Models applying space-for-time substitution, including those projecting ecological responses to climate change, generally assume an elevational and latitudinal equivalence that is rarely tested. However, a mismatch may lead to different capacities for providing climatic refuge to dispersing species. We compiled community data on zooplankton, ectothermic animals that form the consumer basis of most aquatic food webs, from over 1,200 mountain lakes and ponds across western North America to assess biodiversity along geographic temperature gradients spanning nearly 3,750 metres of elevation and 30 degrees latitude. Species richness, phylogenetic relationships, and functional diversity all showed contrasting responses across gradients, with richness metrics plateauing at low elevations but exhibiting intermediate latitudinal maxima. The nonmonotonic/hump-shaped diversity trends with latitude emerged from geographic interactions, including weaker latitudinal relationships at higher elevations (i.e. in alpine lakes) linked to different underlying drivers. Here, divergent patterns of phylogenetic and functional trait dispersion indicate shifting roles of environmental filters and limiting similarity in the assembly of communities with increasing elevation and latitude. We further tested whether gradients showed common responses to warmer temperatures and found that mean annual (but not seasonal) temperatures predicted elevational richness patterns but failed to capture consistent trends with latitude, meaning that predictions of how climate change will influence diversity also differ between gradients. Contrasting responses to elevation- and latitude-driven warming suggest different limits on climatic refugia and likely greater barriers to northward range expansion.

Records of zooplankton occurrence over a 52-year period (1964 to 2015) were compiled from multiple sources, including published articles, technical reports, and government datasets (previously described by Loewen et al., 2019).

Data were standardized to reflect current taxonomy as per Appendix S1 of Loewen et al., (2019), and 119 taxa (identified to genus to higher) were retained analysis.

REFERENCES

Loewen, C.J.G., Strecker, A.L., Larson, G.L., Vogel, A., Fischer, J.M., & Vinebrooke, R.D. (2019). Macroecological drivers of zooplankton communities across the mountains of western North America. Ecography, 42, 791-803.

Data and code for calculating zooplankton biodiversity metrics and replicating analysis of elevational and latitudinal diversity gradients by generalized linear mixed effect/multilevel models.

Sitebysp.csv contains zooplankton occurrence data (site rows and species columns).

Sitebygeo.csv contains site geographic coordinate data (site rows and variable columns), including: ID (lake ID); Lake (lake name); Elevation (masl); Latitude (N); Longitude (E); Area (km2); Depth (m); Sampling.events (estimated/presumed number of sampling events at each site); and Sampling.years (estimated/presumed number of years sampled at each site).

Spbytrait.csv contains species trait data (species rows and trait/taxonomic columns), including: Rownames (taxa names); (Sp (taxa name); Sp (taxa name); Taxa (taxa name with space); Level (taxonomic level); Order (taxonomic order); Length (taxa length, mm); Length.ref (taxa length reference, see Table S1 in manuscript); Length.taxa (length representative taxa); Feed.ord (feeding guild ordinal representation, 1 = substrate-grazing, 2 = seston-filtering, 3 = stationary suspension-feeding with occasional grasping, and 4 = raptorial-feeding); and Full (full taxa name).

coefficients.bio.metrics.csv contains data for plotting coefficients for biodiversity by elevation/latitude models, including: Metric (biodiversity metric), Group (grouping), Parameter (Coefficient); Class (elevational/latitudinal zone); Estimate (metric estimate); Est.Error (metric estimate error); Q2.5 (2.5% quantile); Q97.5 (97.5 % quantile).

coefficients.clim.csv contains data for plotting coefficients for biodiversity by climate modelsi, ncluding: Metric (biodiversity metric), Climate (climate variable); Group (grouping), Parameter (Coefficient); Class (elevational/latitudinal zone); Estimate (metric estimate); Est.Error (metric estimate error); Q2.5 (2.5% quantile); Q97.5 (97.5 % quantile)

Taxalist.rds contains taxonomic ranks of species (rds file).

Bio.metrics.pub.csv contains processed biodiversity data (site rows and biodiversity metric columns) for use with statistical code, including: ID (lake ID); Lake (lake name); Latitude (N); Elevation (masl); lat_class (site latitudinal zone); elev_class (site elevational zone); Area (km2); Depth (m); Sampling.events (estimated/presumed number of sampling events at each site); Sampling.years (estimated/presumed number of years sampled at each site); Taxo.alpha (species richness); Glob.nbsp (number of taxa excluding juveniles); Glob.FRic (functional richness standardized between zero and one); Glob.FDis (functional dispersion); CWM.Bl (community mean body length, mm); CWM.Feed (community mean feeding guilds); Glob.PD (phylogenetic richness); Glob.MPD (phylogenetic mean pairwise distance); Glob.MPD.beta (phylogenetic mean pairwise distance divided by 100 to fit bounds of zero and one); pd.ntaxa.lat (number of taxa used in calculations for standardized effect sizes of phylogenetic richness in latitudinal zones); pd.obs.lat (observed value used in calculations for standardized effect sizes of phylogenetic richness in latitudinal zones); pd.rand.mean.lat (mean of null communities used in calculations for standardized effect sizes of phylogenetic richness in latitudinal zones); pd.rand.sd.lat (standard deviation of null communities used in calculations for standardized effect sizes of phylogenetic richness in latitudinal zones); pd.obs.rank.lat (rank of observed value used in calculations for standardized effect sizes of phylogenetic richness in latitudinal zones); pd.obs.z.lat (standardized effect sizes of phylogenetic richness in latitudinal zones); pd.obs.p.lat (P-values for observed value used in calculations for standardized effect sizes of phylogenetic richness in latitudinal zones); pd.runs.lat (number of randomizations used in calculations for standardized effect sizes of phylogenetic richness in latitudinal zones); pd.ntaxa.ele (number of taxa used in calculations for standardized effect sizes of phylogenetic richness in elevational zones); pd.obs.ele (observed value used in calculations for standardized effect sizes of phylogenetic richness in elevational zones); pd.rand.mean.ele (mean of null communities used in calculations for standardized effect sizes of phylogenetic richness in elevational zones); pd.rand.sd.ele (standard deviation of null communities used in calculations for standardized effect sizes of phylogenetic richness in elevational zones); pd.obs.rank.ele (rank of observed value used in calculations for standardized effect sizes of phylogenetic richness in elevational zones); pd.obs.z.ele (standardized effect sizes of phylogenetic richness in elevational zones); pd.obs.p.ele (P-values for observed value used in calculations for standardized effect sizes of phylogenetic richness in elevational zones); pd.runs.ele (number of randomizations used in calculations for standardized effect sizes of phylogenetic richness in elevational zones); mpd.ntaxa.lat (number of taxa used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in latitudinal zones); mpd.obs.lat (observed value used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in latitudinal zones); mpd.rand.mean.lat (mean of null communities used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in latitudinal zones); mpd.rand.sd.lat (standard deviation of null communities used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in latitudinal zones); mpd.obs.rank.lat (rank of observed value used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in latitudinal zones); mpd.obs.z.lat (standardized effect sizes of phylogenetic mean pairwise distance in latitudinal zones); mpd.obs.p.lat (P-values for observed value used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in latitudinal zones); mpd.runs.lat (number of randomizations used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in latitudinal zones); mpd.ntaxa.ele (number of taxa used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in elevational zones); mpd.obs.ele (observed value used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in elevational zones); mpd.rand.mean.ele (mean of null communities used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in elevational zones); mpd.rand.sd.ele (standard deviation of null communities used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in elevational zones); mpd.obs.rank.ele (rank of observed value used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in elevational zones); mpd.obs.z.ele (standardized effect sizes of phylogenetic mean pairwise distance in elevational zones); mpd.obs.p.ele (P-values for observed value used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in elevational zones); mpd.runs.ele (number of randomizations used in calculations for standardized effect sizes of phylogenetic mean pairwise distance in elevational zones); FRic.ntaxa.lat (number of taxa used in calculations for standardized effect sizes of functional richness in latitudinal zones); FRic.obs.lat (observed value used in calculations for standardized effect sizes of functional richness in latitudinal zones); FRic.rand.mean.lat (mean of null communities used in calculations for standardized effect sizes of functional richness in latitudinal zones); FRic.rand.sd.lat (standard deviation of null communities used in calculations for standardized effect sizes of functional richness in latitudinal zones); FRic.obs.rank.lat (rank of observed value used in calculations for standardized effect sizes of functional richness in latitudinal zones); FRic.obs.z.lat (standardized effect sizes of functional richness in latitudinal zones); FRic.obs.p.lat (P-values for observed value used in calculations for standardized effect sizes of functional richness in latitudinal zones); FRic.runs.lat (number of randomizations used in calculations for standardized effect sizes of functional richness in latitudinal zones); FRic.ntaxa.ele (number of taxa used in calculations for standardized effect sizes of functional richness in elevational zones); FRic.obs.ele (observed value used in calculations for standardized effect sizes of functional richness in elevational zones); FRic.rand.mean.ele (mean of null communities used in calculations for standardized effect sizes of functional richness in elevational zones); FRic.rand.sd.ele (standard deviation of null communities used in calculations for standardized effect sizes of functional richness in elevational zones); FRic.obs.rank.ele (rank of observed value used in calculations for standardized effect sizes of functional richness in elevational zones); FRic.obs.z.ele (standardized effect sizes of functional richness in elevational zones); FRic.obs.p.ele (P-values for observed value used in calculations for standardized effect sizes of functional richness in elevational zones); and FRic.runs.ele (number of randomizations used in calculations for standardized effect sizes of functional richness in elevational zones); FDis.ntaxa.lat (number of taxa used in calculations for standardized effect sizes of functional dispersion in latitudinal zones); FDis.obs.lat (observed value used in calculations for standardized effect sizes of functional dispersion in latitudinal zones); FDis.rand.mean.lat (mean of null communities used in calculations for standardized effect sizes of functional dispersion in latitudinal zones); FDis.rand.sd.lat (standard deviation of null communities used in calculations for standardized effect sizes of functional dispersion in latitudinal zones); FDis.obs.rank.lat (rank of observed value used in calculations for standardized effect sizes of functional dispersion in latitudinal zones); FDis.obs.z.lat (standardized effect sizes of functional dispersion in latitudinal zones); FDis.obs.p.lat (P-values for observed value used in calculations for standardized effect sizes of functional dispersion in latitudinal zones); FDis.runs.lat (number of randomizations used in calculations for standardized effect sizes of functional dispersion in latitudinal zones); FDis.ntaxa.ele (number of taxa used in calculations for standardized effect sizes of functional dispersion in elevational zones); FDis.obs.ele (observed value used in calculations for standardized effect sizes of functional dispersion in elevational zones); FDis.rand.mean.ele (mean of null communities used in calculations for standardized effect sizes of functional dispersion in elevational zones); FDis.rand.sd.ele (standard deviation of null communities used in calculations for standardized effect sizes of functional dispersion in elevational zones); FDis.obs.rank.ele (rank of observed value used in calculations for standardized effect sizes of functional dispersion in elevational zones); FDis.obs.z.ele (standardized effect sizes of functional dispersion in elevational zones); FDis.obs.p.ele (P-values for observed value used in calculations for standardized effect sizes of functional dispersion in elevational zones); FDis.runs.ele (number of randomizations used in calculations for standardized effect sizes of functional dispersion in elevational zones); pd.obs.p.lat.fix (P-values for observed value used in calculations for phylogenetic richness SES in latitudinal zones replacing values of 1); pd.obs.p.lat.qnorm (probit-transformed quantile P-values for phylogenetic richness SES in latitudinal zones); pd.obs.p.ele.fix (P-values for observed value used in calculations for phylogenetic richness SES in elevational zones replacing values of 1); pd.obs.p.ele.qnorm (probit-transformed quantile P-values for phylogenetic richness SES in elevational zones); mpd.obs.p.lat.fix (P-values for observed value used in calculations for phylogenetic mean pairwise distance SES in latitudinal zones replacing values of 1); mpd.obs.p.lat.qnorm (probit-transformed quantile P-values for phylogenetic mean pairwise distance SES in latitudinal zones); mpd.obs.p.ele.fix (P-values for observed value used in calculations for phylogenetic mean pairwise distance SES in elevational zones replacing values of 1); mpd.obs.p.ele.qnorm (probit-transformed quantile P-values for phylogenetic mean pairwise distance SES in elevational zones); FRic.obs.p.lat.fix (P-values for observed value used in calculations for functional richness SES in latitudinal zones replacing values of 1); FRic.obs.p.lat.qnorm (probit-transformed quantile P-values for functional richness SES in latitudinal zones); FRic.obs.p.ele.fix (P-values for observed value used in calculations for functional richness SES in elevational zones replacing values of 1); FRic.obs.p.ele.qnorm (probit-transformed quantile P-values for functional richness SES in elevational zones); FDis.obs.p.lat.fix (P-values for observed value used in calculations for functional dispersion SES in latitudinal zones replacing values of 1); FDis.obs.p.lat.qnorm (probit-transformed quantile P-values for functional dispersion SES in latitudinal zones); FDis.obs.p.ele.fix (P-values for observed value used in calculations for functional dispersion SES in elevational zones replacing values of 1); FDis.obs.p.ele.qnorm (probit-transformed quantile P-values for functional dispersion SES in elevational zones); pd.ntaxa.glob (number of taxa used in calculations for standardized effect sizes of phylogenetic richness with global pools); pd.obs.glob (observed value used in calculations for standardized effect sizes of phylogenetic richness with global pools); pd.rand.mean.glob (mean of null communities used in calculations for standardized effect sizes of phylogenetic richness with global pools); pd.rand.sd.glob (standard deviation of null communities used in calculations for standardized effect sizes of phylogenetic richness with global pools); pd.obs.rank.glob (rank of observed value used in calculations for standardized effect sizes of phylogenetic richness with global pools); pd.obs.z.glob (standardized effect sizes of phylogenetic richness with global pools); pd.obs.p.glob (P-values for observed value used in calculations for standardized effect sizes of phylogenetic richness with global pools); pd.runs.glob (number of randomizations used in calculations for standardized effect sizes of phylogenetic richness with global pools); mpd.ntaxa.glob (number of taxa used in calculations for standardized effect sizes of phylogenetic mean pairwise distance with global pools); mpd.obs.glob (observed value used in calculations for standardized effect sizes of phylogenetic mean pairwise distance with global pools); mpd.rand.mean.glob (mean of null communities used in calculations for standardized effect sizes of phylogenetic mean pairwise distance with global pools); mpd.rand.sd.glob (standard deviation of null communities used in calculations for standardized effect sizes of phylogenetic mean pairwise distance with global pools); mpd.obs.rank.glob (rank of observed value used in calculations for standardized effect sizes of phylogenetic mean pairwise distance with global pools); mpd.obs.z.glob (standardized effect sizes of phylogenetic mean pairwise distance with global pools); mpd.obs.p.glob (P-values for observed value used in calculations for standardized effect sizes of phylogenetic mean pairwise distance with global pools); mpd.runs.glob (number of randomizations used in calculations for standardized effect sizes of phylogenetic mean pairwise distance with global pools); FRic.ntaxa.glob (number of taxa used in calculations for standardized effect sizes of functional richness with global pools); FRic.obs.glob (observed value used in calculations for standardized effect sizes of functional richness with global pools); FRic.rand.mean.glob (mean of null communities used in calculations for standardized effect sizes of functional richness with global pools); FRic.rand.sd.glob (standard deviation of null communities used in calculations for standardized effect sizes of functional richness with global pools); FRic.obs.rank.glob (rank of observed value used in calculations for standardized effect sizes of functional richness with global pools); FRic.obs.z.glob (standardized effect sizes of functional richness with global pools); FRic.obs.p.glob (P-values for observed value used in calculations for standardized effect sizes of functional richness with global pools); FRic.runs.glob (number of randomizations used in calculations for standardized effect sizes of functional richness with global pools); FDis.ntaxa.glob (number of taxa used in calculations for standardized effect sizes of functional dispersion with global pools); FDis.obs.glob (observed value used in calculations for standardized effect sizes of functional dispersion with global pools); FDis.rand.mean.glob (mean of null communities used in calculations for standardized effect sizes of functional dispersion with global pools); FDis.rand.sd.glob (standard deviation of null communities used in calculations for standardized effect sizes of functional dispersion with global pools); FDis.obs.rank.glob (rank of observed value used in calculations for standardized effect sizes of functional dispersion with global pools); FDis.obs.z.glob (standardized effect sizes of functional dispersion with global pools); FDis.obs.p.glob (P-values for observed value used in calculations for standardized effect sizes of functional dispersion with global pools); FDis.runs.glob (number of randomizations used in calculations for standardized effect sizes of functional dispersion with global pools); pd.obs.p.glob.fix (P-values for observed value used in calculations for phylogenetic richness SES with global pools replacing values of 1); pd.obs.p.glob.qnorm (probit-transformed quantile P-values for phylogenetic richness SES with global pools); mpd.obs.p.glob.fix (P-values for observed value used in calculations for phylogenetic mean pairwise distance SES with global pools replacing values of 1); mpd.obs.p.glob.qnorm (probit-transformed quantile P-values for phylogenetic mean pairwise distance SES with global pools); FRic.obs.p.glob.fix (P-values for observed value used in calculations for functional richness SES with global pools replacing values of 1); FRic.obs.p.glob.qnorm (probit-transformed quantile P-values for functional richness SES with global pools); FDis.obs.p.glob.fix (P-values for observed value used in calculations for functional dispersion SES with global pools replacing values of 1); and FDis.obs.p.glob.qnorm (probit-transformed quantile P-values for functional dispersion SES with global pools).

ClimateNA_input_elev_1964-2015Y.csv contains annual climate data across the entire study period (1964-2015) at each sampling location obtained using ClimateNA v6.40 (Wang et al. 2016).

MAT refers to mean annual temperature (°C) and TD refers to temperature difference between mean warmest month temperature and mean coldest month temperature, or continentality (°C). Year (year); ID1 (lake ID); ID2 (lake region); Latitude (N); Longitude (E); and Elevation (masl). See Wang et al. (2016) for details of other variables.

REFERENCES

Wang, T., Hamann, A., Spittlehouse, D. & Carroll, C. (2016). Locally downscaled and spatially customizable climate data for historical and future periods for North America. PLoS One, 11, e0156720.