Female fecundity traits in wild populations of African annual fish: the role of the aridity gradient

The evolution of life history is shaped by life expectancy. Life‐history traits coevolve, and optimal states for particular traits are constrained by trade‐offs with other life‐history traits. Life histories contrast among species, but may also diverge intraspecifically, at the level of populations. We studied the evolution of female reproductive allocation strategy, using natural populations of two sympatric species of African annual fishes, Nothobranchius furzeri and Nothobranchius orthonotus. These species inhabit pools in the Mozambican savanna that are formed in the rainy season and persist for only 2–10 months. Using 207 female N. furzeri from 11 populations and 243 female N. orthonotus from 14 populations, we tested the effects of genetic background (intraspecific lineage) and life expectancy (position on the aridity gradient determining maximum duration of their temporary habitat) on female fecundity traits. First, we found that variation in female body mass was small within populations, but varied considerably among populations. Second, we found that fecundity was largely defined by female body mass and that females spawned most of their eggs in the morning. Third, we found that the trade‐off between egg size and egg number varied among lineages of N. furzeri and this outcome has been confirmed by data from two separate years. Overall, we demonstrate that local conditions were important determinants for Nothobranchius growth and fecundity and that eggs size in arid region was less limited by female fecundity than in humid region.     

Influence of genetic and environmental factors on egg and clutch sizes among populations of Paratya australiensis Kemp (Decapoda: Atyidae) in␣upland rainforest streams, south-east Queensland

Headwater populations of the common shrimp Paratya australiensis were sampled to examine the factors influencing egg and clutch size. Much of the spatial variation in these reproductive traits was associated with differences in altitude, with upper sites having larger eggs and smaller clutches. Mean egg size at high-altitude sites was higher than that previously reported for this species. Temporal variation in egg and clutch size was also observed, with significant increases in egg size during the breeding season. At most sites, this increase was accompanied by a decrease in clutch size. The combined effect of the reciprocal patterns in egg volume and clutch size resulted in relatively little spatial and temporal variation in reproductive effort. In a field experiment, using a unique genetic marker, shrimps were translocated between two sites with significantly different egg sizes. After one generation (18 months), the mean egg size of translocated females was the same as that of females from the `source' population. In contrast, clutch size changed towards that of the resident females. This suggests that egg size is under strong genetic control, while clutch size is influenced by the environment. Received: 14 July 1997 / Accepted: 9 March 1998     

Life history traits related to resource partitioning between synhospitalic species of Colocasiomyia (Diptera, Drosophilidae) breeding in inflorescences of Alocasia odora (Araceae)

Abstract.

• 1 Two closely related species of Colocasiomyia alocasiae (Okada) and C.xenalocasiae (Okada) (Diptera, Drosophilidae) breed in inflorescences of Alocasia odora C. Koch (Araceae), a hermaphroditic understorey clonal herb.
• 2 The two drosophilid species form a synhospitalic pair in Okinawa with alocasiae breeding in the upper half of the inflorescence and xenalocasiae breeding in the lower half.
• 3 C.alocasiae also has the following combination of life history traits: small body size, many eggs, and early reproductive maturity. In contrast, xenalocasiae can be described as having larger body size, fewer eggs, and delayed reproductive maturity.
• 4 Resource partitioning between the two species on the same host may be affected by these life history traits which are associated with their larval habitats.

     

Variation in fecundity and other reproductive traits in freshwater mussels

1. Life histories of the highly diverse and endangered North American freshwater mussel fauna are poorly known. We investigated reproductive traits of eight riverine mussel species in Alabama and Mississippi, U.S.A.: Amblema plicata, Elliptio arca, Fusconaia cerina, Lampsilis ornata, Obliquaria reflexa, Pleurobema decisum, Quadrula asperata and Q. pustulosa, and compare our results with existing life history information for other species. 2. These eight species had reproductive traits characteristic of large, outcrossing populations: hermaphrodites were rare, we found no evidence of protandry, and sex ratios were even or slightly male‐biased. 3. Age at sexual maturity varied among species, ranging from <1 to 2 years for L. ornata to 3–9 years for Q. asperata. In all species, most mature females participated in reproduction and fertilisation success was high. 5. Fecundity was related positively to both length and age, but length was the best predictor. In six species, fecundity increased exponentially with increasing size; in two species the rate of increase in fecundity declined in larger animals. In four species, fecundity declined in older animals. These latter results indicate weak reproductive senescence; however, in all species, older individuals continued to produce large numbers of offspring. Mean annual fecundity differed widely among species ranging from 9647 to 325 709. Within‐species differences in fecundity were found among rivers and among populations within a river. 6. The wide variation in reproductive traits among species indicates the existence of widely divergent life history strategies in freshwater mussels.     

The role of fecundity and reproductive effort in defining life‐history strategies of North American freshwater mussels

Selection is expected to optimize reproductive investment resulting in characteristic trade‐offs among traits such as brood size, offspring size, somatic maintenance, and lifespan; relative patterns of energy allocation to these functions are important in defining life‐history strategies. Freshwater mussels are a diverse and imperiled component of aquatic ecosystems, but little is known about their life‐history strategies, particularly patterns of fecundity and reproductive effort. Because mussels have an unusual life cycle in which larvae (glochidia) are obligate parasites on fishes, differences in host relationships are expected to influence patterns of reproductive output among species. I investigated fecundity and reproductive effort (RE) and their relationships to other life‐history traits for a taxonomically broad cross section of North American mussel diversity. Annual fecundity of North American mussel species spans nearly four orders of magnitude, ranging from < 2000 to 10 million, but most species have considerably lower fecundity than previous generalizations, which portrayed the group as having uniformly high fecundity (e.g. > 200000). Estimates of RE also were highly variable, ranging among species from 0.06 to 25.4%. Median fecundity and RE differed among phylogenetic groups, but patterns for these two traits differed in several ways. For example, the tribe Anodontini had relatively low median fecundity but had the highest RE of any group. Within and among species, body size was a strong predictor of fecundity and explained a high percentage of variation in fecundity among species. Fecundity showed little relationship to other life‐history traits including glochidial size, lifespan, brooding strategies, or host strategies. The only apparent trade‐off evident among these traits was the extraordinarily high fecundity of Leptodea, Margaritifera, and Truncilla, which may come at a cost of greatly reduced glochidial size; there was no relationship between fecundity and glochidial size for the remaining 61 species in the dataset. In contrast to fecundity, RE showed evidence of a strong trade‐off with lifespan, which was negatively related to RE. The raw number of glochidia produced may be determined primarily by physical and energetic constraints rather than selection for optimal output based on differences in host strategies or other traits. By integrating traits such as body size, glochidial size, and fecundity, RE appears more useful in defining mussel life‐history strategies. Combined with trade‐offs between other traits such as growth, lifespan, and age at maturity, differences in RE among species depict a broad continuum of divergent strategies ranging from strongly r‐selected species (e.g. tribe Anodontini and some Lampsilini) to K‐selected species (e.g. tribes Pleurobemini and Quadrulini; family Margaritiferidae). Future studies of reproductive effort in an environmental and life‐history context will be useful for understanding the explosive radiation of this group of animals in North America and will aid in the development of effective conservation strategies.     

Patterns and drivers of intraspecific variation in avian life history along elevational gradients: a meta‐analysis

Elevational gradients provide powerful natural systems for testing hypotheses regarding the role of environmental variation in the evolution of life‐history strategies. Case studies have revealed shifts towards slower life histories in organisms living at high elevations yet no synthetic analyses exist of elevational variation in life‐history traits for major vertebrate clades. We examined (i) how life‐history traits change with elevation in paired populations of bird species worldwide, and (ii) which biotic and abiotic factors drive elevational shifts in life history. Using three analytical methods, we found that fecundity declined at higher elevations due to smaller clutches and fewer reproductive attempts per year. By contrast, elevational differences in traits associated with parental investment or survival varied among studies. High‐elevation populations had shorter and later breeding seasons, but longer developmental periods implying that temporal constraints contribute to reduced fecundity. Analyses of clutch size data, the trait for which we had the largest number of population comparisons, indicated no evidence that phylogenetic history constrained species‐level plasticity in trait variation associated with elevational gradients. The magnitude of elevational shifts in life‐history traits were largely unrelated to geographic (altitude, latitude), intrinsic (body mass, migratory status), or habitat covariates. Meta‐population structure, methodological issues associated with estimating survival, or processes shaping range boundaries could potentially explain the nature of elevational shifts in life‐history traits evident in this data set. We identify a new risk factor for montane populations in changing climates: low fecundity will result in lower reproductive potential to recover from perturbations, especially as fewer than half of the species experienced higher survival at higher elevations.     

Phenotypic Variations in the Life History of Two Clones of Macrobiotus richtersi (Eutardigrada, Macrobiotidae)

A comparative study of life history traits of two clones (CDMr01 and CDMr02) of a triploid thelytokous apomictic population of the eutardigrade Macrobiotus richtersi has been carried out. Both clones were reared under the same lab conditions: temperature of 14 °C, photoperiod of 12 h/12 h (L/D), and nematodes ad libitum as food. Statistical analysis of the life history traits considered has indicated interclonal variability. The two clones were significantly different in the number of eggs per clutch (fertility), number of eggs laid per female per life span (fecundity), hatching percentage of eggs and hatching time. Similarities between clones have been observed with regard to life span, total number of ovipositions per life span, and age at first oviposition. In addition, significant differences in hatching time, with eggs hatched over a long period, were found within each clone. Interclonal variability in life history traits indicated the effects of genetic factors, whereas intraclonal variability reflected the effects of environmental factors. The evolutionary and adaptive significance of the life history phenotypic variations is discussed.     

Biofilms as food for decapods (Atyidae,Palaemonidae) in the River Murray,South Australia

Stable water levels and turbidity associated with flow regulation in the River Murray have promoted the growth of filamentous green algae and Cyanobacteria in biofilms on submerged wood. We investigated the assimilation of biofilm algae by two dominant consumers, the decapod crustaceans Macrobrachium australiense (Palaemonidae) and Paratya australiensis (Atyidae), in two river reaches differing in the extent of floodplain development, hence wetland connectivity. Filamentous Cyanobacteria, a major part of the biofilms assimilated in combination with other foods, were up to 83% of the algal component of the gut content volume of P. australiensis and 44% that of M. australiense. Cyanobacteria have not previously been reported as a major source of nutrition for adult decapods. There was little difference between the stable isotopic signatures (¹³C/¹²C, ¹⁵N/¹⁴N) of the two decapod species, or between decapods in the two reaches. Coarse and fine particulate organic matter from the gorge had similar isotopic signatures to those from upstream and so were likely derived from macrophyte detritus rather than local willows. Red gum leaves and wood were too depleted in both ¹³C and ¹⁵N to register in the diets of either decapod in gorge or floodplain reaches. The most likely food sources for the decapods are littoral plants in the gorge reach and fine particulate organic matter material processed upstream. This is consistent with current hypotheses of organic matter flux in large river systems.     

Effects of variation in resource acquisition during different stages of the life cycle on life‐history traits and trade‐offs in a burying beetle

Individual variation in resource acquisition should have consequences for life‐history traits and trade‐offs between them because such variation determines how many resources can be allocated to different life‐history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life‐history traits and trade‐offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade‐off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life‐history traits but limited effects on trade‐offs. This suggests that in species that acquire a fixed‐sized resource at the onset of breeding, the size of this resource has larger effects on life‐history trade‐offs than resources acquired at earlier stages.     

Size specific demography of three species of Anastrepha fruit flies

Individuals of three Anastrepha species: A. obliqua, A. ludens, and A. serpentina (Diptera: Tephritidae), were sorted according to pupal weight in cohorts of large and small flies. Demographic parameters and reproductive patterns and heterogeneity were determined for each cohort. Large flies of the three species presented greater expectation of life and gross fecundity rates. A. ludens was the species with the longest life span (expectation of life of large adults was 110 days) and the greatest gross fecundity rates (1597 eggs/female for the large flies). While, A. obliqua had the shortest mean age of reproduction (33 days), and the greatest daily egg production (14 eggs/female/day). Net fecundity was similar in these two. A. serpentina had lower fecundity rates.Reproductive information for each size and each species include: age-by-parity relations, fraction of sexually mature life in which females lay eggs, and frequency distribution of individual egg production. Results demonstrate that even under constant laboratory conditions and using standard artificial hosts, there is a great deal of life history variation among these Anastrepha species and among other tephritid fruit flies.     

Life history influences the vulnerability of New Zealand galaxiids to invasive salmonids

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Latitudinal gradients and the shaping of life‐history traits in a gregarious caterpillar

The present study aimed to investigate how the impact of several factors linked to geography would shape life‐history traits in a gregarious species, using the pine processionary moth (PPM) Thaumetopoea pityocampa as a model system. PPM has a wide geographical distribution over the Mediterranean Basin, and it is a strictly gregarious species throughout larval development, where the total reproductive output of each female forms a colony. We reviewed both published and unpublished data on PPM from all over its distribution in the Mediterranean Basin and extracted data on fecundity, egg size, egg parasitoid mortality, flight period, and development time. These life‐history traits were then related to location, expressed as latitude and altitude, local average temperatures, and host tree species. We found that PPM fecundity increaseed with latitude, concomitant with an increase in the length of development and an earlier onset of adult flight. These results are the opposite of that found in other Lepidoptera species with a wide geographical distribution, as well as in insects in general. We propose that a large colony size in PPM is important at higher latitudes because this confers an advantage for thermoregulation and tent building in areas where larvae have to face harsher conditions during the winter, thus shifting the optimal trade‐off between the number and size of eggs with latitude. However, host tree species also affected the relationship between egg number and size and the optimal outcome of these traits is likely a compromise between different selection pressures. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 224–236.     

Do pioneers have r-selected traits? Life history patterns among colonizing terrestrial gastropods

We examine whether pioneer species of terrestrial gastropods (snails and slugs) possess particular life history traits commonly associated with r-selection, using data on gastropod colonization in four areas in north-west Europe (the Kvarken and Tvärminne archipelagos in the Baltic, polder woods in IJsselmeer, and a rehabilitated quarry near Maastricht). Data on age at first reproduction, longevity, clutch size, egg size and lifetime fecundity were gathered from the literature. In order to control for potentially confounding effects of body size on life history traits, we compared the residuals from the allometric relations between life history traits and body size for pioneers and non-pioneers. In snails, all life history traits examined were related to body size. In slugs, all traits except age at first reproduction scaled with body size. Body sizes did not differ between pioneers and non-pioneers in any area. In all four areas, there were no significant differences between pioneers and non-pioneers in any of the life history traits examined, after body size had been taken into account. This indicates that pioneer terrestrial gastropods generally cannot be regarded as r-selected. Pioneer species may possess any of several life history strategies, and the combinations of traits shown by them may have little in common with the r-K selection concept.     

Landscape‐scale life‐history gradients in New Zealand freshwater fish

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Comparing and contrasting life history variation in four aphid hyperparasitoids

1. In primary parasitoids, significant differences in life history and reproductive traits are observed among parasitoids attacking different stages of the same host species. Much less is known about hyperparasitoids, which attack different stages of primary parasitoids. 2. Parasitoids exploit hosts in two different ways. Koinobionts attack hosts that continue feeding and growing during parasitism, whereas idiobionts paralyse hosts before oviposition or attack non‐growing host stages, e.g. eggs or pupae. 3. Koino‐/idiobiosis in primary parasitoids are often associated with different expression of life history trade‐offs, e.g. endo‐ versus ectoparasitism, high versus low fecundity and short versus long life span. 4. In the present study, life history parameters of two koinobiont endoparasitic species (Alloxysta victrix; Syrphophagus aphidivorus), and two idiobiont ectoparasitic species (Asaphes suspensus; Dendrocerus carpenteri) of aphid hyperparasitoids were compared. These hyperparasitoids attack either the parasitoid larva in the aphid before it is killed and mummified by the primary parasitoid or the parasitoid prepupa or pupa in the dead aphid mummy. 5. There was considerable variation in reproductive success and longevity in the four species. The idiobiont A. suspensus produced the most progeny by far and had the longest lifespan. In contrast, the koinobiont A. victrix had the lowest fecundity. Other developments and life history parameters in the different species were variable. 6. The present results reveal that there was significant overlap in life history and reproductive traits among hyperparasitoid koinobionts and idiobionts, even when attacking the same host species, suggesting that selection for expression of these traits is largely association specific.     

Reproductive tactics of four sympatric sticklebacks (Gasterosteidae)

Synopsis The life history traits (longevity, growth, age at maturity, fecundity, egg size and length of breeding season) were compared for four sympatric sticklebacks (Gasterosteidae). Significant differences occurred for growth, longevity, age at maturity and length of breeding season, but not for egg diameters. Differences in life history traits are interpreted in terms of mechanisms permitting the coexistence of these closely-related territorial fish. Egg numbers were lowest inApeltes quadracus and highest inGasterosteus aculeatus withGasterosteus wheatlandi andPungitius pungitius having similar and intermediate fecundities.     

Reproductive bionomics and life history traits of three gammaridean amphipods, Cymadusa filosa Savigny, Ampithoe laxipodus Appadoo and Myers and Mallacoota schellenbergi Ledoyer from the tropical Indian Ocean (Mauritius)

The reproductive bionomics and life history traits of two corophiid amphipods (Ampithoe laxipodus, Cymadusa filosa) and one melitid (Mallacoota schellenbergi) were studied in Mauritius (Indian Ocean) for the period March 1999 to February 2000. Results on the population structure, monthly size class variations, sex ratio, female reproductive states and fecundity are presented. The study demonstrates multivoltinism and continuous reproduction in the three species. Increase in number of juveniles was observed in warmer months for C. filosa and A. laxipodus. Sexual maturity was attained at smaller sizes in warmer months in the three species. Linear relationship on body length and number of eggs in brood pouch are presented. Size-independent analysis of egg number revealed a decrease in number of eggs in cooler months. Sex ratio is male skewed in M. schellenbergi and female skewed in C. filosa and A. laxipodus. Some of the plausible explanations for the reproductive strategies adopted by these three species in a tropical system are discussed.     

Inter- and intraspecific differences in the life history and growth of Caridina spp. (Decapoda: Atyidae) in Hong Kong streams

1. The life‐history pattern of three populations of Caridina cantonensis (Atyidae) and one population of Caridina serrata were studied in four Hong Kong streams: Pak Ngau Shek (PNS), Kap Man Hang (KMH), Pak Tam Chung (PTC), and Lung Fu Shan (LFS). Caridina cantonensis occurred at PNS, KMH and PTC while C. serrata was present at LFS only. Monthly quantitative sampling was carried out at each study site over 2 years to investigate life‐history patterns and cohort‐specific growth rates. 2. Breeding by both species was mainly restricted to the wet season, and there was an increase in the incidence of ovigerous females as water temperatures rose at the start of the summer monsoon. The number of cohorts and the frequency of recruitment of C. cantonensis (one to three times each year) were highly site‐specific, but consistent between years. 3. Shrimp growth was size‐dependent, and decreased with increasing body size. Caridina cantonensis at PTC had the highest cohort‐specific growth rates (0.05–0.14 mg ash‐free dry weight (AFDW) mg^?1 day^?1), followed by C. serrata at LFS (6% lower), and C. cantonensis at KMH and PNS (16–39% lower). Shrimp life spans exceeded 1 year (17–22 months), depending on site and species. 4. Sexual maturity occurred at an earlier age and at a smaller size in populations with higher growth rates. Shrimps at PTC and LFS matured at approximately 4 months old; this was 3 months earlier than at KMH and 5 months earlier than at PNS. Females of C. cantonensis at PTC and C. serrata at LFS may have bred twice during each breeding season, while females in the other two populations bred once only. The number of cohorts produced by each population in each year did not vary between the years of the study. Most sexually mature individuals only survived long enough (for 10–12 months after maturation) to breed during one season. 5. The mean brood size of C. cantonensis varied among streams and was 25–28% larger at PNS and KMH than at PTC. Egg size did not differ among populations. Larger brood sizes at PNS and KMH may have been a consequence of increased availability of algal food in these unshaded stream sites. Caridina serrata had similar egg and brood sizes to C. cantonensis, but females were smaller and reproductive investment was higher. Thus, the degree of intraspecific variation in atyid breeding in Hong Kong was at least equal to, if not greater than, the extent of interspecific variation. Interspecific differences in life history may reflect the frequency of droughts and spate intensity experienced by C. serrata at LFS.     

The interaction effect between intraspecific competition and seed quality on the life‐history traits of the seed‐feeding beetle Acanthoscelides macrophthalmus

It has been shown that intraspecific competition and resource quality may affect life‐history traits of insects, such as body size, fecundity, and survival. However, intraspecific competition and resource quality may interact with each other. The study of such interacting effects is crucial for understanding the influence of these ecological variables on the selection of specific life‐history traits. Here, we investigated whether the interaction between intraspecific larval competition and variation in resource quality affects adult emergence and survival, egg size, fecundity, body size, and sexual size dimorphism (SSD) of the seed‐feeding beetle Acanthoscelides macrophthalmus (Schaeffer) (Coleoptera: Chrysomelidae: Bruchinae) when infesting Leucaena leucocephala (Lam.) De Wit (Fabaceae), its host plant. In the laboratory, beetles were reared on seeds that differed in quality (e.g., different hardness, seed size, water content), in the presence or absence of larval competition. Body size and SSD did not differ between treatments (with and without competition), nor were they affected by varying resource quality. Females subjected to competition during the larval stage and females emerging from seeds of higher quality, displayed the highest fecundity. The proportion of emergent adults was higher in the absence of competition. In addition, larger eggs were laid on the low‐quality resource in the absence of competition, showing a trade‐off between egg size and egg number. Adult survival differed among treatments and resource qualities, suggesting a higher investment in adult survival for individuals emerging from seeds of low quality in the presence of competition. Whether changes in specific traits could be selected for in detriment of others will depend on the strength of intraspecific competition, the variation in resource quality, and the plasticity in the life‐history traits investigated. This needs further clarification.     

A comparative analysis of population structuring and genetic diversity in sympatric lineages of freshwater shrimp (Atyidae: Paratya): concerted or independent responses to hydrographic factors?

1. We determined whether two sympatric mitochondrial DNA (mtDNA) lineages of freshwater shrimp (Decapoda: Atyidae: Paratya australiensis) represent biological species and if they had concerted or independent population responses to hydrographic factors in small streams (the Granite Creeks) in southeastern Australia. 2. Allozyme data indicated the presence of two gene pools at sites where the P. australiensis lineages were co‐occurring and the gene pools were statistically assigned with high probability to each respective lineage. This indicated that these mtDNA lineages in P. australiensis were reproductively isolated and thus biological species. 3. Populations of both lineages were genetically homogeneous among lowland sites within streams, but were isolated by steep stream gradients among upland sites and for lowland–upland site comparisons. However, the magnitude of differentiation was markedly different between the two lineages. Allozyme diversity also differed between the two lineages, suggesting that they have different effective population sizes. Thus, differences in the magnitude of genetic divergence among populations were probably because of different life‐history characteristics, including dispersal ability and population size. 4. Genetic population structure was mostly temporally stable, despite the extreme effects of drought during the first year and substantial stream‐flow during the second. However, stable isotope analyses revealed greater local movement in both lineages during the second year, as greater hydrological connectivity provided more opportunities for dispersal. Thus, although lowland populations within streams were genetically homogeneous, stable isotope data indicated that connections may be sporadic and result from accumulated small‐scale movements among refugial pools. 5. Both lineages were therefore found to have similar small‐scale population responses to the unstable habitats of the Granite Creeks. Results highlight the importance of refugia for the capacity of biota to recover from drought and the need for multiple restored patches to reinstate natural population processes (e.g. resilience, recolonization) in degraded systems.