Calling behavior of mass-reared and wild Anastrepha serpentina (Diptera: Tephritidae)

The calling behavior of mass-reared and wild males of Anastrepha serpentina (Wiedemann) (Diptera: Tephritidae) was studied both in the laboratory and in field cage tests. In the laboratory, density (1, 5, and 10 males per container), age, and hour of day significantly affected calling behavior. Mass-reared males called independently of density, whereas wild males only called at densities of 5 and 10 individuals. Males of both strains started calling when they were 5-7 d old. The daily pattern of male calling was similar in both strains, starting at 0730 hours, and reaching a peak at 1330-1630 hours. Field cage tests showed that mass-reared males started calling when they were 5d old; the period of peak calling was when males were 8-9 d old. In contrast, wild males began calling when they were 10 d old, reaching peaks when males were 13, 15, and 18 d old. Wild males tended to form leks to call during each day of the experiment, whereas mass-reared males only formed leks during 2 d, both strains displaying very low levels. During field cage tests, males, independently of strain, displayed two calling peaks, one peak in the morning and one peak in the afternoon, whereas males observed in the laboratory only showed a single calling peak. The results are discussed in view of the effects of mass rearing A. serpentina males in relation to potential use of the sterile insect technique.     

Lek Site Selection by Male Mediterranean Fruit Flies

The mating system of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann), is based on leks. In field cage choice experiments we examined lek site selection as influenced by the presence of fruit and the presence of conspecific flies. Males were allowed to choose between artificial trees containing fruit and trees without fruit. Males showed a significant preference for the tree baited with fruit. To determine which fruit-related stimuli were important, males were allowed to choose between visual and/or olfactory fruit-derived stimuli. Males preferred to lek on trees presenting both stimuli. To determine the influence of conspecifics on lek site selection, males were allowed to choose between trees containing male and/or female conspecific flies. The presence of pheromone-emitting males was preferred. In addition, both in the field and on artificial trees, we examined the effect of leaf size on lek site selection. In the field, males preferred to lek on leaves of a particular size. Moreover, leaf integrity was important, as males tended to select undamaged leaves as lek sites. In field cage experiments, males consistently chose to lek on trees containing the largest leaves. We conclude that the choice of lek site is influenced by the presence of fruit and of other lekking males, while leaf size and integrity determine male location within a lek.     

Nutritional reserves regulate male participation in Mediterranean fruit fly leks

1. The mating system of Mediterranean fruit flies ( Ceratitis capitata , Diptera: Tephritidae), is based on male leks that form on the foliage of trees. Following observations that not all males participate in leks, the hypotheses that (a) small males are absent from leks and (b) males with inadequate nutrient reserves are unable to participate in leks were examined.
2. The size and weight of lekking males ( n = 183) and resting males captured at the same time in the vicinity of leks ( n = 148) were established. In addition, using biochemical techniques, the amounts of sugar, glycogen, lipid and protein in each individual were established quantitatively.
3. There was no significant size difference between lekking and resting males. However, lekking males were significantly heavier and contained significantly more sugars and protein than resting males.
4. In conclusion, leks are exclusive, and only males with adequate nutritional reserves may join. Thus, in this species, reproductive success is closely linked to foraging success.     

Male signalling and lek attractiveness in the Mediterranean fruit fly

I tested the null hypothesis that females of the Mediterranean fruit fly, Ceratitis capitata, were equally likely to visit leks composed of males with high versus low mating success. In laboratory trials, the observed distribution of matings among males differed significantly from that expected by chance, owing primarily to the higher than expected numbers of individuals with low (mated 0-1 days over 6 consecutive observation days) or high (mated 4 or more days) mating scores. I termed these two groups as 'low' and 'high' maters, respectively. In the field, greater numbers of female sightings were made at artificial leks of high maters than low maters. This result apparently reflected a greater calling propensity among high maters. Slopes of female sightings versus calling level did not differ significantly between leks of low and high maters, suggesting that the observed relationship between calling activity and female sightings was independent of male mating status. Following the same protocol, I conducted a second experiment to examine whether males used the signals of conspecific males to locate lek sites and, if so, whether signal attractiveness varied with male mating ability. Attraction of males to calling conspecifics was far weaker than that observed for females, and over five different trials a total of only seven male sightings were made at any of the established leks. Copyright 2000 The Association for the Study of Animal Behaviour.     

Impact of solar radiation in disinfecting drinking water contaminated with Giardia duodenalis and Entamoeba histolytica/dispar at a point-of-use water treatment

Aims:  To determine the impact of natural sunlight in disinfecting water contaminated with cysts of Giardia duodenalis and Entamoeba histolytica/dispar using plastic containers.
Methods and Results:  Known quantities of Giardia duodenalis and Entamoeba histolytica/dispar cysts in sterile water were exposed to the sun. Containers were made of polyethylene terephthalate, eight painted black on one side, one not painted and another cut open at the top and the last was a high density polypropylene container. Viability testing was performed using vital and fluorescent dyes. The same assays were conducted under cloudy conditions. Thermal control tests were also performed using heat without ultra violet light from the sun. Results show that 99·9% of parasites was inactivated when water temperatures reached 56°C after sunlight exposure.
Conclusion:  Both solar radiation and heat produced by the sun have a synergistic effect in killing cysts of Giardia duodenalis and Entamoeba histolytica/dispar when temperatures rise above 50°C, with complete death at 56°C, using painted 2-l PET containers.
Significance and Impact of the Study:  Solar disinfection system using PET containers painted black on one side can be used to disinfect water against Giardia duodenalis and Entamoeba histolytica/dispar using natural sunlight.     

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.     

Measurement of three-dimensional structure of plants with a simple device and estimation of light capture of individual leaves

1. A simple device called a 'pocometer' (POlar COordinate METER) was developed to measure three-dimensional structure of plants. It consists of a tape-measure to measure distance and two protractors to measure zenith angle and azimuth angle.
2. The pocometer can determine locations of points within a few metres distance with a resolution of less than 1cm. Location of any point on a plant can be measured in 10 to 30s depending on the ease of pulling the tape measure to the point of interest.
3. A system to use data obtained with the pocometer to calculate plant light capture was developed. The degree of shading at any point on a plant is estimated by checking obstruction by other plant parts of the view toward the sky at that point.
4. Photon flux density (PFD) on leaf surfaces was estimated for Aucuba japonica , a broad-leaved evergreen shrub, using the above system. The estimated PFDs for individual leaves of a plant corresponded to the sensor-measured PFDs with correlation coefficients of 0·67 to 0·92.     

Heat resistance of Cronobacter species (Enterobacter sakazakii) in milk and special feeding formula

Aim:  To determine D - and z -values of Cronobacter species ( Enterobacter sakazakii ) in different reconstituted milk and special feeding formula and the effect of reconstitution of powdered milk and special feeding formula with hot water on the survival of the micro-organism.
Methods and Results:  Five Cronobacter species (four C. sakazakii isolates and C. muytjensii ) were heated in reconstituted milk or feeding formula pre-equilibrated at 52–58°C for various times or mixed with powdered milk or feeding formula prior to reconstitution with water at 60–100°C. The D -values of Cronobacter at 52–58°C were significantly higher in whole milk (22·10–0·68 min) than in low fat (15·87–0·62 min) or skim milk (15·30–0·51 min) and significantly higher in lactose-free formula (19·57–0·66 min) than in soy protein formula (17·22–0·63 min). The z -values of Cronobacter in reconstituted milk or feeding formula ranged from 4·01°C to 4·39°C. Water heated to ≥70°C and added to powdered milk and formula resulted in a > 4 log₁₀ reduction of Cronobacter .
Conclusions:  The heat resistance of Cronobacter should not allow the survival of the pathogen during normal pasteurization treatment. The use of hot water (≥70°C) during reconstitution appears to be an effective means to reduce the risk of Cronobacter in these products.
Significance and Impact of the Study:  This study supports existing data available to regulatory agencies and milk producers that recommended heat treatments are sufficient to substantially reduce risk from Cronobacter which may be present in these products.     

Antimicrobial effect and shelf-life extension by combined thermal and pulsed electric field treatment of milk

Aims:  The impact of a combined hurdle treatment of heat and pulsed electric fields (PEF) was studied on native microbiota used for the inoculation of low-fat ultra-high temperature (UHT) milk and whole raw milk. Microbiological shelf-life of the latter following hurdle treatment or thermal pasteurization was also investigated.
Methods and Results:  UHT milk was preheated to 30°C, 40°C or 50°C over a 60-s period, pulsed for 50  μ s or 60  μ s at a field strength of 40 kV cm⁻¹ or for 33  μ s at 50 kV cm⁻¹. Heat and PEF reduced the microbial count by a maximum of 6·4 log in UHT milk (50°C; 50 kV cm⁻¹, 33  μ s) compared to 6·0 log ( P  ≥ 0·05) obtained by thermal pasteurization (26 s, 72°C). When raw milk was treated with a combination of hurdles (50°C; 40 kV cm⁻¹, 60  μ s) a 6·0 log inactivation of microbiota was achieved and microbiological milk shelf-life was extended to 21 days under refrigeration (4°C) vs 14 days in thermally pasteurized milk. Native microbiota was decreased by 6·7 log following conventional pasteurization.
Conclusions:  The findings suggest that heat and PEF achieved similar inactivation of native microbiota in milk and longer stabilization of microbiological shelf-life than thermal pasteurization.
Significance and Impact of the Study:  A hurdle approach of heat and PEF could represent a valid milk processing alternative to conventional pasteurization. Hurdle treatment might also preserve native milk quality better due to less thermal exposure.     

Resistance of Escherichia coli grown at different temperatures to various environmental stresses

Aims:  To study the influence of growth temperature on the resistance of Escherichia coli to three agents of different nature: heat, pulsed electric field (PEF) and hydrogen peroxide.
Methods and Results:  Escherichia coli cells were grown to stationary phase at 10°C, 20°C, 30°C, 37°C and 42°C. Survival curves to a heat treatment at 57·5°C, to a PEF treatment at 22 kV cm⁻¹ and to 40 mmol l⁻¹ hydrogen peroxide were obtained and fitted to a model based on the Weibull distribution to describe and compare the inactivation. Time to inactivate the first log cycle of the population at 57·5°C of cells grown at 42°C was sixfold higher than that corresponding to cells grown at 10°C. On the contrary, cells grown at 10°C and 20°C were more resistant to PEF and hydrogen peroxide treatments.
Conclusions:  The influence of growth temperature on bacterial resistance depends on the stress applied. Cells grown at higher temperatures were more heat resistant, but more sensitive to PEF and hydrogen peroxide.
Significance and Impact of the Study:  Results obtained in this investigation help in understanding the physiology of bacterial resistance and the inactivation mechanisms of different technologies.     

Calling behaviour of the potato aphid Macrosiphum euphorbiae oviparae under laboratory and field conditions

Abstract.  1. The calling behaviour of virgin oviparae of the potato aphid Macrosiphum euphorbiae (Thomas) (Homoptera: Aphididae) was studied at three different constant temperatures under laboratory conditions. The mean age of calling for the first time decreased with a decrease in temperature from 2.9 days at 20 °C to 2.1 days at 10 °C. At all temperature regimes, the mean onset time of calling advanced from about 5 to 3 h after the onset of the photophase, and the mean time spent calling increased by > 4 h over the 8 days.
2. Cohorts of oviparae were also observed at two different periods in late summer–early autumn in the field, to examine the effects of fluctuating abiotic conditions (temperatures, wind velocity, rain) and age on calling behaviour. As under constant laboratory conditions, the mean age of calling for the first time declined with declining temperature, from 3.7 days in early September to 1.6 days at the end of September. Age-related changes in the mean onset time of calling and the mean time spent calling were much less evident under field conditions, due to the inhibitory effects of low temperatures, high winds, and rain on female calling activity.
3. The results are discussed within the context of reproductive success and address a previously proposed hypothesis suggesting that species-specific calling windows may serve as a reproductive isolating mechanism for sympatric aphid species.     

The impact of nitrogen fertilization of tomato on feeding site selection and oviposition by Trialeurodes vaporariorum

The sexual calling incidence of wild and mass-reared Mediterranean fruit fly males, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), was studied at 25 °C and a photophase of 14 h starting at 0600 h. Wild males fed on yeast hydrolysate and sugar attained sexual maturity earlier and exhibited higher calling activity than males fed on sugar alone. In both sugar and yeast hydrolysate-fed wild males, calling levels were high during the first hours of the photophase, declined at 0900–1000 h, then increased again reaching high levels between 1200 and 1500 h, and finally declined and ceased after 1600 h. Food deprivation for 8 h or more caused a significantly lower frequency of calling. Irradiated (sterile) and unirradiated mass-reared males fed on either sugar and yeast or sugar alone did not differ in the age of attaining sexual maturity (maximum levels of calling occurred about 8 days earlier than in the wild males). However, irradiated males fed on sugar called less frequently than males in the other treatments of mass-reared males. The daily rhythm of calling in protein and sugar-fed irradiated males differed somewhat from that of wild males. Calling was recorded at high levels from the beginning of the photophase until 1300 h without any decrease at 0900–1000 h. It then decreased to very low levels from 1500–1800 h and increased slightly at 1900 h. These findings are discussed in relation to the mating behavior of the fly and to the effectiveness of released sterile males in Sterile Insect Technique programs.     

Modelling the growth of Weissella cibaria as a function of fermentation conditions

Aims:  To investigate the effect of pH, water activity ( a _w) and temperature on the growth of Weissella cibaria DBPZ1006, a lactic acid bacterium isolated from sourdoughs.
Methods and Results:  The kinetics of growth of W. cibaria DBPZ1006 was investigated during batch fermentations as a function of pH (4·0–8·0), a _w (0·935–0·994) and temperature (10–45°C) in a rich medium. The growth curve parameters (lag time, growth rate and asymptote) were estimated using the dynamic model of Baranyi and Roberts (1994. A dynamic approach to predicting bacterial growth in food. Int J Food Microbiol 23, 277–294). The effect of pH, a _w and temperature on maximum specific growth rate (μ_max) were estimated by fitting a cardinal model. μ_max under optimal conditions (pH = 6·6, a _w = 0·994, T  = 36·3°C) was estimated to be 0·93 h⁻¹. Minimum and maximum estimated pH and temperature for growth were 3·6 and 8·15, and 9·0°C and 47·8°C, respectively, while minimum a _w was 0·918 (equivalent to 12·2% w/v NaCl).
Conclusions:  Weissella cibaria DBPZ1006 is a fast-growing heterofermentative strain, which could be used in a mixed starter culture for making bread.
Significance and Impact of the Study:  This is the first study reporting the modelling of the growth of W. cibaria , a species that is increasingly being used as a starter in sourdough and vegetable fermentations.     

Effect of temperature and water activity on in vitro germination of Monilinia spp.

Aims:  This study evaluated the effect of temperature (0–38°C) and water activity ( a _w: 0·87–0·99) on the lag phase prior to germination and the percentage of germination over time for Monilinia laxa , Monilinia fructicola and Monilinia fructigena .
Methods and Results:  More than 80% of viable conidia germinated at 25°C and 0·99 a _w within 2 h for M. fructicola and M. fructigena and 4 h for M. laxa . There was no germination at 38°C, and all three Monilinia spp. germinated at 0°C. At the lowest a _w (0·87), none of the Monilinia spp. was able to germinate at any of the incubation temperatures studied. Whereas at 0·90 a _w, conidia were only able to germinate at 15, 25 and 30°C for the three species studied, except for M. fructicola at 15°C. In contrast, at 0·95, 0·97 and 0·99 a _w, germination occurred at all studied temperatures less 38°C. Generally, the lag phase was longer at low levels of a _w (0·90–095), and differences were more evident as temperatures were far from the optimum (0–5°C).
Conclusions:  Germination and lag phase period were markedly influenced by temperature and a _w, and in general when conditions of temperature and a _w were suboptimal, the lag phase was longer and the percentage of germination was lower.
Significance and Impact of the Study:  Knowledge of the germination requirements of this fungus is important in order to understand their behaviour in natural situations and to provide baseline data required for the construction of new prediction models. Our study might be used to develop a predictive model to understand and control the disease caused by Monilinia spp.     

Interactions of Salmonella enterica with lettuce leaves

Aims:  To investigate the interactions of Salmonella enterica with abiotic and plant surfaces and their effect on the tolerance of the pathogen to various stressors.
Methods and Results:  Salmonella strains were tested for their ability to form biofilm in various growth media using a polystyrene plate model. Strong biofilm producers were found to attach better to intact Romaine lettuce leaf tissue compared to weak producers. Confocal microscopy and viable count studies revealed preferential attachment of Salmonella to cut-regions of the leaf after 2 h at 25°C, but not for 18 h at 4°C. Storage of intact lettuce pieces contaminated with Salmonella for 9 days at 4°C resulted only in small changes in population size. Exposure of lettuce-associated Salmonella cells to acidic conditions (pH 3·0) revealed increased tolerance of the attached vs planktonic bacteria.
Conclusions:  Biofilm formation on polystyrene may provide a suitable model to predict the initial interaction of Salmonella with cut Romaine lettuce leaves. Association of the pathogen with lettuce leaves facilitates its persistence during storage and enhances its acid tolerance.
Significance and Impact of the Study:  Understanding the interactions between foodborne pathogens and lettuce might be useful in developing new approaches to prevent fresh produce-associated outbreaks.     

Strategies of mate finding in the European field cricket (Gryllus campestris) at different population densities: a field study

Abstract. 1. Members of a field population of Gryllus campestris L. varied in their walking and calling activity. In both sexes, some individuals occupied burrows whereas others walked around in the observation area. Males at burrows could be either silent or calling.
2. In the course of one summer, population density decreased and the initial balanced sex ratio changed to a large surplus of males.
3. At high population density, there were equal numbers of non-calling males at burrows, calling males at burrows and walking males, while walking females predominated over females at burrows. Non-calling males at burrows achieved more encounters with females than did calling and walking males. Females met males by walking through the population and by waiting at burrows. Thus, calling and phonotaxis were not essential for mate finding and calling was less effective than previously thought.
4. At low population density calling males predominated. Calling males at burrows achieved the most encounters with females. Females met males only by walking around in the population area. Calling was thus more important in mate finding than at high population density.
5. Changes in sex ratio and population density may cause the flexibility in mate finding behaviour of individual crickets.     

Sexual behavior and male volatile compounds in wild and mass‐reared strains of the Mexican fruit fly Anastrepha ludens (Diptera: Tephritidae) held under different colony management regimes

We compared the calling and mating behavior and volatile release of wild males Anastrepha ludens (Loew) with males from 4 mass‐reared strains: (i) a standard mass‐reared colony (control), (ii) a genetic sexing strain (Tap‐7), (iii) a colony started from males selected on their survival and mating competitiveness abilities (selected), and (iv) a hybrid colony started by crossing wild males with control females. Selected and wild males were more competitive, achieving more matings under field cage conditions. Mass‐reared strains showed higher percentages of pheromone calling males under field conditions except for Tap‐7 males, which showed the highest percentages of pheromone calling males under laboratory cage conditions. For mature males of all strains, field‐cage calling behavior increased during the last hour before sunset, with almost a 2 fold increase exhibited by wild males during the last half hour. The highest peak mating activity of the 4 mass‐reared strains occurred 30 min earlier than for wild males. By means of solid phase microextraction (SPME) plus gas chromatography‐mass spectrometry (GC‐MS), the composition of volatiles released by males was analyzed and quantified. Wild males emitted significantly less amounts of (E,E)‐α‐farnesene but emitted significantly more amounts of (E,E)‐suspensolide as they aged than mass‐reared males. Within the 4 mass‐reared strains, Tap‐7 released significantly more amounts of (E,E)‐α‐farnesene and hybrid more of (E,E)‐suspensolide. Differences in chemical composition could be explained by the intrinsic characteristics of the strains and the colony management regimes. Characterization of calling behavior and age changes of volatile composition between wild and mass‐reared strains could explain the differences in mating competitiveness and may be useful for optimizing the sterile insect technique in A. ludens.     

Elevated atmospheric [CO2] promotes frost damage in evergreen tree seedlings

Growth under elevated [CO₂] promoted spring frost damage in field grown seedlings of snow gum ( Eucalyptus pauciflora Sieb. ex Spreng.), one of the most frost tolerant of eucalypts. Freezing began in the leaf midvein, consistent with it being a major site of frost damage under field conditions. The average ice nucleation temperature was higher in leaves grown under elevated [CO₂] (– 5·7 °C versus – 4·3 °C), consistent with the greater incidence of frost damage in these leaves (34% versus 68% of leaves damaged). These results have major implications for agriculture, forestry and vegetation dynamics, as an increase in frost susceptibility may reduce potential gains in productivity from CO₂ fertilization and may affect predictions of vegetation change based on increasing temperature.     

Influence of pregnancy on the thermal biology of the lizard, Sceloporus jarrovi: why do pregnant females exhibit low body temperatures?

1. Selected body temperatures of female lizards, Sceloporus jarrovi , were measured on a photothermal gradient during late pregnancy and again when postpartum, and pregnant females were subjected to one of three fluctuating temperature regimes that simulated body temperatures of (1) pregnant females, (2) postpartum females or (3) allowed normal thermoregulation.
2. Overall, females selected lower body temperatures when pregnant (mean = 32·0°C) than when postpartum (mean = 33·5°C).
3. Females regulated body temperature more precisely when pregnant than when postpartum as judged by their smaller variances in body temperature throughout the day.
4. When pregnant, females selected a lower mean maximum body temperature (mean: pregnant = 32·8°C; postpartum = 34·5°C) than when postpartum, but selected mean minimum body temperatures did not differ.
5. None of the experimental temperature treatments was detrimental to pregnant females. Female body length increased during pregnancy but the rate of increase did not differ among treatments. Moreover, length-adjusted body mass of postpartum females did not differ among treatments.
6. Pregnant females that experienced postpartum body temperatures produced neonates that were smaller in body mass and length than pregnant females that experienced pregnant body temperatures and females that were allowed to thermoregulate.
7. For neonates resulting from the postpartum body temperature treatment, the disparity in the body length, but not mass, was still observed at 9 days of age, although survival and growth of neonates was high and did not differ among treatments.
8. The results demonstrate that pregnant females could maintain higher postpartum body temperatures without compromising their physical condition, but select relatively low body temperatures, presumably to avoid decrements in offspring fitness.     

Influence of food quality and temperature on life history characteristics of the parthenogenetic mayfly, Cloeon triangulifer

SUMMARY. 1. Laboratory and field data indicate that Cloeon triangulifer McDunnough has at least three generations per year in White Clay Creek (Pennsylvania, U.S.A.).
2. The duration of the egg stage ranged from 5 days at 30°C to about 90 days at 10°C.
3. Larvae completed development (i.e. first instar to adult) in 27 days at 25°C, 45 days at 20°C, and 179 days at 10°C on an algal diet dominated by diatoms.
4. Larvae reared on hickory leaves completed development in 30 days at 25°C but died prior to metamorphosis at 10, 15 and 20°C.
5. Adult size (i.e. body length, wing length and dry mass) and fecundity were inversely related to rearing temperature for all laboratory and field experiments.
6. The significant interaction of food quality and temperature suggest that these factors may be important in understanding geographic variation in the life history of C. triangulifer.