Interference by the mantispid Mantispa uhleri with the development of the spider Lycosa rabida

Abstract. 1. The larvae of Mantispa uhleri Banks (Neuroptera: Mantispidae) board spiders to await the production of an egg sac containing their obligate developmental food. While aboard the spider, larvae maintain themselves by feeding on spider blood. This parasitic behaviour was investigated by allowing larvae to board sixth instar Lycosa rabida Walckenaer (Araneae: Lycosidae). Larval parasitism has a direct and indirect effect on the developmental physiology of the spider.
2. The direct effect, equal in both spider sexes, is an increase in development time and a decrease in adult size.
3. The indirect effect on development time and adult size is brought about by the loss of an instar in female spiders only. Parasitized females were mature at nine or ten instars; control females at ten or eleven. Male instar number was not affected; both control and parasitized males were mature at nine or ten instars.
4. The net result is that parasitized female spiders are even smaller than would be predicted from the direct effect alone, but actually mature faster than control females. In males there is only the direct effect. The adaptive significance of this sexually dimorphic response is discussed.     

Phenotypic variability in nesting success among Osmia lignaria propinqua females in a glasshouse environment: (Hymenoptera: Megachilidae)

Abstract. 1. The phenotypic variability of several estimates of fitness among seventeen nesting female O.lignaria Cresson was examined in a glasshouse provided with abundant resources.
2. Females exhibited wide variation in: (a) number of nests and cells made and in their rate of construction, (b) the incidence of mortality of offspring, and (c) the percentage of male offspring produced (52–100%).
3. In confirmation of earlier field studies: (a) more offspring of both sexes were produced during the first half of the nesting season, (b) most female offspring were produced early in the nesting season, (c) most parental investment during the last half of the flight season was made in male offspring, (d) there was a decrease in offspring size with season, with females decreasing more than males, and (e) the ratio of female/male body weight was = 1.8. These results appear to be due to ageing rather than to a reduction in resource availability.
4. No relation between female size and fecundity was evident.
5. There was no evidence that small females tended to produce a greater proportion of the smaller sex (males) than did large females.
6. Smaller offspring of both sexes had a much greater probability of dying overwinter than did large offspring. However, where the sexes overlapped in size (large males, small females), almost all females died and almost all males survived. We hypothesize that as size of offspring produced declines with season, a greater proportion of males are produced because they have a much greater probability of surviving at small body sizes.     

Sex ratio in the spider Pityohyphantes phrygianus affected by winter severity

The sex ratio of subadult Pityohyphantes phrygianus was recorded before and after five winters in a natural population in coniferous forest in south-west Sweden. In autumn, the proportion of males was on average 33–8% (range 28–7–40–1) and the proportion of females was 66–2% (59–9–71–3). In each of the winters 1981 82, 1982–83, 1984–5, the proportion of males decreased significantly. The proportion of males decreased more the lower the February mean temperature. Field experiments showed that low ambient temperature in winter caused high mortality among spiders. Experimental data also suggest that males are more vulnerable to low winter temperatures than are females. Indirect evidence indicates that neither predation pressure nor starvation alone are likely to cause the observed changes in sex ratio in the three winters. However, the combined effect of sex differences in predation by birds and cold-induced mortality may explain why males disappear faster than females in certain winters.     

Influence of sex, body size, and reproduction on overwinter lipid depletion in brook trout

Expressed as percentages of total fresh body weight, lipids of brook trout Salvelinus fontinalis declined between October and April: reproductive males from 2·89 to 1·22%, reproductive females from 3·19 to 1·84%, and non-reproductive males and females from 2·75 to 2·08%. The absolute and proportional overwinter reduction in lipids among reproductive trout was more than twice that of non-reproductive trout, with reproductive males losing significantly more lipids than reproductive females. Larger reproductive individuals lost more lipids during winter, relative to body size, than smaller individuals, although such an effect was not evident among non-reproductive trout. The average overwinter reduction in lipids for reproductive males (58%), females (42%), and non-reproductive trout (24%) was negatively associated with mark-recapture estimates of overwinter survival probabilities of 0·27, 0·36, and 0·58, respectively, providing support for the hypothesis that energy is allocated to reproduction to the detriment of post-reproductive survival. Our emergent hypothesis that reproductive costs differ between sexes, and the life history consequences thereof, merit further study.     

On the life history of the lesser gurnard (Scorpaeniformes: Triglidae) inhabiting the Agulhas Bank, South Africa

Growth analysis based on sectioned sagittal otoliths revealed the lesser gurnard Chelidonichthys queketti on the Agulhas Bank to be relatively fast growing and long lived, with ages of up to 7 years being recorded. Total length at age (mm) was described best by the specialized von Bertalanffy growth model as L _T=306·1 (1 − e^{0·53(t+0·18)}). First approximations of total, natural and fishing mortality rates were determined at 0·73, 0·38 and 0·35 year⁻¹ respectively. The adult population was male dominated with a sex ratio of 1 female: 1·2 males with the mean size of males and females being similar. The lesser gurnard is an iteroparous species with females maturing by the end of the first year of life (195 mm L _T), thereafter spawning throughout the year with reproductive activity peaking over spring and late summer. The lesser gurnard appears to exhibit similar life-history patterns to other triglid species in that it can be classified as a generalist. Generalistic life-history characteristics such as a fast growth rate, early sexual maturity at a relatively large size, a non-seasonal spawning pattern, feeding on a variety of prey organisms and the ability to inhabit various substrata could all contribute to it maintaining a high biomass on the Agulhas Bank.     

The functional relationship between peak spring floods and survival and growth of juvenile Atlantic Salmon (Salmo salar) and Brown Trout (Salmo trutta)

1. The effects of high spring floods on survival and growth of juvenile Atlantic Salmon, Salmo salar, and Brown Trout, Salmo trutta , are explored, using data from a long-term study in the River Saltdalselv, northern Norway. The flow regime in this river is typical for northern rivers.
2. There was considerable variation in year class strength of both species.
3. Mortality of Atlantic Salmon increased significantly in years with high discharge during the alevin stage as well as the first week after emergence. High discharge during the egg stage and more than 1 week after emergence seemed to be of minor importance. Water temperature at emergence was rather high (average 10·5 °C) and did not significantly affect year class strength.
4. Brown Trout emerged earlier than Atlantic Salmon at an average water temperature of 8·2 °C. Highest mortality was observed in years with low water temperatures at emergence as well as high discharge during the alevin stage.
5. For 1-year-old fish or older, the size of the spring peak flood did not influence mortality significantly.
6. Growth of Atlantic Salmon parr was diminished in years with a high peak spring flood. A similar effect on Brown Trout was not detected.     

Comparative nesting and feeding ecology of skylarks Alauda arvensis on arable farmland in southern England with special reference to set-aside

1. A study of skylark Alauda arvensis L. breeding ecology in relation to crop type was carried out from April to August 1992 on arable land in southern England. Set-aside land was included in this comparative study.
2. Territory density averaged 0·15 ha⁻¹. It was 2–3 times higher in fields of set-aside and grass, especially permanent pasture, than in winter and spring-sown cereals.
3. Territory size was nearly twice as large in fields of winter cereals (4·5 ha) than in other crop types (2·5 ha). Where set-aside was present on one farm, territory size in set-aside (1·7 ha) was a third lower than in cereals and grass.
4. Nesting began in set-aside and permanent pasture in April and peaked in late May. Nesting was not detected in spring barley until late May and in silage grass until early June. The density of successful nests in set-aside fields was more than double that in any of the arable crop types.
5. Average clutch size at hatching was 3·91 eggs in fields of set-aside, over 15% higher than in silage grass (3·40) and in spring barley (3·27).
6. Fledging success did not differ according to crop type, but productivity, expressed as the number of fledglings produced per hectare, was 0·50 in set-aside, 0·13 in silage grass, and 0·21 in spring barley. Nests with chicks were not found in fields of winter cereals. The causes of chick death were thought to be predation in set-aside fields, farming practices in silage grass fields, and suspected starvation in spring cereals.
7. The potentially high nesting success of skylarks in set-aside implies that sympathetic set-aside management could play an important part in reversing its decline across the European Union.     

Local adjustment of sex ratio in the spider Pityohyphantes phrygianus

The sex ratio in five populations of the sheetweb spider Pityohyphantes phrygianus in southwest Sweden was investigated in July and September, the spiders being juveniles and subadults, respectively. In July, no heterogeneity between the five populations was found and the pooled samples revealed a sex ratio of 36% males and 64% females. A significant heterogeneity in sex ratio was observed in the September samples; the proportion of males varied between 23% and 40%. In several cases, male mean weight differed significantly between the five populations, comparing the July and September samples separately. This suggests that the prey availability varied considerably between the sites. In September, dead spiders were collected in two sites. This mortality was probably caused by starvation and the mortality rates differed significantly between the two sites. Indirect evidence suggested that the local sex ratio was affected by the prey availability, males being at a relative disadvantage in poor sites. Local environmental conditions may govern the adult sex ratio in this spider by causing differential mortality and/or migration.     

Male mating success and risk of predation in a wolf spider: a balance between sexual and natural selection?

1. Traits that benefit males through sexual selection are simultaneously expected to impair males by provoking costs through natural selection. If we consider the two male fitness components, mating success and viability, then we may expect that the increase in male mating success resulting from a larger trait size will be counterbalanced by an increase in viability costs.
2. We studied the benefits and costs of male mate searching and sexual signalling activity in the wolf spider Hygrolycosa rubrofasciata . In the field, males search females actively and court them by drumming dry leaves with their abdomen. Females have been shown to prefer males with high drumming rate. Male moving and especially drumming is energetically highly demanding and drumming results in significant mortality costs.
3. Our objective in this study was to determine whether male mate-searching activity or drumming activity affect male mating success and the risk of males being predated.
4. It was evident that both higher mate-searching activity and higher drumming activity benefited males by increasing their mating success. Higher mate-searching activity clearly impaired males by causing direct increase in predation risk. There was also a slight tendency that more actively drumming males had higher risk of predation and from all of the predated males 13.3% were caught directly after they had drummed. Furthermore, male drumming activity decreased drastically in the presence of the predator.
5. We conclude that in H. rubrofasciata both increased mate-searching activity and drumming activity benefit males through sexual selection, but at the same time natural selection provokes direct balancing costs on the same traits.     

Asymmetrical costs of sexual conflict in the seaweed fly, Coelopa frigida

Abstract.  1. Sexual conflict can play an important role in the evolution of animal life-history characteristics, including lifespan. Seaweed flies show an increase in mortality rates when exposed to brown algae. The seaweed stimulates females to oviposit and males to mount females. Females typically respond to male mounts by performing a violent rejection response.
2. Here the contribution of sexual conflict to the increase in mortality seen in the presence of seaweed was determined. The survival of single and mixed sex pairs of flies was followed in the presence and absence of seaweed.
3. The two sexes showed differential survival rates, with females living longer in the absence of seaweed. The presence of seaweed reduced survival in both sexes. In the presence of seaweed, female survival was lower when paired with a male. Over 40% of the reduction in survival in females in the presence of seaweed appears to be attributable to sexual conflict.
4. The presence of a female did not significantly affect male survival. Thus the mortality cost of being in the presence of the opposite sex and seaweed appears highly asymmetric.
5. In the presence of seaweed, female survival was lower when females were paired with small males. Small males exhibit higher levels of harassment of females, thus it is argued that pre-copulatory sexual conflict is the probable cause of the increased mortality cost to females of being in the presence of both males and seaweed.     

Demography of the vegetable ivory palm Phytelephas seemannii in Colombia, and the impact of seed harvesting

1. The demography of the vegetable ivory palm Phytelephas seemannii was studied on the Pacific coast of Colombia and a female-based matrix model was used to determine the proportion of seed that can be sustainably harvested from the population.
2. The density of the palm stands ranged from 240 to 420 adult palms ha⁻¹. The sex ratio was 1:1 and palms of both sexes produced leaves at the same rate. Seedlings produced 1·2 leaves per year on average, juveniles 1·8, and adults 6·1–7·4.
3. Adult females had fewer leaves than males (18·5 vs. 21·3 on average). Leaves of females lasted about 2·7 years in the crown, those of males about 3·2 years.
4. Stems creep on the ground, growing at the apex and often dying behind, the extant portion not always reflecting the palm's total age. The longest stem of a female palm in the study plots was 2·5 m, corresponding to an 'extant' age of 85 years; the longest stem recorded was that of a male (outside the plots) 10·5 m long, corresponding to an 'extant' age of 184 years. These figures reflect differences in growth habit, not in longevity.
5. Reproduction began at about 24 years, when the palm still lacked an above-ground stem.
6. The population growth rate λ was 1·059, and was most sensitive to changes in survival of juveniles and adults, and relatively insensitive to changes in fecundity and growth.
7. River channel migration is the most important cause of adult mortality. Phytelephas seemannii is apparently an efficient colonizer of the understorey in the late phases of riverine forest succession.
8. The population can tolerate a harvest intensity of 86% of all seeds before λ decreases to the equilibrium level of 1·00. Monitoring of the populations under intensive harvesting is required.     

The influence of sex and body size on food habits of a giant tropical snake, Python reticulatus

1. In many animal species, dietary habits shift with body size, and differ between the sexes. However, the intraspecific range of body sizes is usually low, making it difficult to quantify size-associated trophic shifts, or to determine the degree to which sex differences in diet are due to body-size differences. Large snakes are ideal for such a study, because they provide a vast range of body sizes within a single population.
2. More than 1000 Reticulated Pythons ( Python reticulatus ) from southern Sumatra were examined, with specimens from 1·5 to > 6 m in snout–vent length, and from 1 to 75 kg in mass. Females attained much larger body sizes than did conspecific males (maxima of 20 vs 75 kg, 5 vs 7 m), but had similar head lengths at the same body lengths.
3. Prey sizes, feeding frequencies and numbers of stomach parasites (ascarid nematodes) increased with body size in both sexes, and dietary composition changed ontogenetically. Small snakes fed mostly on rats, but shifted to larger mammalian taxa (e.g. pangolins, porcupines, monkeys, wild pigs, mouse deer) at 3–4-m body length.
4. Adult males and females showed strong ecological divergence. For some traits, this divergence was entirely caused by the strong allometry (combined with sexual size dimorphism), but in other cases (e.g. feeding frequency, dietary composition), the sexes followed different allometric trajectories. For example, females shifted from rats to larger mammals at a smaller body size than did conspecific males, and feeding frequencies increased more rapidly with body size in females than in males. These allometric divergences enhanced the degree of sex difference in trophic ecology induced by sexual size dimorphism.     

Predation, sensitivity, and sex: why female black rhinoceroses outlive males

Among sexually dimorphic, polygynous mammals, adult femalestend to outlive males and respond more strongly to predatorsthan males. We asked whether a monomorphic, polygynous speciesvirtually immune to predation due to large size (black rhinoceros,Diceros bicornis) conforms to this pattern. Data on 193 interactionswith lions (Panthera leo) and spotted hyenas (Crocuta crocuta)in two nonhabituated populations in Namibia studied from 1991to 1993 revealed that: (1) females were more vigilant or aggressivethan males to either of the potential predators; and (2) whethersolitary or with calves, females attacked more often than males.Although solitary females tended to be more aggressive to lionsthan to hyenas, neither females with calves or males seemedto discriminate between the two carnivores. We also simulatedthe behavior of human predators (poachers) during 69 encounterswith rhinoceroses. While both sexes abandoned local sites becauseof our presence, females ran farther than males, covering upto 40 km in a day. These findings implicate a behavioral mechanismto explain why secondary sex ratios favor females - males aremore prone to human predation, a prediction consistent withdata from 12 populations throughout Africa. Black rhinocerosesappear to be an unanticipated exception to the well-establishedpattern of male-biased mortality in polygynous mammals; in theabsence of intense human predation (a recent event), male mortalityfails to exceed that of females, suggesting that intrasexualcompetition in a polygynous mammal: may not be the primary causeof unbalanced secondary sex ratios. Our results on the causesof sex differences in mortality and in responsiveness to differentpredators reinforce the relevance of behavioral ecology to conservation;such information is necessary for planning how best to minimizenegative human influences on the few remaining wild Africanrhinos.     

Age, sex ratio and timing of the catch of kelts and ascending Atlantic salmon in the subarctic River Teno

By 15 June, 82% of the catch of Atlantic salmon Salmo salar kelts had been taken from the middle part of River Teno, northern Scandinavia. The median date of capture was 4 June for males and 8 June for females. Salmon of 1–4 sea–winters (SW) of both sexes survived spawning to return to sea as kelts. Among males, 1 SW kelts were caught earliest in the spring and 3 SW latest, but among females 4 SW were earliest, then 3 SW and finally 1 and 2 SW. There were 17 river and sea–age combinations among the kelts compared with 23 among the ascending salmon. The smolt age distribution and the mean smolt age differed significantly only between female 2 SW ascending salmon (3·97 years) and kelts (4·14 years). The proportion of 1 SW females was higher and that of 3 SW males lower among kelts than among ascending salmon. The proportion of males among 1 SW ascending salmon was 80% but among kelts only 57%. Similarly, the proportion of males among 3 SW fish was 21% for ascending salmon but only 7% for kelts. Hence overwinter mortality was higher among males. Male and female kelts of 1 and female kelts of 2 SWhad a greater mean length than ascending salmon in corresponding groups indicating a better survival of larger fish within an age group. Grilse ascend rivers after most kelts have left, but the main catch of ascending 2–3 SW salmon takes place concurrently with kelts leaving the river, inadvertently targeting kelts in the fishery.     

Size-dependent predation risk in tree-feeding insects with different colouration strategies: a field experiment

1. Body size is positively correlated with fecundity in various animals, but the factors that counterbalance the resulting selection pressure towards large size are difficult to establish. Positively size-dependent predation risk has been proposed as a selective factor potentially capable of balancing the fecundity advantage of large size.
2. To construct optimality models of insect body size, realistic estimates of size-dependent predation rates are necessary. Moreover, prey traits such as colouration should be considered, as they may substantially alter the relationship between body size and mortality risk.
3. To quantify mortality patterns, we conducted field experiments in which we exposed cryptic and conspicuous artificial larvae of different sizes to bird predators, and recorded the incidence of bird attacks.
4. The average daily mortality rate was estimated to vary between 4% and 10%. In both cryptic and conspicuous larvae, predation risk increased with prey size, but the increase tended to be steeper in the conspicuous group. No main effect of colour type was found. All the quantitative relationships were reasonably consistent across replicates.
5. Our results suggest that the size dependence of mortality risk in insect prey is primarily determined by the probability of being detected by a predator rather than by a size-dependent warning effect associated with conspicuous colouration. Our results therefore imply that warningly coloured insects do not necessarily benefit more than the cryptic species from large body size, as has been previously suggested.     

Differences in size and growth rates of male and female migrating Japanese eels in Pearl River, China

The Sr/Ca ratios in otoliths of silver Japanese eels Anguilla japonica , in Pearl River, China, indicated that both sexes did not stay in brackish water and grew in fresh water from the glass eel stage until spawning migration. This did not support the hypothesis that females tended to distribute upstream and males might be restricted to estuaries. The back-calculated total length of males at glass eel stage was not significantly different from that of females, indicating that the hypothesis that small glass eels became males and larger ones became females may not be true. The mean (±S.D.) age and total length of males at migration were 6·4±1·6 years and 48·3±4·5 cm, which were significantly smaller than for females, 8·3±1·6 years and 61·4±4·1 cm. The age of migration was related inversely to growth rate for both sexes. Growth parameters of the von Bertalanffy growth equation were K =0·21 cm year°1, L _∞=55·7 cm and t _o=-0·55 year for males and K =0·14 cm year⁻¹, L _∞=77·5 cm and t _o=-0·60 year for females. The difference in asymptotic length ( L _∞) between males and females may be because females postpone migration to achieve larger size for maximizing reproductive success.     

Predation, group size and mortality in a cooperative mongoose, Suricata suricatta

1. In social mammals where group members cooperate to detect predators and raise young, members of small groups commonly show higher mortality or lower breeding success than members of large ones. It is generally assumed that this is because large group size allows individuals to detect or repel predators more effectively but other benefits of group size may also be involved, including reduced costs of raising young and more effective competition for resources with neighbouring groups.
2. To investigate the extent to which predation rate affects survival, we compared mortality rates in two populations of suricates ( Suricata suricatta ), one living in an area of high predator density (Kalahari Gemsbok Park) and one living in an area of relatively low predator density (neighbouring ranchland). Most aspects of feeding ecology and growth (including time spent feeding, daily weight gain, growth, adult body weight, breeding frequency and neonatal mortality) were similar in the two populations. In contrast, mortality of animals over 3 months old was 1·7 times higher in the Park than on ranchland.
3. Mortality of juveniles between emergence from the natal burrow and 6 months of age was higher in small groups than large ones in the Park but significantly lower in small groups than large ones on ranchland. Adult mortality declined in larger groups in both areas.
4. The tendency for survival to be low in small groups had far-reaching consequences for the risk of group extinction. During a year of low rainfall in the Park, all groups of less than nine animals became extinct and population density declined to around a third of its initial level. We argue that high group extinction rates are to be expected in species where survival declines in small groups and mortality rates are high.     

Blackcaps Sylvia atricapilla stopping over at the desert edge; inter- and intra-sexual differences in spring and autumn migration

Spring and autumn Palaearctic-African migration patterns of Blackcaps Sylvia atricapilla during stopover at Elat, Israel, showed that males appeared significantly earlier than females during spring but not during autumn migration, suggesting that in males there is a stronger drive to reaching breeding territories early. The difference in mean appearance dates between sexes in spring tended to be greatest in years when the males appeared earliest. Longer spread of passage (the dates between which the central 50% of individuals were captured) for each sex in spring was found in years with an early mean passage datebut was significant only for females. These observations suggest that the timing of Blackcap migration is governed not only by endogenous factors but also by exogenous factors, and when the environmental conditions are unfavourable, the differences in passage dates between sexes decrease and the passage lengths shorten. The early individuals (both males and females) that stopped over at Elat in spring were those with relatively small body size (as indicated by relatively short wings) and relatively large fat reserves and in good body condition (as indicated from fat score and body mass/wing-length ratio). No differences in body size between early and late transients were detected during the autumn migration, but late birds of both sexes carried larger fat reserves. These phenomena may be explained either by leap-frog migration or by differential fitness among wintering males and females or both, with only the fittest Blackcaps being capable of an early departure. These individuals probably face much less intensive intra- and interspecific competition with residents and other transients in stopover sites than do later transients.     

Size and mortality rates of Glossina pallidipes in the semi-arid zone of southwestern Kenya

Seasonal changes in the mean size of tsetse, Glossina pallidipes Austen, as indicated by wing vein length, were monitored during 1983-86 at Nguruman, southwestern Kenya. Changes in size of nulliparous females and wing fray category 1 males were shown to be correlated with the relative humidity 2 months before they were captured. Soil temperature when flies were in the pupal stage had much less effect. Size dependent mortality was demonstrated, with the mean size of flies emerging from pupae significantly less than that of field-caught flies. This mortality must occur at emergence, since there was no evidence of size-dependent mortality once the flies became available to the trap. Size was correlated with density-independent mortality acting on the parent population 2 months previously. It might therefore be possible to use size as an index of the intensity of such mortality. This could be useful when assessing the level of additional mortality required to suppress tsetse populations.     

Risky mate search and mate preference in the golden orb-web spider (Nephila plumipes)

Mate searching is a risky behavior that decreases survival byincreasing predation risk and the risk of energy depletion.However, few studies have quantified actual mortality duringmate search, making it difficult to predict mate searching andmating strategies. Using a mark and recapture study, we examinedmate-searching success in a highly sexually dimorphic species,the golden orb-web spider (Nephila plumipes). We show that despitethe high-density aggregations of this species, male survivalduring mate searching is extremely low (36%) and is phenotypeindependent. Surprisingly, males that survived mate search werein better condition after recapture than prior to release, mostlikely due to kleptoparasitism on females' webs. In a complementaryrelease experiment in a field enclosure, we show that malesare choosy and adjust their choice of female depending on theirown condition and weight. Thus, the high mortality rate of searchingmales in the field may be a cost of choosiness because releasedmales traveled further than necessary to settle on females.Although males were choosy about female phenotypes, they didnot avoid webs with rival males already present. This suggeststhat the cost of continued searching outweighs the cost of competitionbut not the cost of mating with certain females. Further examinationsof mate-searching risk in other species in reference to theirmating system and environmental conditions are necessary todetermine the occurrence and effects of high mortality ratesduring searching.