Glacier influence on bird assemblages in habitat islands of the high Bolivian Andes

Climate projections for the upcoming decades predict a significant loss of ice mass particularly critical for glaciers in tropical mountains. In the dry landscapes of the southern Andes (from Southern Peru to Chile), this global trend has strong ecological impacts on high‐altitude wetlands that support a unique avifauna for feeding, roosting and nesting. As glacier runoffs are expected to affect the area and the quality of wetland habitats, these changes may potentially affect bird communities. To address this issue, we studied the structural and functional diversity of bird assemblages in glacier‐fed high‐altitude wetlands (>4500 m).


| INTRODUC TI ON
Glaciers are sensitive climate indicators and rapidly react to climatic variations with alteration of their mass (Oerlemans, 2001). Glacier retreat has dramatically accelerated during the last decades as a response to a rapid rise of temperatures (Zemp et al., 2019). Worldwide, glaciers have lost ice mass at an average rate of 220 ± 30 Gt/year during the 2006-2015 period (Intergovernmental Panel on Climate Change -IPCC, 2019) and projections for the 21st century indicate a continuing downward trend in glacier cover in almost all regions (Huss & Hock, 2018;IPCC, 2019;Dussaillant et al., 2019). According to the various IPCC scenarios, by 2100, the glacier mass is expected to decrease by 35% to 55% at the global scale and by up to 90% in both Europe and the tropical regions (Huss et al., 2017). A more rapid decline is expected for small glaciers with relatively small accumulation zones (Bosson et al., 2019;IPCC, 2019). Ice loss is the most visible facet of a more profound shift currently underway, affecting multiple components of arctic and alpine systems, from the cryosphere, hydrosphere and pedosphere to the biosphere. Among others, ice loss has vital consequences on the hydrological cycle (flooding/drought), wild and agricultural biodiversity, and human livelihoods Huss et al., 2017).
In mountains, glaciers greatly influence downstream systems, through cold meltwater supply, substrate movement and environmental heterogeneity . Glacier extinction is expected to have both negative and positive consequences on biodiversity (Cauvy-Fraunié & Dangles, 2019). In particular, glacier loss threatens organisms developing on ice (e.g. microorganisms associated with cryoconites; Cauvy-Fraunié & Dangles, 2020), and those dependent on glacier meltwater, such as freshwater microorganisms, invertebrates and fishes (Vincent, 2010, Brighenti et al., 2019 but see Muhlfeld et al., 2020). In turn, recently deglacierized areas offer new colonizable habitats, particularly important for organisms whose lower altitudinal distribution is altered by thermophilization and competition with more competitive taxa (Cauvy-Fraunié et al., 2015;Zimmer et al., 2018).
Birds are prominent and charismatic representatives of mountain biodiversity, in particular in the tropical region (Monasterio & Vuilleumier, 1986), yet we know very little on the potential impacts of glacier retreat on avian communities. While bird richness tends to decrease with elevation (McCain, 2009), they are still abundant and diverse in high-elevation ecosystems in the tropical region (Fjeldså et al., ) and even show peaks of taxonomic and functional diversity at mid-to-high elevations (Altamirano et al., 2020). Birds have adapted their metabolism and behaviour to high-elevation stress related to flight cost, energetic stress and temporal and spatial variation in resource availability (Lague et al., 2017;Wolf & Gill, 1986). In the high tropical Andes, wetland and grassland habitats provide both food resources and stopover sites during bird migration (Fjeldså, 1985;. On glaciers, birds feed on prey that are highly visible, hence easy to detect and catch. Moreover, once the bird dies, feathers and droppings may provide nutrients for ice-dwelling microorganisms (Rosvold, 2016). Birds also fulfil several roles as herbivores, pollinators, predators and dispersers of seeds and invertebrates . Their presence and activity is therefore crucial for maintaining key ecological functions and ecosystem services (e.g. provisioning and cultural) and reciprocally benefit from habitat type and vegetation heterogeneity interfacing with anthropogenic factors (Araneda et al., 2018).
The decrease in glacier cover and its consequences on water availability may have both direct and indirect effects on mountain birds. Direct effects may concern birds that, regularly or occasionally, nest directly on ice (e.g. the White-winged Diuca-Finch Idiopsar speculifer and the White-fronted Gound-Tyrant Muscisaxicola albifrons; Figure 1e), roost in crevasses or voids beneath the glacier, foraging on nearby cushion plants (e.g. the seedsnipe Attagis gayi) or feed on glacier invertebrates (Hardy et al., 2018;Hotaling et al., 2020).
Indirect effects may concern bird assemblages through three main mechanisms: (a) change in habitat area (either an increase in glacier forefield terrestrial habitats or a reduction in wetlands, streams and lakes due to decrease in glacier runoff, Jacobsen & Dangles, 2017), (b) a reduction in habitat heterogeneity (micro-habitats with different characteristics, in particular aquatic ones, Quenta et al., 2016), and (b) a modification of habitat quality (e.g. productivity, temperature and humidity; Huss et al., 2017). The direction and intensity of those effects on bird assemblages would depend on species' life history and functional traits.
Here, we investigated the potential indirect effects of retreating glaciers on avian biodiversity in high-altitude wetlands in the Bolivian Andes (>4500 m). These ecosystems are ideal to look into the effects of changing habitat area, heterogeneity and quality associated with glacier retreat on biodiversity based on the spatial comparisons. First, being "wetlands in drylands," they form welldelimited habitat islands that allow building predictive models linking biological diversity and habitat area (species-area relationships; Anthelme et al., 2014;Rosvold, 2016). Second, the environmental heterogeneity, productivity and moisture of these wetlands are strongly coupled with glacier influence (Cooper et al., 2019;Dangles et al., 2017). It is therefore possible to infer future glacier retreat of these systems from contemporary spatial patterns in wetlands showing a gradient of glacier influence (i.e. space-for-time substitution, Blois et al., 2013). Third, a large number of bird species present on these wetlands are absent from the surrounding habitats and at lower elevation. Finally, these systems are of global conservation relevance as they support important populations of bird species, in particular during the dry season (Gibbons et al., 2016;Tellería et al., 2006). They also serve as stepping-stone ecosystems during the migration and provide connectivity between structurally similar habitats (Venero, 1987;Vuilleumier, 1970). In particular, the role of high-altitude wetlands as migration corridors increases as the quantity and quality of low-elevation riparian habitats decrease . The specific objectives of this study were to: (a) determine the relationship between habitat area and bird diversity using different metrics (species richness, evenness, functional and phylogenetic diversity) across a range of wetlands with different glacial influence; (b) assess the influence of environmental variables (e.g. elevation, glacier cover in catchment, productivity, heterogeneity) on bird richness at the wetland scale; and (c) determine specific bird assemblages associated with different wetland habitat quality in a context of glacier retreat.

| Study sites
The study area was located in the western side of the Cordillera Real of Bolivia (between 15°45′-16°45′S, and 67°40′-68°40′W, Figure 1a). This mountainous area lengthens over 130 km from south-southeast to north-northwest, with an approximate width of 15 ± 20 km. Several summits rise to over 5000 m a.s.l. and are dominated by relatively small glaciers (80% of glaciers <0.5 km 2 ; Dangles et al., 2017). Glaciers of the Cordillera Real represented an area of approx. 324 km 2 in 1975 but have lost about 48% of their surface and 43% of their volume between 1975 and 2006 (Soruco et al., 2009, see also deglacierization maps in the study area: Zimmer et al., 2018 and Figure S1). During the last decades, glacier melting has been more intense in this region than outside the tropics (Rabatel et al., 2013) with a relative ice loss rate reaching 3%/year at low latitude between 2006 and 2016 (Zemp et al., 2019). Present and future climatic features in the study region have been described in details by Dangles et al. (2017). Briefly, annual mean temperature is around 5°C and incident solar radiation remains intense all year long due to low latitude. In contrast, seasonal variation in moisture and precipitation produces drier at- patterns as a function of elevation and latitude Rabatel et al., 2013).
High-altitude wetlands of the tropical Andes, locally named bofedales, can be described as heterogeneous waterscapes that enclose meadows, marshes, peatlands, ponds and streams perennial or temporary depending on precipitation patterns Squeo et al., 2006). In the Andes, reported trends indicate an overall increase in wetlands cover, strongly correlated with precipitation trends Pauca-Tanco et al., 2020). Proportion of different sources of water is highly variable among wetlands as function of topography, size of the watershed, extent of the glacier, infiltration vs. surface circulation of ice runoff (see Cooper et al., 2019;Squeo et al., 2006). There are differences among seasons, as during the dry season up to 40% of the water production in the Tuni-Condoriri valley is of glacier origin (see Soruco et al., 2015). Like other alpine ecosystems, high Wetlands are scattered with ponds, varying in size, shape and depth . They sustain a high aquatic biodiversity (algae, macrophytes, macroinvertebrates, zooplankton and phytoplankton), with some taxa being highly sensitive to environmental changes and therefore potentially indicators of glacier changes (Quenta et al., 2016).

| Bird survey
We surveyed 40 wetlands in five valleys of the Cordillera Real: Hichu Khota, Palcoco, Huayna Potosí, Tuni and Condoriri ( Figure 1a). We chose wetlands above 4500 m in order to exclude impacts from human activities such as villages, mining activities, overgrazing, water collection and artificial lakes. We selected wetlands across a wide range of areas (from 0.3 to 42.3 ha) and glacier influence (from 0 to 22% of glacier cover in the catchment) to estimate the indirect effects of glacier retreat through changes in habitat area and quality. All wetlands were separated by at least 500 m and therefore considered as independent units (Ralph, 1993). Bird census was carried out between October 2013 and July 2014 to encompass the wet, transitional and dry seasons.
Bird richness and abundance were determined by the point count method (Bibby et al., 2000), with unlimited observation distance and three equidistant observation points and six temporal repetitions per wetland. Count duration was 10 min at each observation point, that is 30 min per wetland per visit. In view of the openness of the habitat, this sampling effort allows to survey different area keeping a satisfactory detectability of species present (Hutto et al., 1986;Savard & Hooper, 1995;Wunderle, 1994). In total, we obtained 720 observation points over the 40 wetlands. We alternated the order of visit to cover a broad daytime range in each wetland and minimize potential bias resulting from daily variation in bird occurrence ).

| Environmental variables
We characterized wetlands using both field measurements and database compilations. First, we evaluated environmental features in the field by recording habitat types along 10 randomly oriented transects of 50-m long for small wetlands (<5 ha) and 20 transects for larger ones. Every 1 m in each transect, 21 categories were considered as representative of habitat diversity and recorded in a total of 27,000 points from 540 transects, corresponding to a sampling effort of 240 observer-hour . We regrouped habitat types in 11 resuming categories as follows: Distichia cushion, Oxychloe cushion, Phylloscirpus/Aciachne cushion, Pycnophyllum cushion, forbs, graminoid peatland, bryophyte, marsh (marsh and macrophytes), water bodies and flowing water (see Table S1).
Second, area and perimeter of the 40 wetlands were extracted from 2013 PLEIADES images using ArcGis 10.6 (see Dangles et al., 2017). In addition, we considered the following environmental distance to the nearest lake, calculated in meters with the distance tool in ArcGis10.6; (d) productivity index, using the median normalized difference vegetation index (NDVI) as a proxy (Bonthoux et al., 2018;Paruelo et al., 1997) and calculated from Pleaides satellite images near infra-red and red bands, ranging from −1 to 1 (see Kraemer, 2014); (e) shape index (SI), a proxy of stability that quantifies irregularly shaped fragments of wetlands that would be more susceptible to changes . SI was calculated by dividing the unit perimeter by that of an equal-sized circle SI = P/2√πA, P being the perimeter of each wetland and A the area; (f) wetland environmental heterogeneity, calculated as the Simpson diversity index (Lande, 1996;Yoshioka et al., 2017).
Heterogeneity index was calculated on 11 reclassified groups of habitat types from the 21 original categories measured on the field transects (see Table S1). Ranging from 0 to 1, this index reflects the inverse probability of each habitat types to be present in the same wetland: the highest value meaning a higher diversity of habitats and thus a lower probability to come across the same habitat within a given wetland.

| Bird taxonomic, phylogenetic and functional diversity
We calculated bird diversity as the total number of bird species per wetland and quantified the evenness using the probability of interspecific encounter (PIE, Hurlbert, 1971). Data were log-transformed to meet normality criteria. We assessed phylogenetic and functional diversity following the mean pairwise distance (MPD) approach from Swenson (2014) using the Picante R package (v1.8 Kembel et al., 2010). On one side, MPD is a measure of phylogenetic diversity based on the average of all distances connecting species together in a sample (Cadotte et al., 2012;Webb et al., 2002). For phylogenetic data, we used a subset of phylogenetic trees downloaded from birdtree.org (Jetz et al., 2012). These trees were constructed from the wetlands speciesˈ total pool and pruned to include corresponding species present in each wetland. We calculated phylogenetic diver- Foraging stratum and body weight were not part of the trait analysis as there is no significant vertical structure in the studied ecosystems and because weight incorporates variation in both size and fitness and may therefore be redundant with other traits (Gosler, 2004).
We calculated the dominant principal components of the functional distance matrix with the Gower distance measure (Gower, 1971) and performed a principal coordinates analysis (PCoA) to reduce dimensionality of functional diversity (Maire et al., 2015).
We then performed the FD MPD analysis on the subset of the PCoA as described on Swenson (2014) and obtained a FD MPD value for each wetland. MPD represents a measure of functional diversity weighted by species relative abundance in the community (Laliberté & Legendre, 2010;Mouchet et al., 2010), that is the deviation of trait values from the centre of the functional space.

| Species-area relationship
Considering the classical biogeography study on the relationship be-  Burnham & Anderson, 2002). Model validation was also based on tests of the normality and homoscedasticity of the residuals (see Table S2).
Additionally, we tested for differences in the SAR among models using analyses of covariance (ANCOVAs) on species richness as a function of season while keeping area as a covariate (Fattorini et al., 2017). Finally, we tested the small-island effect (SIE) using the sar_threshold function of the sars R package v1.3.5 (Matthews et al., 2019). This pattern describes SAR changing below a certain threshold area at which species richness varies independently of area (Lomolino & Weiser, 2001), and the pattern of species richness increasing at lesser rate in small than in larger islands (Dengler, 2010;Wang et al., 2018).
We suggesting trait-based assembly (Petchey et al., 2007;Ross et al., 2019). SES also allows evaluating the relative strength of the assembly processes (i.e. the effect of area on diversity), with the significance threshold value fixed to 1.96 corresponding to the 95% confidence interval for the null distribution (Veech, 2012). Thus, in the case of phylogenetic assembly, species might occur with closely related species more than expect by chance (Webb 2000). Likewise, non-random functional distribution indicates assembly processes like environmental filtering or competitive exclusion (Concepción et al., 2017).

| Generalized additive models
We used generalized additive models (GAMs) to explore the links between bird diversity and the environmental factors associated with wetlands (Hastie & Tibshirani, 1987). These models allow considering environmental factors through a range of nonparametric trends of increasing complexity, leading to understand relationships patterns between predictors and response variables (Wood, 2015). We ran the GAMs with the gam function of the mgcv R package (Wood & Wood, 2015), using species richness as a response variable and environmental data as predictors, including the following: elevation, nearest distance to lakes, glacier cover (%GCC), productivity, heterogeneity and shape index (see index description above and Table S5).
After controlling for multicollinearity among predictors (concurvity function; Wood 2017), we set different GAMs under Poisson distribution and log link functions with four variables: elevation, glacier cover, heterogeneity and productivity. After evaluating correlation coefficients between predictor variables using pairwise diagnostic tools, two variables (shape index and nearest distance to lakes) were excluded for having marginal contribution (i.e. variables not bringing additional significant information to the model).

| Canonical correspondence analysis
We assessed the response of bird species and guilds (see below) to changes in specific habitat variables (21 habitats within wetlands) by performing a canonical correspondence analysis (CCA) (Ter Braak, 1986). We first transformed bird abundances into presence/absence data to better capture the influence of environmental predictors on species composition among and within wetlands (Cushman & McGarigal, 2004). We defined seven groups of birds (guilds) based on their use of environmental resources, that is "aquatic" and "terrestrial" and their feeding guilds (see Table S4). We then used the cca function of the Vegan package (Oksanen et al., 2019) to pair the matrix of presence/absence of bird species with six environmental variables (area, glacier cover, elevation, productivity, heterogeneity, distance to lake), then with the 11 habitats type matrix of each wetland. We tested CCA significance with an ANOVA. For the habitat matrix, we included the 11 habitat types regrouping the 21 coverage recorded on field transects (see above, Environmental variables).

| RE SULTS
Across the 40 study wetlands, we recorded a total of 2858 individuals from 41 species, representing 22 families of birds (Table S4). Among the SAR models fitted to our data (see Tables S2.a  slope values (see Figure S3 and Table S3).
Among the four environmental variables considered, GAMs showed significant non-linear relationship between species richness and glacier cover and elevation ( Figure 3; Table 1). The first model, built with one parametric term, productivity and three smooth terms,

| DISCUSS ION
Following the objectives of our study (see introduction), we found Hereafter, these results are discussed in light of the relationship between wetland area and bird communities in melting Andes and the extent and quality of habitats. Last, we discuss the implications of our study in terms of bird conservation in the region.

| Wetland area and bird communities in melting Andes
Building models depicting species-area relationship (SAR) is a key preliminary step to assess the potential effect of wetland area changes on bird communities. In the Bolivian Andes, an increase in high-elevation wetland surfaces over the last 30 years has coincided with an increase in water availability due to glacier melting rates . This observed positive trend in wetland cover is likely to be reversed in a near future as the current increase in glacier runoff is expected to slow until it reaches its maximum and then start a pronounced decline diminishing glacier influence on outflows (Baraer et al., 2012;Huss & Hock, 2018). Wetland area will also depend on future trends in precipitations, which are highly uncertain in the tropical Andes (Vuille et al., 2008, also see Jacobsen . While waterbirds may benefit glacier melting and associated increase in wetland area, annual glacial runoff is expected to decrease after a critical threshold of reduction in ice volume (Baraer et al., 2012;Rabatel et al., 2013). This would induce an increase in low-flow period frequency and intensity followed by continuous low flows until the complete loss of glacier outflow when the glacier disappears. In this case, the future of waterbirds in bofedales will tightly associate with precipitation patterns.
Our study revealed that wetland area is a good predictor of spe-   (6) Family: Poisson s(heterogeneity) Link fun: log Note: Per cent deviance explained refers to the whole model scores, the p-value is indicated for each descriptor and k refers to the basis dimension of each parameter. The models tested species richness as response for the follow predictor variables: glacier cover (percentage of glacier in the catchment area), elevation (m.a.s.l.), productivity (NDVI index) and heterogeneity (heterogeneity index).

F I G U R E 4
Canonical correspondence analysis (CCA). The biplot displays the ordination of Andean wetlands bird species (points) along the first two canonical axis and their correlation with local habitat variables (arrows). Arrows' direction and length show the degree of correlation between bird's species and habitat types. Point size and colour reflect species total abundances on log(1+x) scale and birds' guilds, respectively. Feeding groups are listed in Supporting Information (Table S4). First axis present habitat productivity gradient and second axis water availability gradient. The blue arrow represents the glacier loss gradient. Decrease in water availability display changes in vegetation, with partial drying, more compact soils and less water storage capacity of Phylloscirpus and Aciachne cushions and their associated plants. Changes in wetland vegetation affects diversity indirectly through changes in dominant species and resource availability less isolated compared to oceanic islands (Matthews et al., 2016). It was also lower compared to previous studies on birds in the northern Andes páramo (0.28; Vuilleumier, 1970) and Argentinian Sierras Pampanas (0.36;Nores, 1995). These SAR concern different spatial scales (within mountain-top habitats) and ecological realities (viable populations, habitat selection), which implies that z values can be expected to vary greatly. These lower values may also reflect the overall low species richness in this part of the Andes, compared to other elevations (Araneda et al., 2018). SAR can also be influenced by the taxonomic group (Triantis et al., 2012, Sólymos & Lele 2012, with different responses to SAR between specialist and generalist species (Lehikoinen et al., 2019;Matthews et al., 2016). Thus, our results suggest that wetland area reduction resulting from glacier retreat would have higher consequences for the conservation of aquatic birds, as reflected by its higher z value (0.32) (see Figure   S2). Despite slight differences among fitted SAR models, the log-log linear parameters are more easily interpretable (Rosenzweig 1995) and can help potential subsequent comparisons with other environmental contexts and taxonomic groups on regional or global scales.
Last, SAR can also be influenced by the scale of the area parameter, notably on smaller islands deviating from the overall pattern (SIE; Lomolino, 2000). However, we found no conclusive evidence of such effect for the studied wetlands (see Figure S3).

Standardized effect size (SES) and slope values on FD and PD
MPD analyses suggest that bird communities of high Andean wetlands were not phylogenetically different from randomly assembled communities nor that functional assembly was non-random, however suggesting a trend towards trait-driven arrangement.
The scaling of functional diversity with area might reflect different assembly processes that are expected to occur at different spatial scales (Concepción et al., 2017). As differences in food preferences and in use of habitat space may reduce competition and permit coexistence , competition among ecologically similar species might involve stronger selection pressure in smaller wetlands because of lower niche availability within wetlands (Si et al., 2016;Zhang et al., 2020). This may affect range-restricted species as those restricted to glacier and periglacier margins (e.g. the White-winged Glacier Finch, Idiopsar speculifer). At a larger scale, increasing proportions of certain traits could lead to amplify the homogenization of high Andean bird communities (increasing levels of functional redundancy; Altamirano et al., 2020;Mouillot et al., 2013;Petchey et al., 2007). Thus, rising abundance of species related to puna gramineae habitat and feeding on seeds (e.g. the Sierra Finches Geospizopsis unicolor and G. plebejus) might indicate the progression of drier habitats and loosing of typical wetland conditions. When related to glacier retreat, this situation suggests that ongoing changes in functional diversity might be part of the environmental filtering on some specific traits such as those related to high glacier influenced environments (e.g. aquatic plants and invertebrate-feeding species or ice-breeding). This calls attention to the importance of ecosystem resilience assured by functional diversity adjusting shared similar functions among species in response to environmental change and hence buffer ecosystem function diminution (Concepción et al., 2017).

| Extent and quality of habitats
While area appears to be a key driver of bird assemblages in highaltitude wetlands, other environmental variables may come into play. In particular, environmental factors related to temperature and system stability (e.g. water provision by the glacier) may influence species composition through resource constraints and habitat complexity (McCain, 2009;Worm & Tittensor, 2018;Zhang et al., 2020). NDVI has been long used as proxy of productivity of Andean wetlands as it is sensitive to the greenness of photosynthetically active biomass (Anderson et al., 2021;Chávez et al., 2019;Moreau et al., 2003). However, these sat-

| Conservation implications in melting Bolivian Andes
Bird survey in high-altitude wetlands might be an initial step for longterm ecosystem monitoring under climate change context. We hypothesized that glacier retreat may affect bird assemblages through changes in wetland area and habitat quality. Larger wetlands showed greater number of individual bird species, although with different occurrences (see Figure S7). Within wetlands, some habitats such as ponds or cushions (e.g. Distichia muscoides) may contribute more to total bird diversity than other habitats (Josens et al., 2017;Servat et al., 2018). We indeed found that local habitat complexity (i.e. forbs, Furthermore, facing an increasing dominance of birds of the drier habitats, rare or low-dispersion species might deeply affect wetland functioning with potential cascading effects at higher and lower trophic levels (Franzén et al., 2012;Holt, 2010). Under this scenario, the interplay between habitat productivity (as a response to increasing temperature) and humidity (dependent on future precipitation patterns) will be key to predict the fate of bird communities in high Andean wetlands with decreasing runoff from glaciers. In this context, it is important to document ongoing changes in alpine plant communities (such as tussock and cushion plants; Meneses et al., 2014;Zimmer et al., 2018) as vegetation provides fundamental feeding resources for many birds, both in terrestrial and in aquatic habitats. Andean bogs also increase habitat suitability for specific wetland bird species (aquatic invertebrates and plants feeding gilds) and might help several species to resist extinction (Gibbons et al., 2016;Tellería et al., 2006).
All this advocates for a more integrating and across-taxa analysis of the ecological consequences of glacier retreat.

CO N FLI C T O F I NTE R E S T
The authors declare no conflicts of interest.

PEER R E V I E W
The peer review history for this article is available at https://publo ns.com/publo n/10.1111/ddi.13458.

DATA AVA I L A B I L I T Y S TAT E M E N T
Data are openly available in Supplementary Information.

B I OS K E TCH
The authors are part of a French-Bolivian research team with various expertise (ornithology, botany, entomology, glaciology, hydrology, geography, statistics) that has been working for the last eight years on biodiversity and conservation issues in highaltitude wetlands of the tropical Andes.
Author contributions: KN, RIM, FA and OD conceived the ideas and monitoring design; KN, CML MIG, RIM, FA, and OD collected data; TC, QS, KN, SC-F organized and analysed data; TC and OD led the writing of the manuscript. All authors contributed critically to drafts and gave final approval for publication.

S U PP O RTI N G I N FO R M ATI O N
Additional supporting information may be found in the online version of the article at the publisher's website.