Biodiversity at multiple trophic levels is needed for ecosystem multifunctionality.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Assessing the Viability of the Methylsulfinyl Radical-Ozone Reaction.

Although integral to remote marine atmospheric sulfur chemistry, the reaction between methylsulfinyl radical (CH3 SO) and ozone poses challenges to theoretical treatments. The lone theoretical study on this reaction reported an unphysically large barrier of 66 kcal mol-1 for abstraction of an oxygen atom from O3 by CH3 SO. Herein, we demonstrate that this result stems from improper use of MP2 with a single-reference, unrestricted Hartree-Fock (UHF) wavefunction. We characterized the potential energy surface using density functional theory (DFT), as well as multireference methodologies employing a complete active-space self-consistent field (CASSCF) reference. Our DFT PES shows, in contrast to previous work, that the reaction proceeds by forming an addition adduct [CH3 S(O3 )O] in a deep potential well of 37 kcal mol-1 . An O-O bond of this adduct dissociates via a flat, low barrier of 1 kcal mol-1 to give CH3 SO2 +O2 . The multireference computations show that the initial addition of CH3 SO+O3 is barrierless. These results provide a more physically intuitive and accurate picture of this reaction than the previous theoretical study. In addition, our results imply that the CH3 SO2 formed in this reaction can readily decompose to give SO2 as a major product, in alignment with the literature on CH3 SO reactions.

Decline in territory size and fecundity as a response to carrying capacity in an endangered songbird.

Density-dependent processes are fundamental mechanisms for the regulation of populations. Ecological theories differ in their predictions on whether increasing population density leads to individual adjustments of survival and reproductive output or to dominance and monopolization of resources. Here, we use a natural experiment to examine which factors limit population growth in the only remaining population of the endangered pale-headed brush finch (Atlapetes pallidiceps). For three distinct phases (a phase of population suppression, 2001-2002; expansion due to conservation management, 2003-2008; and equilibrium phase, 2009-2014), we estimated demographic parameters with an integrated population model using population size, the proportion of successfully breeding pairs and their productivity, territory size, and mark-recapture data of adult birds. A low proportion of successful breeders due to brood parasitism (0.42, 95% credible interval 0.26-0.59) limited population growth before 2003; subsequent culling of the brood parasite resulted in a two-fold increase of the proportion of successful breeders during the 'expansion phase'. When the population approached the carrying capacity of its habitat, territory size declined by more than 50% and fecundity declined from 1.9 (1.54-2.27) to 1.3 (1.12-1.53) chicks per breeding pair, but the proportion of successful breeders remained constant (expansion phase: 0.85; 0.76-0.93; equilibrium phase: 0.86; 0.79-0.92). This study demonstrates that limiting resources can lead to individual adjustments instead of despotic behavior, and the individual reduction of reproductive output at high population densities is consistent with the slow life-history of many tropical species.

Morphology predicts species' functional roles and their degree of specialization in plant-frugivore interactions.

Species' functional roles in key ecosystem processes such as predation, pollination or seed dispersal are determined by the resource use of consumer species. An interaction between resource and consumer species usually requires trait matching (e.g. a congruence in the morphologies of interaction partners). Species' morphology should therefore determine species' functional roles in ecological processes mediated by mutualistic or antagonistic interactions. We tested this assumption for Neotropical plant-bird mutualisms. We used a new analytical framework that assesses a species's functional role based on the analysis of the traits of its interaction partners in a multidimensional trait space. We employed this framework to test (i) whether there is correspondence between the morphology of bird species and their functional roles and (ii) whether morphologically specialized birds fulfil specialized functional roles. We found that morphological differences between bird species reflected their functional differences: (i) bird species with different morphologies foraged on distinct sets of plant species and (ii) morphologically distinct bird species fulfilled specialized functional roles. These findings encourage further assessments of species' functional roles through the analysis of their interaction partners, and the proposed analytical framework facilitates a wide range of novel analyses for network and community ecology.

Investment in attending to cues and the evolution of amplifiers.

Signals and cues are extensively used in social interactions across diverse communication systems. Here, we extend an existing theoretical framework to explore investment by emitters and perceivers in the fidelity with which cues and signals associated with the former are detected by the latter. Traits of the emitter that improve cue or signal fidelity without adding information are termed 'amplifiers'. We assume that each party can invest in improving fidelity but that it is increasingly costly the more fidelity is improved. Our model predicts that evolution of amplifier traits of a pre-existing cue occurs over a broader range of circumstances than evolution of signalling in situations where the emitter offered no pre-existing cue to the perceiver. It further predicts that the greater the intrinsic informational value of a cue, the more likely it is that the perceiver (and not the emitter) will invest in the fidelity of detecting that cue. A consequence of this predicted asymmetry is that true communication with reciprocal adaptations in emitters and perceivers to improve signal fidelity is likely to occur predominantly for traits of intermediate reliability. The corollary is that uncertainty of the perceiver will then be a key feature of communication. Uncertainty can arise because perceivers misinterpret signals or do not perceive them correctly, but here we argue that uncertainty is more fundamentally at the root of communication because traits that are intrinsically highly informative will induce only the perceiver and not the emitter to invest in improved fidelity of perception of that trait.

Energetics and transition-state dynamics of the F + HOCH3 → HF + OCH3 reaction.

The F + HOCH3 → HF + OCH3 reaction is a system with 15 internal degrees of freedom that can provide a benchmark for the development of theory for increasingly complex chemical reactions. The dynamics of this reaction were studied by photoelectron-photofragment coincidence (PPC) spectroscopy carried out on the F-(HOCH3) anion, aided by a computational study of both the anion and neutral potential energy surfaces, with energies extrapolated to the CCSDT(Q)/CBS level of theory. Photodetachment at 4.80 eV accesses both the reactant and product channels for this reaction. In the product channel (HF + OCH3 + e-) of the neutral potential energy surface, vibrationally excited HF products in addition to the stable product-channel hydrogen-bonded complex (FH-OCH3) are observed in the PPC and photoelectron spectra. In addition, experimental evidence is observed for the reactant-channel van der Waals complex (F-HOCH3), in good agreement with the theoretical predictions. The relative stability of these long-lived complexes was probed by reducing the ion beam energy, increasing the product time-of-flight, indicating lifetimes on the microsecond timescale for the reactant- and product-channel complexes as well as providing evidence for long-lived vibrational Feshbach resonances associated with the HF(v > 0) + OCH3 product states. This system will provide a model for extending full-dimensionality quantum dynamics to larger numbers of degrees of freedom.

Photodissociation of Cerium Oxide Nanocluster Cations.

Cerium oxide cluster cations, CexOy(+), are produced via laser vaporization in a pulsed nozzle source and detected with time-of-flight mass spectrometry. The mass spectrum displays a strongly preferred oxide stoichiometry for each cluster with a specific number of metal atoms x, with x ≤ y. Specifically, the most prominent clusters correspond to the formula CeO(CeO2)n(+). The cluster cations are mass selected and photodissociated with a Nd:YAG laser at either 532 or 355 nm. The prominent clusters dissociate to produce smaller species also having a similar CeO(CeO2)n(+) formula, always with apparent leaving groups of (CeO2). The production of CeO(CeO2)n(+) from the dissociation of many cluster sizes establishes the relative stability of these clusters. Furthermore, the consistent loss of neutral CeO2 shows that the smallest neutral clusters adopt the same oxidation state (IV) as the most common form of bulk cerium oxide. Clusters with higher oxygen content than the CeO(CeO2)n(+) masses are present with much lower abundance. These species dissociate by the loss of O2, leaving surviving clusters with the CeO(CeO2)n(+) formula. Density functional theory calculations on these clusters suggest structures composed of stable CeO(CeO2)n(+) cores with excess oxygen bound to the surface as a superoxide unit (O2(-)).

Evaluating the Impact of Performance Improvement Initiatives on Transplant Center Reporting Compliance and Patient Follow-Up After Living Kidney Donation.

With the changing demographics of the living donor population and increased regulatory oversight, it is important that transplant centers report outcomes accurately. The aim of our retrospective cohort study of 312 living donors who underwent nephrectomy between 2008 and 2013 was to evaluate the impact of living donor program performance improvement initiatives on: (i) transplant center program reporting compliance; (ii) patient compliance with postdonation follow-up and its associated factors; and (iii) overall financial costs to the transplant center. The effect of the initiatives (donation eras 2008-2010 and 2011-2013) on compliance at key reporting points (6 months, 1 year, 2 years) was analyzed using correlation coefficients, χ(2) and Fisher's exact tests. Multivariable logistic regression models tested the initiatives' effect on the likelihood of patient follow-up. The initiatives were associated with significant improvement in form reporting compliance (r ≥ 0.862, p ≤ 0.027; 1 and 2 year Fisher's Exact p ≤ 0.002) and patient follow-up (χ(2) p ≤ 0.009) with acceptable transplant center costs. Multivariable analyses demonstrated that donation era was consistently and significantly (p < 0.001) associated with increased likelihood of postdonation patient follow-up. Institution of performance improvement initiatives with dedicated program resources is financially feasible and leads to more accurate and complete form reporting and improved patient follow-up after nephrectomy.

Collective Modes and Structural Modulation in Ni-Mn-Ga(Co) Martensite Thin Films Probed by Femtosecond Spectroscopy and Scanning Tunneling Microscopy.

The origin of the martensitic transition in the magnetic shape memory alloy Ni-Mn-Ga has been widely discussed. While several studies suggest it is electronically driven, the adaptive martensite model reproduced the peculiar nonharmonic lattice modulation. We used femtosecond spectroscopy to probe the temperature and doping dependence of collective modes, and scanning tunneling microscopy revealed the corresponding static modulations. We show that the martensitic phase can be described by a complex charge-density wave tuned by magnetic ordering and strong electron-lattice coupling.

Bicolored display of Miconia albicans fruits: Evaluating visual and physiological functions of fruit colors.

PREMISE OF THE STUDY: Most bird-dispersed fruits are green when unripe and become colored and conspicuous when ripe, signaling that fruits are ready to be consumed and dispersed. The color pattern for fruits of Miconia albicans (Melastomataceae), however, is the opposite, with reddish unripe and green ripe fruits. We (1) verified the maintenance over time of its bicolored display, (2) tested the communicative function of unripe fruits, (3) tested the photoprotective role of anthocyanins in unripe fruits, and (4) verified whether green ripe fruits can assimilate carbon. METHODS: Using a paired experiment, we tested whether detection of ripe fruits was higher on infructescences with unripe and ripe fruits compared with infructescences with only ripe fruits. We also measured and compared gas exchange, chlorophyll a fluorescence, and heat dissipation of covered (to prevent anthocyanin synthesis) and uncovered ripe and unripe fruits. KEY RESULTS: Although the bicolored display was maintained over time, unripe fruits had no influence on bird detection and removal of ripe fruits. Ripe and unripe fruits did not assimilate CO2, but they respired instead. CONCLUSIONS: Since the communicative function of unripe fruits was not confirmed, seed dispersers are unlikely to select the display with bicolored fruits. Because of the absence of photosynthetic activity in ripe and unripe fruits and enhanced photoprotective mechanisms in ripe fruits rather than in unripe fruits, we could not confirm the photoprotective role of anthocyanins in unripe fruits. As an alternative hypothesis, we suggest that the bicolored fruit display could be an adaptation to diversify seed dispersal vectors instead of restricting dispersal to birds and that anthocyanins in unripe fruits may have a defense role against pathogens.

[Can Postoperative Accelerated Lens Opacification be Limited by Lying in "Face-Down Position" after Vitrectomy with Gas as Tamponade?]. / Kann eine postoperativ beschleunigte Linsentrübung nach einer Pars-plana-Vitrektomie mit Gas als Tamponade durch eine konsequente "Gesicht-nach-unten-Lagerung" vermieden werden?

BACKGROUND: Developing a post-operative cataract after pars-plana vitrectomy (ppV) or core pars-plana vitrectomy (cppV) with gas as tamponade within 6 to 12 months is a common complication and is mostly accepted as unavoidable. Often a combined cataract-ppV surgery in the first place is recommended. The main goal of this study was to analyse the effects of a "face-down positioning" ("fdp") on the lens transparency and the phaco rate. By using the positive experience of an ergonomic body positioning it should be possible to improve the compliance during the period of "fdp" after surgery. METHODS: During the study period of up to 24 months, we observed in a prospective, controlled, clinical and randomised pilot study 30 patients who all had undergone a ppV/cppV with sulfur hexafluoride gas tamponade (SF6 25 %). All patients of the supported group (SG) and the control group (CG) were requested to keep their face consequently downwards until the gas bubble was resorbed completely in order to avoid a direct contact with the lens. The real time in "fdp" in hours per day (24 hours) and the grade of lens opacification was documented pre- and postoperatively using a Pentacam HR (Oculus, Wetzlar). RESULTS: The compliance of patients tested measuring the real time of "fdp" in hours per day varied largely (SG: 19.5/24; CG: 5.5/24; p < 0,0001). Cataract development was found to be directly related to the real time of "fdp". A cataract surgery was necessary mainly in the non-supported group (SG: 4/15; CG:12/15; p = 0.012). There was no lens opacification in the supported group after performing the "fdp" for at least 20 hours daily until the resorption of the gas bubble was completed. This result did not correlate with the age of the patient. CONCLUSION: With the help of sufficient support at the disposal of the patients, it is possible to significantly improve the compliance during the period of "fdp" after surgery. In order to get a safe and painless "fdp" in bed, an ergonomic body positioning is necessary (Schaefer 2012). By practicing the "fdp" until the gas bubble is completely resorbed, the risk of developing a postoperative cataract can be reduced significantly. Provided there is a good compliance to "fdp", the gas bubble can cause the desired tamponade effect even when using shorter acting gases. By performing a consistent "fdp" it is possible to accelerate the healing process and avoid reoperations. Hereby it should even be possible to use an SF6-air mix or optionally simply air as tamponade.

A randomized study evaluating cinacalcet to treat hypercalcemia in renal transplant recipients with persistent hyperparathyroidism.

Persistent hyperparathyroidism (HPT) after kidney transplantation (KTx) is associated with hypercalcemia, hypophosphatemia and abnormally high levels of parathyroid hormone (PTH). In this randomized trial, cinacalcet was compared to placebo for the treatment of hypercalcemia in adult patients with persistent HPT after KTx. Subjects were randomized 1:1 to cinacalcet or placebo with randomization stratified by baseline corrected total serum calcium levels (≤11.2 mg/dL [2.80 mmol/L] or >11.2 mg/dL [2.80 mmol/L]). The primary end point was achievement of a mean corrected total serum calcium value<10.2 mg/dL (2.55 mmol/L) during the efficacy period. The two key secondary end points were percent change in bone mineral density (BMD) at the femoral neck and absolute change in phosphorus; 78.9% cinacalcet- versus 3.5% placebo-treated subjects achieved the primary end point with a difference of 75.4% (95% confidence interval [CI]: 63.8, 87.1), p<0.001. There was no statistical difference in the percent change in BMD at the femoral neck between cinacalcet and placebo groups, p=0.266. The difference in the change in phosphorus between the two arms was 0.45 mg/dL (95% CI: 0.26, 0.64), p<0.001 (nominal). No new safety signals were detected. In conclusion, hypercalcemia and hypophosphatemia were effectively corrected after treatment with cinacalcet in patients with persistent HPT after KTx.

Birds see the true colours of fruits to live off the fat of the land.

Communication is a characteristic of life, but its reliability and basic definition are hotly debated. Theory predicts that trade among mutualists requires high reliability. Here, we show that moderate reliability already allows mutualists to optimize their rewards. The colours of Mediterranean fleshy-fruits indicate lipid rewards (but not other nutrients) to avian seed dispersers on regional and local scales. On the regional scale, fruits with high lipid content were significantly darker and less chromatic than congeners with lower lipid content. On the local scale, two warbler species (Sylvia atricapilla and Sylvia borin) selected fruit colours that were less chromatic, and thereby maximized their intake of lipids-a critical resource during migration and wintering. Crucially, birds were able to maximize lipid rewards with moderate reliability from visual fruit colours (r(2) = 0.44-0.60). We suggest that mutualisms require only that any association between the quality and sensory aspects of signallers is learned through multiple, repeated interactions. Because these conditions are often fulfilled, also in social communication systems, we contend that selection on reliability is less intense than hitherto assumed. This may contribute to explaining the extraordinary diversity of signals, including that of plant reproductive displays.

Why fruit rots: theoretical support for Janzen's theory of microbe-macrobe competition.

We present a formal model of Janzen's influential theory that competition for resources between microbes and vertebrates causes microbes to be selected to make these resources unpalatable to vertebrates. That is, fruit rots, seeds mould and meat spoils, in part, because microbes gain a selective advantage if they can alter the properties of these resources to avoid losing the resources to vertebrate consumers. A previous model had failed to find circumstances in which such a costly spoilage trait could flourish; here, we present a simple analytic model of a general situation where costly microbial spoilage is selected and persists. We argue that the key difference between the two models lies in their treatments of microbial dispersal. If microbial dispersal is sufficiently spatially constrained that different resource items can have differing microbial communities, then spoilage will be selected; however, if microbial dispersal has a strong homogenizing effect on the microbial community then spoilage will not be selected. We suspect that both regimes will exist in the natural world, and suggest how future empirical studies could explore the influence of microbial dispersal on spoilage.

Adaptation of flower and fruit colours to multiple, distinct mutualists.

Communication in plant-animal mutualisms frequently involves multiple perceivers. A fundamental uncertainty is whether and how species adapt to communicate with groups of mutualists having distinct sensory abilities. We quantified the colour conspicuousness of flowers and fruits originating from one European and two South American plant communities, using visual models of pollinators (bee and fly) and seed dispersers (bird, primate and marten). We show that flowers are more conspicuous than fruits to pollinators, and the reverse to seed dispersers. In addition, flowers are more conspicuous to pollinators than to seed dispersers and the reverse for fruits. Thus, despite marked differences in the visual systems of mutualists, flower and fruit colours have evolved to attract multiple, distinct mutualists but not unintended perceivers. We show that this adaptation is facilitated by a limited correlation between flower and fruit colours, and by the fact that colour signals as coded at the photoreceptor level are more similar within than between functional groups (pollinators and seed dispersers). Overall, these results provide the first quantitative demonstration that flower and fruit colours are adaptations allowing plants to communicate simultaneously with distinct groups of mutualists.

Genetic depletion at adaptive but not neutral loci in an endangered bird species.

Many endangered species suffer from the loss of genetic diversity, but some populations may be able to thrive even if genetically depleted. To investigate the underlying genetic processes of population bottlenecks, we apply an innovative approach for assessing genetic diversity in the last known population of the endangered Pale-headed Brushfinch (Atlapetes pallidiceps) in Ecuador. First, we measure genetic diversity at eleven neutral microsatellite loci and adaptive SNP variation in five Toll-like receptor (TLR) immune system genes. Bottleneck tests confirm genetic drift as the main force shaping genetic diversity in this species and indicate a 99 % reduction in population size dating back several hundred years. Second, we compare contemporary microsatellite diversity with historic museum samples of A. pallidiceps, finding no change in genetic diversity. Third, we compare genetic diversity in the Pale-headed Brushfinch with two co-occurring-related brushfinch species (Atlapetes latinuchus, Buarremon torquatus), finding a reduction of up to 91% diversity in the immune system genes but not in microsatellites. High TLR diversity is linked to decreased survival probabilities in A. pallidiceps. Low TLR diversity is thus probably an adaptation to the specific selection regime within its currently very restricted distribution (approximately 200 ha), but could severely restrict the adaptive potential of the species in the long run. Our study illustrates the importance of investigating both neutral and adaptive markers to assess the effect of population bottlenecks and for recommending specific management plans in endangered species.

Linking signal fidelity and the efficiency costs of communication.

The handicap principle has been the overarching framework to explain the evolution and maintenance of communication. Yet, it is becoming apparent that strategic costs of signalling are not the only mechanism maintaining signal honesty. Rather, the fidelity of detecting signals can itself be strongly selected. Specifically, we argue that the fidelity of many signals will be constrained by the investment in signal generation and reception by the signaller and perceiver, respectively. Here, we model how investments in signal fidelity influence the emergence and stability of communication using a simple theoretical framework. The predictions of the model indicate that high-cost communication can be stable whereas low-cost intermediates are generally selected against. This dichotomy suggests that the most parsimonious route to the evolution of communication is for initial investment in communicative traits to be driven by noncommunicative functions. Such cues can appeal to pre-existing perceptual biases and thereby stimulate signal evolution. We predict that signal evolution will vary between systems in ways that can be linked to the economics of communication to the two parties involved.

Self-supplementation and effects of dietary antioxidants during acute thermal stress.

Thermal stress leads to increased production of reactive oxygen species. If an organism is not able to simultaneously mount an efficient antioxidant defense system, this may lead to increased oxidative damage, which is potentially deleterious in terms of health and fitness. Exposure to cold or heat is therefore expected to be associated with a high demand for antioxidants. In agreement, several studies have shown that supplementing the diet of thermally stressed organisms with antioxidants leads to a reduction of oxidative damage. However, whether organisms can actively supplement their diet with antioxidants to alleviate temperature-induced oxidative damage is unknown. Here, we show that captive Gouldian finches (Erythrura gouldiae) supplement their diet more with seeds rich in antioxidants below than within their thermoneutral zone. Moreover, having access to seeds rich in antioxidants at temperatures below thermoneutrality decreases their oxidative damage. These results indicate that, when facing a thermal challenge, animals are able to take advantage of the antioxidant properties of their food to improve their oxidative balance. Having access to food resources rich in antioxidants may therefore be of primary importance for organisms in their natural habitat, as it may help them to cope with oxidative constraints due to challenging temperature regimes.

Fenestration: a window of opportunity for carnivorous plants.

A long-standing but controversial hypothesis assumes that carnivorous plants employ aggressive mimicry to increase their prey capture success. A possible mechanism is that pitcher plants use aggressive mimicry to deceive prey about the location of the pitcher's exit. Specifically, species from unrelated families sport fenestration, i.e. transparent windows on the upper surfaces of pitchers which might function to mimic the exit of the pitcher. This hypothesis has not been evaluated against alternative hypotheses predicting that fenestration functions to attract insects from afar. By manipulating fenestration, we show that it does not increase the number of Drosophila flies or of two ant species entering pitchers in Sarracenia minor nor their retention time or a pitcher's capture success. However, fenestration increased the number of Drosophila flies alighting on the pitcher compared with pitchers of the same plant without fenestration. We thus suggest that fenestration in S. minor is not an example of aggressive mimicry but rather functions in long-range attraction of prey. We highlight the need to evaluate aggressive mimicry relative to alternative concepts of plant-animal communication.

Limits of selection against cheaters: birds prioritise visual fruit advertisement over taste.

The concept of biological markets aims to explain how organisms interact with each other. Market theory predicts that organisms choose the most rewarding partner in mutualisms. However, partner choice may also be influenced by advertisement which may not be reliable. In seed dispersal mutualism, we analysed whether seed dispersers prioritise taste cues over visual advertisement to select the most rewarding fruits and whether they select against partners with unreliable advertisement. We conducted experiments on black elder (Sambucus nigra), a species of which the colours of the peduncles match the sugar content of their fruits. We created infructescences the colours of which matched or mismatched the sugar content of their fruits. There was no selection against cheaters in the field or by captive blackcaps (Sylvia atricapilla) as seed dispersers. Blackcaps were constrained to select against unreliable advertisement because they swallowed fruits entirely and thus did not obtain an immediate feedback by taste. Instead, blackcaps selected fruits according to the colour variation of red peduncles. Overall, we suggest that the concept of constraints should be incorporated into biological markets. We further contend that biological markets can be more complex than currently acknowledged because a moderate degree of reliability occurred in black elder even in the absence of selection against cheaters.