Whether environmental conditions –in particular energy and water availability– are sufficient to account for species richness gradients (e.g. Currie 1991), or the effects of other biotic and historical or regional factors need to be considered as well (e.g. Ricklefs 1987), was the subject of debate during the 1990s and 2000s (e.g. Francis & Currie 2003; Hawkins et al. 2003, 2006; Currie et al. 2004; Ricklefs 2004). The metabolic theory of ecology (Brown et al. 2004) provided a solid and well-rooted theoretical support for the preponderance of energy as the main driver for richness variations. As any good piece of theory, it provided testable predictions about the sign and shape (i.e. slope) of the relationship between temperature –a key aspect of ambient energy– and species richness. However, these predictions were not supported by empirical evaluations (e.g. Kreft & Jetz 2007; Algar et al. 2007; Hawkins et al. 2007a), as the effects of a myriad of other environmental gradients, regional factors and evolutionary processes result in a wide variety of richness–temperature responses across different groups and regions (Hawkins et al. 2007b; Hortal et al. 2008). So, in a textbook example of how good theoretical work helps advancing science even if proves to be (partially) wrong, the evaluation of this aspect of the metabolic theory of ecology led to current understanding that, while species richness does respond to current climatic conditions, many other ecological, evolutionary and historical factors do modify such response across scales (see, e.g., Ricklefs 2008; Hawkins 2008; D’Amen et al. 2017). And the kinetic model linking mean annual temperature and species richness (Allen et al. 2002; Brown et al. 2004) was put aside as being, perhaps, another piece of the puzzle of the origin of current diversity gradients.

Segovia (2021) puts together an elegant way of reinvigorating this part of the metabolic theory of ecology. He uses quantile regressions to model just the upper parts of the relationship between species richness and mean annual temperature, rather than modelling its central tendency through the classical linear regression family of methods –as was done in the past. This assumes that the baseline effect of ambient energy does produce the negative linear relationship between richness and temperature predicted by the kinetic model (Allen et al. 2002), but also that this effect only poses an upper limit for species richness, and the effects of other factors may result in lower levels of species co-occurrence, thus producing a triangular rather than linear relationship. The results of Segovia’s simple and elegant analytical design show unequivocally that the predictions of the kinetic model become progressively more explanatory towards the upper quartiles of the relationship between species richness and temperature along over 10,000 tree local inventories throughout the Americas, reaching over 70% of explanatory power for the upper 5% of the relationship (i.e. the 95% quantile). This confirms to a large extent his reformulation of the predictions of the kinetic model. 

Further, the neat study from Segovia (2021) also provides evidence confirming that the well-known spatial non-stationarity in the richness–temperature relationship (see Cassemiro et al. 2007) also applies to its upper-bound segment. Both the explanatory power and the slope of the relationship in the 95% upper quantile vary widely between biomes, reaching values similar to the predictions of the kinetic model only in cold temperate environments ­–precisely where temperature becomes more important than water availability as a constrain to plant life (O’Brien 1998; Hawkins et al. 2003). Part of these variations are indeed related with changes in water deficit and number of frost days along the XXth Century, as shown by the residuals of this paper (Segovia 2021) and a more detailed separate study (Segovia et al. 2020). This pinpoints the importance of the relative balance between water and energy as two of the main climatic factors constraining species diversity gradients, confirming the value of hypotheses that date back to Humboldt’s work (see Hawkins 2001, 2008). There is however a significant amount of unexplained variation in Segovia’s analyses, in particular in the progressive departure of the predictions of the kinetic model as we move towards the tropics, or downwards along the lower quantiles of the richness–temperature relationship. This calls for a deeper exploration of the factors that modify the baseline relationship between richness and energy, opening a new avenue for the macroecological investigation of how different forces and processes shape up geographical diversity gradients beyond the mere energetic constrains imposed by the basal limitations of multicellular life on Earth.

References

Algar, A.C., Kerr, J.T. and Currie, D.J. (2007) A test of Metabolic Theory as the mechanism underlying broad-scale species-richness gradients. Global Ecology and Biogeography, 16, 170-178. doi: https://doi.org/10.1111/j.1466-8238.2006.00275.x

Allen, A.P., Brown, J.H. and Gillooly, J.F. (2002) Global biodiversity, biochemical kinetics, and the energetic-equivalence rule. Science, 297, 1545-1548. doi: https://doi.org/10.1126/science.1072380

Brown, J.H., Gillooly, J.F., Allen, A.P., Savage, V.M. and West, G.B. (2004) Toward a metabolic theory of ecology. Ecology, 85, 1771-1789. doi: https://doi.org/10.1890/03-9000

Cassemiro, F.A.d.S., Barreto, B.d.S., Rangel, T.F.L.V.B. and Diniz-Filho, J.A.F. (2007) Non-stationarity, diversity gradients and the metabolic theory of ecology. Global Ecology and Biogeography, 16, 820-822. doi: https://doi.org/10.1111/j.1466-8238.2007.00332.x

Currie, D.J. (1991) Energy and large-scale patterns of animal- and plant-species richness. The American Naturalist, 137, 27-49. doi: https://doi.org/10.1086/285144

Currie, D.J., Mittelbach, G.G., Cornell, H.V., Field, R., Guegan, J.-F., Hawkins, B.A., Kaufman, D.M., Kerr, J.T., Oberdorff, T., O'Brien, E. and Turner, J.R.G. (2004) Predictions and tests of climate-based hypotheses of broad-scale variation in taxonomic richness. Ecology Letters, 7, 1121-1134. doi: https://doi.org/10.1111/j.1461-0248.2004.00671.x

D'Amen, M., Rahbek, C., Zimmermann, N.E. and Guisan, A. (2017) Spatial predictions at the community level: from current approaches to future frameworks. Biological Reviews, 92, 169-187. doi: https://doi.org/10.1111/brv.12222

Francis, A.P. and Currie, D.J. (2003) A globally consistent richness-climate relationship for Angiosperms. American Naturalist, 161, 523-536. doi: https://doi.org/10.1086/368223

Hawkins, B.A. (2001) Ecology's oldest pattern? Trends in Ecology & Evolution, 16, 470. doi: https://doi.org/10.1016/S0169-5347(01)02197-8 

Hawkins, B.A. (2008) Recent progress toward understanding the global diversity gradient. IBS Newsletter, 6.1, 5-8. https://escholarship.org/uc/item/8sr2k1dd

Hawkins, B.A., Field, R., Cornell, H.V., Currie, D.J., Guégan, J.-F., Kaufman, D.M., Kerr, J.T., Mittelbach, G.G., Oberdorff, T., O'Brien, E., Porter, E.E. and Turner, J.R.G. (2003) Energy, water, and broad-scale geographic patterns of species richness. Ecology, 84, 3105-3117. doi: https://doi.org/10.1890/03-8006

Hawkins, B.A., Diniz-Filho, J.A.F., Jaramillo, C.A. and Soeller, S.A. (2006) Post-Eocene climate change, niche conservatism, and the latitudinal diversity gradient of New World birds. Journal of Biogeography, 33, 770-780. doi: https://doi.org/10.1111/j.1365-2699.2006.01452.x

Hawkins, B.A., Albuquerque, F.S., Araújo, M.B., Beck, J., Bini, L.M., Cabrero-Sañudo, F.J., Castro Parga, I., Diniz-Filho, J.A.F., Ferrer-Castán, D., Field, R., Gómez, J.F., Hortal, J., Kerr, J.T., Kitching, I.J., León-Cortés, J.L., et al. (2007a) A global evaluation of metabolic theory as an explanation for terrestrial species richness gradients. Ecology, 88, 1877-1888. doi:10.1890/06-1444.1. doi: https://doi.org/10.1890/06-1444.1

Hawkins, B.A., Diniz-Filho, J.A.F., Bini, L.M., Araújo, M.B., Field, R., Hortal, J., Kerr, J.T., Rahbek, C., Rodríguez, M.Á. and Sanders, N.J. (2007b) Metabolic theory and diversity gradients: Where do we go from here? Ecology, 88, 1898–1902. doi: https://doi.org/10.1890/06-2141.1

Hortal, J., Rodríguez, J., Nieto-Díaz, M. and Lobo, J.M. (2008) Regional and environmental effects on the species richness of mammal assemblages. Journal of Biogeography, 35, 1202–1214. doi: https://doi.org/10.1111/j.1365-2699.2007.01850.x

Kreft, H. and Jetz, W. (2007) Global patterns and determinants of vascular plant diversity. Proceedings of the National Academy of Sciences USA, 104, 5925-5930. doi: https://doi.org/10.1073/pnas.0608361104

O'Brien, E. (1998) Water-energy dynamics, climate, and prediction of woody plant species richness: an interim general model. Journal of Biogeography, 25, 379-398. doi: https://doi.org/10.1046/j.1365-2699.1998.252166.x

Ricklefs, R.E. (1987) Community diversity: Relative roles of local and regional processes. Science, 235, 167-171. doi: https://doi.org/10.1126/science.235.4785.167

Ricklefs, R.E. (2004) A comprehensive framework for global patterns in biodiversity. Ecology Letters, 7, 1-15. doi: https://doi.org/10.1046/j.1461-0248.2003.00554.x

Ricklefs, R.E. (2008) Disintegration of the ecological community. American Naturalist, 172, 741-750. doi: https://doi.org/10.1086/593002

Segovia, R.A. (2021) Temperature predicts the maximum tree-species richness and water and frost shape the residual variation. bioRxiv, 836338, ver. 4 peer-reviewed and recommended by Peer community in Ecology. doi: https://doi.org/10.1101/836338

Segovia, R.A., Pennington, R.T., Baker, T.R., Coelho de Souza, F., Neves, D.M., Davis, C.C., Armesto, J.J., Olivera-Filho, A.T. and Dexter, K.G. (2020) Freezing and water availability structure the evolutionary diversity of trees across the Americas. Science Advances, 6, eaaz5373. doi: https://doi.org/10.1126/sciadv.aaz5373

Many bird species participate in mobbing in which individuals approach a predator while producing conspicuous vocalizations (Magrath et al. 2014). Mobbing is interesting to behavioral ecologists because of the complex array of costs of benefits. Costs range from the obvious risk of approaching a predator while drawing that predator’s attention to the more mundane opportunity costs of taking time away from other activities, such as foraging. Benefits may involve driving the predator to leave, teaching relatives to recognize predators, signaling quality to conspecifics, or others. An added layer of complexity in this system comes from the inter-specific interactions that often occur among different mobbing species (Magrath et al. 2014).

This study by Salis et al. (2023) explored the responses of a local bird community to mobbing calls produced by individuals of two common mobbing species in European forests, coal tits, and crested tits. Not only did they compare responses to these two different species, they assessed the impact of the number of mobbing individuals on the stimulus recordings, and they did so at two very different times of the year with different social contexts for the birds involved, winter (non-breeding) and spring (breeding). The experiment was well-designed and highly powered, and the authors tested and confirmed an important assumption of their design, and thus the results are convincing. It is clear that members of the local bird community responded differently to the two different species, and this result raises interesting questions about why these species differed in their tendency to attract additional mobbers. For instance, are species that recruit more co-mobbers more effective at recruiting because they are more reliable in their mobbing behavior (Magrath et al. 2014), more likely to reciprocate (Krams and Krama, 2002), or for some other reason? Hopefully this system, now of proven utility thanks to the current study, will be useful for following up on hypotheses such as these. Other convincing results, such as the higher rate of mobbing response in winter than in spring, also merit following up with further work.

Finally, their observation that playback of vocalizations of multiple individuals often elicited a more mobbing response that the playback of vocalizations of a single individual are interesting and consistent with other recent work indicating that groups of mobbers recruit more additional mobbers than do single mobbers (Dutour et al. 2021). However, as acknowledged in the manuscript, the design of the current study did not allow a distinction between the effect of multiple individuals signaling versus an effect of a stronger stimulus. Thus, this last result leaves the question of the effect of mobbing group size in these species open to further study.

REFERENCES

Dutour M, Kalb N, Salis A, Randler C (2021) Number of callers may affect the response to conspecific mobbing calls in great tits (Parus major). Behavioral Ecology and Sociobiology, 75, 29. https://doi.org/10.1007/s00265-021-02969-7

Krams I, Krama T (2002) Interspecific reciprocity explains mobbing behaviour of the breeding chaffinches, Fringilla coelebs. Proceedings of the Royal Society of London. Series B: Biological Sciences, 269, 2345–2350. https://doi.org/10.1098/rspb.2002.2155

Magrath RD, Haff TM, Fallow PM, Radford AN (2015) Eavesdropping on heterospecific alarm calls: from mechanisms to consequences. Biological Reviews, 90, 560–586. https://doi.org/10.1111/brv.12122

Salis A, Lena JP, Lengagne T (2023) Acoustic cues and season affect mobbing responses in a bird community. bioRxiv, 2022.05.05.490715, ver. 5 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2022.05.05.490715

Whether individuals are able to cope with new environmental conditions, and whether this ability can be improved, is certainly of great interest in our changing world. One way to cope with new conditions is through behavioral flexibility, which can be defined as “the ability to adapt behavior to new circumstances through packaging information and making it available to other cognitive processes” (Logan et al. 2023). Flexibility is predicted to be positively correlated with innovativeness, the ability to create a new behavior or use an existing behavior in a few situations (Griffin & Guez 2014). 
The post-study manuscript by Logan et al. (2023) proposes to test flexibility manipulability, and the relationship between flexibility and innovativeness. The authors did so with an experimental study on great-tailed grackles (Quiscalus mexicanus), an expanding species in the US, known to be flexible. 
The authors used serial reversal learning to investigate (1) whether behavioral flexibility, as measured by reversal learning using tubes of different shades, is manipulable; (2) whether manipulating (improving/training) behavioral flexibility improves flexibility and innovativeness in new contexts; (3) the type of learning strategy used by the individuals throughout the serial reversals.
The study described in this manuscript was pre-registered in Logan et al. (2019) and received in-principle recommendation on 26 Mar 2019 (Coulon 2019). One hypothesis from this original preregistration will be treated in a separate manuscript.
Among several interesting results, what I found most striking is that flexibility, in this species, seems to be a trait that is acquired by experience (vs. inherent to the individual). This opens exciting interrogations on the role of social learning, and on the impact of rapid environmental changes (which may force the individuals to experiment new ways to access to resources, for example), on individual flexibility and adaptability to new conditions. 
 
REFERENCES

Coulon A (2019) Can context changes improve behavioral flexibility? Towards a better understanding of species adaptability to environmental changes. Peer Community in Ecology, 100019. https://doi.org/10.24072/pci.ecology.100019

Griffin, A. S., & Guez, D. (2014). Innovation and problem solving: A review of common mechanisms. Behavioural Processes, 109, 121–134. https://doi.org/10.1016/j.beproc.2014.08.027

Logan C, Rowney C, Bergeron L, Seitz B, Blaisdell A, Johnson-Ulrich Z, McCune K (2019)
Is behavioral flexibility manipulatable and, if so, does it improve flexibility and problem solving in a new context? In Principle Recommendation 2019. PCI Ecology. http://corinalogan.com/Preregistrations/g_flexmanip.html

Logan CJ, Lukas D, Blaisdell AP, Johnson-Ulrich Z, MacPherson M, Seitz B, Sevchik A, McCune KB (2023) Behavioral flexibility is manipulable and it improves flexibility and innovativeness in a new context. EcoEcoRxiv, version 5 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.32942/osf.io/5z8xs

Continuous observation of animal behaviour could be quite a challenge in the field, and the situation becomes even more complicated with aquatic species mostly active at night. In such cases, biologging techniques are real game changers in ecology, behavioural ecology or eco-physiology. An accelerating number of methodological applications of these tools in natural condition are thus published each year [1]. Biologging is not limited to movement ecology. For instance, fine grain information about energy expenditure can be inferred from body acceleration [2], and accelerometers has already proven useful in monitoring reproductive costs in some fish species [3,4]. The first part of the study by Tentelier et al. [5] is in line with this growing literature. It describes measurements of energy expenditure during reproduction in a fish species, Allis shad (Alosa Alosa), based on tail beat frequency and occurrence of spawning acts. The study has been convincingly conducted, and the results are important for fish biologists. But this is not the whole story: the authors added to this otherwise classical study a very original and insightful analysis which deserves closer interest.
Tentelier et al. propose to use static accelerometer to monitor change in body roundness through the reproductive season. These semelparous fish first mature and built up reserves in the Atlantic Ocean and migrate into fresh water to reproduce. Contrary to iteroparous species, female shads do not have to strategically preserve energy for future reproduction. The females die few days after spawning having exhausted their energetic reserves: they typically lose almost half of their body mass during the spawning season. The beautiful idea in this study was to track down information about this dramatic slimming in the accelerometer data. Indeed, the accelerometer was attached on the side of the fish (close to the dorsal fin). A change in its angle with the vertical plane could be correlated with the change in roundness, the angle declining with the female thinning. Accelerometers have already been used to record body posture [6] but, in the present study, the novelty was to monitor the change in body shape.
Unfortunately, the data by Tentelier et al. are inconclusive so far. Broadly speaking, the accelerometer angle recorded declined through the spawning season, indicating an average slimming of the females, but there was no correlation between the change in angle and the mass loss at the individual level. This was partly due to the fact that the dorsal position of the accelerometer was not optimized to measures egg laying whose effects are mostly observable on ventral side.
Yet, this nice idea deserves more scrutiny. The method seems to be sensitive enough to detect inflation of swim bladder, the gas-filled organ helping the fish to control their position in the water column, as the accelerometer angle increased when the fish stayed close to the water surface. Additional works and proper calibration are certainly needed to validate the use of accelerometer angle as a proxy for body roundness. The actual data were not strong enough to justify a standalone publication on the subject, but it would have been shame to lose traces of such analysis and keep it in the file drawer. This is why I strongly support its report as a side question in a broader study. Science progresses not only with neat conclusive studies but also when unexpected (apparently anecdotal) observations stimulate new researches.

References

[1] Börger L, Bijleveld AI, Fayet AL, Machovsky‐Capuska GE, Patrick SC, Street GM and Vander Wal E. (2020) Biologging special feature. J. Anim. Ecol. 89, 6–15. 10.1111/1365-2656.13163
[2] Wilson RP et al. (2020) Estimates for energy expenditure in free‐living animals using acceleration proxies: A reappraisal. J. Anim. Ecol. 89, 161–172. 10.1111/1365-2656.13040
[3] Tsuda Y, Kawabe R, Tanaka H, Mitsunaga Y, Hiraishi T, Yamamoto K and Nashimoto K. (2006) Monitoring the spawning behaviour of chum salmon with an acceleration data logger. Ecol. Freshw. Fish 15, 264–274. 10.1111/j.1600-0633.2006.00147.x
[4] Sakaji H, Hamada K, Naito Y. 2018 Identifying spawning events of greater amberjack using accelerometers. Mar. Biol. Res. 14, 637–641. 10.1080/17451000.2018.1492140
[5] Tentelier C, Bouchard C, Bernardin A, Tauzin A, Aymes J-C, Lange F, Récapet C, Rives J (2020) The dynamics of spawning acts by a semelparous fish and its associated energetic costs. bioRxiv, 436295. doi: 10.1101/436295 ver. 7 peer-reviewed and recommended by PCI Ecology. 10.1101/436295
[6] Brown DD, Kays R, Wikelski M, Wilson R, Klimley AP. 2013 Observing the unwatchable through acceleration logging of animal behavior. Anim. Biotelemetry 1, 20. 10.1186/2050-3385-1-20

Even though all living organisms are, at the same time, involved in host-parasite interactions and embedded in complex food webs, the indirect effects of parasitism are only beginning to be unveiled.

Prosnier et al. investigated the direct and indirect effects of parasitism making use of a very interesting biological system comprising the freshwater zooplankton Daphnia magna and its highly specific parasite, the iridovirus DIV-1 (Daphnia-iridescent virus 1). Daphnia are typically semitransparent, but once infected develop a white phenotype with a characteristic iridescent shine due to the enlargement of white fat cells.

In a combination of infection trials and comparison of white and non-white phenotypes collected in natural ponds, the authors demonstrated increased mortality and reduced lifetime fitness in infected Daphnia. Furthermore, white phenotypes had lower mobility, increased reflectance, larger body sizes and higher protein content than non-white phenotypes. As a consequence, total energy content was effectively doubled in white Daphnia when compared to non-white broodless Daphnia. 

Next the authors conducted foraging trials with Daphnia predators Notonecta (the backswimmer) and Phoxinus (the European minnow). Focusing on Notonecta, unchanged search time and increased handling time were more than compensated by the increased energy content of white Daphnia. White Daphnia were 24% more profitable and consistently preferred by Notonecta, as the optimal foraging theory would predict. The authors argue that menu decisions of optimal foragers in the field might be different, however, as the prevalence – and therefore availability - of white phenotypes in natural populations is very low.

The study therefore contributes to our understanding of the trophic context of parasitism. One shortcoming of the study is that the authors rely exclusively on phenotypic signs for determining infection. On their side, DIV-1 is currently known to be highly specific to Daphnia, their study site is well within DIV-1 distributional range, and the symptoms of infection are very conspicuous. Furthermore, the infection trial – in which non-white Daphnia were exposed to white Daphnia homogenates - effectively caused several lethal and sublethal effects associated with DIV-1 infection, including iridescence. However, the infection trial also demonstrated that part of the exposed individuals developed intermediate traits while still keeping the non-white, non-iridescent phenotype. Thus, there may be more subtleties to the association of DIV-1 infection of Daphnia with ecological and evolutionary consequences, such as costs to resistance or covert infection, that the authors acknowledge, and that would be benefitted by coupling experimental and observational studies with the determination of actual infection and viral loads.​​​

References

Prosnier L., N. Loeuille, F.D. Hulot, D. Renault, C. Piscart, B. Bicocchi, M, Deparis, M. Lam, & V. Médoc. (2023). Parasites make hosts more profitable but less available to predators. BioRxiv, ver. 4 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2022.02.08.479552