investigate whether queens of a perennial social insect (ant) are able to boost their reproduction following infection with an obligate killing pathogen. Social insect queens are special with regard to reproduction and aging, as they outlive conspecific non-reproductive workers. Moreover, in the ant Cardiocondyla obscurior, fecundity increases with queen age. However, it remained unclear whether this reflects negative reproductive senescence or terminal investment in response to approaching death. Here, we test whether queens of C. obscurior react to infection with the entomopathogenic fungus Metarhizium brunneum by an increased egg-laying rate. We show that a fungal infection triggers a reinforced investment in reproduction in queens. This adjustment of the reproductive rate by ant queens is consistent with predictions of the terminal investment hypothesis and is reported for the first time in a social insect.

Testing the Terminal Investment Hypothesis in California Oaks.

The terminal investment hypothesis-which proposes that reproductive investment should increase with age-related declines in reproductive value-has garnered support in a range of animal species but has not been previously examined in long-lived plants, such as trees. We tested this hypothesis by comparing relative acorn production and radial growth among 1,000+ mature individuals of eight species of California oaks (genus Quercus) followed for up to 37 years, during which time 70 trees died apparently natural deaths. We found no significant differences in the radial growth, acorn production, or index of reproductive effort, taking into consideration both growth and reproduction among dying trees relative to either conspecific trees at the same site that did not die or growth and reproduction from earlier years for the focal trees that did eventually die. Furthermore, we found no consistent trade-off between growth and reproduction among trees that died, nor did dying trees significantly alter their relative investment in reproduction even as they underwent physical decline. Trees approaching the end of their lives are often in poor physical condition but do not appear to differentially invest more of their diminished resources in reproduction compared with healthy trees.

Influence of predator cues on terminal investment in courtship by maleSchizocosa ocreata(Hentz, 1844) wolf spiders (Araneae: Lycosidae)

Abstract Sexual signals play a critical role in mate attraction, but fitness benefits of signal production depend on a number of external and internal influences. Sexual signaling can be energetically expensive, and has potential to attract unwanted attention from predators. Male brushlegged wolf spiders, Schizocosa ocreata (Hentz, 1844) (Araneae: Lycosidae), actively signal to females in the leaf litter habitat during their spring breeding season, but face a tradeoff between current and future reproduction as the season progresses. The terminal investment hypothesis predicts that with fewer available females, increasing risk of predation, and stronger influence of senescence as the season progresses, males should take greater risks to secure mating. We explored this idea by exposing males of increasing ages to female cues alone or female cues combined with predator cues. We found little or no direct evidence to support the terminal investment hypothesis in this species, in that males across all ages esse...