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Having thus given an account of the manner in which he filled his public offices both civil and military, and his conduct in the government of the empire, both in peace and war; I shall now describe his private and domestic life, his habits at home and among his friends and dependents, and the fortune attending him in those scenes of retirement, from his youth to the day of his death. He lost his mother in his first consulship, and his sister Octavia, when he was in the fifty-fourth year of his age. 1 He behaved towards them both with the utmost kindness whilst living, and after their decease paid the highest honours to their memory.
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Learning objectives:
- Describe how feed is manufactured and the typical components of feed and premixes.
- Describe the nutritional requirements of broilers and layers.
- Describe and plan an optimal feeding regimen for broiler and layer chickens.
- Describe poultry problems arising from nutritional defects and how to remedy these problems.
- Describe the main benefits of supplementing feed with enzymes.

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195 Bellevue Dr
$3,475,000, 5 Beds 6 Baths 5,769 Sqft .45 Acres 3 Garages
About This Property
Stunning new home on coveted Bellevue Drive. Tucked into the hillside w/ expansive foothills, city & plains views. Tons of natural light & outdoor living space. Unsurpassed finishes throughout. Open floor plan w/formal living & dining rooms, magnificent chef's kitchen w/ butler's pantry & casual dining area, main floor master w/spacious luxury bath, second main floor bedroom suite. Three more beds up along w/ loft/family room area.Huge unfinished basement for future rec room. One of a kind!
Essential Information
Community Information
Foothills View, Plains View, City View
Interior Features
Eat-in Kitchen, Separate Dining Room, Cathedral/Vaulted Ceilings, Open Floor Plan, Pantry, Walk-in Closet, Washer/Dryer Hookups, Wood Floors, Kitchen Island
Gas Range/Oven, Self-Cleaning Oven, Double Oven, Dishwasher, Refrigerator, Bar Refrigerator, Microwave, Disposal, Smoke Alarm(s)
Hot Water, Multi-zoned Heat, Radiant Heat, 2 or more Heat Sources, 2 or more H20 Heaters
Central Air Conditioning, Ceiling Fan
2+ Fireplaces, Gas Fireplace, Master Bedroom Fireplace, Great Room Fireplace
Exterior Features
Lawn Sprinkler System, Balcony, Patio, Deck, Oversized Garage
Full Basement, Unfinished Basement, Walk-out Basement
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195 Bellevue Dr, Boulder (MLS® #833271) Property History
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Information is provided exclusively for consumers' personal, non-commercial use and may not be used for any purpose other than to identify prospective properties consumers may be interested in purchasing.
Information deemed reliable but not guaranteed by the MLS.
Listing information last updated on March 29th, 2020 at 7:04pm MDT.


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Rebellion is currently in the process of remastering their classic Rogue Trooper as Rogue Trooper Redux. Not everyone will be familiar with the 2000AD character so Rebellion wants to make sure you're up to speed.
This is a remaster of their 2006 release which stars the one and only Rogue, a genetic infantryman (GI). He's a full squad because he carries the consciousness of three fallen GIs via biochips. He's now on a mission to avenge his fallen comrades.
The video also comes with a first look at the Rogue character who has been redesigned from the ground up for his remaster. He's looking good.
You can also follow the remaster news via the official website.


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Participation is open to children 3-14 with a wide variety of communication challenges, including:
Clinical sessions are half-day (full day optional for ages seven through fourteen). Participants must be capable of working in small groups of two to four individuals. Participant age, nature, and severity of the communication disorder are considered prior to acceptance. Final enrollment is based on the availability of services needed by each prospective applicant. Application acceptance is now open. The deadline is April 1st to assure consideration. Late applications will be processed on a space available basis. Due to limited space, early application is highly recommended. Pre-clinic evaluations are conducted at Central Michigan University during May and June to determine intervention goals and approaches for each applicant, as well as the appropriateness of our summer program. Camp dates for 2020 are from July 1 through August 4, 2020. Children attend camp Monday - Thursday only. There is no camp on Friday. Clinicians will be engaged in workshops, planning and administrative tasks.
Enrollment Application Form Application Forms


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Pirate Basketball Shooting Drill Series (Videos Included)
In June, we released the 3 main parts of our Pirate Basketball Shooting Drills series, in which Pro Skills Basketball Denver Director Ross Schraeder teaches and demonstrates layups in part 1, "runners" in part 2, and reverse layups in part 3.
The drill is called the "Pirate" Basketball Shooting Series because it teaches all one-handed basketball shooting.
Get it?
One-handed … like a pirate!
Still no?!?
Haha well, regardless, we wanted to give it a unique name that the kids will remember while working on an important aspect of the game.
Ideally, it is meant to work on weak-handed development, but, in fact, the work is fantastic for both strong-hand and weak-hand development.
In performing this basketball shooting drill, we are looking for players to shoot with proper form, right or left-handed, without using the off-hand in the shot.
Especially on weak-handed layups, kids will begin to push with their off hand, and never really fully develop with their weak hand.
Check out layups in part 1 of the Pirate Basketball Shooting Drills Series video below:
By eliminating one hand with these Pirate Series drills, we force young basketball players to use their weak hand only. With a young team, at the beginning, they are a mess in trying to control the ball with one hand.
However, in just a few minutes, you can see them begin to improve and start to come close to getting the proper form.
Also important in performing this shooting drill is the fact that a vital aspect of being a good basketball player is having good hands and controlling the ball effectively.
See "runners" in part 2 of the Pirate Basketball Shooting Drills Series video below:
We start our Pirate Basketball Shooting Drills Series by having the kids cradle the ball, but then turn it so that they are in a sense shooting a shot (even though it is a layup) rather than shooting an underhand layup.
The cradle plus turn of the ball with one hand is quite challenging at first for players of any age who have not performed a one-hand basketball shooting drill before, but young players begin to see their ball control and coordination develop rapidly in continuing the session.
Check out part 3 and reverse layups in the Pirate Basketball Shooting Drills Series video below:
The Pirate Series doesn't have to just be for shooting a basketball though! It can also be used for a basic two-man passing drill with making one-hand passes off the dribble.
Again, this helps weak-hand development, helps coordination, and speeds up their overall development.
One thing to stress in the pirate session is that we do not want them making one-handed plays in actual gameplay.
We are simply speeding up their weak-hand development in practice and training sessions, and thus making them more coordinated youth basketball players.
Make sure to check out the rest of our youth basketball drills videos and subscribe to your YouTube Channel!
Do you want to see more drills?
Let us know what skills you would like us to cover in our basketball drill videos! Simply drop us a note on our contact page here.


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Sarkar 3 trailer: Amitabh Bachchan is angrier than ever in RGV's film
The trailer of Ram Gopal Varma's Sarkar 3 starring Amitabh Bachchan and Manoj Bajpayee is out now, and it looks like the dark film we wanted for so long.bollywood Updated: Mar 01, 2017 21:03 IST
Ram Gopal Varma, who has been struggling to find his lost form in Bollywood for some time, is relying heavily on Amitabh Bachchan's charisma to rise and shine again.
The trailer of Sarkar 3, the third film in the Sarkar franchise that started in 2005, is all about the return of Subhash Nagre (Amitabh Bachchan), who has already lost his sons, Vishnu (Kay Kay Menon) and Shankar (Abhishek Bachchan).
To set the tone right, Varma has used a similar background score and lighting to that of the first two films. Of course, Subhash Nagre's get-up will also be the same.
Amitabh Bachchan in and as SARKAR in SARKAR 3 Trailer.... May the POWER be with you... https://t.co/ZuUM8b0lJa— Ram Gopal Varma (@RGVzoomin) March 1, 2017
He has also brought in some top actors like Manoj Bajpayee, Ronit Roy and Jackie Shroff to flesh out his screenplay effectively. If Bajpayee is playing an idealist rebel in the film, Ronit Roy will be replacing Chander (Ravi Kale) as Sarkar's most trusted ally.
Amit Sadh and Yami Gautam are also in the film and they are expected to carry the Sarkar legacy forward.
However, the most striking thing about this trailer is its announcement that Amitabh Bachchan will be seen in an angrier, meaner avatar. In fact, he says that he hasn't changed over the years.
But all said and done, Sarkar 3 is probably Varma's last shot to reclaim his space in Bollywood as the most sought after noir filmmaker.
Sarkar 3 will hit the screens on April 7, 2017.


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STRATFORD, Mark Russell
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Warrant Officer Mark Stratford, UK Landing Force Command Support Group, died in a helicopter crash.
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Just an FYI:
Due to my present overload of writing responsibilities and other various pressures and stresses, I've decided to surrender my participation at CT Movies. My latest Through a Screen Darkly column, "Feasting on Films", will be my last contribution there for the foreseeable future (although I certainly won't close the door, as circumstances may change).
It's a sad day for me, in some ways: I've contributed to CT's online endeavors regularly since sometime around the beginning of 2001. I remember that one of the editors at CT told me, about a year-and-a-half into it, that he was surprised I hadn't "burned out" writing about the turbulent territory of religious media and Christian engagement with film. As it is, I still love the topic, and am as passionate about writing about it now as I was eight years ago… and even eighteen years ago, when I wrote about it for Seattle Pacific's student newspaper.
It was tough to make this decision. But I've been taking a lot of steps lately to reclaim a healthy lifestyle. Cutting back on major projects is a part of that, even though that means I have to give up some things that have given me great pleasure in the past.
I've learned so much, made so many new friends, and had so many memorable conversations and debates in my years writing for CT. I'll never forget Film Forum, or my first tour of the CT offices, or the weekends when I met other fine reviewers like Steven Greydanus and Steve Beard, and enjoyed interviewing the casts of Return of the King, Constantine, Firefly, Big Fish, and other movies. I hope that someday I'll find myself in a position to contribute there again.
When I look at how the cultural conversation about Christian engagement with art and entertainment has changed in the last 20 years, I am again impressed with the spirit of courageous engagement at CT. CT has blazed trails that are so much more rewarding than those paths of cultural condemnation and fear-mongering still traversed by so many in religious media.
I'm especially grateful to Steve Lansingh, who first recommended me to take over his Film Forum column in 2000, and CT's Ted Olsen who agreed to take me on, and coached me along the way. That was my introduction to CT, and my first real "job" as a reviewer. My life would be very, very different today had that not occurred. I'm also grateful to David Neff, Mark Moring, and John Wilson who have all been generous with me. I'd like to think I've learned a little about writing along the way. I plan to stay in touch with them, and with all of the friends who have enriched this experience and taught me so much.
But no, I'm not tired of writing about these things. I'll keep on, so long as there are folks interested in reading. This journey's still in the early stages. Thanks so much to all of you, too, that have been encouraging along the way.


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Cross-ecosystem movements of material and energy are ubiquitous. Aquatic ecosystems typically receive material that also includes organic matter from the surrounding catchment. Terrestrial-derived (allochthonous) organic matter can enter aquatic ecosystems in dissolved or particulate form. Several studies have highlighted the importance of dissolved organic carbon to aquatic consumers, but less is known about allochthonous particulate organic carbon (POC). Similarly, most studies showing the effects of allochthonous organic carbon (OC) on aquatic consumers have investigated pelagic habitats; the effects of allochthonous OC on benthic communities are less well studied. Allochthonous inputs might further decrease primary production through light reduction, thereby potentially affecting autotrophic resource availability to consumers. Here, an enclosure experiment was carried out to test the importance of POC input and light availability on the resource use in a benthic food web of a clear-water lake. Corn starch (a C-4 plant) was used as a POC source due to its insoluble nature and its distinct carbon stable isotope value (delta C-13). The starch carbon was closely dispersed over the bottom of the enclosures to study the fate of a POC source exclusively available to sediment biota. The addition of starch carbon resulted in a clear shift in the isotopic signature of surface-dwelling herbivorous and predatory invertebrates. Although the starch carbon was added solely to the sediment surface, the carbon originating from the starch reached zooplankton. We suggest that allochthonous POC can subsidize benthic food webs directly and can be further transferred to pelagic systems, thereby highlighting the importance of benthic pathways for pelagic habitats.
The effect of increased habitat heterogeneity in tidal areas on coexistence between Palaemon prawns was studied at eight sites along the European Atlantic coast. Two species which are sympatric in non-tidal areas, Palaemon adspersus Rathke and P. squilla (L.) are largely allopatric in tidal areas, and the sympatric size difference decreases in allopatry. In tidal areas the smaller species, P. squilla, is restricted to brown algal belts and rockpools. A third species, P. serratus (Pennant), larger than the others, occurs under oceanic salinities in subtidal brown algal belts and there has forced P. squilla to restrict its habitat distribution to adjacent intertidal rockpools. At estuarine salinities, however, P. squilla also inhabits the brown algal belts. A larger diel variation in stomach fullness index in P. squilla than in P. adspersus persists in non-tidal areas. Abiotic factors probably restrict the two largest species, viz. P. adspersus and P. serratus, to subtidal environments; these species are sensitive to the extremes in salinity, temperature or O2 levels characteristic of the intertidal zone. Competition and/or predation probably relegates P. squilla to the intertidal zone. In non-tidal areas, where this zone is reduced, P. squilla increases its niche width and coexists with P. adspersus, and the size differentiation associated with sympatry may reduce interspecific competition.
The reasons behind the absence of the prawn Palaemon adspersus and the presence of P. squilla in rockpools and on bare sand bottoms were studied. Some maximal abundances in different habitats are given. Introduction experiments into natural and artificial rockpools and measurements of tolerance towards low oxygen levels showed that nocturnal hypoxia excluded P. adspersus which was significantly more sensitive to oxygen depletion. Respiration rates measured by the closed-bottle method showed no interspecific difference. On bare sand bottoms P. aspersus was probably excluded by predators, since predator exclusion experiments in cages and predator inclusion experiments in containers showed that P. adaspersus was more vulnerable to predation than P. squilla.The costs for being able to cope with a wide array of habitat in P. squilla are probably balanced by the benefits of access to habitats such as intertidal rockpools, very shallow bottoms and deeper sand bottoms. Tolerance towards abiotic factors extends its habitat range upwards into shallower waters, and tolerance towards biotic factors, i.e. predation, extends it downwards.
Ecosystem processes of northern peatlands are largely governed by the vitality and species composition in the bryophyte layer, and may be affected by global warming and eutrophication. In a factorial experiment in northeast China, we tested the effects of raised levels of nitrogen (0, 1 and 2 g m(-2) year(-1)), phosphorus (0, 0.1 and 0.2 g m(-2) year(-1)) and temperature (ambient and +3A degrees C) on Polytrichum strictum, Sphagnum magellanicum and S. palustre, to see if the effects could be altered by inter-specific interactions. In all species, growth declined with nitrogen addition and increased with phosphorus addition, but only P. strictum responded to raised temperature with increased production of side-shoots (branching). In Sphagnum, growth and branching changed in the same direction, but in Polytrichum, the two responses were uncoupled: with nitrogen addition there was a decrease in growth (smaller than in Sphagnum) but an increase in branching; with phosphorus addition growth increased but branching was unaffected. There were no two-way interactions among the P, N and T treatments. With increasing temperature, our results indicate that S. palustre should decrease relative to P. strictum (Polytrichum increased its branching and had a negative neighbor effect on S. palustre). With a slight increase in phosphorus availability, the increase in length growth and production of side-shoots in P. strictum and S. magellanicum may give them a competitive superiority over S. palustre. The negative response in Sphagnum to nitrogen could favor the expansion of vascular plants, but P. strictum may endure thanks to its increased branching.
Carotenoids in the egg yolks of birds are considered to be important antioxidants and immune stimulants during the rapid growth of embryos. Yolk carotenoid composition is strongly affected by the carotenoid composition of the female's diet at the time of egg formation. Spatial and temporal differences in carotenoid availability may thus be reflected in yolk concentrations. To assess whether yolk carotenoid concentrations or carotenoid profiles show any large-scale geographical trends or differences among habitats, we collected yolk samples from 16 European populations of the pied flycatcher, Ficedula hypoleuca. We found that the concentrations and proportions of lutein and some other xanthophylls in the egg yolks decreased from Central Europe northwards. The most southern population (which is also the one found at the highest altitude) also showed relatively low carotenoid levels. Concentrations of beta-carotene and zeaxanthin did not show any obvious geographical gradients. Egg yolks also contained proportionally more lutein and other xanthophylls in deciduous than in mixed or coniferous habitats. We suggest that latitudinal gradients in lutein and xanthophylls reflect the lower availability of lutein-rich food items in the northern F. hypoleuca populations and in montane southern populations, which start egg-laying earlier relative to tree phenology than the Central European populations. Similarly, among-habitat variation is likely to reflect the better availability of lutein-rich food in deciduous forests. Our study is the first to indicate that the concentration and profile of yolk carotenoids may show large-scale spatial variation among populations in different parts of the species' geographical range. Further studies are needed to test the fitness effects of this geographical variation.
In predator-prey interactions, the efficiency of the predator is dependent on characteristics of both the predator and the prey, as well as the structure of the environment. In a field enclosure experiment, we tested the effects of a prey refuge on predator search mode, predator efficiency and prey behaviour. Replicated enclosures containing young of the year (0+) and 1-year-old (1+) perch were stocked with 3 differentially sized individuals of either of 2 piscivorous species, perch (Perca fluviatilis), pike (Esox lucius), or no piscivorous predators. Each enclosure contained an open predator area with three small vegetation patches, and a vegetated absolute refuge for the prey. We quantified the behaviour of the predators and the prey simultaneously, and at the end of the experiment the growth of the predators and the mortality and habitat use of the prey were estimated. The activity mode of both predator species was stationary. Perch stayed in pairs in the vegetation patches whereas pike remained solitary and occupied the corners of the enclosure. The largest pike individuals stayed closest to the prey refuge whereas the smallest individuals stayed farthest away from the prey refuge, indicating size-dependent interference among pike. Both size classes of prey showed stronger behavioural responses to pike than to perch with respect to refuge use, distance from refuge and distance to the nearest predator. Prey mortality was higher in the presence of pike than in the presence of perch. Predators decreased in body mass in all treatments, and perch showed a relatively stronger decrease in body mass than pike during the experiment. Growth differences of perch and pike, and mortality differences of prey caused by predation, can be explained by predator morphology, predator attack efficiency and social versus interference behaviour of the predators. These considerations suggest that pike are more efficient piscivores around prey refuges such as the littoral zones of lakes, whereas perch have previously been observed to be more efficient in open areas, such as in the pelagic zones of lakes.
Bumble bee abundance in agricultural landscapes is known to decrease with increasing distance from seminatural grasslands, but whether the pollination of bumble-bee-pollinated wild plants shows a similar pattern is less well known. In addition, the relative effects of landscape composition (landscape heterogeneity) and landscape configuration (distance from seminatural grassland) on wild plant pollination, and the interaction between these landscape effects, have not been studied using landscape-level replication. We performed a field experiment to disentangle these landscape effects on the pollination of a native herb, the sticky catchfly (Lychnis viscaria), while accounting for the proportion of oilseed rape across landscapes and the local abundance of bee forage flowers. We measured pollen limitation (the degree to which seed set is pollen-limited), seed set, and seed set stability using potted plants placed in landscapes that differed in heterogeneity (composition) and distance from seminatural grassland (configuration). Pollen limitation and seed set in individual plants did not respond to landscape composition, landscape configuration, or proportion of oilseed rape. Instead, seed set increased with increasing local bee forage flower cover. However, we found within-plant variability in pollen limitation and seed set to increase with increasing distance from seminatural pasture. Our results suggest that average within-plant levels of pollen limitation and seed set respond less swiftly than the within-plant variability in pollen limitation and seed set to changes in landscape configuration. Although landscape effects on pollination were less important than predicted, we conclude that landscape configuration and local habitat characteristics play larger roles than landscape composition in the pollination of L. viscaria.
The preference-performance hypothesis posits that the host plant range of plant-feeding insects is ultimately limited by larval costs associated with feeding on multiple resources, and that female egg-laying preferences evolve in response to these costs. The trade-off of either using few host plant species and being a strong competitor on them due to effective utilization or using a wide host plant range but being a poor competitor is further predicted to result in host plant specialization. This follows under the hypothesis that both females and offspring are ultimately favoured by utilizing only the most suitable host(s). We develop an experimental approach to identify such trade-offs, i.e. larval costs associated with being a host generalist, and apply a suite of experiments to two sympatric and syntopic populations of the closely related butterflies Pieris napi and Pieris rapae. These butterflies show variation in their level of host specialization, which allowed comparisons between more and less specialized species and between families within species. Our results show that, first, the link between female host preference and offspring performance was not significantly stronger in the specialist compared to the generalist species. Second, the offspring of the host plant specialist did not outperform the offspring of the generalist on the former's most preferred host plant species. Finally, the more generalized species, or families within species, did not show higher survival or consistently higher growth rates than the specialists on the less preferred plants. Thus, the preference and performance traits appear to evolve as largely separated units.
Increased N deposition in Europe has affected mire ecosystems. However, knowledge on the physiological responses is poor. We measured photosynthetic responses to increasing N deposition in two peatmoss species (Sphagnum balticum and Sphagnum fuscum) from a 3-year, north-south transplant experiment in northern Europe, covering a latitudinal N deposition gradient ranging from 0.28 g N m(-2) year(-1) in the north, to 1.49 g N m(-2) year(-1) in the south. The maximum photosynthetic rate (NPmax) increased southwards, and was mainly explained by tissue N concentration, secondly by allocation of N to the photosynthesis, and to a lesser degree by modified photosystem II activity (variable fluorescence/maximum fluorescence yield). Although climatic factors may have contributed, these results were most likely attributable to an increase in N deposition southwards. For S. fuscum, photosynthetic rate continued to increase up to a deposition level of 1.49 g N m(-2) year(-1), but for S. balticum it seemed to level out at 1.14 g N m(-2) year(-1). The results for S. balticum suggested that transplants from different origin (with low or intermediate N deposition) respond differently to high N deposition. This indicates that Sphagnum species may be able to adapt or physiologically adjust to high N deposition. Our results also suggest that S. balticum might be more sensitive to N deposition than S. fuscum. Surprisingly, NPmax was not (S. balticum), or only weakly (S. fuscum) correlated with biomass production, indicating that production is to a great extent is governed by factors other than the photosynthetic capacity.
Sphagnum, the main genus which forms boreal peat, is strongly affected by N and S deposition and raised temperature, but the physiological mechanisms behind the responses are largely unknown. We measured maximum photosynthetic rate (NPmax), maximum efficiency of photosystem II [variable fluorescence (F v)/maximum fluorescence yield (F m)] and concentrations of N, C, chlorophyll and carotenoids as responses to N and S addition and increased temperature in Sphagnum balticum (a widespread species in the northern peatlands) in a 12-year factorial experiment. NPmax did not differ between control (0.2 g N m−2 year−1) and high N (3.0 g N m−2 year−1), but was higher in the mid N treatment (1.5 g N m−2 year−1). N, C, carotenoids and chlorophyll concentration increased in shoot apices after N addition. F v/F m did not differ between N treatments. Increased temperature (+3.6°C) had a small negative effect on N concentration, but had no significant effect on NPmax or F v/F m. Addition of 2 g S m−2 year−1 showed a weak negative effect on NPmax and F v/F m. Our results suggest a unimodal response of NPmax to N addition and tissue N concentration in S. balticum, with an optimum N concentration for photosynthetic rate of ~13 mg N g−1. In conclusion, high S deposition may reduce photosynthetic capacity in Sphagnum, but the negative effects may be relaxed under high N availability. We suggest that previously reported negative effects on Sphagnum productivity under high N deposition are not related to negative effects on the photosynthetic apparatus, but differences in optimum N concentration among Sphagnum species may affect their competitive ability under different N deposition regimes.
Ecological stoichiometry has been successful in enhancing our understanding of trophic interactions between consumer and prey species. Consumer and prey dynamics have been shown to depend on the nutrient composition of the prey relative to the nutrient demand of the consumer. Since most experiments on this topic used a single consumer species, little is known about the validity of stoichiometric constraints on trophic interactions across consumers and ecosystems. We conducted a quantitative meta-analysis on grazer-periphyton experiments to test (1) if benthic grazers have consistent effects on the nutrient composition of their prey, and (2) whether these effects can be aligned to the nutrient stoichiometry of grazer and periphyton, other environmental factors, or experimental constraints. Grazers significantly lowered periphyton C:N and C:P ratios, indicating higher N- and P-content of grazed periphyton across studies. Grazer presence on average increased periphyton N:P ratios, but across studies the effect size did not differ significantly from zero. The sign and strength of grazer effects on periphyton nutrient ratios was strongly dependent on the nutrient content of grazers and their food, but also on grazer biomass, the amount of biomass removal and water column nutrients. Grazer with low P-content tended to reduce periphyton P-content, whereas grazers with high P-content increased periphyton P-content. This result suggests that low grazer P-content can be an indication of physiological P-limitation rather than a result of having relatively low and fixed P-requirements. At the across-system scale of this meta-analysis, predictions from stoichiometric theory are corroborated, but the plasticity of the consumer nutrient composition has to be acknowledged.
While phenotypic responses to direct species interactions are well studied, we know little about the consequences of indirect interactions for phenotypic divergence.In this study we used lakes with and without the zebra mussel to investigate effects ofindirect trophic interactions on phenotypic divergence between littoral and pelagic perch. We found a greater phenotypic divergence between littoral and pelagic individuals inlakes with zebra mussels and propose a mussel-mediated increase in pelagic and benthic resource availability as a major factor underlying this divergence. Lakes withzebra mussels contained higher densities of large plankton taxa and large invertebrates. We suggest that this augmented resource availability improved perch foraging opportunities in both the littoral and pelagic zones. Perch in both habitats could hence express a more specialized foraging morphology, leading to an increased divergence of perch forms in lakes with zebra mussels. As perch do not prey on mussels directly, we conclude that the increased divergence results from indirect interactions with the mussels. Our results hence suggest that species at lower food web levels can indirectlyaffect phenotypic divergence in species at the top of the food chain.
Environmental variation connected with seasonality is likely to affect the evolution of life-history strategies in ectotherms, but there is no consensus as to how important life-history traits like body size are influenced by environmental variation along seasonal gradients. We compared adult body size, skeletal growth, mean age, age at first reproduction and longevity among 11 common frog (Rana temporaria) populations sampled along a 1,600-km-long latitudinal gradient across Scandinavia. Mean age, age at first reproduction and longevity increased linearly with decreasing growth season length. Lifetime activity (i.e. the estimated number of active days during life-time) was highest at mid-latitudes and females had on average more active days throughout their lives than males. Variation in body size was due to differences in lifetime activity among populations-individuals (especially females) were largest where they had the longest cumulative activity period-as well as to differences between populations in skeletal growth rate as determined by skeletochronological analyses. Especially, males grew faster at intermediate latitudes. While life-history trait variation was strongly associated with latitude, the direction and shape of these relationships were sex- and trait-specific. These context-dependent relationships may be the result of life-history trade-offs enforced by differences in future reproductive opportunities and time constraints among the populations. Thus, seasonality appears to be an important environmental factor shaping life-history trait variation in common frogs.
Climate change was simulated by increasing temperature and nutrient availability in an alpine landscape. We conducted a field experiment of BACI-design (before/after control/impact) running for five seasons in two alpine communities (heath and meadow) with the factors temperature (increase of ca. 1.5-3.0A degrees C) and nutrients (5 g N, 5 g P per m(2)) in a fully factorial design in northern Swedish Lapland. The response variables were abundances of plant species and functional types. Plant community responses to the experimental perturbations were investigated, and the responses of plant functional types were examined in comparison to responses at the species level. Nutrient addition, exclusively and in combination with enhanced temperature increase, exerted the most pronounced responses at the species-specific and community levels. The main responses to nutrient addition were increases in graminoids and forbs, whereas deciduous shrubs, evergreen shrubs, bryophytes, and lichens decreased. The two plant communities of heath or meadow showed different vegetation responses to the environmental treatments despite the fact that both communities were located on the same subarctic-alpine site. Furthermore, we showed that the abundance of forbs increased in response to the combined treatment of temperature and nutrient addition in the meadow plant community. Within a single-plant functional type, most species responded similarly to the enhanced treatments although there were exceptions, particularly in the moss and lichen functional types. Plant community structure showed BACI responses in that vegetation dominance relationships in the existing plant functional types changed to varying degrees in all plots, including control plots. Betula nana and lichens increased in the temperature-increased enhancements and in control plots in the heath plant community during the treatment period. The increases in control plots were probably a response to the observed warming during the treatment period in the region.
Local flower density can affect pollen limitation and plant reproductive success through changes in pollinator visitation and availability of compatible pollen. Many studies have investigated the relationship between conspecific density and pollen limitation among populations, but less is known about within-population relationships and the effect of heterospecific flower density. In addition, few studies have explicitly assessed how the spatial scales at which flowers are monitored affect relationships. We investigated the effect of floral neighborhood on pollen limitation at four spatial scales in the self-incompatible herbs Armeria maritima spp. maritima and Ranunculus acris spp. acris. Moreover, we measured pollen deposition in Armeria and pollinator visits to Ranunculus. There was substantial variation in pollen limitation among Armeria individuals, and 25% of this variation was explained by the density of compatible and heterospecific flowers within a 3 m circle. Deposition of compatible pollen was affected by the density of compatible and incompatible inflorescences within a 0.5 m circle, and deposition of heterospecific pollen was affected by the density of heterospecific flowers within a 2 m circle. In Ranunculus, the number of pollinator visits was affected by both conspecific and heterospecific flower densities. This did not, however, result in effects of the floral neighborhood on pollen limitation, probably due to an absence of pollen limitation at the population level. Our study shows that considerable variation in pollen limitation may occur among individuals of a population, and that this variation is partly explained by floral neighborhood density. Such individual-based measures provide an important link between pollen limitation theory, which predicts ecological and evolutionary causes and consequences for individual plants, and studies of the effects of landscape fragmentation on plant species persistence. Our study also highlights the importance of considering multiple spatial scales to understand the spatial extent of pollination processes within a population.
Invasive plants may compete with native species for abiotic factors as light, space and nutrients, and have also been shown to affect native pollination interactions. Studies have mainly focused on how invasive plants affect pollinator behaviour, i.e. attraction of pollinators to or away from native flowers. However, when an invasive plant provides resources utilized by native pollinators this could increase pollinator population sizes and thereby pollination success in natives. Effects mediated through changes in pollinator population sizes have been largely ignored in previous studies, and the dominance of negative interactions suggested by meta-analyses may therefore be biased. We investigated the impact of the invasive Lupinus polyphyllus on pollination in the native Lotus corniculatus using a study design comparing invaded and uninvaded sites before and after the flowering period of the invasive. We monitored wild bee abundance in transects, and visit rate and seed production of potted Lotus plants. Bumblebee abundance increased 3.9 times in invaded sites during the study period, whereas it was unaltered in uninvaded sites. Total visit rate per Lotus plant increased 2.1 times in invaded sites and decreased 4.4 times in uninvaded sites. No corresponding change in seed production of Lotus was found. The increase in visit rate to Lotus was driven by an increase in solitary bee visitation, whereas mainly bumblebees were observed to visit the invasive Lupinus. The mechanism by which the invasive increases pollinator visit rates to Lotus could be increased availability of other flower resources for solitary bees when bumblebees forage on Lupinus.
Marginal grassland fragments, such as road verges and field margins, may act as important supplemental habitats for grassland plants in the modern agricultural landscape. However, abundance of pollinators in such fragments has been found to decline with distance to larger natural and semi-natural habitats, and this could have corresponding effects on plant pollination. In this study, we performed a field experiment on road verges with three insect-pollinated grassland herbs to examine the relationship between distance to semi-natural grassland and plant reproductive success in two landscapes with contrasting farming intensities. In Lychnis viscaria and Lotus corniculatus, seed production tended to decrease with increasing distance to semi-natural grassland, but only in the landscape with high farming intensity. Seed production in Armeria maritima spp. maritima decreased with distance in both landscapes. Although many studies have investigated effects of natural habitat on crop pollination, little is known about the impact on pollination in native plants. The results from this study indicate that management of semi-natural grasslands improves not only biodiversity within the actual grassland but also pollination of native plants in the surrounding agricultural landscape.
Foraging differentiation within a species can contribute to restricted gene flow between ecologically different groups, promoting ecological speciation. Galapagos sea lions (Zalophus wollebaeki) show genetic and morphological divergence between the western and central archipelago, possibly as a result of an ecologically mediated contrast in the marine habitat. We use global positioning system (GPS) data, time-depth recordings (TDR), stable isotope and scat data to compare foraging habitat characteristics, diving behaviour and diet composition of Galapagos sea lions from a western and a central colony. We consider both juvenile and adult life stages to assess the potential role of ontogenetic shifts that can be crucial in shaping foraging behaviour and habitat choice for life. We found differences in foraging habitat use, foraging style and diet composition that aligned with genetic differentiation. These differences were consistent between juvenile and adult sea lions from the same colony, overriding age-specific behavioural differences. Our study contributes to an understanding of the complex interaction of ecological condition, plastic behavioural response and genetic make-up of interconnected populations.
Variation in seasonal time constraints and temperature along latitudinal gradients are expected to select for life history trait differentiation, but information about the relative importance of these factors in shaping patterns of divergence in embryonic traits remains sparse. We studied embryonic survival, growth and development rates in the common frog (Rana temporaria) along a 1,400-km latitudinal gradient across Sweden by raising embryos from four populations in the laboratory at seven temperatures (9 degrees C, 12 degrees C, 15 degrees C, 18 degrees C, 21 degrees C, 24 degrees C, 27 degrees C). We found significant differences in mean values of all traits between the populations and temperature treatments, but this variation was not latitudinally ordered. In general, embryonic survival decreased at the two highest temperatures in all populations, but less so in the southernmost as compared to the other populations. The northernmost population developed slowest at the lowest temperature, while the two mid-latitude populations were slowest at the other temperatures. Hatchling size increased with increasing temperature especially in the two northern populations, whereas the two southern populations showed peak hatchling size at 15 degrees C. Analyses of within-population genetic variation with a half-sib design revealed that there was significant additive genetic variation in all traits, and egg size-related maternal effects were important in the case of hatchling size. Overall, our results indicate that unlike larval growth and development, variation in embryonic development and growth in R. temporaria cannot be explained in terms of a latitudinal gradient in season length. While adaptation to a latitudinal variation in temperature might have contributed to the observed differentiation in embryonic performance, the effects of other, perhaps more local environmental factors, seem to have overridden them in importance.
Differences among plant species in visitation rate and seed set within a community may be explained both by the species' floral traits and the community context. Additionally, the importance of species' floral traits vs. community context on visitation rate and seed set may vary among communities. In communities where the pollinator-to-flower ratio is low, floral traits may be more important than community context, as pollinators may have the opportunity to be choosier when visiting plant species. In this study we investigated whether species' floral traits (flower shape, size and number, and flowering duration) and community context (conspecific and heterospecific flower density, and pollinator abundance) could explain among-species variation in visitation rate and seed set. For this, we used data on 47 plant species from two Norwegian plant communities differing in pollinator-to-flower ratio. Differences among species in visitation rate and seed set within a community could be explained by similar variables as those explaining visitation rate and seed set within species. As expected, we found floral traits to be more important than community context in the community with a lower pollinator-to-flower ratio; whereas in the community with a higher pollinator-to-flower ratio, community context played a bigger role. Our study gives significant insights into the relative importance of floral traits on species' visitation rate and seed set, and contributes to our understanding of the role of the community context on the fitness of plant species.
Herbivory can negatively affect several components of plant reproduction. Yet, because of a lack of experimental studies involving multiple populations, the extent to which differences in herbivory contribute to among-population variation in plant reproductive success is poorly known. We determined experimentally the effects of insect herbivory on reproductive output in nine natural populations of the perennial herb Lythrum salicaria along a disturbance gradient in an archipelago in northern Sweden, and we quantified among-population differentiation in resistance to herbivory in a common-garden experiment in the same area. The intensity of leaf herbivory varied >500-fold and mean female reproductive success >400-fold among the study populations. The intensity of herbivory was lowest in populations subject to strong disturbance from ice and wave action. Experimental removal of insect herbivores showed that the effect of herbivory on female reproductive success was correlated with the intensity of herbivory and that differences in insect herbivory could explain much of among-population variation in the proportion of plants flowering and seed production. Population differentiation in resistance to herbivory was limited. The results demonstrate that the intensity of herbivory is a major determinant of flowering and seed output in L. salicaria, but that differences in herbivory are not associated with differences in plant resistance at the spatial scale examined. They further suggest that the physical disturbance regime may strongly influence the performance and abundance of perennial herbs not only because of its effect on interspecific competition, but also because of effects on interactions with specialized herbivores.
The potential interactions of grazing, nutrients and light in influencing autotroph species diversity have not previously been considered. Earlier studies have shown that grazing and nutrients interact in determining autotroph species diversity, since grazing decreases species diversity when nutrients (i.e. N or P) limit autotroph growth, but increases it when nutrients are replete. We hypothesized that increased light intensities would intensify the interactions between grazing and nutrients on algal species diversity, resulting in even stronger reductions in algal species diversity through grazing under nutrient–poor conditions, and to even stronger increases of algal species diversity through grazing under nutrient-rich conditions. We studied the effects of grazing (absent, present), nutrients (ambient, N + P enriched) and light (low light, high light) on benthic algal diversity and periphyton C:nutrient ratios (which can indicate algal nutrient limitation) in a factorial laboratory experiment, using the gastropod grazer Viviparus viviparus. Grazing decreased algal biomass and algal diversity, but increased C:P and N:P ratios of periphyton. Grazing also affected periphyton species composition, by decreasing the proportion of Spirogyra sp. and increasing the proportion of species in the Chaetophorales. Grazing effects on diversity as well as on periphyton N:P ratios were weakened when nutrients were added (interaction between grazing and nutrients). Chlorophyll a (Chl a) per area increased with nutrient addition and decreased with high light intensities. Light did not increase the strength of the interaction between grazing and nutrients on periphytic algal diversity. This study shows that nutrient addition substantially reduced the negative effects of grazing on periphytic algal diversity, whereas light did not interact with grazing or nutrient enrichment in determining periphytic algal diversity.
Alpine plant communities are predicted to face range shifts and possibly extinctions with climate change. Fine-scale environmental variation such as nutrient availability or snowmelt timing may contribute to the ability of plant species to persist locally; however, variation in nutrient availability in alpine landscapes is largely unmeasured. On three mountains around Davos, Switzerland, we deployed Plant Root Simulator probes around 58 Salix herbacea plants along an elevational and microhabitat gradient to measure nutrient availability during the first 5 weeks of the summer growing season, and used in situ temperature loggers and observational data to determine date of spring snowmelt. We also visited the plants weekly to assess performance, as measured by stem number, fruiting, and herbivory damage. We found a wide snowmelt gradient which determined growing season length, as well as variations of an order of magnitude or more in the accumulation of 12 nutrients between different microhabitats. Higher nutrient availability had negative effects on most shrub performance metrics, for instance decreasing stem number and the proportion of stems producing fruits. High nutrient availability was associated with increased herbivory damage in early-melting microhabitats, but among late-emerging plants this pattern was reversed. We demonstrate that nutrient availability is highly variable in alpine settings, and that it strongly influences performance in an alpine dwarf shrub, sometimes modifying the response of shrubs to snowmelt timing. As the climate warms and human-induced nitrogen deposition continues in the Alps, these factors may contribute to patterns of local plants persistence.
Host trees for obligate epiphytes are dynamic patches that emerge, grow and fall, and metacommunity diversity critically depends on efficient dispersal. Even though species that disperse by large asexual diaspores are strongly dispersal-limited, asexual dispersal is common. The stronger dispersal limitation of asexually reproducing species compared to species reproducing sexually via small spores may be compensated by higher growth rates, lower sensitivity to habitat conditions, higher competitive ability or younger reproductive age.
We compared growth and reproduction of different groups of epiphytic bryophytes with contrasting dispersal (asexual vs sexual) and life-history strategies (colonists, short- and long-lived shuttle species, perennial stayers) in an old-growth forest stand in the boreo-nemoral region in Eastern Sweden.
No differences were seen in relative growth rates between asexual and sexual species. Long-lived shuttles had lower growth rates than colonists and perennial stayers. Most groups grew best at intermediate bark pH. Interactions with other epiphytes had a small, often positive effect on growth. Neither differences in sensitivity of growth to habitat conditions nor differences in competitive abilities among species groups were found. Habitat conditions, however, influenced the production of sporophytes, but not of asexual diaspores. Presence of sporophytes negatively affected growth, whereas presence of asexual diaspores did not. Sexual species had to reach a certain colony size before starting to reproduce, whereas no such treshold existed for asexual reproduction.
The results indicate that the epiphyte metacommunity is structured by two main trade-offs: dispersal distance vs reproductive age, and dispersal distance vs sensitivity to habitat quality. There seems to be a trade-off between growth and sexual reproduction, but not asexual. Trade-offs in species traits may be shaped by conflicting selection pressures imposed by habitat turnover and connectivity rather than by species interactions.
Diaspore banks are crucial for the maintenance and resilience of plant communities, but diaspore banks of bryophytes remain poorly known, especially from tropical ecosystems. This is the first study to focus on the role of diaspore banks of bryophytes in tropical rain forests. Our aim was to test whether microhabitat (substrate type) and species traits (breeding system, phylum) are important in explaining the diaspore bank composition. Using samples cultivated in the laboratory, we assessed the number of species and shoots emerging from bark, decaying wood and soil from two sites of the Atlantic rain forest (montane and sea level) in Brazil by comparing the contribution of species by phylum (mosses, liverworts) and breeding system (monoicous, dioicous). More species emerged from bark (68) and decaying wood (55) than from soil (22). Similar numbers of species were found at both sites. Mosses were more numerous in terms of number of species and shoots, and monoicous species dominated over dioicous species. Substrate pH had only weak effects on shoot emergence. Species commonly producing sporophytes and gemmae had a high contribution to the diaspore banks. These superficial diaspore banks represented the extant vegetation rather well, but held more monoicous species (probably short-lived species) compared to dioicous ones. We propose that diaspore bank dynamics are driven by species traits and microhabitat characteristics, and that short-term diaspore banks of bryophytes in tropical rain forests contribute to fast (re) establishment of species after disturbances and during succession, particularly dioicous mosses investing in asexual reproduction and monoicous mosses investing in sexual reproduction.
Although it is well-documented from theoretical studies that pathogens have the capacity to generate cycles, the occurrence and role of pathogens and disease have been poorly empirically studied in cyclic voles and lemmings. In screening for the occurrence of disease in cyclic vole and lemming populations, we found that a high proportion of live-trapped Clethrionomys glareolus, C. rufocanus, Microtus agrestis and Lemmus lemmus at high collective peak density, shortly before the decline, suffered from diabetes or myocarditis in northern Scandinavia. A high frequency of animals had abnormal blood glucose (BG) levels at the time of trapping (5-33%). In contrast, C. rufocanus individuals tested at a much lower overall density, and at an earlier stage relative to the decline in the following cycle, showed normal BG concentrations. However, a high proportion (43%) of a sample of these individuals kept in captivity developed clinical diabetes within five weeks, as determined by BG levels and a glucose tolerance test performed at that later time. A new picornavirus isolated from the rodents, Ljungan virus (LV), was assumed to cause the diseases, as LV-induced diabetes and myocarditis, as well as encephalitis and fetal deaths, were observed in laboratory mice. We hypothesize that LV infection significantly affects morbidity and mortality rates in the wild, either directly or indirectly, by predisposing the rodents to predation, and is at least involved in causing the regular, rapid population declines of these cyclic voles and lemmings. Increased stress at peak densities is thought to be an important trigger for the development of disease, as the occurrence of disease in laboratory mice has been found to be triggered by introducing stress to LV-infected animals.
Antipredator behaviour is an important fitness component in most animals. A co-evolutionary history between predator and prey is important for prey to respond adaptively to predation threats. When non-native predator species invade new areas, native prey may not recognise them or may lack effective antipredator defences. However, responses to novel predators can be facilitated by chemical cues from the predators' diet. The red swamp crayfish Procambarus clarkii is a widespread invasive predator in the Southwest of the Iberian Peninsula, where it preys upon native anuran tadpoles. In a laboratory experiment we studied behavioural antipredator defences (alterations in activity level and spatial avoidance of predator) of nine anurans in response to P. clarkii chemical cues, and compared them with the defences towards a native predator, the larval dragonfly Aeshna sp. To investigate how chemical cues from consumed conspecifics shape the responses, we raised tadpoles with either a tadpole-fed or starved crayfish, or dragonfly larva, or in the absence of a predator. Five species significantly altered their behaviour in the presence of crayfish, and this was largely mediated by chemical cues from consumed conspecifics. In the presence of dragonflies, most species exhibited behavioural defences and often these did not require the presence of cues from predation events. Responding to cues from consumed conspecifics seems to be a critical factor in facilitating certain behavioural responses to novel exotic predators. This finding can be useful for predicting antipredator responses to invasive predators and help directing conservation efforts to the species at highest risk.
Spatial and temporal heterogeneity in the environment is a common feature affecting many natural populations. For example, both the resource levels and optimal habitat choices of individuals likely change over time. One way for organisms to cope with environmental variation is to display adaptive plasticity in traits such as behavior and morphology. Since trait plasticity is hypothesized to be a prerequisite for character divergence, studies of mechanisms behind such plasticity are warranted. In this study, we looked at the interaction of two potentially important environmental variables on behavioral and morphological plasticity in Eurasian perch (Perca fluviatilis L.). More specifically, the plastic responses in activity and morphology of perch exposed to different resource levels and simulated habitat types were studied in an aquarium experiment. The resource level experienced had a large influence on plasticity in both activity and morphology. Behavioral adaptations have been thought to mediate morphological transitions, and we suggest that the morphological response to the resource level was mediated by differences in activity and growth rates. The habitat type also affected morphological plasticity but to a lesser extent, and there was no effect on activity from habitat type. Based on these results, we suggest that it is essential to include several environmental factors acting in concert when studying mechanisms behind trait plasticity. We also propose that variation in resource levels might play a key role in fostering trait plasticity in at least fish populations, while other environmental variables such as divergent habitat complexities and prey types might be less influential. Dynamics in resource levels and optimal habitat choices might thus be important factors influencing character divergence in natural populations.
Seasonal time constraints can pose strong selection on life histories. Time-constrained animals should prioritise fast development over predation risk to avoid unfavourable growing conditions. However, changes in phenology could alter the balance between anti-predator and developmental needs. We studied variation of anti-predator strategies in common frog (Rana temporaria) tadpoles in four populations from the two extremes of a latitudinal gradient across Sweden. We examined, under common conditions in the laboratory, the anti-predator responses and life histories of tadpoles raised with predatory Aeshna dragonfly larvae in two consecutive years with a difference of 20 days in breeding time in the north, but no difference in breeding time in the nouth. In a year with late breeding, northern tadpoles did not modify their behaviour and morphology in the presence of predators, and metamorphosed faster and smaller than tadpoles born in a year with early breeding. In the year with early breeding, northern tadpoles showed a completely different anti-predator strategy by reducing activity and developing morphological defences in the presence of predators. We discuss the possible mechanisms that could activate these responses (likely a form of environmentally-mediated parental effect). To our knowledge, this is the first study to show that a vertebrate modifies the anti-predator strategy of its offspring in response to natural variation in reproductive phenology, which highlights the need to consider phenology in studies of life-history evolution.
Organisms normally grow at a sub-maximal rate. After experiencing a period of arrested growth, individuals often show compensatory growth responses by modifying their life-history, behaviour and physiology. However, the strength of compensatory responses may vary across broad geographic scales as populations differ in their exposition to varying time constraints. We examined differences in compensatory growth strategies in common frog (Rana temporaria) populations from southern and northern Sweden. Tadpoles from four populations were reared in the laboratory and exposed to low temperature to evaluate the patterns and mechanisms of compensatory growth responses. We determined tadpoles' growth rate, food intake and growth efficiency during the compensation period. In the absence of arrested growth conditions, tadpoles from all the populations showed similar (size-corrected) growth rates, food intake and growth efficiency. After being exposed to low temperature for 1 week, only larvae from the northern populations increased growth rates by increasing both food intake and growth efficiency. These geographic differences in compensatory growth mechanisms suggest that the strategies for recovering after a period of growth deprivation may depend on the strength of time constraints faced by the populations. Due to the costs of fast growth, only populations exposed to the strong time constraints are prone to develop fast recovering strategies in order to metamorphose before conditions deteriorate. Understanding how organisms balance the cost and benefits of growth strategies may help in forecasting the impact of fluctuating environmental conditions on life-history strategies of populations likely to be exposed to increasing environmental variation in the future.
Herbivores may damage both leaves and reproductive structures, and although such combined damage may affect plant fitness non-additively, this has received little attention. We conducted a 2-year field experiment with a factorial design to examine the effects of simulated leaf (0,12.5, 25, or 50% of leaf area removed) and inflorescence damage (0 vs. 50% of inflorescences removed) on survival, growth and reproduction in the perennial herb Arabidopsis lyrata. Leaf and inflorescence damage negatively and independently reduced flower, fruit and seed production in the year of damage; leaf damage also reduced rosette size by the end of the first season and flower production in the second year. Leaf damage alone reduced the proportion of flowers forming a fruit and fruit production per plant the second year, but when combined with inflorescence damage no such effect was observed (significant leaf × inflorescence damage interaction). Damage to leaves (sources) caused a greater reduction in future reproduction than did simultaneous damage to leaves and inflorescences (sinks). This demonstrates that a full understanding of the effects of herbivore damage on plant fitness requires that consequences of damage to vegetative and reproductive structures are evaluated over more than 1 year and that non-additive effects are considered.
Landscape characteristics and social behavior can affect the foraging patterns of seed-dependent animals. We examine the movement of acorns from valley oak (Quercus lobata) trees to granaries maintained by acorn woodpeckers (Melanerpes formicivorus) in two California oak savanna-woodlands differing in the distribution of Q. lobata within each site. In 2004, we sampled Q. lobata acorns from 16 granaries at Sedgwick Reserve in Santa Barbara County and 18 granaries at Hastings Reserve in Monterey County. Sedgwick has lower site-wide density of Q. lobata than Hastings as well as different frequencies of otherQuercus species common to both sites. We found acorn woodpeckers foraged from fewer Q. lobata seed source trees (K g = 4.1 ± 0.5) at Sedgwick than at Hastings (K g = 7.6 ± 0.6) and from fewer effective seed sources (N em* = 2.00 and 5.78, respectively). The differences between sites are due to a greater number of incidental seed sources used per granary at Hastings than at Sedgwick. We also found very low levels of seed source sharing between adjacent granaries, indicating that territoriality is strong at both sites and that each social group forages on its own subset of trees. We discovered an interesting spatial pattern in the location of granaries. At Sedgwick, acorn woodpeckers situated their granaries within areas of higher-than-average tree density, while at Hastings, they placed them within areas of lower-than-average tree density, with the outcome that granaries at the two sites were located in areas of similar valley oak density. Our results illustrate that landscape characteristics might influence the number of trees visited by acorn woodpeckers and the locations of territories, while woodpecker social behavior, such as territoriality, shapes which trees are visited and whether they are shared with other social groups.
Environmental stress, such as acidification, can challenge persistence of natural populations and act as a powerful evolutionary force at ecological time scales. The ecological and evolutionary responses of natural populations to environmental stress at early life-stages are often mediated via maternal effects. During early life-stages, maternal effects commonly arise from egg coats (the extracellular structures surrounding the embryo), but the role of egg coats has rarely been studied in the context of adaptation to environmental stress. Previous studies on the moor frog Rana arvalis found that the egg coat mediated adaptive divergence along an acidification gradient in embryonic acid stress tolerance. However, the exact mechanisms underlying these adaptive maternal effects remain unknown. Here, we investigated the role of water balance and charge state (zeta potential) of egg jelly coats in embryonic adaptation to acid stress in three populations of R. arvalis. We found that acidic pH causes severe water loss in the egg jelly coat, but that jelly coats from an acid-adapted population retained more water than jelly coats from populations not adapted to acidity. Moreover, embryonic acid tolerance (survival at pH 4.0) correlated with both water loss and charge state of the jelly, indicating that negatively charged glycans influence jelly water balance and contribute to embryonic adaptation to acidity. These results indicate that egg coats can harbor extensive intra-specific variation, probably facilitated in part via strong selection on water balance and glycosylation status of egg jelly coats. These findings shed light on the molecular mechanisms of environmental stress tolerance and adaptive maternal effects.
A cost of reproduction in terms of reduced future performance underlies all life-history models, yet costs have been difficult to detect in short-term experiments with long-lived plants. The likelihood of detecting costs should depend on the range of variation in reproductive effort that can be induced, and also on the shape of the cost function across this range, which should be affected by resource availability. Here, we experimentally examined the effects of both reduced and increased fruit production in two populations of the long-lived orchid Gymnadenia conopsea located at sites that differ in length of the growing season. Plants that were prevented from fruiting produced more flowers in the population with a longer growing season, had higher survival in the other population, and grew larger compared to control plants in both populations. Fruit production was pollen-limited in both populations, and increased reproductive investment after supplemental hand-pollination was associated with reduced fecundity the following year. The results demonstrate that the shape of the cost function varies among fitness components, and that costs can be differentially expressed in different populations. They are consistent with the hypothesis that differences in temporal overlap between allocation to reproduction and other functions will induce among-population variation in reproductive costs.
To evaluate whether leaf spot disease and related effects on photosynthesis are influenced by increased nitrogen (N) input and elevated atmospheric CO2 concentration ([CO2]), we examined disease incidence and photosynthetic rate of Solidago rigida grown in monoculture under ambient or elevated (560 mu mol mol(-1)) [CO2] and ambient or elevated (+4 g N m(-2) year(-1)) N conditions in a field experiment in Minnesota, USA. Disease incidence was lower in plots with either elevated [CO2] or enriched N (-57 and -37%, respectively) than in plots with ambient conditions. Elevated [CO2] had no significant effect on total plant biomass, or on photosynthetic rate, but reduced tissue%N by 13%. In contrast, N fertilization increased both biomass and total plant N by 70%, and as a consequence tissue%N was unaffected and photosynthetic rate was lower on N fertilized plants than on unfertilized plants. Regardless of treatment, photosynthetic rate was reduced on leaves with disease symptoms. On average across all treatments, asymptomatic leaf tissue on diseased leaves had 53% lower photosynthetic rate than non-diseased leaves, indicating that the negative effect from the disease extended beyond the visual lesion area. Our results show that, in this instance, indirect effects from elevated [CO2], i.e., lower disease incidence, had a stronger effect on realized photosynthetic rate than the direct effect of higher [CO2].
Increased levels of dissolved carbon dioxide (CO2) drive ocean acidification and have been predicted to increase the energy use of marine fishes via physiological and behavioural mechanisms. This notion is based on a theoretical framework suggesting that detrimental effects on energy use are caused by plasma acid-base disruption in response to hypercapnic acidosis, potentially in combination with a malfunction of the gamma aminobutyric acid type A (GABA(A)) receptors in the brain. However, the existing empirical evidence testing these effects primarily stems from studies that exposed fish to elevated CO2 for a few days and measured a small number of traits. We investigated a range of energetic traits in juvenile spiny chromis damselfish (Acanthochromis polyacanthus) over 3months of acclimation to projected end-of-century CO2 levels (similar to 1000 mu atm). Somatic growth and otolith size and shape were unaffected by the CO2 treatment across 3months of development in comparison with control fish (similar to 420 mu atm). Swimming activity during behavioural assays was initially higher in the elevated CO2 group, but this effect dissipated within similar to 25min following handling. The transient higher activity of fish under elevated CO2 was not associated with a detectable difference in the rate of oxygen uptake nor was it mediated by GABA(A) neurotransmitter interference because treatment with a GABA(A) antagonist (gabazine) did not abolish the CO2 treatment effect. These findings contrast with several short-term studies by suggesting that end-of-century levels of CO2 may have negligible direct effects on the energetics of at least some species of fish.
Among-individual diet variation is common in natural populations and may occur at any trophic level within a food web. Yet, little is known about its variation among trophic levels and how such variation could affect phenotypic divergence within populations. In this study we investigate the relationships between trophic position (the population's range and average) and among-individual diet variation. We test for diet variation among individuals and across size classes of Eurasian perch (Perca fluviatilis), a widespread predatory freshwater fish that undergoes ontogenetic niche shifts. Second, we investigate among-individual diet variation within fish and invertebrate populations in two different lake communities using stable isotopes. Third, we test potential evolutionary implications of population trophic position by assessing the relationship between the proportion of piscivorous perch (populations of higher trophic position) and the degree of phenotypic divergence between littoral and pelagic perch sub-populations. We show that among-individual diet variation is highest at intermediate trophic positions, and that this high degree of among-individual variation likely causes an increase in the range of trophic positions among individuals. We also found that phenotypic divergence was negatively related to trophic position in a population. This study thus shows that trophic position is related to and may be important for among-individual diet variation as well as to phenotypic divergence within populations.
Whether fluctuating asymmetry (FA) provides a useful metric indicator of the degree of environmental stress experienced by populations is still a contentious issue. We investigated whether the degree of FA in skeletal elements is useful in elucidating the degree of environmental stress experienced by frog populations, and further, tested the proposition that a trait's sensitivity to stress—as reflected in the degree of FA—is related to the degree of directional selection experienced by the given trait. We compared the degree of FA in four bilateral skeletal elements of male and female moor frogs (Rana arvalis) originating from low (acidified) and neutral pH populations. While the degree of uncorrected FA was unrelated to the degree of acidity, the growth rate and age of the individuals, the size-corrected FA was significantly higher in low than in neutral pH populations and decreased with individual ages and growth rates. In addition, both measures of FA were significantly higher in males and in particular in traits presumably under high sexual selection as indicated by the degree of sexual size dimorphism. All in all, the results indicate that individuals from acidified localities are smaller, younger and exhibit a significantly higher degree of FA than individuals from neutral pH populations. These results constitute the first assessment of FA in amphibians and suggest that the degree of FA in skeletal traits can be a useful indicator of the degree of environmental stress experienced by amphibian populations.
Recent advances in stochastic demography provide tools to examine the importance of random and periodic variation in vital rates for population dynamics. In this study, we explore with simulations the effect of disturbance regime on population dynamics and viability. We collected 7 years of demographic data in three populations of the perennial herb Primula farinosa, and used these data to examine how variation in vital rates affected population viability parameters (stochastic growth rate, lambda(S)), and how vital rates were related to weather conditions. Elasticity analysis indicated that the stochastic growth rate was very sensitive to changes in regeneration, quantified as the production, survival, and germination of seeds. In one of the study years, all seedlings and mature plants in the demography plots died. This extinction coincided with the driest summer during the study period. Simulations suggested that a future increase in the frequency of high-mortality years due to climate change would result in reduced population growth rate, and an increased importance of survival in the seed bank for population viability. The results illustrate how the limited demographic data typically available for many natural systems can be used in simulation models to assess how environmental change will affect population viability.
Changes in growing season temperature and duration may have profound effects on the population dynamics of arctic and alpine plant species in snow-bed and fell-field habitats. We examined how a typical herbaceous pioneer species, Ranunculus glacialis, responded to experimental climate change in open-top chambers for three seasons at an alpine site in southern Norway. Warming had no significant effect on any reproductive, growth or phenological variables, except for seed weight, which increased significantly during the first 2 ears. Despite large differences in average date of snow-melt among years, average reproductive output and ramet size differed little among years. Within-year variation in date of snowmelt had no impact on seed number or weight in either control or warmed plots. Leaf width and ramet leaf number decreased significantly with later snowmelt within a year. Experimental warming reduced the negative effect on ramet size of late snowmelt within a year to some extent. In general, R. glacialis reacts contrary to most other arctic/alpine species to experimental warming. Species with such low responsiveness to environmental conditions may be particularly vulnerable to climatic change, especially if their habitat is invaded by other species with higher phenotypic plasticity and a better competitive ability.
Spatio-temporal variation in seed predation may strongly influence both plant population dynamics and selection on plant traits. The intensity of seed predation may depend on a number of factors, but the relative importance of previous predator abundance ("local legacy"), spatial distribution of the host plant, environmental factors and plant characteristics has been explored in few species. We monitored seed predation in the perennial herb Primula farinosa, which is dimorphic for scape length, during 5 consecutive years, in a 10-km × 4-km area comprising 79 P. farinosa populations. A transplant experiment showed that the seed predator, the oligophagous tortricid moth Falseuncaria ruficiliana, was not dispersal limited at the spatial scale corresponding to typical distances between P. farinosa populations. Correlations between population characteristics and incidence and intensity of seed predation varied among years. The incidence of the seed predator was positively correlated with host population size and mean number of flowers, while intensity of seed predation in occupied patches was positively related to the frequency of the long-scaped morph in 2 years and negatively related to host population size in 1 year. In both scape morphs, predation tended to increase with increasing frequency of the long morph. There was no evidence of a local legacy; incidence and intensity of seed predation were not related to the abundance of the seed predator in the population in the previous year. Taken together, the results indicate that among-population variation in seed predation intensity is determined largely by patch selection and that the seed predator's preference for tall and many-flowered inflorescences may not only affect selection on plant traits within host plant populations, but also the overall intensity of seed predation.
Many related species share the same environment and utilize similar resources. This is surprising because based on the principle of competitive exclusion one would predict that the superior competitor would drive the other species to extinction; coexistence is only predicted if interspecific competition is weaker than intraspecific competition. Interspecific competition is frequently reduced by differential resource use, resulting in habitat segregation. In this paper, we use the closely related collared and pied flycatcher to assess the potential of habitat differences to affect interspecific competition through a different mechanism, namely by generating temporal differences in availability of similar food resources between the two species. We found that the tree species composition of the breeding territories of the two species differed, mainly by a higher abundance of coniferous species around nest-boxes occupied by pied flycatchers. The temporal availability of caterpillars was measured using frass traps under four deciduous and two coniferous tree species. Deciduous tree species showed an early and narrow peak in abundance, which contrasted with the steady increase in caterpillar abundance in the coniferous tree species through the season. We subsequently calculated the predicted total caterpillar biomass available in each flycatcher territory. This differed between the species, with biomass decreasing more slowly in pied flycatcher territories. Caterpillar biomass is strongly correlated with the reproductive success of collared flycatchers, but much less so with pied flycatchers. However, caterpillar availability can only partly explain the differences in seasonal decline of reproductive success between the two species; we discuss additional factors that may contribute to this species difference. Overall, our results are consistent with the suggestion that minor habitat differences between these two species may contribute to promoting their coexistence.
Alpine dwarf shrub communities are phenologically linked with snowmelt timing, so early spring exposure may increase risk of freezing damage during early development, and consequently reduce seasonal growth. We examined whether environmental factors (duration of snow cover, elevation) influenced size and the vulnerability of shrubs to spring freezing along elevational gradients and snow microhabitats by modelling the past frequency of spring freezing events. We sampled biomass and measured the size of Salix herbacea, Vaccinium myrtillus, Vaccinium uliginosum and Loiseleuria procumbens in late spring. Leaves were exposed to freezing temperatures to determine the temperature at which 50 % of specimens are killed for each species and sampling site. By linking site snowmelt and temperatures to long-term climate measurements, we extrapolated the frequency of spring freezing events at each elevation, snow microhabitat and per species over 37 years. Snowmelt timing was significantly driven by microhabitat effects, but was independent of elevation. Shrub growth was neither enhanced nor reduced by earlier snowmelt, but decreased with elevation. Freezing resistance was strongly species dependent, and did not differ along the elevation or snowmelt gradient. Microclimate extrapolation suggested that potentially lethal freezing events (in May and June) occurred for three of the four species examined. Freezing events never occurred on late snow beds, and increased in frequency with earlier snowmelt and higher elevation. Extrapolated freezing events showed a slight, non-significant increase over the 37-year record. We suggest that earlier snowmelt does not enhance growth in four dominant alpine shrubs, but increases the risk of lethal spring freezing exposure for less freezing-resistant species.
Grazing reduces litter thickness and vegetation height and may thereby indirectly affect reproductive success and selection on floral characters in plants with prostrate growth. Reductions in litter thickness and vegetation height should influence both the resource status of plants with leaves positioned close to the ground and the significance of inflorescence height for interactions with pollinators and seed predators. We experimentally examined how simulated grazing of surrounding vegetation affected pollen limitation, fruit predation and fecundity of short-scaped and long-scaped Primula farinosa, which differ markedly in floral display and therefore in expected attractiveness to pollinators. Litter removal and pruning of surrounding vegetation increased fruit and seed production per plant in the year of the treatment and the probability of flowering in the following year. Pollen limitation of fruit and seed production was stronger in the short-scaped morph than in the long-scaped morph, but was not significantly affected by litter removal and simulated grazing of surrounding vegetation. Supplemental hand-pollination reduced seed size in the year of the treatment and flowering probability in the second year, and these effects did not differ among scape morphs or grazing treatments. The results suggest that grazing indirectly favours seed production in P. farinosa, mainly because it increases the resource status of plants that escape damage. Contrary to expectation, there was no strong evidence that litter accumulation and tall vegetation increase the severity of pollen limitation or reduce the relative performance of the short-scaped morph.
Intraspecific positive relationships between abundance and occupancy are observed for many species, suggesting that the same processes drive local and regional species dynamics. Two main groups of mechanisms explain this relationship: spatiotemporal variation in local population growth rates due to variation in habitat quality, or dispersal effects that increase occupancy of a species when locally abundant. Several studies show that spatiotemporal variation in population growth rates causes positive abundance-occupancy relationships, but few have shown dispersal effects. It is believed that such effects should be more evident for species whose dispersal is limited, e.g. metapopulations, but those studies are limited. This study investigates abundance-occupancy relationships in three Daphnia metapopulations in rock pools and the degree to which dispersal or habitat quality affect their local abundances and occurrence. Daphnia longispina and Daphnia magna showed positive abundance-occupancy relationships, but not Daphnia pulex. No single ecological factor could explain the abundance-occupancy relationships of any given species. Instead, dispersal processes and rock pool quality (mainly salinity and depth) seem to act together to shape the abundance-occupancy relationships. Such a conclusion is also supported by an immigration experiment in natural rock pools. This study suggests that although positive abundance-occupancy relationships may be commonly found for metapopulations, both dispersal processes and variation in habitat quality can be factors determining the abundance-occupancy relationship of metapopulations experiencing habitat heterogeneity.


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Regarded by many as the "Nobel Prize for Teaching", the Global Teacher Prize is an initiative that rewards teachers that display works of excellence in the field of education. In Portugal, supported by Galp Foundation, the project initiates its second edition and will once again recognise the Portuguese teacher who is chosen as the best teacher of the year, with a prize of 30 thousand Euros.
Galp Foundation believes that investing in education and bringing the issue to the agenda is investing in a more sustainable future. Therefore, GTP focuses attributing merit to the teachers in society, because it is due to them to transform and impact the lives of young people.
Since education is a central pillar in Galp and its Foundation's sustainability strategy, we have embraced this partnership with the Global Teacher Prize from the very first moment. The first Portuguese edition, last year, demonstrated that Portuguese people adhere to projects that value the educational ecosystem.
Teachers are one of the most trusted professions that the Portuguese value.
According to a study by the Varkey GEMS Foundation, Portugal ranks 14th out of 21 countries in a ranking that evaluates the social status of teachers.
Nevertheless, national teachers achieve better results than their counterparts in countries such as Switzerland, Germany, or Italy.
Teachers are placed among the three professions that the Portuguese trust the most, closely following doctors and firemen. According to a study by GfK on the first edition of the Global Teacher Prize, 91% of respondents stated that teachers were important to their school career and 86% acknowledged the importance teachers had in their lives overall. However, 89% of Portuguese people also admit that it is an exhausting profession and almost 70% of parents discourage their children from pursuing a teaching career. 92% of Portuguese people consider a good teacher to be one who can motivate their students.
Galp Foundation considers it essential to invest in initiatives that contribute to the appreciation and recognition of a profession that enriches the future generations and builds a more sustainable future. The Global Teacher Prize contributes to this goal and that is why it is included in the educational projects promoted by the Foundation.
You can follow the progress of the project through Galp Foudation´s and the Global Teacher Prize´s Facebook
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Adaptive Track and Field Regional Training for
Ohio Middle/High School Coaches and Students (Grades 7-12)
- North: Sunday, February 23, 2020 (Deadline February 16) – Midwest Community Center, Dalton, Ohio
- Northeast: Sunday, March 1, 2020 (Deadline February 23)– Austintown Fitch High School, Ohio
- Brett Followay, USATF Level 1, CDSS
- Casey Followay, Wheelchair Racer, 7x State Champion, US Paralympics All-American
Download the flyer here.
1. Coaches/Athletic Directors
What: Get hands on training and learn about adaptive track and field from experienced coaches.
Time: 2:00-5:30 p.m.
Who: Middle and High School Coaches and Athletic Directors
- Chair parts/maintenance
- Chair propulsion/technique
- OHSAA wheelchair events guidelines/eligibility
- Coaching students with physical disabilities
Cost: $25.00
Register: RSVP required no later than Sunday prior. No onsite registrations accepted. Online here.
2. Students
What: Get fitted for a track chair, learn the basics and get ready to join your school's track team.
Time: 12:30-4:30 p.m.
Who: Male and female students grades 7-12 with a primary physical disability. Not restricted to wheelchair users; appropriate for individuals who are ambulatory and have orthopedic disabilities. Students must be able to independently propel the track chair.
- Equipment fittings
- Chair parts/maintenance
- Chair propulsion/technique
- OHSAA wheelchair events guidelines/eligibility
Cost: $100 for students borrowing equipment for the season, payable on the day of the training. ASPO multi-sport discount eligible.
Guidelines for student attendees:
- Equipment Loan:
- Attendance at one of the regional trainings is mandatory for equipment loan.
- Students who borrow a track chair from ASPO are considered an ASPO athlete and subject to ASPO policies and procedures.
- Students and parents/guardian must arrive promptly at 12:30 for equipment fittings.
- ASPO will do its best to make sure that a chair is available at the training that will work for students that register timely. However, track chairs are not one size fits all and additional modifications may be necessary. If ASPO is unable to fit the student with a track chair at the training, the student will be responsible for picking up a chair at our headquarters in Dalton, Ohio.
- A parent/guardian must be present to sign equipment release form.
- Student is responsible at their own expense for replacing track chair tires as necessary.
- Students should come with a vehicle appropriate for transporting a racing wheelchair as only the rear wheels can be removed. Chairs are approximately 6 feet long.
- Interested in purchasing your own track chair? ASPO is a dealer for Invacare Top End and we can assist you with your order.
- Non-Equipment Loan Attendees: Students who have their own equipment are still eligible to attend the training and should arrive at 2:00 instead of 12:30.
- Survey: Student attendees will be required to complete a survey at the conclusion of the track and field season.
- What to bring: Student attendees should bring a bicycle helmet and wear non-baggy athletic clothing for chair fittings..
Registration: Required no later than Sunday prior. No onsite registrations accepted. Before you register, have student's height, weight and hip width (not circumference, just hip to hip) handy. Complete online registration here.
Sponsors: Support for each regional training is provided by:


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"Doing No Harm" in an Age of Medical Repatriations: Challenges and Opportunities for Health Professionals – Juliana Morris
Juliana Morris
Harvard Medical School
How often do doctors cause harm to their patients when they discharge them from the hospital? For a sizeable group of immigrant patients who are "discharged" to their countries of origin each year, the answer may be: more often than not. The story of Quelino Ojeda Jimenez, an immigrant from Oaxaca, Mexico, who became quadriplegic after a fall at his roofing job in Chicago, Illinois, is case in point.
S-Comm Immigration Initiative Is Bad for Our Health – Karen Hacker, Jennifer Kasper, and Juliana Morris
Karen Hacker,1,2 Jennifer Kasper,1,3 and Juliana Morris1
1Harvard Medical School, 2Institute for Community Health, 3Massachusetts General Hospital
After much deliberation, Massachusetts Governor Deval Patrick decided this month not to sign the Secure Communities (S-Comm) program, following the lead of New York, Illinois, and other states. While this is a significant step in protecting immigrants' rights, health care providers need to be vigilant. It is important to understand the potential negative health implications of this program, which could affect all communities in Massachusetts and across the United States.


text: Taking the Lead: International Student Commencement Speaker Zinzile SibandaZinzile Sibanda grew up in Bulawayo, Zimbabwe. She chose to attend Hamline over other schools after a personable Skype interview with the admissions officer and because it offered a unique opportunity to experience the United States from the midwest. When she arrived in Saint Paul with her mother, they found a green campus and a welcoming community that provided Sibanda a home for four years. That community became critically important when, during her junior year, her mother passed away unexpectedly. She could have abandoned her studies; but, Hamline faculty, staff and students rallied around her, provided support, and ensured that she continued her education and graduated. In her last semester, after a professor encouraged her, Sibanda entered the competition to be the 2018 student commencement speaker. She was selected from among a pool of entrants by the committee. Her speech focussed on how special the Hamline community is, and also how important it is that graduates take what they've learned into the wider world. She challenged her classmates to build and strengthen the communities they find themselves in. "Go out into the world and take the lead...build and strengthen your communities so that they withstand the tests of time and the pressures of the world." An accomplished student while at Hamline, Sibanda received the Karen Vogel Prize in Political Science, a recognition for high achievement and student activity in international organizations, diplomacy and public policy. Sibanda's additional campus engagement included memberships in the Hamline African Students Association and the PRIDE Black Student Alliance. She also served as a global mentor and president to Hamline's Model United Nations. Since 2016, Sibanda has attended the annual National Model United Nations Conference in New York and served as chair to the International Telecommunications Union at the 2018 conference. Following graduation, Sibanda will pursue her graduate degree in political science with a focus on international relations and transnational government at Vrije Universiteit in Amsterdam, The Netherlands. Written by staff. 5/23/18