Clutch size and malarial parasites in female great tits

life-history models predict an evolutionary trade-off in theallocation of resources to current versus future reproduction.This corresponds, at the physiological level, to a trade-offin the allocation of resources to current reproduction or tothe immune system, which will enhance survival and thereforefuture reproduction. For clutch size, life-history models predicta positive correlation between current investment in eggs andthe subsequent parasite load. In a population of great tits,we analyzed the correlation between natural clutch size of femalesand the subsequent prevalence of Plasmodium spp., a potentiallyharmful blood parasite. Females that showed, 14 days after hatchingof the nestlings, an infection with Plasmodium had a significantlylarger clutch (9.3 eggs ± 0.5 SE, n = 18) than uninfectedfemales (8.0 eggs ± 0.2 SE, n = 80), as predicted bythe allocation trade-off. Clutch size was positively correlatedwith the prevalence of Plasmodium, but brood size 14 days afterhatching was not. This suggests that females incur higher costsduring laying the clutch than during rearing nestlings. Infectionstatus of some females changed between years, and these changeswere significantly correlated with a change in clutch size aspredicted by the trade-off. The link between reproductive effortand parasitism may represent a possible mechanism by which thecost of egg production is mediated into future survival andmay thereby be an important selective force in the shaping ofclutch size     

The influence of maternal thermal environments on reproductive traits and hatchling traits in a Lacertid lizard,Takydromus septentrionalis

The thermal environment can induce substantial variation in important life-history traits. Experimental manipulation of the thermal environment can help researchers determine the contribution of this factor to phenotypic variation in life-history traits. During the reproductive season, we kept female northern grass lizards, Takydromus septentrionalis (Lacertidae), in three temperature-controlled rooms (25, 28 and 32 °C) to measure the effect of the maternal thermal environment on reproductive traits. Maternal thermal environment remarkably affected reproductive frequency and thereby seasonal reproductive output, but had little effect on reproductive traits per clutch or hatchling traits. Females kept at 32 °C produced more clutches and thus had shorter clutch intervals than females from 28 to 25 °C. Clutch size, clutch mass, relative clutch mass, egg size and hatchling traits did not vary among the three treatments. The eggs produced by the females were incubated at 27 °C and the traits of hatchlings were measured. The result that egg (offspring) size was independent of maternal thermal environments is consistent with the prediction of the optimal egg size (offspring) theory. The eggs produced by low temperature females (28 and 25 °C) took longer time to complete their post-oviposition development than did eggs produced by high temperature females (32 °C). This suggests that the eggs from low temperatures might have been laid when the embryos were at relatively early stages. Therefore, maternal thermal environment prior to oviposition could affect post-oviposition development in T. septentrionalis.     

In situ egg production and hatching success of the marine copepod Pseudocalanus newmani in Funka Bay and adjacent waters off southwestern Hokkaido, Japan: associated to diatom bloom

. Clutch size and egg viability were examined for wild femalesof Pseudocalanus newmani collected from four different stationsinside and outside Funka Bay, southwestern Hokkaido, Japan,during pre-phytoplankton-bloom conditions in 1997, and froma single station outside the bay during the period from pre-to post-blooming seasons in 1998, to examine spatio-temporalvariability of the reproductive parameters and to detect deleteriouseffects of diatom blooms on in situ copepod reproduction. Clutchsize and hatching success in P.newmani were relatively similarat the four stations in 1997. The eggs almost always hatchedcompletely, but deformed nauplii occurred in 20–40% ofthe cases. The proportion of deformed nauplii gradually decreasedwhen the females were fed on non-diatom Pavlova sp. in the laboratory,suggesting that the occurrence of deformed nauplii was relatedto the quality of in situ food particles, including diatoms.Clutch size varied from 12 to 30 eggs female^–1 duringthe study period in 1998; more than 90% of the variation couldbe explained by the body size of the females which dependedon ambient water temperature. Hatching success also varied,from 15 to 80%, but was not related to either clutch size, femalebody size, water temperature or diatom biomass. These resultssuggest that not only egg production but also hatchability shouldbe measured routinely when estimating recruitment of the copepodinto the planktonic population, and show that neither clutchsize nor egg viability are directly affected by diatom biomass.Although we could not obtain clear evidence of in situ deleteriouseffects of diatoms on abnormal embryos and nauplii, this mightbe related to methodological problems and properties or characteristicsof prey–predator interactions. Also, deformities of copepodnauplii have not yet been observed for reasons other than feedingon diatoms.     

Egg load is a cue for offspring sex ratio adjustment in a fig‐pollinating wasp with male‐eggs‐first sex allocation

Fig‐pollinating wasps (Agaonidae) only reproduce within fig tree inflorescences (figs). Agaonid offspring sex ratios are usually female‐biased and often concur with local mate competition theory (LMC). LMC predicts less female‐bias when several foundresses reproduce in a fig due to reduced relatedness among intra‐sexually competing male offspring. Clutch size, the offspring produced by each foundress, is a strong predictor of agaonid sex ratios and correlates negatively with foundress number. However, clutch size variation can result from several processes including egg load (eggs within a foundress), competition among foundresses and oviposition site limitation, each of which can be used as a sex allocation cue. We introduced into individual Ficus racemosa figs single Ceratosolen fusciceps foundresses and allowed each to oviposit from zero to five hours thus variably reducing their eggs‐loads and then introduced each wasp individually into a second fig. Offspring sex ratio (proportion males) in second figs correlated negatively with clutch size, with males produced even in very small clutches. Ceratosolen fusciceps lay mainly male eggs first and then female eggs. Our results demonstrate that foundresses do not generally lay or attempt to lay a ‘fixed’ number of males, but do ‘reset to zero’ their sex allocation strategy on entering a second fig. With decreasing clutch size, gall failure increased, probably due to reduced pollen. We conclude that C. fusciceps foundresses can use their own egg loads as a cue to facultatively adjust their offspring sex ratios and that foundresses may also produce more ‘insurance’ males when they can predict increasing rates of offspring mortality.     

Resting egg production and oviducal cycling in two sympatric species of alpine diaptomids (Copepoda: Calanoida) in relation to temperature and food availability

Resting egg production and oviducal cycling were investigatedfor the calanoid copepods Arctodiaptomus alpinus and Acanthodiaptomusdenticornis both in the laboratory and in a small karstic alpinelake by making a census of the number of eggs produced and theproportion of females in each of four morphologically distinguishedreproductive conditions each day in the laboratory or duringa 2–3 week period in lake enclosures. In the laboratory,individuals were maintained on a mixed diet of natural phytoplanktonat constant temperatures of 4, 10, 15 and 20C, respectively.Both species differed considerably in their temperature requirementsfor reproduction. Lifetime fecundity was highest at 10C in A.alpinusand at 20C in A.denticornis, with up to 327 eggs ^–1 spawnedin the former and up to 582 eggs ^–1 in the latter species.Unfavorable temperatures were further reflected in an increasein egg mortality and the allocation of time spent in a post-reproductivephase, as well as in a decrease of longevity. Increasing temperaturesenhanced egg production rates due to decreasing clutch productionperiods, although clutch size was negatively correlated withtemperature. Maximum rates reached 5.88 and 7.98 eggs ^–1day^–1 in the laboratory, and 0.73 and 0.55 eggs ^–1day^–1 in enclosures in A.alpinus and A.denticornis, respectively.Egg production rates and clutch size were clearly governed bynutritional conditions in the lake, but were less affected byfood supply in the laboratory. Here, rates of egg productionwere adapted to improving food supply by increasing the frequencyof spawning events, rather than the number of eggs per clutch.No correlation was found between female body size and reproductiveparameters in the laboratory. A very low proportion of totalclutch production resulted in clutches composed of subitaneouseggs, i.e. 0.14% in A.denticornis and 1.20% in A.alpinus. Oviducalphase duration allocations indicate that there exists a temperatureoptimum for gamete maturation.     

Is mating of copepods associated with increased risk of predation?

In laboratory experiments, we tested whether the cyclopoid copepodCyclops vicinus is more vulnerable to predation by juvenile(young-of-the-year) roach (Rutilus rutilus) and/or larvae ofthe phantom midge Chaoborus flavicans during mating than individualadult copepods of the same species. We also tested whether C.vicinusavoided mating, displayed shorter mating times and/or loweractivity when a predator (Chaoborus) was present, or when exposedto water in which predators (Chaoborus) were previously held.Fish ate the same quantity of pairs in copulation and individualegg-bearing females but significantly fewer females withouteggs. Chaoborus ate similar quantities of pairs and femalesbut fewer males. Pairs in copulation and egg-bearing femaleswere detected earlier by fish than individual non-ovigerousfemales or males, and pairs in copulation were captured mosteasily by fish. Cyclops vicinus mated less often when Chaoboruslarvae were present than in water in which Chaoborus was previouslyheld, or in pure tap water. However, predator presence or exposureto predator-treated water had no effect on (i.e. did not reduce)mating duration. Activity of C.vicinus pairs during copulationwas significantly lower when a predator was present, or in predator-treatedwater, than in pure tap water, and activity of pairs decreasedwith increasing activity of the predator. Although our experimentswere carried out under artificial conditions which cannot betransferred to the complex conditions in the field, they suggestthat mating in copepods is dangerous and that copepods haveevolved particular strategies to reduce the risk of predationduring mating.     

The problem of optimal clutch size in a tritrophic system: the oviposition strategy of the thistle gallfly Urophora cardui (Diptera,Tephritidae)

The problem of optimal clutch sizes is a central theme in life history theory. Optimal allocation of eggs is especially complicated for insects in tritrophic systems. In this study we analyze some of the processes determining clutch sizes of the thistle gallfly Urophora cardui, a monophagous tephritid fly associated with Cirsium arvense. U. cardui forms multilocular shoot galls, which vary broadly in their size and number of their gall cells. We investigate various fitness consequences of gall size. An analysis of the number of cells per gall (which is correlated with gall diameter and gall weight) showed that in U. cardui there is mutual facilitation rather than larval competition. Increasing numbers of larvae per gall led to a decreasing mortality and increasing larval weight. Larval weight in turn was positively correlated with the probability of survival to adulthood and with adult weight and fecundity. Thus, all fitness parameters measured favoured large galls. Clutch sizes in oviposition experiments were distinctly larger than the number of gall cells of field populations and in cage experiments, suggesting high mortality of eggs and/or early larval instars. There was a significant relationship between the internal structure (i.e., the size of the growing point) of the bud and clutch size, suggesting that U. cardui females are able to measure bud quality and adapt clutch sizes accordingly. Clutch size was positively correlated with the female's age at first oviposition and negatively with the number of previous ovipositions and previously laid eggs. Since the potential egg capacity per female is higher than the average number of larvae it is likely to produce during its short adult lifespan, U. cardui females tend to be time-limited rather than egglimited, which might favour large clutches once an appropriate oviposition site has been located. As the development of the gall and hence the fate of a clutch depends on a number of unpredictable factors, exclusive concentration of eggs in a few large clusters would involve risks which could be avoided by increasing the number of clutches. Therefore we interpret the high variation of clutch sizes in U. cardui as a mixed strategy of bet hedging and gambling.     

On the reproductive biology of the herbivorous spiny-tailed agamid Uromastyx philbyi in western Saudi Arabia

Abstract

The reproductive biology of the Spiny-tailed Agamid Uromastyx philbyi, a herbivorous desert lizard, was studied in western Saudi Arabia. Reproduction is seasonal, with mating in early spring (March), oviposition in late spring (May–June), and hatching in summer (July). The mean clutch size was 6.67 eggs. Eggs are large (mean mass = 7.48 g) and relative clutch mass (RCM) averaged 0.49. Clutch mass and RCM increased with increasing egg mass. Clutch size, egg size, clutch mass, and RCM were significantly correlated with maternal body size.     

Variations in fecundity over catchment scales: Implications for caddisfly populations spanning a thermal gradient

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The influence of competition between foragers on clutch size decisions in an insect parasitoid with scramble larval competition

The effect of competition between ovipositing females on theirclutch size decisions is studied in animals that lay their eggsin discrete units of larval food (hosts). In such species theeffect of competition depends on the form of the larval competitionwithin such units. In insect parasitoids, there might eitherbe contest (solitary parasitoids) or scramble competition (gregariousparasitoids) between larvae within a host For gregarious parasitoids,a decreasing clutch size with increasing competition betweenforagers is predicted. This prediction is tested in experimentsusing the parasitoid Aphaertta minuta. Parasitoids were eitherkept alone or in groups of four before the experiment, in whichthey were introduced singly in a patch containing unparasitizedhosts. Animals kept together laid on average clutches of 0.74eggs smaller than females kept alone (average clutch is 5.3),thereby confirming the prediction. Clutch size decreased withencounter number, which might be due to the adjustment of thefemale's estimate of the encounter rate with hosts. Finally,the results are compared with those reported for solitary parasitoids(that have scramble larval competition), for which it is predictedthat the clutch size will increase with increasing levels ofcompetition between females.     

Clutch size for Mediterranean loggerhead turtles (Caretta caretta)

We investigated the number of eggs laid and the total clutch volume for loggerhead turtles ( Caretta caretta L.) nesting on the Greek island of Cephalonia (38° 09' N, 20° 33' E) during 1990. Both the number of eggs laid and clutch volume were positively related to female body size, and the mean explained variation for these relationships (36·6% and 37·7%, respectively) were not significantly different. Clutch volume decreased as the season progressed because individuals laid successively smaller clutches. The inter-nesting interval (range 15–19 days) did not significantly influence the number of eggs laid or clutch volume. For individuals seen laying more than once, female size in combination with the number of a clutch in relation to all the clutches observed for that individual, together explained 65·5% of the variation in clutch volume.     

Egg production of the marine,planktonic copepod,Calanus pacificus Brodsky: Laboratory observations

When acclimated to a continuous, superabundant food supply and constant temperature, Calanus pacificus Brodsky females produce eggs at a weight-specific rate ranging from 0.13 · day^?1 at 8°C to 0.21 · day^?1 at 15°C. Maximum weight-specific egg production rates do not change with seasonal changes in female body size. The relationship between egg production rate and food concentration is hyperbolic, with threshold and critical concentrations that are high relative to other species for which data are available. Food concentration and temperature influence spawning frequency (i.e., the time required for oocytes to mature) much more than the number of eggs in a single spawning event (i.e., clutch size). Clutch size is significantly related to female body size.     

Strong evidence for selection for larger brood size in a great tit population

We measured the selection pressure on brood size in a recentlyestablished population of great tits (Parus major L.) in thenorthern Netherlands by manipulating brood size in three years(1995: n = 51, 1997: n = 66, 1998: n = 51), and we estimatedfitness consequences in terms of local survival of both offspringand parents. Enlarged broods had highest fitness; the offspringfitness component was positively affected by manipulation andthe parental fitness component was unaffected. Parental survivaland the probability that parents produced a second clutch werenot affected by the treatment. However, parents that had raisedenlarged broods produced their second clutch later in the season.Clutch size, brood size, and laying date of birds recapturedin the next breeding season were largely independent of thetreatment. We conclude that there is strong evidence for selectionfor larger brood size and reject the individual optimizationhypothesis for this population because the number of young inthe nest predicts fitness independently of the manipulationhistory.     

Observations on some reproductive traits of Triops granarius (Lucas) (Crustacea: Notostraca)

Reproductive traits of Triops granarius females were examined. Fertilization by males was a prerequisite for oviposition. Clutch size ranged from 1 to 154, and clutch frequency (per 24 hours) from 1 to 9. Clutch size generally increased with increasing body size and increasing age. Total reproductive output was dependent upon life span, and estimated at around 200 eggs for a life span of 25 days. Clutch data indicate that T. granarius may have r-selected traits.     

Maternal brooding in the children’s python <Emphasis Type="Italic">(Antaresia childreni)</Emphasis> promotes egg water balance

Parental care provides considerable benefits to offspring and is widespread among animals, yet it is relatively uncommon among squamate reptiles (e.g., lizards and snakes). However, all pythonine snakes show extended maternal egg brooding with some species being facultatively endothermic. While facultative endothermy provides thermal benefits, the presence of brooding in non-endothermic species suggests other potential benefits of brooding. In this study we experimentally tested the functional significance of maternal brooding relative to water balance in the children’s python, Antaresia childreni, a small species that does not exhibit facultative endothermy. Clutch evaporative water loss (EWL) was positively correlated with clutch mass and was much lower than expected values based on individual eggs. The conglomerate clutch behaved as a single unit with a decreasing surface area to volume ratio as clutch size increased. Maternal brooding had a dramatic impact on evaporation from eggs, reducing and possibly eliminating clutch EWL. In a separate experiment, we found that viability of unattended eggs is highly affected by humidity level, even in the narrow range from 75 to 100% relative humidity at 30.5°C (20–33 mg m⁻³ absolute humidity). However, the presence of the brooding female ameliorated this sensitivity, as viability of brooded clutches at 75% relative humidity was higher than that of non-brooded eggs at either the same absolute humidity or at near-saturated conditions. Overall, these results demonstrate that brooding behavior strongly promotes egg water balance (and thus egg viability) in children’s pythons.     

Multiple predator defence strategies in Daphnia pulex and their relation to native habitat

Daphnia may respond with an array of anti-predator defences(behavioural, morphological and life history) to a chemicalcue (kairomone) exuded by its predators: fish and Chaoborus.Given the wide array of potential responses, it is an interestingquestion whether anti-predator defences are coupled or independentof each other. Since anti-predator responses are costly andeven possessing the genetic information to respond to a certainpredator might involve a cost, clones may only react to predatorsthey co-occur with in nature. In this study, we provide evidencefor an uncoupling of responses by Daphnia pulex in several anti-predatordefences against Chaoborus. We were unable to detect a correlationbetween behavioural (migration), morphological (neck-spine induction)and life history [growth rate, neonate size and size at firstreproduction (SFR)] responses. Furthermore, anti-predator responsesdid not always comply with what is commonly believed. We foundthat Daphnia clones can migrate up or down when exposed to fishor Chaoborus kairomone and that population growth rate, neonatesize and SFR can increase or decrease in response to Chaoboruskairomone. We also show patterns in anti-predator defences thatseem to relate to the habitat from which clones were derived.Daphnia clones that were collected in habitats with Chaoborusas the dominant predator tended to react strongly to Chaoboruskairomone by migrating upward and producing neck-spines. Themigration behaviour against fish kairomone in these clones wasoften an unexpected upward migration. The Daphnia clone thatco-existed with fish predators showed a downward migration inthe presence of fish as well as Chaoborus kairomone. Clonesthat had occurred with either both or no predators had mixedresponses. We sometimes found an upward migration in combinationwith smaller body size as a response to Chaoborus kairomone.This may be interpreted as a behavioural defence against Chaoborusand a life-history defence against fish. Daphnia seem not toexhibit defence behaviour against predators they do not co-occurwith. It might be costly for Daphnia to maintain genetic informationto respond to these predators and protect that information fromgenetic drift.     

Complex effects of a kairomone of Chaoborus and an insecticide on Daphnia pulex

Daphnia pulex were reared in Chaoborus-conditioned water containingthe insecticide carbaryl, and their life history parametersand morphologies were investigated. The insecticide inhibitedthe animals' growth and reproduction and delayed their maturationtime more intensely in the chaoborus-conditioned water thanin the control Chaoborus-free water, indicating that a kairomoneof Chaoborus made the Daphnia more sensitive to the insecticide.The Chaoborus conditioned water induced neckteeth formationof D.pulex in instars 1–2 and elongated the intermoultingperiod of juveniles. The moulting to the spined morphs and elongationin duration of juvenile stages seemed to increase the risk ofdamage from the insecticide. The potential population growthrate of D.pulex in treatments was estimated as a possible fitnessindicator of the animals. It was reduced synergistically bythe kairomone of Chaoborus and the insecticide. Some individualskept neckteeth until the third or fourth instar stage when theywere exposed to sublethal concentrations of the insecticidein the Chaoborus-conditioned water. This was considered as aresult of synergistic effects of both the kairomone and theinsecticide. Insecticides may be a factor inducing further developmentof protuberant structures in cyclomorphic Daphnia in naturalwater bodies.     

Reproductive success and foraging of the crab spider Misumena vatia

Summary Reproductive success and growth rate data were collected for individually marked crab spiders Misumena vatia (Clerck) in 1980, 1981, and 1982. All measures of reproductive success were found to be quite variable between individuals within years, but did not differ between years. Reproductive effort (mass of clutch/prereproductive mass of female) was the least variable measurement and was not correlated with female weight at reproduction. Clutch weight and number of eggs per clutch were highly correlated with female reproductive weight. Egg weight was not correlated with the number of eggs per clutch. Hatching success did not vary with clutch size and averaged 94.5%. Growth rates of spiders were highly variable, indicating large variation in feeding rate. In 1981 and 1982, approximately 20% of female spiders were unable to capture enough prey to grow and reproduce. Primary prey species differed in weight and in their contribution to spider egg production. Spiders attacked a larger percentage of bumblebees but captured a larger percentage of honeybees. There was no simple relationship between diet choice and reproductive success. Spiders which selected suboptimal umbels to forage on some or all of the time, however, had significantly lower reproductive success than spiders choosing the best umbels.     

安徽滁州雌性丽斑麻蜥繁殖特征

在安徽滁州地区丽斑麻蜥(Eremias argus)年产两窝卵。窝卵数及窝卵重与雌体体长呈正相关,相对窝卵重与雌体体长无关,卵重与窝卵数无关。窝卵数、窝卵重及卵重在窝序间无明显的差异。卵长径与卵短径呈正相关,卵长径与窝卵数呈负相关,而卵短径与窝卵数无关。雌体主要通过增加窝卵数增加繁殖输出。     

Nest desertion by a cowbird host: an antiparasite behavior or a response to egg loss?

Natural selection can favor songbirds that desert nests containingeggs of the parasitic brown-headed cowbird (Molothrus ater).However, the high variability in desertion of parasitized nestswithin species is perplexing in light of the typically highcosts of parasitism. Because nest desertion can also be a responseto partial clutch predation, we first asked if Bell's vireos(Vireo bellii) deserted nests in response to the presence ofcowbird eggs (antiparasite response hypothesis) or to egg removalby predators and female cowbirds (egg predation hypothesis).Second, we asked whether variation in nest desertion was dueto intrinsic differences among individuals or to variation innest contents. We monitored a large number of nests (n = 494)and performed a clutch manipulation experiment to test thesehypotheses. The number of vireo eggs that remained in a nestwas a strong predictor of desertion both within and among pairs.Neither the presence of a single cowbird egg, which leads tonest failure for this host, nor the number of cowbird eggs receivedin a vireo nest influenced nest desertion. Furthermore, vireosdid not desert experimental nests when we immediately exchangedcowbird eggs for vireo eggs but deserted if we removed vireoeggs and replaced them with cowbird eggs the following morning.Desertion of parasitized nests by Bell's vireos can be almostentirely explained as a response to partial or complete clutchloss and does not appear to have been altered by selection frombrood parasitism.