text:
Present address: Wageningen IMARES, PO Box 68, 1970 AB Ijmuiden, The Netherlands
Invasion success depends on invader body size in a size-structured mixed predation–competition community
Article first published online: 22 JUL 2009
DOI: 10.1111/j.1365-2656.2009.01590.x
© 2009 The Authors. Journal compilation © 2009 British Ecological Society
Journal of Animal Ecology
Volume 78, Issue 6, pages 1152–1162, November 2009
Additional Information
How to Cite
Schröder, A., Nilsson, K. A., Persson, L., Van Kooten, T. and Reichstein, B. (2009), Invasion success depends on invader body size in a size-structured mixed predation–competition community. Journal of Animal Ecology, 78: 1152–1162. doi: 10.1111/j.1365-2656.2009.01590.x
Publication History
- Issue published online: 13 OCT 2009
- Article first published online: 22 JUL 2009
- Received 26 September 2008; accepted 17 June 2009 Handling Editor: Philip Warren
- Abstract
- Article
- References
- Cited By
- coexistence;
- competitive recruitment bottleneck;
- life-history omnivory;
- ontogenetic niche shift;
- productivity gradient
1. The size of an individual is an important determinant of its trophic position and the type of interactions it engages in with other heterospecific and conspecific individuals. Consequently an individual's ecological role in a community changes with its body size over ontogeny, leading to that trophic interactions between individuals are a size-dependent and ontogenetically variable mixture of competition and predation.
2. Because differently sized individuals thus experience different biotic environments, invasion success may be determined by the body size of the invaders. Invasion outcome may also depend on the productivity of the system as productivity influences the biotic environment.
3. In a laboratory experiment with two poeciliid fishes the body size of the invading individuals and the daily amount of food supplied were manipulated.
4. Large invaders established persistent populations and drove the resident population to extinction in 10 out of 12 cases, while small invaders failed in 10 out of 12 trials. Stable coexistence was virtually absent. Invasion outcome was independent of productivity.
5. Further analyses suggest that small invaders experienced a competitive recruitment bottleneck imposed on them by the resident population. In contrast, large invaders preyed on the juveniles of the resident population. This predation allowed the large invaders to establish successfully by decreasing the resident population densities and thus breaking the bottleneck.
6. The results strongly suggest that the size distribution of invaders affects their ability to invade, an implication so far neglected in life-history omnivory systems. The findings are further in agreement with predictions of life-history omnivory theory, that size-structured interactions demote coexistence along a productivity gradient.
The majority of animal populations are characterized by size-structure, where individuals differ in body size and grow throughout their life cycle (Peters 1983; Calder 1984; Werner 1988). As ecologically important traits such as attack rate, metabolic costs and vulnerability to predation scale with body size (Peters 1983; Werner 1994; Persson et al. 1998; Kooijman 2000), growing individuals often experience different predation risks (Paine 1976; Chase 2003) or feed on different resources (Polis 1984; Olson 1996; Cisneros & Rosenheim 1997) over the course of their life (Werner & Gilliam 1984; Ebenman & Persson 1988). Such ontogenetic niche shifts lead to that individuals occupy different trophic positions at different phases of their life cycle and engage in ontogenetically variable mixed interactions (Neill 1975, 1988;Polis 1984; Werner & Gilliam 1984, Wilbur 1988; Persson & Greenberg 1990; Olson, Mittelbach & Osenberg 1995; Byström, Persson & Wahlström 1998; Cameron et al. 2007). In case the interactions change from competition to predation (life-history omnivory; Pimm & Rice 1987), competitive recruitment bottlenecks can be imposed on juvenile predators by their competitively superior, later prey (Werner & Gilliam 1984; Neill 1988; Persson & Greenberg 1990; Olson et al. 1995; Byström et al. 1998; Walters & Kitchell 2001). On the other hand, large predators, when present, diminish prey densities and may thus release their own offspring from competition (Neill 1988; Wilbur 1988; Walters & Kitchell 2001).
The dynamics of trophic configurations with mixed predation–competition interactions has mainly been investigated within the theoretical framework of intraguild predation (IGP; Polis, Myers & Holt 1989; Holt & Polis 1997). This research has led to a number of important insights regarding the dynamics of communities with omnivory, including (i) the necessary condition of competitive superiority of the IG-prey for its persistence, (ii) the exclusion of the IG-prey at high productivity when competition effects are mitigated and predation effects are relatively strong, (iii) the possibility of alternative stable states contrasting in number and identity of species, (iv) the overall smaller scope for coexistence of both species compared with tri-trophic food chains and (v) the pattern of community shifts along gradients of productivity (Holt & Polis 1997; Morin 1999; Diehl & Feißel 2000, 2001; Mylius et al. 2001).
The implications of size-structure, recruitment bottlenecks and ontogenetic niche shifts for the dynamics of growing predator and prey have been discussed already in early literature on mixed predation–competition systems (Werner & Gilliam 1984; Ebenman & Persson 1988; Polis et al. 1989). In addition, numerous short-term studies demonstrate size-dependent mixed interactions (Rice, Crowder & Rose 1993; Crumrine 2005; Griffin & Byers 2006; Rudolf & Armstrong 2008). Still, these features have only rarely been incorporated into IGP theory (Pimm & Rice 1987; Mylius et al. 2001; Van de Wolfshaar, de Roos & Persson 2006). While these studies have corroborated several of the findings of unstructured IGP theory, they have also led to new insights. Size-structure, particularly in co-occurrence with the ubiquitous feature of food-dependent individual development, demotes coexistence of IG-prey and IG-predator and expands the parameter region of mutual exclusion along a productivity gradient compared with the unstructured IGP systems (Mylius et al. 2001; Van de Wolfshaar et al. 2006).
Notwithstanding this progress, the understanding of ontogenetically variable trophic interactions in terms of their impacts on community dynamics remains sketchy. One aspect that has been neglected so far is that the body size of invaders may matter for their invasion success. When trophic interactions and ecological roles change with body size over ontogeny, different sized invaders may face different biotic environments, either beneficial or detrimental to their ecological performance and thus for the invasion outcome. As the biotic environment is also influenced by the productivity of the system, invasion outcome may also depend on productivity.
Here it was tested how invader body size and productivity affect the invasion outcome in a life-history omnivory system using two poeciliid fish species, Heterandria formosa [Agassiz 1855] or Least Killifish, and Poecilia reticulata [Peters 1859] or Common Guppy. H. formosa is the smaller species, so it was assumed that it would function as the IG-prey and P. reticulata as the IG-predator (Fig. 1). The experiment involved the invasion of small numbers of either large or small P. reticulata into resident populations of H. formosa at different productivities. First, it was predicted that large P. reticulata would establish persistent populations with the likelihood of invasion success increasing with productivity as they would benefit from the increasing density of the IG-prey. Second, it was predicted that small invaders would fail irrespective of productivity as they would always face competition by the resident IG-prey without being able to benefit from predation.
Materials and methods
Study species
H. formosa and P. reticulata are small viviparous, sex-dimorphic poeciliid fishes native to the coastal plains in southeastern USA or the Lesser Antilles and the coastal regions of northern South America, respectively. They live in freshwater ponds and streams under a wide range of abiotic conditions (Rosen & Bailey 1963).
H. formosa juveniles are born at a size of 5–8 mm. Size at maturation is 12 and 14 mm for males and females, respectively. Males reach a maximum size of 18 mm while females can attain a size of 36 mm (Fraser & Renton 1940). Females simultaneously carry embryo litters of different developmental stages and they give birth to litters of 1–7 young in intervals of 7–21 days (Cheong et al. 1984). The generation time is about 7 weeks (Travis & Henrich 1986). H. formosa founder individuals for the stock in Umeå were wild caught in June 2003 in Lake Jackson, Florida.
P. reticulata juveniles are born at a size of 6–9 mm. Males mature at a size of 14 mm and females reach maturity at a size of 16 mm. Maximum size is 19 and 41 mm for males and females, respectively. Litters consist of 10–30 young; the brood interval is 25 days. The generation time is c. 10–20 weeks (Reznick & Miles 1989). P. reticulata founder individuals were caught wild in 2002 in the Turure River, Trinidad.
Individuals of both species can easily be distinguished from each other at any age and sex. H. formosa individuals are in general smaller and always show dark, vertical stripes on body and tail. Older individuals have partially orange-coloured back fins. P. reticulata individuals are of lighter colour; mature males have horizontal orange ornamental stripes.
Experimental set-up
The experiment was performed between May and December 2007. The set-up was a semi-closed circulation system of 32 aquaria each with a volume of 80 L plus a reservoir with a volume of 600 L in which the water was aerated, warmed and mixed. Water was exchanged by a constant inflow of 20 L h−1 tab water into the reservoir. Water turnover in each aquarium was four times each day. The water was untreated well water with a hardness of 3·8°dH and a pH of 8·4. To prevent infections with ectoparasites salt was added daily to achieve a salinity of 0·5‰ and an ultraviolet (UV) water sterilizer (Wiegandt HW 4000; Wiegandt, Krefeld, Germany) was installed. Three bio-filters (Eheim 2080; Eheim, Deizisau, Germany) and one activated carbon filter were included to facilitate bacterial denitrification, detritus breakdown and waste product removal. Water temperature was held at 25 °C. The aquaria were lit from above by 15 W neon lights with a 14 h light/10 h dark regime. No refuge was offered.
The fish were fed pelletized food (Sera Microgran). Pellet feeding was accomplished by custom-made high-precision microfeeders (9·45 mg ± 0·41 SD per feeding event) centrally controlled from a computer.
The experimental design was full-factorial with productivity (low, medium, high) and invader body size (large, small) as factors. Replication was fourfold. Productivity was manipulated by varying the semi-chemostatic food supply rate. The daily amounts of food supplied were: 37·6 mg day−1 (low), 75·3 mg day−1 (medium) and 150·6 mg day−1 (high). Invader size treatment levels were either four P. reticulata adults (large; two females and two males, mean length ±1 SE 28·3 mm ± 1·1 and 17·5 mm ± 1·4, respectively) or five P. reticulata juveniles (small; mean length ±1 SE 6·8 mm ± 0·4). These low numbers were used to relate the study to: (i) real-world scenarios where invasive species usually enter new habitats in low densities and (ii) theory where the positive (or negative) growth of an invading population at low densities is used to infer about the invasion success. Two individuals of each sex were used to create balanced sex ratios and to account for potential demographical stochasticity in potential mortality. Five juveniles were used because the sex ratio at maturation is slightly biased towards males and to account for this demographical stochasticity. Only females which were not highly pregnant as judged by the form of their belly and the visibility of embryos through the transparent skin were chosen for invasions. Treatment combinations were semi-randomly assigned to aquaria as the feeder-controlling units made it necessary to group four aquaria of the same productivity regime together. Low-, medium- and high-productivity treatments were stocked with 56 (12 ♀♀, 12 ♂♂, 32 juveniles), 112 (26 ♀♀, 26 ♂♂, 60 juveniles) and 224 (52 ♀♀, 52 ♂♂, 120 juveniles) H. formosa individuals, respectively. All individuals were chosen and assigned randomly to a replicate. The initial size-structure and density of the resident populations chosen were based on the long-term average over 1½ years of eight control populations in a previous experiment in the same aquarium system, with food amount corresponding to the intermediate productivity level used here (Schröder, Persson & de Roos 2009). P. reticulata individuals were added the following day. Invading juveniles had been allowed to get acquainted to the presence of H. formosa for the prior 24 h in small breeding cages inside the stocked aquaria. A separate experiment was performed to investigate whether (i) P. reticulata could invade and persist in the system alone and (ii) assess the background mortality and maturation time of P. reticulata. The same numbers of P. reticulata individuals as before were added to four empty aquaria without H. formosa for each invader size class. These eight aquaria received 37·6 mg of food per day.
Duration of the experiments was determined a priori. The end point was set by the maturation of the offspring of the small P. reticulata invaders. Thus, the experiments covered more than a complete life cycle of P. reticulata.
One juvenile and one male P. reticulata died during the first six days of the experiment and were replaced with new, similar sized individuals the same day as such early mortality may reflect handling stress. If during the experiment only P. reticulata males were left of the initial invader individuals, half of them (or all when it was a single male) were replaced with similar sized females from stocking tanks to account for stochastic effects in maturation outcome.
The experiment and the usage of vertebrates were approved of and followed the guidelines by the Swedish Animal Welfare Agency (DSM), applications Dnr A 95-04 and Dnr A 75-06.
Size-dependent capture rates
Different sized juveniles (victims; range 6–12 mm) of each species were exposed to different sized females (consumer; range 20–35 mm) of the other species to quantify size-dependent predatory capture rates. For each size combination trial, ten victims of the same size were placed in small plastic aquaria without refuge. Two similar sized consumers were placed in transparent plastic tubes inside the aquarium. Individuals had 24 h to get visually acquainted with the presence of consumers before the consumers were released. After another 24 h the remaining living victims were counted. Species-specific cannibalistic capture rates were obtained in similar ways.
Sampling, measurements and maintenance
Sampling intensity was initially higher because it was expected that especially invasion failures would manifest already at the beginning of the experiment. The initial sampling intervals differed slightly between the large and the small invader size treatments. The entire population of an aquarium was captured by repeated hand netting until no individual could be seen during visual inspection of the aquarium. The individuals were then sorted into species and into females, males and juveniles. These subpopulations were placed in a transparent plastic bowl filled with just enough water for the individuals to stay upright and were then photographed. The individuals were counted and measured on a computer screen using an image analysis software (Optimas; Media Cybernetics, Silver Spring, MD, USA). The individuals were measured from the tip of the snout to where the cone-shaped, pigmented body merges into the straight and transparent tail fin (a close approximation to standard length). Relative lengths were transformed into metric units by using reference objects as internal standards. During the samplings, the aquaria were cleaned by netting out debris and the aquaria walls were scraped free of algae. The feeders were cleaned approximately every fourth week, checked for proper functioning and refilled.
Calculations and statistical analyses
A necessary condition for invasion success was the production of recruits by invaders, that is, small invaders had to mature and produce offspring, while the offspring of large invaders had to mature and produce recruits. Invasion success, failure and coexistence were evaluated at the termination of the experiment by noting the presence of individuals of the two species. The invasion was counted as a success when at least one female or juvenile of P. reticulata was present. If only one or more males were left the invasion was judged as a failure. Similarly, populations of H. formosa with at least one female or juvenile present were evaluated as persistent whereas populations with only one or more males left were viewed as extinct. Coexistence was defined as the presence of at least one female or juvenile of each species. In cases where males had been replaced by females, invasion outcome was also evaluated at the termination of the experiment. To test for differences in invasion success between treatments, a Generalized Linear Model was fitted to these binary success/failure data with invader size and productivity as explanatory factors, specifying a binomial error distribution and logit link function. The significance of the factors and their interaction was examined in an analysis of deviance by using conditional F-statistics to correct for overdispersion (Crawley 2002).
To quantify the invasion success of P. reticulata, a model of the form y = a · exp (− rmax· time) with a as the initial density and rmax as the maximum population growth rate at low density was fitted to the time series of the species' total abundance. For the large invader treatment and the replicates in the absence of H. formosa, the model fitting was restricted to the first 100 days of the experiment. This allowed ignoring the precise form of the population growth over ranges of densities. As the densities of H. formosa also, in the majority of the small invader replicates, unexpectedly declined near-monotonically throughout the experiment the same procedure was used to estimate the decline rates of total H. formosa abundances and of abundances of juvenile and adult H. formosa. Similar subdivision of P. reticulata founder populations into juveniles and adults was not meaningful because populations were, at least initially, homogenous in their life-history stage and size structure.
The absolute and log-transformed (to cure non-normality and heteroscedasticity to meet test assumptions) parameter estimates for rmax were then used as the dependent variable in anova. A first analysis tested for differences in rmax of total abundances in relation to species, invader size and productivity levels. A second analysis tested for differences in rmax between H. formosa juveniles and adults, depending on the invader size and productivity. A third analysis tested for differences in rmax of the total P. reticulata and total or stage-specific H. formosa abundance between productivities in the small invader treatments. Standard diagnostic plots were checked for non-normality, heteroscedasticity and the presence of highly influential points. The residuals were reasonably well behaved. In each analysis, two highly influential data points were removed; these outliers mainly concerned data points from aquaria where population dynamics were against the treatment trend.
To yield information about the potential cannibalistic and predatory pressure the average size of vulnerable juveniles (≤12 mm) present was plotted against the maximum size of predatory adults (≥25 mm) present for each replicate and sampling during the first 100 days of the experiment when the population change was most pronounced. The size limits are based on the capture rates. This was done for each species separately to quantify potential cannibalism and for large P. reticulata vs. small H. formosa and vice versa to quantify potential interspecific predation. The maximum piscivore size was used to not lose information on extreme size combinations that may have disproportional large effects. Note that this makes the conclusion on the absence of predation/cannibalism conservative. Predation/cannibalism between size-structured consumer and victim is generally constrained by a lower victim : consumer size ratio below which the predator cannot detect the prey and an upper victim : consumer size ratio above which the consumer cannot handle the prey anymore (Rice, Crowder & Marschall 1997; Claessen, de Roos & Persson 2000). Because the juveniles of both species used here are born at a size far above the lower boundary, size combination data were compared with only the upper boundary of predation or cannibalism size windows. These boundaries were obtained from size-dependent capture rate data by regressing to the size of the largest victim successfully attacked by a consumer of a certain size over the consumer size.
To obtain information about the intensity of potential competition that small invading P. reticulata individuals were experiencing, their body growth rate at different productivities and in the controls were compared using a mixed-effect ancova with the mean growth rate over all present individuals of a given replicate of the small invader treatments and sampling as the dependent variable. As explanatory variables, a single fixed treatment factor consisting of the three productivity levels and the controls was specified and time (the first 50 days of the experiment) was used as the covariate. The replicate was added as a random factor (Crawley 2002).
Hypothesis testing was always two-sided and the α-level used was always 0·05. All computations and statistical analyses were performed in R2·7·0 (R Development Core Team 2008).
In the absence of H. formosa, P. reticulata individuals invaded all empty aquaria and established persistent populations. This was independent of the invader size. The populations were dominated by juveniles (48·3 ± 4·9 juveniles vs. 14·3 ± 1·8 adults, mean of eight replicates over the last four samplings ± 1 SE). Invading adults reproduced already during the first 18 days. The invading juveniles experienced no background mortality in these aquaria at least until their maturation (Fig. 2). rmax values were higher for the small invaders than for the large ones (Fig. 5b; Wilcoxon's W = 0, P = 0·03).
Large P. reticulata successfully invaded and established persisting populations in 10 out of 12 of their trials, while invading small P. reticulata mostly died and were successful in only 2 out of 12 of their trials (Figs 3 and 4). Productivity had no significant influence on invasion outcome (two-way analysis of deviance, invader size: F1,18 = 15·63, P < 0·001; productivity: F2,18 = 1·55, P = 0·22; invader size × productivity: F2,18 = 0·07, P = 1·0). In three out of the four cases when single males were replaced by females, these secondary invasions also failed. When invaders were successful, the resident populations usually went extinct over the course of the experiment. H. formosa populations also declined when the invaders were unsuccessful. Several of these populations showed, however ambiguous, signs of stabilization or even recovery towards the end of the experiment (Figs 3a,f,i and 4c,f,g,h). Only in three replicates, the two species coexisted at the end of the experiment. In one of these replicates, H. formosa densities were still declining while P. reticulata densities still increased (Fig. 3k), whereas in another replicate, only one P. reticulata female remained (Fig. 4a). Large invaders usually died before maturation of their offspring that later successfully reproduced themselves. Small invaders usually went extinct during the first 60–100 days of the experiment before they matured.
Total P. reticulata invader populations had positive initial population growth rates when consisting of large individuals but negative rmax when consisting of small individuals. Total H. formosa population growth rates were always negative but total H. formosa populations declined faster in the large than in the small invader treatments. Productivity had no effect on total population growth rates (Fig. 5a, three-way-anova, species: F1,34 = 8·8, P < 0·005; species × invader size: F1,34 = 9·2, P = 0·005; all other sources: F1 or 2,34 ≤ 3·1, P > 0·05). Juvenile as well as adult H. formosa subpopulations declined faster when large P. reticulata invaded than when invaded by small P. reticulata. Juvenile H. formosa declined consistently faster than adults. Productivity had no effect on these decline rates (Fig. 5a, three-way anova, stage: F1,34 = 155·9, P < 0·001; invader size: F1,34 = 47·1, P <0·001; all other sources: F1 or 2,34 ≤2·5, P > 0·05). In case of the small invader treatment, P. reticulata declined faster than either the total H. formosa populations or H. formosa juveniles or adults (Fig. 5a, two-way anova, species/stage: F3,34 = 26·9, P < 0·001; productivity: F2,34 = 4·9, P =0·013; species/stage × productivity: F6,34 = 2·3, P = 0·06), apart from the low productivity treatment where this difference showed the same trend but was not statistically significant as revealed by treatment contrasts (P > 0·05).
The potential for both predation and cannibalism during the first 100 days of the experiment was higher in the large than in the small invader treatments (Fig. 6). This was mainly because of the presence or absence of large P. reticulata individuals. H. formosa individuals were rarely large enough to prey on P. reticulata juveniles or to cannibalize their own offspring in both invader size treatments (Fig. 6).
Small invading P. reticulata grew more slowly in the presence of H. formosa than when invading the empty aquaria (mean body growth rates in mm day–1 ± CI95%: empty aquaria = 0·28 ± 0·04; high productivity = 0·10 ± 0·06; medium productivity = 0·19 ± 0·06; low productivity = 0·18 ± 0·06; mixed-effect ancova, time: F1,38 = 395, P < 0·001; treatment: F3,12 = 8·8, P = 0·002; time × treatment: F3,38 = 11·7, P < 0·001). Treatment contrasts showed that the treatment effect was because of differences in growth rate between the aquaria with and without the presence of H. formosa (P < 0·05) and not because of differences between the productivity levels (P > 0·05).
The community
The trophic configuration in the experiment was close to the simplest case of a size-structured, ontogenetically variable mixed predation–competition community (Fig. 1), similar to the life-history omnivory systems discussed by Werner & Gilliam (1984), Wilbur (1988) and Walters & Kitchell (2001) and mathematically analysed by Van de Wolfshaar et al. (2006). Large P. reticulata could prey on small H. formosa, while the reciprocal predation link with large H. formosa feeding on small P. reticulata was negligible as the corresponding size combinations were virtually absent. For similar reasons, the potential cannibalistic link within H. formosa was not realized whereas cannibalism in P. reticulata was present in the large invader treatment. Cannibalism may promote coexistence between IG-prey and IG-predator (Van de Wolfshaar 2006; Rudolf 2007) when the cannibalistic rate is stronger than the IGP rate (Rudolf 2007). However, here the reverse seems to hold, as is suggested by the (although statistically not significantly) lower intercept and slope of the upper predation and cannibalism boundaries (Fig. 6) and the higher capture rates by large P. reticulata of small H. formosa compared with small P. reticulata (data not shown). Thus, interaction links that could have increased the trophic complexity and influence the invasion outcome (cf. HilleRisLambers & Dieckmann 2003; Rudolf 2007) were at most weak. These findings led us to conclude that P. reticulata and H. formosa functioned as IG-predator and IG-prey, respectively. The higher density of P. reticulata alone at low productivity results from reporting abundances instead of biomass. P. reticulata populations were dominated by juveniles while initial H. formosa populations had a more balanced size-structure with a high fraction of adults.
Invasion outcome and size-dependent mechanisms
Body size determined the invasion success of IG-predators as large invaders had positive initial population growth rates and successfully established persistent populations whereas small invaders had negative initial population growth rates and failed in their invasions. In correspondence with this pattern, the resident populations declined more rapidly when large invaders entered the aquaria than when small invaders came in (Fig. 5a). Furthermore, a strong potential predation pressure on resident IG-prey juveniles was observed when large IG-predators invaded. The capture rates and size combinations indicate that this predation was exclusively exerted by the large invader individuals and not by large residents (Fig. 6a). These results imply that IGP on the resident IG-prey was actually taking place. In contrast, small invaders experienced a faster decline in abundance than the resident IG-prey (Fig. 5a). In addition, small invaders had positive initial population growth rates when invading empty aquaria (Fig. 5b). Together, this strongly suggests a detrimental and, more importantly, an asymmetric effect of resident IG-prey individuals on small IG-predators. A density effect where small invaders went extinct first (with the residents following) simply because of their low initial numbers is not consistent with this observed asymmetry. The detrimental effect is also indicated by the lower body growth rates of small IG-predators in the presence of IG-prey. Predation by resident IG-prey on small invaders can be ruled out as the cause behind the asymmetry in population growth rates between species in the small invader treatments (Fig. 6b).
In conclusion, we suggest that large invaders entered the residential populations in their ecological role as predators whereas small IG-predators invaded in their ecological role as inferior competitors. Small invaders suffered from a competitive recruitment bottleneck imposed by the IG-prey and consequently were out-competed before maturation and before reaching sizes that allowed them to feed on the IG-prey. In contrast, predation allowed the large invaders to successfully establish persistent populations by driving the IG-prey extinct and thereby diminishing its negative effect on the survival and growth of the IG-predator's offspring. Overall, this scenario is consistent with the presence of a juvenile competitive recruitment bottleneck and an ontogenetic niche shift in the IG-predator (Neill 1988; Wilbur 1988; Walters & Kitchell 2001; Van de Wolfshaar et al. 2006).
The mode of interspecific competition that limited the recruitment of small IG-predator is less clear. Individuals showed pronounced aggressive behaviour towards each other and the slower decline of small invaders in the low productivity replicates with their lower density of residents and thus lower frequency of interspecific encounters may speak for interference competition. On the other hand, the otherwise similar population growth rates and the similarity of body growth rates of small invaders across all productivity treatments indicate that food availability limited food intake, and hence points towards exploitation competition.
Productivity, coexistence and alternative communities
Productivity had no influence on the outcome of size-dependent invasion and stable coexistence between IG-predator and IG-prey was virtually absent across the tested range of productivities. This observed pattern of exclusion is in agreement with the results of theoretical analyses of life-history omnivory systems (Mylius et al. 2001; Van de Wolfshaar et al. 2006). In comparison with unstructured IGP systems, coexistence is demoted and the parameter space for mutual exclusion of IG-predator and IG-prey along the productivity gradient is expanded in communities with life-history omnivory. One mechanism behind this pattern is that in life-history omnivory systems, individuals only hold one trophic position at a time and change their ecological role by food-dependent transitions between life-history stages (Werner & Gilliam 1984; Pimm & Rice 1987; Mylius et al. 2001; Van de Wolfshaar et al. 2006). Van de Wolfshaar et al. (2006) showed that this separation can establish a positive feedback between size-dependent IGP, relaxed interspecific competition and faster body growth of IG-predators, thereby demoting coexistence. This process of increasing predation pressure as a result of food-dependent body growth eventually leads to the extinction of the IG-prey. In our case, the mechanism for IG-prey extinction is likely to be different as the large size of the invading IG-predator adults immediately exposed IG-prey to a high predation pressure. This led to IG-prey exclusion already within a generation and a direct, concomitant improvement of the food conditions of IG-predator offspring. Although we did not find any evidence for productivity dependence, at sufficiently low productivity levels invasion of large IG-predators is expected to fail.
In contrast to the invasion of large IG-predators, invasion failure of small IG-predators irrespective of productivity was expected. This can be explained by that interspecific competition with the resident IG-prey always limits their recruitment. Thus, small IG-predators in their ecological role as inferior competitors will never benefit from the increase in IG-prey with productivity. For this, food-dependent survival and/or body growth are essential processes because only these processes can establish the competitive recruitment bottleneck, unless mortality is caused by a food-independent process such as, for example, predation. This latter factor was, however, not likely in our study as is indicated by the absence of mutual IGP.
Despite the wide productivity gradient used in our experiment with a fourfold difference between the lowest and the highest levels, productivity had no influence on what community established. Other studies have demonstrated major shifts in community structure along similar wide gradients (Morin 1999; Diehl & Feißel 2000, 2001). This difference in results can be related to that the latter studies used protozoan communities where population size-structure is much less pronounced. In contrast, a similar independence of invasion success of IG-predators on productivity along an even wider productivity gradient has recently been observed in a size-structured mite community (Montserrat et al. 2008). The lack of coexistence of IG-predator and IG-prey in a life-history omnivory community has also recently been demonstrated in a whole-lake invasion experiment (Persson, de Roos & Byström 2007).
Whether the contrasting single-species states found here represent alternative stable communities predicted by life-history omnivory theory for intermediate productivity (Mylius et al. 2001; Van de Wolfshaar et al. 2006) is unclear. While there clearly is a priority effect of H. formosa over small P. reticulata and contingency on initial invader body size in what species establishes, the non-invasibility of the P. reticulata-only state has not been tested. However, we see it as unlikely that H. formosa could invade a resident P. reticulata population as predation by large residents would prevent any recruitment. Further, because of the density decline of the resident IG-prey populations also in most of the small invader replicates we cannot unambiguously argue for the stability of the H. formosa-only state. Other studies (Schröder et al. 2009; Schröder, Persson & Reichstein, unpublished data) in the same set-up demonstrated that H. formosa populations fluctuate in densities, rebounding after 200–250 days. These conclusions are based on populations with refuge offered but a field study shows that H. formosa can also persist when vegetation cover is low (Richardson, Gunzburger & Travis 2006). We are therefore confident that H. formosa populations in the absence of P. reticulata, despite their decline, viably persist.
General implications
The majority of animal taxa are size-structured (Peters 1983; Calder 1984;Werner 1988) and ontogenetic niche shifts or recruitment bottlenecks are frequently observed (e.g. Woodward & Hildrew 2002). Size-dependent IGP and life-history omnivory may therefore be a common phenomenon (e.g. Dick, Montgomery & Elwood 1999; Turner, Turner & Ray 2007) and hence also the dependence of invasion outcome on invader body size. Size-dependent invasion success has not been considered in earlier theoretical or experimental studies but may explain unexpected results of previous IGP experiments. For example, Montserrat et al. (2008) demonstrated consistent exclusion of IG-prey after IG-predator invasion along a productivity gradient. They did not report the size-structure of the invader populations but our results suggest that these might have been dominated by adults. Size-dependent invasion success can also explain the non-recovery of overexploited predatory fish stocks (Hutchings 2000) when after the collapse increased forage fish populations reduce the shared resource and limit predator recruitment (Walters & Kitchell 2001). Such a scenario and the associated ecosystem-wide regime shifts have also recently been proposed for the Baltic Sea cod population (Casini et al. 2009).
Overall, our results show that ontogenetically variable mixed interactions can demote coexistence and imply that the size-structure of an invading population can affect its invasion success. Further empirical and theoretical research is needed to investigate the necessary and sufficient conditions and the generality of this phenomenon. In particular, the interaction between the size distribution of the invader population and system productivity deserves further study. For example, how more intermediate-sized invaders than used here – which would need to grow in body size to undergo the ontogenetic niche shift but might be less vulnerable to competition – would fare at different productivities remains to be investigated.
The authors thank Blair Daniel for help with the experiment. Emma Göthe measured the fish and helped maintaining the set-up. Joseph Travis and David Reznick sent the fish from Florida and California, respectively. Lars Lundmark, William Larsson and Lars-Ola Westlund provided technical support for the feeders. André M. de Roos, Zlatko Petrin and three anonymous reviewers gave valuable comments on earlier drafts of the manuscript. The research was supported by grants from the Memorial of J.C. Kempe Foundation to A. Schröder and L. Persson and the Swedish Research Council to L. Persson.
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text: DontTalkNonsense August 10, 2014 10:21 AM Check out the youtube video of Jenson Button at Bathurst. A small insight of how awesome it would be.from what I understand is it's not the surface that would prevent F1 at Bathurst, it's the runoff areas and safety aspect. Reply Report abuse 0 agree 0 disagree

subset: commoncrawl


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US Ecology Announces First Quarter 2013 Results Operating Income up 49%; Adjusted EBITDA up 31% BOISE, ID -- (Marketwired) -- 04/25/13 -- US Ecology, Inc. (NASDAQ: ECOL) ("the Company") today reported financial results for the quarter ended March 31, 2013. Net income for the first quarter of 2013 was $5.4 million, or $0.29 per diluted share, up from $4.5 million, or $0.25 per diluted share, in the first quarter of 2012. Excluding foreign currency translation gains and losses and business development expenses, adjusted earnings per diluted share grew 52% to $0.32 in the first quarter of 2013, up from $0.21 in the first quarter of 2012. Operating income for the first quarter of 2013 was $9.7 million, up 49% from $6.5 million in the first quarter last year. Adjusted EBITDA for the first quarter of 2013 was $13.9 million, up 31% from $10.6 million in the same quarter last year. A reconciliation of earnings per diluted share to adjusted earnings per diluted share and net income to adjusted EBITDA is attached as Exhibit A to this release. Total revenue for the first quarter of 2013 was $42.9 million, up 30% from $33.0 million in the same quarter last year. Treatment and disposal ("T&D") revenue increased 21% quarter over quarter. Transportation revenue increased 114% quarter over quarter. Excluding US Ecology Michigan, which was acquired in May 2012, T&D revenue growth in the quarter reflected a 54% increase in project based work (Event Business) partially offset by a 1% decrease in recurring, Base Business revenue compared to the first quarter of 2012. Total quarterly waste volume disposed or processed at our Idaho, Michigan, Nevada, Texas and Quebec facilities was 223,000 tons in the first quarter of 2013, up 4% from 215,000 tons in the first quarter of 2012. Average selling price ("ASP") for the first quarter of 2013 increased 19% compared to the first quarter of 2012, reflecting a more favorable service mix. For the first quarter of 2013, gross profit was $15.4 million, up 27% from $12.1 million in the first quarter of 2012. Total gross margin was 36% in the first quarter of 2013 compared to 37% in the first quarter of 2012. T&D gross margin for the first quarter of 2013 was 42%, up from 41% in the first quarter of 2012. The year-over-year increase in T&D gross margin reflects a more favorable service mix in 2013 partially offset by the addition of US Ecology Michigan which operates at lower gross margins than our other business lines. Selling, general and administrative ("SG&A") expense for the first quarter of 2013 was $5.7 million, or 13% of revenue, compared to $5.6 million, or 17% of revenue in the same quarter last year. The Company's effective income tax rate for the first quarter of 2013 was 37.1%, down from 39.1% for the first quarter of 2012. This decrease reflects a higher proportion of earnings from our Canadian operations, which are taxed at a lower corporate tax rate, partially offset by higher U.S. state income taxes. At March 31, 2013, cash on hand was $4.5 million. Total borrowings on our lines of credit were $41.0 million, down from $45.0 million at December 31, 2012. At March 31, 2013, $47.2 million was available for future borrowings. "Strong growth in our Event Business drove waste volumes in the quarter, helping us benefit from the inherent operating leverage in the business," commented Vice President, Acting Chief Financial Officer and Chief Accounting Officer, Eric Gerratt. "We are pleased with the strong quarterly performance driven by solid execution at all of our facilities." 2013 Outlook "Continuing business momentum in the first quarter reaffirms management's confidence that US Ecology will deliver solid growth in 2013 on top of record 2012 results," commented Acting President and Chief Operating Officer, Jeff Feeler. "Growth in industrial production in North America is expected to drive increases in recurring Base Business. Our Event Business continues to be healthy with multiple contracts awarded and shipping along with a solid pipeline of opportunities to backfill completed projects. In addition, robust Gulf Coast refining markets continue to support demand for our thermal recycling services. We continue to expect full year earnings per share between $1.45 and $1.55 per diluted share excluding any foreign currency gains or losses, and project 2013 Adjusted EBITDA between $62 and $65 million." Dividend On April 1, 2013, the Company declared a quarterly dividend of $0.18 per common share for stockholders of record on April 19, 2013. The $3.3 million dividend is to be paid on April 26, 2013. Conference Call US Ecology, Inc. will hold an investor conference call on Thursday, April 25, 2013 at 10a.m. Eastern Daylight Time (8:00 a.m. Mountain Daylight Time) to discuss these results and its current financial position and business outlook. Questions will be invited after management's presentation. Interested parties can join the conference call by dialing 866-713-8563 or 617-597-5311 and using the passcode 96536035. The conference call will also be broadcast live on our website at www.usecology.com. An audio replay will be available through May 2, 2013 by calling 888-286-8010 or 617-801-6888 and using the passcode 12662606. The replay will also be accessible on our website at www.usecology.com. About US Ecology, Inc. US Ecology, Inc., through its subsidiaries, provides radioactive, hazardous, PCB and non-hazardous industrial waste management and recycling services to commercial and government entities, such as refineries and chemical production facilities, manufacturers, electric utilities, steel mills, medical and academic institutions and waste brokers. Headquartered in Boise, Idaho, the Company is one of the oldest radioactive and hazardous waste services companies in North America. This press release contains forward-looking statements as defined in the Private Securities Litigation Reform Act of 1995 that are based on our current expectations, beliefs and assumptions about the industry and markets in which US Ecology, Inc. and its subsidiaries operate. Because such statements include risks and uncertainties, actual results may differ materially from what is expressed herein and no assurance can be given that the Company will achieve its 2013 earnings estimates, successfully execute its growth strategy, increase market share, or declare or pay future dividends. For information on other factors that could cause actual results to differ materially from expectations, please refer to US Ecology, Inc.'s December 31, 2012 Annual Report on Form 10-K and other reports filed with the Securities and Exchange Commission. Many of the factors that will determine the Company's future results are beyond the ability of management to control or predict. Readers should not place undue reliance on forward-looking statements, which reflect management's views only as of the date such statements are made. The Company undertakes no obligation to revise or update any forward-looking statements, or to make any other forward-looking statements, whether as a result of new information, future events or otherwise. Important assumptions and other important factors that could cause actual results to differ materially from those set forth in the forward-looking information include a loss of a major customer or contract, compliance with and changes to applicable laws, rules, or regulations, access to cost effective transportation services, access to insurance, surety bonds and other financial assurances, loss of key personnel, lawsuits, labor disputes, adverse economic conditions, government funding or competitive pressures, incidents or adverse weather conditions that could limit or suspend specific operations, implementation of new technologies, market conditions, average selling prices for recycled materials, our ability to replace business from recently completed large projects, our ability to perform under required contracts, our ability to permit and contract for timely construction of new or expanded disposal cells, our willingness or ability to pay dividends and our ability to effectively close and integrate future acquisitions. Investors should also be aware that while we do, from time to time, communicate with securities analysts, it is against our policy to disclose to them any material non-public information or other confidential commercial information. Accordingly, stockholders should not assume that we agree with any statement or report issued by any analyst irrespective of the content of the statement or report. Furthermore, we have a policy against issuing or confirming financial forecasts or projections issued by others. Thus, to the extent that reports issued by securities analysts contain any projections, forecasts or opinions, such reports are not the responsibility of US Ecology, Inc. US ECOLOGY, INC. CONSOLIDATED STATEMENTS OF INCOME (in thousands, except per share data) (unaudited) Three Months Ended March 31, -------------------------- 2013 2012 ------------ ------------ Revenue $ 42,899 $ 33,013 Direct operating costs 21,084 17,638 Transportation costs 6,433 3,299 ------------ ------------ Gross profit 15,382 12,076 Selling, general and administrative expenses 5,726 5,605 ------------ ------------ Operating income 9,656 6,471 Other income (expense): Interest income 5 5 Interest expense (221) (224) Foreign currency gain (loss) (938) 1,091 Other 97 80 ------------ ------------ Total other income (expense) (1,057) 952 Income before income taxes 8,599 7,423 Income tax expense 3,193 2,900 ------------ ------------ Net income $ 5,406 $ 4,523 ============ ============ Earnings per share: Basic $ 0.30 $ 0.25 Diluted $ 0.29 $ 0.25 Shares used in earnings per share calculation: Basic 18,320 18,218 Diluted 18,407 18,254 Dividends paid per share $ - $ 0.18 ============ ============ US ECOLOGY, INC. CONSOLIDATED BALANCE SHEETS (in thousands) (unaudited) March 31, December 31, 2013 2012 ------------ ------------ Assets Current Assets: Cash and cash equivalents $ 4,511 $ 2,120 Receivables, net 32,254 33,947 Prepaid expenses and other current assets 2,510 3,161 Deferred income taxes 597 1,276 ------------ ------------ Total current assets 39,872 40,504 Property and equipment, net 113,319 109,792 Restricted cash 4,111 4,111 Intangible assets, net 39,536 40,771 Goodwill 22,617 23,105 Other assets 491 411 ------------ ------------ Total assets $ 219,946 $ 218,694 ============ ============ Liabilities and Stockholders' Equity Current Liabilities: Accounts payable $ 6,842 $ 6,333 Deferred revenue 5,147 3,919 Accrued liabilities 7,093 7,322 Accrued salaries and benefits 4,402 7,570 Income tax payable 3,947 426 Current portion of closure and post-closure obligations 978 1,913 ------------ ------------ Total current liabilities 28,409 27,483 Long-term closure and post-closure obligations 15,608 15,449 Reducing revolving line of credit 41,000 45,000 Other long-term liabilities 102 114 Unrecognized tax benefits 470 467 Deferred income taxes 16,482 18,159 ------------ ------------ Total liabilities 102,071 106,672 Contingencies and commitments Stockholders' Equity Common stock 184 184 Additional paid-in capital 64,537 63,969 Retained earnings 53,830 48,424 Treasury stock (554) (1,183) Accumulated other comprehensive income (loss) (122) 628 ------------ ------------ Total stockholders' equity 117,875 112,022 ------------ ------------ Total liabilities and stockholders' equity $ 219,946 $ 218,694 ============ ============ CONSOLIDATED STATEMENTS OF CASH FLOWS (in thousands) (unaudited) Three Months Ended March 31, -------------------------- 2013 2012 ------------ ------------ Cash Flows From Operating Activities: Net income $ 5,406 $ 4,523 Adjustments to reconcile net income to net cash provided by operating activities: Depreciation and amortization of property and equipment 3,439 3,223 Amortization of intangible assets 367 350 Accretion of closure and post-closure obligations 307 335 Unrealized foreign currency loss (gain) 1,057 (1,201) Deferred income taxes (716) 338 Stock-based compensation expense 146 203 Unrecognized tax benefits 3 3 Net (gain) loss on sale of property and equipment (41) 22 Changes in assets and liabilities: Receivables, net 1,473 7,624 Income tax receivable - 42 Other assets 732 (66) Accounts payable and accrued liabilities (556) (967) Deferred revenue 1,269 (295) Accrued salaries and benefits (3,106) (1,526) Income tax payable 3,528 896 Closure and post-closure obligations (1,043) (82) ------------ ------------ Net cash provided by operating activities 12,265 13,422 Cash Flows From Investing Activities: Purchases of property and equipment (6,752) (2,386) Proceeds from sale of property and equipment 52 66 ------------ ------------ Net cash used in investing activities (6,700) (2,320) Cash Flows From Financing Activities: Payments on reducing revolving line of credit (4,000) (8,500) Proceeds from stock option exercises 1,050 - Deferred financing costs paid (177) - Proceeds from reducing revolving line of credit - 3,000 Dividends paid - (3,282) Other 2 (243) ------------ ------------ Net cash used in financing activities (3,125) (9,025) Effect of foreign exchange rate changes on cash (49) 32 Decrease in cash and cash equivalents 2,391 2,109 Cash and cash equivalents at beginning of year 2,120 4,289 ------------ ------------ Cash and cash equivalents at end of year $ 4,511 $ 6,398 ============ ============ EXHIBIT A Non-GAAP Results and Reconciliation US Ecology reports adjusted EBITDA and adjusted earnings per diluted share results, which are non-GAAP financial measures, as a complement to results provided in accordance with generally accepted accounting principles in the United States (GAAP) and believes that such information provides analysts, shareholders, and other users information to better understand the Company's operating performance. Because adjusted EBITDA and adjusted earnings per diluted share are not measurements determined in accordance with GAAP and are thus susceptible to varying calculations they may not be comparable to similar measures used by other companies. Items excluded from adjusted EBITDA and adjusted earnings per diluted share are significant components in understanding and assessing financial performance. Adjusted EBITDA and adjusted earnings per diluted share should not be considered in isolation or as an alternative to, or substitute for, net income, cash flows generated by operations, investing or financing activities, or other financial statement data presented in the consolidated financial statements as indicators of financial performance or liquidity. Adjusted EBITDA and adjusted earnings per diluted share have limitations as analytical tools and should not be considered in isolation or a substitute for analyzing our results as reported under GAAP. Some of the limitations are: -- Adjusted EBITDA does not reflect changes in, or cash requirements for, our working capital needs; -- Adjusted EBITDA does not reflect our interest expense, or the requirements necessary to service interest or principal payments on our debt; -- Adjusted EBITDA does not reflect our income tax expenses or the cash requirements to pay our taxes; -- Adjusted EBITDA does not reflect our cash expenditures or future requirements for capital expenditures or contractual commitments; and -- although depreciation and amortization charges are non-cash charges, the assets being depreciated and amortized will often have to be replaced in the future, and Adjusted EBITDA does not reflect cash requirements for such replacements. Adjusted EBITDA The Company defines Adjusted EBITDA as net income before interest expense, interest income, income tax expense, depreciation, amortization, stock based compensation, accretion of closure and post-closure liabilities, foreign currency gain/loss and other income/expense, which are not considered part of usual business operations. The following reconciliation itemizes the differences between reported net income and Adjusted EBITDA for the three months ended March 31, 2013 and 2012: Three Months Ended (in thousands) March 31, -------------------------- 2013 2012 ------------ ------------ Net Income $ 5,406 $ 4,523 Income tax expense 3,193 2,900 Interest expense 221 224 Interest income (5) (5) Foreign currency (gain)/loss 938 (1,091) Other income (97) (80) Depreciation and amortization of plant and equipment 3,439 3,223 Amortization of intangibles 367 350 Stock-based compensation 146 203 Accretion and non-cash adjustments of closure & post-closure liabilities 307 335 ------------ ------------ Adjusted EBITDA $ 13,915 $ 10,582 ============ ============ EXHIBIT A Non-GAAP Results and Reconciliation, continued Adjusted Earnings Per Diluted Share The Company defines adjusted earnings per diluted share as net income plus the after tax impact of non-cash, non-operational foreign currency gains or losses ("Foreign Currency Gain/Loss") plus the after tax impact of business development cost divided by the diluted shares used in the earnings per share calculation. The Foreign Currency Gain/Loss excluded from the earnings per diluted share calculation are related to intercompany loans between our Canadian subsidiary and the U.S. parent which have been established as part of our tax and treasury management strategy. These intercompany loans are payable in CAD requiring us to revalue the outstanding loan balance through our consolidated income statement based on the CAD/USD currency movements from period to period. We believe excluding the currency movements for these intercompany financial instruments provides meaningful information to investors regarding the operational and financial performance of the Company. Business development costs relate to expenses incurred to evaluate businesses for potential acquisition or costs related to closing and integrating successfully acquired businesses. Business development costs in 2012 include the acquisition of Dynecol, Inc. which closed on May 31, 2012 and other business development and strategic planning activities. We believe excluding these business development costs provides meaningful information to investors regarding the operational and financial performance of the Company. The following reconciliation itemizes the differences between reported net income and earnings per diluted share to adjusted net income and adjusted earnings per diluted share for the three months ended March 31, 2013 and 2012: (in thousands, except per share data) Three Months Ended March 31, ------------------------------------------ 2013 2012 --------------------- -------------------- per share per share Net income / earnings per diluted share $ 5,406 $ 0.29 $ 4,523 $ 0.25 Business development costs, net of tax $ - $ - 41 $ - Non-cash foreign currency (gain)/loss, net of tax $ 595 $ 0.03 (702) $ (0.04) ---------- ---------- --------- --------- Adjusted net income / adjusted earnings per diluted share $ 6,001 $ 0.32 $ 3,862 $ 0.21 ========== ========== ========= ========= Shares used in earnings per diluted share calculation 18,407 18,254 ========== ========= Contact: Alison Ziegler Cameron Associates (212) 554-5469 email@example.com www.usecology.com
US Ecology Announces First Quarter 2013 Results
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Posts from Other Sweeteners
Why I Love to Bake With Natural Sugars
I got a text message from my mom a few days ago: Demerara sugar?! I inquired as to what she was referring to so early on a Tuesday morning, and she mentioned that a new pie she was experimenting with for Thanksgiving called for the ingredient. What the heck? she followed, Why not just white sugar like I've done for the last three centuries?
Corn Syrup vs. High-Fructose Corn Syrup: There Is a Difference
The holidays are coming, which means it's time for pecan pie, homemade marshmallows and caramel candies — all recipes that traditionally use corn syrup as an ingredient. More and more, I've been hearing people say they are reluctant to use corn syrup in their baking because of the negative health effects associated with high-fructose corn syrup. But the truth is that corn syrup and high-fructose corn syrup are two different products.
Say Goodbye to Grade B Maple Syrup: Vermont Changes Its Maple Syrup Labeling
Grade B, Grade A... have you ever been confused by maple syrup labeling? Grade "A" sounds like a higher-quality product but this is misleading; personally, I prefer Grade B, which is just as pure but darker and more robust in flavor.
Well, things are about to get less confusing. Vermont is changing their maple syrup labeling, so say goodbye to Grade B.
Help Me Find a Recipe to Make the Most of Maple Sugar!
Good Questions
Q: My supervisor recently gave me a smallish container (maybe a cup total) of maple sugar he bought on vacation, saying that I'm the baker in the office so he thought I would know what to do with it. I've never seen maple sugar before, but I'd like to feature it in a dish, preferably a baked good, so I can bring it to the office to share. Any ideas?
What Are Some Great Recipes with Honey?
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I was wondering if you could suggest some great recipes that include honey.
Sent by Danielle
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Joanne Weir, a San Francisco chef, tequila expert, and the subject of one
of our early kitchen tours, makes a margarita at her Bay Area restaurant Copita
that is known as one of the best in the region. She says — and I agree
whole-heartedly — that the biggest mistake people make with their margaritas
is that they use too many ingredients like Triple Sec, Grand Marnier, or Cointreau.
It's all about the balance of sweet and tart in a glass with just a few
ingredients: 100% agave tequila (Weir prefers blanco), freshly squeezed lime
juice and agave nectar.
Alternative Sweetener: Monk Fruit
Ingredient Spotlight
Monk fruit is the latest darling of the alternative sweetener world, but what is this exotic fruit, and how does it compare to sugar and other substitutes?
No-Bake Snacking: 7 Bars, Cookies and Bites to Pick You Up at 3pm
Even though I work largely from home, I'm still plagued by the 3 pm energy plummet like many of my friends and old coworkers. There's just something about that hour, especially in the winter when the days are colder and darker. Instead of reaching for a soda or another cup of coffee, a small snack with a bit of protein is a great picker-upper, and lately I've been mixing up a batch of quick no-bake snacks to keep on hand when the energy lows strike.
Baker's Ingredient Spotlight: What to Do With Maple Sugar
I don't use white sugar much at home these days. Now I reach for a more unrefined cane sugar like turbinado, or experiment with maple syrup or honey as sweeteners. But the problem with this, for bakers in particular, is that you have to be careful about adding extra liquid to a recipe because it changes the entire composition and structure. So what's a good solution if you want to add a deep, dark sweetener like maple syrup? Pick up a bag of maple sugar!


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In her native tongue, a mother began telling a tale.
"Thousands of years ago, on a small continent hidden from the others, there was a small peninsula at the very bottom of the mass, called the Forbidden This land was a sight to see, filled with majestic forests, plains, and lakes. Besides the birds, no other wildlife lived there. It was tranquil, very silent. There were no sounds, not even the familiar singing of the birds in the morning. No, the only thing that had lived in the land were the sixteen colossi that inhabited each region of the gorgeous peninsula, guarding their respective sectors and living peacefully. However, after hundreds of years, the ancient deity Dormin found the Forbidden Land and deemed it his realm, establishing a massive temple in the middle of the peninsula that dwarfed everything, even the colossi. Connecting the giant structure, named the Temple of Worship, to the highly elevated entrance to the land, Dormin built a narrow bridge. This bridge was miles long, and if one was to fall off it, they'd be facing a thousand foot drop.
"After constructing the Temple of Worship and the bridge leading to it, Dormin inhabited the Land for a hundred years, doing what he did best from his little safe haven - corrupting people, torturing people, and killing people. Not any specific kind of person, no. Dormin didn't have a taste; with is vast power, the god killed anyone he saw fit, and most of the people he killed was for his own interest, their souls empowering him. Being the God of Death, every person killed was a soul put inside of him. However, having the souls of the dead, he was able to put them back in the body, therefore resurrecting that person. However, the laws of mortality didn't allow him to do that. After a while, though, he had so many souls inside of him he became even more corrupted than he already was. With his newly corrupted mind, he decided to make an army of the dead/
"After a few years, Dormin had the bodies of thousands of men and women, the souls put inside them twisted so they followed his orders. And Dormin's ordered his newly-found army to annihilate the civilized nation of Shaiman. Dormin's army marched down the bridge, through the mountains, and into the country, slaughtering everybody village by village. Women, children, men, anything moving was brutally killed. Eventually, the nation took arms, fighting back. Bloody battles took place, the country's army winning some and Dormin's army winning others. However, after seven long years, Dormin's army was slowly killed off, but not after wiping out forty percent of Shaiman's population. Unfortunately, one of the people killed in the war was the king of Shaiman, his throat slit in battle. His successor was his nephew, Emon, for his son was also killed in battle.
"When the war was over, Lord Emon hunted Dormin down to the Forbidden Land. However, instead of killing the deity, Emon split the god into sixteen parts, each part of his soul going into a large statue lining the sides of the temple's main chamber. To make these statues invulnerable, Emon made a spiritual tie between each statue and a colossus, the massive figurines not breaking until the colossus corresponding to the stone statue was dead. To make things even harder for the god, the Lord then made it to where the colossi couldn't be killed with anything other than a single sword. This sword was a blade inherited by each king, passed down royally for ten generations. Each king had carved a rune into the blade, and because of this, people rumored the "Ancient Sword" to be magical, which in fact it was.
"After containing Dormin in the temple and the colossi, Emon and his men left, building a large door at the entrance to the lands and making it clear to all who passed the gate that this land was forbidden, and that trespassers would be killed. People understood this message, and after a while, it the story of Emon, Dormin, and the Ancient War became myth, and to some, all they understood from the story was that Dormin could bring back the dead. One of the people who understood this was a boy named Wander, who was raised on the story. With Emon still ruling after fifty years, Wander believed too little time had passed for the story to be completely false. He knew that at least half of it had to be true, and one of the things that were true was the existence of Dormin and all his powers.
"Wander was always classified as a lover, not a fighter. Everyone in his village knew that Wander was in love with the maiden Mono, and everyone in his village thought that Mono was cursed. Her birthday was March 14th, the same day the Ancient War started. Because of this, people believed she had been scarred by Dormin and the dead from memories brought on by that day. After someone thought she was going to be a killer because of her birthday, and because of a birthmark on her arm that resembled Dormin's elaborate symbol, which looked like a small circle with lines surrounding it to form an x-like shape, they called the local shaman, who deemed her evil and sacrificed her in a religious ritual. Infuriated, Wander dug Mono's body from the grave, and rode his horse, Agro, to the capital city, where he infiltrated the Lord's house and stole the Ancient Sword based on his memory of the Emon-Dormin ordeal. After taking the sword, he used a map of the continent to navigate through the rocky, mountainous terrain of Shaiman, he got to the Forbidden Land's gate. After reading all the signs, he happily rode past the gate and onto the bridge, surprised at the land's beauty and existence but relieved as well.
"He had gotten to the Temple of Worship, where he met the shattered, genderless, and mysterious Dormin, who told him he had to kill all sixteen colossi in order to break all the statues and free Dormin, so he could resurrect Mono. Wander said yes, dismissing Dormin's warning to the cost of the resurrection and killing the first eight colossi, climbing the massive beasts and stabbing them in their brains and such. However, at those eight, when Dormin's voice was becoming more masculine and powerful, the god noticed that Wander was physically deteriorating, his skin becomine pale and blotchy and his hair darkening. But Wander continued to kill four more with ease, and after the twelfth colossus fell, its released soul destroying its statue, Dormin sensed Wander becoming more primative and cruel in his movements. Dormin grinned at this however, as Wander eventually killed the last colossus, destroying the last colossus. When the boy returned to the temple, his skin and eyes were completely white, and he had small horns portruding from his head. All the statues were destroyed, and finally, Dormin was whole again. Seeking a physical body, he possessed the already corrupted Wander, just as Lord Emon and his men had reached the Forbidden Lands. The king decided to kill Wander, knowing he was corrupted beyond repair. His men shot Wander, and the boy slowly fell to the ground, black blood erupting from his body. A death suffered by the colossi.
"With Dormin's death, the Forbidden Land began to fall apart, and the temple began to collapse. Emon and his men fled immediately, not caring about anything but their lives. They left just as the bridge to the temple collapsed to the ground. When they left, everything was left in ruins. The trees had all fallen, the cliffs had all fallen in, the bridge had fallen in segments and littered the ground, and half the temple had collapsed. However, past all the destruction, beside the altar where Mono's body lay during the whole process, stood a beautiful girl. It was Mono, reincarnated. After all that Wander had gone through, his death was not in vein. He went to the ends of the Earth for the woman he loved, and in the end, died, but he got what he wanted...at a great price, like Dormin warned."
"But what happened to Dormin?" a young boy asked as she ended her story. "I mean, if he died when Wander did, then how could Mono've been revived?"
His mom thought for a moment, then shook her head. "I don't know...I'd never really thought of it. It's been a thousand years since this all happened, my son. If Dormin did survive, then he's been dormant for this whole time...maybe he reformed, turned into a good being. But maybe...maybe he's still as evil as he always has been, and maybe he hunts down little boys who don't go to bed," she replied with a smirk. It was true, the boy listened to this entire tale while in his room. It was midnight, yet he wasn't tired. He had thought that his favorite story would've tired him, but it engrossed him instead. His mom's way with words, how she could spin such an intricate web, using details and her voice to make him feel a certain emotion. With this story, he felt sad for Wander, for he never got to see his love live again.
"Tell me about the colossi," the boy asked, ignoring his mom's words about Dormin and not sleeping. "I want to know about them. My tutor at the academy mentioned them before when we were studying myths...this myth."
His mom chuckled. "My boy, this is no myth. Our family's been tied into this story for a thousand years. Did your teacher ever say how our line is supposed to be related to Wander's? We live in the same village he did. Maybe your teachers will tell you about that as well. As for the colossi? I thought I already told you about them when I told you the story before?" he gave her a, Refresh my memory face, and she sighed, then chuckled. "However, ha. They were peaceful beings. Taller than most buildings, they were nearly invulnerable. The only weapon that could kill them was the Ancient Sword Wander wielded, and the legend says that he could only kill a colossus by stabbing them in their most vital organs, like the brain and heart. It also says that when they fell, it was the saddest thing ever. Seeing the light leaving their eyes, their bodies slowly collapse...it must've been extremely depressing."
The boy thought about it a little bit, and then his mom got up from her chair. "I'll see you tomorrow," she said. "Now get in bed." He slowly got under the quilts of his bed and put his hands under his pillow, which for him was the only way he could fall asleep. His mom went to his door, then squeezed his candle's flames out with two fingers. "Good night...I love you, my Seeker."
With that, she left, leaving the boy, named Seeker, to think about the story. He had always wondered, since his sixth birthday a month ago, how Dormin could've been dead, even when he resurrected Mono after Emon left. Maybe it was just something grown-ups understood. Or was it? He didn't know. And as for Seeker being related to Wander? He didn't know what to think of that. Wander, who's story has been one of the most told legends of all time? the story of Wander, Mono, Agro, Dormin, and the sixteen beasts of old were second only to the story that sparked Wander's story; the tale of the Ancient War. To think that he was related to the man - well, boy; he was a teenager when the story unfolded - who was the very essence of "going to the ends of the earth for the one you love" made Seeker feel humbled. It made him feel special. It made him feel...confused. Very confused. Maybe when he was a grown-up, he'd understand. He just couldn't grasp the concept of being related to a legend. He didn't even know if it was possible. Maybe Wander and Mono had kids? Maybe after he died, Wander was...no, he couldn't of been reborn? Dormin never promised to bring Wander back. But maybe, if Dormin possessed Wander at the end, and if he wasn't dead to revive Mono, maybe...
Seeker didn't know. He knew he was too young to be thinking of such things. He knew that in the end, he'd confuse himself to insanity before he turned eight. He rolled around on his bed, trying to find a comfortable position to fall asleep in. However, when he did, he knew he wouldn't fall asleep for another hour. It had always been a problem with him. Sleeping just...wasn't his thing. So, after an hour of just laying there and thinking about things all young children thinking about, like eating and playing around the village and disliking academy, he gave way to the weight gaining on his eyelids. His eyes slowly shut, and before he knew it, he was in the celestial realm of his dreams and subconsciousness.
A/N And so ends the first chapter of this story. Just to let you know, I don't know when I'll update next so don't be mad if it's in a week or a month. I don't know. Anyways, yeah. Read & Review. I know this isn't the most popular subject to write on, but if you find it, comment!


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After being available on iOS for quite some time, Sega has released Sonic Dash into the Play Store. In true Sonic fashion, he's running; however, unlike other editions of the popular franchise, this one doesn't have the hedgehog traveling from left to right. Sonic Dash is like Temple Run in that you are running forward, dodging enemies and obstacles that enter your path. It is free-to-play, so you can give this a whirl without having to pay anything.
Hit the break for the trailer, gallery, and download links. Read more