The interactive effects of climate, life history, and interspecific neighbours on mortality in a population of seed harvester ants

Abstract.  1. Fifteen years of census data from a population of the red harvester ant ( Pogonomyrmex barbatus ), 5 years of census data from populations of Aphaenogaster cockerelli and Myrmecocystus mimicus at the same site, and long-term precipitation data from a nearby weather station were used to examine how nearest interspecific neighbours, climate, and life history influence P. barbatus colony mortality.
2. It was found that more P. barbatus colonies die following dry summers than following wet summers, but there is no relationship between the establishment of new colonies and rainfall.
3.  Aphaenogaster cockerelli , but not M. mimicus , nearest neighbours influence P. barbatus colony mortality in some years and for some ages.
4. The effects of nearest A. cockerelli neighbours are usually most severe following dry summers, when resource availability is reduced.     

Density-dependent population dynamics in clonal organisms: a modelling approach

1. Density dependence may act at several stages in an organisms life-cycle (e.g. on mortality, fecundity, etc.), but not all density-dependent processes necessarily regulate population size. In this paper I use a density manipulation experiment to determine the effects of density on the transition rates between different size classes of the clonal zoanthid Palythoa caesia Dana 1846. I then formulate a density-dependent matrix model of population dynamics of Palythoa , and perform a series of sensitivity analyses on the model to determine at what stage in the life-cycle regulation acts.
2. Seven of the 16 transition probabilities decreased with density, most of them being shrinkage (due to loss of tissue or fission) and stasis (the self–self transition) of medium and large colonies. The only probability to increase was for the stasis of large colonies. Recruitment was quadratically dependent on density, peaking at intermediate densities.
3. Equilibrium cover in the model was 84% and was reached in ≈40 years. To determine which density-dependent transitions were involved in population regulation, the strength of density dependence was varied in each independently. This sensitivity analysis showed that only changes in the probabilities of large colonies remaining large and producing medium colonies, were regulating.
4. These results suggest that regulation is primarily acting on fission of large colonies to produce intermediate-sized colonies, in combination with size specific growth rates. Fission rates decrease greatly with density, resulting in a greater proportion of large colonies at high densities and large colonies grow more slowly than small. Overall, this behaviour is very similar to that of clonal plants which have a phalanx type life history.     

Queen lifespan and colony longevity in the ant Harpegnathos saltator

Abstract.  1. The longevity of field colonies was investigated in the ponerine ant Harpegnathos saltator (Jerdon) in which either reproductive workers (gamergates) or a single queen reproduce.
2. Data from 3 years were used to calculate the ratio between queen-right ( n  = 50) and gamergate ( n  = 12) colonies that can be used to derive the colony mortality of gamergate colonies. Using the survival rates of queens in the laboratory, extrinsic and intrinsic mortality rates of queen-right colonies were calculated.
3. No significant differences in the sizes of queen-right and gamergate colonies above 14 workers was found, suggesting that mortality of established colonies is not size related.
4. The mortality of gamergate colonies is 4.17 times higher than the intrinsic mortality of queen-right colonies.
5. In the laboratory, mean survival time of queens in colonies of more than 14 workers was 1.79 years.
6. Estimated mean survival time of queen-right and gamergate colonies in the field varies between 0.78 and 0.43 years respectively, when no costs of conflict during the replacement of queens occur; however, when the latter costs increase colony mortality to a level similar to extrinsic mortality, the calculated longevity of queen-right colonies would increase to 1.02 years.     

The effects of simulated rainfall on cochineal insects (Homoptera: Dactylopiidae): colony composition and survival on cactus cladodes

ABSTRACT. 1. A rainfall simulator is described that was used to investigate the effects of rainfall on the cochineal insect Dactylopius opuntiae (Cockerell). This species occurs in discrete colonies that are protected by a copious covering of opaque'woolly'wax, and is the most important biological control agent of the prickly pear cactus, Opuntia ficus-indica (L.) Miller, in South Africa.
2. We record the composition of cochineal colonies (stage and number of individuals) in relation to colony size to allow accurate predictions of colony composition from size measurements of the intact, wax-covered colonies before their exposure to rainfall.
3. Even short exposures (15 min) to simulated rainfall (delivered at a rate of about 50 mm/h) resulted in wax erosion, and the effect was most marked in small, recently established colonies. Coverings of compacted wax protected the mature colonies from erosion.
4. The number of colonies that were washed off the plant was a function of colony size and the duration of exposure to simulated rainfall. Most of the small, recently established colonies were washed from the plants after 120 min. All of the large, mature colonies, albeit badly mutilated and containing far fewer individuals, remained on the plants even after 240 min exposure.
5. Survival of individuals, in colonies of all sizes, was less than about 40% after short bouts of rainfall (30 min). First- and second-instar individuals accounted for most of the observed mortalities, and mortality of all stages increased with increasing rainfall duration.
6. These observations provide a partial explanation for the poor performance of D. opuntiae as a biological control agent of cacti in higher-rainfall areas.     

Recruitment in the harvester ant, Pogonomyrmex occidentalis: effects of experimental removal

We examined the importance of experimental removal of mature colonies on colony recruitment in the western harvester ant Pogonomyrmex occidentalis. To test the common assumption that established colonies suppress the establishment of new colonies we removed all colonies in ten 0.25 ha plots in 1996 and an additional five plots in 1997 and measured new colony recruitment in 1997, 1998, and 1999. Using a blocked, paired plot design we found that removal of colonies increased new colony recruitment in some areas of the site, but not others. Spatial variation in the importance of established colonies to recruitment was consistent across years; blocks in which density dependence was important in one year exhibited density dependent recruitment in following years. We estimated that in the blocks where recruitment was affected by established colonies, they accounted for less than 10% of the mortality of foundress queens. The increase in the number of new recruits (on average two additional new colonies) was considerably less than the number of colonies removed; average colony density in the removal plots was 14 colonies per 0.25 ha plot. The consistent lack of importance of established colonies to recruitment in one block and the relatively small response to colony removal in the other blocks suggests that the number of new colonies in a year may not be equivalent to the number of deaths of established colonies in that year. Space limitation is an important influence on recruitment in P. occidentalis, but the magnitude of the limitation varies spatially.     

Oviposition site preference and larval mortality in a leaf-mining moth

Abstract. 1. The univoltine leaf-mining moth, Lithocolletis quercus Ams., is endemic to Israel, where it spends its 10.5 month larval period feeding only in the leaves of Quercus calliprinos Webb.
2. We compared patterns of egg deposition and sources of larval mortality to test whether oviposition patterns and site preferences confer an enhanced likelihood of larval survival.
3. Dominant sources of larval mortality were premature leaf abscission and death from unknown causes, whereas predation, parasitism and intraspecific interference accounted for relatively little larval mortality.
4. Eggs, and thus mines, were aggregated among leaves of host trees even though premature leaf abscission was positively correlated with density of mines per leaf. Interference competition among larvae was the only other density-dependent mortality factor.
5. Oviposition patterns within leaves mitigated the probability of death from larval interference, and probably also from early leaf abscission.
6. Despite these density-dependent mortality factors, overall probability of larval survival to pupation was independent of initial density of mines on a leaf.
7. The long larval period allows synchrony between oviposition flights and times of predictable resource availability.     

The life cycle and growth of Sialis lutaria L. (Megaloptera) in a drainage channel under different methods of plant management

Abstract. 1. The life cycle of Sialis lutaria in two experimental reaches, one treated with the herbicides dalapon and 2,4-D (Chemical) and the other subjected to a conventional method of plant control (Hand), of a drainage channel in Essex took about 2 years.
2. A maximum density of 815 m^-2 was recorded and the density of each year class in both reaches generally decreased exponentially with age.
3. At certain times of the year the mean individual weight of each year class increased exponentially with age. Annual production was estimated to be 985 (Hand) and 1171 (Chemical) mg dry weight m^-2, with ratios of annual production to mean standing crop (P: ) of 1.6 and 1.8 respectively.
4. There was no evidence that the use of herbicides had any discernible effect on the mortality, growth or production of S.lutaria.     

Population studies of the beech leaf mining weevil (Rhynchaenus fagi) in Ireland and Scotland

Abstract. 1. Population budgets for Rhynchaenus fagi, based for the most part on samples of beech leaves taken in June, were constructed for a 10-year period (Northern Ireland) and a 3-year period (Scottish borders).
2. There were years of sustained population growth and sustained decline, but not enough evidence to suggest a cycle. Similar population levels were found at both sites in all but one year.
3. Key factors were associated with the adult stage, although egg mortality may also make a significant contribution to population change.
4. Unknown sources of mortality embodied in the residual mortality were also suspected of acting in a density dependent manner, compensating for variation in parasitism.
5. Death rate of larvae within mines was related to May temperature in Northern Ireland, but not so clearly in Scotland.     

Competition, survivorship and growth in macrophyte communities

SUMMARY. 1. Transplant experiments tested the proposition that the intensity of competition within macrophyte communities varies with standing crop in a freshwater marsh in Ontario, Canada.
2. Transplants of three species ( Carex lasiocarpa, Juncus brachycephalus, Scripus validus ) were grown at sixty-six locations chosen to represent a range of standing crop values. At each location, transplants were grown in plots with all neighbours removed and in adjacent plots with all neighbours present.
3. The effect of neighbours on transplant growth did not vary with standing crop, soil, or water depth, with the exception of one species which was most suppressed by neighbours in shallow water. In contrast, the effect of neighbours on the survivorship of transplants was greatest on organic soils supporting high values of standing crop in shallow water, and least on mineral soils supporting low values of standing crop in deeper water.
4. The results indicated that establishment and survivorship of vegetative propagules was most likely to be affected by neighbours in shallow water, but that the growth of established plants was not influenced by neighbours anywhere. Growth results corroborate earlier findings that competition has little effect on aquatic macrophytes.     

Demography of Bouteloua gracilis in a mixed prairie: analysis of genets and individuals

1 We analysed a data set containing the locations and sizes of individual plants from five 1-m² permanent quadrats that had been mapped annually from 1934 to 1972. We digitized each map and used geographical information software to evaluate annual recruitment and mortality, age structure, survivorship and longevity of genets and individuals of Bouteloua gracilis .
2 Ninety-one per cent of the years between 1938 and 1972 had at least one seedling recruited. An average of 10.3 genets died annually, although the variability among years was high (CV = 70%). The number of seedlings recruited each year was significantly correlated with the number of genets dying. Most genets died before age 10 and the average life span was 3.7 years, although some genets were much longer lived (> 38 years).
3 The average number of individuals per genet was 1.3. Excluding seedlings, 56% of the genets were represented by a single individual. The annual number of individuals in the 5 m² ranged from 29 to 278. Average mortality of individuals was 25.8 per year and the variability among years was high (CV = 91%).
4 The plant community appeared to be relatively stable and openings created by mortality were recolonized by recruitment. Climatic variability did not have a large influence on demography.     

Nourishment affects colony demographics in the paper wasp Polistes metricus

Abstract  1. Colony survivorship and numbers of nest cells, pupae, and adult females were monitored throughout the nesting season for a cohort of 78 colonies of the paper wasp Polistes metricus Say. Thirty-nine colonies received a twice-weekly nourishment supplement of honey during pre-emergence and early emergence phases of the colony cycle; 39 colonies were unsupplemented controls.
2. Colony survivorship was unaffected by the supplemental nourishment. Loss of colonies to predation differed among three sites but was unaffected by supplementation.
3. Honey-supplemented colonies constructed more nest cells than did control colonies but this effect was not expressed until after supplementation had ceased.
4. Honey-supplemented colonies produced more pupae than did control colonies but the number of adult females at nests did not differ between supplemented and control colonies. Because honey-supplemented colonies had more offspring but fewer of them remained as workers at the nest, honey supplementation led to a lower frequency of workers and corresponding higher frequency of reproductives than in control colonies.
5. In a second year of study, colony survivorship and numbers of nest cells, pupae, and adult females were monitored from late pre-emergence until the end of the nesting season for a cohort of 32 colonies of Polistes metricus . In 16 colonies, trophallactic saliva was taken from final-instar larvae on nine dates in the late pre-emergence and early emergence periods; 16 colonies served as controls.
6. Saliva-diminished colonies had lower survivorship, fewer nest cells, fewer pupae, and fewer adult females at the nest than did control colonies.
7. These results show that variation in nourishment in the early to mid phases of the colony cycle can have significant effects on the subsequent colony demographics of Polistes metricus paper wasps.     

Seasonal spatial dynamics and causes of nest movement in colonies of the invasive Argentine ant (Linepithema humile)

Abstract.  1. Colony organisation and movement behaviour of the Argentine ant ( Linepithema humile ) was studied over 3 years in field populations in California and in captive colonies in the laboratory. This invasive species is highly polydomous and unicolonial; colonies consist of expansive and fluid networks of nests and trails. The spatial and temporal organisation of colonies may contribute to ecological dominance.
2. Argentine ant nests and inter-nest trails shift in size, abundance, and location, so that colony networks are spatially contracted in the winter and expanded spring to autumn. Colonies occupy permanent sites; ants migrated to and from the same winter nest locations year after year, and occupied 30% of the same nests repeatedly during seasonal migrations.
3. Nests were moved on average 2–3 m. Forty-two per cent were occupied less than 1 month, 4% the entire study, and the other 54% lasted 3.9 ± 2.3 months (mean ± SD).
4. Nests were located within 2–4 m of woody plants, in warm sites in the winter and cool sites in the summer. Both humidity and food availability influenced nest-site choice in laboratory colonies. However, when faced with a trade-off between factors, the ants chose humid nest boxes over nest boxes near food, and ants moved nests only in response to changes in humidity and not distance to food.
5. The results indicate that L. humile colonies are seasonally polydomous, and that nest movements are driven by changes in microclimate. Colony organisation maintains high local density and increases food supply, which may improve the competitive ability of L. humile colonies and reduce opportunities for species coexistence.     

Intraspecific variation in size and density of Avocet colonies: effects of nest-distances on hatching and breeding success

In many colonial bird species there is considerable intraspecific variation in colony size and inter‐nest distance (colony density). Possible causes of this variation and its effects on hatching success (survival of eggs) and breeding success (probability of a pair raising chicks) were studied in 48 Avocet Recurvirostra avosetta colonies in Schleswig‐Holstein (Germany) between 1991 and 1996. Colony density was influenced by time of year and habitat (categories: island or mainland, close to or far from feeding grounds). Colonies on islands had the highest densities. When all available space at a colony site was used, colonies became very dense (mean nearest‐neighbour nest distance less than 1 m). Colony size (number of clutches) was influenced by time of year, but not by habitat. Hatching success was low in high density colonies and in very low density ‘colonies’ (single nests) and high over a broad range of intermediate nest densities. The low success rate of single nests was caused by a very high predation rate, whereas the low success rate in very dense colonies was caused by a high rate of nest abandonment. Nest abandonment in very dense colonies was associated with a high level of aggressiveness among Avocets during the egg‐laying period. Due to territorial behaviour, Avocets seemed to be expelled from the densest breeding sites. In very dense colonies, high frequencies of clutches of unusual size occurred due to conspecific nest parasitism. The number of Avocets taking part in attacks on potential egg predators was small and (in colonies of more than one clutch) depended neither on colony size nor on colony density. Despite a low hatching success in very dense colonies, individuals breeding in the densest colonies had significantly better chances of raising chicks than Avocets breeding in less dense colonies. Coloniality seemed to be obligatory for Avocets in order to ensure hatching success. The size and density of colonies seemed to be associated with the availability of suitable nesting habitats (islands).     

Effects of reproductive timing and colony size on the survival, offspring colony size and drone production in the honey bee (Apis mellifera)

ABSTRACT. 1. The effects of colony size and time of reproduction on the survival and size of offspring colonies and on drone production were examined for honey bees, Apis mellifera L. Drone and worker production and survival of parental and offspring colonies were monitored following swarming. Also, the temporal patterns of drone emergence and availability of unmated queens were examined.
2. Colony size at swarming was positively correlated with the number of workers invested in offspring colonies and the number of queens produced. However, colony size at swarming was not correlated with the number of offspring colonies produced.
3. Swarm size was positively correlated with drone and worker production after swarms were hived. Worker production of hived swarms was positively correlated with colony survival. Offspring queens which inherited a parental nest survived longer than queens in either primary swarms or afterswarms, presumably due to the advantage of inheriting a nest.
4. Drone emergence peaked just prior to swarming, the time when unmated queens were available. High drone production by colonies initiated by swarms probably reflected an attempt to reproduce prior to winter. The probabilities of a second swarming cycle within the same year and of surviving the winter were low for colonies initiated from swarms.     

Comparative study on the consequences of culling badgers (Meles meles) on biometrics, population dynamics and movement

1.  Capture–mark–recapture data were used to describe the process of recovery from a typical badger removal operation (BRO) at North Nibley, Gloucestershire, UK, which was carried out as part of the government's strategy to control bovine tuberculosis. Data on biometrics, demographics and movement from this low-density disturbed population were compared with those of two nearby high-density undisturbed populations (Wytham Woods and Woodchester Park, UK) in order to study fundamental principles of population dynamics and density-dependence.
2.  Badgers moved more between social groups at North Nibley than in the other study areas, particularly in the immediate aftermath of the removal operation.
3.  Recolonization of the vacated habitat occurred in the first instance by young females.
4.  Although in the first year after the BRO no cubs had been reared in any of the culled groups, and although the shortage of sexually mature boars may have limited the reproductive output of sows in the following year, the population took only 3 years to recover to its (already lowered) preremoval density.
5.  Losses from the adult (and cub) population due to mortality or emigration were smaller at North Nibley than at the other sites.
6.  There was much evidence that during 1995 and 1996 density-dependent effects constrained the reproductive output of the high-density populations, and some support for the hypothesis that badgers exhibit the non-linear 'large mammal' type of functional response to density.
7.  Badgers at North Nibley were younger, heavier and in better condition than badgers at Wytham Woods and Woodchester Park.
8.  We argue that the disease dynamics are likely to be different in disturbed compared with undisturbed badger populations, and that this could affect the effectiveness of BROs.     

Effects of resource availability and social parasite invasion on field colonies of Bombus terrestris

Abstract.  1. The survival, growth and fecundity of bumblebee colonies are affected by the availability of food resources and presence of natural enemies. Social parasites (cuckoo bumblebees and other bumblebees) can invade colonies and reduce or halt successful reproduction; however, little is known about the frequency of invasion or what environmental factors determine their success in the field.
2. We used 48 experimental colonies of the bumblebee Bombus terrestris , and manipulated both resource availability at the landscape scale and date of colony founding, to explore invasion rates of social parasites and their effect on the performance of host colonies.
3. Proximity to abundant forage resources (fields of flowering oilseed rape) and early colony founding significantly increased the probability of parasite invasion and thus offset the potential positive effects of these factors on bumblebee colony performance.
4. The study concludes that optimal colony location may be among intermediate levels of resources and supports schemes designed to increase the heterogeneity of forage resources for bumblebees across agricultural landscapes.     

Abundance and survival of a seed-infesting weevil, Pseudanthonomus hamamelidis (Coleoptera: Curculionidae), on its variable-fruiting host plant, witch-hazel (Hamamelis virginiana)

Abstract. 1. The relative importance of seed abundance and of mortality factors in the population biology of Pseudanthonomus hamamelidis , a hostspecific predispersal seed predator on Hamamelis virginiana , is evaluated by analysis of weevil life tables and abundance estimates in three years with markedly different magnitudes of fruit production.
2. Fruit production of marked Hamamelis individuals averaged 336 m^-2 in 1978, increased three- to four-fold in 1979, and declined again in 1980. More individuals fruited in 1979 than in the other two years.
3. The early larval stage of P.hamamelidis incurred the greatest mortality, with 42–58% dying from often undeterminable causes. Losses in the egg stage ranged from 22% to 31%. Three parasitoid species each accounted for minor mortality. Generation survival was similar in all three years, ranging from 21% to 28%.
4. The percentage of Hamamelis fruit crops infested by P.hamamelidis was high in 1978, averaging 76%, but was much lower in 1979, the year of abundant fruit. In 1980, when fruit were less abundant again, infestation was nearly complete. Egg densities on fruits were lower in 1979 than in the other two years.
5. The fluctuating fruit crops of Hamamelis are responsible for limitation of P.hamamelidis numbers during poor fruiting years. The occasional highly productive fruiting year results in satiation of weevil populations and concomitant seed escape by Hamamelis . Mortality factors appear less important in influencing weevil population dynamics.     

Late deaths and survival after childhood cancer: implications for cure.

OBJECTIVES--To investigate causes of death and survival in subjects who had survived at least five years after diagnosis of childhood cancer; to compare observed mortality with that expected in the general population; and to compare results with a corresponding cohort diagnosed earlier. DESIGN--Retrospective cohort study. SETTING--Population based National Register of Childhood Tumours. SUBJECTS--9080 five year survivors of childhood cancer diagnosed in Britain during 1971-85, of whom 793 had died. Comparison with corresponding cohort diagnosed during 1940-70. MAIN OUTCOME MEASURES--Cause of death established from all available sources of information (including hospital and general practitioner records and postmortem reports) and underlying cause of death coded on death certificate. RESULTS--Of the 781 deaths for which sufficient information was available, death was attributed to recurrent tumour in 578 (74%) cases, treatment related effect in 121 (15%), second primary tumour in 52 (7%), and other causes in 30 (4%). Comparison of observed mortality with that expected in the general population indicated a fourfold excess of deaths from non-neoplastic causes. The risk of dying of recurrent tumour in the next 10 years after surviving five years from diagnosis during 1940-70 and 1971-85 fell from 12% to 8%. The risk of dying from a treatment related effect increased slightly from 1% to 2%. CONCLUSION--Improvements in five year survival after childhood cancer have been accompanied by a reduction in risk of dying from recurrent tumour during the subsequent 10 years and by a slight increase in risk of dying from treatment related effects. The results provide information relevant to decisions concerning balance between effective treatments and their potentially harmful effects.     

Climate, social factors and research disturbance influence population dynamics in a declining sociable weaver metapopulation

Population trends are determined by gains through reproduction and immigration, and losses through mortality and emigration. These demographic quantities and resulting population dynamics are affected by different external and internal drivers. We examined how these demographic quantities were affected by weather, research-induced disturbance, local density, colony site and year in a metapopulation of 17 sociable weaver (Philetairus socius) colonies over 17 years of study (4 years for reproduction). Most colonies declined, but at different rates. The four demographic quantities were related to different drivers. Survival strongly varied among years and colonies and was positively related to rainfall and negatively related to extreme temperature (together explaining 30 % of variation) and disturbance (measured as number of captures conducted at a colony; 7 %). There was a trend for a positive relationship between reproduction and rainfall (50 %). Movement was mainly related to local density: individuals were more likely to emigrate from small to large colonies and from colonies that were either well below or above their long-term mean. They were more likely to immigrate into colonies that were nearby, and below their mean size. We then quantified the effects of these relationships on metapopulation dynamics using a multi-site matrix projection model. Rainfall was potentially a strong driver of metapopulation dynamics. In addition, field-work disturbance might have contributed to the decline of this metapopulation but could not explain its full magnitude. Hence, through a combination of analytical methods we were able to obtain information on the main drivers affecting dynamics in a declining metapopulation.     

Long-term numerical changes and regulation in a river stock of European eel Anguilla anguilla

1. The objective was to explore the regulatory mechanisms underlying the numerical dynamics of a river stock of European eel Anguilla anguilla based on a purely observational study. We used a 21-year data set on the Rio Esva (Asturias, northwestern Spain) stock of eels, encompassing eel density at an estuary site and at nine sites distributed among three tributaries selected to differ in the distance from the tidal limit and fishery data on glass eels from the nearby Rio Nalón estuary.
2. The abundance of glass eels and estuarine and in-stream densities declined simultaneously over a 15-year period. However, following a historical minimum in the abundance of glass eels around the year 2000, the density continued to decline in the estuary whereas density in the streams began an increase that has continued to the present.
3. In-stream cohort mortality rate ( Z ) varied from 0.0016 day⁻¹ to only 0.0003 day⁻¹ (on an annual basis from 0.57 to 0.099 year⁻¹). The negative effects of intra-cohort and inter-cohort densities on mortality rate emphasized the responses of the riverine stock to compensatory density-dependence. As a consequence, the number of eels surviving to commence the silvering stage was similar in magnitude in the strongest and weakest cohorts.
4. These results suggest that, even if the low abundance of glass eels persists, in-stream density may increase towards a stable value regulated by in-stream density-dependence.