Living up to thermal potentials
Nikki Moore
09/07/2021
Data coverage
| Realm | Number of species |
|---|---|
| Terrestrial | 364 |
| Marine | 51 |
| Intertidal | 21 |
| Thermal limit | Number of species |
|---|---|
| tmax | 206 |
| tmax, tmin | 186 |
| tmin | 44 |
| Class | n |
|---|---|
| Reptilia | 275 |
| Amphibia | 60 |
| Actinopterygii | 26 |
| Bivalvia | 21 |
| Insecta | 15 |
| Arachnida | 11 |
| Malacostraca | 8 |
| Gastropoda | 6 |
| Echinoidea | 4 |
| Archelosauria | 2 |
| Ascidiacea | 2 |
| Articulata | 1 |
| Asteroidae | 1 |
| Lepidosauria | 1 |
| Neogastropoda | 1 |
| Polyplaclophora | 1 |
| Rhynchonellata | 1 |
Species traits data
| Trait | Variable type | Values | Number of species missing trait |
|---|---|---|---|
| Maximum body length | continuous | length (cm) | 7 |
| Trophic position | categorical | carnivore, omnivore, herbivore, fungivore, insectivore | 0 |
| Migratory? | categorical | Non-migratory, Migratory | 0 |
| Dispersal distance category | categorical, but numeric | 10-100, 0-1, 1-10, 100+, NA | 4 |
| Dispersal ability category | categorical | pelagic development, walking, slithering, pelagic development and crawling adults, pelagic development and sessile adults, pelagic development and swimming adults, non-pelagic development and crawling adults, non-pelagic development and sessile adults, NA, non-pelagic development and swimming adults, swimming, crawling | 3 |
| Cold season dormancy? | categorical | No, Yes | 0 |
| Hot season dormancy? | categorical | No, Yes | 0 |
| Latitudinal midpoint of realized range | continuous | latitude (degrees) | 0 |
| Trait | Variable type | Values | Number of species missing trait |
|---|---|---|---|
| Pelagic habitat type | categorical | benthic, NA, demersal, benthopelagic, reef-associated, pelagic, coral associated | 0 |
| Upper depth limit | continuous | depth (m below sea level) | 8 |
| Lower depth limit | continuous | depth (m below sea level) | 7 |
| Coastal or oceanic | categorical | coastal, NA, coastal-oceanic?, coastal_oceanic, coastal?, coastal-oceanic, oceanic? | 0 |
| Trait | Variable type | Values | Number of species missing trait |
|---|---|---|---|
| Upper depth limit | continuous | depth (m below sea level) | 9 |
| Lower depth limit | continuous | depth (m below sea level) | 9 |
| Trait | Variable type | Values | Number of species missing trait |
|---|---|---|---|
| High-elevation species (>1000m) | categorical | NA, non-highland specialist, , highland specialist, No | 0 |
| Lower elevation limit | continuous | elevation (m above sea level) | 164 |
| Upper elevation limit | continuous | elevation (m above sea level) | 161 |
| Trait | Number of species with inferred trait |
|---|---|
| Hot season dormancy | 352 |
| Cold season dormancy | 188 |
| Dispersal distance category | 39 |
Model fitting
Cold niche filling
filling_value ~ abs_lat_mp*realm + metric
random = ~1|Class/Order/Family/Genus
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | 1.4599766 | -5.1400566 | 8.0600097 | 3.3322965 | 116 | 0.4381292 | 0.662 | |
| abs_lat_mp | 0.1917942 | 0.0808339 | 0.3027545 | 0.0560229 | 116 | 3.4234991 | 0.001 | *** |
| realm: Intertidal | -18.1209468 | -38.4362615 | 2.1943679 | 8.3024302 | 6 | -2.1826076 | 0.072 | . |
| realm: Marine | -1.3151807 | -17.1360707 | 14.5057093 | 6.4656559 | 6 | -0.2034103 | 0.846 | |
| metric: lt | -1.8710476 | -7.9337422 | 4.1916470 | 3.0226632 | 53 | -0.6190063 | 0.539 | |
| abs_lat_mp:realmIntertidal | 0.4693985 | 0.1338021 | 0.8049950 | 0.1694396 | 116 | 2.7703003 | 0.007 | ** |
| abs_lat_mp:realmMarine | -0.1218230 | -0.4132322 | 0.1695862 | 0.1452872 | 53 | -0.8384976 | 0.406 |
Results:
- Cold niche overfilling increases with latitudinal midpoint of the realized range across all realms
- Largest effect size for intertidal species: ~0.66 degrees C per degree latitude
- Terrestrial species: ~0.19 degrees C per degree latitude
- Not significant for marine species: ~0.07 degrees C per degree latitude
- Cold niche filling is not significantly different for critical versus lethal limits
filling_value ~ abs_lat_mp*realm + log_range_area_km2*realm +cold_season_dormancy_ + log_maximum_body_size + dispersal_distance_continuous + Trophic_position + metric
random = ~1|Class/Order/Family/Genus
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | -27.3154631 | -38.0961275 | -16.5347988 | 5.4382339 | 107 | -5.0228556 | <0.001 | *** |
| abs_lat_mp | 0.2784471 | 0.1836787 | 0.3732155 | 0.0478053 | 107 | 5.8246101 | <0.001 | *** |
| realm: Intertidal | -124.2980165 | -200.2480824 | -48.3479507 | 31.0391508 | 6 | -4.0045560 | 0.007 | ** |
| realm: Marine | 2.6568881 | -43.3382056 | 48.6519819 | 18.7972009 | 6 | 0.1413449 | 0.892 | |
| cold_season_dormancy_: Yes | -5.5022243 | -7.7854568 | -3.2189918 | 1.1517613 | 107 | -4.7772261 | <0.001 | *** |
| log_maximum_body_size | -0.2192102 | -1.5347168 | 1.0962963 | 0.6635985 | 107 | -0.3303356 | 0.742 | |
| dispersal_distance_continuous | -0.0002722 | -0.0068482 | 0.0063039 | 0.0033173 | 107 | -0.0820490 | 0.935 | |
| Trophic_position: omnivore | -1.8202767 | -5.1111250 | 1.4705717 | 1.6600464 | 107 | -1.0965216 | 0.275 | |
| Trophic_position: insectivore | -0.8062955 | -4.9470689 | 3.3344779 | 2.0887854 | 107 | -0.3860117 | 0.7 | |
| Trophic_position: carnivore | -1.0709172 | -5.6322998 | 3.4904655 | 2.3009589 | 107 | -0.4654221 | 0.643 | |
| log_range_area_km2 | 2.2103452 | 1.8098935 | 2.6107969 | 0.2020052 | 107 | 10.9420215 | <0.001 | *** |
| metric: lt | -1.1998429 | -7.4888271 | 5.0891412 | 3.1326136 | 51 | -0.3830166 | 0.703 | |
| abs_lat_mp:realmIntertidal | 0.1249786 | -0.2225437 | 0.4725009 | 0.1753053 | 107 | 0.7129197 | 0.477 | |
| abs_lat_mp:realmMarine | -0.2304109 | -0.4938098 | 0.0329879 | 0.1312019 | 51 | -1.7561555 | 0.085 | . |
| realmIntertidal:log_range_area_km2 | 8.5501454 | 4.0506236 | 13.0496672 | 2.2697536 | 107 | 3.7669927 | <0.001 | *** |
| realmMarine:log_range_area_km2 | -0.3517969 | -2.5905707 | 1.8869769 | 1.1151583 | 51 | -0.3154681 | 0.754 |
Results:
- Cold niche overfilling increases with latitudinal midpoint of the realized range across all realms
- Largest effect size for intertidal species but not significant: ~0.40 degrees C per degree latitude
- Terrestrial species: ~0.28 degrees C per degree latitude
- Not significant for marine species: ~0.05 degrees C per degree latitude
- Cold niche overfilling increases with increasing realized range area across all realms
- Largest effect size for intertidal species: ~10.76 degrees C per log(km2)
- Terrestrial species: ~2.21 degrees C per log(km2)
- Not significant for marine species: ~1.86 degrees C per log(km2)
- Cold niche filling is not significantly different for critical versus lethal limits
- Species that undergo cold season dormancy have significantly less cold niche overfilling
- Cold niche overfilling decreases with increasing body length, but not significantly
- ~ -0.22 degrees C per log(cm)
- Effect of trophic position not significant, but cold niche overfilling is greatest in: herbivore, insectivore, carnivore, omnivore
Warm niche filling
filling_value ~ abs_lat_mp*realm + metric
random = ~1|Class/Order/Family/Genus
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | 1.8022343 | -3.0016194 | 6.6060881 | 2.4343267 | 178 | 0.7403420 | 0.46 | |
| abs_lat_mp | -0.1523705 | -0.2159041 | -0.0888369 | 0.0321953 | 178 | -4.7326919 | <0.001 | *** |
| realm: Intertidal | 0.1274967 | -10.7258227 | 10.9808162 | 5.4998604 | 178 | 0.0231818 | 0.982 | |
| realm: Marine | -6.9484877 | -13.6055573 | -0.2914182 | 3.3734337 | 178 | -2.0597671 | 0.041 | * |
| metric: lt | -0.2922067 | -3.7196973 | 3.1352839 | 1.7368622 | 178 | -0.1682383 | 0.867 | |
| abs_lat_mp:realmIntertidal | 0.0417772 | -0.1751521 | 0.2587065 | 0.1099277 | 178 | 0.3800425 | 0.704 | |
| abs_lat_mp:realmMarine | 0.2175485 | 0.0978068 | 0.3372901 | 0.0606784 | 178 | 3.5852692 | <0.001 | *** |
| Results: |
- Warm niche underfilling increases with latitudinal midpoint of the realized range in terrestrial and intertidal systems, decreases with latitude in marine systems
- Largest effect size for terrestrial species: ~ -0.15 degrees C per degree latitude
- Intertidal species: ~ -0.11 degrees C per degree latitude
- Not significant for marine species: ~0.07 degrees C per degree latitude
- Warm niche filling is not significantly different for critical versus lethal limits
filling_value ~ abs_lat_mp*realm + log_range_area_km2*realm + hot_season_dormancy_ + log_maximum_body_size + dispersal_distance_continuous + Trophic_position + metric
random = ~1|Class/Order/Family/Genus
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | -8.8929583 | -15.1577535 | -2.6281630 | 3.1733574 | 168 | -2.8023815 | 0.006 | ** |
| abs_lat_mp | -0.1623640 | -0.2231235 | -0.1016046 | 0.0307770 | 168 | -5.2755039 | <0.001 | *** |
| realm: Intertidal | 14.8206178 | -26.6655047 | 56.3067402 | 21.0143013 | 168 | 0.7052634 | 0.482 | |
| realm: Marine | -18.6022748 | -34.3757049 | -2.8288446 | 7.9898432 | 168 | -2.3282403 | 0.021 | * |
| hot_season_dormancy_: Yes | -2.9092083 | -5.9590563 | 0.1406397 | 1.5448642 | 168 | -1.8831483 | 0.061 | . |
| log_maximum_body_size | 0.1670771 | -0.5889396 | 0.9230938 | 0.3829513 | 168 | 0.4362881 | 0.663 | |
| dispersal_distance_continuous | 0.0019233 | -0.0011587 | 0.0050052 | 0.0015611 | 168 | 1.2319693 | 0.22 | |
| Trophic_position: omnivore | 1.5088632 | -1.1638446 | 4.1815710 | 1.3538283 | 168 | 1.1145159 | 0.267 | |
| Trophic_position: fungivore | -0.4651843 | -9.6126678 | 8.6822992 | 4.6133816 | 105 | -0.1008337 | 0.92 | |
| Trophic_position: insectivore | 1.0153286 | -2.0846131 | 4.1152703 | 1.5702386 | 168 | 0.6466078 | 0.519 | |
| Trophic_position: carnivore | 0.9644588 | -2.2552882 | 4.1842057 | 1.6309245 | 168 | 0.5913571 | 0.555 | |
| log_range_area_km2 | 0.7555783 | 0.5142037 | 0.9969530 | 0.1222654 | 168 | 6.1798190 | <0.001 | *** |
| metric: lt | -1.6329004 | -4.6925464 | 1.4267455 | 1.5498272 | 168 | -1.0536016 | 0.294 | |
| abs_lat_mp:realmIntertidal | 0.1033910 | -0.1548866 | 0.3616685 | 0.1308274 | 168 | 0.7902853 | 0.43 | |
| abs_lat_mp:realmMarine | 0.2065461 | 0.0986104 | 0.3144819 | 0.0546736 | 168 | 3.7778060 | <0.001 | *** |
| realmIntertidal:log_range_area_km2 | -1.3636071 | -4.4269757 | 1.6997615 | 1.5517129 | 168 | -0.8787754 | 0.381 | |
| realmMarine:log_range_area_km2 | 0.6982404 | -0.2540206 | 1.6505014 | 0.4823565 | 168 | 1.4475609 | 0.15 |
Results:
- Warm niche underfilling increases with latitudinal midpoint of the realized range in terrestrial and intertidal systems, decreases with latitude in marine systems
- Largest effect size for terrestrial species: ~ -0.16 degrees C per degree latitude
- Not significant for intertidal species: ~ -0.06 degrees C per degree latitude
- Marine species: ~0.04 degrees C per degree latitude
- Warm niche underfilling decreases with increasing realized range in terrestrial and marine systems, increases in intertidal systems
- Not significant for marine species: ~ 1.45 degrees C per log(km2)
- Terrestrial species: ~0.76 degrees C per log(km2)
- Not significant for intertidal species: ~0.61 degrees C per log(km2)
- Warm niche filling is not significantly different for critical versus lethal limits
- Species that undergo hot season dormancy have more warm niche underfilling (not significant)
- Warm niche underfilling decreases with increasing body length, but not significantly
- ~ 0.17 degrees C per log(cm)
- Effect of trophic position not significant, but warm niche underfilling is greatest in: omnivore, insectivore, carnivore, herbivore, fungivore
Sensitivity analysis: acclimatization
Cold niche filling
filling_value ~ abs_lat_mp*realm + metric
random = ~1|Class/Order/Family/Genus
Subset without acclimation adjustment:
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | -7.1684912 | -19.3468117 | 5.0098292 | 6.0903065 | 61 | -1.1770329 | 0.244 | |
| abs_lat_mp | 0.3723804 | 0.2203590 | 0.5244018 | 0.0760250 | 61 | 4.8981314 | <0.001 | *** |
| realm: Intertidal | -17.2392861 | -47.8995205 | 13.4209482 | 11.9273520 | 5 | -1.4453574 | 0.208 | |
| realm: Marine | 7.5567011 | -23.0519651 | 38.1653673 | 11.9072911 | 5 | 0.6346281 | 0.554 | |
| metric: lt | 0.8290770 | -6.5527062 | 8.2108602 | 3.6092686 | 29 | 0.2297078 | 0.82 | |
| abs_lat_mp:realmIntertidal | 0.2769613 | -0.1317900 | 0.6857127 | 0.2044141 | 61 | 1.3549028 | 0.18 | |
| abs_lat_mp:realmMarine | -0.3636750 | -0.6962535 | -0.0310965 | 0.1626118 | 29 | -2.2364613 | 0.033 | * |
Subset with acclimation adjustment:
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | -8.3886704 | -21.8470961 | 5.0697553 | 6.7304796 | 61 | -1.2463704 | 0.217 | |
| abs_lat_mp | 0.3595773 | 0.2030725 | 0.5160820 | 0.0782671 | 61 | 4.5942326 | <0.001 | *** |
| realm: Intertidal | -24.0828520 | -57.2925670 | 9.1268630 | 12.9191433 | 5 | -1.8641214 | 0.121 | |
| realm: Marine | 4.2386904 | -30.2207542 | 38.6981350 | 13.4053093 | 5 | 0.3161949 | 0.765 | |
| metric: lt | 0.3824132 | -7.4539678 | 8.2187943 | 3.8315409 | 29 | 0.0998066 | 0.921 | |
| abs_lat_mp:realmIntertidal | 0.3805119 | -0.0423870 | 0.8034107 | 0.2114892 | 61 | 1.7992021 | 0.077 | . |
| abs_lat_mp:realmMarine | -0.2754273 | -0.6240850 | 0.0732304 | 0.1704736 | 29 | -1.6156594 | 0.117 |
Warm niche filling
filling_value ~ abs_lat_mp*realm + metric
random = ~1|Class/Order/Family/Genus
Subset without acclimation adjustment:
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | 3.7747406 | 0.1475452 | 7.4019359 | 1.8239856 | 84 | 2.0695013 | 0.042 | * |
| abs_lat_mp | -0.2264156 | -0.2959904 | -0.1568407 | 0.0349867 | 84 | -6.4714795 | <0.001 | *** |
| realm: Intertidal | -8.7049371 | -17.5598866 | 0.1500124 | 4.4528344 | 84 | -1.9549205 | 0.054 | . |
| realm: Marine | -9.5709003 | -14.6811103 | -4.4606902 | 2.5697401 | 84 | -3.7244623 | <0.001 | *** |
| metric: lt | 0.9873362 | -1.9178853 | 3.8925577 | 1.4464207 | 50 | 0.6826065 | 0.498 | |
| abs_lat_mp:realmIntertidal | 0.2425043 | 0.0450168 | 0.4399918 | 0.0993094 | 84 | 2.4419079 | 0.017 | * |
| abs_lat_mp:realmMarine | 0.2310607 | 0.1235864 | 0.3385350 | 0.0540449 | 84 | 4.2753429 | <0.001 | *** |
Subset with acclimation adjustment:
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | 0.0114807 | -4.1046982 | 4.1276596 | 2.0698777 | 84 | 0.0055465 | 0.996 | |
| abs_lat_mp | -0.1865879 | -0.2574396 | -0.1157361 | 0.0356288 | 84 | -5.2369975 | <0.001 | *** |
| realm: Intertidal | -0.9371823 | -10.1219189 | 8.2475542 | 4.6186724 | 84 | -0.2029116 | 0.84 | |
| realm: Marine | -4.2157528 | -9.6811897 | 1.2496841 | 2.7483709 | 84 | -1.5339097 | 0.129 | |
| metric: lt | -0.8799398 | -4.1645794 | 2.4046998 | 1.6353214 | 50 | -0.5380837 | 0.593 | |
| abs_lat_mp:realmIntertidal | 0.0793966 | -0.1201735 | 0.2789667 | 0.1003566 | 84 | 0.7911447 | 0.431 | |
| abs_lat_mp:realmMarine | 0.1682771 | 0.0565719 | 0.2799823 | 0.0561725 | 84 | 2.9957185 | 0.004 | ** |
Sensitivity analysis: preferred temperatures
Warm niche filling
filling_value ~ abs_lat_mp + metric
random = ~1|Class/Order/Family/Genus
Subset without Tpref/Tb adjustment:
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | 3.2526418 | 0.5188066 | 5.9864769 | 1.3806552 | 119 | 2.3558683 | 0.02 | * |
| abs_lat_mp | -0.1764834 | -0.2485732 | -0.1043935 | 0.0364072 | 119 | -4.8474863 | <0.001 | *** |
| metric: lt | 1.1540592 | -6.6309450 | 8.9390633 | 3.9316220 | 119 | 0.2935326 | 0.77 |
Subset with Tpref/Tb adjustment:
| Value | CI (lower) | CI (upper) | Std.Error | DF | t-value | p-value | ||
|---|---|---|---|---|---|---|---|---|
| (Intercept) | 1.4928324 | -0.4729743 | 3.4586392 | 0.9927816 | 119 | 1.5036866 | 0.135 | |
| abs_lat_mp | -0.0974660 | -0.1482081 | -0.0467239 | 0.0256260 | 119 | -3.8033995 | <0.001 | *** |
| metric: lt | 0.7691801 | -4.5643622 | 6.1027225 | 2.6935724 | 119 | 0.2855613 | 0.776 |
Range filling
underfilling ~ abs_lat_mp + realm + metric
random = ~1|Class/Order/Family/Genus
## Warning: Some predictor variables are on very different scales: consider
## rescaling## Warning in eval(family$initialize, rho): non-integer #successes in a
## binomial glm!## boundary (singular) fit: see ?isSingular## Warning in vcov.merMod(object, use.hessian = use.hessian): variance-covariance matrix computed from finite-difference Hessian is
## not positive definite or contains NA values: falling back to var-cov estimated from RX## Warning in vcov.merMod(object, correlation = correlation, sigm = sig): variance-covariance matrix computed from finite-difference Hessian is
## not positive definite or contains NA values: falling back to var-cov estimated from RX## Generalized linear mixed model fit by maximum likelihood (Laplace
## Approximation) [glmerMod]
## Family: binomial ( logit )
## Formula: D_under ~ abs_lat_mp + range_area_km2 + realm + metric + (1 |
## Genus)
## Data: under
## Control: glmerControl(tolPwrss = 0.001)
##
## AIC BIC logLik deviance df.resid
## 121.5 147.4 -52.8 105.5 178
##
## Scaled residuals:
## Min 1Q Median 3Q Max
## -11407870 -1 0 0 375
##
## Random effects:
## Groups Name Variance Std.Dev.
## Genus (Intercept) 0 0
## Number of obs: 186, groups: Genus, 98
##
## Fixed effects:
## Estimate Std. Error z value Pr(>|z|)
## (Intercept) 2.982e+01 1.541e+06 0.000 1.000
## abs_lat_mp -2.810e-04 2.840e-02 -0.010 0.992
## range_area_km2 -1.220e-06 2.435e-07 -5.012 5.38e-07 ***
## realmMarine 1.443e+02 1.915e+07 0.000 1.000
## realmTerrestrial -2.725e+01 1.541e+06 0.000 1.000
## metriclt 3.554e+02 1.828e+07 0.000 1.000
## metricmixed 1.787e+00 1.262e+00 1.416 0.157
## ---
## Signif. codes: 0 '***' 0.001 '**' 0.01 '*' 0.05 '.' 0.1 ' ' 1
##
## Correlation of Fixed Effects:
## (Intr) abs_l_ rng__2 rlmMrn rlmTrr mtrclt
## abs_lat_mp 0.000
## range_r_km2 0.000 0.026
## realmMarine -0.056 0.000 0.000
## relmTrrstrl -1.000 0.000 0.000 0.056
## metriclt -0.062 0.000 0.000 -0.388 0.062
## metricmixed 0.000 -0.484 -0.140 0.000 0.000 0.000
## fit warnings:
## Some predictor variables are on very different scales: consider rescaling
## convergence code: 0
## boundary (singular) fit: see ?isSingular
underfilling ~ abs_lat_mp + range_area_km2 + realm + dispersal_distance_continuous + dispersal_ability_category + maximum_body_size_SVL_HBL_cm_ + hot_season_dormancy_ + cold_season_dormancy_ + Trophic_position + metric
random = ~1|Class/Order/Family/Genus