Climate-driven declines in arthropod abundance restructure a rainforest food web

May combine two claims.

Geotrygon montana showed no decline in abundance between 1990 and 2015 at El Verde, while Todus mexicanus experienced a 90% reduction in catch rate.

“For example, the ruddy quail dove (Geotrygon montana), which eats grains and fruits almost exclusively, showed no decline in abundance between 1990 and 2015 at El Verde, while Todus mexicanus, the Puerto Rican tody, whose diet is composed entirely of insects (26), experienced a 90% reduction in catch rate.”

• cites [26] Reagan D Waide R 1996 The Food Web of a Tropical Rain Forest Univ of Chicago Press ChicagoNot checked yet

  Not checked in the free run.

• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.

May combine two claims.

From a single species of walking stick to a community of over 120 canopy taxa, long-term declines have occurred in parallel with rising temperatures.

“First, from a single species of walking stick to a community of over 120 canopy taxa, long-term declines have occurred in parallel with rising temperatures.”

• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.
• AssumptionRests on an assumption: The meteorological stations at El Verde and Bisley adequately represent the temperature exposure of the arthropod and insectivore populations studied.

May combine two claims.

While average temperatures increased 8-10% over the same period, the proportion of extreme temperatures increased 14 times.

“While average temperatures increased 8–10% over the same period, the proportion of extreme temperatures increased 14 times.”

• AssumptionRests on an assumption: The meteorological stations at El Verde and Bisley adequately represent the temperature exposure of the arthropod and insectivore populations studied.

May combine two claims.

Inclusion of mean daily rainfall as a covariate improved the unanimity index for all analyses except for walking sticks, which had a UI of 100 without the covariate, and for the E. coqui at the Sonadura Old study site, which retained a low UI and indeterminate causal path with and without the covariate.

“Inclusion of mean daily rainfall as a covariate improved the UI for all analyses except for walking sticks, which had a UI of 100 without the covariate, and for the E. coqui at the Sonadura Old study site, which retained a low UI and indeterminate causal path with and without the covariate.”

May combine two claims.

Across all six populations, the average independent effect of MnMaxT on abundance was 25%, while the averages for four other temperature-related variables ranged from 11.4 to 14.3%.

“Across all six populations, the average independent effect of MnMaxT on abundance was 25%, while the averages for four other temperature related variables ranged from 11.4 to 14.3%.”

May combine two claims.

Quasi-Poisson regressions of walking stick abundance against time and against MnMaxT during the census periods were both negative and significant.

“Employing Willig et al.’s (24) census data, we also conducted quasi-Poisson regressions of walking stick abundance against time and against MnMaxT during the census periods (Fig. 5 C and D). Both regressions were negative and significant.”

• cites [24] Willig MR Presley SJ Bloch CP 2011 Long-term dynamics of tropical walking sticks in response to multiple large-scale and intense disturbances Oecologia 165 357 368 20677015 sourceNot checked yet

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May combine two claims.

Canopy sticky-trap catch rates declined from 37 mg per trap per day in July 1976 to 5 mg per trap per day in July 2012, and from 21 mg per trap per day in January 1976 to 8 mg per trap per day in January 2013.

“In July 1976, the canopy trap catch rate was 37 mg per trap per day compared with 5 mg per trap per day in July 2012, and 21 mg per trap per day in January 1976 vs. 8 mg per trap per day in January 2013.”

• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.

May combine two claims.

In Woolbright's El Verde E. coqui study area, the population had a significant negative trend in phase 1, a rapid increase from 6 to 57 individuals between January and October 1990, and then a steady decline in phase 3 at about the same rate as phase 1 until the census ended 6 y later.

“In the first phase, E. coqui numbers had a significant negative trend followed by a rapid increase during phase 2 from 6 to 57 individuals between January and October 1990, most likely due to immigration from surrounding areas following Hurricane Hugo. In phase 3, the population resumed a steady decline at about the same rate as phase 1, until the census ended 6 y later.”

• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.

May combine two claims.

Between 1978 and 2015, mean maximum yearly temperature at El Verde rose by 2.0 °C at an average rate of 0.050 °C per year.

“Between 1978 and 2015, MnMaxT at El Verde rose by 2.0 °C at an average rate of 0.050 °C per year.”

• AssumptionRests on an assumption: The meteorological stations at El Verde and Bisley adequately represent the temperature exposure of the arthropod and insectivore populations studied.

May combine two claims.

The number of dry days at El Verde has been decreasing since 1975, while the variation in daily rainfall has been increasing.

“However, the number of dry days (rainfall less than 1 mm) has been decreasing since 1975 (SI Appendix, Fig. S8G), while the variation in daily rainfall has been increasing (SI Appendix, Fig. S8H).”

• AssumptionRests on an assumption: The meteorological stations at El Verde and Bisley adequately represent the temperature exposure of the arthropod and insectivore populations studied.

May combine two claims.

Quasi-Poisson regressions of E. coqui numbers against time and MnMaxT for Stewart's Activity Transect were both negative and significant.

“The quasi-Poisson regressions of E. coqui numbers vs. time and MnMaxT for Stewart’s Activity Transect were both negative and significant (Fig. 7 A and B) (28).”

• cites [28] Stewart MM 1995 Climate driven population fluctuations in rain forest frogs J Herpetol 29 437 446 sourceNot checked yet

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• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.

May combine two claims.

For the Sonadura Old E. coqui population, only the regression of numbers on time was significant (P = 0.013), while the regression of abundance on temperature was marginally insignificant (P = 0.067).

“For the Sonadura Old population, only the regression of numbers on time was significant (P = 0.013), while the regression of abundance on temperature was marginally insignificant (P = 0.067).”

• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.

May combine two claims.

Arthropod biomass for July 2011-2012 and January 2012-2013 fell four and eight times, respectively, compared with earlier samples.

“Compared with the earlier samples, arthropod biomass for July 2011–2012 and January 2012–2013 fell four and eight times, respectively.”

• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.

May combine two claims.

Regressions of E. coqui numbers against time and temperature for the Sonadura West study area were both negative and significant.

“The regressions of E. coqui numbers vs. time and temperature for the Sonadura West study area were also negative and significant (SI Appendix, Fig. S3A).”

• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.

May combine two claims.

Between 1993 and 2015, the rate of temperature increase at Bisley Tower was 0.055 °C per year, not significantly different from the rate at El Verde.

“Between 1993 and 2015, the rate of temperature increase at Bisley Tower was 0.055 °C per year, not significantly different from the rate at El Verde.”

• AssumptionRests on an assumption: The meteorological stations at El Verde and Bisley adequately represent the temperature exposure of the arthropod and insectivore populations studied.

May combine two claims.

The density of Anolis gundlachi dropped twice, and the density of Anolis evermanni declined 10 times.

“The density of A. gundlachi, the most common species, had dropped twice, and the density of A. evermanni declined 10 times.”

• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.

May combine two claims.

From 1975 to 2015, rainfall at the El Verde station averaged 3,653 mm/y with no significant positive or negative trends.

“From 1975 to 2015, rainfall at the El Verde station averaged 3,653 mm/y with no significant positive or negative trends.”

• AssumptionRests on an assumption: The meteorological stations at El Verde and Bisley adequately represent the temperature exposure of the arthropod and insectivore populations studied.

May combine two claims.

Regressions of E. coqui numbers on time for the Bisley Stream and Bisley forest study areas were both negative and significant.

“Regressions on time were both negative and significant (SI Appendix, Fig. S3 C and D).”

• AssumptionRests on an assumption: The historical data from the 1970s and other earlier censuses are comparable to the authors’ later samples despite the time gap and any differences in observers or methods.

May combine two claims.

The unanimity indices equaled 100 in five of the six groups, indicating that the causal path Temperature → Abundance is supported by all three of Vinod's criteria for causality.

“The unanimity indices equaled 100 in five of the six groups, indicating that the causal path Temperature → Abundance is supported by all three of Vinod’s criteria for causality.”

• AssumptionRests on an assumption: The statistical associations between temperature and abundance are not confounded by unmeasured variables that could produce the observed relationships.

May combine two claims.

The proportion of temperatures greater than 29 °C had both negative (El Verde E. coqui) and positive (walking sticks, E. coqui at Sonadura West and Sonadura Old) effects.

“the proportion of temperatures greater than 29 °C had both negative (El Verde E. coqui) and positive (walking sticks, E. coqui at Sonadura West and Sonadura Old) effects (SI Appendix, Table S5).”

• cites SI Appendix, Table S5Not checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

The canopy and walking stick censuses spanned three La Niña and three El Niño episodes.

“the canopy and walking stick censuses spanned three La Niña and three El Niño episodes.”

• cites SI Appendix, Fig. S9Not checked yet

  Not checked in the free run.

Classified as probably not a substantial claim. May combine two claims.

Tropical species that evolved in comparatively aseasonal environments should have narrower thermal niches, reduced acclimation to temperature fluctuations, and exist at or near their thermal optima.

“As Janzen (3) pointed out, tropical species that evolved in comparatively aseasonal environments should have narrower thermal niches, reduced acclimation to temperature fluctuations, and exist at or near their thermal optima.”

• cites [3] Janzen D 1967 Why mountain passes are higher in the tropics Am Nat 101 233 249 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

Stewart and Woolbright estimated that the E. coqui population alone harvested 114,000 prey items per hectare per night in the Luquillo forest.

“Stewart and Woolbright (47), for example, estimated the E. coqui population alone harvested 114,000 prey items per hectare per night in the Luquillo forest, and Reagan and Waide (26) calculated that A. stratulus, A. gundlachi, and A. evermanni collectively consumed ∼443,300 invertebrates per hectare per day at El Verde.”

• cites [47] Stewart M Woolbright L 1996 Amphibians The Food Web of a Tropical Rain Forest Reagan DP Waide RB Univ of Chicago Press Chicago 363 398Not checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

Deutsch et al. predicted that tropical insects should decline by as much as 20% under climate warming.

“Deutsch et al. (5) predicted that, for insects living at mid-to-high latitudes, rates of increase should grow as climate warms, while in the tropics insects should decline by as much as 20%.”

• cites [5] Deutsch CA 2008 Impacts of climate warming on terrestrial ectotherms across latitude Proc Natl Acad Sci USA 105 6668 6672 18458348 PMC2373333 sourceNot checked yet

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May combine two claims.

The counter cascade of arthropod declines despite predator decreases conflicts with decades of research on trophic dynamics and with studies documenting the major impact of anoles, frogs, and birds on arthropod abundance.

“This “counter cascade” conflicts with decades of research on trophic dynamics (44), as well as studies documenting the major impact of anoles, frogs, and birds on arthropod abundance (45, 46).”

• cites [44] Terborgh J Estes J 2010 Trophic Cascades: Predators, Prey, and the Changing Dynamics of Nature Island Press Washington, DC 2nd EdNot checked yet

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• cites [45] Pacala S Roughgarden J 1984 Control of arthropod abundance by Anolis lizards on St. Eustatius (Neth. Antilles) Oecologia 64 160 162 28312332 sourceNot checked yet

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• cites [46] Marquis R Whelan C 1994 Insectivorous birds increase growth of white oak through consumption of leaf-chewing insects Ecology 75 2007 2014 sourceNot checked yet

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May combine two claims.

Positive SOI values (La Niña episodes) have an immediate positive impact on the abundance of canopy invertebrates, and negative SOI values (El Niño episodes) have an immediate negative impact.

“This indicates that positive SOI values (La Niña episodes) have an immediate positive impact on the abundance of canopy invertebrates, and negative SOI values (El Niño episodes) have an immediate negative impact.”

Probably not a substantial claim: may not pass S3 (load-bearing).

Predictions of tropical ectotherm vulnerability have been verified in a variety of tropical reptiles, amphibians, and invertebrates.

“These predictions have been verified in a variety of tropical reptiles, amphibians, and invertebrates (4–8).”

• cites [4] Whitfield SM 2007 Amphibian and reptile declines over 35 years at La Selva, Costa Rica Proc Natl Acad Sci USA 104 8352 8356 17449638 PMC1895953 sourceNot checked yet

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• cites [5] Deutsch CA 2008 Impacts of climate warming on terrestrial ectotherms across latitude Proc Natl Acad Sci USA 105 6668 6672 18458348 PMC2373333 sourceNot checked yet

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• cites [6] Huey RB 2009 Why tropical forest lizards are vulnerable to climate warming Proc Biol Sci 276 1939 1948 19324762 PMC2677251 sourceNot checked yet

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• cites [7] Sinervo B 2010 Erosion of lizard diversity by climate change and altered thermal niches Science 328 894 899 20466932 sourceNot checked yet

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• cites [8] Sunday J Bates A Dulvy N 2011 Global analysis of thermal tolerance and latitude in ectotherms Proc Biol Sci 278 1823 1830 21106582 PMC3097822 sourceNot checked yet

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May combine two claims.

The simultaneous declines of all 10 arthropod taxa in the sweep samples, the 10 most-abundant taxa in Schowalter's canopy samples, and arthropods occupying all strata point to an overriding environmental factor that has had ubiquitous, adverse effects on forest arthropods regardless of taxonomic affiliation, stratum occupied, or type of niche exploited.

“Second, the simultaneous declines of all 10 arthropod taxa in our sweep samples, the 10 most-abundant taxa in Schowalter’s (23) canopy samples, and arthropods occupying all strata of our forest study area, all point to an overriding environmental factor that has had ubiquitous, adverse effects on forest arthropods regardless of taxonomic affiliation, stratum occupied, or type of niche exploited.”

• cites [23] Schowalter TD 1994 Invertebrate community structure and herbivory in a tropical rain forest canopy in Puerto Rico following Hurricane Hugo Biotropica 26 312 319 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

Reagan and Waide calculated that A. stratulus, A. gundlachi, and A. evermanni collectively consumed approximately 443,300 invertebrates per hectare per day at El Verde.

“Stewart and Woolbright (47), for example, estimated the E. coqui population alone harvested 114,000 prey items per hectare per night in the Luquillo forest, and Reagan and Waide (26) calculated that A. stratulus, A. gundlachi, and A. evermanni collectively consumed ∼443,300 invertebrates per hectare per day at El Verde.”

• cites [26] Reagan D Waide R 1996 The Food Web of a Tropical Rain Forest Univ of Chicago Press ChicagoNot checked yet

  Not checked in the free run.

May combine two claims.

Climate warming has emerged as a dominant force affecting arthropod populations in the Luquillo forest and, most likely, in other tropical forests experiencing significant increases in ambient temperature.

“We suggest that climate warming has emerged as such a dominant force affecting arthropod populations in the Luquillo forest and, most likely, in other tropical forests experiencing significant increases in ambient temperature.”

• AssumptionRests on an assumption: The statistical associations between temperature and abundance are not confounded by unmeasured variables that could produce the observed relationships.

May combine two claims.

Hallman et al.'s statistical models largely eliminated land-use change as an explanatory variable but did not thoroughly analyze climate-change variables.

“Their statistical models largely eliminated land-use change as an explanatory variable but did not thoroughly analyze a number of climate-change variables.”

• cites [61] Hallmann CA 2017 More than 75 percent decline over 27 years in total flying insect biomass in protected areas PLoS One 12 e0185809 29045418 PMC5646769 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Decreases in Puerto Rico's Eleutherodactylus populations began in the 1970s, with three species classified as threatened by 1985.

“Decreases in Puerto Rico’s Eleutherodactylus populations began in the 1970s, with three species classified as threatened by 1985 (64).”

• cites [64] Joglar R Burrowes P 1996 Declining amphibian populations in Puerto Rico. Contributions to West Indian Herpetology: A Tribute to Albert Schwartz, Contributions to Herpetology, eds Powell R, Henderson RW (The Society for the Study of Amphibians and Reptiles, Ithaca, NY), Vol 12, pp 371–380Not checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

Increases in average temperature had a positive influence on insect biomass in Hallman et al.'s analysis.

“Increases in average temperature had a positive influence on insect biomass, an effect predicted by Tewksbury et al. (62) for temperate insects.”

• cites [61] Hallmann CA 2017 More than 75 percent decline over 27 years in total flying insect biomass in protected areas PLoS One 12 e0185809 29045418 PMC5646769 sourceNot checked yet

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• cites [62] Tewksbury JJ Huey RB Deutsch CA 2008 Ecology. Putting the heat on tropical animals Science 320 1296 1297 18535231 sourceNot checked yet

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Classified as probably not a substantial claim.

Huey et al. proposed that thermal-conforming lizards inhabiting shaded tropical forests should be especially vulnerable to climate warming.

“Huey et al. (6) proposed that thermal-conforming lizards inhabiting shaded tropical forests should be especially vulnerable to climate warming.”

• cites [6] Huey RB 2009 Why tropical forest lizards are vulnerable to climate warming Proc Biol Sci 276 1939 1948 19324762 PMC2677251 sourceNot checked yet

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May combine two claims.

At Monteverde, Costa Rica, Anolis tropidolepis and Anolis altae populations were diminishing by 4.4% and 3.1% per year, respectively, for 11 and 13 y before both species became locally extinct.

“At Monteverde, Costa Rica, populations of Anolis tropidolepis and Anolis altae were diminishing by 4.4% and 3.1% per year, respectively, for 11 and 13 y before both species became locally extinct.”

• cites [63] Pounds J Fogden M Campbell J 1999 Biological response to climate change on a tropical mountain Nature 398 611 615 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

Sinervo et al. reached broadly similar conclusions for baskers and thermal-conformers in temperate and tropical habitats.

“In an encompassing study spanning local extinctions of lizard populations on five continents, Sinervo et al. (7) reached broadly similar conclusions for baskers and thermal-conformers in temperate and tropical habitats.”

• cites [7] Sinervo B 2010 Erosion of lizard diversity by climate change and altered thermal niches Science 328 894 899 20466932 sourceNot checked yet

  Not checked in the free run.

May combine two claims.

This vulnerability results from lower critical thermal maxima than basking species, limited acclimation via thermoregulation, and reduced thermal safety margins.

“This vulnerability results from lower critical thermal maxima than basking species, limited acclimation via thermoregulation, and reduced thermal safety margins.”

• cites [6] Huey RB 2009 Why tropical forest lizards are vulnerable to climate warming Proc Biol Sci 276 1939 1948 19324762 PMC2677251 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

Other studies have documented declines in arthropod abundance across a broad range of geography, habitats, and taxa.

“Other studies have also documented declines in arthropod abundance across a broad range of geography, habitats, and taxa (10–12, 54, 55, 56–58).”

• cites [10] Conrad M Warren B Fox R Parsons M Woiwoda I 2006 Rapid declines of common, widespread British moths provide evidence of an insect biodiversity crisis Biol Conserv 132 279 291 sourceNot checked yet

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• cites [11] Forister ML 2010 Compounded effects of climate change and habitat alteration shift patterns of butterfly diversity Proc Natl Acad Sci USA 107 2088 2092 20133854 PMC2836664 sourceNot checked yet

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• cites [12] Brooks D 2012 Large carabid beetle declines in a United Kingdom monitoring network increases evidence for a widespread loss in insect biodiversity J Appl Ecol 49 1009 1019 sourceNot checked yet

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• cites [54] Kotze DJ O’Hara RB 2003 Species decline—But why? Explanations of carabid beetle (Coleoptera, Carabidae) declines in Europe Oecologia 135 138 148 12647113 sourceNot checked yet

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• cites [55] Thomas JA 2004 Comparative losses of British butterflies, birds, and plants and the global extinction crisis Science 303 1879 1881 15031508 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

Ongoing losses of anuran biodiversity on Puerto Rico and other West Indian islands are part of a global amphibian decline that has continued largely unabated since at least the 1970s.

“The ongoing losses of anuran biodiversity on Puerto Rico and other West Indian islands (66, 67) are part of a global amphibian decline that has continued largely unabated since at least the 1970s (68–72).”

• cites [66] Burrowes P Joglar R Green D 2004 Potential causes for amphibian declines in Puerto Rico Herpetologica 60 141 154 sourceNot checked yet

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• cites [67] Hedges S 1993 Global amphibian declines: A perspective from the Caribbean Biodivers Conserv 2 290 303 sourceNot checked yet

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• cites [68] Pounds JA 1990 Disappearing gold BBC Wildlife 8 812 817Not checked yet

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• cites [69] Pounds JA 1991 Amphibian watch: New clues in the case of the disappearing amphibians Wildlife Conservation 94 16 18Not checked yet

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• cites [70] Barinaga M 1990 Where have all the froggies gone? Science 247 1033 1034 17800052 sourceNot checked yet

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• cites [71] Crawford AJ Lips KR Bermingham E 2010 Epidemic disease decimates amphibian abundance, species diversity, and evolutionary history in the highlands of central Panama Proc Natl Acad Sci USA 107 13777 13782 20643927 PMC2922291 sourceNot checked yet

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• cites [72] Pounds JA Fogden MPL Savage J Gorman G 1997 Tests of null models for amphibian declines on a tropical mountain Conserv Biol 11 1307 1322 sourceNot checked yet

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Probably not a substantial claim: may not pass S4 (based).

The authors propose that reduced consumption and declines in abundance result directly from climate-driven declines in food abundance.

“while we propose that reduced consumption and declines in abundance result directly from climate-driven declines in food abundance.”

May combine two claims.

As climate warms, lizards should increasingly become nutritionally challenged, resulting in reduced body condition, growth rates, egg production, fat body mass, physical performance, and immune system response.

“As climate warms, lizards should increasingly become nutritionally challenged, resulting in reduced body condition, growth rates, egg production, fat body mass, physical performance, and immune system response.”

Probably not a substantial claim: may not pass S3 (load-bearing).

Pounds and coworkers have presented a wide-ranging body of evidence indicating that climate warming is a central factor promoting outbreaks of Batrachochytrium and the mass die-offs that follow.

“Pounds and coworkers (63, 73, 77–79) have presented a wide-ranging body of evidence indicating that climate warming is a central factor promoting outbreaks of Batrachochytrium and the mass die-offs that follow.”

• cites [63] Pounds J Fogden M Campbell J 1999 Biological response to climate change on a tropical mountain Nature 398 611 615 sourceNot checked yet

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• cites [73] Pounds JA 2006 Widespread amphibian extinctions from epidemic disease driven by global warming Nature 439 161 167 16407945 sourceNot checked yet

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• cites [77] Pounds JA Crump ML 1994 Amphibian declines and climate disturbance: The case of the golden toad and the harlequin frog Conserv Biol 8 72 85 sourceNot checked yet

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• cites [78] Pounds JA Carnaval AC Puschendorf R Haddad CFB Masters KL 2006 Responding to amphibian loss Science 314 1541 1542 17158306 sourceNot checked yet

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• cites [79] Pounds JA 2001 Climate and amphibian declines Nature 410 639 640 11287931 sourceNot checked yet

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May combine two claims.

B. dendrobatidis has been detected in E. coqui populations at higher elevations in Puerto Rico, but the fungus is restricted to areas above 600 m.

“While B. dendrobatidis has been detected in E. coqui populations at higher elevations in Puerto Rico (66, 75), the fungus is restricted to areas above 600 m (76)”

• cites [66] Burrowes P Joglar R Green D 2004 Potential causes for amphibian declines in Puerto Rico Herpetologica 60 141 154 sourceNot checked yet

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• cites [75] Burrowes P Longo A Joglar R 2008 Geographic distribution of Batrachochytrium dendrobatidis in Puerto Rico Herpetol Rev 39 321 324Not checked yet

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• cites [76] Longo AV Burrowes PA Joglar RL 2010 Seasonality of Batrachochytrium dendrobatidis infection in direct-developing frogs suggests a mechanism for persistence Dis Aquat Organ 92 253 260 21268989 sourceNot checked yet

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May combine two claims.

Death rates from starvation, predation, and disease should increase along with concomitant decreases in reproduction.

“Consequently, death rates from starvation, predation, and disease should increase along with concomitant decreases in reproduction.”

Probably not a substantial claim: may not pass S3 (load-bearing).

Most research has focused on the fungus Batrachochytrium dendrobatidis, which has precipitated mass-mortality events and catastrophic reductions in over 200 frog species world-wide.

“Most research has focused on the fungus Batrachochytrium dendrobatidis, which has precipitated mass-mortality events and catastrophic reductions in over 200 frog species world-wide (63, 73, 74).”

• cites [63] Pounds J Fogden M Campbell J 1999 Biological response to climate change on a tropical mountain Nature 398 611 615 sourceNot checked yet

  Not checked in the free run.

• cites [73] Pounds JA 2006 Widespread amphibian extinctions from epidemic disease driven by global warming Nature 439 161 167 16407945 sourceNot checked yet

  Not checked in the free run.

• cites [74] Catenazzi A 2015 State of the world’s amphibians Annu Rev Environ Resour 40 91 119 sourceNot checked yet

  Not checked in the free run.

May combine two claims.

If climate warming is not driving arthropod declines, then whatever the factors may be, they must completely override the reduction in mortality from lowered predation levels.

“If climate warming is not driving arthropod declines, then whatever the factors may be, they must completely override the reduction in mortality from lowered predation levels.”

Probably not a substantial claim: may not pass S3 (load-bearing).

Hallman et al. found a 75% decline in flying insect biomass between 1989 and 2016 at 63 protected reserves in Germany.

“Overall, they found a 75% decline in biomass between 1989 and 2016.”

• cites [61] Hallmann CA 2017 More than 75 percent decline over 27 years in total flying insect biomass in protected areas PLoS One 12 e0185809 29045418 PMC5646769 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S4 (based).

No studies have explored reductions in arthropod prey as a possible causal mechanism behind amphibian declines.

“To our knowledge, no studies have explored reductions in arthropod prey as a possible causal mechanism.”

Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Currently, 3 of the original 16 species of Puerto Rican Eleutherodactylus are considered extinct, and 13 species are listed as critically endangered, endangered, vulnerable, or near threatened.

“Currently, 3 of the original 16 species of Puerto Rican Eleutherodactylus are considered extinct, and 13 species are listed as critically endangered, endangered, vulnerable, or near threatened (65).”

• cites [65] IUCN 2017 IUCN Red List of Threatened Species. Available at www.iucnredlist.org. Accessed May 25, 2017Not checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

Between 1923 and 1998 in Singapore, insectivores experienced the highest level of extinctions of any foraging guild.

“Between 1923 and 1998 in Singapore, insectivores experienced the highest level of extinctions of any foraging guild (101)”

• cites [101] Castelletta M Sodhi NS Subaraj R 2000 Heavy extinctions of forest avifauna in Singapore: Lessons for biodiversity conservation in Southeast Asia Conserv Biol 14 1870 1880 35701951 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

Over a 20-y period, 54% of bird species native to Britain have also decreased.

“Over a 20-y period, 54% of bird species native to Britain have also decreased (55).”

• cites [55] Thomas JA 2004 Comparative losses of British butterflies, birds, and plants and the global extinction crisis Science 303 1879 1881 15031508 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

In many studies of passerine declines, insectivores are often the group exhibiting the largest decreases in abundance, as appears to be the case in the Luquillo forest.

“We also note that in many studies of passerine declines, such as Mac Nally et al. (98), insectivores are often the group exhibiting the largest decreases in abundance, as appears to be the case in the Luquillo forest.”

• cites [98] Mac Nally R 2009 Collapse of an avifauna: Climate change appears to exacerbate habitat loss and degradation Divers Distrib 15 720 730 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Galapagos finches provide a classic example of how climatic events associated with El Niño can lead to major fluctuations in food supplies, which in turn drive reductions in abundance of up to 40% per year in Geospiza populations.

“Galapagos finches provide a classic example of how climatic events associated with El Niño can lead to major fluctuations in food supplies, which in turn drive reductions in abundance of up to 40% per year in Geospiza populations (108).”

• cites [108] Grant B Grant P 1993 Evolution of Darwin’s finches caused by a rare climatic event Proc R Soc Lond 251 111 117 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

We found only two articles that ascribed the collapse of avian populations to the impact of climate warming on prey abundance.

“We found only two articles that ascribed the collapse of avian populations to the impact of climate warming on prey abundance.”

Probably not a substantial claim: may not pass S3 (load-bearing).

In the Guanica dry forest in southwestern Puerto Rico, mist-net data indicate that dry season bird populations dropped by two-thirds over two decades.

“in the Guanica dry forest in southwestern Puerto Rico mist-net data indicate that dry season bird populations dropped by two-thirds over two decades (86).”

• cites [86] Faaborg J 2013 Long-term decline of a winter-resident bird community in Puerto Rico Biodivers Conserv 22 63 75 sourceNot checked yet

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May combine two claims.

From data gathered over 55 y and 16 locations in Puerto Rico, the National Weather Service found that temperatures during La Niña events averaged ∼2.0 °C lower during the dry season and 1.5 °C lower during the wet season compared with El Niño episodes.

“From data gathered over 55 y and 16 locations in Puerto Rico, the National Weather Service found that temperatures during La Niña events averaged ∼2.0 °C lower during the dry season (December to August), and 1.5 °C lower during the wet season (May to November) compared with El Niño episodes.”

• cites National Weather ServiceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

In general, chytrid epidemics are often associated with repeated warm years and drier weather.

“The relationship between chytrid outbreaks and climate is complex, but in general epidemics are often associated with repeated warm years and drier weather.”

Probably not a substantial claim: may not pass S3 (load-bearing).

In montane habitats, upward migrations into cooler refugia can spell the difference between survival and extinction as temperatures rise at lower elevations.

“In montane habitats upward migrations into cooler refugia can spell the difference between survival and extinction as temperatures rise at lower elevations (119).”

• cites [119] Colwell RK Brehm G Cardelús CL Gilman AC Longino JT 2008 Global warming, elevational range shifts, and lowland biotic attrition in the wet tropics Science 322 258 261 18845754 sourceNot checked yet

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Classified as probably not a substantial claim. May combine two claims.

Based on our literature searches, no studies have explored reductions in arthropod prey as a possible causal mechanism behind amphibian declines, nor the possibility that malnutrition could interact with rising temperature and drought, compounding the impacts of disease through depression of immunocompetence.

“Based on our literature searches, no studies have explored reductions in arthropod prey as a possible causal mechanism behind amphibian declines, nor the possibility that malnutrition could interact with rising temperature and drought, compounding the impacts of disease through depression of immunocompetence (83).”

• cites [83] Blaustein AR 2012 Ecophysiology meets conservation: Understanding the role of disease in amphibian population declines Philos Trans R Soc Lond B Biol Sci 367 1688 1707 22566676 PMC3350657 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Altitudinal shifts can cause rapid changes in community structure, disrupting coevolved interactions, creating "no-analogue communities," and contributing to community disassembly via accelerated extinctions.

“However, altitudinal shifts can also cause rapid changes in community structure, disrupting coevolved interactions, creating “no-analogue communities,” and contributing to community disassembly via accelerated extinctions (120, 121).”

• cites [120] Sheldon KS Yang S Tewksbury JJ 2011 Climate change and community disassembly: Impacts of warming on tropical and temperate montane community structure Ecol Lett 14 1191 1200 21978234 sourceNot checked yet

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• cites [121] Urban MC Tewksbury JJ Sheldon KS 2012 On a collision course: Competition and dispersal differences create no-analogue communities and cause extinctions during climate change Proc Biol Sci 279 2072 2080 22217718 PMC3311897 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

In Canada and the northeastern United States, the total reduction in aerial insectivores between 1970 and 2010 was greater than 60%, the largest decline of all passerines analyzed.

“In Canada and the northeastern United States, the total reduction in aerial insectivores between 1970 and 2010 was greater than 60%, the largest decline of all passerines analyzed (84, 99).”

• cites [84] Berlanga H 2010 Saving Our Shared Birds: Partners in Flight Tri-National Vision for Landbird Conservation Cornell Lab of Ornithology Ithaca, NYNot checked yet

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• cites [99] North American Bird Conservation Initiative Canada 2012 The State of Canada’s Birds, 2012. Available at stateofcanadasbirds.org/. Accessed October 1, 2016Not checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

If these predictions are realized, the impact on food webs and ecosystems will be even more rapid than expected from the steady, linear increases in mean temperatures that are currently ravaging the natural world.

“If these predictions are realized, the impact on food webs and ecosystems will be even more rapid than expected from the steady, linear increases in mean temperatures that are currently ravaging the natural world.”

Probably not a substantial claim: may not pass S3 (load-bearing).

Bird populations in many tropical and temperate ecosystems have been declining in parallel with those in Puerto Rico.

“Bird populations in many tropical and temperate ecosystems have been declining in parallel with those in Puerto Rico.”

May combine two claims.

ENSO impacts rainfall across Puerto Rico, increasing dry season precipitation by 13% during El Niño years and causing a 14% increase in wet season precipitation during La Niña years.

“ENSO also impacts rainfall across Puerto Rico, increasing dry season precipitation by 13% during El Niño years and causing a 14% increase in wet season precipitation during La Niña years.”

• cites National Weather ServiceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

Shifts in altitudinal distributions of arthropods, insectivores, producers, and top predators may also play an important role in the observed declines in abundance.

“While we have focused on population collapse driven by climate change, shifts in altitudinal distributions of arthropods, insectivores, producers, and top predators may also play an important role in the observed declines in abundance.”

Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Diffenbaugh et al. predicted a two- to threefold increase in the probability of historically unprecedented maximum daily temperatures between 2018 and 2036, and a three- to fivefold increase between 2036 and 2055.

“Diffenbaugh et al. (115) predicted a two- to threefold increase in the probability of historically unprecedented maximum daily temperatures between 2018 and 2036, and a three- to fivefold increase between 2036 and 2055.”

• cites [115] Diffenbaugh NS Singh D Mankin JS 2018 Unprecedented climate events: Historical changes, aspirational targets, and national commitments Sci Adv 4 eaao3354 29457133 PMC5812734 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Anomalous rises in seawater temperature associated with ENSO have caused severe reductions in food supplies, leading to starvation and mass mortality of marine birds.

“Anomalous rises in seawater temperature associated with ENSO have also caused severe reductions in food supplies, leading to starvation and mass mortality of marine birds (109–111).”

• cites [109] Piatt J Van Pelt T 1997 Mass-mortality of Guillemots (Uria aalge) in the Gulf of Alaska in 1993 Mar Pollut Bull 34 656 662 sourceNot checked yet

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• cites [110] Ayala L Paz-Soldan L Garate P 2013 A mass mortality event of sooty shearwaters (Puffinus griseus) on the central coast of Peru Notornis 60 258 261 sourceNot checked yet

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• cites [111] Fuentes H Antonietti E 1989 Efectos del Fenómeno El Niño en las aves marinas del litoral peruano Boletín Estudio Regional del Fenómeno El Niño 29 19 26Not checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Ocean warming associated with climate change has precipitated bottom-up trophic cascades that have restructured both benthic and pelagic food webs and driven declines in shearwaters and auklets.

“More recently, ocean warming associated with climate change has precipitated bottom-up trophic cascades, analogous to those proposed here for terrestrial systems, which have restructured both benthic and pelagic food webs and driven declines in shearwaters and auklets (112–114).”

• cites [112] Veit R McGowan J Ainley D Wahland T Pyle P 1997 Apex marine predator declines ninety percent in association with changing oceanic climate Glob Change Biol 3 23 28 sourceNot checked yet

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• cites [113] Baduini C 2001 Mass mortality of short-tailed shearwaters in the south-eastern Bering Sea during summer 1997 Fish Oceanogr 10 117 130 sourceNot checked yet

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• cites [114] Springer A Byrd G Iverson S 2007 Hot oceanography: Planktivorous seabirds reveal ecosystem responses to warming of the Bering Sea Mar Ecol Prog Ser 352 289 297 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

In Canada, 44% of 460 bird species have had negative trends since 1970, and 42 species common to the United States, Canada, and Mexico have lost 50% or more of their populations in 40 y.

“In Canada, 44% of 460 bird species have had negative trends since 1970, and 42 species common to the United States, Canada, and Mexico have lost 50% or more of their populations in 40 y (84).”

• cites [84] Berlanga H 2010 Saving Our Shared Birds: Partners in Flight Tri-National Vision for Landbird Conservation Cornell Lab of Ornithology Ithaca, NYNot checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

Mac Nally et al. attributed the ongoing declines to the impact of increasing temperatures and decreasing rainfall on arthropods and other food resources.

“They attributed the ongoing declines to the impact of increasing temperatures and decreasing rainfall on arthropods and other food resources.”

• cites [98] Mac Nally R 2009 Collapse of an avifauna: Climate change appears to exacerbate habitat loss and degradation Divers Distrib 15 720 730 sourceNot checked yet

  Not checked in the free run.

May combine two claims.

Pearce-Higgins et al. calculated a climate-sensitivity index for prey of 17 upland, insectivorous bird species in the United Kingdom and showed that the index was correlated with recent negative population trends.

“Pearce-Higgins et al. (97) also analyzed the diets of 17 upland, insectivorous bird species in the United Kingdom, calculated a climate-sensitivity index for their prey, and showed that the index was correlated with recent negative population trends.”

• cites [97] Pearce-Higgins J Dennis P Whittingham M Yalden D 2010 Impacts of climate on prey abundance account for fluctuations in a population of a northern wader at the southern edge of its range Glob Change Biol 16 12 23 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

In a lowland tropical rainforest at La Selva, Costa Rica, Sigel et al. found that at least 50% of declining bird species were insectivorous.

“In a lowland tropical rainforest at La Selva, Costa Rica, Sigel et al. (85) also found that at least 50% of declining bird species were insectivorous.”

• cites [85] Sigel BJ Sherry TW Young BE 2006 Avian community response to lowland tropical rainforest isolation: 40 years of change at La Selva Biological Station, Costa Rica Conserv Biol 20 111 121 16909664 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

Stapley et al. reached a similar conclusion concerning the effects of ENSO on Anolis populations in Panama.

“Stapley et al. (20) reached a similar conclusion concerning the effects of ENSO on Anolis populations in Panama.”

• cites [20] Stapley J Garcia M Andrews RM 2015 Long-term data reveal a population decline of the tropical lizard Anolis apletophallus, and a negative affect of El Niño years on population growth rate PLoS One 10 e0115450 25671423 PMC4325001 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Given ongoing climate change, the elevational range of Batrachochytrium will almost certainly change, as well as its impact on Puerto Rican frogs, especially juveniles.

“Given ongoing climate change, the elevational range of Batrachochytrium will almost certainly change, as well as its impact on Puerto Rican frogs, especially juveniles (80).”

• cites [80] Langhammer PF Burrowes PA Lips KR Bryant SB Collins JP 2014 Susceptibility to the amphibian chytrid fungus varies with ontogeny in the direct-developing frog Eleutherodactylus coqui J Wildl Dis 50 438 446 24807186 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Reductions in habitat area as species move up mountains increase extinction rates and can erode genetic diversity, further reducing fitness and response to ongoing changes in climate.

“The reductions in habitat area as species move up mountains also increase extinction rates (87), and can erode genetic diversity, further reducing fitness and response to ongoing changes in climate (122, 123).”

• cites [87] Sekercioglu C Primack R Wormworth J 2012 The effects of climate change on tropical birds Biol Conserv 148 1 18 sourceNot checked yet

  Not checked in the free run.

• cites [122] Arenas M Ray N Currat M Excoffier L 2012 Consequences of range contractions and range shifts on molecular diversity Mol Biol Evol 29 207 218 21778191 sourceNot checked yet

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• cites [123] Rubidge EM 2012 Climate-induced range contraction drives genetic erosion in an alpine mammal Nat Clim Chang 2 285 288 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

Pearce-Higgins et al. showed that trends in golden plover population dynamics were closely associated with temperature-based reductions in prey abundance.

“Pearce-Higgins et al. (97) analyzed the demographics of the golden plover in Europe and showed that trends in population dynamics were closely associated with temperature-based reductions in prey abundance.”

• cites [97] Pearce-Higgins J Dennis P Whittingham M Yalden D 2010 Impacts of climate on prey abundance account for fluctuations in a population of a northern wader at the southern edge of its range Glob Change Biol 16 12 23 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

In the rainforest at La Selva, Costa Rica, 10-20% of resident birds declined between 1960 and 1999.

“In the rainforest at La Selva, Costa Rica, 10–20% of resident birds declined between 1960 and 1999 (85)”

• cites [85] Sigel BJ Sherry TW Young BE 2006 Avian community response to lowland tropical rainforest isolation: 40 years of change at La Selva Biological Station, Costa Rica Conserv Biol 20 111 121 16909664 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

Numerous studies have documented the impact of ENSO on all major taxa considered here, including invertebrates, frogs, lizards, and birds.

“Numerous studies have documented the impact of the ENSO on all of the major taxa considered here, including invertebrates (116), frogs (63, 73, 77, 117), lizards (1, 5), and birds (118).”

• cites [116] Gutiérrez-Fonseca PE Ramírez A Pringle CM 2018 Large-scale climatic phenomena drive fluctuations in macroinvertebrate assemblages in lowland tropical streams, Costa Rica: The importance of ENSO events in determining long-term (15y) patterns PLoS One 13 e0191781 29420548 PMC5805265 sourceNot checked yet

  Not checked in the free run.

• cites [63] Pounds J Fogden M Campbell J 1999 Biological response to climate change on a tropical mountain Nature 398 611 615 sourceNot checked yet

  Not checked in the free run.

• cites [73] Pounds JA 2006 Widespread amphibian extinctions from epidemic disease driven by global warming Nature 439 161 167 16407945 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Nebel et al. concluded that these declines resulted from impoverishment of flying insect populations, and suggested acid rain as a possible mechanism.

“Nebel et al. (100) concluded that these declines resulted from impoverishment of flying insect populations, and suggested acid rain as a possible mechanism.”

• cites [100] Nebel S Mills A McCracken J Taylor P 2010 Declines of aerial insectivores in North America follow a geographic gradient Avian Conserv Ecol 5 1Not checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

An alternative to the climate-warming hypothesis for population declines in Luquillo and other tropical montane habitats is a mechanism proposed by Lawton et al.

“An alternative to the climate-warming hypothesis for population declines in Luquillo and other tropical montane habitats is a mechanism proposed by Lawton et al. (126).”

• cites [126] Lawton RO Nair US Pielke RA Sr Welch RM 2001 Climatic impact of tropical lowland deforestation on nearby montane cloud forests Science 294 584 587 11641496 sourceNot checked yet

  Not checked in the free run.

May combine two claims.

Our analyses provide strong support for the hypothesis that climate warming has been a major factor driving reductions in arthropod abundance, and that these declines have in turn precipitated decreases in forest insectivores in a classic bottom-up cascade.

“Our analyses provide strong support for the hypothesis that climate warming has been a major factor driving reductions in arthropod abundance, and that these declines have in turn precipitated decreases in forest insectivores in a classic bottom-up cascade.”

Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Because all dispersion tests rejected the null hypothesis, only quasi-Poisson regressions were used.

“Because all dispersion tests rejected the null hypothesis, only quasi-Poisson regressions were used.”

• cites [148] Kleiber C Zeileis A 2008 Applied Econometrics with R (Springer, New York), Available at https://cran.r-project.org/web/packages/AER/index.html. Accessed June 1, 2016 sourceNot checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

Climate variables are often highly correlated and can generate spurious results when used in regression.

“Climate variables are often highly correlated and can generate spurious results when used in regression (34, 134).”

• cites [34] Mac Nally R 2002 Multiple regression and inference in ecology and conservation biology: Further comments on identifying important predictor variables Biodivers Conserv 11 1397 1401 sourceNot checked yet

  Not checked in the free run.

• cites [134] Knape J de Valpine P 2011 Effects of weather and climate on the dynamics of animal population time series Proc Biol Sci 278 985 992 20880886 PMC3049023 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

We did not utilize automated regression techniques such as stepwise selection and backward elimination.

“We did not utilize automated regression techniques, such as stepwise selection and backward elimination, due to a variety of issues including erroneous type I error rates and inflated R2 measures (33).”

May combine two claims.

This hypothesis also provides a parsimonious explanation for why similar cross-taxa, concordant decreases in reptiles, anurans, and birds have occurred in Costa Rican rainforests and are likely occurring across a broad range of tropical ecosystems.

“This hypothesis also provides a parsimonious explanation for why similar cross-taxa, concordant decreases in reptiles, anurans, and birds have occurred in Costa Rican rainforests (3, 63), and are likely occurring across a broad range of tropical ecosystems.”

• cites [3] Janzen D 1967 Why mountain passes are higher in the tropics Am Nat 101 233 249 sourceNot checked yet

  Not checked in the free run.

• cites [63] Pounds J Fogden M Campbell J 1999 Biological response to climate change on a tropical mountain Nature 398 611 615 sourceNot checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

All multivariate quasi-Poisson regressions were checked for multicollinearity using the Variance Inflation Factor and Bartlett's Test of Sphericity.

“All multivariate quasi-Poisson regressions were then checked for multicollinearity using the Variance Inflation Factor and Bartlett’s Test of Sphericity.”

Classified as probably not a substantial claim. May combine two claims.

Given that the ruddy quail dove is an exclusive granivore and could confound patterns in insectivore abundance, we excluded this species from counts of captured birds.

“Given that the ruddy quail dove (G. montana) is an exclusive granivore (SI Appendix, Fig. S4) and could confound patterns in insectivore abundance, we excluded this species from counts of captured birds.”

• cites SI Appendix, Fig. S4Not checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

We selected models having the smallest deviance residuals and the lowest χ2 test statistic P values because Akaike's Information Criterion cannot be calculated for quasi-Poisson models.

“Because Akaike’s Information Criterion cannot be calculated for quasi-Poisson models, we selected models having the smallest deviance residuals and the lowest χ2 test statistic P values.”

May combine two claims.

Consequently, the degree of shade, cooling, and humidification of the habitats surrounding the Luquillo Mountains have all increased.

“Consequently, the degree of shade, cooling, and humidification of the habitats surrounding the Luquillo Mountains have all increased (128).”

• cites [128] Otero LM Huey RB Gorman GC 2015 A few meters matter: Local habitats drive reproductive cycles in a tropical lizard Am Nat 186 E72 E80 26655361 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

Since 1998, 8 of 33 Luquillo forest bird species and 10 of 12 resident frog species have undergone elevational shifts.

“Since 1998, 8 of 33 Luquillo forest bird species (124) and 10 of 12 resident frog species (125) have undergone elevational shifts.”

• cites [124] Campos-Cerqueira M Arendt WJ Wunderle JM Jr Aide TM 2017 Have bird distributions shifted along an elevational gradient on a tropical mountain? Ecol Evol 7 9914 9924 29238525 PMC5723601 sourceNot checked yet

  Not checked in the free run.

• cites [125] Campos-Cerqueira M Aide TM 2017 Lowland extirpation of anuran populations on a tropical mountain PeerJ 5 e4059 29158987 PMC5694215 sourceNot checked yet

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Probably not a substantial claim: may not pass S3 (load-bearing).

For Woolbright's census data at the Sonadura Old study area, we removed one outlier for the 1995 census data when E. coqui numbers increased from 29 to 236 and returned to 45 individuals in the next census.

“For Woolbrights’s (29, 30) census data at the Sonadura Old study area, we removed one outlier for the 1995 census data when E. coqui numbers increased from 29 to 236 and returned to 45 individuals in the next census.”

• cites [29] Woolbright L 1991 Impact of Hurricane Hugo on forest frogs in Puerto Rico Biotropica 23 462 467 sourceNot checked yet

  Not checked in the free run.

• cites [30] Woolbright L 1996 Disturbance influences long-term population patterns in the Puerto Rican frog, Eleutherodactylus coqui (Anura: Leptodactylidae) Biotropica 28 493 501 sourceNot checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

We constructed correlation matrices of the HP predictors and eliminated variables having correlations of 0.7 or higher to guard against effects of multicollinearity.

“To guard against such effects of multicollinearity, we constructed correlation matrices of the HP predictors and eliminated variables having correlations of 0.7 or higher.”

Probably not a substantial claim: may not pass S3 (load-bearing).

Pounds et al. have presented a detailed rebuttal of the Lawton scenario with respect to montane forests in Costa Rica.

“Pounds et al. (63) have presented a detailed rebuttal of the Lawton scenario with respect to montane forests in Costa Rica.”

• cites [63] Pounds J Fogden M Campbell J 1999 Biological response to climate change on a tropical mountain Nature 398 611 615 sourceNot checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

Following the same procedures and using the same study area, arthropod abundances were again estimated during July 2011 and January 2012 using both sticky-traps and sweep netting.

“Following the same procedures and using the same study area, arthropod abundances were again estimated during July 2011 and January 2012 using both sticky-traps and sweep netting.”

• cites [22] Lister BC 1981 Seasonal niche relationships of rainforest anoles Ecology 62 1548 1560 sourceNot checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

We conducted our field work in the same study area utilized by Lister in the 1970s.

“We conducted our field work in the same study area utilized by Lister (22) in the 1970s.”

• cites [22] Lister BC 1981 Seasonal niche relationships of rainforest anoles Ecology 62 1548 1560 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

Luquillo forest birds and frogs are known to exhibit elevational range shifts similar to those encountered on other tropical mountains.

“However, Luquillo forest birds and frogs are known to exhibit elevational range shifts similar to those encountered on other tropical mountains.”

• cites [124] Campos-Cerqueira M Arendt WJ Wunderle JM Jr Aide TM 2017 Have bird distributions shifted along an elevational gradient on a tropical mountain? Ecol Evol 7 9914 9924 29238525 PMC5723601 sourceNot checked yet

  Not checked in the free run.

• cites [125] Campos-Cerqueira M Aide TM 2017 Lowland extirpation of anuran populations on a tropical mountain PeerJ 5 e4059 29158987 PMC5694215 sourceNot checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

Hence, we eliminated 2 of the 11 climate variables that had the highest correlations with the other climate variables.

“Hence, we eliminated 2 of the 11 climate variables that had the highest correlations with the other climate variables.”

Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

In theory, clear cutting of lowland forests raises surface temperature, which in turn increases conductive heat flux from exposed ground to the atmosphere, reducing both evapotranspiration and evaporative cooling and lowering humidity.

“In theory, clear cutting of lowland forests raises surface temperature, which in turn increases conductive heat flux from exposed ground to the atmosphere. This reduces both evapotranspiration and evaporative cooling (126) and lowers humidity.”

• cites [126] Lawton RO Nair US Pielke RA Sr Welch RM 2001 Climatic impact of tropical lowland deforestation on nearby montane cloud forests Science 294 584 587 11641496 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

If a given variable did not make at least a 5% independent contribution to the explanation of the deviance, it was not included in the regression analyses.

“If a given variable did not make at least a 5% independent contribution to the explanation of the deviance, it was not included in the regression analyses.”

Classified as probably not a substantial claim.

To compare Anolis densities with Lister's estimates, we sampled anoles within the same 15 × 15-m quadrat during July 2011 and January 2012.

“To compare Anolis densities with Lister’s (22) estimates from July 1976 and January 1977, we sampled anoles within the same 15 × 15-m quadrat during July 2011 and January 2012.”

• cites [22] Lister BC 1981 Seasonal niche relationships of rainforest anoles Ecology 62 1548 1560 sourceNot checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

Recaptures of the same individuals within a given sampling period were eliminated from the analysis.

“Recaptures of the same individuals within a given sampling period were also eliminated from the analysis.”