DOI: 10.5061/dryad.905qfttpb
title: Data from: Biodiversity patterns diverge along geographic temperature gradients

description: 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.
description: 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. <em>Ecography</em>, 42,
 791-803.
description: 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. <em>PLoS One</em>, 11,
 e0156720.