It is my pleasure to recommend the paper by Raynal et al. [1] about using random forest for parameter inference. There are two reviews about the paper, one review written by Dennis Prangle and another review written by myself. Both reviews were positive and included comments that have been addressed in the current version of the preprint.

The paper nicely shows that modern machine learning approaches are useful for Approximate Bayesian Computation (ABC) and more generally for simulation-driven parameter inference in ecology and evolution.

The authors propose to consider the random forest approach, proposed by Meinshausen [2] to perform quantile regression. The numerical implementation of ABC with random forest, available in the abcrf package, is based on the RANGER R package that provides a fast implementation of random forest for high-dimensional data.

According to my reading of the manuscript, there are 3 main advantages when using random forest (RF) for parameter inference with ABC. The first advantage is that RF can handle many summary statistics and that dimension reduction is not needed when using RF.

The second advantage is very nicely displayed in Figure 5, which shows the main result of the paper. If correct, 95% posterior credibility intervals (C.I.) should contain 95% of the parameter values used in simulations. Figure 5 shows that posterior C.I. obtained with rejection are too large compared to other methods. By contrast, C.I. obtained with regression methods have been shrunken. However, the shrinkage can be excessive for the smallest tolerance rates, with coverage values that can be equal to 85% instead of the expected 95% value. The attractive property of RF is that C.I. have been shrunken but the coverage is of 100% resulting in a conservative decision about parameter values.

The last advantage is that no hyperparameter should be chosen. It is a parameter free approach, which is desirable because of the potential difficulty of choosing an appropriate acceptance rate.

The main drawback of the proposed approach concerns joint parameter inference. There are many settings where the joint parameter distribution is of interest and the proposed RF approach cannot handle that. In population genetics for example, estimation of the severity and of the duration of the bottleneck should be estimated jointly because of identifiability issues. The challenge of performing joint parameter inference with RF might constitute a useful research perspective.
 

References
 

[1] Raynal L, Marin J-M, Pudlo P, Ribatet M, Robert CP, Estoup A. 2017. ABC random forests for Bayesian parameter inference. arXiv 1605.05537v4, https://arxiv.org/pdf/1605.05537
[2] Meinshausen N. 2006. Quantile regression forests. Journal of Machine Learning Research 7: 983-999. http://www.jmlr.org/papers/v7/meinshausen06a.html

Recent advances in next-generation sequencing are allowing us to uncover the evolution of sex chromosomes in non-model organisms. This study [1] represents an example of this application to birds of two Sylvioidea species from the genus Zosterops (commonly known as white-eyes). The study is exemplary in the amount and types of data generated and in the thoroughness of the analysis applied. Both male and female genomes were sequenced to allow the authors to identify sex-chromosome specific scaffolds. These data were augmented by generating the transcriptome (RNA-seq) data set. The findings after the analysis of these extensive data are intriguing: neoZ and neoW chromosome scaffolds and their breakpoints were identified. Novel sex chromosome formation appears to be accompanied by translocation events. The timing of formation of novel sex chromosomes was identified using molecular dating and appears to be relatively recent. Yet first signatures of distinct evolutionary patterns of sex chromosomes vs. autosomes could be already identified. These include the accumulation of transposable elements and changes in GC content. The changes in GC content could be explained by biased gene conversion and altered recombination landscape of the neo sex chromosomes. The authors also study divergence and diversity of genes located on the neo sex chromosomes. Here their findings appear to be surprising and need further exploration. The neoW chromosome already shows unique patterns of divergence and diversity at protein-coding genes as compared with genes on either neoZ or autosomes. In contrast, the genes on the neoZ chromosome do not display divergence or diversity patterns different from those for autosomes. This last observation is puzzling and I believe should be explored in further studies. Overall, this study significantly advances our knowledge of the early stages of sex chromosome evolution in vertebrates, provides an example of how such a study could be conducted in other non-model organisms, and provides several avenues for future work.

References

[1] Leroy T., Anselmetti A., Tilak M.K., Bérard S., Csukonyi L., Gabrielli M., Scornavacca C., Milá B., Thébaud C. and Nabholz B. (2019). A bird’s white-eye view on neo-sex chromosome evolution. bioRxiv, 505610, ver. 4 peer-reviewed and recommended by PCI Evolutionary Biology. doi: 10.1101/505610

150 years after Darwin published ‘Descent of man and selection in relation to sex’ (Darwin, 1871), the evolutionary mechanism that he laid out in his treatise continues to fascinate us. Sexual selection is responsible for some of the most spectacular traits among animals, and plants, and it appeals to our interest in all things reproductive and sexual (Bell, 1982). In addition, sexual selection poses some of the more intractable problems in evolutionary biology: Its realm encompasses traits that are subject to markedly different selection pressures, particularly when distinct, yet associated, traits tend to be associated with males, e.g. courtship signals, and with females, e.g. preferences (cf. Ah-King & Ahnesjo, 2013). While separate, such traits cannot evolve independently of each other (Arnqvist & Rowe, 2005), and complex feedback loops and correlations between them are predicted (Greenfield et al., 2014). Traditionally, sexual selection has been modelled under simplifying assumptions, and quantitative genetic approaches that avoided evolutionary dynamics have prevailed. New computing methods may be able to free the field from these constraints, and a trio of theoreticians (Chevalier, De Coligny & Labonne 2020) describe here a novel application of a ‘demo-genetic agent (or individual) based model’, a mouthful hereafter termed DG-ABM, for arriving at a holistic picture of the sexual selection trajectory. The application is built on the premise that traits, e.g. courtship, preference, gamete investment, competitiveness for mates, can influence the genetic architecture, e.g. correlations, of those traits. In turn, the genetic architecture can influence the expression and evolvability of the traits. Much of this influence occurs via demographic features, i.e. social environment, generated by behavioral interactions during sexual advertisement, courtship, mate guarding, parental care, post-mating dispersal, etc.
The authors provide a lengthy verbal description of their model, specifying the genomic and behavioral parameters that can be set and how a ‘run’ may be initialized. There is a link to an internet site where users can then enter their own parameter values and begin exploring hypotheses. Back in the article several simulations illustrate simple tests; e.g. how gamete investment and preference jointly evolve given certain survival costs. One obvious test would have been the preference – courtship genetic correlation that represents the core of Fisherian runaway selection, and it is regrettable that it was not examined under a range of demographic parameters. As presented the author’s DG-ABM appears particularly geared toward mating systems in ‘higher’ vertebrates, where couples form during a discrete mating season and are responsible for most reproduction. It is not clear how applicable the model could be to a full range of mating systems and nuances, including those in arthropods and other invertebrates as well as plants.
What is the likely value of the DG-ABM for sexual selection researchers? We will not be able to evaluate its potential impact until readers with specialized understanding of a question and taxon begin exploring and comparing their results with prior expectations. Of course, lack of congruence with earlier predictions would not invalidate the model. Hopefully, some of these specialists will have opportunities for comparing results with pertinent empirical data.

References

Ah-King, M. and Ahnesjo, I. 2013. The ‘sex role’ concept: An overview and evaluation Evolutionary Biology, 40, 461-470. doi: https://doi.org/10.1007/s11692-013-9226-7
Arnqvist, G. and Rowe, L. 2005. Sexual Conflict. Princeton University Press, Princeton. doi: https://doi.org/10.1515/9781400850600
Bell, G. 1982. The Masterpiece of Nature: The Evolution and Genetics of Sexuality. University of California Press, Berkeley.
Chevalier, L., De Coligny, F. and Labonne, J. (2020) A demogenetic individual based model for the evolution of traits and genome architecture under sexual selection. bioRxiv, 2020.04.01.014514, ver. 4 peer-reviewed and recommended by PCI Evol Biol. doi: https://doi.org/10.1101/2020.04.01.014514
Darwin, C. 1871. The Descent of Man and Selection in Relation to Sex. J. Murray, London.
Greenfield, M.D., Alem, S., Limousin, D. and Bailey, N.W. 2014. The dilemma of Fisherian sexual selection: Mate choice for indirect benefits despite rarity and overall weakness of trait-preference genetic correlation. Evolution, 68, 3524-3536. doi: https://doi.org/10.1111/evo.12542

Historically neglected or considered a rare aberration in animals under the biological species concept, interspecific hybridisation has by now been recognised to be taxonomically widespread, particularly in rapidly diversifying groups (Dagilis et al. 2021; Edelman & Mallet 2021; Mallet et al. 2016; Seehausen 2004). Yet the prevalence of introgressive hybridizations, its evolutionary significance, and its impact on species diversification remain a hot topic of research in evolutionary biology. The rapid increase in genomic resources now available for non-model species has significantly contributed to the detection of introgressive hybridization across taxa showing that reticulated evolution is far more common in the animal kingdom than historically considered. Yet, detecting it, quantifying its magnitude, and assessing its evolutionary significance remains a challenging endeavour with constantly evolving methodologies to better explore and exploit genomic data (Blair & Ané 2020; Degnan & Rosenberg 2009; Edelman & Mallet 2021; Hibbins & Hahn 2022).

In the marine realm, the dearth of geographic barriers and the large dispersal abilities of pelagic species like cetaceans have raised the questions of how populations and species can diverge and adapt to distinct ecological conditions in face of potentially large gene-flow, the so-called marine speciation paradox (Bierne et al. 2003). Contemporaneous hybridization among cetacean species has been widely documented in nature despite large phenotypic differences (Crossman et al. 2016). The historical prevalence of reticulated evolution, its evolutionary significance, and how it might have impacted the evolutionary history and diversification of the cetaceans have however remained elusive so far. Recent phylogenomic studies suggested that introgression has been prevalent in cetacean evolutionary history with instances reported among baleen whales (mysticetes) (Árnason et al. 2018) and among toothed whales (odontocetes), especially in the rapidly diversifying dolphins family of the Delphininae (Guo et al. 2021; Moura et al. 2020).

Analysing publicly available whole-genome data from nine cetacean species across three families of Delphinoidae – dolphins, porpoises, and monondontidae – using phylogenomics and demo-genetics approaches, Westbury and colleagues (2022) take a step further in documenting that evolution among these species has been far from a simple bifurcating tree. Instead, their study describes widespread occurrences of introgression among Delphinoidae, drawing a complex picture of reticulated evolutionary history. After describing major topology discordance in phylogenetic gene trees along the genome, the authors use a panel of approaches to disentangle introgression from incomplete lineage sorting (ILS), the two most common causes of tree topology discordances (Hibbins & Hahn 2022). Applying popular tests that separate introgression from ILS, such as the Patterson’s D (a.k.a. ABBA-BABA) test (Durand et al. 2011; Green et al. 2010), QuIBL (Edelman et al. 2019), and D-FOIL (Pease & Hahn 2015), the authors report that signals of introgression are present in the genomes of most (if not all) the cetacean species included in their study. However, this picture needs to be nuanced. Most introgression signals seem to echo old introgression events that occurred primarily among ancestors. This is suggested by the differential signals of topology discordance along the genome when considering sliding windows along the genome of varying sizes (50kb, 100kb, and 1Mb), and by patterns of excess derived allele sharing along branches of the species tree, as captured by the f-branch test (Malinsky et al. 2021; Malinsky et al. 2018). The authors further investigated the dynamic of cessation of gene flow (and/or ILS) between species using the F1 hybrid PSMC (or hPSMC) approach (Cahill et al. 2016). By estimating the cross-coalescent rates (CRR) between species pairs with time in light of previously estimated species divergence times (McGowen et al. 2020), the authors report that gene flow (and/or ILS) most likely has stopped by now among most species, but it may have lasted for more than half of the time since the species split from each other. According to the author, this result may reflect the slow process by which reproductive isolation would have evolved between cetacean lineages, and that species isolation was marked by significant introgression events.

Now, while the present study provides a good overview of how complex is the reticulated evolutionary history of the Delphinoidae, getting a complete picture will require overcoming a few important limitations. The first ones are methodological and related to the phylogenomic analyses. Given the sampling design with one diploid genome per species, the authors could not phase the data into the parental haplotypes, but instead relied on a majority consensus creating mosaic haploidized genomes representing a mixture between the two parental copies. Moreover, by using large genomic windows (≥50kb) that likely span multiple independent loci, phylogenetic analyses in windows encompassed distinct phylogenetic signals, potentially leading to bias and inaccuracy in the inferences. Thawornwattana et al (2018) previously showed that this “concatenation approach”  could significantly impact phylogenetic inferences. They proposed instead to use loci small enough to minimise the risk of intra-locus recombination and to consider them in blocks of non-recombining loci along the genome in order to conduct phylogenetic analysed, ideally under the multi-species coalescent (MSC) that can account for ILS (e.g. BPP; Flouri et al. 2018; Jiao et al. 2020; Yang 2015). Such an approach applied to the diversification of the Delphinidae may reveal substantial changes compared to the currently admitted species tree.

Inaccuracy in the species tree estimation may have a major impact on the introgression analyses conducted in this study since the species tree and branching order must be supplied in the introgression analyses to properly disentangle introgression from ILS. Here, the authors rely on the tree topology that was previously estimated in McGowen et al. (2020), which they also recovered using their consensus estimation from ASTRAL-III (Zhang et al. 2018). While the methodologies accounted to a certain extent for ILS, albeit with potential bias induced by the concatenation approach, they ignore the presumably large amount of introgression among species during the diversification process. Estimating species branching order while ignoring introgression can lead to major bias in the phylogenetic inference and can lead to incorrect topologies. Even if these MSC-based methods account for ILS, inferences can become very inaccurate or even break down as gene flow increases (see for ex. Jiao et al. 2020; Müller et al. in press; Solís-Lemus et al. 2016). Dedicated approaches have been developed to model explicitly introgression together with ILS to estimate phylogenetic networks (Blair & Ané 2020; Rabier et al. 2021) or in isolation-with-migration model (Müller et al. in press; Wang et al. 2020). Future studies revisiting the reticulated evolutionary history of the Delphinoidae with such approaches may not only precise the species branching order, but also deliver a finer view of the magnitude and prevalence of introgression during the evolutionary history of these species.

A final part of Westbury et al. (2022)'s study set out to test whether historical periods of low abundance could have facilitated hybridization among Delphinoidae species. During these periods of low abundance, species may encounter only a limited number of conspecifics and may consider individuals from other species as suitable mating partners, leading to hybridisation (Crossman et al. 2016; Edwards et al. 2011; Westbury et al. 2019). The authors tested this hypothesis by considering genome-wide genetic diversity (or heterozygosity) as a proxy of historical effective population size (Ne), itself as a proxy of the evolution of census size with time. They also try to link historical Ne variation (from PSMC, Li & Durbin 2011) with their estimated time to cessation of gene flow or ILS (from the CRR of hPSMC). However, no straightforward relationship was found between the genetic diversity and the propensity of species to hybridize, nor was there any clear link between Ne variation through time and the cessation of gene flow or ILS. Such a lack of relationship may not come as a surprise, since the determinants of genome-wide genetic diversity and its variation through evolutionary time-scale are far more diverse and complex than just a direct link with hybridization, introgression, or even with the census population size. In fact, genetic diversity results from the balance between all the evolutionary processes at play in the species' evolutionary history (see the review of Ellegren & Galtier 2016). Other important factors can strongly impact genetic diversity, including demography and structure, but also linked selection (Boitard et al. 2022; Buffalo 2021; Ellegren & Galtier 2016). 

All in all, Westbury and coll. (2022) present here an interesting study providing an additional step towards resolving and understanding the complex evolutionary history of the Delphinoidae, and shedding light on the importance of introgression during the diversification of these cetacean species. Prospective work improving upon the taxonomic sampling, with additional genomic data for each species, considered with dedicated approaches tailored at estimating species tree or network while accounting for ILS and introgression will be key for refining the picture depicted in this study. Down the road, altogether these studies will contribute to assessing the evolutionary significance of introgression on the diversification of Delphinoides, and more generally on the diversification of cetacean species, which still remain an open and exciting perspective. 

References

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Bierne N, Bonhomme F, David P (2003) Habitat preference and the marine-speciation paradox. Proceedings of the Royal Society of London. Series B: Biological Sciences 270, 1399-1406. https://doi.org/10.1098/rspb.2003.2404

Blair C, Ané C (2020) Phylogenetic Trees and Networks Can Serve as Powerful and Complementary Approaches for Analysis of Genomic Data. Systematic Biology 69, 593-601. https://doi.org/10.1093/sysbio/syz056

Boitard S, Arredondo A, Chikhi L, Mazet O (2022) Heterogeneity in effective size across the genome: effects on the inverse instantaneous coalescence rate (IICR) and implications for demographic inference under linked selection. Genetics 220, iyac008. https://doi.org/10.1093/genetics/iyac008

Buffalo V (2021) Quantifying the relationship between genetic diversity and population size suggests natural selection cannot explain Lewontin's Paradox. e-Life 10, e67509. https://doi.org/10.7554/eLife.67509

Cahill JA, Soares AE, Green RE, Shapiro B (2016) Inferring species divergence times using pairwise sequential Markovian coalescent modelling and low-coverage genomic data. Philos Trans R Soc Lond B Biol Sci 371, 20150138. https://doi.org/10.1098/rstb.2015.0138

Crossman CA, Taylor EB, Barrett‐Lennard LG (2016) Hybridization in the Cetacea: widespread occurrence and associated morphological, behavioral, and ecological factors. Ecology and Evolution 6, 1293-1303. https://doi.org/10.1002/ece3.1913

Dagilis AJ, Peede D, Coughlan JM, Jofre GI, D’Agostino ERR, Mavengere H, Tate AD, Matute DR (2021) 15 years of introgression studies: quantifying gene flow across Eukaryotes. biorXiv, 2021.1106.1115.448399. https://doi.org/10.1101/2021.06.15.448399

Degnan JH, Rosenberg NA (2009) Gene tree discordance, phylogenetic inference and the multispecies coalescent. Trends Ecol Evol 24, 332-340. https://doi.org/10.1016/j.tree.2009.01.009

Durand EY, Patterson N, Reich D, Slatkin M (2011) Testing for ancient admixture between closely related populations. Mol Biol Evol 28, 2239-2252. https://doi.org/10.1093/molbev/msr048

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Edelman NB, Mallet J (2021) Prevalence and Adaptive Impact of Introgression. Annual Review of Genetics 55, 265-283. https://doi.org/10.1146/annurev-genet-021821-020805

Edwards CJ, Suchard MA, Lemey P, Welch JJ, Barnes I, Fulton TL, Barnett R, O'Connell TC, Coxon P, Monaghan N, Valdiosera CE, Lorenzen ED, Willerslev E, Baryshnikov GF, Rambaut A, Thomas MG, Bradley DG, Shapiro B (2011) Ancient hybridization and an Irish origin for the modern polar bear matriline. Curr Biol 21, 1251-1258. https://doi.org/10.1016/j.cub.2011.05.058

Ellegren H, Galtier N (2016) Determinants of genetic diversity. Nat Rev Genet 17, 422-433. https://doi.org/10.1038/nrg.2016.58

Flouri T, Jiao X, Rannala B, Yang Z (2018) Species Tree Inference with BPP Using Genomic Sequences and the Multispecies Coalescent. Mol Biol Evol 35, 2585-2593. https://doi.org/10.1093/molbev/msy147

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The Latitudinal Diversity Gradient (LDG) has fascinated natural historians, ecologists and evolutionary biologists ever since [1] described it about 200 years ago [2]. Despite such interest, agreement on the origin and nature of this gradient has been elusive. Several tens of hypotheses and models have been put forward as explanations for the LDG [2-3], that can be grouped in ecological, evolutionary and historical explanations [4] (see also [5]). These explanations can be reduced to no less than 26 hypotheses, which account for variations in ecological limits for the establishment of progressively larger assemblages, diversification rates, and time for species accumulation [5]. Besides that, although in general the tropics hold more species, different taxa show different shapes and rates of spatial variation [6], and a considerable number of groups show reverse patterns, with richer assemblages in cold temperate regions (see e.g. [7-9]).
Understanding such complexity needs integrating ecological and evolutionary research into the wide temporal and spatial perspectives provided by the burgeoning field of biogeography. This integrative discipline ¬–that traces back to Humboldt himself (e.g. [10])– seeks to put together historical and functional explanations to explain the complex dynamics of Earth’s biodiversity. Different to quantum physicists, biogeographers cannot pursue the ultimate principle behind the patterns we observe in nature due to the interplay of causes and effects, which in fact tell us that there is not such a single principle. Rather, they need to identify an array of basic principles coming from different perspectives, to then integrate them into models that provide realistic –but never simple– explanations to biodiversity gradients such as LDG (see, e.g., [5; 11]). That is, rather than searching for a sole explanation, research on the LDG must aim to identify as many signals hidden in the pattern as possible, and provide hypotheses or models that account for these signals. To later integrate them and, whenever possible, to validate them with empirical data on the organisms’ distribution, ecology and traits, phylogenies, fossils, etc.
Within this context, Meseguer & Condamine [12] provide a novel perspective to LDG research using phylogenetic and fossil evidence on the origin and extinction of taxa within the turtle, crocodile and lizard (i.e. squamate) lineages. By digging into deep time down to the Triassic (about 250 Myr ago) they are able to identify several episodes of flattening and steepening of the LDG for these three clades. Strikingly, their results show similar diversification rates in the northern hemisphere and in the equator during the over 100 Myr long global greenhouse period that extends from the late Jurassic to the Cretaceous and early Neogene. During this period, the LDG for these three groups would have appeared quite even across a mainly tropical Globe, although the equatorial regions were apparently much more evolutionarily dynamic. The equator shows much higher rates of origination and extinction of branches throughout the Cretaceous, but they counteract each other so net diversification is similar to that of the northern hemisphere in all three groups. The transition to a progressively colder Earth in the Paleogene (starting around 50 Myr ago) provokes a mass extinction in the three clades, which is compensated in the equator by the dispersal of many taxa from the areas that currently pertain to the Holarctic biogeographical realm. Finally, during the coldhouse Earth’s climatic conditions of the Neogene only squamates show significant positive diversification rates in extratropical areas, while the diversity of testudines remains, and crocodiles continue declining progressively towards oblivion in the whole world.
Meseguer & Condamine [12] attribute these temporal patterns to the so-called asymmetric gradient of extinction and dispersal (AGED) framework. Here, the dynamics of extinction-at and dispersal-from high latitudes during colder periods increase the steepness of the LDG. Whereas the gradient flattens when Earth warms up as a result of dispersal from the equator followed by increased diversification in extratropical regions. This idea in itself is not new, for the influence of climatic oscillations on diversification rates is well known, at least for the Pleistocene Ice Ages [13], as is the effect of niche conservatism on the LDG [14]. Nevertheless, Meseguer & Condamine’s AGED provides a synthetic verbal model that could allow integrating the three main types of processes behind the LDG into a single framework. To do this it would be necessary to combine AGED’s cycles of dispersal and diversification with realistic models of: (1) the ecological limits to host rich assemblages in the colder and less productive temperate climatic domains; (2) the variations in diversification rates with shifts in temperature and/or energy regimes; and (3) the geographical patterns of climatic oscillation through time that determine the time for species accumulation in each region.
Integrating these models may allow transposing Meseguer & Condamine’s [12] framework into the more mechanistic macroecological models advocated by Pontarp et al. [5]. This type of mechanistic models has been already used to understand the development of biodiversity gradients through the climatic oscillations of the Pleistocene and the Quaternary (e.g. [11]). So the challenge in this case would be to generate a realistic scenario of geographical dynamics that accounts for plate tectonics and long-term climatic oscillations. This is still a major gap and we would benefit from the integrated work by historical geologists and climatologists here. For instance, there is little doubt about the progressive cooling through the Cenozoic based in isotope recording in sea floor sediments [15]. Meseguer & Condamine [12] use this evidence for separating greenhouse, transition and coldhouse world scenarios, which should not be a problem for these rough classes. However, a detailed study of the evolutionary correlation of true climate variables across the tree of life is still pending, as temperature is inferred only for sea water in an ice-free ocean, say earlier half of the Cenozoic [15]. Precipitation regime is even less known. Such scenario would provide a scaffold upon which the temporal dynamics of several aspects of the generation and loss of biodiversity can be modelled. Additionally, one of the great advantages of selecting key clades to study the LDG would be to determine the functional basis of diversification. There are species traits that are well known to affect speciation and extinction probabilities, such as reproductive strategies or life histories (e.g. [16]). Whereas these traits might also be a somewhat redundant effect of climatic causes, they might foster (i.e. “extended reinforcement”, [17]) or slow diversification. Even so, it is unlikely that such a model would account for all the latitudinal variation in species richness. But it will at least provide a baseline for the main latitudinal variations in the diversity of the regional communities (sensu [18]) worldwide. Within this context the effects of recent ecological, evolutionary and historical processes, such as environmental heterogeneity, current diversification rates or glacial cycles, will only modify the general LDG pattern resulting from the main processes contained in Meseguer & Condamine’s AGED, thereby providing a more comprehensive understanding of the geographical gradients of diversity.

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